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B HaCTOHIII.eii pa60Te aBTopOM IICnOJlb30BaH 60Jlbllloii .pm,TIIQe-
CRlIii MaTeplIaJl II 060CHOBHH CBoii B3rJlH]:( Ha .pYHRQIIOH3J1bHYIO 3BO-
JlIOn;IIIO xpOMa.p.pHHflOii CHCTeMbl H ec CTPYRTYPY Y MJleI\OnIIT3IOIII.IIX,
B TOM 'IRCJlC H 'Ie.TIOBel\a. OTp;3BaH ce6e OT'ICT B CJlOiKHOCTll no-
CTaBJICHHOH 33]:(a'IU, aBTOp TeM He MeHee lIa]:(eeTCH, qTO Bce H3-
JlOmeHHOe nOCJJymuT nOBop;OM 1\ ncppCMOTPY YlwpeHIIBIIHlXCH npe]:(-
cTaB.TIeHIIII 0 .pYHI{QIIOH3:JhIlOM 311aQeHllII napHraHrmlOH3pHOII CU-
CTeMbl.
Hc06xo]:(I1MOrTh Tal>oro nepecMoTp3 ]J.I1I>TyeTcH HC TO.lbI>O Tpe60-
BaHIIHMJI 61I0.10fll'll'Cl,IIX HayR, HO II MC]:(IIn;IIHCI\IIMII ]:(I1Cn;nn.'IIIHaMII.
B pH]:(e cJlYQaeB xlIpypnpICCl\aH npal\TIIKa IICnOJIb3yeT nplleMbl, RO-
TOpbIe OCHOBaHhl lIa HenpaBH.'IbHbIX npep;cTHH.'ICHIIHX 0 .pn:UIO:lOrlI-
qeCI\OM Ha3Ha'H'1I1I1I IlIciiIlO-rpYP;lIhIX naparanrJlnen, 3 naTO.'IOrOaHa-
TOMHqeCRaH HOMeHKJIaTypa HeROTopblX HOBo06pa30BaHHii 6a3HpyeTcH
Ha OillIl60qHhiX Tpal\TOBI\3X npnpO]:(LI ::JTUX orryxOJleii.
B OCHOBY onllcaHlIU: c06CTBeHHLIX HCCJle]:(OBalmii 1I0:IOiHeH .p3RTU-
qecl-\lIii MaTepIla.'I aBTop3, a T3l-\me TP pa60Thl, IWTOpblC BblnO.'IHeHbI
rro;rJ: ero PYKOBO]:(CTBOIII II B COBMeCTnLIX I1CC.'Il']:(OBaHIIHX C 8. X. npnii-
Mal> (I3.:ra]:(oc), H. JL IOIllI,eBHq, JI. n. l\[a:IIIHOECIWii n acnllpanToM
A. 10. Ey]:(allQeBbllll. H BhlpaiI,aIO 11M 60.'IhIllYIO npII3HaTe.'IhHOCTh 3a
rrO,D;,D;cpmI\Y BO Beex HaQIIH3111IHX H p;pymeclwe COAeiicTlHlc.
TenJlo 6J1arop;aplO npo.peccopa T. II. I3HuorpaAOBY 33 I enllhle CO-
BeTLI npII nop;roTonKe PYROIIIICH R IIeq3TII.
HaCTOHIII.aH pa60Ta Ill' 6LIJIa 6bl BbInOJIlIeHH II He YHII;:Ie:m CBC1a.
eCJlH 6bl He rrOBCe,IIHeBrr3H nOMOlIl.b, yq3CTHe II MOpaJlbHaH nO)I]:(epa\l>a
MOerO ]:(pyra n Myma A. A. EOJlOTlIlICl>oro. npUHOIllY eMY r.'Iy601,YIO
OJlaro]:(apHOCTb.
OCHOBHOii TeRCT MOHorpa.pIDI HarrllC3H ]:(0 1968 r., HO Bce JlIITep3-
TypHble MaTepH3J1bl, OTHOCHIII.IleCH R H3.;r0i1,eHHbIM B Hcii rrp06.TIeMaM
II orry6J1HRoBaHHLle HOCJle ec HaHIlcaHHH, nRJIIO'ICllhl B TCI{CT II KpHTU-
qeCI\II p3306paHLI.
J\lapT - Maif, 1971 1'.
BBE,I:(EfHIE
HaqaJIo pa3BHTIIH IIpe.n:CTaBJIeHHH 0 CIICTeMe xpoMaiP.pIIHHblX,
IlJIII naparaHrJlHOHapHbIX, OpraHOB, ROTOpblMH Mbi paCnOJIaraeM B
naCTOHlI{ee BpeMH, OTHOCHTCH R cepe]:(lIHe npOillJloro CTOJleTHH.
B 1856 r. Vulpian npII MaRpOCROIIHqeCROM Ha6mop;eHHII oUHapyamJl
xapaRTepHYro RpaCO'lllYro peaRD;Hro M03rOBoii cy6cTaHD;1l1I Hap;nOqeq-
HHKOB npH 06pa60TRe HX IIoJlYTopaxJloplIcTbIM meJle30M. Op;HaRo nep-
!ioe OnIIC3HUe cneD;HiPIPIeCROii peaR H M03rOBoro CJlOH Hap;nOqeqHH-
Ka Ha RJleTOqHOM ypOBHe IIpHHap;JlemIIT IIpo.peccopy KHeBclwro YHH-
BepCllTeTa B. A. EeD;Y (1864), ROTOpblii, ]:(eHcTBYH Ha meJle3Y pac-
TBOpOM xpmlOBblX cOJleii, nOJlyqllJl meJlTo-6ypyro ORpaCRY ee R.'IC-
TOIi. JTa peaKD;I1H, OCIIOBaUHaH, l\ali 0I>a3aJlOCb BnOCJlep;CTBHII, Ha
BOCCTaHOBJIeHIIII OKIIC.TInTeJJHMII COJJ;epmHMoro KJleTOR - ap;peHaJlHHa,
nOJlyqHJla B p;aJlbHeii:meM caMoe IlIHpOROe pacnpOCTpaHeHHe cpep;H
Bcex rIICTOXIIMlIqeCRIIX peaRD;IIH, YIIoTpe6J1HeMblx IIPH IICCJle]:(OBaHHH
naparaHrJllIOUapHOii TKaHH. CTo.'leTHHH p;aBHOCTb OTp;eJlHeT Hac OT ee
nepEoro onucalillH. C Tex nop olla UPOqHO BOIllJla B npaRTHRY HC-
CJlep;OBRTe:lbChOH pa60ThI U I1CnOJlb3yeTcH, HapH]:(y C ]:(pyrlIMII MOp-
q)OJlOrlI'IeChIlJlIlI II D;IITO:IOrUqeCI,IIMU MeTo]:(aMII, P;JlH HilyqeHUH :,}TOrO
CBoeoopa3Horo TIIIIa K.'IeTOIi, oupa3yrolI{UX RPYUHbIe u MeJlhlle CRon-
.l"eHIIH B pa3HblX y'racTKax TeJla, pa36pocanHblX B OJ];IIHOqKY II He-
60JlbillIIMII rpYIIIIaMII BHYTPH opraHOB, no XO.D:Y hpOBenocnblX COCYP;OB
II cpep;1I 3.l"eMeIlTOB BereTaTllBHoH HepBlloi'1 CIICTeMbI.
1'0]:( cnycTH IIOCJle pa60Tbi B. A. Bel\a II, BepoHTHO, He3aBIICIIMO
OT Hero, IIop;06HaJl peaRn;HH B M03roBOM cJloe Hap;IIOqeqHlIROB 6bIJIa
YCTaHOB.'IeHa HeJlleD;RIIJ\I yqeHblM Henle (1865).
Peahl\lIH EeD;a - feH.:Ie, 06ycJloBJlennaH BOCCTanOBHTe.JIbIlOii cno-
c06nOCTblO a peHa.'IUHa, BnepBbW UbIJIa p;OKa3ana ill vitro Mulon
(1904a), liOTOpblll O]:(HOBpeMeIlIlO nO.'Iyqll.'I nOJlOmnTeJlbnbIe rHCTOXH-
JlmqeChm> peahD;lIIl Ha ap;pena.'II1H npH IIOMOlI{n ]:(pyrIIx coe]:(unenllH..
npll o6paooThe nOJlYTopax.'IOpIlCTbIM meJle30M I\IITon:ra3J1Ia RJleTOR
npnoopeTa.'Ia 3e.'IeHOBaTblii OTTen OK, nplI p;eiiCTBlIII annmplI]:(OM OC-
JlIlIH - qepHblii (MuIol1, 1903, 1904b). nOqepneHHe rpaHYJl D;HTO-
n:la3l1I{,I elI{e 3a]:(OJIrO )];0 :nIIX pa60T 6bIJIO nOJlY'leno B onblTax
A. C. ,Aore.1H (1894) npn 06pa60Tlw naparanrmwnapnoii TRanll
830THO-hIICJlbIM cepeopoM. nce ::JTII peaKD;HH UbIJIll cneUII.plIqllblMII
llJIH napanUIrJlllOliapHI>IX K';WTOII: H 3aBIICeJIII OT COCCTallOBlITeJlbHOii
)J
moc06HOCTII aApeHaJIHHa (Laignel-Lavastino, 1905; Kingsburg.
1911; Ogata a. Ogata, 1917; Weymann, 1922).
Ha Py6eme ABYX CTOJIeTllll IIOHBIIJIIICb .pYH]:(aIlIeHTaJIbHbIe pa60Tbi
Stilling (1890, 1898, 1899), Vinccnt (1896, 1897a, b, c, 1900),
Kohn (1898, 1900, 1902, 1903), Kose (1902, 1904, 1907), Zucker-
kandl (1901, 1912), Wiesel (1901, 1902, a, b) II II03]:(Hee Poll
(1906), B ROTOpbIX AaHO ]:(CTaJIbHOe onHcaHHe MIIRPOCROIIlIqenWU
c'rpYRTypbI xpoMa.p.pIlHHoll TRaHH, YCTanOBJIeHO pa3BlITIIC ee H3 OA-
Horo TRaHeBoro I1CTOqHHRa C CHMnaTHqeCKOll HepBHoii. clH'TeMoii., 110-
Ka3ano Tonorpa.pUqeCIWe pacnpeAcnCIIIle no XOAY HCpBHblX CTBOJIOB
H Y3JIOB STOll CHCTeM.bI, a TaKme 6JIll3KHll ROHTaRT C KpOBeHOCHbiMII
COCYAaMH. BbiJIlI BbICJ\a3aHbI co06pameHIIH II 0 npllpo]:(e RJIeTOR, 06-
pa3YIO:m;nx xpoMa.p.pIIHHYIO TRaHb. OCHoBblBaHcb Ha reHeTlI'IeCROll
CBH3II naparaHrJIIIOHapHOii CIICTeMbI C CIiMnaTlIqeCKOll II UCnOJIb3YJI
TeXHlI'IeCRHe IIpHeMbI, ]:(OCTYIIHble AnH D;HTOJIOrUqeCROro anaJIII3a Toro
BpeMeHII, aBTopbl npeAnOJIOmHJIH cy:m;eCTBOBaHne OAHOii MaTepllH-
CROll KJIeTKH (CIIMna TorOHHH), II3 KOTOpOll B npOD;ecce pa3BIITHH AII.p-
.pepeHD;IIPYIOTCH ABa caMOCTOHTeJIbHblX SJIeMeHTa - CllMrraT06JIaCT
(CIIMIIaTlI'IeCRaH KJIeTRa) II xpOMa.p.pIlH06JIaCT (xpOMa.p.pIlHHaJl
RJIeTRa). HaqIlHaH C STllX pa60T, TKanb, ROTopaH HeU3MeHHO II Bcer-
]:(a O]:(HOTHnHO pearIlpyeT na AenCTBHC COJIen xpOMa, nOJIyqHJIa na-
3BaHIIe «XPOllW.pIlJIbHOll» (Stilling, 1890), «XpoMa.p.pIIIIIIOii» (Kohn,
1898), II.'IH «.pCOxpOMHOn» (Poll, Sommer, 1903). B pauOTax rrOC.JIe-
AYIOIII.IIX JIeT YKopelIlIJIHCb II IIOJIyqHJIU Hau60JILllIee pacnpOCTpaHeHue
ABa 1I0CJIeAHHX naHMeHonaHIIH - .peoxpOMHaH, HJIlI xpOMa.p.plInHaH_
HapHAY C STUMH Ha3BalUIHMH 1I3-3a reHeTH'lCcIWH CBH311 C CIIMna-
TlI'IeCROll HepBHOll cIIcTeMon II anaTOMlIqeCROrO pacnOJIOmeHIIH Y
BbICilllIX n03BOHOqnbiX B6ml3I1 HepBHblx raHrJIHeB xpOMa.p.pHHHaJl
TRaHb IIpH06peJIa BTopoe na3BaHue - IIaparanrJIlIOHapHOll (Kohn.
1903) (rpe1J:. cnapc1» - OKOJIO).
BoraToe COCYAIICToe cHa6menue IIaparaHrJIHOHapnol1 TKaHII, Ha-
JIlIqlIe B Hell RaIIIIJIJIHpOB II CIIHYCOIlAOB, nJIOTHO OnJIeTalOlII.I1X KJIeT-
KII, npIICYTCTBHC B D;HTOnJIa3Me UOJIblllCro UJIII Mcnblllero RO-1IHqeCTBa
xpOMa.p.pHHHbIX, apreHTo.p.pHHHbIX, IIJIH OCMno.pHJIbHblX, rpaHYJI H B.
TO me BpeMH OTCYTCTBIIe B 3TlIX opraHaX BbIBOAHblX npOTOROB AaJIO
OCHOBaHHe IICCJIeAOBaTeJIHM BblCRa3aTb MblCJIb 06 3HAOKpIIHHon IIplI-
pOAe xpoMa.p.pIlHHbIX 3JIellleHTOB, BbI]:(eJIHIO:m;IIX CBOll ceRpeT - a]:(pe-
HaJIIIH - HenocpeAcTBeHHO B RpOBb.
IIepBble IICCJIeAOBaTeJIII xpOMa.p.pHHHOll TRaHII (Wiesel, Vin-
cent, Kohn, Kose, Poll, Zuckerkandl II AP.) pacclIIaTpIIBaJIH ee He
TOJIbKO B Mop.pOJIOrHqeCROM acneRTe, HO II CTpeMHJIIICb AaTb ell (KaK
II Bo06:m;e BceM HccJIeAyeMbIM CTpYRTypaM opraHlI3Ma) .pYHRD;nOnaJIb-
HYIO oD;eHRy II oIIpe]:(eJIHTb IIX .pH3HOJIOrIIqCCROe Ha3HaqeHHe. He-
MHOrOQIICJIeHHbIMII pa60TaMH TOro BpeMeHH 6LIJI YCTaHOBJICH .pIl3I10-
JlOrIIqeCRIIll 3.p.peRT Ha BBc]:(eHIIe B l{pOBenOcHoe PYCJIO mIlBOTHbIX
3KcTpaRToB II3 OpraHOn, cOAepma:m;IIx aApeHamlH: II3 M03rOBoro C.'lOJl
HaAIIOqeqmmOB (Oliver, Schafer, 1895, U;bI6Y.'1bCRHll, 1896) II U3 op-
laHOB U;YRepRaHAJIH, npeACTaBJIHIOIII.IIX c060n 061l1l1pHble CROIIJIeHIIJI
10
naparaHrJIHOHapHon TRaHH B 3aOplOIDIIHHOM IIpOCTpaHCTBe 3apop;LI-
men (Biedl, Wiesel, 1902). II B TOM II B p;pyroM CJIY'IaHX Ha-
6JIIOp;aJIH 1I0BblIIIeHIIe TOHyca rJIa)J;RUX Mblllln; II RpOBHHoro p;aBJIeHIIJI.
Pe3YJIbTaTbl paOOT 3TIIX aBTopOB II aHaJIOrIPIHbIe 3RCIIeplIMeHTbi p;py-
rux IICCJIep;OBaTeJIen] HMeJIII cyr:u;eCTBeHHoe 3HaqeHIIe B TOM OTHOllle-
HHH, 'I:TO, BO-lIepBLIX, 1I0)l;TBep)l;HJIII MopcpOJIOrHqeCRlIe HaOJIIOp;eHIIH 0
:rraJIHqHII a]:(peHaJIHHa B RJIeTRaX xpOMacpcpHHHon TRaHH, BO-BTOPbIX,
YCTaHOBHJIH HaHOOJIOO xapaRTepHble qepTbl CPH3HOJIOrlIqeCROrO p;en-
CTBHJI IIpOP;YKTOB IIX ceRpeD;HII,. ROTOpbIe II IIO HaCTOHlIJ;ee BpeMH CqlI-
TalOTCH OCHOBHbIMH.
B pe3YJIbTaTecpH3IIOJIorlIqeCRIIX paOOT, OlIpep;eJIIIBIlIHX ap;peHa-
.JIHHOBYIO cpYHKD;UIO lIaparaHrJIHOHapHon TRaHH, BOllleJI B YlIoTpe6-
.JIeHHe TepMHH «ap;peHaJIQB8H» TRaHb. C Tex 1I0p 3TO Ha3BaHIIe IIplI-
OopeJIO IIpaBO rpamp;aHCTBa II P;O cero BpeMeHH HCIIOJIb3yeTcH B JIII-
TepaType HapH]:(y C TepMIIHaMII «xpOMacpcpIIHHaJl» H «lIaparaHrJIIIO-
lIapHaJl».
llIupoKoe paClIpOCTpaHeHHe xpOMacpcpHHHon TRaHH B opraHlI3Me,
llaJIlIqHe ee He TOJIbRO y MJIeROIIIITalOr:u;IIX, rp;e OHa BlIepBbIe 6bIJIa
o6HapymeHa, 110 II Y IIpep;CTaBHTeJIen p;pyrlIx KJIaCCOB 1I03BOHOqHblX
1f\IIBOTHbIX, rp;e ec Ha3bIBalOT cYlIpapeHaJIOBOn 2 (Balfour, 1877,
1881-1882; Vincent, 1896, 1897a, b, C, d, 1900; Kohn, 1898;
Giacomini, 1898-1920; Grynfeltt, '1902a, b, 1903a, b, 1904a, b),
YRa3blBaeT Ha YHHBepCaJIbHOCTb ee 3HaqeHHH. 06r:u;HOCTb pa3BIITIIJI
C 3JIeMeHTaMH BereTaTHBHon HepBHon CHCTeMbI II CXO]:(CTBO cp1I31I0JIO-
TII'IeCROrO p;encTBIIH ceRpeTa (ap;peHaJIHHa), IIpO]:(yD;HpyeMoro IIapa-
raHrJIHOHapHblMH RJIeTKaMH, ]:(aJIO IIOBO]:( R OO'be]:(HHeHIIIO opraHoB,
pa36pocaHHblx B pa3Hblx yqaCTKaX TeJIa, B e]:(IIHYIO CHCTeMY. 8Ta
CHCTeMa 1I0JIyqHJIa TO me Ha3BaHHe xpOMacpcpHHHon, lIaparaHrJIIIO-
HapHon (Kohn, 1903), HJIII ap;peHaJIOBon. TIoCJIep;HHn TepMHH Hap;o
npH3HaTb ep;HHCTBeHHO IIpaBHJIbHbIM II YHHBepCaJIbHbIM, TaR RaR
TOJIbRO OH OTpamaeT Cyr:u;HOCTb CHCTeMbl - ee CIIoc06HOCTb R Bblpa-
OOTKe ap;peHaJIHHOIIO]:(OOHbIX Ber:u;CCTB.
TaRHM oopa30M, B pe3YJIbTaTe KJIaCCH'IeCKHX HCCJIep;OBaHHn Toro
neplIop;a H MHOrOJIeTHHX P;IICRYCClIn R HaqaJIY TeRYlIJ;ero CTOJIeTIIJI
O.blJIH YCTaHOBJIeHLI CTPYRTYPHo-cpYHKn;HOHaJIbHbIe CBonCTBa xpOMacp-
· CBop;Ka JIIITepaTypld no BTOMY Bonpocy npllBep;eHa Y M. ll. Qe60KcapoBa
(1910,1949).
TepMIIHId «cYllpopeHaJIOBaH)} II «!HHTeppeHaJlOBaH» TRaHII COOTBeTCTBYIOT 060-
3Ha'ieHIIHM «M03roBOH)} II <<K0PKOBIdH)} CJIOH Hap;nO'le'lHIIROB BIdCillllX n03BO-
HO'lIIIdX. OHII IIpllH3p;JIelliaT EaJIbcJ>yPY (Balfour, 1877, 1881-1882), ROTOpldH
OllllCaJI Y xpH eBldx pld6 p;Be cJ>YHKl\1I0HaJIbHO pa3JIIIQHlde CTPYKTypld, op;Ha
113 KOTOpldX IIpep;CTaBJIeHa lIapHIdMII cerMeHTapHIdMIl TeJIaMII IIa KpaHllaJIbHOM
.KOHl\e rOJIOBHIdX 1I0'leK (CYllpa HaJlOBaH), a P;PyraH HellapHIdM opraH OM, KO-
"TOpIdII JIelliIlT MelliP;Y p;Op3aJlbHOH aopTOH II Kayp;aJIbHOH BeHOH II rpaHII'lllT
C 3ap;HIlMH 'laCTHMH nO'IeK (IlHTeppeHaJlOBaH). 8TlI TIa3BaHlIH YRpenllJIIICb B
JIlITepaType II lICIIOJIb3YIOTCH p;o HacTOH ero BpeMeHlI npll lI3Y'leHlIlI HlI3illllX
1I03BOHU'lHIdX, XOTH TOlIorpacJ>lIH BTIIX TKaHeH, TaR llie KaR «KOPKOBIdH» II
-«.M03roBOH)} CJIOll Hap;nO'le'lHIlKOB, He Bcerp;a COOTBeTcTByeT TepMlIHOJIOrll'le-
CKIIM BldpameHHHM.
11
cf>IIHHOH TRaHlI II C03]]:aHO CTpOHHOe yqeHlIe () CIICTeHe 8lU>eH8no-
BbIX 3H]]:OKpIlHHbIX OpraHOB. BLuIII BbIHCHeHbI aHaTOMO-Tonorpaqm-
qeCRlIe OC06eHHOCTII xpoMaqJllmHHblx (.peOXpOMHbIX) OpraHOB Y pa3-
HblX rpYlI1I 1I03BOHO'IHbIX iKIIBOTHbIX, OCHOBHble qepTbl ilK rIlCTOJIO-
rlIqeCROH OpraHll3an;1I1I II IJ;IITOJIOrllqeCKIIe OC06eHHOCTII napeHXIIMa-
T03HblX RJIeTOR. BblJIO ]]:ORa3aIl0, qTO rOpMOH3JIbHOe CO]]:epiKIIMOe IIX
OTHOCIITCH K a;xpeHa.TIIIHY II B .pYHhIJ;IIOHa.TIbHOM OTHOmeHIIII ]:(eUCTBY-
eT nO]]:06HO .p1l3110JIOrIlQeCROMY 3.p.peRTY CIIMlIaTII'IeChOH HepBHOD:
CIICTeMbi II qTO pa3BIIBaeTCH Xpm.Ia.p.pIlHH3H CIICTeMa 113 o;XUOrO IIC-
'IO'IHIffia C CIIMlIaTIlqeCROH HepBHOll CIICTeMOH II ]]:OJIiKHa paCCMaTplI-
mirbCH RaK COCTaBHaH ee qaCTb.
- lke 113JIOiKeHHOe 3aBepllllIJIO ]:(.TIIlTe.TIbHbIU II II.'IO]:(OTBOpHblii 3TaII
IICCJIe]]:OBaHIiH a]]:peHaJIOBOH CIICTeMbJ, IICTORII ROTOpblX OTHOCHTCH K
cepe]]:IIHe XIX B.
AaJIbHeIlllIee pa3nllTIIe pa60T no napar3Hr.'lnOHapnoii cnCTeMe,
ROTOpLIMIi 03HaMeHOBaJIIICb nepBLIe ]]:eCHTIIJIeTIIH TCRYlIJ;erO CTo;rr:eTIIH,
xapaKTepll3YIOTCH yr.TIy6JIeHlleM Tex 3H3HIIII, ROTOpbIe YiKe 6blJIII ;Xo-
6blTbl no Mop.pOJIOrlill II 3apo]]:LlllieBoMY pa3BIITllIO naparanr;IIIIoHap-
HbiX OpraHOB II paCllIlIpeHlIeM 06'beRTOB IIcc.TIe]]:OBaHIIH.
Hall60JIblllerO pa3BIITIIH ]]:OCTlIrJIO 1I3yqeHIIe xpoMa.pqJlIRRou CII-
CTeMbl BLJCIlIIIX 1I03BOHO'IHl>IX. r.rraBHLJM 06pa30M MJJeKonIlTaIOll.{IIX.
npOBO]]:IIBllIeeCH B cpanUIITeJIbIlO-MOp.pO,TIOrIPieCKOM, rIlCTO.'lOrlI'IeCROM
Ii 3M6pIIOJIOrll'IeCROM aCII(,I Tax. HpoMe Toro, B ncpn0ii leTBepTIf
XIX B. Haqaml IIOHBJUITbCH IICCJIC]:(OBaHIIH, BblllOJIHeHHLIC Ha OpraHaX
qeJIOBeRa. B qlICJIe IIX 3aCJIYiKIIB31OT BHIIMaHIIH IICCJIe]]:OBaHIIH HalllllX
OTeqeCTBeHHLlX aBTopOB - M. 11. t.le60RcapoBa (1909, 1910).
It M. flxoHToBa (1913a, 6, 1914), r. <1>. IIBaHoBa (1930).
B 3KCIIcpIIMeHTaJIbROM IICCJIe]]:onaRIIII M. H. Qe60I{CapOBa (1909,
1910), B ROTopm.r O]]:HOBpeMeHHO npe]]:CT3B.'1eHa CBO]]:Ra Hcero Toro,
qTO 6bIJIO c]:(e.TIaHO ]]:0 Hero no IIHRpeTopHoii .pYHRn;1I1I H3]]:nO'Ie'IHll-
ROB, IIOKa3aHO 3Ha'IeHIre CIIMnaTII'IeCROrO qpeBHOro HepBa, IIIIHepBII-
PYIOI.4ero iKeJIe3Y II IIrpalOI.4ero pOJIb ceRpeTopHoro nYChOBOrO Me-
X3RII3Ma.
CepIIH pa60T l\. M. HXOHTOBa IIOCBHI.4eH3 rIlCTO:IOnr'IeCKmry
CTpoeH1l1O II pa3BIITllIO naparaHrmleB qe.'10BeI\a. ConOCTaB.'lHH MOp.pO-
JIOrII'IeCKYIO opraHlI3aIJ;IIIO lIaparaRrJIlIeB Y B3pOC.TIbIX JIIO]]:cii C xapah-
TepHbIMII CTPYRTypHLIMII OC06CHIIOCTRMII IIX B 3l\16pIlOrenC3e, aBTOp
c]]:eJIaJI PH]]: n;eHHbIX Ha6JIIO]]:eHllii II BbIBO]]:OB, He 1I0TepHBllIIIX 3Ha're-
HIIH ]:(0 HaCTOHI.4erO BpeMeHII.
OqeHb IIHTepeCHO ]]:eTaJIbHCiilllee onlicaHlie HXOHTOBbIM nap3-
raHrJIIIH, paCnOJIOiKenHOrO B 3]:(HeHTIIIJ;II11 COHROll apTepllIl Y MeCTa
eJIeHIIH ee Ha BnYTpCHHlO1O II HapYiKHYIO. rpYlIlIa h:ICTOh, O'IeRb
pe]]:KO ]]:alOIIJ;IIX 1I0.JJOiKIITeJIbHYIO xpOMa.pqnIHHYIO pcaRQlllO, HaXO]]:IIT-
CH B TeCHO ROHTaRTe C TRanblO CTeHOK RpOBeHOCHOrO cocy]]:a II lIe
IIMeeT HCRO BblpaiKeHHOll RanCYJIbl, KOTopaH OT]]:eJIH.TI3 6LI IIX OT
oRpyiKalOI.4IIX 3JIeMeHTOB COCY]]:IICToii CTeHKII. BMeCTe C a]]:BCHTIIl.{IIeU,
C ROTOpOll 3TII RJIeTRJI COCTaBJIHIOT e]]:IIRoe IJ;eJIoe, 01111 npOHII3aH"I
MHROTHblMII II 6e3MHlWTHLIMII nepnaMII, Kamm.TIHpaMII II MeJIKII U
f2
cocy]:(aMII. TaRoe CTpoeBlle He 1I03BOJIHeT HXOHTOBY Ha3bIBaTb 8TO
06pa30BaHlIe opraHOM B CTporOM CMblCJIe CJIOBa, II OB 0603BaqaeT
ero RaR ROMIIJIeRC TRaneBbiX meJIe311CTbiX RJIeTOR. OlIpeAeJIeHlIe 8TOB:
CTpYRTYPbl RaR TRaHeBOU, a He opraHHOH, OqeHb xopomo corJIacyeT-
CH, RaR Mbl 1I0CTapaeMCH IIORa3aTb ]:(a.TJbllle, C IIpeACTaBJIeHIIHMII 0
cpllJIOreHlIlf BceR lIaparaHrJIIIOBapHOII CIICTeMbl.
B pa36IIpaeMblH lIepllOA HOBbIe AaHHbIe 110 3M6plIoreHe3Y XpO-
MacpcpllHHblX opraHOB qeJIOBeRa 0lIy6JI1IROBaJIlI Hammar (1925), HeU
(1925), Kohno (1925), Keene, Hewer (1927), )J.. M. rOJIy6 (1936)
II no aM6pllorene3Y nTlln; - Lutz, Case (1925), Miller E. (1926),
Okuda (1928), Brauer (1932, 1934), )J.. M. ro.TJy6 (1936). EoraTblR
MaTeplla.'I 110 cpaBHIITeJIbHOH rlICTOJIOrllll II cpaBHIITeJIbHOH 8M6p1I0-
JIOrlllI lIaparaBrJIlleB y MJIeROlllITalOll\lIX 6b1JI IIpCACTaBJIeH B pa60Tax'
Celestino da Costa (1917-1939) II Kohno (1925). nOCJIeAHIIR 01I1I-
caJI CTPYRTYPY HaAlIoqeqHIIROB Y npcAcTaBIITe cR pa3HWX ceMeHCTB
II pOAOB ]:(eBHTII OTpHAOB MJIeROIIIITalOll\IIX: Y eXIIAHbl, 3eMJIepoiffilI,
BO]:(HOH RYTOPbl, RpOTa, JIeTyqCII c06alm, JIeTyqeU Mbllllll, IIOAROBOHO-
ca, YlliaHa, oHAaTpbl, CYCJIIIRa, MYllIJIOBRII, 3aHn;a-6eJIHRa, MblllllI,
UpblCbl, MypaBbeAa, Hocopora, CJIOHa, hOCYJIII, CCpHbl, oBn;e6bIua, uap-
ca, MeABeAH-ry6aqa II Y HeROTopblX 06e3bHH (6a6YIIHa, MauaRlI, Map-
TbIIllRII, qepHOrO lIaBllaHa). 06a aBTopa - Celestino da Costa II Koh-
no, Ha OCHOBaHlIII rllCTOXlIMllqeCI{IIX hapTlIH, rOllopll 1I 0 CYll\eCTBO-
BaHlIII n;IIRJIllqeCROH IIHRpeTopnoH AeHTeJlbHOCTII lIaparaHrJIIIOBapHblx
UJIeTOU.
TaulIM 06pa30M, HaROIIJIeHHblH B 20-30 rOAbI Halliero CTOJIeTIIH
HOBblH 60JIblllOH cpaRTllqCCRIIH MaTepllaJI no MopcpOJIOrlllI II aM6plIO-
JIOrlllI xpOMacpcpllHHOH CIICTeMbI MJIeROlllITalOll\lIX 3HaqlITeJIbHO pac-
mllpllJI Hallill npe]:(CTaBJIeHIIH 0 pacnpOCTpaHeHIIII naparaHrJIlIeB Y
BblCIlIIIX 1I03BOHOQHbIX, 06 IIX CTPYhTypHbIX B3alIMOOTHOllleHHHX C
APyrllMII TRaHHMII, 06 nx IIHHepBan;lIl1 II BaCRYJIHpII3an;IIII, rlIcTocplI-
3IIOJIOrllll II ;)M6pIIOHaJIbHOM pa3BlITli1I. 1I3yqeHIIe aM6plloreHe3a II
CTpYRTypHOH opraHH3an;I111 naparaHrJInCB qeJIOBeRa CbIrpaRO HeCOM-
HeHBYIO 1I0JIOmllTCJIbHYIO pOJIb B nOHlIMaHlII1 HC TOJIbRO CP1I31I0JIOrll-
QeCRIIX npOn;eCCOB, npOTeRalOll\IIX IIpll HenocpeACTBeHHOM yqaCTlI1I
rOpMOHaJIbHblX B03Aei'rcTBIIH co CTOpOlIbl xpoMacpcpllHHblX opraHOB,
BO ORa3aJIO BJIIIHHlle II Ha 1I0HIIMaHlIC pH;J.a lIaTOJIOrll'IeCRIIX HBJIe-
HIIH. napaJIJIe.TJbHO BceCTopOHHeMY 113Y'leHUIO naparaHrJIlIeB B 3AO-
pOBOM opraHH3Me pa3BepHYJIHCb IICCJIeAOBaBIIH 110 naTOJIOrHII, B qaCT-
HOCTII, no pa3BIITllIO ollyxOJIeH lIaparaHrJIIIOHapHoH IIPIlPOAbl (CM.
MOHorpacpIlII Terracol, 1956; Gastpar. 1961; MaccoHa, 1965).
COBCplllCHHO eCTeCTBeHIIO, QTO HOBbIe MeTO]:(lIqeCRlle npHeMbl IIC-
C.'IeAOBaHHH ClIMlIaTo-aApeHaJIOBOH CIICTeMbl, MHoro06pa311e 06'1:eRTOB,
IIO]:(BeprHYTbIX IICCJIeAOBaHHIO, II pa3Ho06pa3IIe opraHOB, ROTOpbIe
ClIen;lIa. T lbHO 113yqaJIIICb IIJIII B ROTOpbIX CJIy<raiiHO 6bIJIII HaiiAeml
lIaparaHrJIIIII, BChpbIJIIi HehOTopble HOBbie cpahTbI II 1I0pOAIIJIII PHA
]:(IICRYCCIIOHHLIX BonpOCOB. r JIaBllbIMII 113 HIIX, CYAH no 1I0CJIe]:(CTBII-
HM, BaJIOmllBllIlIM OTlIeqaTOR Ha IICCJIe.n:OBaHIIH B TeqeHlIe MHorlIX
nOCJIeAYIOll\I1x AeCHTIIJIeTIIH, HBJIHIOTCH aTHIIII3M cTpOeHIIJI pHAa na-
ta
paraHrJIlIeB, rHCTOXHMH'leCKlIe CBOnCTBa H pa3BIITHe KOTOpblX He
YKJIa]:(bIBaJIHCb B Te cxeMbI, KOTOpbIe 6bIJIH YCTaHOBJIeHbI H IIpe)];onpe-
)];eJIeHbI HCCJIe)];OBaTeJIHMH )];JIH lIaparaHrJIHeB BblCIlIHX n03BOHO'lHbIX.
ATlInH'leCKHe KJIeTKH pa3Hbie aBTopbI Ha3bIBaJIH JIH60 «OC06bIMII»
JIH60 «rJIOMYCHbIMII». llO]:(XO)];HIIJ;HM TepMHHOM, OTpamalOIIJ;IIM CTPYR-
TYPY TaKHX aTHIIH'lHbIX KJIeTOK, 06bl'lHO 60JIee MeJIKHX, 'leM B M03-
rOB OM CJIoe Ha)];IIOqe'lHHlm, MomHO C'llITaTb «aTpo.pH'lHble». XapaK-
TepHoe HX CBonCTBO 3almlOqaJIOCb B HenOCTOHHCTBe IIJIII nOJIHOM OT-
CYTCTBIIII B HHX rOpMOHOB, )];OCTYIIHbIX rHCTOXHMH'leCKOMY BbIHBJIe-
HRIO MeTO)];aMH XpOMRpOBaHHH, OCMHpOBaHIIH II cepe6peHIIJI. KpOMe
Toro, 6bI.'I0 npH3HaHO, qTO «rJIOMYCHbIe» KJIeTKH HK06bI pa3B1lBalOT-
CH He II3 CHMnaTH'leCI\On HepBHon CHCTeMbl, IWR 3TO 6bIJIO YCTaHoB-
JIeHO elIJ;e OCHOBOIIOJIOmHHlWMH Y'leHHH 0 naparaHrJIHHX II 1I0]:(TBepm-
)];eHO B nOCJIe)];YIOIIJ;IIx pa60Tax, a H3 napacHMlIaTlI'leCIWn CIICTeMbI.
K TaRHM CIIOpHblM lIaparaHrJIHHM OTHeceHbl Y3eJIKII xpOMa.p.pIIHHbIX
RJIeTOK, paCIIOJIOmeHHbIe B 6Hq,YPKaIJ;HII 06:rn:en COHHOil apTeplIlI,
B 06JIaCTII )];yrH aopTbl, Mem)];y aopTon 11 JIerO'lHOn apTeplIen, ORO-
JIOCep)];e'lHble H )];PyrlIe MeJIKHe 1I3paraHrJIHOHapHbIe CTPYI\TYpbl rpy)];-
Hon 06JIaCTH. B ]:(a.TIbllenllleM 1I0)];06Hb1X aTHlIlI'lHblX lIaparaHrJIlIeB
OKa3aJIOCb 3Ha'lHTeJIbHO 60JIbme II Bce 60JIbmee 'lIICJIO HX CTaHOBIIT-
CH II3BeCTHbiM - HX HaXO)];HT II B HacTOHlIJ;ee BpeMH B pa3Hblx opra-
H3X BblCllIliX 1I03BOHO'lHbIX. OHII BCTpe'laIOTCH, I<3R IIOKa3aHO B no-
CJIe)];YIOIIJ;IIX rJIaBaX, II Y HII3RoopraHH30BaHHblx .pOpM n03BOHO'lHbIX.
TpY)];HO CI\a3aTb, 1\ 'leMY MOrJIII 6bl CBeCTHCb pe3YJIbTaTbl pa-
60T Toro IIepHO)];a, B IWTOPblX aBTopbl B Ton IlJIH HHon .popMe KOH-
CTaTHpOBaJIIi HJIIi BCKOJIb3b YIIOMHHaJIII 0 CTPYI\TypHO-XHMHqeCKIIX
II rJICToreHeTHqeCKIIX oc06eHHOCTHX HeKOTopblX xpoMacpcplIHHblX 06-
pa30B3HIIn Y IITHIJ; II MJIeKOmITaIOIIJ;HX. B03MomHo, HalIpaBJIeHHe 3TIIX
IICCJIe)];OBaHHii BOIllJIO 6bI B 60JIee npaBHJIbHOe, 060cHoBaHHoe peaJIb-
HbIMH .paRTopaMII PYCJIO, B CMbICJIe H3Y'leHHH 3H)];OKpHHHoro xapaK-
Tepa lIaparaHr.'IHeB, eCJIH 6bI 1I0)];HHTbIe BOlIpOCbl He CJIIIJIIICb II He
nepelIJIeJIHCb C )];Pyroii, OqeHb 60JIblliOi'I, HO cOBepllleHHo caMOCTOH-
TeJIbHOn IIp06JIeMoH. Mbl nMeeM B BH)];Y IIp06JIeMY pecpJIeR OreHHbIx
iKeMOpeIJ;elITOpHblX 30H.
OTl\pblTHe xeMopeqenIJ;UH cHHoKapOTII)];HbIX II JIerO'lHO-aOpTaJlb-
HblX 30H 1I0cJIymlIJI0 TOJIbl\O nepBblM TOJl'IKOM R lIepeCMoTpy Bcen
CIICTeMbI naparaHrJlHeB Y BbICIlIHX II03ROHO'lHbIX, paHee IIpH3H8HHblX
B Ra'leCTBe ;m]J:oRpIlHHbIX. 3aTeM B03HHKJlII pa3HOpe'llIBbIe ROHqen-
qllR, B ROTOpbIX lIaparaHrmlOHapHoH CHCTeMe JlH60 IIplIIIIICbIB3J13Cb
BTopaH cpYHKIJ;HH - pen;enTopHaH, Jlll60 CTaBHJlOCb no]:( COMHeHlle
cpYHRIJ;HOHaJIbHOe e]:(IIHo06pa3IIe ee OpraHOB B CMblCJIe Bbl]:(eJIeHIIH
MpeHaJlHHOII0)];06HbIX ropMoHoB. AIICKYTHpyeMbIe Bonpocbl cnoc06-
CTBOBaJlII 1I0HBJleHHIO HOBOH RJIaCclIcplIKaIJ;IIH naparaHrJllIeB H )];eJle-
HHIO IIX Ha a]:(peHaJIHHOreHHbIe, xpoMa.p.pHHHbIe, HJlII CIIMlIaTuqeCRlIe
( «IICTHHHbIe») H Ha JIHllIeHHbIe 3THX CBOHCTB - HexpoMacp.pHHHble,
1IJ111 napaclIMnaTH'leCRlIe ( «HeIICTIIHHble» ). O)];HOBpeMeHHo IIOHBII-
JlOCb II TpeTbe nOHHTHe - «CeHCOpHbIX» 1I3paraHrJllIeB. llpe)];CTaBJle-
HlIe 0 lIapaCHMnaTII'leCKIIX lIaparaHrJlIIHX 1I0)];KpeIIJlJlJlOCb 3M6plIOJlo-
14
I'lI'IeCKIIMII IICCJleAOBaHIIJlMII, B KOTOpblX HR'laJlII nOJlBJlJlTbCJI YKa3a-
HIIJI Ha rlICTOreHeTlPlecKyro CBJl3b HeKOTOpblX II3 3THX 00pa30BaHIIH
He C Y3J1aMII H BOJIOKHaMII CIIMnaTII'leCKIIX CTBOJlOB, a C CIICTeMOH
oJlymAaror:u;ero HepBa. BOJlblllYro pOJIb B oocymAeHlI1I Bcex CIIOpHblX
BonpOCOB CbIrpaJlII HCCJleAOBaHIIJI a.p.pepeHTHOH II 3.p.pepeHTHOH HH-
HepBaQHH 3TIIX opraHOB II Mop.pOJlOrlI'leCKOH TpaKTOBKII HHHepBaQH-
OHHbIX MexaHH3MOB.
IIepecMoTp caMoro nOHHTHH naparaHrJlIIOHapHOH TKaHH, Ha'laB-
mIIiIcH B 20-30-e rOAbl XX B., 3aTHHYJlCH II, He IIplIBeAJI K KaKHM-
JIHOO 00lIJ;elIplI3HaHHblM oooor:u;eHHHM, AomHJI B TaKOM HeYCTOHBllIeM-
CH, nOJIHOM IIpOTIIBOpeqIIH, BHAe AO HallilIX AHeH.. B HaCTOJlr:u;ee Bpe
MH, OJlaro]:(apH HCKJlro'lIITeJlbHOMY paCQBCTY HOBblX TeXHH'leCKIIX nplI-
eMOB H 1I0HBJleHllro B03MomHOCTII npHHQlInHaJlbHO HHblX nO]:(XOAOB K
IICCJleAOBanllro OIlOJlOrII'leCKlIX IIpOQeCCOB, npeACTaBJleHHH 00 opraHax
aApeHaJlOBOH CliCTeMbI YCTapeJlU, TaK KaK 1I0TepHJlII TY .paKTlI'leCKYIO
OCHOBY, Ha KOTOpOH CTPOUJlIICb. )J;OCTaTO'IHO CKa3aTb, qTO MeTOAbi
rlICTOXliMUU II 3J1eKTpOHHOH MIIKpOCKonIIII IIOKa3aJlII HeCOCTOJITeJlb-
HOCTb KOHn;enn;IIH 0 cyr:u;eCTBOBaHIIII (<HeXpOMa.p.pIlHHbIX», He co]:(ep-
mar:u;IIX KaTeXOJlaMIIHOB KJleTOK, a TaKOH MeTO]:(, KaK aBTopaAlIorpa-
.pIIJl, n03BOJIlIJi nO-HOBOMY OQeHIITb Te KapTIIHbl rlICToreHe3a, KOTO-
pble IIpemAe UbIJIU OlIucaHbl C HeOObl'laHHOH Tr:u;aTeJlbHOcTbro. HapHAY
C 3TIIM 1I0HBIIJlaCb B03MmKIIOCTb paCllIu.ppOBaTb UHHepBan;IIOHHble
CTPYKTypbl Ha naparaHrJllIOHapHblX h.'ICThaX, nOJlY'leHHbIe B CBoe
BpeMH caMblM COBepmeHHblM II cnCQII.pIl'lHbIM AJlJI HepBOB MPTOAOM
cepeopeHIIH, KOTOpblH CbIrpaJi orpOMHYro nporpeCCIIBHYro pOJIb B
IICCJle]:(OBaHUHX QeHTpaJlbHOH II ncplI.pepU'leCKOH HepBHOH CIICTeMbI,
HO TaK me, KaK II JlroOOH APyroH MeTOAII'leCKIIH npueM, nopoAIIJI
cnopHble BonpOCbI, OCTaBllIueCH OTKpblTbIMII.
He]:(OYMeHHbIe BonpOCbI U HeCOBMeCTHlIlble MHeHIIH cyr:u;eCTBOBaJlII
U B IIpOmJlOM BeKe. XapaKTepHbIM nplIMepOM 3Toro MomeT CJfY-
mUTb KapOTHAHblH naparaHrJlIIH, KOTOpbIH, oy]:(yqII BnepBbIe onlIcaH
MaRpOCKOnlI'leCKII Taube B 1743 r. KaK raHrJlIIH, CBH3aHHbIH C Bepx-
BIIM meHHblM Y3J10M t (lliiT. no Kohn, 1900), paCCMaTplIBaJiCH nOTOM
TO KaK meJle3IICTbIH opraH C 31IIITeJlIIOnOAOOHblM pacnOJlOmeHlIeM KJle-
TOK II C ooraTblM HepBHblM CHaomeHIIeM U3 BepXHero meihIoro
CIIMnaTII'leCKOrO raHrJlIIH - «Nervendriise» (Luschka, 1862), TO KaK
COCYAIICTaH CTpYKTypa: JlIIOO II3BIIJlIICTaH, 00pa3yror:u;aH rJIOMepyJfY-
\Obi - «Glomeruli arteriosi intercarotici» (Arnold, 1865; Pfortner,
1869; Waldeyer, 1875), JlII60 pYAIIMeHTapHaH, nOCTpOeHHaJl II3 KJle-
TOK KpOBeHOCHbIX COCYAOB (Marchand, (891), JlIIOO, HaKOHeQ, COCY-
AUCTaH KpOBHHaJl meJle3a co CTPYKTypoii, aHaJlOrH'lHOH M03rOBoH
qaCTII Ha]:(IIO'le'lHIIKa (Stilling, 1892, 1898). PHAOM C TaKIIMII onpe-
AeJleHIIHMII KapOTIIAHoro naparaHrJlIIH 1I0'lTU BcerAa CTOJlJl KOn'lIIKO-
BblH naparaHrJlIIH, CTPYKTYpHO O'leHb CXOAHblH C KapOTIIAHbIM.
I IIo IiICTOpll'leCKOH cnpaJIKe, KOTOpylO npllBop;IiIT Adams (1958), Ha o, BII IIMO.
C'lIlTaTb, qTO npliloplilTeT nplilHap;J1ellililT He TOJIbRO Taube, HO II Haller (1742)
II ero P;BYM Y'leHIIKaM: Taube (1743) II Berckelmann (1744).
i5
«Habent sua fata libelli)} - (<lmllrlI IIMelOT CBOIO cYJJ;b6y)),- KaK
rOBOpllJIII JJ;peBHlle pIlMJIHHe. HaY'IHbIe np06JleMbl, Kal{ II KHIIrll, TaR-
me, O'ieBIIJJ;HO, IIMelOT CBOIO cYJJ;b6y. He60JIbIDOH Y3eJIOK, He BblJJ;eJI:HIO-
r:u;nHC:H Hll'IeM 3aMe'IaTeJIbHb1M IIO CBoeH CTpYRType, YiKe CBblllle ]:(BYX
CTOJIeTIIH IIplIBJIeRaeT BHIIMaHlIe ClIe lIaJInCTOB pa3HblX IIpo.pIIJIeH II
- 06cymJJ;aeTCH C TO'IlHI 3peHII:H IIp06JIeM, JJ;IIaMeTpaJIbHO npOTIIBOIIO-
JIOmHbIX ]:(pyr ]:(pyry.
U,eJIb lIaCTOHll(eM pa60Tbi He TOJIbKO B OlIpOBepmeHlI1I Bcex Tex
«lIaCJIOBUIIih), KOTOpbIe BO;,HlH{JI1I 3a 1I0CJIeJJ;HlIe JJ;eCHTIIJIeTIIH B MOp-
cpOcpYHlHJ;IIOHa.TIbHOii ou,cml:e a]:(peHaJIOBblX opraHOB II B meJIaHIIII Bep-
HYTb 11M YTepHHHYIO penYTau,nlO 9HJJ;OKPIlHHbIX, HO rJIaBHOe B TOM,
'IT06b1 IIOJJ;OMTII R IIpo6.TICMe HX .pIIJlOreHe3a II C HOBbIX MeTOJJ;OJIOrll-
qeCKIIX 1I03I1U,IIH npOaHaJIII3IIpOBaTb lIaparaHrJInOHapHYIO CIICTeMY C
( y'IeTOM Beero HaKOnJIeHHOrO apceHa.TIa <sHaHIIH.
B03 BpalIJ; aHc . K___'i:eMopeu,eIITopHbIM 30HaM, HaliOMHIIM OCHOBHble
MOMenTbl II3 IICTOpll1l pa3BIITIIH t)TOM np06JIeMbl.
- B CPII31I0JIoru'IeCI{HX pa60Tax 20-30-x rOJJ;OB [Hering (1924,
1927), C. Heymalls, J. Heymans (1926), C. Heymans (1934),
C. Heymans, Bouckaelt (1930), Bouckaert, Dautrebande, C. Heymans
(1931), C. Heymans, Bouckaert, Dautrebande (1932), Heymans,
Bouckaert, Regniers (1933)] II B noc.TIC]:(YIOll(HX 6blJIO nOlia3aHO pecp-
JIeKTOpHoe BJIIIHHlle JJ;aBJIellllH (Hering, 1924) II XIIMuqeCRoro COCTaB8
RpOBII Ha JJ;blxaTeJlblIbIii II cOcy]:(oJJ;BnraTeJIbHbIH lIeHTpbl C IIHTepOpe-
qelITopoB Rap]:(lIoaOpTaJIbHOll II cIIHoKapoTIIJJ;HOll 06JIaCTeH.
OTRpblTlle pe.pJIel{C-OH'HHoro 311a'IeHIIH aTIiX 30H 1I0JIOiKIIJIO Ha'Ia-
JIO u,eJIeYeTpeMJIeHHoMY 113Y'IeHllIO MexaHII3MOB xeMopeu,elIu,1I1I n pe-
rYJIHu;nll I{POB006palIJ;eHIIH, IIHTeHC-IIBHO II IIJIOJJ;OTBopno pa3pa6aTbIBa-
eMOMY JJ;O HaC-TOHlIJ;ero npeMCHIi. Mop.poJIom, 3aHHBllIlleCH rIlCTOJIO-
rlI"leCKIIM 113Y'IeHlleM 611.pypl',au,lI11 COHHOH apTepll1l II IIHHepBau;ueH
aToH 06JIaCTH (Dl'iiner, 1925; de Castro, 1926, 1928), enoc-06CTBOBaJIII
YKpelIJIeHlllO pa3BIIBaeMbIX .p1l3110JIOraMII npeJJ;cTan.TIeHllii 0 ee 'IYBCT-
BnTeJIbHOCTII K MexaHIPleCIUIM U XIIMlI'IeCKHM pa3JJ;paiKIITeJIHM. B no-
IIC-KaX Mop.pOJIOrll'IeC-KOrO c-y6eTpaTa, OCYlIJ;eCTBJIHlOlIJ;erO .pYHKU;1I1O
BOClIpllHTIIH pa3]:(pameIlIlM, de Castro ]:(aJI IICqepnbIBalOr:u;ee 0lIHC8Hlle
acp.pepeHTHblx IIpll60pOB B THaHHX t)TOH 06JIaC-TlI II ]J:ORa3aJI B aKC-
IIepllMeHTe IIpIlHa]:(JlemHOCTb IIX I{ 6.TIY}I{]:(alOll(eMY II H3bIKOrJIOTOq-
HOMY HepBaM. all no.TIaraJI, 'ITO CHHYCHblM HepB XepIlHra, KOTOpbIH
HBJIHeTCH BeTBblO H3bIKO-rJIOTO'IHOrO HepBa, 3aKaH'IllBaHCb peU;enTOp-
HblMII IIpll60paMil na «rJIOMYC-HbIX)) KJIeTImX HapOTIIJJ;HOH apTeplIlI,
06pa3yeT xeMopeu;enTopbl, a 3aKaH'IIlBaHC-b B C-TeHKax KapOTIIJJ;Horo
cIIHyc-a,- npec-coppu,enTOpbl. TaKIiM 06pa30M, B pe3YJIbTaTe .p1l3110JIO-
rn"leCI{IIX II Mop.pOJIOrnqCCHIIX I1cc.TIe]:(OBaHIIH 6bIJlO KOHCTaTupOBaHO
JJ;Ba Tlliia peU;elIU;lIl1, OJJ;I1H 113 IWTOPblX pearllpOBaJI Ha U3 IeHeHIIH
KpOBHHoro .o;aBJIeIUIH n C-OHHOii apTepHII, ]:(pyroH Boc-npmmMaJI OT-
I\JIOHeHIIH B XUMII3Me RpOBII. CTPYKTypHbIH aTIIII1I3M KJIeTOK KapOTII -
Horo lIaparanrJIIIH, 0 KOTOpOM rOBOpUJIOCb BbIllIe, OTC-YTC-TBIIe B HIIX
ropMOHOB II IIpIlCYTC-TJHIC Ha KamJJ;OH 1I3 HIIX KOHn;eBbIX peu;eIlTOp-
HblX IIJIaCTIIHOK, coeJJ;I1HeHHbIX C "lYBCTBIITeJIbHbIMII HepBllblMII BO-
c..
-
N
1(j
JIOImaMU, ]J.aJIO OCHOhilHIIC de Cast.ro OTpllu;aTb IIX :JH]J.OKpIlHHbIe
CBoHcTBa II YTBepm]:(aTb, qTO IIMeHIIO 3TJIM «rJIOMYCHbIM» K.TJeTKaM
upUHa]J.JIemlIT q,YHKU;IIH BocnpUHuMalOlIJ;erO annapaTa, perucTpupylO-
r:u;ero xeMo]J.uHaMuKY KpOBII.
BCJIe]:( 3a Mopq,OJIOrllqeCKUM1I I1CC.TJe]:(OB3HUHMU de Cast. I 0 nOHBU-
JlUCb pa60TbI Penitschka (1931), Palmc (1934), Nonidez (HJ35a, b,
1936), Murat.ori" (1935, 1937), KOTOpbIC, U3yqllB naparaHr.TJIIII B
06JIaCTII CCp U;3, aOpTaJIblIOH II .TJerO'lHOH ]:(yr, OTHeCJIlI IIX no CXO]:(-
HbIM CTPYI>TypHblM II rUCTOXIIMIPWCKIIM npll3H3KaM I> XCMopeu;en-
TOpHblM «r::IOMYCHbIM» oupa30naHlIRM II He npu:maBaJIU IIX :In]:(OKpUH-
HYIO npllpo]J.Y. «fJIOMYCHbIe» K.'ICTIHI npllcep]J.eqHOH 06JIaCTII Tal>me
1II1le.-UI 60raTYIO qYBCTBUTCJIbHYIO lIHHCpBaU;lIlO, OCYIl(eCTB.TJHeMYIO ]J.e-
npeCCOpHblM HepBOM. no aHa.TJorllll C l>apOTII]:(HbIM «rJIOMYCOM» 6bI.TJO
BbICI>a2aHO npe]:(nO.TJOmeHUe, 'lTO BO.10IiHa ]:(enpeccopa, 331>3UqUnanCb
na «r.'IOIIIYCUbIX» K'IeTK3X ]J.yrll aopTbI, .'ICrO'lHOll II nO]J.li:llOqll'lHOll
apTepllRX. 06JIa]:(a1OT xeMopeu;enTopHoii q,YHRU;lIell, a em OI\oHqallUH
B CTeHKax 3TUX apTepUH - npeccopeu;enTopHOH.
Tal>on 6bl!I OCHOBIIOll IITor 11CCJIP]J.OB3UIIH 0 Mopq,o:wrUqeCKOM
cy6cTpaTe, OTBeTCTBeHHOM 3a BocnplIRTlIe XIIMU'IeCKIIX C]J.BJIrOB KpO-
BII. J3eCqIlCJIeHHbJe nOCJle]J.ylOllJ,lIe IICCJIe]J.OBaHUH npenpaTUJIU
KOHu;enu;ulO 0 CYJ1:{ecTBOB8HIIII oc06L1x «rJIOMYCHLlX» RJIeTOK C npIIHa]J.-
JlcmalIJ;pii 11M xeMopeu;cnTopHoii: q,YlmU;IICH B He3bI6JICMblii q,aRT.
COIwpmCHHO eCTceTBCUUO, 'lTO BonpOCbI peq,JICI\Topuoii pery JIR-
:U;IIII I>poBo06palIJ;eHIIR U TCM 60JIec ellH33HHoro C HpOl30CHa6meHIICM
M03ra, IIMeH orpOMHoe TCopCTu'reeIWC II npaHTUqeeHOe 3UaqCUlfe,
IIpll06pc:m llIllPOKYIO nonYJIHpHOCTb II nOJIyqu.'1n npll3HaHlle B l\a-
-qeCTBe aHTyaJIbHbIX np06JIeM no MIIOrIIX 06JIaCTHX HayqHblX 3Ha-
Huii - B q,1I3UOJIOrUn, q,apMaKOJIOrnn, RJIlIllIIqeCIWH Me]J.UU;UHe. 13 qa-
CTHOCTn, npll JIeqeHUII pH]J.a 3a6onenallllii 6b1JIa c]J.e.'Iaua nOlIblTHa
OnepaTJIBHOro BIIlelllaTeJIbCTBa B o6.'IaCTII I\apOTII]:(Horo clIuyca B nu]:(e
nCpepe31>1I HepBOB II :JI,CTupnaU;lIlI «r.-IOMYCOB» (Leriche, FonLaine
et Froehlich, 1935). Onepau;uR Y]J.a:leUIIH H3pOTU]:(HOrO «f.loMyca»
BOIll.'Ia B xllpypnlqeCKYIO npaHTJlhY II UpOU3BO]J.UTCR IIllOr]:(a II Tencpb
B Tex C:1yqaHX, 1>0r]J.a MP]:(UI>II IIllJ,YT ('uoco6a CHHTb nORblllleHHYIO
qYBCTBUTe.lbHOCTb II npepBaTb nOTOI> aq,q,epeUTHbIX UIImYJILCOB (Ha-
npmlep, npll 6pOHXUaJlbHOii 3CTMC).
!J:a.l1,11 e iilllee pa3BIITIIe np06m MbI UHTepopeu;enU;UII B IlIUpOI>OM
CMblC.le C.1UBa, II B qaCTHOCTII xeMopeu;enU;UII, CBH33HO C IIMen3MlI
HalllUX OTeqeCTBeHHbIX UCCJIc]:(onaTcneii - C q,U3uo.'IOruqeCIWII 1111\0-
.'1011 H'. l\L Bbll>olJa II D. H. llepUllrOBCI\Oro II C Mopq,OJIOrnqCCKOU
1lI1>0.1011 B. II. JIaBpeHTbeBa. Ha I1tOeii naMHTII B JIa60p3TOpllIf
B. n. JIaBpeHTbcBa 6bIJIO nOJIO/HeHO IUI'IaJIO uayqelllIlO 3q,<I>CPCHTuLIX
Heiipouon ImK CUCTeMbI, CIII'H3JHI:mpYlOlIJ;eii 0 COCTOHHIIlI BUYTpeHIIIIX
Opr3UOn. Yme HC33]J:0.rrrO ]:(0 CMCpTU IIIoii nOIwiinbIll Y'IlITC.lJb C :JHTY-
3ua3MOM II YBJle'IeHUCIIt, cnoiicTnCUHhIIltIl CIIIY npn p33pa60Tl\C Tex
IlJIII IlHblX np06JIeIlt, npl1CTYIIIIJI K :JTJIM IlCCJH']J.OB3HllHM. Cpc;:I,n pH]:(a
opranon C ;)TUX n03IIU;UH B. II. JIanpeuTbcn3 IIHTepCCOIJ3ml II uap3-
raHrmUI. Do 3naJIOrUU C ]J.PyrUMII 'IyncTBlITeJIbHL1MJI npu60paIlIJI Oil
17
I
npe.n;nOJIaraJI, 'lTO OHII HBJIHIOTCJI XeMOpeIJ;eIITOpaMII, B 1\0TOpblX «rJIo-
MYCHbIe» I\JIeTI\II, 6y.n;Y'l1l rJIIIaJIbHOH IIpIIpO.n;bl, IIrpaIOT pO:Ib ClIeIJ;II-
cpII'leCI\IIX I\JIeTOI\ - TpaHCcpOpMaTOpOB IIMIIYJIbCa, CBOnCTBeHHYIO He-
CB060.n;HblM HepBHbIM OI\OH'laHIIHM (JIaBpeHTbeB, 1948). O]:(HaI\O, Y'lII-
TblBaH IIX 06IIJIbHYIO CIIMIIaTII'IeCI\YIO IIHHepBaIJ;IIIO, lIepepOlli]:(eHlIe.
CIIMlIaTU'leCI\IIX OIWH'laHlln Ha I\JIeTl\aX l\apOTII.n;Horo lIaparaHrJIIIH
IIPII 3l\CTllplIaIJ;1I1I BepXHero llIenHOro Y3JIa, JlaBpeHTbeB He OTl\a3b1-
BaJICH II OT ]:(PyroH lIX cpYHI\IJ;lIl1 - B l\a'leCTBe meJIe3 BHYTpeHHeH
Cel\peIJ;lIl1. -----
AaJIbHeiinIlIe IICCJIe]:(OBaIIIIH IIo.n;TBepJ{RJIII MHoroe 113 lIepBOHa-
qaJIbHblX IIpe]:(IIOJIomeH-uii, - l.;oe-'lTO HalllJIO ]:(Pyroe OU'bHCHeHUC. Ho
caMoe BamHoe, Ha 'lTO MbI 06paIII;aeM BHIIMaHlIe - 3TO TO, 'lTO pa-
60Tbl lio peIJ;elIIJ;lI11 BHYTPCIIHIIX opraHOB paCmllpllJIII IIpe CTaB-
JIeHIIH II 0 JIOl\aJIII3aIJ;IIII COCy.n;IICTbIX pecpJIeI{COB, He OrpaHII'leHHblX
TOJIbl\O l\ap.n;1I0aOpTaJIbHOn II l\apOTII.n;Hon 06JIaCTHMII, I\al\ 1I0JIaraIOT
Heyma os, Neil ( 1958). XeMOpeIJ;elITOpHaH cpYIII\IJ;IIH 6bl.'Ia 06Hapy-
meHa BO Bcex BHYTpeHHIIX opraHax - B CeJIeaeHl\e, IIO'll\aX, MOqe-
BOM IIY3bIpe, B 1I0JIOBblX meJIeaax, B meJIcaax BHYTpeHHen Cel\peIJ;1I11
(Ha]:(IIO'leqHIIRaX, IIJ;IIToBII.n;Hon), B JIerKIIX, BO Bcex OT]:(eJIaX IIIIIII;eBa-
pllTeJIbHOro TpaKTa (IIHln;eBo.n;e, meJIy.n;Ke, TOIlIWM II TOJICTOM Imllle'l-
HilKe) II B IIIIIII;eBapIlTeJIbHbIX meJIeaax (CJIIOHHbIX, 1I0]:(meJIY]:(0'lHOH,
lIe'leHII) (CM. MOHorpacplIlI: EbiKOB, 1947, qepHIIrOBCKlIii, 1960 II
C60PHIIK pa60T 1I0.n; pe.n;. E. It IlJIe'lIWBOn, 1947).
O.n;HaIw ]:(a.'IeIW He Bea]:(e _cpe.n;1I 06CJIe.n;OBaIlHbIX. TaR HaabIBae-
MblX XeMOpeIJ;eIrropm:,IX:--CTrch M IIMceTCH «rJIOMyc.naH» TRaHb B TOM
1I0HIIMaHIIII, KaK ee onlIcaJI dc Castro B l\apoTll.n;Hon aOHe, XOTH II
BCTpeqaIOTCH lIaparaHrJIIIOHapHbIe Y3eJII\II (oc06eHHo B opraHax
6pIOlliHOH IIOJIOCTII) , HO OHII qaCTO 06.TJa.n;aIOT BceMII IIpIl3Hal\aMII
xpoMacpcplIHHoH TKaHII, CO BceMII IIPllCYIII;IIMII en CTPYKTypHblMII II
rllCTOXIIMlI'leCl\lIMII CBonCTBaMll. C ]:(pyroii CTOpOHbl, Bce «aTIIIIII'l-
HbIe» lIaparaHrJIlI1I 0l\a3aJIIICb «TIIIIII'lHbIMII» B CMblCJIe IIpllHap:JIem-
HOCTII IIX K 3H.n;OKpIIHHon CIICTeMe. HOBbIe BblCORoqyBcTBlITeJIbHbie
MeTO]:(bl rIICTOXIIMIIII, cplI3110JIOrllIT II 6I10XIIMIIII, a TaIDKe 3JIeI\TpOH-
HaH MIIKpOCI\OIIIIJI 1I03BOJIIIJIIT BblHBIITb B HIIX ClIeIJ;n4in'leCRlle rop-
MOHbl - ap:peHaJInH II Hopa.n;peHaJIIIH (npIIHMal\, 1959; RO$s, 1959;
Lever et aI., 1959; Muratori et aI., 1965a, b, 1966; Muratori.
1968) .
M03roBon CJIon Ha]:(IIO'le'lHIIKOB, ROTOpbIH, no lIepBOHa'laJIbHOMY
IIpe]:(IIOJIOmeHITIO It M. EbIKoBa (1947) II E. II. JIaBpeHTbeB3
(1947), OTHOCII.TJCH K XeMOpeIJ;enTOpHbIM opraH aM, 0l\a3aJICH B aTOM
OTHOllieHIIII He e.n;IIHCTBeHHbIM, a O]:(HIIM II3 opraHOB, Kam]:(bln II3
KOTOpblX 06JIa]:(aeT 'lYBcTBIITeJIbHOCTbIO CO cBoeii c06cTBeHHoH iJ!YHK-
IJ;IIOHaJIbHOn OKpaCKon (qepHIIrOBCI\IIH, 1940 a, 6, B, 1943, 1947.
1960; EbIKOB, 1947: JIaBpeHTbeB, 194.7; Jle6e.n;eBa, 1965). 06III;e:n
cpYHI\IJ;I1en ]:(JIH Bcex aHrllOpeIJ;eIITOpOB, 06HapymeHHbix B opraHax
6pIOlliHon IIOJIOCTII, KaK II .n;JIH KapOTII.n;HbIX II aOpTaJIbHblX 06JIaCTeH.
HBJIHeTCH xeMO- II IIpeCCOpeIJ;elIIJ;IIH (EbiKOB, 1947), ORa3bIBaIOIII;aJl
peq)JIeKTOpHoe BJIllHHIIe Ha BblCOTY I\pOBHHOro .n;aBJIeHlIH, pllTM cep-
18
c;;)
e'IHblX CORpa eHIIll, ]:(bIXaHIIC. Ha OCHOBaHlIH aHaml3a OrpOMHorO
.MaTeplIaJIa 110 CPlI3UOJIOnIH UHTepOpen;enn;IIU (II XeMOpen;enn;IIII), no-
.JIyqeHHoro B CBOeH JIa60paTopuu, It M. EbiROB AeJIaeT O'IeHb Bam-
Hoe 0606r:u;eHue. OH nUIlICT, 'ITO UHTepopen;elITopbl BHYTpeHHIIX opra-
HOB HeJIb3H OrpaHU'IUBaTb TOJIbRO anrnOpCn;eIITOpaMll, npeAnOJIaraH
.nX cymecTBoBaHlIe B CaMlIX TRaHHX, a He TOJIbRO B Tex COCYAax,
ROTOpbIe IIX BaCRYJIHpIl311PYIOT.
o JIORaJIU3an;UU xeMopen;enTopOB BHe COCYAllCTbIX CTCHOR, llMeH-
HO B TRaHHX, nllllleT II B. H. qepHllroBclmH (1943, 1947, 1960).
CTOJIb me CHCnTll'IeCROe OTHOllleHlle R orpaHlI'IeHlI1O COCYAIICTOH
'IYBCTBIITeJIbHOCTJI TO.TIhRO ABYMH OUJIaCTHMU - cIIHoRapoTll]:(HOH II
RapAlIoaopTa.TIbHOii - 6bIJIO Bb1CRa3aHO II MopcpOJIOraMll, 60raTblH
.cpaI\TUqeCI\UII MaTCpUaJI ROTOpbIX 0 CTpYRType COCYAIICTbIX pen;enTO-
pOB onpOBepraJI 3TY 1\0Hn;Cnn;UIO (T. A. fpuropbeBa, 19;j4) , IlJIII,
BO BCHROM c.TIy'rae, orpaHU'IUBaJI ee 311a'IeHlIe (AOJIro-Ca6YPOB, 1945,
1958; IlBaHoB, 1949). B MOHorpacpull T. A. fpllropbeBoH (1954)
J.(aH aHaJIll3 np06JIeMbI UIiHepBan;UU RpOBeHOCHbIX COCY]:(OB II rOBopUT
CH, B 'IaCTHOCTU, 'ITO tIYBCTBUTCJIbHbIMII allIIapaTaMU, perUCTplIpylO-
r:u;IlIlIH XIfMU'IeCRUe C]:(BllrU B opraHU3Me, HBJIHIOTCH COCYAIfCTO-TRa-
HeBbIe pCn;enTopbl RalIIIJIJIHpHOrO PYCJIa. C 3TOH TO'IROII 3peHIIH IIOJI-
HOCTblO COrJIalll3eTCH B. A. AOJlro-CaGypon (1956). T. A. fplIropbe-
BOll RpIITII'IeCRll paCCMOTpeHbl U BonpOCbI 0 «rJIOMYCHOH» II napa-
raHrmIOHapHOH THaHU II BbICHa3aIIO OTplln;aTeJIbHOe OTHOllieHue H IIX
yqaCTUIO B pen;elITopHOH cpYHHn;uU. ct>aRTll'IeCHlle A3HHble T. A. fplI-
ropbeBOll (1954), xapaRTepU3YIOr:u;ue cHa6meHIIe BereTaTIIBHblMU II
acpcpepeHTHbIMII HepBaMII RpoBeHocHoH CIICTeMbl MJIeI\OIIIITalO IIX,
1I03BO:IHIOT CYJJ.I1Tb 0 ABIIraTeJIbHb1X II 'IYBCTBIITeJIbHbIX IIHHepBan;II-
()HHbIX IIpu60pax Ha HpOBeHOCHblX cocYAax Bcex HaJIII6pOB, nORa3bI-
BalOT llX WYHRn;UOHaJIblibIe B03MOlliHOCTU H BocnpUHTIIIO XUMU'IeCHIIX
II MeXaHll'IeCHUX pa3]:(pameHuH II H pCWJIeRTOpHoH peaRn;UU rJIaAHllX
:JIIbIIlIn; COCYAUCTbIX CTeHOH Ha 3TII B03AciicTBllH. CJIOmHO YCTpoeH-
HbIe U IlIUpOHO paCllpOCTpaHeHHble HcpBHbIe OI\OH'raHllH Ha caMIIX
COCYAIICTbIX CTeHl\aX cnoc06Hbl, O'ieBIIAHO, OUCCne'IUTb pecpJIel{TOp-
Hhle peaRn;IIU, BblpamalOlIJ;ueCH B cocY.lUICToii nO]:(BJIlliHOCTlI, 6e3 Y'Ia-
CTUH «r.TIOMycHOH» TRaHll. 13 TO me BpeMH cpaBHUTCJIbHaH orpaHII-
"IeHHOCTb B pacnpOCTpaHCHUII «r.TIOMYCOB» OCTaeTCH CJIa6b1M MeCTOM
]];allie ]:(o'IH Tex llCCJIeAoBaTe.TIeH, ROTopble np1l3HalOT 3a HIIMII .pYHR-
n;UIO COCY]:(llcTo-TRaHeBon XlIMlI'IeCROH 'IYBCTBlITeJIbHOCTII (I\.ynpIIH-
HOB, 1959).
IIcCJIeAOBaHue T. A. fpuropbeBon, npoBeAeHHoe Ha M,'IeROIIIITalO-
II{IIX, He paCRpblBaeT, OAHaRO, IIpU'Ilill II 3Ha'ieHIIH HaJIU'IIIH «rJIO-
MYCOB» B COCY]:(IICTbIX CTCHI\aX TaIUlX apTepllH, Han: COHHbIC, JIerO'I-
RbIe lIJIlI aopTa. ATunll3M CTpoeHllH II HeROTopbIX rUCTOXIIMU'IeCRIIX
CBonCTB «rJIOMYCOB» OCTaeTCH HenOHHTHblM II npOAOJImaeT CJIymllTb
()AHllM 113 IIOBO]:(OB AJIH On;eHRII IIX RaR CTPYRTYP, BocnpIIHIIMalOll(IIX
pa3]:(pameHIIe.
B HaCTOHr:u;ee BpeMH HaROlIlIJIOCb AOCTaTO'IHO .paliTOB B nOJIb3Y
3HAORpuIlHoro xapaRTepa «rJIOMYCHbIX» 06pa30B3HIIH II IIMeeTCH
19
.n;OCTaTOqHO OCIIOBaHHll .n;.TIR TOrO, qT06bI CqlITaTb IIX lIaparaHrJllIO-
HapHLIMH. :Ka3aJIOCb 6bI, «rJIOMYCbJ», RaK lIaHau;eH Xel\lOpeu;em::{HII
RpOBeHOCHblX COCY]:(OB, JJ;O.'1iKHbl 6bI 6blTb OTBeprHYTbI, o.n;HaKO IIOTOK
JIHTepaTypbl 0 CHHOKapOTII.n;HbIX H RapAIIOaOpTaJIbHblX «rJIOMycax-
xeMopeU;enTOpaX» He y6blBaeT. IToCJIe.n;OBaTeJIII 3Toro HanpaBJIeHIIR,
B npOTHBOBec orpOMHOMY qHC.;IY .paKTOB, KOTOpbIe He YRJIa.n;bIB3IOTCH
yrlW B KOHU;enU;I1IO XeMOpeu;enTopHoii: .pYHKU;IIII «rJIOMYCHbIX» R:'IeTOR.
IIbITaIOTCH 06'bRCHHTb ]:(3HHble .paKTbl co CBOHX 1I03HU;UH H C03.n;aIOT
HOBbIe He060CHOBaHHble nmOTe3bI. :KapOTH]:(Hble H aOpTaJIbHble «rJIO-
MYCHble» K.ileTRH npo]:(O.'1If,aIOT H3yqaTbCH IICKJIIOqIITe.'1bHO C TOqRU
3peHIIR IIX XeMOpeu;enTOpnOll .pYHKn;1I11 (CM. 0630p Comroe, 1964).
B I\IOnorpa.pIlH W. E. Adams «The comparative morphology of
the carotid body and carotid sinus» (1958) aBTOp YTBepiK.n;aeT,
qTO pa60Tbl de Castro (1926, 1928) O:JHaMeHOBaJIH c060ii: HOBYIO
3noxy B IICCJIe]:(onaHIfII n HaCblTIIJIII co.n;epiKaHlieM «rJIOMYClIyIO»
CTPYKTYPY 6H.pypKaU;IIII 06I.qeii COHHOii: 3pTepllH.
B HalllY 2a.n;aqy He BXO.n;IIT paCCMOTpeHIle COCTOHHIJH Bcero :norn-
Bonpoca II ou;eHI;:a MaTepllaJIOB, CBR3aHIIbIX C np06.'1eMOH xeMopeU;en
U;I1I1. :KocnYTbCH ee OKa3aJIOCb neII36eiKHblM TOJIbKO B TOH II:IOCROCTU.
B KOTOpoii OHa 3aTparIIBaeT naparanrJIIIOnapnyIO CIICTeMY II TeCHO
IIepen.TIeTaeTCH B HeROTopLIX pa:J.n;e.TIax C ee Mop.po.pynRu;nOHa.'1bHbIM
n03namIeM. ITo CYI.qeeTBY npllxo.n;lITCH RacaTbCH ne B U;eJIOM Bceii:
30nbl, BoenpllnlIMalOI.qen XIIMII3M KpOBlI, a TOJIbKO Tex K'IeTOqHbIX
CTpYRTYP naparaHrmlOnapHOH TI,alllI, KOTOpbIe Hlw6bl HB.'1RIOTCR OT-
BeTCTBeHHbIMH 3a OCYI.qeeTB.rrenIIe xeMopeu;enTopnoro IIpou;ecca. :Kpo-
Me Toro. C03pe.TIa HaCTOHTeJIbnaR He06xo]:(I1MOCTb CIICTeMaTII3aU;I1I1 Ha-
KonllBIlIIIXCH .paKTOB B 06JIaCTU 113yqeHIIR naparaHr.'1l1eB, ROTOpaJl
1I0MOrJIa 6bI e]:(eJIaTb na]:(JIerKaI.qlle BbIBO.n;bl, COOTBeTCTBYIOI.qIle CO-
BpeMeHnOMY ypOBHIO HayqHbIX 3ualIIIll 0 naparanrJInonapHoii CIICTe-
Me. TaKaH nOTpe6noCTb ]:(llIiTyeTcH TeM, qTO Mop.pOJIOrll 1I0TepHJIIt
e]:(I1HYIO TOqKy 3penllR na MOp.pOJIorIlIO II rIICTo.pII3IIOJIOrIlIO napa-
raHrmwnapnoii: CIICTeMhI II, B 113BeCTHon Mepe, nBO]:(HT B 3a6.'1YiRJIe-
IIIIe «PII3JIO.TIOrOB, .papMaKo.'101'OB II naTO.TIOrOB. ITOqTU KaiK.n;oe .p1I-
3JIOJIOfllqeeKOe II.TIII .papMaIWJIOfllqeCKOe IICCJIe]:(OBanIJe 0 pe.pJIeRCOren-
nbIX 20nax RapOTII]:(HbIX II aopTa.'1bIlbIX 06JIaCTen naqlIHaeTCH C an3-
TOMo-rUCTOJIOrllqeCROrO BEe.n;eHJlR, B KOTOpOM aBTopbl no BnO.TIne no-
HRTHbIM npllqlllIaM, ne 6y.n;yqJl cneU;IIa.TIJlCTaMJI,- 6epYT rOTOBbI
Mop.pD.1IOrIlQeCKlIe CBe.n;eHIIH. Tal{, Monorpa.pIlH C. B. AHlPIROBa .n
M. JI. BeJIenblw1'O (1962), KOTopaH co]:(epmllT Mnoro IIOBb1X cpn3110-
JIoro-.papMaKQ.1IOrIlQeClillX .palnoB, .n;aiKe 03ar.TIaB.1Jena «<})apMaKOJIO-
rIlH XlmIlopeu;eUTopOB KapOTIJ]:(H01'O KJIy60QKa». <D1I:JIIOJIOrlI n .papMa-
ROJIOrJl .n;06blBaIOT HOBLIe .paKTbl, pa3BIIBaIOT HOELle npe;J.CTaB.'IenIlR.
a npllKJIa.n;blBaIOT I1X K Mop.po:wrIJQeCHIIM CTpYRTypaM, nay'moe 3Ha-
'1enlle KOTOpbIX OrpalllIQlInaCTCR .n;aBIIO YCTapeBllIlIMIJ cne.n;ennRMlI
20-x ro.n;OB TCI,YI.qerO eTOJICTIlH.
EI.qe XYiKe 06CTOIlT Ae.'10 C 3TIIMII BOnpOCaMIJ n 06JIacTiI naTO-
JlOrnll. B OIIKO.'10rllll B03nIlK.TIa U;CJlaH rJIaBa, 03arJIaB.'1eHlIaR «Xe-
MopeU;CIITopIlbIC onyxoJIlI» (CM. 0630p Burman, 1956). B ;)TY rJIaBY
20
BKJIIOqeHbl OnYXOJIH, HCXOp;HlIwe H3 «aOpTaJIbHOrO H RapOTHp;HOrO CH-
HycOm), H3 (<apTepHO-BeH03HbIX H RORU;HrHaJIbHbIX rJIOMYCOB», H3
(<RapOTnp;HbIX rJIOMYCHblX TeJI», H3 (<aOpTHRO-nYJIbMOHaJIbHbIX TeJI»
H H3 P;PyrHx (<HeXpoMa!f>!f>HHHbIX naparaHrmleB (IOrYJIHpHOrO, BaraJIb-
HOrO, U;HJIHapHoro, Rapp;HOaOpTaJIbHoro»). AJIH BCeX :3TnX onYXOJIeR
XapaRTepHO OTCYTCTBHe CeRpeU;HH ap;peHaJIHHa H HOpaJ];peHaJIHHa
H OTpHu;aTeJIbHaH xpoMa!f>!f>HHHaH peaRU;HH. B :3TY me rJIaBY - qTO-
OqeHb nOKa3aTeJIbHO - BKJIIOqPHbl onYXOJIH, npOHCXOp;Hm;He H3 M03ro-
BOro CJIOH Hap;nOqeqHHIWB, 1I3 CROnJIeHHR xpoMa!f>!f>HHHoH TRaHH B
peTponepHTOHeaJIbHOM npOCTpaHCTBe, CBH3aHHble C TOpaKO-JIYM6ap-
HOR CHMnaTHqeCROH u;enOqKOH II C BHCU;epaJIbHbIMH CnJIeTeHHHMH;
Bce OIflI p;aIOT THnHqHYIO xpoMa!f>!f>HHHYIO peaKU;HIO H ceKpeTHpYIOT
ap;peHaJIHH H Hopap;peHaJIHH. llepBaH rpynna onYXOJIeR Ha3bIBaeTCH
xeMop;eRTOHaMII, BTopaH - !f>eoxpoMOU;HTOMaMH. 06'bep;HHeHHe HX B
OP;HY rpynny OCHOBaHO Ha cBoeo6pa3HH HX onYXOJIenoro pOCTa, xa-
paRTe pH oro P;JIH TKaHeR HeRpmmTOp;epMaJIbHOrO rrpOHcxomp;eHHH. Ta-
ROR pOCT 06YCJIOBJIeH rHCToreHC30M naparaHrJIHOHapHOR CHCTeMbl,
pa3BHBaIOm;eRCH H3 He Hpo:mTop;epMbl. Ep;IIHcTBeHHO rrpaBHJIbHblM
6blJIO 6bl CqHTaTb «xeMopeu;enTOpHble» onyxOJIH, HapHP;Y C !f>eoxpo-
MOU;HToMaMH, pa:moBHp;HoCTHMH HeRpO:3RTOp;epMaJIbHOrO 3JIOKaqeCT-
BeHHoro HJIH p;06pOKaqeCTBeHHoro pOCTa, no He Ha3blBaTb HX xeMO-
peu;enTopHbIMH. I1CXOP;H H3 TOro, qTO XeMOpeI enU;HeR 06JIap;aIOT Bce
TKaHeBbIe CIICTeMbl Bcex opraHOB, a Mbl pa3p;eJIHeM TOqRY 3peHHH,
KOTopaH npH3HaeT ;)TO TBepp;o YCTaHOBJIeHHblM !f>aKToM, cym;ecTEoEa-
HHe rJIaBbI 0 xeMopeu;enTopHbIX onYXOJIHX He onpaBp;aHO HH Teope-
THqeCKH, HH npaRTHqeCI{H. Cop;epmaHHeM TaROR rJIaBbl p;OJImHbl 6bI
6blTb onYXOJIH Bcex BHYTpeHHHX opraHOB, IIOCROJIbRY Bce OHH o6JIa-
p;aIOT xeMopeu;elITopHblMH annapaTaMH, qTO, pa3YMeeTCH, 6blJIO 6bI
napap;OI\COM.
focnop;CTBYIOm;He B3rJIHp;bl Ha xeMopeu;enU;HIO KaR 6bl paCCeRJIH
naparaHrJIHOHapHYIo CHCTeMY Ha p;Be rpynnbl opraHOB, H3yqeHHe RO-
TOpblX nOIllJIO no P;BYM pa3HbiM HanpaBJIeHHHM. llo OTHOllleHHIO I)
OP;HOH rpynne HCCJIep;OnaTeJIH aaHHJIHCb nOHCKaMH HOBblX «p;OKa3a-
TeJIbCTB» peu;enTopHoII !f>YHRU;HH KJIeTOK H paCIlIHpeHHeM 06'beRTOB
B cpaBHHTeJIbHO-Mop!f>OJIOrHqeCKOM pa3pe3e. 113Y'leHIIC BTOpOR rpyn-
nbl - (<HCTHHHbIX» xpoMa!f>!f>IIHHbIX naparam'JIHeB, rJIaBIIblM 06pa30M
Hap;nOqeqHHKOBbIX, B0306HOBllJIOCb TOJIbKO B 50-x rop;ax, Rorp;a HH-
Tepec K HHM IIOBbICHJICH B CBH3H C HOBblMH p;aHHbIMH 6IIOXHMHH.
OTRpblTHe Hopap;peHaJIHHa, BblHCHeHlIC ero !f>H3HOJIOrHqeCKOrO
p;eRCTBHH (CM. 0630p Euler, 1956), npHHap:JIemHOCTb K rpynne Ra-
TeXOJIaMHHOB H 6HocHHTe3 no cxeMe Blaschko (1939) If Holtz
(1939) (CM. 0630pbl MaTJIHHOR H MeHbIllHKoBa, 1964, 1967) nOJIY-
qHJIH illHpOKHR pe30HaHC B Mop!f>OJIOrHqeCKHX HCCJIep:OBaHHHX, Bblpa-
3HBllIIIXCH B nOIICKax CTPYKTYP, n IWTOPblX JIOKaJIH3YIOTCH KJICTOq.
Hble aMIIHM.
I1cnOJIb30BaHHe COBpCMCIIHblX MeTop:OI3 rIICTOXIIMIIqeCROrO II :3JIeK-
TpOHHOMHl\pOCKOIIHqeCKOrO IICCJIep:oBaHHR B npIIJIOmeHHH K CIIMnaTO-
ap;peHaJIOBOR CHCTeMe, HapHP;Y C o6HapY1KeHIIeM MHorHX HOBblX !f>aK-'
:H-
TOB H C03p;aHHH HOBblX rHIIOTe3, Bbl3BaJIO pHP; TPYP;HOCTeH, OTHOCH-
II\HXCH R HHTepnpeTa:IWH nOJIyqaeMbIX p;aHHbIX. 3aTpyp;HemIH H npo-
THBOpeqHH CRa3aJIHCb H npH COnOCTaBJIeHHH YJIbTpaCTpYRTypHOro H
rHCTOXHMHqeCROrO aHaJIH3a C pe3YJIbTaTaMH 6HOXHMHqeCRoro H3yqe-
.HHH ap;peHaJIOBbIX OpraHOB. B HaCTOHl ee BpeMH ell\e HeT p;OCTaTOq-
HbIX OCHOBaHHH, qT06bI p;eJIaTb BbIBOP;bI no HeROTopbIM HCCJIep;yeMbIM
.H P;HCRYTHpyeMbIM BonpocaM H npHHTH R RaRHM-JIH60 cepbe3HbiM
06061IJ;eHHHM. :=ho OTHOCHTCH H R o eHRe P;BYX THnOB RJIeTOR B M03-
rOBOM CJIoe Ha.n;nOqeqHHROB H B p;pyrHx naparaHrJIHHX, CHHTe3HpyIO-
III;HX p;Ba ropMOHa - ap;peHaJIHH H Hopap;peH8JIHH.
EOJIblllOH HHTepec Mopq,OJIOrOB R Bonpocy 0 CYlIJ;eCTBOBaHHH
.)];BYX TunOB xpOMaq,q,HHHbIX RJIeTOR Bblpa3HJICH B TOM, qTO, HaqHHaH
C 50-x rop;OB, BO MHorHX CTpaHax EBponbl H B CIIIA 1I0CTeneHHO
B03HHRJIlI HayqHbIe eHTpbl, B ROTOpblX pa3pa6aTbIBaloTcH BonpOCbl
Mopq,OJIOrHH ropMoHo06pa30BaHHH B M03roBOM CJIoe Hap;nOqeqHHROB.
llHoHepaMH B :3TOM p;eJIe 6bIJIH illBep;CRaH illROJIa HCCJIep;OBaTeJIeH
BO rJIaBe C Hillarp H Hokfelt (Hokfelt, 1951; Hillarp, Hokfelt,
1953, 1955) H JIa60paTopHH Eranko, B -<1>HHJIHHP;HH (Eranko, 1951;
Erank6, Hopsu, 1958; Eranko, Palkama, 1959; Eranko, Hau-
ninen, 1960). BCRope nOCJIe ony6J1lmoBaHHH nepBbIX HCCJIep;OBaHHH
nOHBHJIaCb cepHH pa60T H3 JIa60paTOpHH Coupland B AHrJIHH
(Coupland, 1953, 1959, 1965; Coupland, Pyper, Hopwood,
1964; Coupland, Hopwood, 1966), Picard H Vitry BO
<1>paHIJ;HH (1959, 1964), Klein H Kracht B <DPf (1958), Giorda-
no- Lanza B IlTaJIHH (1965), Benedeczky C coaBTopaMH B BeHrpHH
(1964; Benedeczky, Puppi, Tigyi, 1965; Benedeczky et al" 1965;
Benedeczky, 1967a, b), TramezzRni C coaBT. B ApreHTlJHe
(Tramezzani et a!., 1964), Wood C coaBTopaMH B CIIIA (1962,
1963; Wood, Barrnett, 1964). B COBeTcRoM COI03e :3TOH np06JIeMOH
HHTeHCHBHO 3aHHMaeTCH 3. C. Ka HeJIbCOH C COTpyp;HHRaMH (Ka -
HeJIbCOH, 1964, 1966a, 6, 1967a, 6, 1969; llop;ropHaH, 1965,
1967, a, 6, B). KpOMe nepeqUCJIeHHbIX JIa60paTopHH, np06JIeMa p;n-
Mopq,H3Ma xpoMal!Jl!JHHHbIX RJIeTOR u3yqaeTCH H B p;pyrHx MeCTax H
C Ramp;bIM rop;OM qHCJIO HTOJIOrHqeCKHX H HTOXHMHqeCRHX pa60T
B :3TOM HanpaBJIeHHH YBeJIHqHBaeTCH.
Ha COBpeMeHHOM :3Tane pa3BHTHH yqeHHH 06 ap;peHaJIOBOH CHCTe-
Me np06JIeMa RJIeTOqHoro P;HMopI!JH3Ma npeBpaTH.'IaCb B OP;HY H3
eHTpaJIbHblx. HaCTOHIIJ;HH nepHop; MomHO oxapaRTepH30BaTb nepHo-
J];OM HaROnJIeHHH l!JaRTOB. Op;HaRO yme H ceiiqac RpaHHee YBJIeqeHHe
nOHCRaMH npH3HaKoB )]:HMopI!JH3Ma KJIeTOqHblX THnOB xpoMal!Jl!JHHHblx
opraHoB BbI3BRJIO HeRpHTHqeCROe OTHomeHHe HeRoTopbIX aBTopoB RO
MHOfHM nOJIyqaeMbIM l!JaRTaM, nop;qac O,');HOC.TopoHHee TOJIKOBaHHe H
paccMoTpeHHe HX BHe CBH3H C p;pyrJlMH nop;06HbIMJI HBJIeHHHMJI HJIII
IIpO eCCaMH, MorYIIJ;JlMH CJIymHTb OCHOBOH P;JIH 06'bHCHeHHH :3THX
1Cf!aRTOB. HanpHMep, :3TO OTHOCHTCH K Cm{peTOpHbIM IJ;UKJIaM B XPOM-
ftl!Jl!JJlHHOH RJIeTRe II aHaJIII3Y CTPYRTYP, yqaCTBYIOIIJ;JlX B 6J10CJlHTe-
3e MOHOaMJlHOB. C O]J,HOH CTOpOHbI, He Bcerp;a yqHTbIBaIOTCH MOpI!JO-
JIOrH'IeCRHe nORa3aTeJIH I!JYHR HOHaJIbHbIX CTap;HH CeRpe JlH, ROTO-
22
pble MDrYT CDBna]:(aTb C npll3HaRaMII MCTDqHDrD P;IIMDpqJll3Ma, a,
C P;PyrDH, nepep; Mop/f>DJIOraMII CTDIIT CJIDmHaH 3ap;aqa onpep;eJIeHIIJI
He TOJIbRD ap;peHaJIIIH- II HDpa]:(peHaJIHHCDp;epmaIIWX RJIeTDR, a 11
JIDRaJIII3aIJ;I1U BHYTplllmeTDqHDrO 6l1DClIHTe3a rDpMDHDB II erD MDp/f>O-
JIDrllqeCRDrD npDJlBJICHIIH. B nDUCRax CTPYRTYP, xapaKTepll3YIDIIJ;IIX
DTJIII'IllTeJIbHble qepTbl P;BYX Tun DB KJIeTDR, He Bcerp;a yqUTbiBaIDTCJI
JIDRaJIII3aIJ;lIJ1 /f>epMeHTDB, RDTDpaH y'IaCTByeT B MeTUJIHIJ;UII HDpap;pe-
HaJIlIHa, pacnpc]:(eJIeHlIe rrpep;llIeCTBeHHIIRDB, IIX ClIHTe3 B KJIeTRaX II
p;pyrllc npDIJ;CCCbl.
o P;BYX RJIeTOqHblX TIIII8X xpDMa/f>/f>IIHHDH TRaHII Y HlI31l1l1X /f>DPM
nD3BDHDqHbIX yme Dny6mmDBaHD MHDrD pa6DT. B CDDTBeTCTBYIOlIJ;lIX
rJIaBaX rDBDplITCH 0' /f>aKTII'ICCKDU: CTDpDHe :3TIIX IICCJIep;DBaHIIH. Ho
cnDpHble BDnpDCbI ]:(11- II MDHDMDp/f>1I3Ma xpDMa/f>/f>IIHHbIX MeTDR, BD
1I36emaHue nDBTDpeHIIH, paCCMaTpllBaIDTCH B rJIaBe IX, nDCBHlIJ;eHHDR
rllcTD/f>II3I1DJIOrllll MD3rDBDro CJIDH Hap;nDqeqHIIRDB.
B CBH311 C 1I3JIDmeHHbIM, n;eHTpaJIbHDH npD6JIeMOH HalilerD IIC-
CJIep;DBaHIIH JlBJIHCTCJI DIJ;CHRa /f>YHKIJ;IIDHaJIbHDrD 3HaqeHlIH Bcex lIa-
paraHrJIIIDHapHblX DpraHDB BblCllIlIX nD3BDHDqHbIX, B TDM qIlCJIe MJIe-
RDnIITaIDlIJ;lIX II 'ICJIDBCKa, II D60cHDBaHlIe npIlHap;JIemHDCTII ux - K
ep;IIHDM :3HP;DRpUHIIOM ap;peHaJIDBDii CIICTeMe. C :3TDM IJ;eJIbID, HapHP;Y
C I1CIIDJIb3DBaIlIlCM cD6cTBcnHDrD /f>aKTII'IeCKDrD MaTCpllaJIa lID :3M-
6pllDreHe3Y II cpaBRIITCJIbllDll 1.IOp/f>DJIDfllII naparaHrJIlIeB Ha npe]:(-
CTaBIITeJIHX DTp;eJIbHbIX rpynn nD3BDHO'lRbIX (l\pyrJIOpOTbIe, aM/f>II-
61111, penTIIJIIIH, nTIIIJ;bl, MJIeRDnIlTaIOlIJ;lIe), 6b1JIO npllBJIeqeHD nO' B03-
MDmHDCTII Bce, TJ:TO mmeCTIIO B JIIITCpaType nO' :.JTIIM BDnpDcaM, II
1I3Y'ICHa /f>al{TII'IeCRaH CTDpDHa MDp/f>DJIDrllqeCRIIX IICCJICp;DBaHliM Y
Bcex KJIaCCDB IID3BDHDqRblX B /f>UJIO- II DIITDreHe3e. TaKDII cpaBHII-
TeJIbHD-MDp/f>DJIDrllqeCRII:ll II cpaBHIITeJIbHD-:3M6pUDJIDrllqCCKIIII nop;xo-
JIbl, a TaRme 3HaRDMCTBO C /f>YHRIJ;I1DHaJIbHDH :3BOJlIDIJ;I1eH BceH CIlCTe-
Mbi nD3BomWlI nDp;BeprllYTb RplITll'IeCKDMY paccMDTpeHIIID MHDrIle
RDHRpeTHbIC IIDJIDmeHIIH II D61IJ;IIe IWHIJ;enIJ;UII 8BTDPDB, BD3HIIRlillIe
lID 3TIIM BonpDcaM B nDCJIep;HIIe p;eCHTIIJICTIIH.
.lI.JIH ]:(DCTIImenUH HaMe'IeHHDM IJ;eJIII aBTDpOM CDopaH II npDaJTaJIII-
3I1pDBaH JIIITeparypHbI:ll MaTepllaJI lID TpeM HanpaBJICHUHM: nO' q,YHR-
IJ;I1DHaJIbHDII 1.IOp/f>DJIDrIlll CDCYP;IICTDH CIICTeMbI y pa3IIbIX Rm'lCCDB
nD3BOHDqHbIX mIlBDTHbIX - RpyrJIDpDTblX, pbI6, aM/f>I16I1M, penTH]HfK,
IITHIJ;, MJI(mDnIlTaIDIIJ;UX II y 'ICJIDBeRa, lID MDp/f>DJIDrnll nepll/f>pPIl-
qeCRD:ll HcpBnDM CIICTeMbI - ee HeiipDHDB II DT]:(eJIbHbIX 3BeHbeB pe/f>-
JIeRTDpHblX p;yr, II paCCMDTpena C Tex me IID31IIJ;U:ll naparaHrJIIID-
HapHaH CIICTeMa, HaqIlHaH C RpyrJIDpDTblX II KOHqaH MJIeRDIIHTaIO-
IIJ;HMII. IIpDBep;eHHbIH aHaJIII3 p;aJI BD3MDmHDCTb IIpDCJIep;IITb ypDBHII
MDp/f>DJIDrIl'IeCKDrD pa3BlITUH Tpex CIICTeM, IIX /f>YHRn.IIDHa.'IbHble BD3-
MDffiHDCTII II 3ap;aqII B pa3HbiX rpYlInax nD3BOHOqHbIX - DT HII31I1I1X K
BblCIlIUM - II YCTaHDBIITb HeKDTDpble D61IJ;IIC II qaCTHbIe MDp/f>DJIDrIl-
qeCRIIe 3aKDHDMepHDCTII, RDTDpble JIemaT B DCHOBe /f>I13I1DJIDrnqeCROR
]:(eHTeJIbHDCTII cnCTeMbl Ha pa3HbiX ypDBHHX /f>IIJIDreHe3a U Ha pa3HbiX
CTap;I1J1X DHTDreHeTII'IeCRDrD pa3BIITIIJI DpraHII3MDB. B npDIJ;eCCe I1C-
CJIe]:(DBaHII:ll OTp;eJIbnbIX D6"beKTDB - BHe IICTDpll1l pa3BIITIIH mIlBDT-
23
HbIX OpraIUI3MOB - 3TH 3aROHOMepUOCTil OCTanRJIIICb He::>RMe'IeHHbIMH,
no aHaJIH3 cOBoRynHocTH BCCX .o;aHHbIX n03BOJIIIJI HaM 060CHOBaTb
IIpCCMCTBeHHYIO CBH3b Mem.o;y aHaTOMHqeCRHM pacnpe.o;eJIeHHeM II
rHCTOJIOrHqeCRoii: xapaRTepHCTHROH xpoMaiPiPuHHoii: TRanli y HII3ll1HX
n03BOHOqHbIX iHHBOTHbIX, .o;HiPiPY3HO pacceHHHYIO B RpynHbIx apTe-
pHaJIbHbIX H BeH03HbIX COCy.o;HCTblX CTBOJIaX TYJIOBHIIW, B nO]:(RJIIO-
qHqHblX, ma6epHbIX II Mempc6epHblX apTepHHX, C TeMH yqaCTRaMH
COCy.o;HCTOH CIiCTeMbI BbICIlIHX n03BOHOqHLIX II M,TIeROnllTaIOIl\HX, B RO-
TOpbIX 3aJIOmeHa «rJIOMYCHaH» TRaHb. « fJIOMYCbl» BbICllIllX n03BO-
HOqHblX U xpoMaiPiPllHnaH TRaHb npllcep.o;eqHblX COCy.o;OB HalllUX no-
3BOHOqHLIX - rOMOJIOrHqHbIe CTPYRTypbI.
HaR 113BeCTHO, .o;yra aopTbI, JIerOqHbIe apTepIlll H COHHbIe apTe-
pHll pa3BIIBaIOTCH B iPllJIOreHe3e II3 ma6epHbIX apTepUH. BCR CHCTe-
Ma iHa6epHLIX COCy.o;OB npCTepnCBaeT CTPYRTypHLIe npc06pa30BaHIIH,
Be.o;Yll\IIe IIOCTeneHHO JIH60 R nOJIHoii-aTpOlpllll apTepllaJIbHLIX II Be-
H03HbiX .o;yr, JIll60 R 06pa30BaHllIO py.o;llMeHTapHbIX CTPYHTYP B BII]:(C
npOTOROll, COc.o;llHHTeJIbHOTRaHHbIX THmeH II CBH30R. BrrcpBLIe nall-
60Ree pC3Rlle nepeCTpOHRU ma6cpHoro annapaTa HacTynaIoT npll nc.
pexo.o;e Ha Ha3CMHbIH 06pa3 iHll3Hll II pa3BHTllH JIerO'IHOro .o;bIxaHllH.
MopiPOJIOrlIqeCRaH :JBOJIIOD;IIH ma6cpHoro cocy.o;llCToro annapaTa He
MOrJIa HC OTpa3HTbCH Ha ]:(pyrllx TR3HeBbIX 3JIeMCHTax iHa6cpHOH
CllCTCMbl, ROTopble, 6y.o;yqll BOBJIeQCnbl B 3TOT npOD;CCC, npeTCpneJIH
CTOJIb iHe rJIy60Rlle nepCCTpOHRll, Hal\ II C06CTBCHHO COCy.o;llCTOC PYC-
. JIO ma6cp. H TaRHM TRaHCllLIM 3JIeMeHTaM npllHa.o;JICII\aJIll II i\OMn-
JICRCLI xpoMaiPiPllHHLIX HJICTOK.
XpoMaiPiPllHHaH TRaHb rrpllcep.o;CqnOM 06JIaCTU HU31l111X n03BOHOq-
HbIX, RaR H BCCX p;pyrHx 06JIaCTeH TYJIOBIIll\a, TeCHO cnaHHa C CO-
Cy.o;llCTOH CTCHROH II HBJIHCTCH ec COCTaBHOM qaCTbIO. 13 TCX OTpe3-
Rax coeY,lJ,IlCTOro Rnnap3'fR, ROTOpLIe no.o;BcpraIOTCH aTpoiPllll IIJIH
IIpeBpall\aIOTCH B py.o;llMCHTapHbH' 3aqaTl\H, OHa TOII,e llJIll IICqe3aCT,
llJIll CTaHOBllTCH py.o;IlMeHTOM. Yme O.o;I1H :JTOT npon;cce cnOCOOCH
npHBeCTll Ha onpe.o;CJICHHLIX :JTanax :mOJIIOII,IIII no:mOHOqHLIX R B03-
HIIRHOBeHIlIO ne]:(opa3BIITblX OCTpOBlWB xpoMa«I>iPlIHHoM TRaHII, pacno-
,1I0mCHHbIX B cOCy.o;lICTLIX CTeImax IlIci1Hoii: II rpy.o;HOII 06.uaCTeR. Hc-
.o;opa3BIITIIe IIX, RaR II JII060ro py.o;lIMeliTapHoro oprana, MomCT BbI-
pamaTbcH B 1"IICTOJIOrlIqeCIWM cTpoenlIlI, B D;1l'I'OJIOrIlqeCl\OH CTPYRTY-
pc II B D;IITOXIIMII'IeCI\HX CBoIIcTBax, a p;Jm :JII.o;ORpHUHOii: meJIe3UCTOH
RJICTIHI - B ncpBYIO OqCpe.o;b B OTCYTCTBIIU U,TUI CHlIII,eHIIII cnoc06-
HOCTH R BbJpa60TRc cncD;HiPHqeCI\HX ropMOHOB.
HapH.o;y C pcrpeccHBHLIM pa3BHTllcM xpoMa iPlllIHoH Tlia HII , TCC-
HO CBH3aHHI,Il\I e 061l\HMH nYTHMII 3BOJIIOn;HOHHoro npe06pa30BaHHH
npHcep.o;e\lHLIX ma6epHLIx .o;yr, xpoMaq)(JJHHHaH TRaHb UPOXO.o;IIT
. CJIOmUhIH nYTb MopiPOJIOrHqeCRHX 1I3MCHeI-IHM II CMell\eHHH, 06YCJIOB-
JICHHLIX iPYHl\D;IIOHaJIbHOH 3BOJIIOD;IICH BceH a.o;peHaJIOBOii CHCTCMLJ.
B OCHO Be 3'1'011 3BOJIIOn;HII JIemllT TaROC nOBLIIliCHIiC ypOBHH pa3BH-
TlIH HCpBHOH cHcTeMbI, IWTopoe 06CCneqHBaCT rocno.o;cTByIOll\ee BJIHH-
HHC Ha Roop.o;HHaD;HIo Bcex iPYHRD;Hii BbICllIlIX opr3HH3MoB. Y npep;-
CTaBHTf'JIeH pa3HLIx RJIaCCOB n03DOHOQHbIX - OT HH3ll1HX R BbICIlIHM
-24
06pa aeT Ha ce6H BHHMaHlIe rpa]:(HeHT B pacnpe]:(eJIeHHH xpoMaqJ-
4>UHHOii (cynpapeHaJIOBOii) TRaHH B TYJIOBH e mHBOTHbIX. fpa]:(IIeHT-
BbIpamaeTCH B CJIe]:(YIO eM. Y HU3KOOpraHH30BaHHbIX 4>OpM (y
RpyrJIOpOTblX H ceJIaxHii) xpoMa4>4>HHHaH TKaHb cOCpep;OTOqeHa B
RpynHblx MarHCTpaJIbHblX COCYJ];HCTbIX CTBOJIaX TeJIa, a TaRme B
rJIaBHOM n TnbCllpYIO eM yqaCTRe apTepHaJIbHOii CHCTelllbI - cepP;Qe,.
BeH03HOM CHHYCC (KOTOpbIH nOCTeneHHO B npoQecce 3BOJIIOQHOHHO-
ro pa3BHTHH DXOJ];HT D COCTaB npeJ];ccpJ];HH), B apTepHaJIbHOM KOHY-
ce H, no HeKOTopbIM aDTopaM (Giacomini, 1902; Gaskell, 1912),
B KpynHblx apTepHaJIbHbIX OTBeTBJICHHHX, npHHap;JIema HX CHCTeMC
]:(op3aJIbHOH aopTbI. .[(aJIee (y P;BOHKop;bIllIa HX, ROCTHoxpH eBbIx u
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ll\eCTBeHHO coxpaHHeTCH TOJibKO B KpynHbIX BeH03HbiX CTBOJIaX 11'-
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BeTB.'H:'HIlHX. B I1poQecce p;aJILHeHllIero 3BOJIIOQHOHHoro pa3BHTHH H
3Ta JIOKaJIII3al\IIH nOCTeneHHO IIICHHeTCH II cymllBaeTCH, II xpoMa4>-
4>IIIlHaH TIHlJII, !WIIQeHTpIlpYCTCH I1a rpaHHIJ;e cpegHeH H XBOCTOBOll
qacTcii TYJIOBIIllJ,a, B6JIH311 3aJ];HeH CTCHKH TeJIa, no 60RaM OT aopTbi
II I10JIoii BC'Hbl (y alll4>lIolln, PCIITlIJIIIH), Y BLIllieCTOHllJ,HX 4>OPM--,-
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nHTY,IJHI\HII.
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IfJIIIIIIOH TlmHII DLlpalliaeTCH HC TOJlbRO B TOM, qTO OHa nORHJ];aeT
apTepllaJILIIOe PYCJIO II, DOOOllJ,C, Marn:CTpaJIbHbIH RpOBOTOR H nepe-
J];BHraCTCH 1I0CTCneHHO OT OOJIce Rpymlblx COCYP;OB K OOJIee lIIeJIRHM,
HO II B TOM, 'ITO olla lureCT TeHJ];CIIQIIIO R nepeMcrqeHHIO BHYTPIl
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CBeTOIII, J];O Hapyailloro ap;BCHTIIQIIaJIbIlOro II P;O OKOJIOCOCYP;HCTOii
PblXJIoii COC' J];HHIITCJIhIlOii TlHlHlI.
STOT DTOpOl1 nYTb Mop4>OJIOrllqCCROii ncpecTpoiiRll xpoMa4>4>HH-
HhlX opraHoB CDH3aII C I1porpccCllpYIOIl\Cll 3BOJIIOQllen ClllllIIaTllqe-
CKon lIepRlIOll TJ\alIlI, HaXOJ];HllJ,eiicH B 3aqaT04HOM COCTOHHHH Y
RpyrJIOpOTblX II IIpllo6peTalOllJ,PR :ma4eHIIC BhICOKocneQllaJI1l311pOBaH-
HOR CIICTCMbI Y BbIClIlIIX 1I03BOHO'lIIhIX. B CDH311 C pa3BIITlleM CHM-
naTllqcclwii HCpBHOii CIICTe lbl COCY]:(llCTo-rYMopaJIbHble MexaHII3MbI
perYJIHI lIn II KOOp;J,I1HaI\I111 <IJlI3110JIOrII4eCRIIX 4>YHKQIIH YCTynalOT
MCCTO HPpmn,DI IJJIlIHIIIIHIII II BJIeI{YT 3a COOOR 04eHb cyr:u;eCTBeH-
Hble 1I3MeIleHnH B Mop4>OJIOrn4CCIWR opraHlI3aQHII BceH CHCTeMbi.
B .HODbIX YCJIODnHX OJIII30CTb aJ];peHeprH4eCRIIX mCJIe3IICTbiX 3JIeMeH-
TOB K IIlarllCTpaJIbHOMY l<pOBeHOCHOIlIY PyCJIY TepHeT CBoe nepBO-
CTenCIIHoe, ilU13HeHHO DamHoe 3Ha'IeHHe, H Ollll OT Hero OTcTynalOT.
Ha CMeHY OOJIec p;peBlIeH 1fJ0PMC pcrYJIHQHH B xOJ];e 3BOJIIOQlIH npll-
XOJ];llT nell3Mcpnlll0 OOJIee CODeplllCHllaH HcpBHaH perYJIHQHH. Xapal{-
1'epIJ3YH <pTJ31I0JIOrII4PCI\YIO CTOPOHY CMPHbI pcrymlpYIOllJ,HX CHCTeM,
A. A. YXToMclmii (1931, 147 CTp.) nHCaJI: «qeM 60JIee cnYCKaeM-
CH MM 110 4>llJIOrCIIPTll'lCCKOR II OHTorenCT1l4CCKOii JIeCTHHQe, TeM
60JIee C6JIHmalOTcH IemJ];Y c060ii, nO-BIIJ];IIMOMY, II no BpeMeHH, H no
25
tfJYHKIWH HepBHbIe HMnYJIbCbl H rYMOpaJIbHO-Me]:(HaTOpHble EJIHHHlIfl.
Ha060pOT. qeM R 60JIee BbICORHM OpraHH3aIJ:HflM Mbl nOJ];HHMaeMCfl,
'TeM CaMOCTOHTeJIbHee 110 CBOeMY J];eMCTBHIO HepBHble HMnYJIbCbI H
MeJ];HaTOpbI; TeM 60JIblllYIO CaMOCTOJlTeJIbHOCTb npH06peTaeT CpO'IHaJl,
MrHOBeHHaH CHrHaJIH3aIWH ]:(JIH npHHHTHH OqepeJ];HbIX MepOnpIlHTHM
nepeJ]; JIHn;OM TeRy:rn;HX C06b1THH B cpeJ];e».
BTOpoij II3 YRa3aHHbiX IIYTeH MopiPOJIOrHqeCKOH nepeCTpOMRII
xpoMaiPiPlIHHblX OpraHOB CBH3aH He TOJIbRO C COCYJ];HCTbIM annapa-
TOM llIeHHo-rpY]:(Hoi'1 06JIaCTII, 110 IIpeJ];CTaBJIHeT C060H 3aROHOMep-
HblH npon;ecc, xapaKTepHbIH J];JIfl BceH xpoMaiP!fJHHHoM CHCTeMbI B
u;eJIOM. B TO me BpeMH OH BJIHHeT Ha 3BOJlIOU;HIO meMHo-rpYJ];HblX
XpOMacP!fJIIHHbIX opraHOB, flBJIHIO:rn;HXCfl COCTaBHOH qaCTbIO 06:rn;eM
xpoMaiPiPHHHoM CHCTeMbl, B TOM me HanpaBJIeHIIH, KaKoe ORa3bIBaeT
Ha Hee perpeCCHBHafl 3BOJlIOU;HH ma6epHoro anlIapaTa. KapJ];IIo-aop-
TaJIbHble, nplICepp;eqHbIe II ImpOTIIJ];HbIe Y3JIbi XpOlIIaiPiPUHHOH TI\aHII
B pe3yJIbTaTe peJ];YRn;IIII ma6epHbiX COCYJ];OB, l\al\ paBHbIM 06pa30M
B pe3YJIbTaTe 06:rn;HX npou;eCCOB nOCTerreHHoro 060c06JIeHHfl xpOMa<!J-
iPHHHOi'1 TRaHII II3 COCTaBa COCYJ];HCTbIX CTeHOR, 06YCJIOBJIeHlIOrO
!fJYHRU;HOHaJIbHbIMH nOTpe6HOCTJIMH opraHH3MOB, TepflIOT CBH3b C 3TH-
MH COCYJ];aMH, HCqe3aIOT HJIH OTOJ];BIIraIOTCH OT HHX II B rrpon;ecce
HCqe3HOBeHHH HJIH IIepeJ];BHmeHHH OCTaBJIHIOT «CJIe]:(bl» CBoero npe-
6blBaHHH B 3THX MeCTax. «fJIOMYCbI» - 3TO He 'I TO HHoe, RaK «CJIe-
J];bI» iPHJIOreHeTlIqeClwro IIpOrrlJIOro xpoMaiPiPHHllOii TRaHII.
n pou;eCCbl CMe:rn;eHIIfl xpOMacPcPHIIHOH TRaHII flBJIHIOTCH pe3YJIb-
TaTOM u;eJIeC006pa3Horo MOPcPoiPYHKU;IIOHaJIbHOrO IIpIIcnoc06JIeHHH R
HOBbiM B03HHRaIo:rn;HM B IIpou;ecce 3BOJlIOU;HH 3arrpocaM II J];JIH HaII-
60JIee 3iPcPeKTHBHoro pellleHHH HOBbiX iPYImU;HOHaJIbHbIX 3aJ];aq. i.hu
CMe:rn;eHHH BblpamaIOTCJI B u;eHTpaJIII3aU;HH xpoMacPiPHHHOH TRaHH B
OJ];HOM napHOM opraHe - HaJ];rrOqeqHHRe II BcerJ];a 6LIBalOT conpflme-
Hbi C J];PyrHM HBJIeHlIeM - nporpeCCHBHblM pa3BHTHeM CHMnaTlIqe-
CROH HepBHOM CHCTeMbi. 8TII J];Ba rrpou;ecca HJ];YT BMeCTe PYKa 06
PYKY, 6YJ];yqH B3aHMHO He06xoJ];I1MbI II BbIrOJ];HbI. OJ];IIH npou;ecc 6e3
J];PYroro He ocy:rn;eCTBHM. «CJIeJ];bI» me cBoero 6b1BllIerO MeCTonpe-
6bIBaHIIH xpoMaiPiPHHHafl TRaHb OCTaBJIHeT B BH]:(e MeJIRHX rHe3]:(
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ma6epHoro annapaTa H Bcell CIIHYCHo-npeJ];cepJ];HOll 06JIaCTH, C OJ];HOll
CTOpOHbl, H 060c06JIeHHH xpOMaCPcPHHlIOH TRaHII H3 CTeHOR Hpyn-
HbIX CTBOJIOB RpOBeHOCHOll cHcTeMbI,- C APyrOll. 06a 3TII npou;ecca
npHBoJ];HT, B ROHeqHOM HTore, K 0J];H03HaqHblM pe3YJIbTaTaM. B ]:(PY-
rHX qaCTHX TeJIa, rJ];e npeBpa:rn;eHHe xpoMaiPiPHHHblX opraHoB B PY-
J];HMeHTapHble flBJIHeTCH CJIeJ];cTBlIeM TOJIbKO npHCIIOC06HTeJIbHOll 3BQ-
.mOU;HII, B pe3YJIbTaTe ROTOpOll xpoMaiPcPHHHaH TRaHb «oTOJ];BHraeTCJlIt
OT MarHCTpaJIbHOrO RpoBoToKa II cocpep;OTOqHBaeTCH B caMOCTOHTeJIL-
HblX H30JIHpOBaHHblX opraHax (KaK, HalIpHMep, M03rOBOll CJIOH HaA-
1I0qe1JHHROB IITHU; H MJIeROrrHTaIO:rn;HX), Pyp;HMeHTapHble OCTpOBKH ee
26
BCTpeqalOTCH peme. B RaqeCTBe 1I0CJIe]:(HIIX CJIep;yeT, HaIIplIMep, pac-
CMaTpHBaTb KOlIqHROBYIO meJIe3Y, lIaparaHrJIIIH, OIIHCaHHble B CTeH-
RaX BeH03Horo pYCJIa, B 6apa6aHHOn lIepelIOHRe H p;pyrHe. BlIpo-
qeM, HaJIHqHe pyp;HMeHTapHblX lIaparaHrJIHeB B 6apa6aHHOn nepe-
1I0HKe TOme, BHP;HMO, CJIep;yeT CBH3aTb C pe]:(YKl1;Hen ma6epHblX co-
CY]:(OB. H3BecTHo, qTO COCyp;HcTble H p;pyrHe TRaHeBbie 3JIeMeHTbI
ma6epHoro alIlIapaTa IIpHHHMalOT yqaCTHe He TOJIbRO B 06pa30Ba-
HIIH apTepHaJIbHbIX p;yr rpYP;Hon 06JIaCTH II KapOTHp;HbIX COCY]:(OB
mell, HO BXOP;HT B COCTaB pHp;a p;pyrllx opraHHblX II TRaHeBbiX
CTPYRTYP - 1I0P;'bH3blqJIbIX H qeJIIOCTHbIX ROCTen, cnYXOBoro IIPOXo-
p;a HapymHoro yxa, eBCTaXHeBon Tpy6b1 II p;pyrllx, K HX qHCJIY OT-
HOCHTCH H 6apa6aHHaH lIepelIOHRa.
CHCTeMaTH3npOBaHHbIII TaKHM 06pa30M JIHTepaTypHbln MaTeplIaJI,
corrOCTaBJIeHHbIl1 II 1I0p;KpelIJIeHHbln B pHp;e pa3p;eJIOB co6CTBeHHblMH
IICCJIe]J;OBaHIIHMn, n03BOJIHJI IIpHllTII K H3JIOmeHHblM Bblme 0606m;e-
HHHM.
l\fbl TBepAO yoeif\]J;enbI B TOM, qTO OTCYTCTBHe HCTOpHqeCROrO
1I0]J;xo]J;a B nCCJIe]J;OBaHHH H3JIOmeHHOn IIp06JIeMbI HBHJIOCb IIpH 1 m-
Hon, Bbl3BaBllIeii JIOmHble IIpe]J;CTaBJIeHHH 0 iPYHRl1;HOHaJIbHOii pOJIH
lIaparaHrJIHeB II Heo60CHOBaHHble ROHl1;elIl1;lI11 06 HX IIpHHa]J;JIemHOCTH
R BOClIpllHHMalOm;eMY, a He HCIIOJIHHTeJIbHOMY 3BeHY HepBHon p;eJl-
TeJIbHOCTH. OrpOMHaH qYBCTBIITeJIbHaH HHHepBaI(HH XpOI\IaiPiPHHHOB
TRaHII Ha Bcex 3Tallax iPHJIOreHe3a IIpOmmaeT CBeT Ha 06HJIHe pe-
l1;eIITOpHblX npnoopOB B RapOTH]J;HOM lIaparaHrJIHH, C HCqepIIbIBalO-
m;en IIOJIHOTOII OIIHcaHHblX de Castro (1926) y MJIeROIIHTaIOIIJ;HX.
De Castro, na OCHOBaHlIl1 aiPiPepeHTHoii HHHepBal1;HII H OTCYT-
CTBIIH xpoMaiPiPlIHHoH peaRlJ;Hu B ItapOTH)]:HOM lIaparaHrJIHII, BbiCKa-
3aJI MbICJIb 0 XeMOpel1;elITOpHon iPYHRl1;HII 3TOro opraHa. Ha caMOM
me ]:(eJIe 60raTOe Clta6meHHe RapOTH]J;Horo lIaparaHrJIHJI aiPiPepeHT-
HbIMII IIpn60paMII, TaR me RaK 0611JIbHaH BaCKYJIHpII3aI(HH 3Toro
opraHa, HHJIHIOTCH neOT'LeMJIeMLIMII H caMbIMH xapaRTepHbIMII CTPYR-
TypaMH, ROTopble 06eCIIeqllBaIOT 3H)]:pORPIlHHYIO iPYHRI HIO xpoMaiP-
iPHHHL1X opraHOB Ha Bcex 3Tanax HCTOpHqeCROrO pa3BHTHH 1I03BO-
HOqHbIX. OTCYTCTBue me rHCTOXHMllqeCROH peaRl1;HH HBJIHeTCH OP;HIIM
H3 caMLIX rJIaBHbIX nORa3aTeJIeil pY)]:UMellTapHoro xapaRTepa 3Toro
opraHa. ECJIU Ha IIpOTHmeHnn 3BOJIIOl1;1I1i n03BOHOqHbIX OT HH31l111X
iPOpM K BblCIlIHM aiPiPepeHTJlaH JIHHepBal1;HH He IIpeTepneBaeT CY-
m;eCTBeHHbIX RaqeCTBeHHblX IIpe06pa30BaHJIH, a CRO]J;HTCH JIHmb R 1\0-
JIllqeCTBeIlHOMY 113MeneHllIO pel1;enTOpHbIX npn60pOB, cHa6maIOm;HX
cynpapeHaJIOBYIO (xpoMaiPiPHHUYIO) TRaHb, TO BaCRYJIJlpIl3al1;HH ee
HClIbITLlBaeT ROpeHIILIe CTPYRTypHble IIepeCTpOHRH. llOCJIe]J;HHe 06-
YCJIOBJIeHbI 1I0HBJIeHHeM n 1I0CTelIeHHblM yCJIOmHeHHeM B 3BOJIIOl1;II11
3iP epeHTHoro 3BeHa HepBHldX peaKl1;I1H II, CJIe]:(OBaTeJIbHO, CTaHOB-
JIeHIIH nOJIHOl1;eHHOrO peiPJIeRTOpHoro aRTa. Ha CMeHY IIpllMHTIIBHOMY
YCTPOHCTBY, npH KOTOpOM xpoMaiPiPlIHHble RJIeTKH JIemaT B6JIH3H
IIpOCBeTa KpynHblX cocy]:(on TYJIOBHm;a, OMbIBalOTCH HX RpOBblO II
Bbl)]:eJIHIOT ropMOHbl B MarHCTpaJIbHbIll I\pOBOTOR, npHXO]:(HT CJIOmHO
YCTpoeHHble opraHLI, B ROTOpbIX CHCTeMa RpOBo06pam;eHHH YCJIOif\-
27
HlIeTCH napaJIJIeJIbHO YCJIOmHeHHIO HHHepBaIJ;HOHHbIX MeXaHH3MOB,
'CnOC06HbIX 06eClIeTJ:HTb IIX pe!fJJIeRTOpHYIO aRTHBHOCTb. Y BblCIlIHX
n03BOHoqHbiX TaRHM opraH OM HBJIHeTCH MeP;YJIJlpHaH qaCTb Hap;no-
--qeqHHlm. II pHcnoco6uTeJIbHaJl 3BOJIIOIJ;IIH HBJIHeTCH TeM MexaHII:3MOM,
B pe3YJIbTaTe ROToporo CHMnaTHTJ:eCRaH nepBllaH CHCTeMa c ee 6bI-
CTpOH II CTporo JIORaJIH30BaHHOH CHrHaJIH3aIJ;HeH (YXTOMCKHH, 1951)
npHlliJIa Ha CMeHY rYMopaJIbHO-::JHP;ORpHHHOH ]:(eHTeJIbHOCTH XPO-
Ma!fJ!fJHHHoij TKaHII. TOJIbKO 1I0CJIep;OBaTeJIbHOe H p;eTaJIbHOe H3yqe-
HlIe npOHBJIeHHH 3TOH npHcnoc06HTeJIbHOH 3BOJlIOIJ;HH MeTop;OM cpan-
HHTeJIbHOH MOp OJIOrHII II 3M6pllOJIOrHII nOp;BeJIII Hac R npaBHJIbHOH
'OIJ;eHRe xpoMa!fJ!fJHHHoH CHCTeMbi BblCIlIHX 1I03BOHOTJ:HbIX, B TOM qnc-
JIe MJIeROnHTaIOIIJ;IIX iHHBOTHbIX n TJ:eJIOBeRa.
B ,[IaHHOH pR60Te Mbl CTpeMllJIHCb ynoTpe6JIHTb MeTO,[lbl, ap;eK-
BaTHbIe B Ram,[lOM OTp;eJIbHOM CJIyqae nOCTaBJIeHHbIM 3a,[lRqaM. llpn
rHCTOXHMHTJ:eCROM onpe,[leJIeHIIH ropMOHOB npllMenHJIHCb peaKIJ;llII
XpOMllponaHlHI II 1l0,[lllpOnRHIHI no Hillarp, H6kfelt (1953, 195;:)}.
peaRIJ;IIH \V ood (1963) II BbICOKOqyncTBIITeJIbHbIH !fJJIyopeCIJ;eHTHbli'l:
MeTO,[l, OCHOBaHHblH Ha ROH,[IeHCaIJ;llH olloreHHblx aMHHOB C !fJ0pMaJIb-
).l;erHp;HblM ra30M (Falck, Owman, 1965). ]J;JIH npOBe,[leHHH 3TOro
MeTop;a llCnOJIb30BaJIRCb yCTRHoBRa ,[IJIH 6b1CTpOH JIIIO!fJIIJIbHOH CYIliKII
II 3RJIHBRH B rrapa!fJIIH, CROHCTpyHpoBaHHaH P;JIH 3TllX IJ;eJIell
A. 10. BY,[IRHIJ;eBbIM (1967, 1968). ]J;JIH H3yqeHHH ,[IH!fJ!fJepeHIJ;HpOBRH
RJIeTOR cHMnaTO-a,[lpeHaJIOBOH CHCTeMbI H BbIHcneHHH HCTO'IHHRa pa3-
BHTllH naparaHrJIHOHapHbIX 3JIeMeHTOn npHMeHenLI IJ;HTOJIOrHqeCKHe
MeTO,[lbI OKpaCRn no YHHa-Bpallle, «DeJIbreHY H MeTO]:( aBTopap;norpa-
!fJnH. fHcTO!fJH3HOJIOrHTJ:eCROe H 3ROJIOrIITJ:eCROe H3yqeHHH !fJYHRIJ;IIH
iKeJIe3nCTblX naparRHrJIHOHapHbIX RJIeTOK npOBe.n;eHO C 1I0MOlIJ;LIO CBe-
TOBOro H 3JIeRTpoHHOro MIIKpOCRonOB.
00061IJ;eHHe !fJaRTHqeCRoro MaTepHaJIa no a]:(peHaJIOBOii cHCTeMe
n03BOHOqHbIX c n03HIJ;HH CpRBHHTeJIbHOH Mop!fJOJIOrHHH n03BOJIHJIO cp;e-
JIaTb BbIBO,[lbl, OTpamalOlIJ;He TOTJ:KY 3peHllJl aBTopa, H 3THM OOYCJIOBH-
JIO onpe,[leJIeHHYIO n;eJIeHanpaBJIennOCTb BceH pa60TbI, OrpaHHqllB
Rpyr 06cymp;aeMblx BonpOCOB. OCTaJIHCb HeOCBelIJ;eHHblMH B3aUMOOT-
HOllleHHH M03roBOro H RopROBoro CJIOeB Hap;nOTJ:eTJ:HHROB (HJIH cyn-
pa- H IlHTeppeHaJIOBOH THaHeH HH31I1HX n03BOnOTJ:nLlx), xapaKTepH3Y-
IOIIJ;He HeROTOpbIe CTOpOHbI !fJYHKIJ;HOHaJIbHOH 1Il0p!fJOJIOrHH a,[lpeHa-
JIOBOH CHCTeMbi. ::.ha np06JIeMa HMeeT caMOCTOHTeJIbHOe 3HaqeHUe
(CM. 0030p Gorbman, Bern, 1962, H pa60Tbi Chester Jones, 1957,
1963) II HaMH 3aTpOHYTa TOJIbRO B IIpep;eJIax nOCTaBJIeHHOH 3ap;aqn.
qACTb ITEPBAH
CPABHMTEnbHAH MOp OnOrMH
CMMnA TO-A)J:PEHAnOBOII CMCTEMbI
I' .ilU6a r
I pyr JIOPOTbIE
B HaCTOHl ee BpeMH IllbI pacrrOJIaraeM p;OCTaTOqHO nOJIllblMII cne-
p;eHHHIIIII 06 aHaTOMlIqeCROM pacnOJIOmeHHII cynpapOHaJIOBOH (xpo-
MaiPiPHHHoH) TRaHH B TeJIe RpyrJIOpOTbIX. IIopBble orrllcaHIIH :3TOIi
CHCTelllbl, COOTBeTCTBYIO He cOBpeMeHHbIM npe]:(cTaBJIOHHHM 0 HeH,
npuHap;:Ien.aT ]:(ByM anTOpaM - Giacomini (190 a) II Gaskell
(1912). IIocJIe HX HCCJIep;OnaHHH lie npH6aBHJIOCb RaRHX-JIH60 1II0p-
iPoJIorllqeclUlx cBep;eHHH, ROTopble 6bI cy ecTBeHHO 1f3111eHHJIH llJIH
BHeCJIII HOtITO npHH HnllaJIbHO HOBoe B Hallill 3HaHHH n aToll 06-
JIaCTH. Pa60TLI Giacomini, 1I0CBHTHBllIero He O.o;HO .o;eCHTHJIeTIIO 1I3-
yqeHlIIO CIICTelllbl (<Ha,rJ;nOqeqHHROBIt pbI6 II RpyrJIOpOTbIX, TOJIbRO nop;-
RpenJIHlOTCH HOnLIlInI iPaRTaMH II pa3BHBalOTCH H8 HOBOM IIleTOP;H-
qOCRO 1 ypOBHe - HaR n OTHome1I1I1I rllCTOJIOrHqeCROH CTPYRTypbI cy-
IIpapeHa:IOBOU TRaHII, TaR II B OTHomeHHH B3aHlIIOOTHomeHHH ee C
oRpymalO HMH opraHaMH H TRaHeBbIMH aJIeMeHTaMH .o;pyrHx CIICTeM.
Giacomini ( 1902a) , lICIIOJIL3YH clIe IIiPHqHYIO xpoMaiPiPnHHYIO
peaK HIO, p;eTaJIbHO OnHCaJI y B3pOCJIbiX oc06eH IIlOpCI\HX MHHor,
a TaI\me B lIepHOp; HX JIlIqUHHOrO pa3BHTHH (y nocRopoeR), CHCTe-
MY RJIeTOqHbIX aJIellleHTOB meJITo-6yporo BeTa, IlIHpOlW pacnpocT-
paHeHHYIO y RpyrJIOpOTbIX, no IIpeHMy eCTBY Ha p;Op3aJIbHOH CTeHRe
TYJIOBIIII.\8. IIJIaCTbl H oT]:(eJIbHLle rne3.o;a aToH TRaIlH BCTpeqaIOTCR,
HaqHHaH OT BToporo ma6epHoro cerMOHTa p;o aHaJIbHOro oTBepcTHH.
HaH60JIee MaCCHBHble CROnJIeHHJI cyrrpapOHaJIOBOH TRaHH cOCpO.o;OTO-
"IeHbl y cnnHHou aopTbI H Ha ee RpynHblx apTepHaJIbHbIX BeTBHX.
IIJIOTHble CJIOll I\JIeTOR C xpoMaiPiPHHHbIMH CBOHCTBaMH JIemaT no
60RaM aopTbI - Memp;y ee CTeHI\OH H rrpOCBeTOM COOTBeTcTByIO eii
Rapp;HHaJIbHOH BeHbl, conpOBomp;alOT OTXO H He OT aopTbI lIapHe-
TaJIbHble apTeplIH H npOp;OJImaIOTCJI BP;OJIb ]:(op3aJIbHbIX H BeHTpaJIb-
HblX HX OTBeTBJIeHHH. Ha6JIIOp;aeTCJI HepaBHOMepHaJl IlIHpHHa CJIoen
xpolllaiPiPHHHblX RJIeTOR - OHH YTOJI aIOTCH B MeCTax OTxomp;eHIIH
napneTaJIbHbIX apTepHH H YTOHqalOTCH Me)R)];y HHMH, 06pa3YH Me-
29
TaMepHono.o;06HbIe CTPYRTypbl. B COnpOBOm.o;eHHH BeHTpaJIbHblX apTe-
PHH H no.o; 3H.o;OTeJIHeM H.o;ynWX pH.o;OM naplIeTaJIbHbIX BeH nJIa-
CTHHRH xpoMalJ!lJ!HHHbIX RJIeTOR npOHHRalOT rJIy60Ro B 6plOmHYID
1I0JIOCTb. B XBOCTOBOH 06JIaCTH 06a 60ROBbiX aOpTaJIbHblX CJIOH CY-
IIpa peHaJIOBOH TRaHH CJIHBalOTCH .o;pyr C .o;pyroM H IIpOHH3blBalOT
COCy.o;HCTYIO cTeHRy, BbICTYIIaH, C O.o;HOH CTOpOHbl, B npOCBeT cocy-
.o;a, a C .o;pyroH,- yrJIy6JIHIOTCJI B coe]:(HHllTeJIbHYIO TRaHb.
B BepXHeH qaCTH TeJIa ROJIHqeCTBO ee He YMeHbmaeTCH. B 06-
JIaCTH cep.o;D;a, TaM, r.o;e 06e 3a.o;HHe Rap]:(IIHaJIbHbIe BeHb! OTRpbl-
BaIOTCH B BeH03HbIH CHHYC, cynpapeHaJIOBaH TRaHb, conpOBom]:(al3-
maH aopTY, BCTynaeT B npocBeT BeH03Horo cHHyca. B 3TOM MeCTe
OHa co Bcex CTOpOH 06BOJIaRHBaeT OTXO.o;JlID;YIO OT aopTbi qpeBHyro
apTepHIO, IIpOHHRaeT no Xo.o;y nOqeqHblX apTeplIH B napeHXHMaT03-
HYIO TRaHb 1I0qeR, a OTp;eJIbHbIe ee OCTpOBKH yrJIy6JIHIOTCH B CTeH-
Ry aopTbi. llOCJIe.o;Hee Ha6JIIO.o;eHHe Giacomini, onHcaBllIero 3JIeMeH-
Tbl xpoMalJ!IJ!HHHoH TRal1H B nepHRap.o;HaJIbHOH 06JIaCTH RpyrJIOpo-
TbIX, 3aCJIymHBaeT 60JIblllOrO BHHMaHHH, oc06eHHo .o;Ba 06CTOJlTeJIb-
cTBa, ROTOpbJe ROHcTaTHpyeT aBTop. Bo-nepBblx, caMO npHCYTCTBHe
cynpapeHaJIOBOH TRaHH B yqaCTRaX, npHJIelHalD;HX R cep.o;D;y, BO-
BTOpbIX, ee .o;OJIbqaTOe HJIH Y3JIOBaTOe CTpoeHHe II norpymeHHe He-
ROTOpblX Y3eJIROB B CTeHRY aopTbi. fIMeHHO B 3THX MeCTax y BblC-
mHX n03BOHOqHbIX mHBOTnblX pacnOJIaraIOTCH IIpHcep.o;eqHbIe napa-
raHrJIHH (roMOJIOrH CTPYRTYP RpyrJIOpOTbIX), 0 IJ!YHl\D;HH ROTOpbIX
B TeqeHHe nOCJIe.o;HHX .o;eCHTHJIeTHH Be.o;eTCH .o;HCRYCCHH npHHD;HnHaJIb-
Horo xapaRTepa - ocnapHBaeTCH npHHa.o;JIemHOCTb HX R CIICTeMe XpO-
MalJ!lJ!HHHbIX opraHOB. Mem.o;y TeM BbI]:(eJIeHHe npHaOpTaJIbHblX rrpH-
cep.o;eqHbIX naparaHrJIHeB H3 rpymIbI .o;pyrlIx aHaJIOrHqHbIX xpOMaq.-
cI>HHHblX opraHOB n03BOHOqHblX IIpOTIIBope<IHT, RaR MbI nOCTapaeMCH
nORa3aTb B .o;aJIbHeHllIeM 1I3JIOI:IWHHH, BceMY Xo.o;y IJ!YHRD;HOHaJIbHOK
3BOJIIOD;HII 3TOH CHCTeMbI.
llpeRapOTH.o;HYIO 06JIaCTb pacnpOCTpaHeHHH xpoMalJ!lJ!lIHHOH TRa-
HH Giacomini ( 1902a) Ha3bIBaeT 10rYJIHpHoH. llo ero .o;aHHblM, Ha-
qHHaH OT MeCTa Bna.o;eH'HH HpeMHblX BeH B BeH03HbIH ClIHYC, cyn-
papeHaJIOBaH TRaHb, npo.o;BHraHCb R ma6epHoH 06JIaCTH, IIOCTeneHHO
OTXO.o;HT OT aopTbi H H.o;eT B.o;OJIb HpeMHbIX BeH, npHJIeraJl R HH I
C TOH CTOpOHbI, r.o;e BeHbI conpHRacalOTCH C n03BOH01JHblMll apTe- .
pHHMH H .o;pyrHMH apTepHaJIbHbIMH BeTBHMH. 113 OIIHcaHHH ma6pn-
HOH 06JIaCTH peqHoii MHHorH, H3yqeHHOH Gaskell (1912), CJIe.o;yeT,
qTO xpoMalJ!IJ!HHHaH TI\aHb pacnpocTpaHHeTcH II B CTeHRax cerMeH-
TapHbIx BeH ma6epHoH o6JIacTlI, lI.o;ylD;HX BORpyr HOTOXOp.o;bI.
8TH CBe.o;eHHH 0 CTPYRTypHOH opraHH3aD;HH CllCTeMbl xpoMalJ!lJ!llH-
HblX opraHoB y MllHorOBbIX 6blJIII nO]:(TBepm.o;eHbI B nOCJIe.o;HHe ro.o;Ll
IJ!JIIOOpeCD;eHTHo-rHCTOXHMIlQeCRHM MeTO]:(OM (Falck, 1962; Falck,
Owman, 1965), BbICOROqYBCTBHTeJIbHbIM .o;JIH MHHoaMlmOB. BblJIO 110-
Ra3aHO HaJIHqHe cneD;HlJ!llqeCRH IJ!JIIOOpecD;HpYIOID;HX xpoMaIJ!IJ!HHHb!X
BJIeMeHTOB B CTeHRax 3a.o;HHX Rap.o;llHaJIbHbIX BeH, 6pIOmHoH aopTbl
II oc06eHHo B yqaCTRaX, npHJIeraIOlD;lIx R cep.o;D;y (JIeoHTbeBa,
1966a, 1968).
30
Pile. 1. CxeMa. Cpea 'lepea
cpep;HIOIO 'laCTb TYJlOBH a MOp-
CROH MJIHorII (Petromyson ma-
rinus)
CYIIPapeHaJIOl<aH TRaUb - nYHRTHPI
rpYIIIIhI HHTeppeHaJIOBoi!; TRaHH-
'lepHbIe TO'nm; nc - HOTOXOpga; cv-
RapAHHaJIbHhIe BeHhI; ao - aopTa;
k - nO'lHH; sic - cmmHot1: M03r;
pad - AOpSaJIbHhIe napHeTaJIbHhIe
apTeplm; pvd - TO me, BeHhI; pav -
BeHTpaJIbHhIe napHeTaJIbHhIe apre-
111m; pvv - BeHrpaJIbHhIe napHeTaJIb-
1lliIe BeHbI (Giacomin1, . no
Chester Jones, 1957)
TaI>IIM 06pa30M, qpe3Bbl'IaHHO XapaRTepHaH OC06eHHOCTb XPO-
IImiPqnlHHoH TRaHII RpyrJIOpOTblX (II pbl6) - 3TO ee MeCTOIIOJIOme-
HlIe B CIICTeMe MarlICTpaJIbHOI'O COCyp;IICTOrO RpOBOTORa. PerYJIHpHO
II B 60JIblIllIX ROJIlIqeCTBaX OHa cocpep;OTOqeHa B CTeHRax 60JIblIlIIX
COCYP;OB TYJIOBIIIqa, rJIaBHblM 06pa30M B CIICTeMe CIIIIHHOH aopThI
(pIIC. 1) II B HalI60JIee RpYIIHblX apTeplIaJIbHblX CTBOJIaX roJ:!OB-
HOro II XBOCTOBoro OTp;eJIOB. ITJIaCTbI xpoMaiPiPlIHHoH TRaHII, COCTO-
HIqlIe II3 KJIeTOR pa3Ho06pa3HoH iPOpMbI II BeJIlIqlIHbI, BblTHnyTbIe
B P;JIIIHY IIJIII ROJIbn;eo6pa3Ho OXBaTblBaIOIqlIe IIpOCBeT cocyp;a, qaCTO
OTrpaHuqeHbl OT Hero JIlIlIlb TOHRIIM CJIOeM 3Hp;OTeJIlIaJIbHOH BbICTIIJI-
KII (Giacomini, 1902a). Cy63Hp;OTeJIlIaJThHOe pacrrOJIOmeHlIe cYIIpa-
peHaJIbHblX TeJIen; II03me OIIlIcaHO II faCReJIOM (Gaskell, 1912), RO-
TOpblH 06HapymlIJI IIX B BIIp;e CIIJIOIIIHOI'O TOHRoro CJIOJl B MarIICT-
paJIbHblX apTeplIHX Ha IIpOTHme.B1I1I Bcero TYJIOBIIIqa pe'lHOH MIIHorlI.
Ha TO, qTO xpoMaiPiPlIHHble TeJIa OqeHb TeCHO CIIaHHbI C 3HJ];OTe-
JIlIeM, 06paTIIJI BHIIMaHlIe eIqe ITeTTII B Op;HOH II3 CBOIIX paHHIIX
pa60T (Pettit, 1896). OTlIpeIIaplIpoBaB xpoMaiPiPlIHnyIO TRaHb B CTeH-
Kax ma6epHbix COCYP;OB, OH OTMeTIIJI HeB03MomHOCTb ee OTp;eJIeHIIH
31
6e3 nonpemp;eHlIH Bcell COCY]:(IICTOH CTeHl\H. B Tex CJIY IallX, l\Orp,a \
KJIeT01JIILle rpynnbl BCTpe1JaIOTCll B ap;BeHTII lIaJIbHOM CJIOe aopTbl
llJIIi Mem]:(y aopTOH II Rap]:(IIHaJIbHbIMIi BeHaMII, OHII MorYT IIMeTb
O'IpOCT1JaTYIO 1jI0PMY (Giacomini, 1902a) II 6bITb CXOP;HbIMII C co-
oTBeTcTBylO IIMII :meMeHTaMIi CeJIaxHii: II aMIjIII6I1H. PacnoJIarallCb
Bp;aJIIi OT npocBeTa cocyp;a, OHII, O]:(HaRO, 0I\a3blBalOTCll npOll1l3aH-
HblMIi lIepliBacRYJIHpHblMII JIII llfJaTII1JeCKIiMII IIpocTpaHcTBa?>1II II OT-
p;eJIlllOTCll OT HIIX TOJIbKO TOHKIIM CJIOeM 3H]:(OTeJIII1l. RaK npaBlIJIO,
nepliBacHYJIllpHO paCIIOJImlWHHble rpynnbl xpoMaljllfJlIHHblx HJIeTO'\
BCTpe IaIOTCH 3Ha'-l.I1Te.'1bHO peme II lIIeJIb1Je no pa3MepaM.
113 npIlBe]:(eHHLIX HaOJIIOp,eHllii: CJIep;yeT, ITO ynpapeHaJIOBall
]Hallb KpyrJIOpOTLIX IIMeeT TeH]:(eH 1I1O rpynnlIpoBRTbCll B Tex Y1Ja-
CTI{aX cocyp,I1cToii CIICTeMbl, r]:(e reMop;IIHaMII1JeCHIiA YCJIOBlIll HpOBO-
TOHa cllorooHbl 06eClle'-l.IITb HallOOJIee 6LICTPYID p;OCTaBI{y HpoBn II
ee COCTaHIlLlX 1J3CTeii IW nceM ncplilbepll'leCHIiM opraHaM II TKallll 1
I mliBOTHoro.
C 3T0I1 T01JRII 3peHII1l npep,CTaBJIlllOT I1HTepeC B3allMOOTHomeHHH
xpoMalfJlfJHHHblX 3JIeMeHTOB C rJIanllLIM HarHATaTeJIbHblM IIpH60pOM
COCY]:(I1cToii CIICTeMbi - cepp, eM. 0 Tml, ITO XPO?>IaIfJIfJIIHHbIe HJIeTlm:
npOH1I3LIII<llOT BCIO TOJI y CTeHHlI neH03Horo Clmyca cepp, a HpyrJIO-
POTbIX, 1I3BeCTHO II;} pa60T Giacomini (1902a) II Gaskell (1912).
IIpncYTcTnne ;nnx ImeTOn n IIpe,.'J;cep;J:lIllX II }I;eJIy]:(otmax cepp; a ?>1lI-
1I0r 11 MimCHlI II CXO.L\CTRO IlX 1I0 IICIWTOpblM IIpn3HaIial\1 C xpo-
MalfJ4JlIHHLulII KJleTna 1II IHI UIOtIC1JHIIIWB MJleKOnIITalO IIX nOKa3aHO
B pll]:(e (OBpel\1eHIILIX IICCJlep,OBaHllii (A ugustinsson et aI., 1956:
Ostlund ('1, aI., 196U; Johnels, Palmgren, 1960; Bloom et aI.,
1961; HoHmeislel' el aI.. 1961; Jen ell, 1961: rOllhlpUU II JIeonTb-
eBa, 190;), a, 0; JleonTbena, 1968: Caravita, Coscia, 19(6). IIc-
nOJlb3YlI aprellTol}JIIJlbnhle cnoii:cTBa xpo?>ral}J4JlIHHLIX 3JIeMeHTOB, AIl-
gustinsson C coaBTOpa?>lII (195G) oUHapynm:m y lIlHor, a Johnels,
Palmgren (19GO) y MUKCIIH cBoeoupa3nylO CTpyHTYPY cynpapeHa-
JIOBblX HJleTOH II cnc IIlIJlI'1eClme rpanym,1 B IIX IITOnJIa3Me, ORpa-
mennbIe a30THO-KIICJlhlM cepeupoM. Ostlund c coaBTopaMII (t 9GO),
Bloom C coaBTopaMII (19G1), Hoffmeister C coaBTOpaMII (19Gl),
Caravita Ii Coscia (1966) OnlICflJIII 3J1eKTpOllHoMIIHpOCKOnIl1JecHYIO
KapTlIH)' OCMII04J11J1bHbIX rpaHy.lIH lIii B TeJIaX 3TIIX ImeTOK. OHa3aB-
llIyroCH MopljlOJIOrWICCHII CXOp,1I0ii: C rpaHYJIaMlI M03roBoro CJIOH nap;-
nO'Ie t IHIIJ-;')B MJleIWnUTalOl lIX. DUOXIIl\IlI'1eCHOe nCCJIe]:(OBalllle rpa-
nYJI, lI;'OJlllpOBaUHbIX 113 cynpapmw.TlOlJoir TliIlnll (Bloom el al..
1 fJ(1), YCTaHOBIIJIO B IInx HaTeXOJIflMlIlIOBoe cop;epmllMoe, .a BBep;e-
Hlle MIllloraM pe3epmIHa Bbl3BaJIO TaHoe me lICTo eHne rpauy,'1
(Bloom et aI., 1961; Caravita, Coscia, 19(6), Kanoe Ha6JIIOp;aeTC;I
B M03rOROM CJIoe Hap;nO'le IHIIHOB MJIeKOnllTalO IIX (Holzhauer,
Vogt, 1956), 1JTO ]:(aJIO OCHOBanliC npllaHaTb nOJIHYIO anaJIOrlI1O 3TIIX
CTPYHTYp. CJIep;yeT npn 3TOM BCnOMHlITb, ITO e e Gaskell (1912)
nOKa3aJI npllcYTcTBlIC a]:(peUaJIlIna B cynpapenaJIOBblX TeJIaX KpyrJIO-
pOTbIX. Y t lllTbIBall, '11'0 rpynubl HJIeTOK cynpapenaJloBoH TKallll qaCTO
BCTpeqUIOTCll B Tex me MeCTax, rp;e naXOp;llTCll III1TeppeHaJIOBl,Ie
32
Tc.'nJ a, II He ]:(onOJIbCTBY1lCb O]:(HIiMII JIlillib MOpq,OJIOfllqeCInIMll npll-
:maImMIi l:I l:ITOXIIMII'IeCKOii peaK lIeH (c XpOl\laTOM KaJIII1l), Gaskell
BBOP;II:1 B KpoBeIlocHoe pyc.'IO KomeK 3KCTpal\Tbl, 1I0JIyqeHIIbIe II3
aOpTaJIbHblX CTeHOK IIJIIi 113 Kapp;IIHaJIbHbIX BeH MIiHor, II nOJIyqaJI
y HIIX nO]:('be!\[ KpOB1lHOrO p;aBJIeHII1l. TaKIIM 06pa30M, npIlCYTCTBII'3
ap;peHaJIIIHono]:(oGHLIX coe]:(IIHeHIIH cBoiicTBeHHO He TOJIbKO cepp; y
KpyrJIOpOTbIX, IIO BceH cynpapeHaJIOBOH TKaHII, paCCe1lHHoi'l B CTeH-
Rax RPYIIHbIX COCYP;OB II cepp; a II COCTaBJI1lIOlQeH e]:(IIHYIO CYIIpa-
peHaJIOnyIO CilCTeMY HlI3ll1nX n03BOHOqHbIX.
B wpq,OJIOrll'IeCKOH opraHlI3a 1I1i _ X1!.OMaq,!JI HHbIX _ :1JI MeHTOB
cepp; a KpyrJIOpOTblX Ha]:(o IIOp;qepKHYTL ]:(Bl' ocoGeHHocTII, KOTOpbIe
Tome xapaKTepll3YIOT IIX :1H]:(OKpIlHHYlO q,YHK lIIO.
llpemp;e Bcero, pOMaq,q,IIHHbIe KJIeTl\H, pacnOJIOmeHHble l"Iem
Mblme'lHbIMIi BOJIOImaMR, lIMeIOT MnOrOOTpOCT'IaTble BbipOCTbl II pa3-
HeTBJIeHII1l: BCJIe]:(CTBlie 'lero 'mCTO nplloGpeTaIOT npll'IY]:(JIIiBble oqep-
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:leHnii: pa3HOH BeJlIlqllHbl, npellMYlQeCTBeHHO BGJIlI311 IIOJIOCTII cep]:( a.
Y MIIKCIIH OTpOCTKII aHaCTOM03l1PYIOT p;pyr C p;pyroM (Bloom et
al., 1961; Johnels a. Palmgren, 1961) II oGpa3YIOT HeqTO Bpop;e
CIIH IITII1l (pIiC. 2). Y MIiHor, 110 HalliliM p;aHHbIM, xpoMaq,q,IIH1IbIO
:1JIeMeHTbl BbIT1lHYTOH JieHToBII]:(HOii 1jI0pMbi CTeJIIOTC1l B BIIp;e OP;HO-
CJIOHHblX T1lmeH KJIeTOK BP;OJIb Mblme'lHblX BOJIOKOH IIpep;Cepp;IIH II
B MeHbllleM KOJIlIqeCTBe - meJIyp;OqKOB. OHIi Henocpep;CTBeHHO rpa-
HlI'IaT JIIiGO C JIaKYHapHblMIi npOCTpaHCTBaMII, JIIIGO C IIOJIOCTbIO
cep;J; a (pIiC. 3) 1'1, CJIe.u:onaTeJIbHO, OMblBaIOTC1l npoTeKaIOlQeii
J>pOBbIO. EOJIblllllHCTBO K.'IeTOIt p;aIOT nO.'IOil\IfTeJIbHYIO peaK IIIO C co-
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JIIIH) .
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TblX 1lBJIJleTC1l, BII]:(IIMO, IjIYHK 1I0HaJIbHO nplicnocoGJIeHHblM IIp1l3Ha-
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TaKoro npep;IIOJIOmellll1l yGemp;aIOT Hac II :1JIeKTpOHHOMIIKpOCKOIIlIqe- .
CRlIe HaGJIIO]:(eHII1l Caravita II Coscia (1966) Hap; xpoMaljlljlIlHHblMII
KJIeTKaMII cepp; a pe3epIIIIHII3I1pOBaHHbIX MIIHor. O'IeHb TO"IHble 01111-
CaHII1l :1TIIX aBTopOB II IIpIlJIaraeMbie 3JIeKTpOHHorpaMMbi He Bbl3bl-
BaIOT COMHeHlIii B TOM, qTO B IIepIlOp; oCBoGomp;eHII1l ropMOHOB \
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TOpbIe B.ICTYIIaIOT B npOCBCT paCIlIIIpeHHblX MemKJIeTO"IHbIX IIpOCT-
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KpyrJIOpOTblX P;O CIIX nop Bbl3b1BaeT pa3Hoe TOJIKOBaHIIe IIpIlpOP;LI
:1TIIX :1JIeMeHTOB (Augustinsson et al.. 1956; Bloom et aI., 1961).
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gren, 1960)
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me cnoiicTUCJIII3 I\JIJIlIll3H, IWJIO'mOBllp;n3H llJIli ncpeTeHoo6pa3H3f!
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34
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2* 35
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nOJIOCTII BblCTJIaHbl ynJIOII eHHbIMH :1H]:(OTeJIlIaJIbHbTMII KJIeTRaMII C
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xpoMaiPiPHHHblM KJIeTKaM. XapaKTeplI3YH MoPiPOJIOrH IeCKHe IIOKa3a-
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iPHHHblX KJIeTOK. B IIOJIb3y 3Toro rOBopHT H p;pyroe Ha6JIIOp;eHIIe
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MeCTax COlIpnKOCHoneHHH ]:(BYX coce]:(Hnx xpoM3iPiPnHHbix KJIeTOK.
O'leBH]:(HO, TOJIbKO rpaHH bl Mem]:(y xpoMaiPiPHHHoii KJIeTKOH II :1HJJ;O-
TemlaJIbHOH IIJIlI Mem]:(y xpoMaiPiPHHHoH KJIeTKOH II Mblllle IHblM BO.'IOIi-
HOM CJIymaT MeCTaMH Bblp;eJIeHHfI ropMOHOB.
3p;eCb YMeCTHO ncnOMHHTb npemHHe 1Ia6JIIOp;eHnH B cneTOBOM
MIIKpOCKOIIe GrynfelU. (1903a), KOTOpblH C yp;nBHTeJIbHOH TO IHOCTbIO
OIIIICaJI HTOJIOrHIO xpoMaiPiPHHHblX 3JIeMeHTOB B COCYP;HCTbIX CTeH-
Kax KPyrJIOpOThIX H pbl6. Ha OCHOBaHHII OqepTaHHH KJIeTOK, xapaK-
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'l'ex MeCTax, rp;e KJIeTKII 06pa3YIOT Bcero OP;IIH CJIOH BORpyr cocyp;a
H, CJIe]:(OBaTeJIbHO, IIop;BeprHYTbl O]:(HHaKOBOMY BJIHflHHIO iPlIKcaTopa,
HeKOTOpbIe H3 HHX naKYOJIH3HpOBaHbI, p;pyrHe eJIHKOM 3a6nTbi xpo-
MaiPiPllliHblMH rpaHYJIaMH. B HaCTOfl:m;ee BpeMH :1Hp;OKpnHHblH xa-
paKTep IiJIeTOK, IIpHH HIIlIaJIbBO Op;IIHaKOBbIR BO Bcex COCYP;HCTbIX
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IIHqeCKOM ypOBHe.
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CTaBIITeJIen Bcex KJIacCOB II03BOHO lHbIX mHBOTHblX - OT KpyrJIOpo-
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36
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n CaMOH 'fOJIIQe MblllleqHOrO CJlOH npe]l;Cepf];IIH II meJIY]I;OqKOB B6J1H3H
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rJIaBHblX MarHCTpaJIbHblX CTBOJIOB TYJIOBlIlI(a II rOJIOBbl, npOH1I3b1BaeT I
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HOB (Ka TeXOJIaMIIHOB), HaXO]l;HCb B MblillQe cep]l;Qa - QeHTpaJIbHOM
HarHeTaTeJIbHOM aIIIIapaTe COCY]:(HCTOH CHCTeMbl - cnoc06HbI, Bbl6pa-
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mHBOTHoro.
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nOJlYQCHHbIe lIa cep):\eQHOH MbllllQe MHHor II CBII):\eTeJIbCTBYIOIQHe 0
npllCYTCTBIIJI RaTeXOJIaMHHOBbiX ropMOHOB B caMHX MbillIQax. OKa-
3aJIOCb, qTO caMlI MbIllIe IHbIe BOJIOKHa 06JIa]:(aIOT CJIa60H xpOMaw-
WHHIIOi'I peaKQHeH (Bloom et aI., 1961; rOBblpHH, 1967) II co]:(ep- \
maT 8.'ICKTpOHHOnJlOTHble rpaHYJIbl, aHaJIOrH IHbIe TaKOBblM B XPO- I
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1966) .
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8THX <l>aKToB, B QaCTHOCTlI 06CYlli):\BTb BOIIpOCbl 0 TOM, HBJIHeTCH
JIll HaJlH'lHe KaTeXOJIaMIiHOB B MI.IIlIQe cep]l;Qa !lIHHOr pe3YJIbTaTO 1
HaI.OllJlellllH B HHX ropMonOB lIJIIi CJIe]:(CTBHeM HX HOBo06pa30BaHlIH.
3TH BOnpOCI.I HMeIOT caMOCTOHTeJlbllOe 3HaqeHHe II ]l;HCKYTHpYIOTCII
B COOTBeTCTBYIOIqeii: JIHTepaType. OTllIeTIIM 3]1;eCb TOJIbKO, ITO IIPH-
CYTCTBJle RaTeXOJIal\IIIHOB B MbIllIeQUblX BOJlOKHaX cepP;Qa C WYHI\-
n;UmIa:JLHoii: TOqRII 3pelllIH Tome c,ac;lyeT paCCMaTpllBaTb KaK npH-
CIIOCOOUTCJlbllI.lH MexaHII3M, CIIoc06cTBYIOIqHH YCKopeHHOH ]l;OCTaBKC
XllMII IeClm aKTHBHblX BelQeCTB (KaTeXOJIaMHHOBbIX ropMOHOB) K Me-
CTY Ha3HaqeHIIH - K 8iPweKTopHblM opraHaM.
BIIOXIIMH<JeCKHMII onpe]l;eJIeHIIHMlI (Ostlund, 1954; Ostlund et
aI., 19GO; fOBblpHH, 1967) yCTaHOBJleHO He06b1'laii:Ho BI.ICOKOe 110
cpaBHeHHIO C ]l;pyrHMH n03BOHO<JHbIMH cO]l;epmaHlIe KaTeXOJIaMHHOB
B ep;uw KpyrJlopoTbIX. no Ostlund (1954) B ccp]:(Qe MHKCUH Ka-
TeXOJlaMJlHOB B 120 pa3 60Jlbllle, 'leM B cep]:(Qe CeJIaXHH H KOCTII-
CTblX pbI6, H B 54 pa3a 60Jlbllle, QCM B cep]l;n;e MJleKOnHTaIOIqHX.
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CTbl!lUl, llau]:(eHo B cep]:(n;e MllHOr (rOBblpHH, 1967): B IIpe]:(cep]:(JlHX
225 MKZ/Z ]l;JlH a]:(peHaJlHHa H 138 MJIi,Z/Z ]l;JlH HOpa]l;peHaJIHHa,
B mCJIy]:(OqKaX COOTBeTCTBeHHO 54 H 43 Mnz/z; y KapnOBblX, 110
aHHbIM Toro me aBTopa, B npe]:(cep]l;HHx 2 Mnz/ Z a]l;peHaJlHHa
H 1,1 M1iZ/Z Hopap,pellaJIIIHa, B meJlY]l;O<JKax COOTBeTCTBeHHO 1 H
3 Mnz/z. no ]:(aHHbIM E. M. CTa6poBcKoro (1967), 3aIIaCbI a]l;pe-
HaJIHHa H HOpa]l;peHaJlHHa Y MHHor cocpe]:(OTOqeHbl He TOJlbKO B
cep]:(n;e, HO II B KpynHblx cocY]l;ax TYJlOBIIIqa.
37
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XOJIaMIIHbI, HaM KameTCH, qTO nYTb K 06'bHCHCHHIO IIX 60JIbmHX
KOH:qCHTpa:qIIH B cepp;:qe MIIHOrII JIemHT Bce B TOH mc IIOTpe6HOCTJI
6b1CTpOH pCaJI1I3a:qIIII ropMoHOB II, CJIep;OBaTeJIbHO, He06xop;IIMOCTII B
IIX 60JIblllHX 3aIIacax.
B 'leM CJIep;yeT IICKaTb IIplI'lHHY CTOJIb TeCHbiX MOPWOJIOrnqc-
CKIIX B3alIMOOTHOlllCHHH CYIIpapeHaJIOBOH TKaJIlI KpyrJIopOTbiX C OC-
HOBHblMII HalI60JIec KPYIIHblMH cocyp;aMH TYJIOBIII:qa II cepp;:qeM, T. e.
C TeMH y'laCTKaMII COCYP;IICTOH CIICTCMbI, KOTOpbIe 06CCIIe'llIBalOT op-
raHH3MY MarHCTpaJIbHblH KpOBOTOK?
Y BblCIlIHX II03BOHO'IHbIX p;OCTaBKa H pacnpCp;eJIeHlIe no opra-
HaM iPII3IIOJIOrlI'leCKH aKTIIBHblX BCI:qCCTB, B TOM 'lHCJIe MOHoaMII-
HOB, OCYI:qCCTBJIHCTCH IIpII IIOCpCP;CTBC HCpBHOH CIICTeMbi. CJIOmHO
IIOCTpOCHHbIe H TO'lHO OTpcryJIHpOBaHHbIe HCpBHblC McxaHII3MbI y
aTHX mHBOTHblX 06CCIIeqHBalOT MrHOBCHHYIO pcaK:q1I1O aiPiPcKTopHblX
HCHpOHOB B OTBCT Ha aiPiPepcHTHbiC CHrHaJIbI, IIOCTYIIalOI:qHe C IIC-
' pHiPepHH.
Mbi 3HaCM, qTO. qCM BbilliC B anOJIIO:qHOBHOM OTHOIliCHHII opraHlI3a-
lIH mllBOTHOro, TCM COBCplllCHHCC CTPYKTYpHO-WYHK:qHOHaJIbHble
CBOHCTBa HCpBHOH CHCTeM{,I, TCM P;HwiPcpcn:qHpOBaHHCC cc peiPJICKTOP-
HaH ]:(CHTCJIbHOCTb.
A KaK B aTOM OTHomCHHII 06CTOlIT P;CJIO Y KpyrJIOpOTLIX - y iRII-
BOTHbIX, KOTOpbIe no npHMlITHBHOCTII CTpOCHHH BblP;CJICHLI B OCO-
6ylO rpynny 6eCqCJIIOCTHbiX (Agnatha)?
XOTH 0 MOPWOJIOrH'leCKOH opraHlI3a:qHU HepBHoH CHCTCMbI Rpyr-
JIOpOTblX HCT p;OCTaTOqHbIX CBCP;CHIIH, op;HaKO nMelOI:queCH p;aHHbIe
1I03BOJIHIOT paccMaTpHBaTb IIHTCPCCYIOI:qIIH Hac BOIIpOC 06 aIInapaTe
peiPJICKTOpHOH p;yrII H 0 HCHpOHaJIbHblX OTHOIliCHIIHX y KpyrJIO-
pOTbIX.
H3 HCCJIc]:(oBaHHH IICpHiPCPHQCCKOH HcpBHon TKaHII y MHRCIIH
II I\nrnor (Julin, 1887; Giacomini, 1902a; Johnston, 1908; Kuntz.
1911; Allcn, 1917; Brandt, 1922; ]1,. It TpeTLJlKOB, 1927; Ayers,
1930; Whiting, 1948; Nicol, 1952; Augustinsson et at, 1956;
Rybak, 1960; Jcnsen, 1961; MHJIOXIIH, 1953a, 6, 1961a, 6, 1963),
HCKOTOpblC MaTeplIaJIbi KOTOpblX BOmJIII B pyKoBop;cTBa (Krausc.
1923; Hirt, 1934; Goodrich, 1958), MomHO IIpcp;CTaBIITb COCTaB II
paCIIpCp;eJIeHHC aBTOHOMHOH HepBHoH CIICTeMbi KpyrJIOpOTblX B CJIe-
p;YIOI:qeM BlIJ];e. ITorpaHII'IHble CIIMIIaTlI'leCKHC CTBOJIbI, B KOTOpblX
'raHrJIHH 6b1JIII 6b1 CBH3aHbI IIpOP;OJIbHO HP;yI:qUMII KOMlIccypaMH.
OTCYTCTBYIOT.
CIIMIIaTlI'leCKHC HcpBHble KJIeTKII pa36pocaHbI B CTeHKax TCJIa.
B IIaplIeTaJIbHblX cocy]:(ax, llP;OJIb BeHTpaJIbHblX H p;Op3aJIbHb1X pa3-
\ BeTBJIeHHH CIIIIHaJIbHblX HepBOB II P;UWWY3HO CBH3aHbI p;pyr C ]:(py-
: rOM HepBHblMH BOJIOKHaMH. MeTaMeplIJI CHMIIaTII IeCKIIX aJIeMCHTOB.
: CBOHCTBeHHaH BblCllIlIM II03BOHO'IHblM, Y KpyrJIOpOTblX TOJIbKO HaMe-
l 'iaeTcH II 06YCJIOBJIeHa MOPWOJIOrlI'leCKIIMII KOHTaKTaMH C cerMeH-
TapHblMH BCHaMII TYJIOBIII:qa. CIIJIeTeHIIJI Ha MeCTe aTIIX KOHTaKTOB.
KOTOpblC HaqlIHalOTCH II03ap;II ma6cpHOH 06JIaCTII II IIpOp;OJImaIOTCH
38
BJI,OJlb p;JIIIHHOII OCII TYJIOBII a mIlBOTHbIX, paCCMaTpllBalOTCH
(TpeTbHKoB, 1927) Kal{ 3aqaTOK 6YAY lIx IIorpaHllwblx CTBOJIOB
Bcex Gnathostomen. Y 3TIIX mIlBOT.HbIX cerMeHTapHaJl CBH3b 110-
rpaHllqHOrO CTBOJIa co CIIIIHaJIbHblMII HepBaMII B03HHKaeT Ha TOM
MeCTe, r]:(e Y Agnatha CIIMnaTIIKYC KOHTaKTlIpyeT C cerl\leHTapHbI-
MIl neHaMII.
fOMO.'IOrIlqHOCTL JTIIX CTpYKTYP p;aeT HaM IIpaBO C'IIITaTb, qTO Y I
Kpyr:IOpOTbiX raHrJlII03HbIe CIIMIIaTUqeClme CIIJIeTeHIIH COCTOHT 113
HeiipOHOB 2-ro IIopH]:(Ka )J;BYX 1JIeHHoro aiPiPepeHTHoro 3BeHa peiPJIeK-
TOpHOii p;yrII CIIMnaTllqeCKOD: HepBHOD: cnCTeMbl, a IIX OTpOCTKII pac-
CMaTpllBaTb B RaqeCTBe IIOCTraHrJlUOHapHblX BonOKOH, lI]:(y IIX R IIe-
pniPepllqeCRnM opraHaM.
B )lOpiPOJIOrIlqeCKIIX OIIlIcaHIIHX HepBHoii TKaHII IIeplIiPepllqeCKIIX
opraHOB KpyrJlopoTblX rOBopllTCH 0 IlInpOKOM pacnpOCTpaHeHIIII ee B
IIII eBapIITeJlbllOii Tpy6Ke, cepp;:qe, MOqeIIOJIOBOH II p;pyrux CIICTeMaX
upraHlI3Ma. B aTOM OTHomeHHII IIpep;CTaBJIeHllH CTapbIX aBTopOB, OC-
HOBaHHLle Ha Oqellb T aTeJlbHOM II3yqeHIIII HeHporUCTOJlOrllqeCKII-
I\IH )leTo]:(aI\UI nepuiPepll'IeCKOH IIHHepBa:q1l1l opraHOB, He Bbl3bIBalOT
HUKaKIIX COI\me11ll11 B iPaKTII'IeCKOH CTopOHe ]:(ena, HO He Bcer):{a
II03BonHIOT II3-3a HeJl,OCTa TKa 3KCIIepIIMeHTaJIbHO- MopiPOJIOrlI'IeCKIIX
CBe]:(CHIIH OIIpe]:(eJIIITb IIpllUap;JICmHOCTb Tex IIJ111 IIHblX HepBHblX aJIe-
MCHTOB I{ CII maTIIqeCIHIM 'lIJIII IIapaCIIMnaTllqeCKIIM IIpOBop;HIIKaM.
ITo OTHOllieHIIIO R nIIIIJ;eBapIlTeJlbHOMY KaHaJIy, IIHTpaMypaJIbHbIe I
CIIJleTeHIIH KOToporo OTnOCHTCH K Hall60Jlee II3yqeHHOMY pa3p;eJlY
nepIHvepnqeCKOD: HepBHOD: CIICTeMLI MIIHorOBbIX, Momno C YBepeH- \
HOCTblO CKa3aTb, 'ITO B ero I1llHepBa:q1I1I IIpeBaJIllpYIOIIJ;ce BJIIHlIIlW
OKa3bIBaCT 6JIymp;alOIIJ;IIH nepn. H 3TOMY BbIBO]:(y IIpIlXOP;HT CaMB
RBTOpbl, IIOJlaraH, Ha OCIIOBaHIIII cpaBHIITeJlbHo-MopiPOJlorllqCCKIlX
Ha6JIIOp;eHIIII, ITO BaranbHLle aJIeMeHTbl B nIlIIJ;eBapllTeJlbHOii CUCTeMe
Agnatha IIrpalOT pOJIb aY3poaxoBcKoro CnJleTeHIIH Gnathostomen
(Brandt, 1922; TpeTbHKoB, 1927). B pa60Tax A. A. MIIJIOXIIHa
( 19530, 1963) IIOKa3aHa neplliPepll'IeCKaH peiPJlcKTOpHaH ]:(yra,
B CTenKe KIIllIe IHIIRa I\IlIllOTII. B ]:(UiPiPY3HOM lIepBHOM CnJIeTeHIIII,
e e He p;lIiPiPepeH:qnpOBaIlHOM Ha MLlmCqHOe II nOp;CJIII3nCTOe, aB-
TOp onllcaJi HepBHbIe 3J1eMeHTLI, CXO).\HLle no CTpyKType C KJIeTKa!\UI
II TIIlIa J];oreJIH, p;eHJI,pIlTLI IWTOPLIX 06pa3YlOT pe:qCIITOpHble pa3-
BeTB:leHIIH B TKaHHX KIIIlICqHlllm, a aKCOH - IIepll:qeJIJlIOJlHpllblii an-
napaT Ha ]:(BlIraTeJIbHOH IilleTKe I Tlllla J];oreJiH. Hirt (1934) B Ka-
qeCTBe aprYMeHTa B IIOJIb3Y npellMYIIJ;eCTBeHHO IIapacllMnaTllqeCIWH
IIHHepBa:q1I1I CCblJIaeTCH Ha XOP; OHToreHeTU IeCKOrO pa3BIITIIH, B IIe-
pllO]:( ROToporo ClIl\maTll IeCKlle 3JIeMeHTLI OTCTalOT B pa3BUTIIIl cpaB-
HIITeJlbHO C lIapaCUl\maTII IeCKUMII, II llX BOJIOKHa p;OCTllraIOT Kllllle'l-
HOH Tpy6Kll II03me II B MelibIllIIX KO:III'IeCTBax.
CllMnaTllqeCKlIe BOJIORna onllcaHbl B IIO 1KaX (Krause, 1923), MO-
qeTOqHHlm (TpeTbHKoB, 1927), xpOMaiPiPlIHHOH TKaUII a6p;OMlIHaJIb-
HOH 06JIaCTlI (Nicol, 1952). B aTlIX opraHax 60Jlee BepOHTHO, HaM
Ramel'CH, npcp;CTaBIITb CIiMIIaTlIqeCKYIO IIPIIPOP;y TepMUHaJIbHLlX pa3-
BeTBJleHUH, TaR KaK OHII Haxop;HTCH B6J1113II OCHOBHblX cnJleTeHlIH
39
CHMnaTIi'IeCKHX Hell:pOHOB, II nOCJIeJ1;HlIe MOrYT JICrKO BCTynaTb B
KOHT8KT C HHMH, XOTJI II HMeIOT cJIa60 pa3BIITbIe, KOpOTKHe nOCT-
raHrmIOHapHbIe BOJIOKHa.
Cep]:(e'IHaJI MblIlI a KpymopoTbiX H 06IIJIbHaH IIHHepBa I1JI ee
6b1JIa OIIHCaHa ]1... It TpeTbJIliOBbiM (1927) H Augustinsson C co-
aBTopaMH (1956). nepBblii CqlITaJI HepBIIbIe BOJIoKHa MHOKapp;a CHM-
IIaTH'IeCKHMH, a BTOpbIe, Ha OCHOBaHH'H Mopl!>OJIOrnqeCKoro, i1JH3HO-
JIOrIl'IeCKoro H 6HOXHMHqeClwro aHaJIH30B, nplf3HaBaJIII (npeIIMYlQe-
cTBeHHo) HX IIapaclIMlIaTII'lecKYIO npllpoP;Y. C BBe]:(eHHeM I!>.'IIOOpeC-
eHTHo-rnCTOXHMH'IeCKOrO MeTo]:(a, n03BOJIHIOIQerO 1130JIHpOBaHHO
BblJIBJIJITb a]:(peHeprH IeCKlle CTPYKTYPbl H p;I1l!>l!>epeH HpoBaTb IIX OT
jl;pyrHx HepBOB (Falck, 1962; Falck, Owman, 1965), nO]:(TBep]:(H-
JIaCb IIOCJIep;HJIJI TOqKa 3peHHJI (rOBblpllH II JIeOHTbeBa, 1965a). ORa-
3aJIOCb, 'ITO B OTJIH'IHe OT Bcex p;pyrHx KJIaCCOB II03BOHOqHLIX, Y IW-
TOpblX MOHoaMHHbl B cep]:(eqHoii MbIllIqe KOHqeHTpHpyIOTCJI B OOJIaCTIf
KOH eBbiX OTp;eJIOB CHMIIaTH IeCKIIX HeiipoHoB, Y KpyrJIOpOTbIX 3TII
CTpyKTYPbl He 06HapymeHbI. Cep]:(e'IHaJI MblllI a MllHorOBbiX COBceu
JIHllIeHa CIIMnaTlI IeCKOll: IIHHepBa IIII.
no OTHOllleHHIO K HHHepBa HII cepp; a MIIKCHH ]:(aHHble elQe 60-
JIee p;eMOHCTpaTHBHbl. l\'.oMnJIeKCHbIM IICCJIe]:(OBaHlleM rHCTOJIOrnqe-
CHHMH, I!>H3HOJIOrH IeCKHMH H i1JapMaKOJIOrH IeCKJlMH MeTo]:(aMII (Ry-
bak, 1960; Jensen, 1961) B 1\1II0Kap]:(e 3THX iKIIBOTHblX Bo06IQe He
o6HapymeHa IIHHepBa:qHJI 1.
1I3JIomeHHoe 1I03BOJIJIeT npHiiTH K BbIBO]:(y, qTO TepMHHaJIbHbie
HepBHbIe CIIJIeTeHIIJI B nepHilJepHqeCKHX opraHax OTHOCHTCJI rJIaB-
HblM 06pa30M K BeTBJIM 6JIymp;aIOIQerO HepBa II '1TO, HeCMOTpJI Ha
CKY]:(OCTb I!>II3HOJIOrlI'IeCKIIX p;aHHblX, MomHO 1I0JIaraTb, 'ITO MOTOp-
HYIO i1JYHK HIO P;JIJI BHYTpeHHOCTeii HeceT HMeHHO 3TOT HepB. a HO
CHMIIaTH'IeCKHll:. O'IeBHp;HO, Ha 3TOii CTa]:(IIH 3BOJIIO HOHHoro pa3BII-
TIIJI (KpyrJIOpOTble) OTpOCTKH, OTXOp;JIIQHe OT CIIMnaTHqeCHHX Heiipo-
HOB TyJIOBlIlQHblX Y3JIOB (IIOCTraHrJIHOHapHbIe BOJIOKHa), elQe He ]:(0-
CTlIraIOT HJIII He Bcerp;a ]:(OCTHraIOT CBOerO «MeCTa Ha3Ha'ieHIIJI» B
IIepHl!>eplIqeCKOM opraHe H He HBJIJIIOTCJI, CJIep;OBaTeJIbHO, IIOJIHO eH-
HblMH HeiipOHaMH, cnoc06HbiMII OCYIQeCTBJIJITb 3i1JilJeKTopHYIO - ]:(BII-
raTeJIbHYIO HJIH ceKpeTopHYIO - i1JYHK HIO.
TOJIbKO JIll (mep;Opa3BIITHe» nOCTraHrJIHOHapOB CHMIIaTHqeCKOrO
HeiipOHa p;eJIaeT HeIIOJIHoqeHHoii CIIMIIaTlIqeCKYIO IIHHepBaqnIO Y
KpyrJIOpOTblX? lIJIH OCTaIOTCJI He]:(opa3BIITbIMH H ero CBH3n C Hell:-
pOHaMH ClIlIHaJIbHbIX eHTpOB aBTOHOMHOii HHHepBaqIIII H C aiIJilJe-
peHTHoii RJIeTKoii?
HaH60JIee nOJIHOe IIpe]:(CTaBJIeHlIe 0 HeiipOHaJILHblX OTHOllleHHJIX
y KpyrJIOpOTbiX MomHO IIOJIy IHTb II3 pa60T Tpex aBTopOB. l\'. HnM,
I To o6CTOHTeJIbCTBO, 'lTO BJIeKTpOHHOMIIRpOCROnH'leCRII (Ostlung et aI., 1960;
Hoffmeister et aI., 1961; Caravita, Coscia, 1966) He 06HapYllieHId HepIJHlde BOo
JIORHa B cepP;JJ;e MIIHor, He AaeT OCHOBaHIIH nOJIaraTb, 'lTO TaROBlde Bo06 e
OTCYTCTBYIOT, TaR RaR B BJIeRTpOHHOM MIIRpOCKone MOlliHO BHp;eTb op;HOBpe-
\{eHHO B nOJIe apeHIIH JIlImb 'laCTb RJIeTRH.
40
B lIepBYIO Oqepep,b, OTHOCHTCH iPYHp,aMeHTaJIbHbIe HCCJIep,OBaHHH, IIplI-
Hap,JIemaI.qHe 300JIOry Op,eCCIWrO YHHBepCHTeTa . It TpeTbHROBY H
H3JIOmeHHbIe HM B TpeX MOHorpaiPHHx (1910, 1916, 1927). B HID.
J];aHO OIIHCaHHe M03ra HerO lIepHiPepHqeCRHX CBH3eH Y lIeCROpOH-
RII II B3pOCJIOH peqHOH MHHorH. lI3 60JIee paHHHX HCCJIep,OBaHHu
3aCJIymHBaeT BHHMaHHH pa60Ta Johnston (1908), a cpep,H cpaBHH-
TeJIbHO Hep,aBHHX - HCCJIep,OBaHHe CIIHHHoro M03ra JIHqHHRH MIlHOrH
Whiting (1948). B 3TllX pa60Tax OIIHcaHbl OCHOBHble TIIIIbI qYBCT-
BHTeJI.bHbIX H p,BHraTeJIbHbIX HeHpoHoB H HX CBH311 C lIepHiPepHeH.
IIpHBep,eM 60JIee IIOp,p06Hoe HX OIIHcaHHe.
qYBcTBHTeJIbHble HeHpoHbl JIemaT B CIIHHaJIbHblX raHrJIHHX; HX
lIepHcfJepHqeCRHH OTpOCTOR pa3BeTBJIHeTCH B pa3JIHqHbiX qaCTHX TY-
JIOBH:I.Qa MHHorH, o6pa3YH qYBCTBHTeJIbHble OROHqaHlUl (Retzius,
1892; TpeTbHRoB, 1927; MIIJIOXHH, 1953a, 1963); pe elITOpHble OROH-
'IaHIIH nH:I.QeBapHTeJIbHOrO RaHaJIa IIpHHap,JIemaT, BllWIMO, qYBCTBII-
TeJIbHbIM HeHpoHaM CHCTeMbI 6JIymp,alOlI{ero HepBa (MHJIOXHH, 1963).
QeHTpaJIbHblH OTpOCTOR qYBcTBHTeJIbHOH RJIeTRH BXOAHT B CIIHHHOH
. M03r II, B OTJIHQHe OT Toro, qTO HMeeT MeCTO Y p,pyrHx II03BOHOq-
HbIX, He OIIJIeTaeT HelIocpep,CTBeHHO MOTopHbie HJIlI aCCO HaTHBHble
lme TIm , a BXOp,HT B COCTaB 061I{ero IIOBepXHOCTHoro CIIJIeTeHHH M03-
ra. B o6pa30BaHHH CIIJIeTeHHH IIpHHHMalOT yqaCTHe, IIOMHMO aRCO-
HOB qYBCTBHTeJIbHblX HeHpOHOB, p,eHp,pHTbl p,BHraTeJIbHbIX H acco-
HaTHBHblX RJIeTOR. . It TpeTbHRoB (1910) p;aJI IIOp,p06Hoe, IIpe-
RpaCHO HJIJIIOCTpllpOBaHHoe OIIHcaHHe p,BHraTeJIbHOrO p,Op3aJIbHOrO Hp,-
pa B cnHHHOM M03re MIlHOr, He pa3rpaHHqeHHOe TOlIorpaiPlIqeCRH
Ha COMaTHqeCRHH II BHC epaJIbHbIH OTp,eJIbI, H He IIOp,TBepP.1IJI TeM
caMblM YRa3aHHH Johnston (1908) 0 CYlI{eCTBOBaHHH BHC epaJIbHO-
aiPcfJeRTopHbIX 60ROBbiX porOB. BHraTeJIbHble H acco:qnaTHBHble aJIe-
MeHTbl IIO ,D:. It TpeTbHROBY pacceHHbI B CIIHHHOM M03re H He 06pa3Y-
lOT HII porOB, HH 30H, HH OTp,eJIOB. Y MHHor OTCYTCTBYIOT lIyqROBble
RJIeTKH HJIH RaRHe-JIH6o P,PyrHe 3aMRHYTbie B ce6e CHCTeMbl HeHpo-
HOB, KOTopble 6bl Bep,aJIH pa3JIHqHblMH peiPJIeRCaMH. OTCYTCTBYIOT
Y HlIX H MHROTHble 060JIOqRH Ha HepBHblX BOJIORHax. OqeBHp,HO,
ROHTaRTbI 6e3MHROTHbiX HepBOB C p,BHraTeJIbHbIMH p,eHp,pHTaMH OCY-
lI{eCTBJIHIOTCH Ha npOTHmeHHH Bcero BOJIORHa H 06eCIIequBaIOT xa-
paRTepHblH H ep,HHCTBeHHbIH Y lIeCROpOHRH peiPJIeRC Bcero TeJIa
(TpeTbHRoB, 1910) 1. CTOJIb me HeClIe HaJIH3HpOBaHHbIH peiPJIeRC
OCYlI{eCTBJIHeTCH, BIIp,HMO, H C HHTepope eIITHBHbIX OROHqaHHH BHYT-
peHHHX opraHOB. OTCYTCTBHe 60ROBbiX porOB CIIHHHoro M03ra H nepe-
RJIIOQeHHe HMIIYJIbCOB B nOBepXHOCTHOM CIIJIeTeHHH M03ra, IIpIIHHMa-
IOlI{eM B ce6H BHc epaJIbHO-p,BlIraTeJIbHble BOJIORHa, cIIoco6cTByeT
IIOp,OOHOMY p,HiPiPY3HOMY peiPJIeRCY.
CBep,eHHH 0 ROHTaRTax lIepHiPepHqeCROrO CHMnaTHqeCROro HeH-
pOHa C IIperaHrJIHOHapHblMH CIIHHaJIbHblMH HepBaMH MeHee IIOJIHbl,
1 B03MolliHO, TaKoe CTpOeHlIe npaBOMO'IHO cpaBHHTb C Ka6eJIbHldM CTpoeHHeM
HepBoB BO BpeMH pereHepa 1I1I HepBHldx BOJIOKOH, 06 IIH MlIeJIHHOBldii cJ>YT-
JIHp KOTOpldX AaeT «0606 eHHIdH» pecJ>JIeKC (E. 11. JIaBpeHTbeB).
41
He op;H03HaTJHbl Y pa3HbiX aBTopOB :II qaCTO IIpOTllBOpeqnBbl. C YBe-
peHHOCTbIO MomHO CKa3aTb TOJIbKO Op;HO, qTO, eCJIn n nMeIOTCH CBH-
3n 8iPiPeKTopHbiX Heii:poHoB aBToHoMHoii: HepDHoii: cnCTeMbl RpyrJIO-
pOTbiX p;pyr C p;pyroM ( eHTpaJIbHOrO n lIepniPepn'IeCKoro), TO OHa
HellOCTOHHHbl n He Be3p;e perYJIHpHO BCTpeTJaIOTCH.
IIepniPepnTJeCKne cnMrraTnqeCKne KJIeTKn, CDH3aHHble C Bnc e-
paJIbHblM1'I BeTBHMn ClInHaJIbHblX HepBOB TOpaKO-JIyM6aJIbHoii: 06JIa-
CTn, OlIncaHbI BP;OJIb cpe]:(Heii: TJaCTn ]:(op3aJIbHoii: aopTbi n Kap]:(H-
HaJIbHblX BeH (Marcus, 1910; Ayers, 1930; Goodrich, 1958). XOTH
n O'IeBnp;HO, 'ITO TaKue BOJIOKHa, BLlxop;H lIe II3 CIIIIHHoro M03ra.
CYlI{eCTByIOT, HO nHTeplIpeTa UH nx B KaqeCTBe CIIlIlIIaTUqeCKUX 3a-
TpYp;HeHa BCJIep;CTBUe OTCYTCTB:IIH y KpyrJIOpOTblX pa3BuToH MUeJInHO-
BOH 060JIOTJKU CIInHaJIbHblX HepBOB. AniPiPepeH ua uH Memp;y CUM-
lIaTIITJeCKnMn n CIIUHaJIbHblMn HepBaMn B p;aHHOM CJIyqae HaM IIpep;-
CTaBJIHeTCH Hep;OCToBepHoii:. C p;pyroii: CTOpOHbI, TOT iPaKT, qTO lIe-
puiPepUqeCKne CUMrrarnqeCKIIe KJIeTKII IIpelIMyll{eCTBeHHO
paCIIOJIOmeHbI Ha 1J;Op3aJIbHoii: CTeHKe TeJIa (Nicol, 1952) II, B qaCT-
HOCTn, I'PYIIIIIIPYIOTCH B6JIII3I1 )J;op3aJIbHblX KOpCIlIKOB crrIlHaJIbHblX
HepBOB (Goodrich, 1958), 1I0p;TBepm1J;aeT TO, qTO OHII KOHTaKTnpYIOT"
CBOIlMII OTpOCTKaMII C eHTpaJIbHbiMII p;BnraTeJIbHO-BIIC epaJIbHbllln[
Heii:poHaMH. I1MeHHO B 8TIIX MeCTax Han60JIee peaJIbHO IIpep;CTaBHTb
nx COep;I1HeHIIH, TaK KaK I13BeCTHO, qTO BnC epaJIbHO-p;BIIraTeJIbHbIe
BOJIOKHa y KpyrJIOpOTblX BbIXOP;HT 113 CIIIIHHoro M03ra qepe3 ]:(op-
3aJIbHble KOpelllK1I (3aBap3IIH, 1941; Goodrich, 1958). B TO me-
BpeMH IIOJIOmlITeJIbHblX p;aHHblx 0 rrpncYTcTBIm IIpHMblX cBH3eii: Mem-
p;y OTpocTKaMn lIepniPepIlqeCKIIX cnMllaTIIqeCKIIX KJIeTOK co CIIIIHaJIb-
HLIMII HepBaMII lmC epaJIbHoro XapaK'fepa He rrMeeT'CH, a B p;eTaJIb-
HblX I1CCJIep;OBaHIIHX, KaK1IMH HBJIHIOTCH pa60TbI Johnston (1908) D
TpeTbHKoBa (1927), OHII OTpIl aIOTCH.
_ TaKIIM 06pa30M, rrop;blTomIlBaH p;aHHble 0 HeiipOHaJIbHblX OTHO-
llIeHlIHX B aBTOHOMHOH HepBHoH CIICTeMe KpyrJIOpOTbIX, MOffiHO CKa-
3aTb, qTO lIepIliPepIlqeCKne pe elITopHble IIpIl60pbl II Bee 'IYBCTBII-
TeJIbHOe 3BeHO, lIepeKJIIOqaH I1MIIYJIbC D eHTpaJIbHoii: HepBHOH CII-
CTeMe Ha 8iPiPepeHTHoe KOJIeHO peiPJIeKTopHoii: p;yrIl, He p;OCTaBJIHeT
ero, op;HaKO, K BIIC epaJIbHblM opraHaM 113-3a OTCYTCTBnH perYJIHp-
HbiX CBH3eH Mem]:(y rrpe-n 1I0CTraHrJInOHapHblMn BOJIOKHaMII II n3-38
Hep;opa3BIITnH 1I0CTraHrJInOHapOB. ECJIII II B03MomHO OCYlI{eCTBJIeHne
peiPJIeKCa, TO TOJIbKO B OqeHb OrpaHIIqeHHOM Y'laCTKe TyJIOBIIrr a-
fia p;Op3aJIbHOH CTopOHe TOpaKO-JIYMOaJILHOll (a6p;0lIUIHaJILHoi-i) 00-
JIaCTII, B TOM ep;I1HCTBeHHOM MeCTe, rp;e y 8TllX npII:I1HTIIBHbIX mIl-
BOTHblX rpYllrrnpYIOTCH Han60JIee I1blpameHHble CII maTntlCClme CIIJIe-
TeHnH. GHII oKpymaIOT p;Op3aJIbHYIO aopTY, Kapp;I1HaJIbHble II CeOIeH-
"IapHble BeHbl n Haxop;HTCH n6JIH3H KopelIlKOR CnnHaJIbHblX HepBOB.
GCHOBHble me MaCCbI xpoMaiPiPnHHoH TKaHU, cocpep;OTOqeHHblC B Tex
me MeCTax, HBJIHIOTCH op;HnM :113 TKaHeBblX 9JIeMeHTOB, BXO]:(HII{UX n
CTPYKTypHblii: COCTaB CTeHOK aopTbl II Kap]:(I1HaJIbHbIX BeH II BllleCTe
C HnMU, BIIP;I1MO, 1I0JIY'IaIOII{llX CIIMrraTII'IecKyIO IIHHepBa IIIO. B ;)TIIX
MeCTax OCYlI{eCTBJIHeTCH MopiPOJIOrn'IeCKaH CBH3b xpoMaiPiPnHlIblx
42
3J1eMeHTOB C CUMlIaTU'leCI\1IMU t, HO Y KpyrJlopoTblX OHa BbipameHa
.MeHee O eTJlUBO. no Giacomini II Gaskell, xpoMacI>uHHbIe ,me-
MeHTbI y KpyrJlopoTblX TeCHO ClIaHHbl C raHrJlU03HbiMII KJleTKaMII
He CUMIIaTH'IeCKHX, a CIIIIHaJlbHblX Y3J10B, H CBH3aHbi C HepBaMII
ap;HIIX KopelliKoB. 8TH Ha6J1Iop;eHHH IIpep;CTaBJIHIOT uHTepec C TOqKU
3peHIIH rUCToreHe3a o6eux TKaHeM - xpoMacI>cI>uHHoM H CUMlIaTHqe-
CKOH. II Ta H p;pyraH TKaHH, rpY"IIHpYIO HecH Y B3pOCJlblX MHHor
.If JllPIUHOK BOKpyr CIIUHaJlbHblX HepBoB, pa3BHBaIOTCH H3 KaKoM-TO
06mell aKTOp;epMaJlbHOH OCHOBbl 2 .
OCYlI{eCTBJleHlIe cI>YHKquoHaJlbHOll aKTUBHOCTU xpoMacI>cI>HHHbIX
opraHoB Y KpyrJlOpOTblX MomHO IIpep;CTaBHTb P;BOHKUM IIYTeM: JlH60
qepe3 pecI>JleKTOpHbIM IIYTb - Korp;a CUrHaJi C KaKOrO-JlII6o opraHa
(rrepucfJepHqeCKUe peqelITopHble IIpu60pbl p;OCTaTOqHO pa3BHTbI) pecI>-
JleKTOpHO p;OCTaBJIHeTCH He K TO MY me opraHY (acI>cfJeKTopHblX CBJI-
3eH e e HeT) , a K xpoMacI>cI>uHHbIM opraHaM TOpaKO-JlYM6aJlbHOM
06J1aCTU, II xpoMacI>cfJuHHble aJleMeHTbl, Bbl6paCbIBaH ropMOHbl B Ma-
rHCTpaJlbHblH KpOBOTOK, p;OCTaBJIHIOT UX K «lIoTpe6HTeJlIO»; Jl1I6o-
PyroM IIYTb - CTHMYJlHn;UH xpoMacI>cI>IIHHbIX aJleMeHTO IIpOUCXOP;HT
He3aBIICliMO OT HepBHblX HMIIYJlbCOB, a B pe3YJlbTaTe pa3p;pameHIIH
co CTOpOHbl OMblBaIOlI{eH UX KpOBH, Korp;a Bblp;eJleHHe ropMOHa npo-
lICXOP;UT He3aBUCIIMO OT IIOCTYIIJleHIIH UMnYJlbca. JlH xpoMa4>cI>HH-
BOM TKaHH cepp;:qa RpyrJlOpOTblf(, IIOJlHOCTbIO JlHllIeHHOro CHMnaTII-
"IeCKOH HHHepBa:qIIU, BTOpOM IIYTb HBJlHeTCH eWIHCTBeHHblM II B Ta-
ROM cJlyqae xpoMacI>cI>HHHble KJleTKH B MHOKapp;e KpyrJlopoTblX ocy-
lI:\eCTBJlHIOT, BHp;IIMO, p;Be cI>YHK:qHH: Bblp;eJlHIOT B KpOBOTOK B HymHblc
MOMeHTbI ropMOH H 06eCneqHBaIOT aBTOMaTU3M - 6e3HepBHOM (y
MHKCHH) II JlHllIeHHOll ap;peHeprHqeCKOM l1HHepBa:qHH (y MIIHor) -
Cepp;eqHOM Mblm:qbl. CrOJIb BblCOKue Tpe6oBaHUH, npep;'bHBJlHeMble K
xpoMacfJcI>HHHblM aJleMeHTaM co CTOpOHbl MHOKapp;a, BepOHTHO, H 06'b-
JlCHHIOT CTOJIb me CJlOmHOe CTpoeHHe UX y KpyrJlopoTbIX-- B BHp;e
BeTBHCTbIX, aHaCTOM03HPYIOII{IIX p;pyr C p;pyroM (y MUKCHH) 11 OT-
pOCTqaTbIX, OnJleTaIO UX Ramp;oe MblllleqHOe BOJlOKHO (y MUHor)
KJleTOK.
TaKUM 06pa30M, Bce H3J10meHHOe B aToll rJlaBe n03BOJIHeT 3a-
RJlIOqHTb, '{TO Y KpyrJlopoTbiX OCHOBHble MaCCbl xpoMa4>cI>HHHoM TKa-
HH cocpep;OTOqeHbl B CTeHKax cepp;n;a H KpynHblX COCYP;HCTbIX CTBfI
.JIOB TYJlOBm:u;a H llIeu - B CIICTeMe p;Op3aJlbHOM aopTbl, B n03BOHOq-
HblX II ma6epHbiX apTeplIHX H B Kapp;HHaJlbHbIX BeHax;
1 3geCl> He paa611paeTCH BOilpOC 0 I1YTHX TaKOH CBH311 - HBJIHeTCH JIll OHa pe-
aYJIbTaTOM nenocpegcTBeHHoro KOHTaKTa HepBHoro OKOH'laHIIH Ha xpOMacJ>-
cJ>HHHOH KJIeTKe IIJIH onocpegoBaHa rJIagKoMldmeqHIdM BOJIOKHOM COCYJlIICTOH
CTeBKII.
:2 R COlliaJIeHIIIO, npnxogHTcH OTMeTIITb, qTO pa60T, nOCBH eHHIdX !!THM BOnpO-
caM, nO'ITII HeT, a cpegH HMeIO IIXCH rJIaBHOe BHIIMaHHe ygeJIHeTCH JIH60 ca-
MIdM paHHIIM cTagHHM paaBHTHH BM6plIOHaJIbHldX aaKJIagoR HepBHOH CHCTeMId
II AIIcJ>cJ>epeH lIpOBKe BJIeMeHTOB CHMilaTH'leCKIIX y3JIOB nOrpaHII'lHOrO CTBOJIa,
JIIf60 opraHoreHeay naparaHrJIlIeB, rge c06CTBeHHO pa3BHTIIIO IIX ImeTO'lHIdX
3JIeMeHTOB ygeJIHPTCH MeHbllIe BHHMaHHH, 'leM aHaTOMo-rHCTOJIOrnqeCKOMV
(:TpoeHHIO.
43
xpoMa«lJ«lJHlIHaH TKaHb IIpeRMYDWCTBeHHO paCIIOJIOmeHa B6JIR3II
IIpOCBeTa COCY)J;OB - HeIIOCpep;CTBeHHO DO]:( 8Hp;OTeJIlIaJIbHOn BbICTIIJI-
KOn.
poMa«lJ«lJRHHLle KJIeTKR, P;R«lJ«lJY3HO paCCeHHHbie B CTeHKaX CO-
CYP;RCTOn CIICTeMbI R B Cepp;e 1 IHOn Mblmn;e KpyrJIOpOTbIX, Rp;eHTRqHbI
110 «lJopMe, YJIbTpaCTpYKType R rRCTOXIIMRqeCKRM CBOHCTBaM II IIpR-
Hap;JIemaT K ep;nHOH 8Hp;OKpnHHOii CnCTeMe;
CRMrraTnqeCKaH CnCTeMa Y MUKCRH R MRHOr rrpep;CTaBJIeHa CTPYK-
TypaMR, He RMeIOII{RMR o«lJopMJIeHHblX perYJIHpHblX CBH3en co crrR-
HaJIbHblMR HepBaMn R He CIIoc06HblMR 06eCIIeqRTb 1I0JIHOn;eHHYIO
pect>JIeKTopHYIO p;eHTeJIbHOCTb BIIClI,epaJIbHbIX opraHOB 8TRX mRnOT-
HbIX.
r.4a6a II
PbIBbI
CTpDeHHe H ltWpiPDJIDrHtIeCKHC B3aHMDDTHDllIeHHH CDCYP;HCTblX II
ap;peHaJIDBblX DpraHDB pbl6 B qeJIDM DTpamaIOT elI{e (<UpHMHTHDHblH»
CTPYKTYPHblH ypDBeHb, CBDHcTBeHHblH KpyrJIDpDTbIM, HD IIpIlCTIDCD6-
JIeHHblll K o6eCneqeHHIO a<PiPeKTHRuDro iPYHKD;HoHHpDBaHHH Bcell CH-
CTeMbl Ha p;aHHDH cTyneHlI iPHJIDreHe3a. OCD6eHHocTH, KOTDpble CJIe-
p;yeT paccMaTpHBaTb KaK CJIep;YIOII{HH 6DJIee cDBepllleHHblH aTan B
pa'3BHTHH cIIMnaTD-ap;peHaJIDBDll CHCTeMbl pbl6, 'CBOP;H CH K P;BYM 00-
HOBHblM MOMeHTaM. TIepBblH 3aKJIIOqae CH n TDM, qTD DCHDBl1aH Ma'Cca
'CY1IIpapeHaJFOBDll TKaHH 'cDxpaHHCl'CH TDJIbKD Ha BeHTpaJIbHDH 1I0Bepx':
HDCTH aDpTbI, B CTeHKax CerMeHTapHblX Mempe6epHbix ap epHH, B aK-
CHJIHpHblX H .tIpyrHx ee BeT,BHX (y CeJIaXHH H P;BDHKDp;bllllalI{lIx) II CO-
Cpe;IJ;Ol'DqlIB'ae CH B 'CHCTeMe BeHD3HbiX COCYP;DB - B sinus MDnrD,
B Kap]:('HHaJIbHbIX BeHax II HX pa3BeTBJIeJIHHX, BM'eCTe C KDTDpblMH
npDHlI3bl'BaeT BCIO lIapeHxHMy rOJIDBHOH nDtIKH (y KOCTHD-XpHlI{eBbix
H KOCTHCTblX pbI6). BTDpDB DTHOCHl'CH I p;aJIbHeHllIeMY pa3BHTHIO 'II
MDp<pDiPYHKqHDHaJIbHoMY ooBepllleHcTBoDaHHIO cIIMnaTII'leCKDH HepB-
HDB CHCTeMbl.
PacnpDcTpaHeHHe H CTPYKTypHaH OpraH1I3aD;HH cynpapeHaJIDBDll
TKaHH y xpH:I.QeBbiX pbIO DnlIocaHLI ltlHOrDtIlICJIeHHblMH HOCJIep;DBa.TeJIH-
MH (Leydig, 1852; BalfDur, 1877, 1881-1882; Chcvrell, 1889, 1894;
Vincent, 1897, a, b, c, d; 1922; KDhn, 1898, 1903; BDttazzi, 1902;
Grynfeltt, 1903a; CiacciO', 190G; PDll, 1906; Radu, 1932; YDung, 1933;
FancellD, 1937; Olivereau, 1959). Ha DCHoBaHH aTHX pa60T MDp-
iPDJIDrHH xpoMa4>iPlIHHDH 'CHCTCMbl y aToro OTpHP;a /IlpeP;CTaBJIHeTCH B
CJIep;YIO:I.QeM BHp;e.
OCTpDBKH cynpapeHaJIDBDH TKaHU cnp;RT Ha BeTBHX I!I;Dp3aJIbHDH
aDpTbl B 'BHp;e napHblX TeJIeq, KDTDpbItJ THHYTCH B]:(OJIb BceI'O TYJIOBH-
:I.Qa, D6pa3YH qenDqKH nO' D6e 'CTDpOHbI IID3EDHD IHIIKa. CynpapeHaJIO-
BbIe TeJIbqa HatIHHaIOT'CH n03ap;H cepp;D;a, nJIDTHO 1\ HeMY nplmeraH, II
P;DXOP;HT p;03ap;HeI'D KDHqa nDtIeR (puc. 4). OUH TCCHD cnaHHbl C
Mempe6epHbIMII apTepUHMII (pnc. 5, A, B) II MeCTaMlI BHep;pHlOl'CH B
CTelIKY aopTLI. MeTaMepuDe paCnDJIDmemre IlX HapYlllaCTCH B ncpe]:(-
HeM KDHqe TYJIDBII:I.Qa, rp;e B pe3YJIbTaTe CJIHHHHH HeCKDJIbKHX cerMeH-
TapUblX TeJI D6pa3yeTcH n:pynHoe .CKDnJICHUe xpDMaiP(IJIUIHDH THaHH, C
]:(anHIIX nDp Ha3I.lDaCMOC «aI\<clIJIHpHbIM cepAIWM». Ha3BHHlIe 3TD He
CJIyqaHHD, IIDCKDJIbKY «aTWHJIHpHDe cepp;qe» nJIDTHD cnaHHD CD CTeH-
KaMH art. axillaris (-art. subc1avia) (pIfC. 6, A, B) IIJIH C ee BeT-
45
.
P B c. 4. CxeMa pacnpocT-
paHeHHH CIICTeMbl HagnOqCq-
IIIIKOB aRYJIbI (Scyllium ca-
nicula):
AO - aopra; BY - CHMnaTHqCC-
RHe l'aHI'JIHH; A - cynpapeHa-
nOBM THaHb; I - HHTeppeHaJIo.
BaH THaHb; l! N - nO'lHII; OE-
nH eBOA (Bachmann, 1954)
46
BHMII (puc. 6, B) U IIYJIbCllpyeT CllHXPOB-
HO C lIy.'lbca lIeji cepp, 3; B 4>YBK 1I0HaJIb-
HOM OTlIomeUllll eMY p,JIHTeJIbHOe BpeMH
IIplIIIllCbIB8JIll yqaCTUe B I1pKYJIHTOpHOR
CUCTel\le pL16, IIJIU Ifuaqe - pO JIb BCIIO IO-
raTeJIbHoro cepiJ: a. BlIepBbIe Leydig
("1852) BbICHa3aJI npep,nOJIOmeHHe, qTO
«al<CIIJIHpHOe cepp,l.{e» HBJIHeTCH cBoe06-
pa3HbiM H8p,rrOqe'IHUKOM xpm.u;eBblX pbl6,
npU6JIll3RB TOM caMblM 1I0IIllMaHRe 3Toro
OpraHa H COBpeMeHHblM IIpep,CTaBJIeHIIHM
o lleM, a Balfour (1877, 1881-1882),
II3yqUBllIUH ero paSBUTlIe, YBnp,eJI B HeM
aHaJIOr Mep,YJIHpHOH qaCTll Ha nOqeqHllKa
MJIeKOIIUTaIO:I.QIIX. He6e3bIHTepeCHO
cpaBHeHue, KOTOpOC IIpOBOp,llT Leydig
Memp,y aKCUJIJIpHblM cepp, eM aKYJI II CKa-
TOB co cBoe06pa3HblMR rp03p,eB HbIafB
me,lIe3aMll MllKCllH, OIIRcaHHblMll 110ra-
HeCOM MIOJIJIepOM ( IIT. 110 Leydig, 1853)
1I03ap,1I ma6p, IIO 06e CTOpOHbl cepp, a.
Leydig Haxop,RT CXOp,HLIM rllCTOJIOrllqe-
CKoe cTpoeHue meJIes y Tex U p,pyrux Rll-
p,OB pbl6 II IIpllCOep,UHHeTCH K MHeHllIO
MIOJIJIepa 11., KOTOpblH IIpll3HaeT RX Hap,-
nOqeqHRKaMU MUKCRII. B paBHoH Mepe
Leydig nOJIaraeT, qTO meJIe3bI, IIplIRpe-
IlJIeHHbIe K BeH03HbIM CTBOJIaM 3ap,HeH qa-
CTll TYJIOBll:I.Qa y lIeCKOpOeK, HBJIHIOTCH
aHaJIOrUqHblMU oopa30BaHuHMll.
no COBpeMeHH.bIM B033peHRHM (Lutz,
Wyman, 1927; Young, 1933; Nicol, 1950,
a, b, 1952; Coupland, 1965), «aKCIIJIHpHOe
cepp, e» OTHOCDTCH K TRUllqHblM lIapa-
raHrJIUHM BblcmlIX 1I03BOHOqHbIX. C Me-
p,UaJIbHOH CTOpOHbl K HeMY IIpllMblKaeT ra-
CTpaJIbHblH CHMlIaTDqeCKUH raHrJIDH
(Pitzorno, 1910; Nicol, 19S0b), HJIeTKD
I\OToporo qaCTllqHO IIpOHll3bIBaIOT meJIe-
3UCTYIO qaCTb «aKClIJIHpHOrO cepp,.u;a»
(Nicol, 1952) U BMOCTe CHUM IIpRKpe-
IIJIHIOTCH K 3a]:(HeMY Kapp,RHaJIbHOMY C,II-
HYCY. BOJIblllUHCTBO meJIe3UCTbIX ImeTOK
«al{CII.'IHpHOro cepiJ:u;a» p,aeT 1I0JIOmII-
TeJIbHYIO xpoMa4>4>IIIIHYIO peaK.u;mo II 06-
HapYiKllBaCl' HenpaRlf.TIbUYIO, 3Be3p,qaTYIO,
MlioroOTpOCTqaTYIO CPOPMY C MeJIRHMll
TOMHOOKpallUlBaIO:I.QIIMRCH Hp,paMll (Vin-
cent, 1897c; Nicol, 1950a, b). B aRCHJIHpHOM cepp,l e aRYJI Hopa pe-
HaJIlma 60JIbme, qeM B IIOQRaX II cepp,u;e (Peyrin et aI., 1969).
HeOOJILIlIUe xpoMaiP<pIIIlllLlC CI\OIIJICIIUH Y xpHru;enLIX pLIO HaH1J;e-
Hbl BORpyr HaqaJIa 3ap,HIIX Kapp,lIlIaJILHLIX llCH II pellie B Mellipeoep-
HblX BeHax (Fancello, 1937). TIOIia3aTeJIbHO, qTO TeCHoe IIepeIIJIe-
TeHHe, MeCTaMH IIOJIHOe 'CJIHHHUe CIIMnaTllqeCRHX, cynpapeHaJIOBLIX II
COcyp,HCTLIX opraHOB (pnc. 5, A, B), Cll\6 Ha 3ape HCCJIep,OB8UHU :noii
int lc
Sf!
k
nC'
Pile. 5. nOJJycxelllaTlJ1ICClWI' 1130Upaa,I'HIII' nonepl"lHoro (A) II nr:op;om,Horo
(B) Cpl'30B UOJJaCTJI nO'IeK .mYJJLI (Scyllium canicula)
da - AOpaaJIbHaH aopTa H cc MC}HPCOCplIl>ie BCTBH; Ct' - HapnUHaJIbHaH BeHa; 8V - cyl5Kap-
):IIIHaJIbIJaH BeHa; BUp - cynpapCHaJIOBaH THaHb; sg - CHMnaTII'ICCh"He l'aHI'JIHH; nc - HepB-
UbiI' KJIeTHII; nv - BHCI1CpaJIbHbIe HCpBbI; Ic - npOAOJIbUbIe CBHaH; in! - HHTeppeHaJIOBali
THaub; It - nO'lHH (Young, 1933)
47
06JIaCTU Bbl3bIBaJIO CIIOpbI IIO IIOBOP;Y .pYHK UOIIaJIbHOrO 3HaQeIlIUI
CTOJIb TeCHblX KOHTaKTOB i)TUX TKaHeH. MbICJIb 0 TOM, qTO Hap;nOqeq-
HUK (ero «M03rOBOH» CJIon) rrpe]:(CTaBJIHeT C060H qaCTb HepDHon cn-
CTeMbl - CIwpee, qeM (<KpOBHHYIO meJIe3Y», BblCKa3aHHaH Leydig
(1853), 6bJJIa nop;p;epm8na Balfour (1881-1882), Gottschau (1883)
II A. L!:ocToeBcKuM (1886). VincenL me (1897 a, c), Hao6opoT, OT-
C.Q.J7
A
aD.........
4
.
6
P u c. 6. CxeMa p1!3BeTBJIeHUll aOpTbI CRaTa (Torpedo IDarmOl'ata) (A) 11 aRyJIW
(ScylliuID canicula) (B)
ao - aopTa; a. ax.- aHCHJIHpHaH apTepHR; I>d - nOpaaJIbHblC BeTBH aopTbI; C. ax - aHCH-
nHpHoe «cepAI:Ie»; C8 - cynpapeHaJIOBbie TCJIbI:la (Grynfeltt. 1903)
48
CTalIBaJI lIepnOCTeneHIIOe 3Haqemle 6JIH3KUrU pa'CiIIUJIUmeHIUI cyrrpa-
peHaJIUBblX TeJI K CUCYP,IICTUMY PYCJIY II nUJIaraJI IIpeYBeJIlI'IeHHUH
PUJIb, KOTUpylO Balfuur (1881-1882) npIIllUCbIBaJI IB 3TOM CMblCJIe
CUMnaTIIKYCY. YTBepmp,euue Balfour UCHUBbiBaJIOCb, npaBp,a, Ha 113-
yqeHlfII JIUqUHUqHUrO pd3IHITlUI ceJIaxni'I II nU3BUJIliJIU eMY BIIC'pBble
BblCKa3aTb cou6pamCiHiH 0' HOpBHUM IICTUqUUKe BU3HIIKHOBeHIIH CYII-
papeHaJIUBblX TeJI.
Crrup 31'OT ne .'HlllieH uCHuBaHuH.
H3BOoCTHO, qTU napHble cerMeHTapHble OCTpUBKll xpuMacp«!JUHHuH
TKaHU, cnp,Hru;lIe Ha lleTBHX Mempe6epHbix apTepliH (Kuhn, 1898);
(Fancella, 1937, H p,p.), pa3p,eJIeHbl Ha p,aJIbKII caep,HHHTeJIbHaTKaH-
HblMH IIpOCJIOii:KaMH, up,eTbl KalIcYJIOH 1I Ramp,bIB 1I3 HlIX IIMeeT CHCTe-
MY cucyp,HcTara (Vincent, 1897 8, c; Grynfeltt, 1903 a) H HepB-
Hara (Battazzi, 1902; Yaung, 1933; Nical, 1952) cHa6meHlIH. Ta-
KHM o6pa3aM, ,Kam;:J;bIii OCTpUBaK, B H3BeCTHaH Mepe, HMeeT yme op-
raHHYIO CTPYKTYPY.
,l];eTaJIbHaH pa3pa6aTKa cacyp,ncToH Map«!JaJIarHH 'cynpapeHaJIaBbiX
apraHOB xpHru;eBbIX pbl6 IIpHHap,JIemIIT Grynfeltt (1903 a), nUCJIe pa-
6UT KUTopuru IIUJIyqeHa HCHaH KapTHHa 0'6 HX p,OBOJIbHU CJIOmHaH 'Ba{;-
KYJIHpHUH CHCTeMe. Mempe6epHbie apmpUII, IIU Grynfeltt, a6pasYlOT
{;acyp,IIcTbIH KJIy6aK, us KUTUPUro KpOHb IIUCTYlIaeT B BeHbl, OIIUHCbl-
BalOru;He upraH cHapymH H B'lIap,alOru;lIe HJIH HelIocpe;:J;CTBeHHU B Kap-
p,IIHaJIbHYIO BeHY UJIH «!JapMHpYIOru;HC J];.Be IIUlIepeqHble neTBH, RUTa-
pble CJIIIBalOl'CH B BeHbl, rrpoxO]:\Hru;lIe IIU 6uKaM aupTbl. ::ha CJIUmHa
IIUCTpaeHHaH CHCTOMa BeHU3Haru UTTUKa UqeHb p,eMuHcTpaTHBHo 'lIORa-
3aHa Ha CXCMe Grynfeltt (pHC. 7).
TaKIIM 06pa30M, yme y xpHru;elBblX pbl6 Ha6JIIO;:J;aeTcH «IIpUTuTlIm>
upraHHara KpO'lluu6paru;eHIIH B ap,peHaJIOBbiX meJIe3aX, p,ucTHralOII\HH
BblCORUrO Map«!Ju«!JYHKqHuHaJIbHUrU CUBepmeHCTBa y BbICIlI.IIX n03BU-
HUqHbIX. MeCTHue KpaBuu6paru;eHlIe B 3THX meJIe3aX y CeJIaXHH pa3-
BHBaeTCH, HeCMUTpH Ha TO' qTa HX KJIeTKH, IIUMHMU KalIHJIJIHpHUii
ceTH, aMbIBalOTCH apTepHaJIbHaH HJIII BeHaSHUH KpaBblO, IIpaTeKalO-
II eii B rrpUCBeTax KPYIIHblX cacyp,uB, B CTeHKax KUTOPblX UHH JIeiKaT
(pHC. 8, A, B).
E.cTecTBeHHo, qTO xapaKTep KpoBua6paru;eHHH y6ep,HTeJIbHU CBnp,e-
TeJIbCTnyeT 0'6 3H;:J;UKpUHHUii iKeJIe3HCTUii rrplIpu;:J;e cYlIpapeHaJIaBblX
KJIeTOK. HeCMUTpH Ha 3TU, y Grynfeltt 1I Vincent HallleJICH, IIUMU-
Ma Balfuur, p,pyroH UlIlIUHeHT B JIHl{e Kohn (1898, 1900, 1902, 1903,
1930), KUTUPblii IIUp,'IepKIIB8JI HepBHblH xapaKTep BceH CHCTeMbl.
Y6emp,aJIa ero B 3TOM rJIaBHblM o6pa3uM U6ru;HOCTb pa3BlITIIH ;:J;BYX
TKaHeH (Kuhn, 1898).
n03iKe, KaK 1I3BeCTHU, u6a MHeHIIH UKa3aJInCb clIpaBe]:(JIHBbIMR.
Mbl XOpUIliU renepb 3HaeM U6ru;HUCTb IICTUqHIIKa pa3BllTIIH xpuMa«!J-
«!JIIHHUH II ClIMrraTIIqeCKUH CRCTeM lI3 3KTUn;epMaJIbHOrU 3aqaTKa, CBUH-
CTBeHHuru BceM rr03BUHUQHbIM, II B TO' lI;:e BpeMH HIIKeM He UClIapllBa-
eTCH npIIHap,JIemHOCTb K ;m;J;UKpIIHHbIM upraHaM TaKIIX meJIes, KaK
MU3rUBOH CJIUH Hap,IIUQeqmmOB B3pOCJIbiX acu6eH IIJIII upraH U;YKep-
RaHp,JIH y 3apup,blllleii nbICllIlIX IIU3BUHOQHblX mIIBaTHbIX.
49
vi
P II C. 7. Cxellia KpOB006pall\e-
HUH B cyupapeHaJIOBOIll TeJIbl\e
ai aRYJIbI (Scyllium catulus)
r
C8 - cynpapPHlIJIOBoe TeJIbqe; 00-
aopTa; ai - Mcmpe6epHaH apTepHH;
vi - MellipeoepHble BeHbI; r - nOqRH
do - npogOJ1bHble BeHbI (Grynfeltt.
1903)
vi
ao
¥ ROCTHCTblX pbl6 UCCJIep;OBaTeJIU p;JHITeJIbHOe BpeMH He MOrJIII
HaiiTu CTPYRTypbI, rOMOJIOruqHbIe cynpapeHaJIOBbiM TeJIaM ceJIaxuD:
(Pettit, 1896; Vincent, 1897 b, d), Ha OCHOBaRIIII qero 6blJIO cp;e-
JIaHO 3aRJIlOqeHUe 06 OTCYTCTBUU Y HUX ap;peHaJIOBbiX opraHOB. ¥R-
perrJIeHIDO 'aTom .B3rJIHp;a cnoco6CTBOBaJIII 'U iPn3UOJIOrUqe0RUe :3RC-
rrepuMeHTbl. TaR, B Cpa'BHUTeJIbHbIX onblTax 9KCTpaRTbl 113 lIapHblX
cyrrpapeHaJIOBblX TeJI aRYJI U CRaTOB Bbl3blEaJIU y MbillIeii ap;peHaJIU-
Honop;06Hoe p;eiiCT;Bue, a 9RCTpaRTbl, B3HTble II3 TRaHeii aHaJIOrUq-
HblX yqaCTROB Y ROCTUCTblX, p;aBaJIII oTpfIlWTeJIbHble pe3YJIbTaTbi
(Moore, Vincent, 1898). BrrepBble Giacomini (1902 b) 1 OIIIICaJI y
ROCTUCTblX pbl6 llCTIIHHble cynpapeUaJIOBbie TeJIa, p;aIOlI{lIe xpoMaiP-
iPIIHHYIO peaKD;lIIO C 6nxpOMaTOM RaJIIIH, no pacrrOJIOmeHHbIe B OCHOB-
HOM He B pa3BeTBJIeHIIHX aopTbI, a B CTeHRax Rapp;IIHaJIbUbIX 'BeH.
IIo3me cy:npapeHaJIOBble TeJIa 6blJIn Haiip;eHbl TeM me aBTopOM II y
MHorlIX p;pyrux BIIP;OB ROCTIICTblX pbl6 (Amiurus catus L., 1904-
1905; Murenoidi, 1908-.1909, 1909, 1910; Salmonidi, 1909-1910,
1910-1911, 1911; Anguilla vulgaris L., 1908; Argyropelecus II
Scapelus, 1920-1921). PacrrpocTpaHeilI>I: OHn B pa3HbIX y-qaCT.KaX Be-
H03HOii 'CliCTeMbl, rJIaBHblM 06pa30M, B .cnCTeMe RapJJ;I1HaJIbHbIX BeH II
BP;OJIb OTBOP;HII{IIX nOqeqHblX BeH. ¥ HeRoTopblX rrpe.n;cTaBIITeJIeii RO-
CTIICTblX (yrpII) cynpapeHaJIOBaH TRaHb coxpaHHeTCH elI{e B BeH03-
HOM CIIIIY,ce (Giacomini, 1908).
qTO RacaeTCH Mempe6epHblx BeTBell aopTbI, TO Hn Giacomini.
HII nOCJIep;yIOII{IIe IICCJIep;OBaTeJIU (Baecker, .1928; Overbeeke, 1960;
1 Giacomini nplIHap;J1elliaT cJ>YHp;aIlIeHTaJIbHble IICCJIep;OBaHIIH no 1I3yqeHlIIO cTpO-
eHlIR cynpapeHaJIOBOH H IIIITeppeHaJIOBOH TRaHeH p1d6. OJ.{II1IM TOJlbRO ROCTH-
CTIdM pld6aM nocBH eIlo OKOJIO 20 pa60T, OXBaTIdBaIO lIX nepHop; C 1902 no
1926 r. (nepe'leHb HX eM. Bachmann, 1954; Nandi, 1962).
50
Oguri, 1960; Nandi, 1962; Wakisaka, 1963, a, b; Onozato, Yamamo-
to, 19(6) He roBOpHT 0 HHX RaK 0 rJlUBHbIX MOCTax JlOKaJlII3an;1I1I
cynpapellaJlOBOM TRallH y KOCTHCTLIX pL16. B03MomHO, MopiPOJIOrHQe-
CKOC pacnpe]:(eJIeHUC no XO]:(y Ben03Horo pYCJIa H nOCJIymUJIO npl1-
qHHOii MHorHX Hey}:\a(I nepBbIX If.ccJle}:\OlJaTeJleii, ROTopble, OqeBH}:\HO,
nbITaJIHCb HaMTII IIX B Tex me MeCTax, r}:\e OBI! HX llaXO}:\HT y ce-
JlaxlIii.
"'----
- -
A
v
r
PH C. 8. llonepe'lHlde Cpe31d 3ap;neii: a6p;OMlIHaJI],HoH CTeHKH aKymll (Scyllium
canicula) ua ypOBBe sinus Monro (A) - rpynna nepep;HlIx cynpapeBanoBblx
TeJIe II Ha ypoBHe 3ap;Hell qaCTII nO'WK (B)-rpynna 3ap;HIIX cynpapeHaJIOBIdX
TeJIe
cs - cynpapeHaJIOBbie TeJIhqa; ao - aopTa; ai - Mempe6epHble apTepuu; a. cOe-'IpeBHD-
6pblllilliCe>JHan apTeplllI; sin - sinus Monro; r - nO'lHU (Grynfeltt. 1903)
51
I
I
A
I
A
P n c. 9. CynpapeHaJIOBaH II FlHTeppeHaJIOBaH TRaHb B CTeHRe RapAIIHaJIbHOll
BeHId yrpH (Ophisurus - Ophichthys serpens):
A - a,npeH8JIOBbie RJIeTRII; 1 - IIHTeppeHaJIOBbie RJIeTHH (Giacomini, 1910)
CynpapeHaJIOBbIe TeJIa He rrrpe}];CTaBJIHIOT C060ll I\pynHblx Tl\aHe-
BblX Il\OMnJIeRCOB, a BCTpeqalOTCH I\al\ HaOJIHpOBaHHbIe I\JICTOqHble rpyn-
nbl, I\OTOpbIe UJIU o6pa3YlOT rHea}];HbIC CI\OnJIeHHH, HJIH B BU}];e TOH-
KHX nJIaCTOB CTeJIIOTCH B]:(OJIb BeH03HbIX CTeHOI\ (Giacomini, 1902 b;
1908-1909, 1910, 1911; Baecker, 1928; Overbeeke, 1960; Oguri,
1960; Nandi, 1962; Wakisaka, 1963 a, b; Onozato, Yamamoto, 1966).
ABTOpbl nO}];qepl\HBalOT }];Ba Xapal\TCpHbIX CBOll'CTBa xpoMaq)(jlHHHblx
RJIeTOI\ I\OCTIICTbIX pbI6: pacnOJIOmeHHe IIX HenOCpe}];CTBeHHO IIO}]; 8H-
.r\OTeJIHeM (pHC. 9), r}];e OHH, B}];aBJIHBaHCb B npOOBeT oocy}];a, OMbl-
BalOTCH I\pOBblO "II nepellJIeTeHHe IIX B Hel\OTOpblX MOCTax C 8JIeMeH-
TaMil HHTeppeHaJIOBOll TI{aHU. TaIUiMH I..IeCTaMU qam;e Be-ero HBJIHIOT-
CH CTeHI\H nepe}];Hero OTpe31\a 3a}];Hei-i l\ap}];IIHaJIbHOII BeHbI II BeH ro- -
JIOBHOR noql\lI (Giacomini, 1902 b, 1908, 1911; Young, 1931). B 3TH X
;yqaCTl\aX OHU BbIrJIH}];HT yme Hano}];o6lIe Ha}];nOqeqHlIROB aMIjJ1I6HR.
PacnpOCTpaHeHlI1O U rHCTOJIOrHqeCI\OMY CTpoeHHIO cynpapeHaJIO-
BOR TRaHH y oceTpoBblX Giacomini nOCBHTHJI cnen;lIaJIbHOe iHCCJIe-
}];(JBaHlle (1904), Ha I\OTOpOM CJIe}];yeT OCTaHOBIITbCH, Tal\ Ral\ B Hmf
OTMeqeHbl Hel\OTOpble }];CTaJIH, UHTepeCHbIe C TOql\I1 3peHIIH cpaBHII-
TeJIbHOR MopIjJOJIOrIlH.
fHe3}];a H THmll 'cynpapeHaJIOBLIX iI\CJJe3, Ha'IIIHaHCb Ha ypOBHe
cep}];n;a, THHYTCH B}];OJIb 33}];HIIX Rap}];HHaJIbHbiX BeH }];O XBOCTOBOR
OOJIaCTII, nO}];06HO TOMY, "'ITO Ha6mo;n;aeTcH y I\OCTUCTblX pLI6. OHU
pacnOJIOmeHbl no}]; 3H}];OTeJIUeM IIJIIi Bne}];pmOTCH B TOJIm;y HapymHblx
o60JIOQeI\BeH, r}];e 06pa3YlOT COe}];IIHUTeJIbHOTl\aHHLIe YTOJIIlI,eHIIJI,
npOHII3aHHbIe RanHJIJIHpHOll ceTblO. Hau60JIblllee JIX EOJIlIqeCTBO co-
Cpe}];OTO"'IeHO Ha Me}];I1aJIbHbIX CTeHl\aX Rap]:(HHaJIbHbIX BeH, B TOM IIX
OTpe3l\e, r}];e OHII npOXO}];HT "'Iepe3 nOqeqHYrc Tl\aHb. B 8THX YQaCT-
52
Rax xpDMacI>cI>IIHHbIe 3J1eMeHTbt D6pa3YIOT 1]0.,3: 3H]:(DTemIeM l\ap}];IIHaJlb-
HbIX II nDqeqHblX BeH OIlJlDIlIHDH RJleTD1JHbIH CJlDH, I\DTDpblH B BlIAe DT-
;J:e.'lbHblX THmeH IIJlH ,1I3DJlHpDnaHHbIX RJleTDR rrpDHIlKaeT rJly6DI\D B
rrDqeqHYIO napeHXIIMY - B ee I\paHlIaJlbHYIO JUlMcI><m}];HYIO H XBDCTD-
BYIO 31\CRpeTDPHYIO qa'CTII. KDJlHqCCTBD cyrrpapeHaJlDBDH Tl\anH. BD3-
paCTaeT rrD Mepe npH6JUlmeHIIH 1\ XBDCTDBOMY rroJilOcy nDql\H.
CynpapeHaJlDBbIe RJleTl\1I DCCTpOB HMelOT pa3Hoo6pa3HYlO cPOPMY,
pa3JIII1JHbIe pa3Mepbl, DOJIaAalOT cpaBHHTeJIbHD KpynHblM H]:(pDM (1Ja-
e paCrrDJIDmeHHblM ;mCn;eHTpHQHD) II 3epHHCTOH Bal\yomf3Hp::>BaH-
HDH n;IITDrrJIa31'tIOH.
lI'3 rrprrne;a;eHHblX DnHcaHHH GiaCDmini CJle]Q'eT 06paTHTb BHHMa-
HHe Ha Xapal\TepHbIe BHelllHHe DqepTaHIUI cyrrpapeHaJlDBblX RJle1'DI\:
HapMY C npDCTDH, D6bl1JHD nerrpaBHJlbHD MHorDrpaHHDH cI>DpMOH 1Ja-
e BcerD BCTpe1JalO eHCH y BblCIIIHX rrD3BDHDqHbIX, HMelOTCH I\JIeTUH
C ]:(JlHHHbIM'H BblTJlHYTbIMH OTpDCTKaMII.
HpDMe TDrD, GiaCDmini ]:(aCT DrrIlCaHU€ TorrDrpacI>H1JeCI\DrO pac-
npe}];eJleHIIH cynpapeHaJlDBDH Tl\aHII B TCJle 06CJle,uOBaHHbIX pbl6. XD-
TH DCTpOBRH ee JlemaT B}];OJlb npO]:(OJIbHDll DCII BeeI'D TYJlDBH 3, TOM
He Menee HalI60JlblIllIe llX liOJlHqeCT,Ba CDcpeAOTDqeHbl B orpallUqeH-
HDM Y1JaCTl\e - B D6J1aCTII rrDqCI\, T. e. 13 TDll 1JaCTII TeJIa, r}];e B
]:(aJIbHeUllieM pa3BIrBalOTCH Ha;J:nDqeqHHI\H BbICIlI'IlX rrD3'8DHDqHbIX.
B cpaBHHTeJlbHD-MDP4>DJlo.rH1JeCI\OM nJiaHe npe}];eTaBJIHeT HHTepoc
e e D}];IIH cI>aI\T, ROTDpblll Jllllllb BCIWJIb3b YilIOMHHaCT Giacomini
(1904) npH DnlIcaHlIll cyrrpapeHaJlDBDii Tl\aHH y oceTpoBblX. OCTpDB-
Rll ee, nOM liMO. pacrrpDCTpaHeHIIH B CTeHl\aX TYJlDBH HbIX BeH, B He-
RDTDpblX MeCTax CDrrpHRacalOTCH C HHmHIIMH nD3BDHDqHbiMII }];yraM'II.
r]J;e npDXD}];IIT crrIIHHaH aDpTa. HaM ,l\amCTCH 1303MomHbIM paccMaTpH-
BaTb MOC,TonOJlomeHHe 3THX DCTpOBI\DB B l\aqOCTBC {<OCTaTDqHbIXI) cyn-
papeHaJIDBblX meJle3, crn;c coxpaHIIBIIIHX lwrrraRT C apTepHaJlbHDR
CTeHKDll, CBoucTBeHHbill KpyrJIDpDTblIlI H CeJlaXHHM.
YqaCTI\H HenOCpe}];CTBCHHoro I\DHTal\T3 C aDpTaJIbHblM I\pOBDTDI\DM
cDXpaHHJIHCb II y RDCTlfCTblX pbl6 B ]:(QBDJlbHD 6DJlblll0'M MaclIITa6e.
B 3TDM DTHDlIleHllii rrpe}];CTaBJIHCT 1Jpe3Bbl1JaRHD 6DJIblllDR IIHTepec pa-
6DTa Gerard (1943). K COmaJIeHIIIO, Gerard, orry6J1HI\DBaBIlIIIR CBDIO
pa6DTY B 40-x l'O]J;ax, RDr]J;a Yllie rOCnD}];CTBDBaJia TeOpllJl «rJIDMYCOBI),
rrD;(MeHHBllIaH C.o6DR TeDpHIO 3H}];Dl\pIIHHDH npHpO}];bl aOpTaJlbHblX II
KapOTlI)J;HblX naparaHrJlHeB, He 6b1JI cB06D}];eH OT ee BJlIIHlIlIH II no]:(-
1JIIHIIJI eH Tpal\TDBRY CBDero cI>aRTII1Je-CI\DrD MaTepHaJia. Tal\ I\al\ pa6D-
Ta Gerard, rrpe}];CTaBJIHeT }];JlH pa36HpaeMDro BDnpoca lICI\JIlOqIITeJI',-
Hblii IIHTepec, Mbl DCTaHOBHMCH na Heif IID,1;po6Hee.
Gerard rrplI IICCJIe}];DBaHHII Tpex BII}];DB RDCTHCTbIX pbl6 DTpH}];a 1\1)-
JlIOqenepLIX (LepadDgaster GDuanii, Lepadogaster GandDllii H Cal-
liDnumus lyra) OrrHCaJI B CTeHl\e aDpTbl «rJlDMYC», CDCTDH HH II3 ce-
TlI CIiHYCOll}];DB, Brra}];alO HX MHDrD1JIICJIeHHbIMII DTBepCTIIHMlI B rro-
.:IDCTb aDpTbI, II H3 I\pynHblx KJleTOI\ (npDTOnJIa3Ma KDTDpbIX lIHDf}];a
CD}];epmllT pa3JIIIqHbiX pa3MepDB rpaHYJIbl), DRpymcHHblX ceTbIO D1Jf Hb
Y3Knx MemI\JleTD1JHbIX eJlCll, cDD6 alO uxcH }];pyr C ]:(pyrDM II C rrpo-
. CBeTDM aDpTbt. ABTDp rDBoplIT, 'ITD 8TII I\JIeTl\1I IIMelOT MDpcI>DJIDfIiIO
53-
.ceKpeTopHblX H 'ITO 6JIaro.[(apH YCTpoiicTBY rJIOMyea l\alliJ];aH U3 HIIX
-OMblBaeTCJI nJIa3MOH I;:P0B'H H3 aopThI.
Gerard no.[(p06HO OCTaHaBJlHBaeTCH Ha OIlIIcaHHH MeCTOHaxom.[(e-
HHH «rJlOMyea», lIa CTpoeHHII B 3TOM yqaCTKe aOpTbt H Ha CTPYRTY-
pe caMoro rJlOMyca. C HeKOTopblMH BapHaIJ;IIHMH OH O.[(HHaROB y Beex
"Tpex HeCJle.[(OBaHHhIX BH]:(OB phl6. <1>aKTlI'IeCRHH MaTepHaJi RBTopa
OqeHh IIHTepeCCH II I103BOJIHCT eJlaTb OIlpe]:(eJleHHbte BbtBOp;bl.
n03ap;H TOro MeCTa, r.[(e CJIIIBalOTCH :mHf'pllHXlIaJlLHbte ma6epHbic
apTepHH, .oT aOpTbt OTXOP;HT npoIleiPpHqeCKHe ap1'8pHH, I1HTaIOr:u;oe
e.[(IIHcTBeHHbIii HeiPpoH iPYHKIJ;I10HHpYIOr:u;eH Y 3THX pbl6 rOJlOBHOll
IIOqKH. A em.e n03a]J;H OT I1pOHeiPplI'IeCKHX apTepHH OT aopTbl CJleBa
OT.[(eJIHeTCJI nO.[(KJIlOqHqHaH apTepllH, a cnpaBa - I OpOTKHll 06r:u;I1H
.cTBOJI nO.[(li."IlOqIIqHOll II qpet>Hoij apTepliH (pHC. 10). «rJIOMYC» pac no-
Jlarael'CH Ha MeeTe ;norD aOpTaJlhHOrU tmepeRpeCTa», COCTaBJlHH e.[(H-
Hoe II;eJloe C cocY.[(lIcToii cTeHliOii. B D6J1RCTH (<IlepeKpeCTa» CTeHl\a
aopTbl JIMeeT HeCI\OJlbKO JIHOe cTvoeHHe, qeM B pyrHX oTpe3Kax:
tunica intima nocTpoeHa II3 CJlOH 3H.[(OTeJllIaJlbHbtX RJleTDR, paCnDJlO-
meHHblX Ha TOHKOll 3J1aCTHqeCROH MclM6paHe, tunica media - II3
II;lfpKymqmbIx MblllleqHblX RJleTOI\, a tuni.ca adventicia - 113 ROJlJla-
reHOBbIX 'BOJlOKOH, 60raTbiX eocy]:(aMH t vasa vasorum.), rpyrrnaMII
xpoMaiPiPHHHbIx KJleTOR H MeJlRIIMII HepBHblMH CTBOJllmSMII C pa3-
06pocaHHblMII B.[(OJIb HlIX raHrJlH03HbIMH RJIeTKaMH. «rJl.oMyeHOe» YCT
POiiCTBO, ROTopoe COCTOHT 113 nepenJleTeHIIJI HenpaBIIJlbHbIX co06r:u;a-
IDIIJ;IIXCH .[(PYl' C .[(PYl'oM CIIHYCOII.[(HbIX nOJlOCTen H RaHaJlOB, aBTOp
cpaBHHBaeT e ryoKoH. Ha MeCTe BIl'l.[(eHHH RaHaJlOB B npOCBeT aopTbI
membrana elastica interna npepblBaeTcH. B HeKoTopblX MeCTax «rJlo-
MYCHbIih> Jla6IIplIHT ,oo06r:u;aeTcH C neHaMH, IIpOXD.[(Hr:u;HMII B CTeHKe
cocy.[(a.
. «rJlOMYCbI» MorYT 6b1Tb pa3Hbix pa3MepoB II iPOPMbl. "Y L. Gouanii
.1I L. Gandollii OHII ]:(OCTlIraIOT B )J;JlIIHY 115-280 .lit,. II pacnoJiara-
IDTCH Ha BeHTpaJlbHOii CTopOHe aopTbl. «rJlOMycHbte» RJleTKII pa3HDii
BeJllI'IIIHbl H nenpaBHJlbHOii iPOPMbl, OnJleTeHHbIe RRnHJlJlHpHOH ceTblO,
paCIIOJlOmeHbI y 060IIX BIIP;OB Lepadogaster B a.[(BeHTIII HH (pile. 11).
"Y L. Gandollii OHII co]:(epmaT l'paHYJlbl B I.1;HTOnJla3Me RaK TlIIIHq-
HbIe ceRpeTOpHbIe RJleTKII. Y L. Gouanii TaJUIX rpaHYJI HeT, D;I;8al\0
Gerard, cpaBHIIBaH IIX y 060IIX BII)J;OB pbl6, npH3HaeT, qTO HeT nplI3-
naKOB, no ROTOpbIM MomHO 6bl OTBepmYTb TY me ceKpeTopHYIO iPYHI\-
I.1;HIO '<JTIIX RJleTOqHblX 3J1eMeHTOB II Y L. Gouanii. OH MOTn.BllpyeT 3TO
CCblJlKDll Ha ]:(Pyrne 113-BeCTHbIe npllMepbl, Korp;a ceRpeII;IIH npOTeKaeT
003 MopiPoJlOrllqec.I\IIX npOHBJleHIIii, ocTynHblx npIIMeHHBIlIII IeH Me-
TO,JJ;aM.
Er:u;e 60Jlbllle B :nml OTHOIll€1I11I1 MomHO CKa3aTb 0 «rJIOMYCHbIX»
RJleTKaX Callionymus lyra, liOTOpbIe, B OTJIHqHe OT Lepadogaster,
3anHMalOT cpe]:(HIOIO II BHYTpeHHIOIO 060JlO'II\H aopTbI, T. e. paenoJlo-
meHbl 6mlme 1\ ec npOCBCTY II ]J;oeTHralOT B ]:(JlIIHY 350-530 .litH. no
eo06r:u;eHIIIO caMoro aBTopa, IIX «pa3IlTeJlhHbIMH» oeo6eHHocTHMII HB-
JlHIOTCH, Bo-nepBblX, KOabIJ;e06pa3HOtJ (MeCTaMH) paenOJIOil\eHIIe BO-
Rpyr aopTbl (pIlC. 12, B, B) H, 'BO-BTOpblX, HepaBHoMepHaH 3epHu-
.54
CTOCTb :qUTOnJla3!\lbl (puc. 12, A I, B I, B {), ROTopaH 3aCTaBJIHeT
aBTopa p;y-MaTb 0 ceRpeTopHbIX :qulmax B aTUX RJleTRax. Gerard He
cm.meBaeTCJI B TOM, 'ITO B aTUX RJleTRaX npoUCXOP;UT npo:qecc 06pa30-
BaHUJI rpaayJl, ROTopLIe B p;aJlbHeHllIeM BLIp;eJlJlIOTCH. B nOJlb3Y ceR-
peTopHOH nplIpop;LI 3epHUCTbIX RJleTOI\ rOBopUT II ROHTaRT lIX C I\pO-
BbIO: 60JlblllaJl UX qaCTb Haxop;UTCH HenOCpep;CTBeHHO nop; aHp;OTe.lIueM
(puc. 12, B I), a p;pYI'ue JlemaT cpe]:(u MemRJleTOqHbJX eJleH, ROTO-
pbIe, C OP;HOH CTOpOHbl, oRpymaIOT Ramp;ylO II3 HUX, a C p;pyroii:,- npo-
HURaIOT Memp;y 3Hp;OTeJlUaJlbHbIMU RJleTRaMU U Bna aIOT B npocBeT
aopTbl (puc. 12, B 1).
TaRoBa cI>anTlIqeCnaH CTopOHa pa60Tbi Gerard. A RalWBO B caMOM
p;eJle 1I0JlOmeHUe Be:m;eii:?
TIpemp;e Bcero 06pa:m;aeT Da CCOH BllUMaHUC MopcI>oJlOrUqeCRaH xa-
pal\TepllCTHl\a I\JleTOR, p;aHH3H aI3TOpU 1 U nOl\aaaHHaJl Ha ero 'IIpe-
I\paCHbJX UJlJlIOCTpa:qIIHX. fnCTOJlorU'ICCl\aH II :qUTOJIOrUqeCl\aH CTPYR-
Typbl onncaHHblX 06paaoBaHuii - 3CpHUC.T!JCTb, HepaBHoMepHocTb pac-
npe]:(eJlCHllJI 3epeH n pa3HaH HaCbI eHHOCTb .llMU :qIlTOllJla3Mbl, Tan
me l\aR BeJlllqUHa, OqepTaHllH II pa3Ho06pa3HaJl cI>opMa KJleTOK, on-
JleTeHHblX eJIeBUp;HOii RaUl1J1J1HpHOii ceTblO, HBJlHIOTCH xapaHTepHeii-
IlII1MU npU3HaKaMU naparaHr.JIUOH3pHblX RJleTOI\, HaXO]:(H llX'CH B pa3-
HblX CTap;UHX 'CeKpe:qUll (CM. puc. 12).
q TO RacaeTCH cI>ll6pllJlJlHpHb1X CTPYKTYP, KOTOpbIe OTMeqeHLI Ge-
rard II TpaKTYI01'CH l\aR MuocI>u6pu.ITabl U naJiuque 1\0TOpblX IIpOTllBO-
peqnJlO 6bI npn3HaHUIO 3a KJleTKaMII naparaHrJlUOHapHOH npupop;bl, TO
nIl IIJlJlIOCTpa:qIIII (CM. pUC. 11, B n 12, B I), HU TeOpeTIIqeCI\lle'
co06pameHIIH aBTopa, ROTOpbIMlI on, BlIP;llMO, p OBO]:(cTBOBaJICH npn:
TpaKTOBKe IIX B KaqeCTBe MUOcI>1I6pllJIJlHPHbIX, He IIop;TBepmp;aIOT II He
onpaBp;bIBaIOT aTOii o:qeHI\U. Ha llJlJIIOCTpa:qUHX, nplIBep;eHHblX B pa60-
Te, OHn BbIrJlJlP;JlT Hanop;o6ne TOJlCTbIX, KOPOTKIIX, II3BIITblX IIJIll 'IIpH-
MblX OCMIIOcI>IIJlbHbIX (apreHTocI>UJlbHbIX) HUTeii: KOMnJleKCa fOJIbp;mn.
IIHOl1)J;a CBH3aHHbiX 'C HP;pOM, IIHOrp;a CB06o,l{H0 JJema UX B :qUTOnJla3-
Me, HO He noxomII Ha TOHqaiilllue HUTII MuocI>II6pIIJIJI (CM. PIIC. 11, B).
TaKoe cpaBHeHue aBTopa BocnpIIHUMaeTCJI Ran HaTHHYToe II IICRYCCT-
BeHHoe.
Op;HaKO TaKoe cpaBHeHue nOHap;oom1OCb aBTOpy, OqeBU]:(HO, P;JIH
TOro, qTo6bI flblCKa3aTbCH B nOJlb3y reHeTUqeCROH npeeMCTBeHHOCTII
rJlap;I\IIX Mbllll:q U «rJlOMYCHbIX» RJleTOK, TOqHO TaK me RaR cy ecTBo-
BaHne lIepeXO HblX KJleTOqHblX <lJOPM Memp;y TeMII U p;pynIMu, p;eMOH-
cTpupyeMblx aBTopOM Ha pIlC. 12, A I II BI. Pa60Ta Gerard, RaR MbI
rOBOpUJIII, nOCBH eHa p;ORa3aTeJlbcTBy Toro, qTO OTl\pblTbIe llM CTPYR-
TYpbl Y .KOCTIlCTbIX pbl6 aHaJlOrUqHLI aOpTaJlbHbIM «rJloMycaM» BbIC-
IlIIIX n03BOHOqHbIX. OP;HIIM U3 aprYMeHTOB B nOJlb3Y aTOH anaJlorUII
cJlymlIT reHeTUqec.RaH CBH3b P;BYX TKaHeii: - rJlap;ROMJiIllIeqHbIX 'BOJIO-
KOH II «rJloMycHbIX» RJleTOK, JlK06bI IIOK83aHHaH B CBoe BpeMH IIIy-
MaxepOM (Schumacher, 1938) Ha MJlCKOnIITaIO UX. He rOBopH yme
o TOM, qTO ROH:qen:qUII, Ha ROTopbIe onUpaeTCJI Gerard (Goormagh-
tigh, 1935; Schumacher, 1938; Goormaghtigh, Pannier, 1939), 0 cy-
:m;eCTBOBaHIIU xpoMacI>cI>IIHHblx, HexpoMacI>cI>uHHbJX n CMelllaHHbIX na-
5[)
P II C. 10. CX(, H1 p<1CIIOJIO;I'l'lIIIH
apTl'pl1ii ('aOpTa., J.lIO['O IIl'jJ('-
KpecTM II cocl'l\IH'ii IIPOIIl'clJjJII-
q(,cKlIii OOJIaCTII liOCTI1CTOii l'hi-
UbI (Lcpadoga:-;I.;' (;oranii)
Ae - 8IIII6paHXIHl.1J hULIC apTcpUIf.
Ao - aopTa; C - 'IIWnHaR apTc:
pHn; Gt - Imyuo'IcK npoHcdJpoca;
Rd, Rg - npanall II .rrCB3H no'llUI;
BC-nogHJIIO'III'Inan apTl'pliR (Gcrard,
1!t43)
CA ;
\ rc
\ \ \fAC
\.\
R.g
R'ff
PIle. 11. CX('Ma (A) pac\I!J.lIo-
;+;ellIlH rJIolliyca B CTI'I1Kl' aOj)TbI
KOCTI1CTOii pld6bI Lepadogaster
GOl'anii; II CTpYKTypa (B) rJIO-
MYCRoro YCTpoilcTlla, o'lC'p'um-
Rorv Ha CXCIliU IIpH lOyrOJIbHJI-
KO)I:
B _ cynpapCnaJJOBbIC I<JJCTHH; V -
BcnYJ\1d; t - dJl1Upu.T\J\RpnbIc 1'J10MYC-
nbIC KJICTI<H; end - 3HllOTCJ1Hfi (G -
rard, 1943)
:;
;A
E
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r; " \"
. \1C \ t, .
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," s" .
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:--, :. "-! -<. r
''' i .
50 MK '. . ...... ,"-
I -...;' .
A · -..-::
<
\"":.4.....;-;\. ¥' .
,.... ;. t..'.......::. -U ..,, ,,
.. . =: .. .... ;_ :- .. : . .. . _._' . . f.t'9.
A _ - , . _ .:. _.....__... - ..... ;- _ _ r.
<I . ''t. C '''. " . -....... -.;0,'"
'. , ;_. ,,"-"'3: :"::,,; _.
I'. . \! >- . t)OMI< ---i .'
n"' J
1 ) ca:
6 '-_ __-- -
,;
- -y.......
I" )
'1 II'
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-. rf' ==- c:... «)_____
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. ; "'...... V
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'_:_."-;'.._." - - '>(!:I
A.
r " - ....:::
f ..v ;t..-... '-,
------- "',# '3J! ,). -','- ..
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.., ;t! .-o ' t '\' :',})
I . "'" I ; :J.' & J'G' / ,'.
'l . t'; , . <'V' .
\ ( .\0) . .._. _. . # .. . \...l../ -' !
'-_---.J ,, . £., .;;#<'"
-- - - - "," ....;..,.. _"....-r- ....50 "01
8' - ,... r
B
-<,
I'll c. 12 CXl'}lhI (A ,B, B) paCIIOJlOiI>l'IIIIH I"Jlmlyca lIa pa3IIhlX YJlOUIIJlX a0I'TLI
y IWCTlu'Toii phlOhi (Callionymns lyl'a)
.4 1 , £1, HI _ C1PYIITYPbI I'JIOMYCHOI'O YC7)1ofiC'fII" "" YPOIHIIIX CXCM: .40 - aopTa;
Gl - I:JI)60'1H1I npoHcdJpoca; Bg - nOjJ,IIJIIO'Il\'IIIan aJlTcpllH «(H'rard, 1943)
paraBTJlHeB, 06 II}];eHTHqHOCTII HexpoMaiPiPlIBBblx lIaparaHrmIeB C ap-
TepiloBeB03HbIMH aHaCTOM03aMil H 0 IIpOHcxom]:(eHHH Tex II }];PyrlIx 113
MblllleqBO-HIIHTeJlHOH}];HbIX RJleTOK B HaCTOHl1we BpeMJI Ha}];O apH3HaTb
HeCOCTOHTeJlbHbIMH (rpHropbeBa, 1954) - BeIIOHHTHO, nOqeMY OHIi
OblJlH IIPHTHIIYTbI Gerard R ero pa60Te. Be}];b caM Gerard oTpHrt;aeT
Y ROCTHCTblX pbl6 BaJlHqHe apTepnOBeH03BbiX aHaCTOM030B (nx H He
BIL)J;HO Ha pHCYBKax) , B TO BpeMH KaR 110 Schumacher IIMeHBO OHn
HBJlHIOTCH HexpoMaiP<pHHBbIMH lIaparaHrJlHJlMIl II IIPOH3BO}];HLIMH Mbl-
.IIIeqHblX KJIeTOK, a 110 Goormaghtigh II Pannier IIMeHHO IIX IIpU-
cYTCTBlle CBII}];eTeJlbCTByeT 0 «rJloMY<:Boih> CTpYKType, ROTopaH HeceT
qYBCTBHTeJlbBYIO iPymur;mo. HpoMe TOro, Schumacher YTBepm}];aJl 0
IIpOIICXOm}];eHIIII lIS rJla}];ROMbIllIeqBLIX RJleTOK TOJlbKO HexpoMaiPiP:HH-
HblX lIaparaHrJlHeB, BO He IIpnqnCJlHJI K HTOMY xpoMaiPiPnHBbH'. XOTH
Gerard II He CT8BIIJI rIICTOX:HMlIqeCRHX peaRrt;IIH (R COmaJleHHIO), HO
110 xapaRTepy rt;nTOIIJla3MaTnqeCRHX rpaHYJlHrt;HH, nx lIpnCYTCTBll10
IIJlH OTCYTCTBIIIO B RJleTRaX MomBO rOBopIITb 0 HaJllfqnII B OIIHcaHHbIX
IIM CTPYI{Typax KaR xpoMaiPiPnHBbIx, TaK II HexpoMaiPiPlIHHblX 3.:Ie-
MeBTOB. II Te II }];PyrlIe IIPOUCXO}];HT, lIO MHf'HIlIO aBTopa, 113 rJla]:(KO-
MbIllIeqHblX BOJlOROH.
H3 Bcero CKa3aHHoro 06 IIIITepeCHoii pa60Te Gerard MomBO 3al\-
JlIO'UlTb, qTO IThleOTCH OqeHL 60JlblllOe pasHo06pasHe iPOpM cynpapeBa-
JlOnblX (naparaHrJlIWHapBblx) KJIeTOR - OT llIap006pa3Bbix }];O BbITH-
HYTbIX II BepeTeHoo6pa3Hbix - n HeT HIIRaRHX IIepeXO}];HbIX iPoP)[
MeJI{}];y llIap006pa3HbIMn KJIeTKaMU, onncaHHblMH HM 'B aopTe, II rJIap;-
ROMbIlIIeqHblMH ,HJleMeHTaMH ee CTeHOK. PaBBblM 06pa30M B pa60Te
Gerard He IIOKaSaHO II HO }];0I\a3aHO IIpnCYTCTBlle MIwiPH6pnJIJIHpHbiX
CTPYRTYP B KJIeTI\aX. Bce OlIlicanHble Gerard MopiPoJlOr:HqeCKHe IIpH-
SHaKH XapaI{TepU3YIOT 3H}];ORPIIIIHbIii xapaKTep ,HTIIX o6pa30BaHHH, qTO
IIp1l3HaeT H caM allTOp.
B 3TOH CBH3II Hall-I IiameTcn o'reBb nHTepeCHblM npOBeCTn }];B8
.cpaBBemm: 1) Ba pnc. 11, B IIORa3anbl MeJlKHe, Be Co}];epma lIe
sepeH «rJlOMYCHble» I\JIeTRII, a BHU3Y CJIena (o603BaQeBO 6YKBOii «S»)
1I01\a3aBa rpYlIlIa RJleTOK, ROTopbIe caM Gerard CQIITaeT He «rJlOMYC-
HbIMII», a CYlIpapeHaJIOBbiMII; Te n }];pyrne 110 CTpYKTYpe n}];eHTnqHbl
II pa3J1lfqaIOTCH, ITO HallleMY MHeHIIJO, TOJlbRO rpaHYJlIIpOBaHHOCTblO
rt;nTOIIJla3Mbl: lIepBbIe He CO}];epmaT 3epeB (T. e. JlBJIHIOTCH HexpoMaiP-
iPlIHBblMIl OCTpOBl\a:r.m cynpapeBaJIOBOii TRaBII), a BTOpbIe - 3epHO-
CTbie (xpoMaiPiPlIHHbIe OCTpOBl\1I CYlIpapeHaJlOBOH TKaB:H); 2) IIplI
.cpaBBeB:HlI Ba TOM me plIC. 11, B IIOXpoMaiPiPHBHblX CYlIpapeHa-
JlOBblX KJleTOR Y Lepadogaster Gouanii C XpC'MCJ.iPiPIIHHbIMlI Y Cal-
lionymus lyra (CM. pilC. 12) lIepBble IIpOH3BO}];HT BlIeqaTJleHlle
«aTpoiPnqBbIX», T. e., C HallieH TOtIKII .3peHllH, OHII yme 6e3}];eH-
CTBYIOID;He PY}];lIMeBTapHble 06pa30BaHHH :H OqeBb IIOXOmll Ha KJle-
TOqHbIe HJleMeHTbI KapOTII}];BOll me.'Je3bI MJleKOD:HTaIOID;IIX, B TO BpeMJI
RaK BTOpbIe eID;e iPYHRrt;:HOHIIPYIOT II BbI}];eJlHIOT IIpO}];YKTbI ceRpeTa B
I{POBb.
B 3aRJIIOqeHHe paa60pa pa60Tbi Gerard OTMeT:HM, qTO aBTOp IIbl-
TaJiCH naiITII rOMOJlOrIlqeCK:He mpYliTypbI Y }];Pyrllx ROCTIICTbIX pbl6,
-58
HO, RpOMe Lophius piscatorius, [10[1bITI\I1 OCTamICb 6e3pe3YJlbTaT-
HbIMIl. ABTOp crrpaBeMIlBO rOB pIlT, qTO Y pa3HbiX BII OB pbl6 OHa
MorYT pacrrOJIaraTbCH B pa3HbiX Y'iacTI\ax aOpTLI. Cn-paBegmrnocTb
ero rrpe)1;rrOJlOllieHlIH rrO]:(TBeplli)1;aeTCH aHamI30M cpaBHIITeJlbHO-MOPcI>o-
JloruqeClwro MaTeplIaJla. Ha OCHOBamm 113J10llieHHbIX cI>aRTH'IeCRIIX
)1;aHHbIX 0 cyrrpapeHaJlOBblX opraHax BIIYTPII I\JIaCCa pblO 1\lOmHO [0-
BOplITb 06 061IJ;eH 3aROHOMCpHOCTH, 3aRJlIOqalOIIJ;enCH B rrepeMelIJ;eHHII
8TIIX opraHOB OT MarIlClpaJIbIlOro apTepIlaJIbHOrO PycJla R MarIICT-
paJIbHOMY BeH03HOMY. BrrOJIHe eCTeCTBeHHO, 'ITO rrJlaCTbl cyrrpapeHa-
JlOBOll TRaHII, pacrrpOCTpaHeHHblC Y CeJIaXlIll (II Y RpyrJlOpOTbIX) B CII-
CTeMe aopTbl, coxpaHHIOTCU B aOpTaJIbHOii CTeHRe B BIIp;e OCTaTO'l-
HblX o6pa30BaHIlii II MoryI' BCTpe'laTbCH l'aM B JlI060M MeCTe.
B Ra'leCTBe IIMeHHO TaRIIX OCTaTOqHblX OCTpOBROB CJle)1;yeT pac-
CMaTpOBaTb nall)1;eHHbIe Gerard CTPYRTYPbl B aopTe ROCTIICTblX pbl6;
DX cI>YHRn;HOHaJlLllOe 3Ha'limlle nbIHB.:mCTCH rHCTOJIOrlI'IeCRIIMII MeTO-
)1;aMIl.
fnCTOJlOrnqeClme HCCJle;J;OBaHIIH rrOIia3aJIII, '1TO Hap y C RJleTRa-
Mil, pear:npYIOIIJ;IIMII TlIJID'IHOH xpoMacI>cI>lIIIHOn peaKIJ;I1eH Ha )1;ellCTBHe
cOJleii xpOMa, B HeROTopblX yqa-CTRax cyrrpapeHaJIOBOii TRaHIl IIMeIOT-
CH )1;pyrne KJleTRII, ROTopble, 6Yo)J;Y'lll B CTPYRTypHOM OTHOllieHIIH Tll-
rrll'lHblMIl cyrrpapeHaJIOBbIMII, rre BCTyrraIOT, O)1;HaRO, B peaRn;nlO C 61I-
XpO:\IaTOM RaJlIlJl II OCTalOTCH 6eCn;BeTHbIMII (Grynfeltt, 1903, a; Ci-
BCcio, 1906; Radu, 1932). 8TH rrOCTOHHHO BCTpeqBlOlIJ;lIeCJI HexpoMBcI>-
cI>IIHHbIe RJleTIm TPBRTYIOTCH aBTOpal\1II Jl1I60 RaR (<llepeXO)1;Hble» cI>op-
Mbl (Ciaccio, 1906), Jlu60 RaR HBXO)1;H:m;HeCH B COCTOHHHH «JIOROJl».
Young (1933, 1950) OIIHCBJI nx OROJIO nOqeR (y caMn;OB) RBR «HHaR-
TIlBHble» 4>oPMbi 'H pacCMaTpllBaJi B I{B'IeCTBe B03MOlliHoro pe3epB8
)1;.'1H perrpo]:(yRn;nH. HexpoMacI>cI>I1HHOCTb MomeT 6b1Tb 06YCJIOB.1JeHa
)1;BYMJI nplIqlIHBMII. IIepnBJI - n;HR.1JIl'IeCRaH ceRpeTOpHaJl aRTIIBHOCTb.
B rrpon;ecce ROTOpoii RJleTRa BbI6paCblBaeT ropMOH; 8TO JlBJleHHe Bpe-
MeHHoe, Ha6J1IOtJ;aS1ICH BO Bcex opranax a)J;peHBJlOBOii CIICTeMbl H xa-
paRTeplI3yeT O]J;Hy II'3 cI>B3 meJle3HCToii BRTHBHOCTII RJIeTRIl. BTOpBH
npllqHHa - CHHllieHHe crren;IIcI>HqeCROll aRTHBHIJCTIl RJleTKH, C'BJl3aH-
Hoe -C rrOTepen eIO aH)J;ORpIIHHOH cI>YHRIJ;I1H; ,8TO HBJIeHHe rrOCTOHHHoe..
BCTpeqBeTCH 1I36IIpaTeJlbHO TOJIbRO B HeROT<ipbIX opraHax, n ROTOpblX
B 60Jlbllleii IIJlII MeHbmeH CTeneHIl IIMeIOTCH nplI3HaRII MoPcl>ocI>YHRn;H-
OHaJlbHOD pY]:(IlMeHTaI(HII. B 8THX oprBHax ropMOHbl llMelOTCH B HHq-
'I'OlliHbIX ROJlH'IeCT,nax, '1TO 1I0)1;TBeplli)1;aeTCH cI>JIyopeon;eHn;Hell RJleTOR.
HaJlH'IlIe HexpoMacI>cI>IIHHbIX RJleTOR n xpoMacI>cI>IIHHoii TRaHH Pb16.
YCTBHOBJleHHoe B pB3Hoe npeMJI pB3llbiMII aBTopaMII, 3acJlymIIBaeT
npHCTaJlbHOrO BHIIMBHIIH RaR IlMelO:m;ee TeOpeTIIqeCROe 3Ha'leHIIe. Or-
-CYTCTBHe CrreIJ;HcpIl'leCKOii rHCTOXIIMII'IeCROii peaRIJ;HII B HeROTopblX
xpOMacpcI>IIHHbIX iRJleTO'lHblX ROMII.1IeRCBX y MJleROrrHTalOlIJ;HX HBnJlOCb.
RBR Mbl rOBoplIJlII EO BBe)1;eHHII, OCHOBHOH npnqlIHoii pB3HOpeqlIBbiX
On;eHOR 3TIIX CTPYRTYP, ROTOpbIe CTBJlH pBCCMHTpIIBaTbCJI He IB Ra-
qeCTBe rraparBHrJlHeB, a B Ra'leCTBe «rJlOMYCOB - xeMopen;enTOpOB».
IIpHcYTcTBHe HexpoMacI>cI>HHHbIX RJleTOR Y CeJlaXHll B «aR'CHJlHpHOM
cep)1;n;e» - B opraHe, ROTOpblH llOJlY'IUJI Bce06:m;ee rrpH3HaHHe RaR TII-
59
IIHqHbIH naparaHrJIIIH, CJlymuT npUMepOM TOro, qTO HexpoMaiPiPlIH-
HOCTb RJleTOI\ caMa no ce6e He MomeT UCnOJlb30BaTbCH iI RaqeCTBe
;n;OBop;a npOTnB npUHap;JlemHOCTU IIX R CUCTeMe naparaHrJllIeB. KpOl'tIe
'roro, CJIep;yeT OTMeTUTb, qTO HexpoMaiPiPlIHHocTb naparaHrJlUOHap-
HI.IX I\JIeTOR OqeHb pacnpOCTpaHeHHoe II, RaR MI.I YBIIP;I1M Hume, YHU-
BepCaJlbIIOe HBJIeHHe cpe II n03BOIlO'lHbIX. 3p;e.Cb BamHO no]:(qepRHyTb,
qTO OTpIl aTeJIbHaH rIlCTOXIIIIUPIeCIiaH peaR I1J1, HeH3MeHHO HaOJIIOp;ae-
MaH BO MHorux !\neTI\aX «aKCIIJlRpHOrO cepp; a» XpHm;eBbIX pI.16, He
noc, JlymIIJla op;HaIW, IIOBop;ml I. nCRJIIO'lemno ?TIJrO opralla 1I3 CIICTe-
MI.I naparaHrJlUen, T. e. He CJlY'lIlJlOCb Toro, qTO npOIl30mJlO C HeRO-
TOpblMII naparaHrJIIIRMlI y BI.ICIlIIIX rr03BOHOqHbiX (lIpemp;e Bcero C Ra-
pOTIIP;HI.IM «rJIOMYCOM»). Ha06opOT, Ha rrpOTHa\eHUU p;JIIITeJlbHorO
BpeMeHII, Ha'IImaJl IC Balfour (1877) U IWHQaH Young (1950), Olive-
reau (1959), Coupland (1965) U p;pyrnMu UCCJlep;OB3TeJIHMH BbICRa-
"3bIBaeTCH npIIH UIIlIaJlbHO ep;IIHO;J;YlIllIoe OTHOllieHue R 3TOMY opraHY
RaR R :mp;oRpuHHOMY, npUHap;::remam;eMY rraparaHrJIUOHapHOH cnCTeMe.
8H)J;0I{PIlHHI.Ie CBOHCTBa «aRCnJlHpHoro cep..'J; a» nO]:(TBepm]WHI.I II
iPII3I10JlOrU'IeCI\I1MII :mcnepIIMeHTaMII. IIpeccopHI.IH 3iP«peRT m{CTpaR-
TOB 113 «aRCUJIHpHOro cep]:( a» XpHm;eBI.IX pbl6 6b1J1 nOJlY1feH Oliver,
Schafer (1895); Moore, Vincent. (1898); Lutz, Wyman (1927). ITo.
'CJIe Toro I<3R OI.IJl OTRpI.IT BTOpOH TOpMOH xpoMaiPiPIIHHoii TI\aHU-
Hopap;peHaJIIIH, nOHBnJlaCb cepUH pa60T West c coaBTopaMII (West,
1955; Shepherd, West, 1953; Shepherd et aI., 1953, npoBep;eHHblx
Ha XpHm;eBblx pbl6ax. ABTOpbl nORa3aJllI npucYTcTBue B «aI\CUJlHp-
HOM cepp; e» CeJlaXlIH 1I0pap;peHaJIIlHa -II npUnIICaJlU eMY rJIanHYIO
pOJlb B iPu3IIOJlOrU'IeCI\OM ]:(eiiCTBlm, a Coupland (1953) cqeJI Hopa]:(-
peHaJlIlH ep;I1HCTBeHHI.IM aI\TUBHO p;eiIcTBYIOm;1IM HaqaJlOM (<3RCIIJIHp-
HOro cepp;rra». HaM BCTpeTIIJIaCb TOJlbRO O]:(Ha pa60Ta (Christie, 1933b),
B ROTOpOH 1I0I\a3aH runOTeH3UBHblii aiPiPeRT, Bbl3BaHHbIii (!RapOTHp;u-
HOM» - Bem;eCTBOM, aRCTparnpOBaHHblM .113 aToro opraH a U aHa:IOrUQ-
HblM Bem;eCTBY, Bblp;eJleHHOMY aBTopOM (Christie, 1933a) U3 onyxoJlu
RapoTHp;Horo TeJIa qeJIOBeRa. HaM TpY]:(HO CYP;UTb 0 iPII3UOJIOrIlQe-
cRoii qaCTH pa60TbI Christie Ha CeJIaXUHX. MbI MomeM TOJlbRO CRa-
3aTb, qTO OHa corJlacyeTcH C ero MopiPOJIOrnqeCRUMII II 3M6pIIOJlOrn-
qeCRIIMII p;aHHbIMH, rOBop HMII 0 BaraJlbHOM (napaCHMnaTIIqeCROM),
a He CUMnaTlIqeCl\OM npoucxomp;eHUII RJleT01JHbIX aJleMeHTOB «aRCU-
JlHpHOrO cep.D:rra». KpUTU'IeCI\IIH pa360p ROH en I1U 0 napaCIIMnaTII-
qOC,I\OM IIC'fO'IHRRe pa3BUTIIH naparaHrJlIlOHapHOii TI\aHH paCCMaTpIl-
BaeTCH B rJlaBe VIII, rp;e pa36IIpalOTCH BODpOCbI rIlCToreHe3a xpoMaiP-
IjJIIHHon TRaH.II. qTO me RacaeTCH XpHm;eBblx pbl6, TO Mbl OTMeTUM
TOJlbRO, qTO iPaRTIIqeCRaH CTopOHa pa60TbI Christie (1933b) HIIReM
He 6b1JIa DO]:(TBepm]:(eHa, p;ame Adams (1958) - OpTOp;ORCaJIbHbIM
DpOTIIBHHROM lIaparaHrJlnOHapHoH nplIpOp;b1 TaRoro '(<HeXpoMaiPiPnH-
Hora» naparaHrJIHH, RaI\IIM HBJlHeTCH RapOTIlp;Hoe TeJIO Bblcmnx 1103-
BOH01JHbIX.
TaRIiM 06pa30M, «aRCHJlHpHOe cepp; e» ceJlaxnn DO MopiPOJlOrH-
qeCRIIM DpIl3HaRaM, 6UOXIIMHqeCRIIM cBoiicTBaM H 4'H3110JlOrllqeCRnM
IIORa3aTeJIHM JlBJlJleTCH TIIDllqHblM DaparaHrJlHeM, HO, HeCMOTpJl Ha
60
3TO, cOAepmnT KJlCTlHI, IWTopble He pearnpYIOT clIe n.pnqeCKoii: pe-
aR Hell npH rIICTOXIlMII'leCKOii 06pa60TRe COJlHMH xpOMa II, CJleAOBa-
TeJlLHO, He 06HapYllillBaIOT RaTeXOJlaMHHOBbIX ropMOHOB, BblJlBJlHe-
MblX ;JTIIM cnoc06o:\L Pa3HaH CTeneHb rOpMOHaJlbHOrO COAepmHMoro B
ero KJleTRaX HBJlHeTCH, OqCIHlAHO, OTpameHlIeM pa3Horo .pYHR HO-
HaJlbHOrO HX COCTOHHIIH; HaROnJleHHH ropMOHa B nepHOA CHHTe3a II
onYCTOllieHuH II rrepllOA BbIACJIeIHIH. He UCRJIlOqeHa H .puJloreHeTUqe-
CRH ooyc.rrollJleHH,HI PYAIIMeHTa lIH .pYHlu lIii «aRCHJlHpHOrO CepA a)
RaR opraua, IIC'le3alO ero II 3allICHHe!llOro Y ROCTUCTbIX pbI6 xpo-
Ma.p.pIlHHoii TI-<8HbIO B 1I1i.1r1ICTpaJlbHLIX BeHax. ECTecTBeHHo, qTO B na-
paraHr:llIHX ROCTIICTbIX, IWTOpLIe e e BCTpeqaIOTCH B CUCTeMe aopTbI,
PYAInlallTa IIH BblpameHa CIIJlbHec.
J.I.:m o eHRlI cYllpapeHaJlOBOH TliaHII C TO'II{U 3peHUH ;JHAORpHH-
HOII CPYHR I1I1 R.'IeTOli npeACTaBJIHIOT IIHTepeC pa60TbI Baecker (1928)
II Nandi (1962) Ha IWCTHCTbI X lbl6 Baecker IICCJIeAOBa.:I 14 BHAOB
113 7 CeMellCTB ROCTIICTblX pbI ,a -andi - 123 BHAOB U3 55 ce-
MeHCTB (!).
Baccker nOAPo6HO OCTaHallJlIlBaeTCH Ha 'CTpYRType II rUCTOJlOrU-
lIeCIHlX CHoiicTB3X I-\JleTOR 11 06pal1 aeT BHlIlIWHue Ha pa3UYIO CTeneHb
IIX XPOllWiP.pllIlHOCTII.
Y OAHOH u Toii me OCOOll, Ha OJ.j;Ho:\l rU-CTOJlOrU'leCHOM epe3e
Baccker 06Hapym.i.lBaJi <JJleMCHTbl C eJle 3aMeTHLIM meJlTblM ORpaIllH-
BaHIIp.1I1 IITOnJla3Mbl AO llHTeHCIIBIlO 6yporo. He !lICHee IIHTepeCHbI
ero -onllcaHlIH xpOMa.piPllHHoCTU HJleTO'lHbIX rpynrr y pa3HbIX BIIAOB
OTpHAa HOCTIICTLIX pLIO -- OT IIIITenCI1BHO }RCJlTOii:, npe06J1aAaIO eH y
6blQliOB, AO cJla60ii: y RaplIOBLIX IT, lIaKOHe , OTpI1 aTeJlbHOH y CeJlb-
eBbIX, y ROTOpbIX xpOMa.piPuHHoCTh IIHorAa BbIHBJlHeTCH C TPYAOM
mill eOBceM He orrpe]:(e.:IHeTCH. AnTop cnpaBeAJlHBO YTBcpmAaeT, qTO
npIl onpefJ;eJleHIlIl npllua;J;JlenmOCTII -TRaHU R cynpapeHaJloBoH HeJlb3H
OCHOBhlBaTbCH TOJl_ !,O_H!!. fncIo.xHMlI'leCKOH peaR UU, a HaAo yqHTbI-
BaTb cOBoRynH ocT b xapaRTepHbIX Mop.poJlOrnqeCRHX CTPYRTYP. CTe-
lIeHb IIHTeHCUllHOCTII peaI{TUBHOH {)1{paCRH RJleTOR Baecker CBH3bIBa-
eT C cOAepmaHHeM aApeHaJlHHa. Pe3RHe IWJle6aHHH ero MorYT 6b1Tb
06ycJloBJleHbl pHTMlIqeCRIIMlI 1I3MeHeHItHMH B pa3HbIe ce30Hhi rOAa,
a TaRme .pYHH lIoHaJlhRblMlI nepHoAaMII B mll3HII RJleTKH.
QeJlbIO pa60TbI Nandi (1962) HBJlHeTCH FbIHCHeHHe B03MomHblX
!tlOp oJlOrIlqeCRlIX 3aRoHoMepHocTeii: B3aHMHblX ROHTaRTOB cYlIpape-
HaJlOBOH H HHTeppeHaJlOBOH TRaHeii: OTpHJ];a ROCTIICTblX pbI6. ABTOp
He YCTaHOBHJI HII ;JIWJlOrll'leCRIIX, HIi .pU3110JlOrUqeCRIIX 3aROHOMep-
HOCTeH cpeAH OOCJICJ.j;OBaHHbIX ceMeiiCTB II BImOBIi He BbIHBIIJI Ra-
RIIx-.:m60 HanpaBJIeHlIll ;JBOJlIOWUI Han:nOqe'lHHHOB B CMblCJle CTPYR-
TYPHhlX B3aIlMOOTTIOllleHllii: iJ:BYX THaH£ii: me.T:le3bI. TeM He MeHee ero
pa60Ta eHHa B ;J;pyrOl',1 OTHOIIIeHHH. Ha OCHOBaHUH -o6ll1HpHOrO Ma- \
TepHaJia n AeTaJlbHoro onucaHuH Mop .pOJl Onm Nandi AeJiaeT BbiBOA 06
YHUBepCaJlbHOCTH pacnpOCTpaHeHIIH f.TIallHbIX Macc xpoMa.p.pHHHoH
TRaHIi B CJlCTeMe 3aAHnx RapAHHaJlbHblX BeH H JlORaJlH3a IIH ee y
nOAaBJIHIOlI\ero 60JlblllIlHCTBa BII]I;OB 'ROCTHCTbIX pLI6 illOA :m,lJ;OTeJlII-
eM - y caMoro npoc BeTa cocy]:(a.
61
Nandi (1962), no.n;06Ho Baecker (1928), HallleJI B CYDpapeHa-
JIOBOH TRaHlI pbI6 rpynnbI xpoMa.p.piIHHbIX RJIeTOR, He ]:(alO IJX nOJIO-
mHTeJIbHOH xpOMa.p.pIIHHOH pe arm; IIU. IJ;eHHocTb Ha6JIIo.n;eHllii Nandi
3aRJIIOqaeTCH 'B TOM, qTO OH .n;eTaJIhHO OUl1CbIBaeT HCKJIIOqUTeJIbHOe
IIOCTOHHCTBO, C ROTOpbIM OTpU aTeJIbHaH xpoMacl><PHHHaJl peaR UH 3a-
ROHOMepHO DOBTopHeTCH B op;mlx U Tex me yqaCTRaX, y O.n;HIIX II Tex
iI\e BII)];OB pbl6 H qTO 3TII yqaCTRI1 COBlIa.n;aIOT C Ha6JIIo.n;aBIlIHMIICJI
Giacomini (1910, 1911) H OveJ'beeke (1960). HMeHHo TaI\aH CTpO-
raH JIORaml3a HH He n03BOJIHeT aBTopy COrJIaCIITbCH - C Overbeeke.
KOTOpbIH BH,IJ;IIT npHqImy 9TOrO HBJIelUUI B ney.n;aqHOll .plmcawm TRa-
Hell. )J;aJIee Nandi paCllIupHeT apCeHaJI fUCTOXUl\lHqeCRIIX cpe.n;CTB .n;JIH
BbIHCHeHIIH XBMBqeCIHlX CBOllCTB rOpMOHaJIbHOrO co.n;epmIUlOro CYII-
papeHaJIOBOH TRanH IWCTl1CTbiX pbI6 II, KpOMe xpoMa.p.pBHHoii peaK-
D;BH, DpHMeHHeT Ho.n;aTHYIO (HilJarp, H6kfelt, 1953, 1955). O.n;HaRo
06 Bll 'BbIB , KOTOpbIH .n;eJIaCT aBTop 113CBOIIX Ha6JIIo.n;eUlIll, He
TOJIbRO He y6em.n;aeT, HO Bbl3bIBaeT HeIWTOpOe He.n;OYMeHue: Ha OC-
HOBaHUH OTpB aTeJIbHOH peaI\ nlI Ha I10pa!l;peHamlH OH Bo06 e no.n;-
<BepraeT COMHeHHIO H.n;eHTllqHOCTb rOpMOHaJIbHOrO co.n;epmlnIoro xpo-
Ma.p.pHHHbIX KJIeTOR pbl6 C TaROBblMII y MJIeROUl1TalO IIX. l\Ieil, y TeM
MHorHe aBTopbI 06HapymllJIU B HUX nOJIOmIlTeJIbHYIO xpollla.p.pIlHHYIO
(Giacomini, 1904-1921; Baecker, 1928; Oguri, 1960; Nandi, 1962)
B .pJIyopeC eHTHYIO peaR Ull (Olivereau, 1959) U .n;ORa3aJIU 6uoxlI-
MHqeCI{BMll MeTo.n;aMli npBcYTcTBHe a.n;peHamma, Hopa.n;peHamma, .n;o-
lIaMIlHa II '.n;e30KCH.peHllJIaJIaHBHa (Philipport at aI., 1965), a :)JIe
poHHOMBKpOCKODII'leCRH - co.n;epmaHuc B I\JIeTRaX RaTeXOJIal\IUHOBbIX
rpaHYJI (Wakisaka, 1963 a, b; Onozato, Yamamolo, 1966). B OIlLI-
Tax C pe3epDHHHJa lleH nORa3aHO HCTo eHue 3anaCOB RaTeXOJIaMB-
HOB II3 xpoMacl><PuHHoll TriaHII pLI6 (Yaron, 1970; Banerji, 1970).
B pa60Te Onozato II Yamamoto (1966), HapH.n;y C :meKTpOHHOMB-
RpOCRODlIqecKIIM aHaJIII30M, CYDpapeHaJIOBLIe RJIeTI\II 6blJIn o6cJIe.n;o-
BaHbI rHCTOXUMHqeCRH Ha co.n;epmaHlIe RaTeXOJIaMIIHOB. MCnOJIb30Ba-
HHe MeTO.n;a Wood (1963) D03BOJIHJIO aBTopaM YCTaHOBHTb DpnCYT-
CTBBe Hopa;:\peHammcoAepma Hx ImeTOR (65-70 % COOTHOCHTeJIbHO
RO :BC€M xpoMa.p.pHHHbIM KJIeTRaM), a .n;BoiiHaH o6pa60TRa rJl)'TapaJIb-
.n;erH.n;OM - TeTpaRCII)];OCMlleM (Coupland et aI., 1964) .n;aJIa OCHOBa-
HHe BbICKa3aTbCH B DOJIb3Y cy ecTBoBamIH TpCX TIIDOB RJIeTOR B CYIl-
papeHaJIOBOll TRaHB pbl6: CBeTJIbIe (a.n;peHaJInHco.n;epma ne), TeMHO-
ORpameBItble (Hopa.n;peH:1JIBHco.n;epma ne) II YMepeHHOORpallleHHbIe.
-B- "B"Jl.e-RTpOHHOM MnRpocRoDe COOTBeTCTBYIO ne RJIeTO'IHbIe 9JIeMeHTbi
06JIa.n;aJIII :QailHoll y>O.pMOii: OOIIl() «pBJI bHbIX rpaHYJI: C KpymoH cep.n; e-
:BIIHOn (a.n;peHaJIBHco.n;epma He) , C DaJIOqROBH.n;HOll cep.n;n;eBHHoll
(Hopa;:\peHamlHco epma Be) II COCMelllaHHbIM COCTaBOM rpaHyJIHp-
HbiX cI>OpM. -
IIo cy6MHRpOCROnHqeCROll CTpYRType II rHCTOXUMUqeCRHM peaR-
nHM xpoMa.pHHHbIX l\JIeTOI\ 30JIOTOn Pb161{1I Onozato II Yamamoto
BblCRa3aJIHCb B DOJIb3Y cy ecTBoBaHIIJI RJIeTOqHOll CDen;lIaJI1l3aT IIII
BHYTPH xpoMa.pclJlIHHon TRaHH Pbl6, He3aBllCHMoro 3HaqeHIIH Hopa.n;-
peHaJIIIHa, HBJIHlO erocH He TOJIbRO npe.n;meCTBeHHBROM a.n;peHa.'1IlHa,
62
no HrpalOlI(BM, BH]:(HMO, pOJIb caMOCTOJlTeJIbHOro ropMOHa H, na OC-
nOBaHHIl BCerO :JToro,- B IIOJIb3Y nOJIHOn rOMOJIOrllll cYlIpapenaJIO-
BOM: TI\aHII C xpOMa«p«PllHHon TRaHblO .n;pyrn RJIaCCOB n03BOHOqHblX.
B OTJIllqlIe OT aTIiX aBTopOB Yaron (1970), IICnOJIb3YH TOT me Me-
TO.n; Wood (1963), o6napymBJI TOJIbRO O.n;IlH Tlln RJIeTOR, co.n;ep-
malI(lIX Hopa.n;peHaJillH.
fOMOJIOrllqHOCTb OllJlCaHHbIX CTPYRTYP - cYlIpapeHaJIOBOH TRaHH
pbI6 H naparaHrJIlIOHaprrOM: BbIcmllX n03BOHOqHblX - He BbI3b1BaeT
HIiRaRIIX COMneHIIn:. OCHOBaH.!IeM .n;JIH npH3HaHHH rOMOJIOrlIll, HaR
HaM RameTCH, MorYT CIrym HTb rlICTOJIOrIIqeCRlIe, r IlCT OXIIMllq eC Rlle,
6110XIIMlIqeCRile H «p1l3110JIOrlPleCRlle DOHa3aTeJIll, 0 ROTOpbIX peqb
IllJIa BbIllie. qTO me RaCae'fCH cy ecTBOBaHllJl TaR Ha3bIBaeMbIX HOp-
ap;peHamIHOBbiX H a.n;peHaJIIIHOBbIX ImeTOH II CBH3aHHOM: C aTHM
JIORaJIII3a:qIIH MeTllJIHpYlo eii CDoc06HOCTH, TO :WH BonpOCbI Tpe6Y10'f
cne:qllaJIbHOrO pa360pa (06 aTOM no.n;p06no B rJIaBe VI).
AHaJIII3 «paRTHqeCIWro MaTepHaJIa DO a.n;peHaJWBOM: CHCTeMe pbI6
BblHBIIJI OCHOBHYIO 3aROHOMepHOCTb: rJIaBHbIe Mac.CbI xpOMa«P«PHHHOM:
TRaHII cocpe.n;OTOqHBaIOTCH B CTeHRax Imp.n;HHa.T[bHbIX BeH, 1I0RH.n;alO'f
cep.n;:qe II OT'IaCTII CIlCTeMY aopTbl.
B qeM MOiI\CT 3aHJIlOqaTLCH DplIqHHa TaRoro DepeMe eHIIH? qTO
Dpep;CTaB.TIHeT C060H CHMDaTII'IeCRaH HepBHaH CIICTeMa pbI6 H eCTb JIll
Ral,He-JIH60 xapaRTepHbIe oc06eHHOCTII aToM: CIICTeMbi Y paJHbIX OT-
pH.n;OB pbI6?
" Y CeJIaXDM c.erMeHTapHbIe CHMDaTlIqeCRlIe raHr IIII, paCDOJIOmeH-
Hble B.n;OJIb 3a.n;HIIX Rap.n;HHaJIbHblX BeH II CDIIHHOII aopTbI - PMOM
C cynpapeHaJIOBblMII TeJIaMll, CBH3aHbI C :qeHTpaJIbHOH HepBHon Cll-
CTeMOH Docpe.n;CTBOM 6eJIbIX rami communicantes, ROTOpbIe co.n;ep-
i:I\aT nperaHrJIllORapHbIe BOJIORHa; cepbIx me rami communicantes
II, CJIe]:(OBaTe.T[bHO, DOCTraHrJIIIOHapHblX BOJIOROH R Rome II ee .n;epll-
BaTaM HeT, H He HMeeTCH RomHoH CHMDaTHqeCROH HHlIepBa:qHII
(Young, 1933) . Y XpHlI(eBblX pbl6 rre Ha6JI1o.n;aeTCH perYJIHpHblX
MemY3JIOBbIX CBH3eu II, CJIe.n;OBaTeJIbHO, 1JeTRO BbipamelIHOH 1I0rpa-
BHqHOM: CHnaTllqeCROl1 :qenoqlm (Young, 1933; Nicol, 1950a, b,
1952; CRoplI:qRaH, CaQRO, 1967). O.n;HaRo, 6JIaro.n;apH TOMy qTO DO-
rpaHHqHbIe ClIMnaTHqeCRlle Y3JIbl y HHX HMelOT coe.n;IIHllTeJIbHbIe BeT-
Bll co cnllHaJIbHbIMIl HepBaMll, aHaTOMll'leCROe pacnOJIOmeHlle IIX
60JIbme npll6JIllmaeTCH R cerMeHTapHOMY, qeM Y RpyrJIOpOTbIX, II OHIl
IIHnepBllpylOT THanll TOJIbHO COOTBeTCTBYIO IlX cerMeHTOB Te Ia. Bepx-
HIIe raHrJIllH, pacnOJIOmCHHbIe Y 3apo.n;blllleH xpmu;eBbiX pbl6 cer-
MenTapHO, B npo ecce OHToreHe3a CJIllBalOTCH II 06pa3YloT CaMIJM
. RPYIIHbIH raCTpaJIbHbIII CHMDaTll'leCRHH Y3eJI D03a.n;1l ma6epHblx AP-
TepHH II He.n;aJIeRO OT qpeBHOH apTepHH. Y B3pOCJIbIX oc06eM: aRy.lI
II CRaTOB OH CBH3aH C HeCROJIbRHMII CerMeHTaMH CDlIHaJIbHbIX Hep-
BOB (Chevrel, 1889; Young, 1933, 1950).
IIoCTraHrJIIIOHapHble BOJIORHa OT CHMDaTHqeCRIIX Y3JIOB K BHYT-
penHOCTHM Il.n;YT, o.n;HaRO, He co CDHHaJIbHblMH HepBaMII, a DO Xo.n;y
CerMeHTapHbIX, XBOCTOBbiX H ma6epHblX apTeplIH. OHH 06pa3YIOT ry-
GTOe HepBHoe CDJIeTeHHe BORpyr Mblme1JHOH 060JIOqKH 3THX apTe-
63
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, .
. 10 M-K""-" ..
,;;.-
".\,jl-..
'!
. n
r- I
PH c. 13. HepBHaH RJIeTRa Ha rpaHlII{C Mem)lY CIUmaTll'leCKIU\ ml'."lYAOqHbUi
raHrJIJ].eM 11 xpmJacpcpllHHOii TKaHhlO B ('aKCIIJIRpHO 1 cl'p;me» aKY.:IbI (Scyllius
canicula) (no RaXaJIIO)
BUp - cynp3peH3JIOB3H TH3Hb; f - HepDHOC BOJIOHHO C TCpMIIHaJIHMII; gl - A(,lIgpIfTII'IC-
CHHD HJIy60H; n - HAPO ncpBHofi HJICT.nI (YoulIg, 1933)
pIlll, ,lWP3aJlbHOll aopTbi II B Macce cynpapeHa::IOBbIX meJIea. nO YT
BeplliAemno Young ( 1933), ;)TII meJle3bl o6.TJ8AaIOT TaROli 60raTC
pa3BeTBJleHHOH ceTbIO 6e3MHRoTHbiX HepBHbIX BOJIOROH II CIIMnaTlIqe-
CRIIX RJleTOR, Rafwii HeT HII B OAHOM APyroM opraHe CeJlaXIIH, II,
CJleAOBaTeJlbHO, HaXOAHTCH nOA IIOCTOHHHblM ROHTpOJleM CIIMnaTIIqe-
CROH nepBHOH CIiCTeMbI.
Young YAaJlOCb npOCJle]:(IITb XO]:( MOTOpHbIX nOCTraHrmtoHapHblX
BOJlOROH, OTXOAHIqIlX OT CIIMnaTII'IeCRIIX HJleTOR, AO IIX OIiOH'IaHlIii
Ha TeJle xpOMa.p.pIIHHbIX MeTOR (pIle. 13). ::ho Ha6moAeHue IIMee-r
IIpIIH IInIlaJIbHO BamHoe 3naqeHlIe AJiH nonIIMaHIIJI II npaBIIJlbHOii
oqeHRII I1HHepBa 1I0HnbIX OTHOllieHIIH CIIMnaTlIqeeROii II a]:(penaJIoBoii
CIlCTeM y n03BOHOqHblX mIiBOTHbIX. 113BecTHo, qTO nocJle pa60Tbi
Elliott (1913) IlInpOROe paenpocTpaneHIIe nOJlY'IIIJla ero fJmOTe3a
o TOM, qTO xpOMa.p.pIIHHaH RJleTRa HBJlHeTCH BTOphIM (nOCTfaHrmW-
HapHbIM) HeHpOHOM B ABYXqJleHHOH CIIMnaTII'IeCliOH qenO'IKe. 810
npeAnOJlOmeHlle, BHaqaJle IICnOJlb30BaBllIeeCH B :UlTepaType KaK rll-
nOTeTIIqeCIWe, nOCTeneHHO npO'IHO BOIllJlO B 06I1xOA HaY'IHOM .:IIITe-
paTypbI, II 6e3 AOCTaTOqHbIX Ha TO OCHOBaHl-lH npIIHIIMaeTCH B Ha-
CTOHIqee npeMH 60JlblllIIHCTBOM aBTopOB RaR HenpeJlOmHblii .paRT.
Pa360p IIHHepBaqnoHHhIx oTHollieHIIH MemAY HepBHoH H xpm.m.p.pI1lI-
HOH CIiCTeMaMII c]:(eJlaH HaMIl B rJlaBe 06 aApeHaJloBoii CIICTeMe MJIe-
ROnllTaIOIqIIX (rJlaBa VI), rAe Ha npIIMepe M03rOBoro eJlOH HaAno-
qeqHIIROB nOABepraeTCH RpIITIIqeCROMY paccMoTpeHIIIo II rllnOTe3a
Elliott. Mbi OTMe'IaeM 3AeCb .paRT, ROTOpblH 06HapymIIJI Young Ha
CeJIaXIIHX, nOCROJIbRY OH HBJlHeTCH eAIIHCTBeHHbIM, onIIcaHHbIM Y
JlII3ll1IIX n03BOHOQHbIX, II HeonpOBepmIIMbIM ]:(ORa3aTeJlbCTBOM OIllII-
60qHOCTII npeACTaBJIeHIIH Elliott H BceH ROH enqIIII 0 xpoMa.p.pIIH-
HOH RJleTRe RaR BTOpOM HeHpoHe. C TOqRII 3peHIIJI aBOJIIO HOHHbIX
3aRoHoMepHocTeH HeMbICJlIIMO npeACTaBIITb ce6e TaRoe nOJlOmeHlle,
qTO Ha OAHOU 113 cTyneHeH IICTOplIqeCROrO pa3BIITIIH xpOMa.p.pIIH-
HaH RJleTRa HBJlHeTCH BTOpbIM CIIMnaTIIqeCRIIM HeupOHOM, a Ha ]:(py-
rou OHa CTaHOBIITCJI xpOMa.p<);JIIHHOii RJIeTROH sue generis.
J{ COmaJIeHIIIO, HaM He YAaJIOCb HaHTII B JlIITeparype cBeAeHIIll
o qYBCTBIITeJIbHOH IIHHepBaqIIII BHyTpeHHIIX opraHOB xpHIqeobIX Pbl6,
64
ROTOpLIe 6b1JUI I5b1 nOCT8TOqHLT ;;.:m TOro, qT06b1 COnaBlITb no:moe
npe;:J;CTaBJleHIIe 0 CTeneHI! pa311IlTUH Y HHX CHrHaJIbHOll ClICTeMbi.
OAHaRo, Ha OCHOBamIlI oTAeJlbHblX :.JRCnepI!MeHTOB, nOIm3hIHalOllI,Hx,
'ITO l\IeXaHllqeCROe pa3ApamelUw KI!llIe'lllllRa BbI3bIBaeT nocTynaTeJIb-
uoe pew.'1eKTOpnoe CORpalll,eHHe ero (Young, 1933), l\I01l\HO llo.laraTb,
'ITO B KaKoll-To CTeneHII Y ceJIaXUll pa3BHTa MeCTHaH pel}J.leRTop-
HaH ACHTeJIbHOCTb CIlMnUTnqeCKOl1 CHCTeMbI. HallOOJIee AeMOHCTpa-
TliBIIO 3TO nOKa3aHO B onbITaX lIa COCYAIICTbIX HepBax: m..cnepH-
1IleHTa.lbHoe nOBbIllIeHHe lipOBJIHOrO naB.TIeHIIH y aKY.rr. Bbl3BaHHoe
pa3ApameHHeM qYBCTBIITeJIbHblX HCpBOB (Mackay, 1931; Huntsman,
1931) ImH BBe]:(CHlleM aApeHaJIIIHa (Lutz, Wyman, 1932), cUIIAeTeJIb-
cTByeT 0 CYIll,CCTBORaHlIH 1I0JIHO eHHOrO peWJIeKTopHoro OTBeTa.
OAllm..o 1I0CJle ncpepC3liH 6.1Yi1i alOllI,erO HepBa pa3Apafficllue wa-
paAII'IeClilll\l TOROl\l lIepBOrO ClIlIIllUTII'leCliOrO raHr.ll1H 11.111 ClIIlHIIoro
l\I03ra m..YJlbI He LbI3bIBaeT YCRopemm cep]:(eqHoro pHTlIIa (Lutz,
1930a), a 1I0CJIe y]:(aJIeHIIH nepBoro CIIMnaTIIqeCIWro raHI'JIliH 3a-
;I;epmllBalOllI,ee B:IIImme 6JIymAal0lll,ero lIepBa lIa COIipalll,PHUe MbIID-
l\hI cepn a OCTaeTCH (Lutz, 1930b).
Y KOCTIICTbIX pLl6 CIIMlIaTII'leCKIIe 1I0rpaHIIqllble CTBOJILI IIOJIHO-
CTblO CWOpMllpoBaHbI (CKopnqKaH, Caqlw, 1967) H nOCTpoeHbI 110
TOMY me IIJIaHY, ImK II Y 1\1:leROIIIITaIOIll,I1X. IIpoMemYToqHoe IIOJIO-
meHBe B CTpoeHHII CBMnaTIIqeClWX CTBOJIOB 3aHIIMalOT oceTpOBbie
p1>I6b1, Y ROTOpblX raHrmlH 6ecnopH!1;OqHO pacceHHbI IIO coceACTBY
C KapnHHaJIbHbIMH BeHaMH, BAom, JlHMWOHgHOii II :JKCKpeTOpHoii qa-
cTeii Me30HeWPoca. CHMnaTIIqeCRIIe CTBOmlIl.I1 06pa3YIOT Ha CTeHRax
RapAIIHaJIbHbIX H B03BpaTHbIX BeH 1I0qeK CIIJlereHHe, B KOTOpOM pac-
ceHHbI CHMnaTH'leCRIIe IweTKll. B OTHOllleHHH CBH3H I1X C cynpape-
HaJIOBblMH RJIeTRaMH MbI npIlFOAHM OlIllcaHIIH Giacomini ( 1904) ,
ROTOPOMY npHHaAJIemUT HaH60JlIJe lIoAP06Hoe HCCJIenOBaHHC oceTpo-
BhIX. ABTop no]:(qepKIIBaeT, qTO ROHTaRTbI MemAY ABYMH TRaHHMII
OqeHb penRII H qTO, CJIe]:(OBaTeJIbHO, JIORaJIII3a BH xpoMaWWHHHoii
TRaHII pHAOM C raHrJIHHMII II lIepBHbIMU CTBOJIBKaMII He HBJIJleTCJI
HX 06blqHblM MeCTOM. ECJIlI H BCTpeqalOTCH B 3anHeii 06.'1aCTII TY-
JIOBHllI,a eAHHHqHble HJIH 06'beAlIlleHHble B MeJIRHe rHe3na xpOMaw-
WHHHble RJIeTRII BHYTPH CIIMlIaTII'leCliHX raHrJIBeB IlJIlI HepBHbIX CTBO-
JIHROB, COIIYTCTBYIOIll,He Be HaM, TO RaR HCKJUOqeHHe, a He liaR npa-
BIIJIO. 8THM waRTaM Giacomini (1904), Ral, II MHorHe cro npen-
llIeCTBeHHIIRII, IIpBnaBaJI 60JIbIDOe 3HaqeHlIe.
Y ROCTIICTbIX pbl6 raHrJIHII CBH3aIlbl co cnmra;lbHblMH HepBaMII
IIperaHrJIHOHapHbIMII (MHROTHbIMII) I1epBHbIMH BOJIORHaMlI, npOXOAJI-
IlI,HlIUI B 6eJIbix H cepblx rami communicantes (Young, 1931; Ni-
. col, 1952). IIperaHrmlOHapHble BOJlORHa MorYT 3aRaHqIIBaTbCH Ha
CIIMlIaTHqeCRHX RJIeTRaX CBoero cerMeHTa, HO MOryT nepeXOAHTb Ha
RJIeTRII BbiIDe- II HllmeJIemaru;HX CCfMeHTOB. TaRoe CTpoeHHe C03AaeT
60JIee BbIrO]:(HbIe YCJIOBHH ]:(JIH awweRTIIBHoii ROOpnHHa HH WYHRqHII
BHYTpeHHHX opraHOB. AHTarOHHCTlIqeCRaH HHHepBa HH, RaI{ II Y ce-
JlaxBll, He Be3Ae BblpameHa. TaR, Me30Hewpoc II rOHa;I;bI JlHHepBII-
PYIOTCH TOJIbRO cHMnaTHRycoM (Young, 1931), B cepnqe Y ceJIaxlIii
3 H. A. CMilTTeH
65
CHMnaTIPIeCROH MHHepBa:L HII He IIMeeTCJI (Stannius, 1849; Lutz,
1930; Young 1939; Nicol, 1952), a y ROCTHCTblX pbl6, 110 AaH-
HbIM B. A. rOBblpl:lHa II r. P. JIeoHTbeBoH (1965 a) II B. A. rOBhlpH-
Ha (1967), OHa BblpameHa HeJlCHO. OAHaRO B pa60Te Gannon, Burn-
stock (1969) y ABYX BIIAOB ROCTIICTblX pbl6 (Salmo iredius II S.
trutta) 06HapymeHbI RaR XOJIIIHeprlPIeCRlIe, TaR II aApelleprlIqeCI lIe
BOJIORHa (.pJIyopec:qeHTHo--rHCTOXHMHqeCRHM MeTOAoM) B BeH03HOM
cllHyce, npeAcepAHHX, JIYROBIII{e aopTbI, B CTeHRax RopOHapHblX apTe-
plIll II aopTbI. B TO me BpeMH Randall (1968), Ha OCHOBaHHII
3JIeRTpORapAHorpaMM plIAa ROCTHCTJ.,IX pbl6 (.popeJIb, Tepnyra II AP.),
nOJIaraeT, qTO ClIMnaTlIqeCRaH IIFHepBaI{HJI CepAI{a y pbl6 OTCYTCT-
ByeT. ::ho He corJIacyeTcH C npemHIIMII IICCJIeAOBaHIIHMlI, B ROTOpblX
aBTopbl HaIDJIH HepBHbIe BOJIORHa, RaR 3aAepmllBalOlIJ;He, TaR II YCRO-
pHIOII.\Ue cepAe1.IHYIO AemeJIbHOCTb (Clark, 1927; Izquierdo, 1930;
Young, 1931). TaRBM 06pa30M, CJIc]:(yeT OTMeTBTb, qTO OTHOCHTeJIb-
no cBMnaTHqeCROH IIHHepBaI{IIB cepAI{a ROCTHCTblX pblO neT eAHHoro
MHeHHH H TpeOYIOTCJI AOnOJIHHTeJIbHblC HCCJIeAOBaHIIH.
H OTHOllleHHU xapaRTepa HepHHLIX cn:WTeHHll, oopa30BaHHLIX nOCT-
raHI".'IHOHapHbIMH BOJIORHaMlI, II IIX pacnpeACJIeHBH Ha RpynHblx
cocYAax TYJIOBHlI.\a ROCTl:lCTbIX pbIO HeAaBlIO 6b1JIO YCTaHOBJIeHO
r. P. JIeoHneBOH (1966 a), qTO .pmoopec:qeHI{IIH CIIMnaTlIqeCRIIX
BOJIOROH, cHaomaIOlI{lIX MarIlCTpaJIbHble COCYAbl, AOCTilraeT HaUOOJIee
MOII{HOro pa3BHTIlH B aopTe. «1>JIIOOpeCI{eHI{IIH xpoMa.p.pHHHbIX I\Jle-
TOR Ha CTeHRax 3aAHIIX RapAHHaJIbHblX BeH elO He 6bIJIa oOHapy-
mella. 8TH Ha6JI£OAeHHJI JIeoHTbeBOH, C OAHOH CTOpOHbI, nOATBep-
AHJIlI C nOMOlI{bIO OqeHb ClIeI{H.pIlqeCROrO AJIH aApeneprlPIeCRHX Be-
lI{ecTB MeTOAa HaJIHqHe xpOMa.p.pHHHOH TRaHil B RapAHHaJIbHblX Be-
Hax ROCTII.CTblX pbI6. C APyroll CTOpOHbl, OHII nORa3aJIJI HOBbIe
oc06eHHOCTU B opraHH3aI{I111 CHMnaTIlqeCRHX CnJIeTeHlIll TYJIOBlllI{a
y IIpeACTaBl1TCJIeH 3Toro OTpHAa, ROTopble HeB03MomHO Obl.i:rO BbIHC-
"HIITb npemHHMll neHpOrlICTOJIOrlIqeCRIIMn MeTOAaMII, He 1I03BOJIHIO-
II{HMH All.p.pepeH:qIIpOBaHHO BblHBJlHTb aApeHeprHqeCRBe H XOJIIIHep-
I'HqeCRlIe Cl:lCTeMbI. IIpelIMYlI{eCTBeHHoe pa3BHTHe CIIMnaTHqeCRHX BO-
JIOROH Ha CTeHRax aopTbI Hap;O, OqeBlIAHO, paCCMaTplIBaTL B KaqeCTBC
HOBOll, OOJIee COBepmCHHOll cTyneHII B CTPYRTYPHOH opraHH3a:qllll
CHMnaTII'IeCROD: HepBHon CIICTeMbi. MLI paCI{eHIIBaeM 3TH HaOJIIOAe-
HHH RaK COOTBeTCTByIOIJ:J;He TeM Mop.pOJIOrlt'IecIGUI II3MeHeHIIHM, RO-
Topble Ha6JIIOAaIOTCH B 8BOJIIOI{HII cynpapeHaJIOBOn TRaHII. 8Ta no-
CJIeAHJlH, KaR Mbi rOBOpl1JIH, nORII]:(aeT B OCHOBHOii CBoeH Macce
apTepHaJIbHOe PYCJIO.
llyn, 110 ROTOPOMY BAeT lIepeMelI{eHHe cynpapeHaJIOBOH TRaHII,
nORa3bIBaeT nOCTcneHHYIO nOTepIO nOTpeOHOCTH mlIBOTHbiX B OblCTpOl\l
nOCTYllJIeHHH ropMOHOB B I{eHTpaJIbHblH RpOBOTOR. Y RpyrJIOpOTLIX
ropMOHbI BbiAeJIHlOTCH B nOJIOCTb npeAcepAHH H me.lIY]:(O'lROB II, no-
CTOHHHO HaCblII{aH RpOBb B I{eHTpaJIbHbIX MarIlCTpaJIbHblX COCYAax,
YTHJIII3IIPYIOTCH nepH.peplIqeCRBMH opraH aM II. CTeHRH cepAI{a ceJIa-
XIIH yme He HMeIOT B CBoeM COCTaBe xpOMa.p.pHHHOH TRaHII, TaR ilia
RaR II aopTa, HO coxpaHHlOT ee B npilCepAe'lHblX RpynHblx COCYAax
66
II Mempe6epHbIX apTepllHX. Y lWl'TlIO-XpIIlI eBbIX II lWCTIICTblX II ::ITa
JIORa.rrIlaaqIIH TepHeTCH, H xpOMa.p<pIIHllble ::I.TIeMeHTbI pacnO:IaraIOTCH
B CTeHRax MarHCTpaJIbHblX (RapAIIHaJIbHbIX, HpelllHblX) BeH II llX neT-
Beii.
- TaRIIM 06pa30M, xapal<Tepneillllee CBOllCTBO xpOMa.p<pIIHlIOll TW1-
nIl BCTYlIaTb B TeCHbIll IWHTal T C RpOBbIO ;mOJIIOupyeT. BUYTPII
Imacca pbl6 ::IBOJIIOqIIH BblpamaeTCH B nepeMe eHInl OT lIlarllCTpaJIb-
1101'0 lipOBOTORa Ii 60JIee MeJlKII!\1 apTepllHM n OT apTepIIaJIbHOrO
pYCJIa R neH03I10MY. OToABuraHCb OT qeHTpaJIbHOrO RponOTOI a, lIe-
60JIbillne rpynnbl xpOMa.p.pHHDbIX RJIeTOR «3aCTpeBaIOTI) B CTeHRax
nPYIIHbIX COCY]:(OB l'IJIII npHCTeHOqHOH 06JIaCTll. 3TH OCTaTO'IHhIe rpyn-
nbI, 1I0TepHBilllIe CBH3b C OCHOBHOil Maccoil lIepeMeCTHBllIeilCH TI\a-
IIH, OImablBaIOTCfI, RaR 6YAeT BJIAHO AaJIbllle, B COC,TOHHIIII .pYHKIWO-
HaJIbHOrO 6e3AellCTBIIH II npeBpa aIOTCH B PY]:(HMeliTbl. PY]:(IIMeaTap-
Hble OCTpOBRII, coxpaHHBillne He60JIbillOe IWJInqeCTBO ropMOHOB,
MomHO BCTpeTIITb yme y TaRHX HII3RO OprIJHH30BaHHbIX .pOPM, Ran
ROCTll('.TbIe pbI6bI. «Ra cMeay)} oTcTypalO eH OT apTepIlaJIbHoro PYC-
JIa xpOMa.p.pIIHHOH TRaall pa3BJmaeTCH CIIMnaTHqeCRaH HepnnaH eIl-
CTeMa, ROTopaH y pLI6 AOCTIIraeT 60JIee COBepllleHHbIX .poP!\{ H 'W-
CTIIqHO npn06peTaeT perYJIflpHLle pe.pJIeIiTOpnble CBfl3I1. Yme y pbI6
CJIeAOnaTeJIbHO, MOmeT 6blTb pe.pJIeRTOpHoe BbiAeJIeHne rOpMOHOB, IW-
TOpOe B AaJIbHellllleM Y BblCillIlX 1l03BOHO'IHbIX- 3aMeaHeT 60Jlee npH-
MIITIIBHblH cnoc06 cHa6meHlIJI OpraHII3Ma MOnOaMIIHaMII.
TaHoBbI xapaHTepHbIe 'IePTLI pa3BIITIIH CIIMnaTo-aApeHa,;:rOBOH CII-
CTeMbl phlG.
3*"
r.ttU6a III
3EMHOBO HbIE
B HacToHI ee BpeMH B JUlTepaType HaROIIHJUlCb pa3p03HeHHLle,
HO MnorOqHCJIeHHble cBe]:(eHuH 0 cTpYlnypHOH opraHH3a IIIl, rHCTO-
cPB3BOJIOrllil II rHCTOXHMIUI XpOMa.p.pBHHOH CIICTCMbl 3eMHoBoAllbIX.
HeCt.lOTpJl Ha pHA oc06eHHocTeii n aHaTOMHqeCROM pacnpeAeJIeHBII
XpOl\Ia.p.pHHHbIX Y3JIOB Y pa3HLIx OTPHAOn aM.pH6HH, MOilmo BhlAe-
JIHTb 06 He npB3HaRII, 0T.T:II1q81O Ue naparaHrJIHOHapHylO cllcTeMY
Bcero Imacca aM.pH6HH RaR CJIe]:(ylO ero IICTOpHqeCROrO ::lTana n
pa3BuTBII n03BOHOqHbIX. BblHBJIHIOTCH HOBLIe B3aHMOOTHOllleHHH C
cHMnaTHqeCROH HepBHOH CUCTeMOH, a rpaHH bI pacnpocTpaHeHHH
xpOMa.p.pHHHOll TRaHII BAOJIb MarHCTpaJIbHLlX COCYAOB TYJIOBH a npo-
AOJIiRalOT cymHBaTbcH. Hall60JIee RpynHbIe e CROnJIeHHH ROH eHTpIl'
PYIOTCH B 06JIaCTH COCY]:(HcToro annapaTa nOqeR - Ha HX BeHTp8JIb-
HOH nOBepxHocTil II Ha npHJIeil(8 IIX yqaCTRaX 6pIOlliHOH aopTbl II
1I0JIOH BeHbl. 3Aecb .popMllpyeTcH «.pH:IOreHeTHqeCRUii 3aqaTORI) 6y-
Ay ero HaAIIOqeqHBRa, KOTOpblll OROHqaTCJIbHO nLIAe.1JHeTCH B BHAe
BHRaIIcYJIlIpOBaHHoro opraHa yme Y neKoTopbIX OTpHAoB pellTHJIBii
(pIIC. 14). IIpeAcTaBJIHIOT 60JIbllloii HHTepec yqaCTRH xpoMaif».pIlHHoU
CHCTeMbl aM.pIl6I1H, paCIIOJIOmeHHble B 06JIacTII 6bIBllIBX ma6epHbix
COCYAOB.
- II3JIOiI\eHHe MaTeplIaJIOB 110 aM.pH6HHM MLI HaqHeM C OROJIOIIOqeq-
HbIX yqacTRoB xpoMa.p.pHHHoH TRaHH, HrpalO HX pOJIb HaAlIoqeqnn-
ROB, a 3aTeM lIepeHAeM R APyrilM ee OTAeJIaM.
IIepBoe ynoMHHaHlIe 0 HaAnOqeqHHlWBOM OpraHe 3eMHOBOAHbIX
BCTpeqaeTCH y Swallimerdalli (1738), OllilcaBmero em RaR (mplI-
AaTORI) ceMeHHHRa. AaJIbHeHIlIHe CBeAeHHH 0 HaAIIOqeqHIIRaX y pa3-
HblX OTPHAOB aM.pu6HH AaHbI B 0630pax H OpHrHlla.1JbHbIX pa60Tax
Ecker, Wiedersheim (1899), Vincent (1897c), Giacomini (1898a),
Brauer (1902), Hoffmann (1902), Grynfcltt (1930b, 1904a, b),
Biedl (1915), Dittus (1936), Hartman, Brownell (1949), Bach-
mann (1954), Goodrich (1958), Hanke, Weber (1965).
HapHAY C 06 BMH qepTaMH B CTpoeHHH aa]:(1I0qeqHOH TKaHII Y
Bcex OTpHAOB aM.pB6HH y RamAoro H3 HBX IIMeIOTCJI oc06eHHocTH,
CBOHCTBeHHbIe HaAIIOqeqHIIRaM TOJIbKO ]:(aHHoH rpYllnbl mnBOTHbIX,
Mop.pOJIOrBqeCRBe H3MeHeHHH, ROTopble na6JIIOAaIOTcH B RJIaCCe aM-
.pH6HH, OTpamalOT nOCJIeAOBaTeJIbHOCTb ;)BOJIIO 1I0HHbIX IIpe06pa30-
BaHHll B pHAe - 6e3HorHe (Gymnophiona), XBOCTaTbIe (Urodela)
Ii 6eCXBOCTbIe (Anura) 3eMHOBOAHbIe.
68
"Y 6e3HOrUX aMqHI6HH, ROTOpble HBJIHI OT CH H8H60JIee IIpHMHTIIB-
HbIMII II3 Hb!Jle 2lillBy 3eMHOBOMbIX, H86J11o.n;aeTCJI HaII60JIee
-«.n;peBHIIIl» Tlill aH8TOMUqOCKOro. paCIlpOCTpaHeHHH XpOMat}tq>HHHOll
TR8HH. llepe.n;HHH qaCTb Ha.n;IIOqeqHIIROB Y HHX IIpe.n;CTaBJIeHa .n;JIHII-
HbIM O.n;HOpO.n;HbIM TeJIOM, JIemaIII;HM Ha Me.n;BaJIbHOM Rpae noqeK,
a 3a.n;HHH COCTOHT H3 OT.n;eJIbHbIX 06pa30BaHHH (pHC. 15, A), ROTO-
pble MeTaMepHO pacnOJIOmeHbI Ha BeHTpaJIbHOii CTopOHe nOqeR IlJIH
BHe.n;pHIOTCfi B HX napeHxHMY (Brauer, 1902; Berkelbach, 1934;
Dittus, 1936). 0 npHMHTHBHOCTH CTPYRTYPHOH OpraHH3aI{HH xpo-
MaqJlpHHHoH CBCTeMbi 6e3Horllx aMqm6HH MomHO Cy.n;HTb no pJl.n;y
IIpH3HaROB, onHcaHHblX B OqeHb T:rn;8TeJIbHOM IICCJle.n;OBamIII Dittus
( 1936), Y o.n;Horo 1I3 IIpe.n;cTaBHTeJIeii 9Toro OTpH.n;a I{eHJIOHCROii qep-
BHrH. K qIICJIY T8RHX npH3HaKOB OTHOCHTCH B3aHMOOTHOllleHHH C
COCy.n;BCTOii, CBMnaTBqeCKOH II BHTeppeHaJIOBOH CHCTeMaMH, a Tal{me
cPopMa xpoMa.plfJlIHHblx RJIeTOR. Dittus YCTaHoBII.TI Ha cepHMHblX no-
Uepe'-lHbIX cpe3ax TeJIa mHBOTHbIX XapaR'lepHbIe KOHTaRTbi xpoMa.p-
cPlIHlIOii TKaHH C KPYUHblMII cocy.n;aMH TYJIOBB:rn;a, CBOHCTBenHbIe PbI-
6aM. P83.n;eJIlIB BCIO 3TY 06JIaCTb Ha HeCROJIbKO 30H (pHC. 15, A),
OH onucaJI rHe3.n;Hoe pacnpe.n;eJIeHBe lIaparaHrJIIIOHapHoH TRaHH B.n;OJIL
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1 - cynpapeHaJIOB8H THaHb; 2 - HHTeppeH8J10Ball THaHb; a - nO'IHH (Gorbman, Bern,
t 962)
nmllqbl
69
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aopTbl, a TaRme B IIepIlTOHeaJIbHOii
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B). BMecTe C cocy aMII naparaIIr-
JIIIOHapabIe RJIeTRII IIpOHIIRaIOT C
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MeHTL1. H x rHe3]:(a, lIJIII OT]:(eJILIJbIe
ImeTH:II, IIHor a paCIIOJIOmeHbI B
CTeHH:aX aopThl II aa OTXO HII{IIX OT If(e/l'lHbIU {
Hee BeTBHX, B HlimaeH I10JIOH Bene nf/3bf/lb
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CTBOJIOB He MomeT OOeCIIelIlITb MeTa- rOHOo.
MepHYIO CTPYRTYPY.
K qHCJIY IIpH3naHOB, xapaRTepH-
3YIOI:QHX cpanHIITeJIbHO neBblCOH:HH
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TH:anH Y 6e3norHX aMWUOUll:, CJIe]:(y-
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Y qepBHrll RpYIInbie xpOMaWWlInHble
RJIeTH:1I C H:pyrJIblM IIJIII OBaJIbHbIM
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(Ichthyophis glutinosus)
HagnO'le'lHIIHH - 'lepHhIe;
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cpeaoB pilC. B;
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B - nonepe'lHhle CpeSH H8JtIIO'le'IIDIIROB Ha paaHblX YPOBHHX cxeMbi A:
-cynpapeHaJIOB8H THaHb - '1epHaH; HHTeppeHIIJIOBaH - nyHKTlIp; VN -
nepeAHH'I nO'lC'lHaH Beaa; Vc - nOJIaH Bella; D - HHme'lHHK; Le-
ne'lflHb; Ly - JIHM4IaTH'leCKOe npOCTj)aHCTBO; L3 - JIHM4IaTII'lecHHt1: CII_
.BYC; un - BOJIb4l0BO TeJ10; g - I'OH8AbI; Ao - aopTa (Dittus. 1936)
JIll o6JIaAalOT OqeHb cJIa6oii: QBeTHoii: peaRQJlcii: BemplilHa IIX Ht{ep
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IIpeAeJIeHHaJl IIO RJIeTRe. Ha6JIIOAalOTCH O'-laru naparaHr.'lIlOHapHOll
TRaHII II BHe BCHRoii: CBJl31I C IIHTeppeHaJIOBOii, qTO Tome HBJIHeTCH
IIpHMIITIIBHbIM IIORa3aTeJIeM, HalI60JIee OTQeTJIlIBO Bblpail\eHllbE\1 Y ce-
JIaxlIii:.
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paCIIOJIOmeHlIe xpoMaclJclJlIHHoii: TRaHII Ha6JIIOAaeTCH npeIll\IYlQeCTBeUHO
B XBOCTOBbiX OTAeJIax IIO'leK.
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AJllQeH B COCTaB Ha]:(IIO'le'lHbIX meJIe3, IIpOAOJImaeT CYJImBaTbCH B
HaIIpaBJIeHlI1I IIO'leK II IIplIJIemalQIIX Y'laCTKOB. HaAIIO'leIIHIIKII JIemaT
BAOJIb cpeAHeH JIIIHIIII TYJIOBIIIQa II IIplIMbiKalOT K BeHTpO-l\1et{IIa:Ib-
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HOii: TRaHeBoii: MaCCOH, a OTAeJIbHblMII OCTpOBRaMII, pacceHHHblMII B
ABa pHAa MemAY IIOqRaMII II CTBOJIOM HnmHeH IIOJIOH BCHbI, a TaRme
BAOJIb BeTBeii: IIO'le'lHbIX apTeplIii: II BeH (Gaupp, 1899; Hoffmann,
1902; Hartman, Brownell, 1949). Y 6eCXBOCTbiX napHble TeJIa Hat{-
IIO'le'lHIIROB IIpeACTaBJIHIOT co6oii: CnJIOIlIHble nJIaCTIIHRII HepaBHmlep-
HOii: TOJIlQlIHbl (CM. plIC. 14).
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BIITeJIeii: aMclJu61Iii: HBJIHeTCJI IIX CBH3b C OTBOAHIQUMU 1I0'IC'lHbE\1II
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HIIKaMII II BMeCTe C llX BeTBHMII BHeAPHIOTCH B IIOqeIIHYIO napeH-
XIIMY (Biedl, 1915; Bachmann, 1954; Goodrich, 1958; C!\lIITTeH,
1963; Hanke, Weber, 1965).
TaKIIM o6pa30M, Y aMclJII6IIH HaAIIO'leqHllKII Ha'lIIHaIOT OclJOpM-
JIJlTbCH B Tex MeCTax CIIIIHHOii: 06JIaCTlI Ty.1J:OBlIlQa, B KOTOpblX OHU
pa3BlIBalOTCH Y Bcex BblCllinx II03BOH01ffiblX mIIBOTHbiX. Ho Y HIIX
OHII He JlBJIHIOTCJI elQe caMOCTOHTeJIbHblMII opraHaMII (B TOM CMbIC.1J:e.
KaK 3TO IIplIHHTO C'lIITaTb Y BblCllIlIX II03BOHOIIHbIX), a HaxoAHTCfl
B «3aBIICHMOM» IIOJIOmeHll1I OT lIoqeK. OHII 3aKJIIO'leHbl BMeCTe C
HIIMII B 06III;YIO coeAHHIITeJIbHOTKaHHYIO RalICYJIY (pIIC. 17), He mlC-
lOT eIII;e caMOCTOHTeJIbHoii: CllCTeMbl opraHHoro RpOBOTOKa (Bach-
mann, 1954), CBJl3aHbi co CTeHKaMII IIO'leqHbIX BeH (qaCTIlqHO II
apTepllii:) n IIMelOT C HlIMU o6IQee KpoBoo6paIII;eHlIe. Oco6eHHocTb
3Toro KpoBoo6paIII;eHIIH 3aCJIymlIBaeT ClIeIwaJIbHOrO YIIOMIIH:llfHH, TaR
KaK B03HllKaeT B 3BOJIIOQllOHHOM pa3BlITIIII Y aMclJll6uii BlIepBblC.
B TOM MeCTe, r]:(e OTBOAHIII;ne BeHbl nOqeR BnaAalOT R nOJlYIO BeHY,
naAIIOIIeIIHIIRII ROJIbQeo6pa3HO IIX oRpymalOT II nOHpblBalOT BeHTpaJlb-
IlYIO IIOBepXHOCTb V. cava. OTBOAHIII;lIe IIOqeIIUbie BeHbl OT;J;aIOT R
HaAIIO'leqHIIRY BeTBll, ROTOpbIe BHOBb pacnaAalOTCH Ha RalIu."IJIHpbl,
BlIaAalOIII;ue 3aTeM B IIOJIYIO BeHY. ::ha nOpTaJIbHoro Tllna ClICTeM3
RpOBoo6paIII;eHIIH, RlIepBble Ha6JIIOAaeMaJl Y CaJIaMaHAPbl eIII;e B IIpOIlI-
JIOM CTOJIeTIIH Ecker (1846), BIIOCJIeACTBllH MHoroRpaTHo OIIHCblBa-
:raCb (Hoffmann, 1902; Grynfelu., 1904a, b; Bachmann, 1954).
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ep;IIHeHbl p;pyl' C p;pyroM y Anura, qeM y Urodela, qTO BIIOJIHe
IIOHJlTHO C TOqKII 3peHIIH 60JIee BblCOKoro 3BOJIIO IIOHHoro pa3BIITIIH,
KOToporo p;OCTIIraIOT Anura. "Y 1I0CJIep;HnX yme HaqHHalOT C03p;a-
BaTbCH CTpyKTypHble B3aHMOOTHOllieHIIH P;BYX Hap;IIO'leqHIIROBbIX CIl-
CTeM, IIp1I6JIIImalOIII;He IIX K BblcmlIM 1I03BOHOqHbIM.
B 50-60-e rOAbl 1I0HBIIJIIICb pa60Tbl 110 BbIHCHeHHIO BOlIpoca
o MOHD- IIJIII P;IIMop.pII3Me xpoMa.p.pIIHHbIX 3JIeMeHTOB y npep;CTa-
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felt (1955) II B y.p-JIY'Iax IIO Erank6 (1955a) . ROHTpOJIeM CJIY-
mIDI Hap;IIO'leqHIIK Mblmell. B 60JIbmell qaCTH KJIeTOK 1I0CJIe IIpOBe-
p;eHIIH IIOp;aTHOro MeTop;a 06HapymeH oca]:(OK TeMHO-KOpIIqHeBOrO
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8.Jij>eHaJIIIH. Pe3YJIbTaTbl 06pa60TRII B y.p-JIyqax BbIHBIIJIII y TPII-
TOHa II CaJIaMaHp;pbl OqeHb He3Ha'lIITeJIbHYIO pa3HII y B OKpacKe KJIe-
TOK IIO cpaBHeHIIIO C KOHTpOJIeM (Ha Mblmax), HO, 1I0CKOJIbRY ClIeKT-
paJIbHaH xapaKTepnCTHKa Hopap;peHaJIIIJra y MbIlliell II aM.pH6HH ol\a-
3aJIaCb IIp;eHTIIqHOll, TO aBTopbl 1I0JIaralOT, qTO .p.rryopee eHII;1IJI KJIe-
TOK II Y aM.p1I6IIll 06YCJIOBJIeHa IIPIICYTCTBlIeM HeMeTIIJIlIpOBaHHoro
ropMOHa.
TaKIIM 06pa30M, aBTOp IIplI3:HaeT P;IIMOp.pH31I KJIeTOqHbIX CTPYRTYP
y aM.pH6IIll II OTp;aeT IIpep;IIOqTeHIIe B KOJIIIqeCTBeHHOM OTHOille-
HIIII Hopap;peHaJIIIHCOp;epmaIII;IIM KJIeTKaM. AHaJIOrlI'IHOrO MHe-
HIIJI IIpIIp;epmIIBaIOTCJI aBTopbl 3JIeKTpOHHOMIIKpOCROIIIIqeCKIIX IICCJIe-
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aJIbp;er.lI)J;e C IIoCJIep;YIOIII;ell 06pa60TKo:ii COJIblO cepe6pa, 06HapymlIJI
ocap;OK, KOTOpblii: clIe .pII'IeB TOJIbKO P;JIH rpaHYJI Hopap;peHaJIIIHa
II He BbllIap;aeT B rpanyJIax ap;peHaJIIIHa. Piezzi IIOJIY'IIIJI. KapTIIHbl,
xapaKTepII3YIOIII;IIe IIPIICYTcTBlIe ap;peHaJIHH- II Bopap;peHaJIHHCOp;ep-
maIII;IIX KJIeTOK.
HapHp;y C YKa3aHHbIMII pa60TaMII, B KOTOpblX aBTopbl IIOKa3aJIII
IIPIICYTcTBlIe y aM.pII6IIll P;BYX TIIIIOB KJIeTOK, IIMelOTCH II p;pyrne
IICCJIep;OBaHIIJI, p;OCTaTO'IHO 06ocHoBaHHble. qmoy QII-p;a, q3Hb MIIHb-
XY3ll (1 65) IIpOBeJIII nop;po6Hblll IITOXIIMII'IeCKIIii: aHaJIII3 Hap;IIO>-
'm'lHoii: 06JIaCTII IIO'IeI{ y ;JIHrYllieK (Rana nigromaculata). OBII
74
()npC;\C:Ifl.'lll, IIOMUMO cne IUIJlIqeclwll peaKl\III1 Ha RaTeXOJIaMIIHbl,
paClIpep;eJIeHlle B hJleTl,aX PHK, rJIllhOreHa, aClwp611HOBOll RIICJIa-
TbI, XOJIeCTepIlHa, !II1IIIIP;OB, eJIO'IHoii: cPoccl>aTa3b1 II IIplImJIH R BbI-
BO]:(Y, qTO B HajJ.IIOqe'lIumax JIHrYllieR IIMeeTCH OP;IIH TIIII xpoMacl>-
cPlIHHblX R.TleTOR.
llplI'IIIny pa3HOpe'II1I10CTII 1I0JIY'IenHbIX pe3yJIbTaTOB MomeT 6b1TL
GJIep;yeT IIChaTb R ce3011HblX ROJIe6aIIIIHx. TaRoe IIpep;1I0JIOmeHlIe B03-
Bln,aeT lIa OCHonaHlIll pa60Tbi B. 11. COROJIOBa ( 1968), Bblme]:(meii:
H3 .TIa60paTopll1l 3. C. l\a HeJIbCOIH\. AllTOp P;JIH BblHCHeHIIH BOlIpoca
o c ecTBoBalIIIII MOHO- lIJIII P;1IlIWPcPII3Ma XpOMacPcl>IIHHbIX RJIeTOR
aMcPlI6I111 (OH pa60TaJI Ha Rana temporaria) IICIIOJIb30BaJI qeTblpe
l,puTepIlH ;J,!IH IIX OlIpep;eJIeHIIH - rIlCTOXIIMllqeCRlIii:, TIIHRTOpllaJIb-
Hblii, MOP(POJIOflPlecRlIii: II TonorpacPlIqecmIii:. HII TOlIorpacPH'IeCRIIC,
HIl MOP(pO,TIOrIl'IeCInle pa3JIII'IIUl He )".\aJIII Hap;emHbIX RplITeplleB P;JIH
pa:lrpaHIPleHIIH p;nyx I,.'leTO'IHbIX TIInOB y JI1IrYllieR. He 1I0MOrJIII B
::!TOl\f OTHOllleHlIlI II fliCTOXIIMII'IeCRIIe peaRQIIII Hillarp a. H6kfelt.
Do TIIHI,TOpllaJIbHblM clloii:cTBaM a]:(peUaJIIITI- II Hopap;peHaJIIIHCOp;ep-
;na I1e WIeTRII OTJIII'Ia.'lUCb ]:(pyr OT p;pyra TOJIbRO B 3I1MHIlii: lIe-
pIIOP;, a B JIeTHlIii: 6b1JIlI OI-:pameHbl B npoMemYTOqHbIe TOHa. BJIIIH-
nile ce30HHOCTII Ha cBollCTBa xpoMacl>qHIHHblx hJIeTOR CRa3aJIOCb II
Ha pa3Mepax H;:\ep, ROTOpLIe, IIO p;aHHbIM B. 11. COROJIOBa, IIMeIOT
Op:ImaI;OBYIO IJe.'III'InHY, HO OU:Iap;aIOT CTaTIICTlIqeCRlI ]:(OCTOBepHbIMU
('eaOHIIbHlll I';OJIeUaIIIIHMII.
HeROTOpble YRa3aHIIH Ha ce30HHbIe 1I3MeHeHIIH B xpOMacPcPlIHHOll
7haHII aMcl>II61111 IIMeIOTCH II B GOJIee paHHell pa60Te Lakshman
(1964), hOTOpbIii:, HCCJIep;YH rJIaBHbIM 06pa301\1 IIHTeppeIIaJIOBYro
7naHb y pH]:(a BIlP;OB (Rana tigl"ina, Rana hexadactyla, Hana cy-
anophlyctis, Bufo melanosticutus, Upmodon II Kaloula sp.), OT-
MeTIIJI, 'ITO OnpaCRa xpoMacPcl>IIIIJIOH n,aHII OpaHIi{eBbIM JJ{ oCJIa6e-
BaeT BO BpCMH CIIH'II>II.
B Harnell pa60Te, IIpone;J,elIHoii n 1962-1963 ro]:(ax C QeJIbIO
BblHCHeHIUl ap;peHaJIIIH- II Hopap;peHamlHcop;ern,a lIX RJIeTOl\ II CpO-
hOB IIX 1I0HBJIeHIIH B pa3BHTlIII rOJIOBaCTHl\OR (CMIITTeH, 1963, 1964),
Mbl nbITaJIIICb IIpOCJIep;IITb II ce30IIHbIe mmLI 3HP;ORpIlHHOH .pYHRQIIH
xpoMacl>cPlIHHoH TRaHH y B3pOC:IbIX JIHrYlllel{ (Rana viridis) BO Bpe-
MH CIIH'IRII 31I1\1011 II 60]:(pcTBoBaHIIH JIeTOM. 113-3a Hep;OCTaTOqHO qeT-
RlIX p;aHHbIX Mbl B03p;epmllBaeMcH OT lIy6mmalWll 3Toro MaTepllaJIa.
lITO me RacaeTCH IIPIICYTCTBlIH P;BYX RJleTOqHbIX TIIIIOB, pearllpyIO-
I1X C IIOp;aTOM RaJIIIH II CO!lHMII xpOMa (no Hillarp, H6kfelt)
B lIapaaOpTaJIbHbIX Y3JIaX II B Y'IaCTRaX, npllJIeraIO lIX Y rOJIOBa-
CTIIROB R Me30HecPPOCY, TO TaM nOJIY'IeHa 'IeTRaH HapTIIHa, 1I0Ra-
3blBaIO aJl IIPIICYTCTBlIe RaR Tex, TaR II p;pyrllx RJIeTOR, 6eCIIopJl-
P;OqHO pacnpep;CJIeHIIbIX cpe]:(11 IIIITCppeHaJIbHblX (pIIC. 22B, 82B).
llOC.TJep;HIIii: BOIIpOC, ROTOpblH 3aCJIymlIBaeT BHIIMaHIIH C ToqRII
3peHIIJI cpaBHIITeJIbHOH MOPcPOJIOrllll,- 3TO IIHHepBa IIH Hap;IIO'Ie'I-
HIIROB aMcl>1I6I1H. B 3RCIleplIMeHTaJIbHOll pa60Te Inoue (1958) II3yqe-
no BJIIIHHlIe uepepe3RII BeHTpaJIbHbIX II p;Op3aJIbHb1X RopelliROB Ha
pa3HbiX ypOBlliIX CIIIIHHoro M03ra ma6 Ha YMeHbmeHlle cop;epmaHHH
75
ap;peHaJIHHa B Ha)J;IIOqeqHHRaX, Bbl3BaHHOe IIHCYJIIIHOBOll rUIIOrJIIIhe-
MHeii:. 8cl>cl>eRT OT IIHCYJIIIHOBOii: rIIIIOrJIHI{eMIIII (ROTOpaH BbI3blBaeT
IICTOII eHHe ap;peHa.Uma B Hap;II01Ie1JHHRaX) IIOJIHOCTblO CHIma.'ICH ne-
pepe3Roii: 3-7 IIap neHTpaJIbHblX RopelllIWB. ABTOpbl ]:(eJIalOT BbIBO)J;,
qTO ceRpeTopHMe BOJIORHa R Hap;IIOqeqHHRY IIPOXOP;HT Y n,a6 qepe3
Te me ROpCIlIRII, 'Iepe3 ROTopble OHIl IIP;YT Y MJIeROrrIITalO lIx.
HaM IIpep;CTaBJIHe'fCH IUlTepecHblM oTMeTIITb, qTO H3]:(IIOqeqHIIRII
aMcl>1I6IIn, He 6YP;yqlI e e nO HOCTblO Ocl><'pMJICHbl RaR CaI\IOCTOH-
TeJIbHbIe opraHbl (3aRJIIOQeHbl B OP;HY RanCYJIY C nO'lRaMII II CBJl3a-
Hbl C HHMII 06 IIM RpOBoo6pa eHlIeM), OUJIap;aIOT, op;HaRO, IIHHep-
Baqlleii:, CTPYRTypHaH opraHII3aqHJI ROTOpOll nOqTII COBna aeT C Ta-
ROBon Y MJIeROIIHTalO IIX. Y IIOCJIep;HIIX CeI{peTOpHbIe Bo;rIOlma npo-
XOP;JlT qepe3 nepe]:(HIIe RopemRII 4-9-ii: rrapbl (IIJIbima. 19 9).
TaR 06CTOIIT p;eJIo Y 3eMHoBop;HbIX C xpoMacl>cl>IIHHoll TRaHblO
OROJIOnOqeqHOn 06JIaCTII, ROTopaH cooTBeTcTByeT I\leCTonO.IOmeIIlllO
Hap;IIOqeqHHROB Y BbllllecToH 1IX cl>OpM n03BOHO'lIIbIX.
)J:pyrlIe yqaCTRII xpoMacl>cl>IIHHon CIICTeMbl aMcl>1I6I111 C n03HqIlII
cl>IIJIOreHe3a IIpep;CTaBJIHIOT IIHTepec CBOHMII B3aIlMOOTHOllieHIIHMII C
cIIMnaTIIqeCRoii: HepBHon clIcTeMon II C RpoBeHocHblMII cocy]:(aMll.
Y npep;cTaBIITeJIen pa3Hbix OTpHP;OB 3eMHoBop;HbIX - pa3HaJl CTe-
neHb MeTaMeplI1I xpoMacl>cl>IIHHoii: TRaHII. J\aR 6bI.TIO nORa3aHO Bblllle.
y 6e3HOrHX aMcl>II61In (Ichthyophis glutinosl1s L.) cerMeHTapHoe pac-
IIOJIOmeHIIe naparaHrJIIIOHapHblx OCTpOBROIJ B 3aqaTROBOM IIepuo-
p;e pa3BIITIIH p;OCTaTOqHO OTQeTJIIIBO coxpallHeTCJI RaR y MOJIOP;bIX mII-
BOTHbIX, TaR II y B3pOCJIbiX ocoueii:. Y XBOCTaTbIX, ROTOpbIe CTOHT
cTyneHblO Bblllle 6e3HorIIX, cerMeHTapHoe pacnOJIOmeHlIe IIaparallr-
:IIIOHapHoii: TRaHII npaRTIIQeCRII OTCYTCTByeT, B TO BpeMJI RaR y 6ec-
XBOCTblX OHO BHOBb B03HIIRaeT. Y Bachmann (1954) Bbl3bIBaeT He-
)J;oyMeHIIe 60JIblllOe pa3JIHQUe, cy eCTBYIO ee B 3TOM OTHOllleHllII
Me>R)J;y XBOCTaTblMII II 6e3HorHMII 3eMHOBop;HbIMII, TeM oo.:):eo CTpaH-
Hoe, QTO, 6e3 COMHeHIIH, 60JIee BbICOROCTOH I1e 6ecxBocTbie BHonb
o6HapymIIBalOT MeTaMepIIIO, XOTH II B 3aQaTOQHOM COCTOHHIIII.
C Hallien TOQRII 3peHIIH, R 06'bHCHeHIlIO TaRoro, B 113BeCTIIOll
Mepe IIapap;ORCaJIbHOrO cl>aRTa, MomHO nop;ollTII, C OP;HOll CTOpOHbI,
C Tex me n03IIqIIll MoPcl>ocl>YHRquoHaJIbHbIX B3alIMOOTHOllieRIIU xpo-
Macl>cl>IIHHon CHCTeMbl C RponOTOROM, a C ]:(pyroii - C n03lIqnu, y'lII-
TblBalO IIX B03paCTalO YIO CTPYI{TYPHYIO OpraHIl3aqlI1O CIIMIIaTJPIe-
CRon HepBHon CIICTeMbl B PHP;y 3eMHOBO)J;RbIX, reHeTIIQeCRH cnH-
3aHHYlO C xpoMacl>cl>IIHHon.
1I3 cl>aRTlf1IeCRUX CBep;eHuii:, ROTopble IJblTeRalOT 1I3 IICC.'Ie]IOBaIIIIH
Dittus (1936), CJIep;yeT, qTO me3]:(a xpoMacl>cl>IIHHbIX R.'leTOR y
enJIOHcRoii: QepBHrII IIMelOTCH OROJIO aopTbl H B OTXOP;H IIX OT lIee
apTepIIHX II QTO OHM nCTpeqalOTCH Rpanlle pe RO B CIIl\flIaTlIQeCI\1IX
Y3JIaX norpaHIIQHOrO CTBOJIa. 1I3 Clm3aHHOrO Bblllle OqeBII HO, 'ITO
y 6e3HorIIX (BO BCHROM CJIyqae, y qenJIOHCROii qepBHrlI) MeTaMe-
plIH CBH3aHa He C CIiMnaTIJ1IeCROll, a C cerMeHTapHOll apTepIIa;lb-
HOii: CIICTeMOIl. 8TO paccym eHIIc MomHO rrpOp;OJImIITb II p;a.'Ibllle.
Y Urodela, y ROTOpblX OTCYTCTByeT MeTaMeplIJl IIaparaHmIIeB, O;J;HO-
76
BpeMeIIHO TepJleTCH II CBJl3b C apTepIIO:IaMII, 0 '!eM MOmHO 3aRJlIO-
'IIITb 1:13 OIIIICaHIIll 3TOro OTpHp;a aMcl>II6IIii: B pa60Tax Hartman,
Brownell (1949) II Bachmann (1954). Ho IIO'IeMY MeT3Mepnoe
paCIIOJlOmeHl:le BHOBb B03HIII<aeT Y Anura? B IIOIICRaX OTBeTa Ha
3TOT BOlIpOC Mbl 06paTIIJlIICb R CTpoeHIIIO CIIMllaTII'IeCRoii: HepBHoD
CIiCTeMbl, cBoii:cTBeHHoMY TOMy II P;PyrOMY OTPHP;Y aMcl>lIollii. KJlac-
ClI'IeCRlle IICCJlep;OBaHIJJI Giacomini (1898a), PyIWBOp;CTBO Ecker,
Wiedersheim (1899) II IIOCJle]:(YIO IIe pa60TbI Werner (1930), Hirt
(1934), Singer (1942), Nicol (1952) p;alOT ]:(OCTaTO'IHO 1I0JIHOe
IIpep;CTaBJleHUe 0 CTPYRTYPHOll opraHll3aQIIll CIIMlIaTllqeCROll nepBHon:
CllCTeMbl Y 060lIx OTpHp;OB.
Y Urodela napHaH CIIMnaTlI'IeCRaH Qe1I01fl<a JlemlIT no 60RaM
p;Op3aJlbHOii aopTbl. B ee COCTaB BXOP;HT THmII CIIMnaTU'IeCRIIX Bo-
,TIOROH, ROTOpI,Ie cop;epmaT pacceHHHI,le raHrJlII03HbIe RJleTl<II, TO:lbRO
t.IeCTaMII 06'bep;IIHeHHbIe B raHrJllI1l II IIpep;CTaBJlHIO lIe c060ii He-
ocl>opMJleHHble IIJleRcyco06pa3HbIe CTPYRTypbl. llOMIIMO 3TlIX OClIOB-
HblX n;eIIO'Ie'IHbIX TJliReii Y HelwTopbIX XBOCTaTblX aMcl>u6uii: (Nec-
turus) IIMelOTCH ROJlJlaTepaJlbHble TJlmU HepBHblX BOJlOROH, JleiRa IIe
IIO 6m.aM OT OCHOBHI,IX U CBH3aHHbie C HIIMU TOHRIIMU HepBHI,IMII
nOJlORHaMIi. B COCTaBe ROJlJlaTepaJibOLIX TJlmeii: IIMelOTCH c06cTneo-
Hble MeJlRUe raHrJlIIII, ROTOpbIe coep;IIHeHbl rami communicantes co
ClIIIHaJibOblMII lIepBaMU (Hirt, 1934; Nicol, 1952). Singer (1942),
1I0]:(p0600 06CJlep;OBaBIlIIIii: CIiMlIaTII'IeCRYIO CIiCTeMY llIeiiHo-rpYP;HoH
06J1aCTII TpIITOHa, OIIIICaJi 1I0p;RJlIO'III'IHOe CIIJleTeHlIe II paClIpOCTpa-
HeHIIe ero BP;OJlb 1I0P;RJlIO'III'IHbIX II 1I03BOHO'lHbIX apTepliH. OT 1I0P;-
R.'IlOqllqHOro CIIJleTeHIIH OTXOP;HT ]:(Ba HepBa R cepP;QY, a BP;OJlb 110-
3BOOO'IHbIX apTepuH OepBObIe CTBOJIllRII II]:(YT BlIepep; B llIeHHYIO 06-
.'IaCTb II oa3ap; B a6]:(oMIIOaJlboylO qaCTb CIIJIeTeHIIH, rp;e 1I0CTraHr;r1I0-
HapHI,Ie BeTBII ux paClIpOCTpaHHIOTCJI oa cerMeHTapHbIe apTepIIII.
Singer, 1I3yqaH XOP; cnllIHWbHbIX HepBoB, IIX CBH3II C RopeIIlRallHI II
COCTaB rami communicantes, IIpIlXOP;IIT R BbIBOP;Y, 'ITO MHOiReCTBO
1I0CTraOrJlIIOHapHbix CIIMlIaTUqeCRIIX BOJlOROH p;OCTIUaeT II IIHoepBII-
PyeT IIplIJlema lle R aopTe OTpC3RU RpoBeHocoblX COCY;:J;OB, HO OT-
CYTCTnyeT n CMelllaHllblX ClIlIlIaJlbHblX HepBax. Nicol COrJlalliaeTCJI
C Singer OTHOCIITe,TlbHO IIHHepBaQl1U nOCnaHrJlIIOHapaMU 6J1If3J1ema-
IIJ;IIX apTepllaJlbHblX COCYP;OB, 00 p;06aBJIHeT, qTO IIMelOIII;IIeCJI CBe-
]:(eHIIH p;aJIeRO ne p;ocTaTo'lHI,I, qT06b1 Moamo 6I,lJI0 OlIpe]:(eJleHHO CY-
P;IITb 0 OalIpaB,1IeHHII II paclIpocTpalleBlIII npe- II 1I0CTraOrJlIIOHapHbiX
BO:IOROH B BncQepa:lbHblX cocyp;ax II BO ImYTpeoHnx OpraHaX. lloJI-
HOro IIpep;CTaBJIeOIIH 0 CTelIeOII n XapaI{Tepe cHa6iReHIIH 1I0Hepna-
QnOHIII,IMII IIpn60pa1.I11 BHYTpeHHIIX OpraHOB XBOCTaTbIX aMtPII611ii MI,l
lie IlMeCM ]:(0 CIIX nop. Op;HaRO P;JIH IIpOBOp;IIl'.lOro Hal\1lI aHaJlll3a
]:(3HHI,Ie Hirt, Singer II Nicol p;OCTaTO'IHbl, qT06b1 c]:(eJIaTI, onpe-
p;eJleUIlLIe BbIBOP;bl.
Ha OCOOBaHIIII IICCJIe]:(oBaoull :JTIIX aBTopOB, MI,l 1I0;:J;qepRUBaeM
p;na 06CTOHTeJII,CTBa, ROTOpbIe HaM l\amYTCH BamObIMII. llepBoe-
:JTO TO, 'ITO Y XBOCTaTbIX aMcl>n6nii I1MCIOTCH nOrpaHU'IIII.fe CIIMIIa-
TIIlIeCRlle CTBOJlbl, B ROTOphlX OTCYTCTBYIOT CTpOro O'Iep'IeIIIIWe CIIM-
77-
naTII1.J.eCRIIe Y3JIbi II CerMeHTapHOe IIX paCnpep;eJIeHIIe BP;OJIb P;JIIIH-
HOii: OCII TYJIOBIIIIJ;a. BTOpoe OTHOCIITCH R IIHHepBa:I IIII IIpIIJIemaIIJ;IIX
Ii aOpTe apTeplIaJIbHbIX CTBOJIOB, 0 cHa6}I{8HlIH ROTOpblX IIOCTraHrJIlIO-
HapaMII C YBepeHHOCTbIO rOBOpHT aBTOpbI.
Y Anura Mopcl>OJIOrIIqeCRaH opraHIIaalWH CIiMlIaTII1.J.eCRoii: HepB-
HOii: CIICTeMbl, ee paclIpocTpaHeHlIe II CTpYRTypa lIepIIcl>eplI1.J.eCRIIX
HepBHblX IIplI6opOB paapa60TaHbi 3HaqIITeJIbHO IIOJIHee. O IeHb 60JIb-
llIoe BHIIMaHlie Mopcl>OJIOrIlH aBToHoMHoii: HepBHoii CIlCTeMbl JIHrYIliRII
yp;eJIHeTCH B IICCJIep;OBaHIIHX Giacomini (1898a), Ecker, \Vieders-
heim (1899) II Gaupp (1899), a TaRme B pa60Tax Langley (1910),
Langley, Orbeli (1910-1911), Werner (1930), Hirt (1934), Nicol
(1952), Goodrich (1958). Ha OCHOBaHIIII 3TIIX pa60T MomHO CRa-
3aTb, 1.J.TO Y Anura aBTOHOMHaH HepBHaH CIlCTeMa OpraHII30BaHa IIO
TIIny HepBHoii: CIICTeMbl MJIeROIIIITaIOIIJ;I1X. MHorIIe aBTopbI, onIICbIBaH
ee CTPYRTYPY, p;eJIHT CIIMlIaTII1.J.eCHYIO HepBlIYIO CIICTeMY Ha PM OT-
p;eJIOB: 1) llIeiiHo-ma6epHblii:, cop;epmaIIJ;IIH TPII CIiMlIaTlIqeCRIIX raHr-
JIIIH, COOTBeTCTBYIOIIJ;IIX 2-4 CIIIIHaJIbHbIM HepBaM; 2) a6p;OMIiHaJIb-
HbIll, IIMeIOIIJ;nii: 1.J.eTblpe CIIMlIaTII1.J.eCRIIX raHrJIIIH, COOTBeTCTBYIOIIJ;nx
5-8 ClIIlHaJIbHbJM nepBaM, II 3) CaRpaJIbHO-ROII1.J.IIROBblii: C P;BYMH
CIIMlIaTII'IeCRIIMIi raHrJIIIHMII, JWTOpbIe COOTBeTCTBYIOT 9-10 cnll-
HaJIbHbIM HepBaM. B BIICI,epaJIbHbIX opraHax II COCYP;IICTbIX CTeHRax
IIMeeTCH 06IIJIbHaH IIHHepBaqllH IIOCTraHrJIROHapHblMIi BOJIORHaMIi.
Xo]:( BIIcqepaJIbHblX BOJIOROH 60JIee p;eTaJIbHO pa3pa6oTaH Y JIHrYIliRII,
1.J.eM Y ma6bl (Nicol, 1952).
06 YCJIOmHeHHoH CTPYRTYPHOll opraHII3aqIHI CIiMnaTII1.J.eCRoii:
HepBIIOH CIIeTPMbI n COCYP;IICTbIX CTeHRax Anura, no cpaBHeHllIO C
HIImeCTOHIIJ;IIMIi cl>opMaMII CBII]:(eTeJIbCTBYIOT Ha6.TlIO]:(eHIIH r. P. JIe-
OHTbeBoii: (1966a, 1968), a TaI{meMaTepIIaJIbl H. B.BOP;POBOii: (1957),
C1.J.IITaIOIIJ;eii, 1.J.TO CIIMlIaTII'IeCRaH IIHHepBaqllH cepp;IW BnepBbIe Ha-
1.J.IIHaeT Ocl>OpMJIHTbCH Y aMcl>II6nii:. 8TOMY COOTBeTCTBYIOT rIICTOXII-
MlIqeCRHe p;aHHble Falck II coaBTopOB (1963), ROTopble IIORa3aJIII B
COCTaBe lIeperOpop;RII npeJJ:cepAIIii: cl>JIyopecqeHqnIO HepBHbIX BOJIOROH,
a Ha aap;Heii: CTeHRe aTpIIoBeHTpIlRYJIHpHoii: 06JIaCTII cepp;qa ap;pe-
HaJIIiHOBYIO cl>.lIyopecqeHqIIIO HepBHblX RJIeTOR, aRCOHbl ROTOpblX pac-
IIpOCTpaHHIOTCH B meJIyp;OqeR. IIop;TBepmp;eHlIe CIIMnaTII1.J.eCROH
IIpIIpOP;bl 3TIIX RJIeTOR, IIOMnMO cTporoH clIeqIIcl>H'lHocTn caMoro
cl>JIyopecqeHTHOro MeTop;a, aBTopbl BllJJ;HT B pe3YJIbTaTe pe3eplIlI-
HII3aqIIII JIHrymeR, IIOCJIe ROTOpOH CBeqeHlIe IICqe3aeT. Ha OCHOBa-
HIIII CIIeRTpocl>JIyopOMeTplI1.J.eCROrO IICCJIep;OBaHIIH, IIORa3aBmero IIplI-
CYTCTBlIe ap;peHamma B IIpep;CepP;IIHX B ROJIlf'IeCTBe 1,5v / 2 II HOp-
ap;peHaJIIiHa 0,01V / 2, a B meJIyp;OqRaX 1,9 II 0,01V / 2 COOT-
BeTCTBeHHO IIPII IIOJIHOM OTCYTCTBBII cepoToHIIHa II p;OlIaMIIHa,
aBTopbl C1.J.IITaIOT, qTO Mep;IIaTOpOM B ap;peHeprII1.J.eCROii HepnHoii: CII-
CTeMe JIHrymRII CJIymIIT ap;peHaJInH 1.
I CBeAeHIIH 0 ROJIII'leCTBeHHOIII COAeplliaH1I1I RaTeXOJIaMIIHOB II COOTHomeHHII
aApeHaJIHHa II HopaApeHaJIIIHa B cepP;l\e aMcJ>1I61ID: O'leHb HeOp;HmmaqHbI H npo-
TIIBOpeqlIBbl (rOBldpIlIl, 1967).
78
8:IeHTpOHHOMIIRpOClWIIII'IeCRII Y OCHOnaHIIH meJIYJJ;O'IRa cepp;qa
JIHrYIlII>II (Rana esculenta) nORa3aHbl nepDHbIe BOJIORHa II CIIHaIICbl,
cop;epmarn;ne nY3LlpbRII II rpaHYJIbl pa3HbiX pa3MepoB (Rybak et
aI., 1966) . ABTOpLI cpaBHIIBaIOT 3TII OHOnqaHlIH C XOJIHHeprlI'Ie-
CRIIMII II a)];peIIeprII'IeCRIIMII Y P;PyrlIx n03BOHO'IHLIX.
Do aIlHbIM B. A. fOBLlpllHa II f. P. JIeoHTbeBoll BbIJlCHHeT-
CH, qTO ap;peHeprIIqeCRaH IInnepBaqllH IIMeeTCJI TOJIbRO B Tex yqaCT-
Hax COCYJ:!IICTOll CIICTeMbI, rp;e B CTpYRTypHblH COCTaB CTeHOR CO-
CYP;OB BXO]:(JlT rJIa]:(ROMLlllIe'IHbIe ;meMeHTLI. IIo p;aHHbIM
r. P. JIeoHTbeBoii (1966a, 6), I1CCJIe]:(oBaBllIell B 3TOM HaIIpaBJIeHIIII
BHYTpeHHlIe opraHLI RpyrJIOpOTbIX, pL16, aMcl>1I611ll, peIITlIJIlIll, nTIIq
II MJIeROIIIITaIOrn;IIX, II B. A. fOBLlpliHa ( 1965), 113yqlIBllIero ap;pe-
HeprII'IeCRYIO IIHHepBaqIIIO nOrrepeqHOrrOJIOCaTblX Mbllllq, CJIep;yeT, qTO
RamI.TIJIHpbl II BeHbl, JIIIIIIeHHLle rJIa]:(Roll MYCRYJIaTypbl, He IIMeIOT
D CBoeM COCTaBe IlHuepBaqlIoHHI>IX rrplI6opOB CIIMnaTlT'leCROll rrpII-
PO)];bl. f. P. JIeoHTbeBa (1968), cpaBHIIBaH pacnpep;eJIenne ap;peHep-
rlI'IeCRIlX BOJIOROH B apTepIIHX II BeHax JIHrYIliRII C cocyp;aMII PI,I6,
rOBopllT 0 3HaqlITeJIbHOM YCJIOmlIeJllIII CTPYI.TypLI ap;peHeprlI'IeCRIIX
HepBHLIX rrplI60poB D CTeHRax RpoBeHocHbIX COCyp;OB aMcl>"6IIif.
CJIeP;yeT nOp;qepRHYTb, 'ITO MbICJIb 0 TOM, qTO B mlIBOTHOM Mllpe TOJII>-
RO rJlap;ROMblllleqHaH TRaHb o6JIap;aeT clIMrraTIIqeCROll IIHHepBa-
D;IIell, IIplIHap;JIemlIT T. A. fpIIropbeBOll II 1I3JIOmeHa B ee MOHO-
rpacl>IIII (1954). fplIropbeBa rrOJIaraJIa, qTO MexaHII3M CHMrraTII'Ie-
CROll I1HHepBaqlIll Bcex BHYTpeHHIlX opraHOB onocpep;OBaH opraHHLI-
MII RpOBeHOCHblMII cocyp;aMII, Hecyrn;IIMII B CBOIIX CTeHRax rJIap;Ro-
Mblllle'IHble BOJlORHa, II qTO napeHXlIMaT03HbIe 3JIeMeHTbi caMIIX
opraHOB, nOCTpOeHHbIe II3 p;pynlx TRaHell, Herrocpep;CTBeHHoii CIIM-
rraTII'IeCROll IIHHepBaqlIeii He 06JIap;aIOT.
EOJIee COBepmeHHaH opraHlI3aqllH CIIMrraTlI'IeCRoii: HepBHO" ClI-
CTeMbI Anura II 06IIJIbHaH IIHHepBaqIIH eIO TRaHell p;eJIaIOT II3JIIIIII-
HllM nop;po6Hoe onIIcaHlIe ee qeHTpaJIbHblX II ncpIIcl>eplIqeCRIIX nplI-
60POB, o6ecrre\:mBaIOrn;lIx BblCOHIIII ypOBeHb cl>YHRqIIoHaJIbHOll aRTIIB-
HOCTII II HepBHoro ROHTpOJIH Hap; opraHaMII BIIcqepaJlbHblX CIICTeM,
B TOM 'IIICJIe Hap; cepp;qeM. OCTaHOBIIMCH TOJIbRO Ha TOM, 'ITO IIMe-
eT R HaM HenOCpe)];CTBeHHoe OTHOllieHIIe. Y JIHrymRII yme IIOJlHOCTbIO
ccl>opMIIpoBaHbl 6eJIbIe II cepble rami communicantes, ROTOpbIe co-
ep;IIHJlIOT IIX co ClIIlHaJIbHblMII HepBaMII. TaRaH Mopcl>OJIOrlI'IeCRaH op-
raHII3aqlIH CBJl3eii aBToHoMHoii: HepBHoll ClICTeMbl C qeHTpaJIbHoii:
06eCIIe'IlIBaeT 6ecxBocTblM aMcl>II6I1HM CTporo cerMeHTapHoe pacrro-
JlOmeHIIe CIIMIIaTII'IeCRIIX Y3JIOB B nOrpaHlIqHblX CTBOJIaX, OTJllI'IaIO-
rn;ee IIX B aTOM OTHOllleHlIlI OT XBOCTaTbIX. IIocTraHrmlOHapHble
CIIMnaTII'IeCRIIe BOJIORHa, ROTOpbIe B II306UJIIIlI IIHHepBlIpYIOT COCY]:(II-
CTYIO cIIcTeMy, OTXOP;JlT IIJIII OT Y3JIOB norpaHII'IHoro CTBOJIa IIJIlI
OT npeBepTe6paJlbHbIX raHrJIlIeB, Tome yme ccl>opMlIpoBaHHblX y aM-
cl>II6IIii: (Nicol, 1952).
TaRIIM o6pa30M, y Urodela Y3:Jbl nOrpaHII'IHOrO CTBOJIa He IIMe-
lOT cerMeHTapHOll CTPYHTypbl. Y Anura OHII CTporo cerMeHTapHbl.
XpoMacl>cl>IIHHble WIeTRII, ROTopLie y Urodela COCTOJIT II3 pa3p;eJIb-
79
HbIX (He CJIIITbIX) Macc, TeCHO CBH3aHbi C CIiMnaTlIqeCRIIMII raHr-
JIIIJlMII Ha npOTHmeH1I1I Bceii: a6p;OMIIHaJIbHOM 06JIaCTII (Giacomini,
1898a; Noble, 1931; Hirt, 1934; Francis, 1934; Nicol, 1952).
"Y Anura OHII B DII]:(e nOJIOCOR JIemaT lIa l3eHTpaJIbHOM CTopOHe no-
'IeR, HO, I\pOMe Toro, Haxop;HTCH II BHYTPII CIIMnaTII'IeCRIIX raHrJIII-
eB (Giacomini, 1898a; Krause, 1923; Noble, 1931; Nicol, 1952).
Bce II3JIOmeHHOe n03BOJIHeT 3aI\JIIOqIITb, qTO 6ecIIopHp;OtIllOMY
pacrrpep;eJIeHlIIO CIIMnaTll'IeCI\I1X Y3JIOB nOrpaHlIqHbIX CnJIeTeHIIii Y
Urodela II CTporo cerMeHTapHOMY Y Anura COOTBeTcTByeT II pac-
nOJIOiheHIIe Y HIIX naparaHfJIIIOHapHblX Y3eJIROB.
nOCJIe OTcTynJIeHIIH, ROTopoe Mbl cp;eJIaJIII C QeJIblO pa360pa
CTPYRTYPHblX oc06eHHocTeii: CIIMnaTlIqeCKOM HepBHoii: CIICTeMbl 3eM-
HOBOp;HbIX. Mbl B03BpaTIIMcH R xapaRTeplIcTIIRe MeT3MeplIII xpoMaiP-
iPlIHHblX OCTpOBROB Y pa3Hbix OTPJlAOB aMiP1I61IM, 3aRoHoMCpnOCTIJ
IICTOplI'IeCROrO pa3BIlTHH ImTopoii HeJIerKO no]:(p;aIOTcH aHamI3Y
(Bachmann, 1954).
l\Ibl BIIp;eJIII, qTO Y 6e3Horoii: qepBHrH (Dittus, 1936) xpoMaiP-
iPlIHHaH TRaHb npOJlBJIHeT elQe OTqeTmmylO TeHp;eHQIIIO rpynnnpo-
BaTbCH B6JIII31I cerMeHTapHblX COCY]:(OB TYJIODIIrn;a, 'IeM Bbl3BaHa ee
coocTBeHHaH cerMeHTapHOCTb. "Y Urodela II Anura 3BOJIIOQUH xpo-
MaiPiPlIHHoii TRaHII HanpaB.rreHa Ha ROHQCHTpaQlI1O B QeHTpaJII.noii
06JIaCTII Te.-rIa, rp;e OHa ClQe coxpaHHeT npOqHYIO CBH3b C nOIJeq-
HblMII BeHaMII, IIX pa3BeTB.1IeHIIHMII II ROHTaRTlIpyeT C MarIlCTpa.'1b-
HbIMII cocyp;aMII 6pIOIlIHOii: 06JIaCTII, HO TepHeT CBH3b C cerMeHTap-
HI.IMII I\pOBeHOCHI.IMII cocy,rWltUi. nOTepH 3TOM CBH3U BJIeqeT 3a c060n
II 1I0TeplO MeTaMeplI1I B pacnOJIOmCHlIII xpoMaiPiPlIHHblX rpynn, OCO-
6eHHO y XBOCTaTblX aMiP1I61IH. "Y oeCXBOCTbIX me, HapH y C no-
Tepeii MeT3MeplIlI, 06YCJIOB,1IeHHoii pacnpOCTpaneHIIeM xpoMaiPiPlIH-
Hb1X rHe3p; BP;OJIb cerMeHTapHblX COCYP;OB, B03HIIRaeT HOBaH MeTa-
MepuH, 06YCJIOBJIeHHaH HORbIM 3TArrOM 3BOJIIOQHOHHoro pa3BIITIIH
p;aHHoro OTpHp;a 3eMHOBop;HbIX II CBH3aIIHaJl C oOJIee DblCOROH CTpyR-
TypHOii OpranlI3aQUeH ClIl\lIJaTlIqeCROii nepBHoii: CIICTeMbl. CerMeHTap-
HOe CTpOeHlIe Y3JIOB nOrpaHlIqHOrO CTROJIa y JIHrYllieR onpep;eJIHeT
II MeTaMepHoe pacnpep;eJIeHIIe xpoMaiPiPlIHHbIX RJIeTO'IHbiX rpynn,
ROTOpbIe Co1Iep;YIOT 110 nYTHM pacnpOCTpaHeHlIH CIIMnaTII'IeCRIIX 3JIe-
MeHTOB. "Y XBOCTaTblX 3MiPI16IIH HeT CTporo cerMeHTapHoii CTPYR-
Typbl nOrpaHUqHOrO CTBOJI3 II OHII, rrOTepHB «CTapylO COCYP;lICTYIO
MeTaMepUIO), He nplIo6pemi II (<HOBoii clIMnaTUqeCROil MeTaltWplHn.
l\Ibl Ha3bIBaeM MeTaMepIlIO xpoMaiPiPuHHoil TRaHII y 6e3Hornx
aMiPII6IIii (mepBlIqHOih), T. e. IICTOplIqeCRII 60JIee ApenHeii iPopMoii,
ROTopaH 06YC,TIOBJIeH3 TOnOrpaiPUqeCRIIMIi COCY]:(I1CTo-xpoMaiP<I>IIIIHbl-
Mil ROHTaRTaMII. MeTaMepli10 y Anura - (<BTopllqnOH), B03HIIRllIeii
B p;aJII.Heii:llIeM 3BOJIIOl\IIOHHOM npol\ecce aeMHOBop;HbIX, B pe3YJIbTaTe
ROToporo 06pa30BaJIIICb HOBbIe JIOIW,1JbHble CBH:1I1 C CIIMnaTUqeCliOll
HepBHon CllcTeMoii. OTpH]:( Urodel3 no CBonM MoPiPoiPYHlH.{UOHaJIb-
Hblltl Cl30iicTBaM CTOIIT 6mm,e 1\ Anura, xpoMaiPiPlIHHbIe IweTRlI y
HUX 0,111131\0 ROHTaRTnpylOT C CIIl\lIIaTllqeCRIIMII DJIeMeHTaMII II He
IIMelOT MeTaMepHoro paCIIOJIOiheIlIiH. B TO me BpeMH xpoMaiPlt>nH-
80
Hble 3JIeMeHTbl nOTepHJIH CBH3b H C CerMeHTapHblMII cocyp;aMH TYJIO-
BHrqa, a nMeCTe C HHMII II <mepBII'IHYIO» MeTaMeplIlO, CBOHCTBeH-
HYIO 6e3HOfIIM aM<p1I6IIHM.
TaKoBbl HalliH npCp;CTaBneHlIH 0 MeTaMeplIH xpOMa<p<pIIHHOH TRa-
HII BHYTPII Rnacca 3eMHOBop;HbIX.
H'poMe onucaHHblX XpOI\Ia<p<pIIHHbIX KOMnneRCOB Y B3pocnblX nH-
rYllieR BCTpeqalOTCH rpynnbl 3TIIX KneTOR B nepH<pepWIeCRIIX op-
raHax, B6nH3n COCYP;UCTbIX CTeHOR uml HepBHblX 3J1eMeHTOB BereTa-
TIIBHOH CIICTeMbI. GHII onIlcaHbl, HanpUMep, B apTepUH Y ROpHH
.nen>HX (CI>llIpHOB, 1890) H B menyp;OqRaX cepp;II;a B6nH31I IIX BHYT-
peHHeH 060nOqRH (Chakravorti, 1959). MbI HaXOp;IInH IIX B ap;BeH-
TIIII;nn aopTbl, B COCTaBe cnMnaTlIqeCROrO Y3na Y KOpHJI 6pblmmeH-
KII (pIIC. 18) II B P;Pyrux MeCTax. Erqe 60nee mHpORoe pacnpo-
cTpaHeHHe xpoMa4><PIIHHbIX S:WMeHTOB BCTpeqaeTCJI B nepHop;, npep;-
llIeCTBYIOIII;lIil MeTaMop<po3Y JlHrYllieR. XapaRTepHO, 'ITO Y rOJlOBaCTI1-
KOB pa311blx CTap;uH nop;o6Hble rpynnbl xpoMaW4>IIHHI>IX R.neTOli
BCTpeqalOTCH qarqe, qeM y n3pOCJlblX. GHII paCceJIHbI nop; 3ll;I;OTeJIHeM
opraHHblx MeJlRUX COCYP;OB - Benyn (pIlC. 19), apTepUOJl (puc. 20)
II. I.alln.'I."'IHpOB (puc. 21), a TaKme Ha BceM npOTHmeHUU aopTbI
(pnc. 22, A, B) H 3a]:(Heii: nOJIoii BeHLI (puc. 23).
HanlI'lue ap;penamIH- II Hopa]:(peHamIHcop;epmarqux KJleTOR B Hap;-
IIO'IeqHllKax II B p;pyrnx nepll<pepUqeCI\lIX opraHax YCTaHOBneHO y
P;BYX RIlP;OB .'IHrymeK - RaIla arvalis II Rana temporaria; uccne-
]:(OBaHbl rO,TIOBaCTlIKII Ha 23, 24, 25, 26, 27, 28, 29, 31-ii CTa]:(HHX
pa3BIITHH II ceroneTRII I. I1cnO. T lb30Baml MeTop;bI XpOMlIpOBaHIIH II
1I0p;upOBaHUH no Hillarp, Hokfelt (1953, 1955) II <pnyopeCII;eHTHblii
no Fakk, Owman (1965).
Ha XpOI>llIpOBaHHbIX npenapaTax cYMMapHblH COCTaB <peHO,TIbHbIX
aMlIIlOB BnepBbIe o6HapymIlllaeTCH B II;IITOnna3Me KJleTOK lIa 26-ii
CTa]:(IIII n napaCTaeT B nocnep;Hue cpOIm OHTOreHe3a (CMuTTeH,
1963) .
qeM CTapllle B03paCT mlIBOTHoro, TeM 60Jlee OrpaHIPIeHO pac-
npOCTpaHenHe xpoMa<P4>IIHHOH TRaHU B opraHax H TeM 60nbme OHa
KOHII;eHTpIlpyeTcH B 06J1aCTII neplIap;BeHTIIII;lIaJlbHOrO npOCTpaHCTBa
6plOlliHOH aopTbI, nonoH BeHbI H COCYP;OB BeHTpOMep;HaJlbHorO KpaJl
noqeR.
Hop8p;peHamlHoBble KJleTRn yp;aeTcH o6HapymlITb MeTop;OM IIop;n-
pOBaHlIH B II03p;HeM nepHop;e .TIIlqUHOlIHOrO pa3BIITIIH - Ha 29-H CTa-
p;nn. GHn BCTpeqalOTCH B CTeHKax COCY]:(OB pa3Horo RaJlH6pa, Ha.
qIlHaH C aopTbl (pnc. 22, B) H ROHqaH MeJlRIIMH OpraHHblMII cocy-
]:(aMII. RaRon-nlI60 3aKoHoMepHocTH B pacnpep;eJleHHH XPOM- II HO]:(-
nO.'IOmnTenbHbIX KneTOR OTMeTnTb He yp;aeTcH. To me caMoe I1MeeT
leCTO B Ha]:(nOqeqHURax.
TIo.1.yqeHHble p;aHHble 0 cpoKax nepBblx npH3HaRoB CIIHTe38 rop-
MOHon B xpoMa4>4>HHHbIX RneTKax, COOTBeTCTBYIOIqlIe 26-H CT8P;HH
1 Onpep;eJIl'UHe B03paCTa mlrymeK npOll3BOAHJIOCb no nplI3HaKaM, YRa3aHHldM B
Ta6JIHl\aX Rugh (1962).
81
.-
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(',
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'11
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_ .' !t.. A " .- f#' ,(:; R",\, . " I .
,<.. " ",.,r. "I JJ' ; ,..<1". "i; "-"""f ¥ - ...
. .. >+ '* '> '" ": L ....'iI> -,.\', ,. "
f. "''Ih - ." ... ' ,..
J'-" ... ,.',"''''; ' 1f-" ',:'-... r "Ij
.. ... . lr
.j, 1 , .1 Jk ___ ..!! <';;: ' i7 - /'; i
-. ....
:R T '" -..r
::<t
.
:.
.,;IV;,
.,
- -.....
,
.. ..
II II c. 18. llaparanrJ\JlOwqllllolii Y;II'.,1(1I{ ]I c"CTan!' CJI\llIaTlI'[l'CHOrO y;wa Y 1>01)1\11
UIJL!lli}lWIIRII Jwry"nm (R. t!'mporaria)
OHpaCRa MeTHJIOBbIM aeJleUbIM-nIlpOIllIlIOM. 01<. >( 5, OU. X 20
Pile. 1 ). XIJoMa4>41JIIIIII,](' 1\.!l('T1;1I 1I0!! ;III;IOTP.'ljll'\1 y \1l'CTa nna.'l.('JIIIH IWIIJIJI-
JIHpa IJ ]I!'HYJlY n 1I01\;I>!'JlYl\o'lJloii ;],('JI(';II' ro.'IO'IaCTlll'" .!lRryIlI1>JI (H. arYali ).
27-11 cTa,o,TlH no Tafi.!llll{l' Hugh. X]lo la(I) IJlllllaH P('iU"\1U1 no XIIJlJlapny n XeK-
nl'JIbTY
OH. X 5, 06. X 20
,;b
rf
. ]I'J4' ;
' .... _ ""'.. >'JJ, ' 0'_7]#-:"'- ..
'" .,f ; ',...".. .'" 0 . .J ' t . . ",)
, {' -\ ,. it":f, ', '; ,
f", ' ...-.J. d: .,....'.
< :' ''*{ - 'i Jt - .-:-' :-/
.; . 't.
v .
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... , 'I,. '.""" .". !
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h ft th. ".",...",':,, . ..... '..
h.o _ ...C' -...;.' c.,, ' .... ':;;' ..--,,"" -:' J
1 . ;' It: 'I>'.'L>"""'.
'/ '>:" 1: .. ,i £i;:
,
:> ?'.:.- ,'"I
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..... " -,
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<. t"p -,.
<F ,.... . "" '\,' lPiI
)(4.
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.... ,.j:.....A. X!':","
't' .:. "..:- -:.c:-> ".:1--....< )i' V'
. . eC ,,"(... . ": :1:,. j, ;.'
,.. . ".. _,.. <., h.# " " .. ,"i<" '.
<; >/ "i' tIt"" ,,' ., - .;" <;.: " :xT
" ,
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'T
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',: ".. _ >. .h- .Jr;t"t-."....
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<>" - ff' v
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'11: -=-t ." to- ,:
f'> ".- .
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.
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, 'I ....
I'.I C. 20. XplJ Ii\(I'(IIlIllllhll' JlJII'T1\1I II a,l,UIL'I< II'1J1 "PTl'I'ILJ.lhl rO. 0lH1rTl1l\a JlH.
ryulIOI (It. al'yalis). :W-H na;ulH 110 TallJ1111l1' Hugh. .\I'O)H1( ( lnTlTaH pCa1\/Ulfi
IlO XlT.'l:WPIlY II .\Clwe:lI,ry
'1 - npOCBeT apTCpIIOJIbI; 6 - xpOMaep!jJIIIIHbIC HJICTlm; 6'- nlfI'MCHT. OH. x5 OU. x40
l' II c. 21. Xpmlarp( lIIJ\J\<Il' JlJ1I'Tlm Ul'pl'TI<llooop:1alloii .pOpMr.I OKOJIO RamIJIJIHpa
II lliI]:{TTO'II"IIlIII,1' rOJIOnaCTIlI,a !IHrYlllim (II. tl'mporaria). 31-H CTa]:{IIH no Tau-
,TIlII\" Hngh. Xpo la.prplIilllilH III a:;11I11I 1111 '\1I'L'lapny II XelWOJILTY
01'. x 5, oG. X 20
-,.-----r ... \,.; . ;?/
." '- ' '?i.
c, . ; . -..... f,.
":f ;: : :; :'J'
"'" - . ..
/ . -.-.:; -iii ; :
"'
..
> .-. ;-- ';-
.I .s' :.,!..., \--3 . .
.,,"" 10\ ': '<-,,:
t (, '?
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-'t ... )", , i :
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''lh.j-L .,. !"!:' '.
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JIIICb B 06JIaCTU rOJIOBHoii: nOqRlI, 6pIOIlIHOii: 06JIaCTII U 3a6p101lI1IH-
HorO npOCTpaHCTBa cpe]:(Heii: U 3a]:(Heii: TpeTII Ty.'10Bu:m;a, a lIIeJIKII&
ee rpynnM nOJIyqUJIII pacnpOCTpaHeHIIe B pa3JI1I'-IHMX BHYTpeHHIIX
opraHax B COCTaBe CIIMnaTUqeCRUX HepBOB, IIAy:m;ux no XO;J;y apTe-
plIOJI MbIllIe'lHOrO Tuna. 06UJIbHOe CKOnJIeHlIe xpOMa<p<PlIHHoii: TRaHlI
11 napaaOpTaJIbHOii 6pIOillHOii: 06lIaCTlI HBJIHeTCH 3TanOM UCTOpll'le-
CKoro pa3BIITUH naparaHrJIUOHapnoii: TKaHlI, ROTOpblii: OTpamaeT no-
CTeneHHYIO .QeHTpaJIU3a.QIIIO aH;J,QJipllIlllOii a]:(peHaJIOBoii: <PYHK.QIIII, CO-
npHmeHnyIO C nporpeCCIIBHbIM pa3BUTIIeM CUMnaTII'leCKOii HepBHOll
CIICTeMM. HaJIH'-Iue pacceHHHbIX XpOMa<p<PIII-lUbIX KJIeTOK B KpOBeIlOC-
HbIX cocY;J;ax 6eCXBOCTbIX aM<p1I6IIii, O'-IeBIIAHO, TaK me HBJIHeTCH
pe3YJIbTaTOM nporpeccupYlo:m;eii: ;)BOJIIOD;JIII CIIMnB.TlI'leCROii HepBHoii:
CIICTeMbI II 06UJIbHOrO pa3BeTBJIeHlUI ee nOCTraHrJIUOHapHMX BOJIO-
KOH B rJIa]:(KOMbIllIe'lHbIX COCYAIICTbIX CTeHKax.
HaM OCTaeTCH paccMoTpeTb B aToii: rJIaBe e:m;e OAun Bonpoc, Ra-
caIO:m;IIii:cH <paKTUqeCKOrO MaTepUaJIa no naparaHrJIIIHM, pacnOJIO-
meHHbIM B ma6epHbIX cOCYAax :m'lUHOK II IIX rOMOJIOraX Y B3pOC-
JIbIX oco6eii:.
3eMHOBO]:(HMe, BnOJIHe onpaB]:(blBaH CBoe Ha3BaHlle, 3aHUMaIOT
npOMemYTO'lHOe nOJIOmeHUe Mem]:(y CBOUMU BOAHMMII npe]:(KaMU-
pM6aMU U 60JIee BbICOKO opraHII30BaHHbIMU Ha3eMHbIMII n03BOHOq-
HbIMU. Mx COCYAlICTaH CIICTe m OT'laCTU coxpaHHeT qepTM, CBOii:CT-
BeHHMe mllBOTHbIM. Be]:(YI.QIIM nO]:(BOAHblii: 06pa3 m1I3HII, HO npll06-
peTaeT U HOBbIe oc06eHHOCTII, npucnOC06JIeHHbIe R Ha3eMHOMY CY-
IIWCTBOBaHllIO. )1,JIH anaJIII3a HHTepeCYIO:m;ero Hac Kpyra BonpOCOB
o COCYAUCTo-naparaHrmIOHapHbIX YCTpoii:cTBax, 06eCneqUBaIO:m;ux
<PYHR.QUOHaJIbHYIO aRTlIBHOCTb ropMOHOB xpOMa<p<plIHHOll CIICTeMM,.
He06xoAIIMO paCCMOTpeTb CTeneHb pa3BIITIIH II MOP<POJIOrU'leCRU&
oc06eHHOCTl! COCYAIICTOii: CIICTeMbI aM<pu6uii:.
B nepuo]:( pa3BlITIIH JIU'-IUHOH, YCJIOBUH cy:m;eCTBOBaHUH ROTOpbIX
OrpaHII'leHM BO]:(HOii: cpe]:(oii 06IITaHlIH, ]:(bIxamre ocy:m;eCTBJIHeTCH
ma6paMlI, U Bce apTeplIaJIbHMe ]:(yru y HUX coxpaHHIOT e:m;e <PYHK.QUIO
COCY]:(UCToro opraHa, CBOllCTBeHHYIO pbI6allI. IIocJIe MeTaMop<p03a, no,
Mepe nporpeCCIIBHoro pa3BlITlIH opraHOB ]:(bIxaHIIH, qUCJIO apTe-
pUaJIbHbIX Ayr YMeHblllaeTCH. )1,Be nepBble napM pe]:(Y.QupYIOTCH B
paHHeM OHToreHe3e. 3TOT np0.Qecc HaqlmaeTCH e:m;e Y ABOHRO]:(MllIa-
:m;UX pbI6 II 3aKaH'lIlBaeTCH Y aM<plI6uII nOJIHoii aTpO<pueii: ABYX nep-
BbIX nap (puc. 26). M3 TpeTbeii: napbI ma6epHbix COCYAOB II KOp-
Heii: 6pIOIlIHOii: aopTbI pa3BlIBaeTCH CnCTeMa HapymHblx II BHYTpeH-
BUX COHHbIX apTepuii:. M3 qeTBepTOii: napbl B03HUKaIOT IIapHble ]:(yrII
cnuHHoii: aopTM. Y XBOCTaTMX aM<pu6uii: cy:m;eCTByeT TaR Ha3bIBae-
MMB COHHblii npOTOR (ductus caroticus), rOMOJIOrlI1JRMii IIapHMM
89-
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PH C. 26. CxeM8.I'01\ICJIOrIiH apTep:!; JIbHbIX p;yr pa3JUI'lHbIX n03BOHOYJHbIX (Blip;
'C 6plOmHoii: CTOpOHId)
I - IICxogHOe paCnOJIomeHHe Y 3apO bIIIIefi: 1 - (j - apTepHaJIbHblC AYI'H; 7 - 6pIOIIIHaH
aopTa; 8 - CllHHHaH aopTa; II - VII - apTepHaJIbHaH CHCTeMa: II - pbI6a (gBOHHOgbIIIIa-
IilaH): 3 - 6 - npHHOcHII{He H BbIHOCHII1He ma6epHIiIC apTepHH; 9 - JIeI'O'lHaH apTepllH;
III - XBOCTaTaJI aM4J1I6HH: 3 - COHHbIe apTepHH; 4 - gYl'a aopTbI; Ii - 60TaJlJIOB npOTOH;
10:- COHHblt1: npOTOH; IV - 6ecxBocTaH aMdlH6I1H; V - pellTHJII1H; VI - IlTHl\a; VII-
MJleROnHTaIOII1ee
:.:.-::. ,:: .J
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IWpHHM aopTbI MemJl,Y TpeTbeH II qeTBepToH p;yraMu, a TaKme COX-
paHHeTcH nHTaH napa apTepllaJlbHbIX Jl,yr B BlIJI,e TOHRIIX COCYJl,OB,
ROTOpbIe <PYHKlJ;llOHllPYIOT B RaqeCTBe BTOpOH Jl,06aBOqHOH napbI
aOpTaJlbHbIX Jl,yr (pllC. 26). Y 6ecxBocTblX aM<p1I6llH nHTaH napa
apTeplIaJIbHblX ]:(yr IIC1Je3aeT, a COHHbIH IIpOTOK npeBpa:m;aeTCH B
'COHHo-aOpTaJlbHYIO CBH3RY (ligamentum aortico-carotideum). JIe-
rOqHble apTepllII HBJlHIOTCH rOMOJlOraMII meCTOH napbI p;yr, HO B HllX
npeBpa:m;aIOTCH TOJlbKO npORCllMaJlI>HbIe Y1JaCTKII meCTOH: napbl,
a Jl,llCTaJlbHbIe OCTaIOTCH B BlIJI,e 8HaCTOM030B; y XBOCTaT1>IX :'JTll
aHaCTOM03bl npeJl,CTaBJleHbI CPYHRIJ;UOHIIPYIO:m;IIMU 60TaJlJlOBbiMII npo-
:90
8
VI
TOR3!\1II, Y 6eCXBOCTbIX - HeWYHInJ;IIOHlIPYIO nMII 60Ta.rrJIOBbIMIl
CBJl3KaMII (CM. pnc. 26). (llhIMKeBUq, 192 ; IllI\HlJIM3Y3l'H, 1938;
TepeHTbeH, 1950; BOPOHIJ;OB8 II p;p., 1952). BeHo3HbIii CUHYC, KO-
TOphlii Y IITIIQ II M."ICROIIIITaIO IIX BXO.n;I1T B COCT8B IIpe cep.n;nii.
a Y peIITlfJIuii eIIJ;e OT."III'II1M ImI. C31\IOCTOHTe.lIbHOe 06pa30BaHne,
Y 3MqJHonii e e o6oc06JIeH.
ECTccTBeHHo, 'ITO HaCTy"aIO 3H 1I0C:Ie MCT1uIOpw03a pe.n;YKIJ;I1H
ma6epHoro alIlIapaTa .n;omKH3 1I0B:Ie'lb 3a c060ii II IIpon:ecc pe,u:YK-
I.(IIII IlJIII 1I0JIHOrO IIC'Ie3HOBCHIIH xpoMaWWIIIIHbIX IWMIIJJCI{COB, KO-
Topble BXO.n;HT B CTPYKTYPHbIii COCTaB CTeHOK m86epHbix 8pTepIIii.
IIpoQeccy IIX 3TPOWUII cIIoco6cTRyeT e e II TO 06CTOHTeJIbCTBO, qTO
y I1MWII611ii 13 pe3Y.TIbT8Te HcconcpmCHCTRa ;'lero1.JHoro arrrr3paTa lJ;bl-
XaHIIJI C03.n;aeTCH .n;OIIOJIITllTeJIbHaH ,lJ,blxaTe.lbHaH CilCTeM8, oLy ecTB-
JIJleMaJl Komeii II CJIII3IICTOii 060JIOqKOM pOTOBOM rrOJIOCTrr. OTcTaBa-
HIIIO MOPWOJIOrnqeCKOrO p33BIITIIJI JIerKIIX OT WU3IIOJIOrIIqCCRIIX IIO-
Tpe6HocTeii: opr8HII3M8 B KlIc.rropo,u:e (TepeHTbeB, 1961), O'IeBII]:(HO,
COIIYTCTByeT oIIpe]:(eJIeHHblii CTPYKTYPHo-WYHKI\IIOH3JIbHbIll ypOBeHb
COCy.n;IICTOii CIICTeMbl ma6epHoii 06JIaCTII. Co CTOpOHbl M8JIoro Kpyra
KpOBoo6p8 eHIIJI, JlBJIHIO erOCJI p;epIIBaTOM m86epHbix cocy.n;OB, He
Tpe6yeTcH BbIlIOJIHeHIIJI CJIOmHblX mlI3HeHHO B8mHbIX 3ap;8q, ROTo-
pbIe IIpe.n;'bHBJIHJIliCb Obi I{ HeMY OpraH8l\1II JIeroqHoro P;bIX3HIIJI. Ha
:1TOM OCHOB8H111I MomHO yme 38p8Hee rrpe;J;IIOJIOmIITb, qTO CTPYRTypbl,
ROTopbIe JlBJIJlIOTCH rJIaBHblMII IICTOqHIIKaMII 8p;peHeprlI1.JeCKIIX ne-
CCTB y HII3KoopraHII30BaHHbiX mlIBOTHblX (xpOMaWWlInHble opra-
Rbi), MorYT WYHl{n:UOHllpOBaTb rrOCJIe MeTaMOPW03a He CTOJIb H8-
rrpfUKClIHO, IWI{ eCJIII OM IIM npIIIlIJIOCb ::ITO p;eJI8Tb B YCJIOBIIJlX BI.I-
COROpa3BIIToii .n;bIXlne.'IbHOii CIICTeMbl. IIHblMII CJIOB8MII, H8.n;0 rro."Ia-
raTb, 'ITO rrap3r8rrr.'IIIII, P83BUTbie B ma6epHoii CIICTeMe II B OROJIO-
m86epHbIX 80pTaJIbHbIX yqaCTKaX pbIO II KpyrJIopOTblX, Y aMwu6IIll
TepJlIOT CBoe WII3110."IOrIpreCROe 3H8qeHlle B RaqeCTBe nymHbIX op-
rarrOB.
IIpoIJ;ecc pa3BIITIIH KpOBeHOCHoii CIICTeMbI B PHAY rr03BOHO'IIIhIX
IIOKa3bIB8eT, qTO BbIXO.n; aUIBOTHbIX Ha CYIllY - eMeH8 BO]:(IIOii: cpe.n;bI
OOIITaHIIJI Ha H83eMHbIii - C.TIYiI\I1T OCHOBHhJM waRTOpOl\I, OIIpe,u:eJIHIO-
IIM MOPWOJIOrIIqeCKlIe oc06eHHoCTII II rrYTb OHTOrellGTllqeCIWro pa3-
BIITIIJI KpOBeHOCHOll CIICTeMbI 3 l\Icf1n61111. He l\lOmeT 6hlTb COMHeHlUl
B TOM, qTO CTPYKTypHo-wynKQIIonaJIhHbIe rrepccTpoiiIm, H3cTynaIO-
IIe B KpOBeHocHOll CIICTeMe lIa :1TOM :1T8rre :JBomoQIIoHHoro pa3-
BIITIIJI, He MorYT He OTp8311TbCH H8 T8RIIX iKe CTPYKTYPHO-WYHK-
QrrOHaJIbHbIX lIepecTpoiiKax 1I8par8HrmlOH8pHOll cIIcTeMbI, TeCHO cnn-
3aHHoii C cOCy.n;IICTbIM pYC.'IOM. ECTeCTBeHHO, qTO OT.n;eJIbHhIe opraHbl
::ITOM CIICTeMbI rro.n;BepraIOTCH H81I60JIbllleMY BJIIIJlHIIIO aTpOqJlPle-
CKIIX IIpOQeCCOB B Tex yq8CTK8X COCY]:(UCTOll CIICTeMbl, KOTOpbIe C8MII
60JIbme Bcero lIepeCTpaIIB8IOTCH, pe.n;YQllpYIOTCJI II 8 TPOWIIPYIO'lCJI.
T8KIIMII y<I8CTK8MII JlBJIJlIOTCH ma6epHbIe COCy.n;bl. CTerreHb MOPWo-
JIOrlI1.JeCKOM peP;YKQIIII B IIUX xpOMaWWIIlIIIbIX OCTpOBKOB, OqeBII.n;HO,
3aBIICIIT OT YPOBHJI 06 lIx aTpOWII'IeCIHlX IIpOQeCCOB B Tex IIJIII IIHblX
OTpe3K8X ma6epnoii CIICTeMbl: BO npeMJI MeTaMOPW03a BMeCTe C ma-
91
6epHbIMII cocy.n;aMII IICqe3aeT II XpOMacpcpUHHaH TKaHb, a B cocy;(ax,
KOTOpbIe JIII60 coxpaHHIOTCH B Bu.n;e pyp;IIMeHTapHbiX npOTOKOB, JIII60
B BH)J;e coe.n;IIHIITeJIbHOTI<8HHbIX CPU6p03HbiX THmeM, OHa MomeT no]:(-
BepraTbCJI JII060M CTeneHII pe.n;YKQIIII.
MopcpOJIOrlIqeCKlIe oco6eHHOCTII pa3BIITIIH cOCy.n;IICTOM CIICTelllbl
B ma6epHOM 06JIaCTII aMcp1I61IM, OTpamalOlQlIe nepexop;HblM aTan IICTO-
plIqeCKorO pa3BIITIIH, npe.n;CTaBJIJlIOT 60JIbmOM IIHTepec. IIplIMeqa-
TeJIbHbl B paBHOM CTeneHII KaK cpaBHIITeJIbHQ-MopcpOJIOrH'leCKlIe ]:(aH-
HbIe Memp;y B3pOCJIbiMU OC06HMU XBOCTaTbIX II 6ecxBOCTbIX aMcpIl6IIM,
TaK II CBep;eHIIJI 06 IIX JIlIqlIHOqHOM pa3BIITIIII, npOTeKalOlQeM B Bo.n;e.
K COmaJIeHIIIO, xpoMacpcplIHHaJl TKaHb y aMcplI6IIM He II3yqeHa B
TaROM acneKTe, II B JIIITepaType MbI HamJIII TOJIbKO CBe.n;eHIIH 0
_xpOMacpcplIHHbIX KJIeTKaX TpeTbeM napLI ma6epHbIX COCy.n;OB, 1I3 KO-
'TOpbIX pa3BUBaeTCH CIICTeMa COHHblX apTeplIM, II 0 naparaHrJIIIHX
cep.n;Qa. Mem.n;y TeM 3HaqeHIIe aTIIX IIaparaHrmIeB y aMcplI6IIM IIMe-
.eT npHMoe OTHomeHlIe K OQeHKe KapOTn.n;HbIX II Kapp;IIo-aOpTaJIbHbIX
opraHOB y MJIeKOnlITalOlQlIX II qeJIOBeKa. C aTOM TOqKII 3peHIIJI HaII-
OOJIblllIIM IIHTepeC npe.n;cTaBJIJlIOT TpeTbH, qeTBepTaJl II meCTaH IIapLI
ma6epHblx apTeplIM, KOTOpbIe yqaCTBYIOT B CPOpMlIpOBaHlI1I Hapym-
HOM II BayTpeHHeM COHHblX apTeplIM, p;yrII aopTbI II JIerOqHOM apTe-
plIII (pIIC. 27, A, B, B). B JIlIqlIHOqHOM pa3BIITlIII aMcpu6lIM TpIl
nplIHOCJllQlIe ma6epHbIe apTepnlI, II.n;YlQlIe 110 TpeTbeM, qeTBepToii
II nJlTOM ma6epHbIM .n;yraM, paClIa.n;alOTCJI Ha KanlIJIJIJlpbl B Hapym-
HblX ma6pax U nona.n;alOT B BbIHOCHIQIIe ma6epHbIe apTeplI1i II ]:(a.'Jee
B cnIIHHYIO aopTY; Ha MeCTe meCTOM apTeplIaJIbHOM .n;yrll ma6phI
He pa3BUBalOTCJI II OT Hee OTXO.n;JlT JrerOqHbIe apTeplIli.
Ha IIYTII TpeTbeM ma6epHOM .n;yrlI y JIlIqlIHOK pa3BIIBaeTCH rete
mirabile - qyp;ecHaJl cocyp;lIcTaJl ceTb (pIIC. 27, A), Ha MeCTe KO-
TOpOM Y B3pOCJIbiX aMcplI6IIM o6pa3YlOTcJI KapoTII.n;HbIM CIIHYC II Rapo-
TH)J;HbIM JIa6lIplIHT (CM. IIIMaJIbraY3eH, 1953).
B HaCTOJllQee BpeMJI I1MeeTCJI orpOMHoe qIICJIO ony6JIIIKOBaHHbIx
pa60T 0 pa3BIITIIII II CTpoeH1I1I KapOTIIf];HOrO JIa6lIplIHTa aMcpII6IIM.
OCHOBHbIe BonpOCbl, KOTOpbIe aKcneplIMeHTaJIbHO IICCJIe.n;yroTcJI II
.o6cym.n;aIOTcJI aBTopaMII, MomHO pa3AeJIIITb Ha .n;Be rpynlIbI. B 0]:(-
HOM II3 HIiX 6bIJIli 1I3yqeHbI aHaTOMIIJI II rllCTOJIOrH'leCKOe CTpoeHIIe
KapOTH)J;HorO cocy.n;IICToro JIa6IIpIIHTa II 3HaqeHIIe aTOM CTPYKTYPbl B
cpII3IIOJIOrlIli KpOBOo6paIQeHIIJI aMcplI6IIii. B p;pyroM rpynlIe pa60T
rJIaBHOe BHIIMaHlIe aBTopOB yp;e:meTcH nOIICKaM KJIeTOqHhIX CTPYR-
TYP, rOMOJIOrlIqHblX KapOTU]:(HOM meJIe3e IIJIII KapoTII.n;HOMY «rJIOMY-
. cy» BLICIlIIIX n03BOHOqHbIX, B TOM qlICJIe MJIeROIIIITaIOIQIIX. B 60.1Ib-
.mIIHCTBe llCCJIe.n;OBaHllM anTopLI Kaca.JIIiCb 060IIX BonpocoB II qaCTO
.06'he.n;IIHHJIll MOPCPOCPYHK:QIIOHaJIbHOe 3HaqeHUe Tex II ,u:pyrllx
CTpyKTYP ( cocy.n;lIcToro JIa611pUHTa U ImeTOqHLIX aJIeMeHTOB, eCJIII
nOCJIe.n;Hue 6bIJIII IIMII Hall,u:eHbl).
,lJ,eTaJIbHOe uayqeHlIe pacmupeHlIoro yqaCTKa 061Qeii: COHHOII ap-
Tepllu - «rJIaBHOM KaMepbI» JIa6UpllHTa, ee ]:(IIcTaJIbHOro nOJIIOCa, IIO-
cTpoeHHoro II3 rYCTOM KanllJIJIJlpHOM ceTII, a TaKme CTPYKTypbI
BHYTpeHHeii: II HapymHoM cOHHon apTeplIl1 II IIX llHHepBaQuII, HC
92
A
Pile. 27. CxeMa llia6epHoro
RpoBoo6pa eHUH all1clJH611H
A - JIlt'lHHRa xBocTaTot1: aM-
iI6ltlt; B - B3pOCJIaH ca,oaMaHgpa;
B - B3pOCJI8H JIHI'YIIIHa; ao. d.-
CnHHHaH aOpT8; a, e - npHHOCHII1He
JI Bl>IHOCHIqHe apTepltH HaPYlliHblX
ma6ep (br); ei, ee - BUYTpeHHHH It
HaPYlliHaH COHHble apTepHH; pi - CO-
cygHCTOe CnJIeTeHlte, H3 HOTOpOI'O
pa3BIIBaeTCH HapOTHAHblfi napal'aHI'-
JIHA; p - JIeI'0'lH8H apTepHH, Bel-
nOARJlIO'lH'lHaH apTepltH; e. mes-
HllIIIe'lHO-6pLIllilliee'lHaH apTepHH
{IUMaJIbl'aY3eH, 1938)
,
,
I
I
, I
.."
npllBeJlII, o.n;HaIW, K e.n;IIHO]:(YIliHOMY MHeHlI10 0 CPYHKIJ;IIOHaJlbHOM 3Ha-
qeHlI1I ;:noro opraHa (Boyd, 1933; PaIme, 1934; Pischinger, 1934;
Boissezon, 1939a, b; A. CMllpHOB, 1951; Carman 1955 II .n;p.) 1.
::hOT Bonpoc MbI OCTaBJlJleM B CTopOHe KaK He IIMelOlQlIH npJIMoro
<lTHOmeHlIJI K HallllI1\1 3ap;aqa1\I, a OTHOCJlIIJ;UMCJI, CKopee, I{ cneU;II-
cplIKe YCJlOBlIll reMo.n;IIHaMIII{1I Y aMcplI6nll. H'apOTlI]:(HaJl 06J1aCTb lIH-
TepecyeT Hac C TOll TOqRlI 3peHlIJI, qTO JlBJlHeTCH COCY]:(lICTOH
CTPYRTYPOll, 06pa30BaHHOll MHorOQIICJleHHbIMII TOHlWCTeHHbIMlI COCY-
.n;lICTbiMlI aHaCTOM03aMlI, KOTOpbIe coe]:(lIHHIOT nepBYIO apTeplIaJlbHYIO
p;yry C COHHOll apTeplIell. CpeAII PMa npe.n;nOJlOmeHlIll cYlQecTByeT
MHeHlIe 0 rOMOJlOrlIlI KapoTII]:(Horo JlaOlIplIHTa II KapoTlI.n;Horo clIHyca
MJleKOnIITalOlQlIx. OHO OCHOBaHO Ha HeKOTopblX .n;aHHbIX pa3BIITIIH
3TOro 06pa30BaHIIJI, ero CTpoeHIIH II IIHHepBaU;lIlI.
1 JIIITepaTYpa no MopcJ>OJIOrnll RapoTIlp;HOro JIa6npllHTa aMcJ>1I611H Han60JII'e
IIOJIHO IIpnBe)J.eHa B MOHorpacJ>HII Adams (1958).
93
}{apOTII.n;HhIH JIa611pHHT OTC-YTC-TnyeT y 6C3HOfl1X lIa Bcex CTa-
.n;HH.X pa3BHTIIJI (Klump, Egg('rd, 19; 4) II Y JII1'HmOR 6eC-XBOCTLIX II
XBOCTaThIX. Y IIOCJIe]:(HIIX OH (POpMupyeTcH TOJIbl-lO B nepIIO.n; MeTa-
MOPW03a, T. e. nOCJIe pe]:(yRI.{IIII maup II pa3llIITIIH JIerRIIX - B TOM
me MeCTe, r.n;e 06pa3YIOTCJI RapOTII]:(HhIH clmyc MJIelWIIJITaIOIQIIX (Bo-
yd, 1933; Fedele, 1941; Foxon, 1952). ITo rucTOJIOrHqec-n:OH CTPYI,-
Type CTeHRII rJIaBHOH RaMepbl RapOTn,J;Horo JW6IIpnHTa TepJlIOT MI.}-
meqHO-3JIaCTHqeCROe CTpOeHlIe, Xapal{TepHOe ]:(JIH COHHbIX apTepllH,
HCTOHqaIOTCJI Hano.n;06He RapOTH]:(noro cliHyca MJleIWnHTaIOIQHX U CO-
CTOJIT 113 3JIaCTlIqeCRHX :JJIeMeHTOB (Carman, 1955). ::hIaCTlIqeC-Ime
3JIeMeHTI.I H3 RapoTH]:(Horo JW6HpHHTa BMeCTC C TOmnIMil ROJIJIareHO-
BbIMH BOJIORHaMH II coe,n;IIHIITeJIbHOTIiaIIIIhIMlI RJIeTI-\aMII IHIJIeTaIOTC-H
B RanHJIJIHpHYIO ceTb H 06pa3YIOT He}I{HYIO MemRannJIJIHpHYIO TI\aUb
C MeHbllleH (Rau, 1924) IIJIn 60JIbllleU (Palme (1934) npUMeCbIO
rJIa]:(RHX MblllleqHblX BOJIOROH (Boissezon, 1939 a, b; Chowdhary,
1951). COrJIaCHO pa60TaM Fedele (1941) II Neil C c-oaBTopaMII
(1950), RapoTH.n;HbIH JIa6npllHT cHa6maeTc-H MemlmpOTII,D.IIOH BeTBhIO
Jl3bIR0-rJIOTOqHOro HepBa ceH30pHoro xaparnepa II 6JIym.n;aIOIQHM Hep-
BOM, ROTOpbIH nOCblJIaeT BeTBH qepe3 ropTaHHI.lii HepB R JIaUnpIIH-
TY, a Qepe3 rJIOTOQHblH - R .n;yre aopTLI. B C-OCTane 6JIym.n;aIOIIJ;erO
HepBa, B03MomHO, npoxo.n;JlT BOJIORna .n;enpeecopuoii npHpo.n;bI (Niki-
forowsky, 1913), H rJIOTOqHaJl BeTBb ero HBJIJIeTCJI rOMOJIOrOM p,e-
IIpeCCOpHoro HepBa MJICROnHTaIOII(HX (Fedele, 1943). B 60JIee n03,J,-
HeH pa60Te Carman (1955), a .n;o Hero Palme (1934) nOJIaraJIII,
QTO 60raTaJl HHHepBal.{IIH oCYlQeCTBJIJleTCJI npelIMYlQeCTBeHHO B 06-
JIaCTH 06IQeH COHHOH apTeplIH, a c06cTBeHHo RapoTII.n;HbIH JIa6npnHT
IIMeeT MaJIO HepBHblX BOJIOROH. ITplICYTcTBlIe B RapOTII.n;HOM .lw6n-
pHHTe ]:(BnraTeJIbHblX HepBOB, IIO MHeHIIIO Adams (1958), Oqenb
BepOJlTHO, TaI{ RaR B OI{pymaIOlQeH coe]:(IIHUTeJIbHOH TRaHH, RaR npa-
BHJIO, BCTpeqaIOTeJl Mlmp0C-IWlInqeCRlIe raHrJIUH, BnepBLIe onlICaHHbIe
elQe Zillimerlllann (1887).
rOMOJIOrHJI RapOTII.n;Horo JHJUnpHHTa xnOCTaTblX II 6eCXBOCTbiX alll-
clJII6IIH H RapOTH]:(Horo eIluyca MJIeIWlJIITaIOIIJ;IIX, IWTOpbIe pacnOJIO-
meHbl y MeCTa .n;eJIeHHH o6IQeH C-OIIHOit apTepnH H pa3BIIBaIOTC-H
IIPH YQaCTIIH COCY]:(OB TpeTLeii ma6eplIoii .n;yru, n03BOJIHeT .n;OIIYC-
THTb, qTO B RapOTII]:(HOM JIaUnplIRTe I1MClOTC-H HJIeTI{H, no]:(06HbIe TeM,
ROTOpbIe 1I3BeCTHI,l B l{apoTH.n;noH OPJIaCTH BbICIlIHX 1I03BOHOqHbIX
RaR naparaHrJIIIOHapHbIe JIJIH «rJIOMYCHLIe» II HOTophIe OIIHcaHbl RaR
xpoMaclJ4>IIHHbIe B ma6pax RpyrJIopOTl>IX II pbl6 (Giacomini, 1902a,
b; Gaskell, 1912).
Ha npOTWReHlIH MHorHX ]:(ecHTIIJIeTHii aBTopbl, 1I3yqaBllIIIe Hapo-
TII.n;HbIH JIa6HplIHT y pa3HbiX IIpe.n;cTaBHTeJIeii aMWH6HH - TpIITOHOB,
CaJIaMaH]:(p, JIJlrYllieR II ma6, He nm:ryqaJIH B 3TOM OTHOllleHHII O]:(HO-
3HaqHbIX pe3YJIbTaTOB. Pa3HorJIaCIIH RaCaJIIICb caMoro clJaHTa cYlQe-
CTBOBaHIIJI cnel.{HaJIbHLIX RJIeTOR, OIIHCaHHJI rHCTOJIOrHqeCHOH CTpyH-
TypbI H lICTOqHHROB, 113 HOTOpblX 3TH RJIeTHII pa3BHBaIOTCH. C Ha-
llIeH TOqIm 3peHIIJI, pa3HOpeqlIBOCTb pe3YJIbTaTOB HCCJIep;OBaHIIH MO-
meT 6hITb JIOHHTa II 06'bHCHHMa, eCJIH yqeCTb pa3nyIO CTeneHb
94
pe.n;ym lIl1 II nepeCTpOlum B ma6epnoM aun8p8Te y pa3HbIX OT-
pJl.n;OB aM.p1l611II. C ;JTUX rKe n03IIQlIii WUI IiClil(6TCH nOHHTHOII QIITOJIO-
rllqeCliaH neo.n;Hopo.n;HOCTb OlilIcallHblX pa311blMlI 8BTopaMII KJIeTOK.
Boissezon (19398) , Ha oCIIOB8I1IIII cpaBHIITeJIbHO-QIITOJ"IOrnqeCKIIX
nplI3HaRoB - 60JIblllOrO CBeT.'loro H;:t;pa II CBeTJIOII BaRyoml3lIpoBaH-
HOH npOTOnJIa3Mbl, 1i0TOpbIe Oil Haomo;:J;aJI B Ii;leTliaX Ii8pOTII]:(Horo
.TJa61Ip1l1lT8 Anura II B liapOTII]:(HOM 'feJIe MJIeKOIIIITalOll\HX (Bois-
sezon, 1936, 19398), Cp8BHIIBaJI lIX ]:(pyr C .n;pyroM II H83bIBaJI
anIITemIOII.n;HbIMH. B 60JIee n03.n;HIIX IICCJIe]:(OBaH1IJlX HX lI]:(eHTII.pII-
QlIPOB8JIlI JIII60 Tome C KJleTliaMII ImpOTlI]:(Horo TeJIa MJIeIwnlITalO-
lI\IIX (Chowdhary, 1951), JIUOO C cIIMllaTHlIeCIHIMlI IiJIeTIiaMII (Fedele,
1941) . C .n;pyroH CTOpOHLI, lIl\ICIOTCH pa60TbI, B 1i0TOpLIX aBTopbl
JIII60 Bo06lI\e OTplIII,alOT CYll\eCTBOBaHlle KaKIIX-JIII60 cneQH.plIqeCKHX
3::rellleHTOB B 1i8pOTu.n;nOIlI .lC\onpIlIITe (Han, 1924; Boyd, 1933; Hat-
tori, 19; 4; Palme, 1934; Carman, 1955), JIII60 npIlnIlCblBalOT 11M
He aliTo.n;cpM8JIbHOe npOlIcxom.n;eHlIe, 8 pa3BIITlIe 113 3H.n;OTeJIIIJI CO-
Cy.n;OB (Pischinger, 1934).
CJIep;yeT OTMeTIITb, qTO 8TIIII1I3M CTpOCHIIH JlaparanrmWH8pHbIX
RJIeTOK II Hp;ep B K8pOTII]:(HOii 06JIaCTII, napJl.n;y C HeraTlIBHOH .peo-
XpOMHOH peaKQlIeu IIX QIITOn,'la3MLI, HCO]:(HORp8THO CJIyilUIJI nOBo.n;OM
Ii OTpllQaHll1O IIX 1I8paraHr:1II0H8pHOii npllpo.n;bl. 8TO OTHOCIITCJI He
TO."IbIiO K 8M.pu611Hl\I II He TO.lbliO K IIX 1i8pOTII.n;HOMY yqaCTRY, HO II
I. 80pTaJIbHblM, nYJIbMOH8JIbIlbillI II npllcep]:(eqHblM naparaHrJIIIHM
Bcex n03BOHOqHbIX. nplIMepOM MomeT CJIymlITb pa60Ta Milcou
(1930), I{OTOpbln 06CJIe.n;oB8JI R8pOTII.n;HYIO 06JIaCTb y JIJlrYllleR, 6bl-
1i0B, KpLIC, KpOJIIIKOB, KomeK, JIOma.n;eH, c06aK, 06e3bJlH 11 QeJIOBeI{8
(Bcero 103 OC061I). Mop.poJIOrUqeCRIIM IWHTpOJICM aBTOpy CJIymIIJIII
naparaHrJIIIOHapHbIe liJIeTKII 1I103rOBoro CJIOH H8,n:nOqeqHIIROB II 6plOm..
HblX n8p8raHrJIIIeB Tex me mIIBOTHLIX. 06HapymeHHYIO P83HIIQY B
p83MCp8X, .popMe 11 CTpYRType RJIeTOqHLlX JI.n;ep Mem.n;y naparaHrJIIIO-
HapHbIMII IiJIeTRaMII Ha.n;nOqeqHIIKOB II cneQII.pIf'IeCKIIMII RJIeTRaMII
1i8pOTII.n;HOII 06JIaCTII aBTOp paCCMaTplIBaeT KaR 8prYMeHT npOTIIB
IIX OTom.n;eCTBJIeHIIH.
B xo]:(e pa3BUTIIH I1CCJIe]:(oBaHlIu KapOTII]:(Horo lIaparaIlrmIH y
MJIeliOnIlT81OIl\IIX II no Mepe IIpll3H8HIIJI ero ImK XeMOpell,enTOpHou
CTPYKTYPbI IIOJlBHJI8Cb Teup;CHQIIH I1CI{8Tb B K8pOTIIAHOM JI86IIplIHTe
aM.p1l61IH COOTBeTCTBYIOlI\YIO cPYHKQlllO, a RJIeTRaM, CYlI\eCTBOBaHlIe
KOTOpLIX B TOIl 11.'111 IIHoii qJOpMe npll3HaeTCJI IICCJIe.n;OBaTeJIJlMII (IipO-
Me Fedele, 1941) nplIlIIIC8Tb xeMopeQenTopHYIO pOJIb (Boissezon,
1939B; Chowdhary, 1951). B CBoeH IIIOHorpa.pIlIl Adams (1958),
CTOHIII;IIH Ha n03I1QIIII B830-CeH30pHOrO TOJIROB8HIIH Ii8pOTII.n;HOll
30Hbl MJIeKonIIT81OIII;IIX, pa3RIIB8eT :JTY me TOqKY 3peHHH II B OTHO-
llIeHlI1I aM.p1l6IIn. C O]:(HOH CTOpOHbl, OH npIl3HaeT, qTO BpJl.n; JIll
RapoTII.n;HbIH JIa6IIplIHT aMIlIII6IIH B QeJIOM MomHO, no 8HaJIorlIlI C
MJIeKOIIllTalOIII;IIMII, np1l3H8Tb 3a xeMopeQelITopHblH opraH - CJIHm-
KOM OTJIIPIHO ero CTpoeHlle. C .n;pyroH CTOpOHbl, OH BblCKa3blBaeTCH
38 xeMopeQenTopHYIO nplIpo.n;y cneQllllmqeCRIIX KJIeTOR. Adams He
MomeT COrJI&CIITbCH C TeM, qTO OHII OTCYTCTBylOT B K8pOTIIAHOM JIa-
95
6npllHTe (Palme, 1934; Carman, 1955 II .n;p.) II nOJIaraeT, qTO,
HapJl.n;y C KJIeTKaMIi cocy.n;I1CToro npOllcxom.n;eHlIJI (Pischinger, 1934),
B 3TO MeCTO n03me BHe.n;pHIOTCH xeMopeQenTOpHbIe R."1eTKII II f{ame
He IICKJIlOqaeT B03MomHOCTII 06pa30BaHlIJI IIX in situ (11). 8TO no-
CJIe.n;Hee npe.n;nOJIOmeHlIe JlBJIJleTCJI nOCJIe.n;OBaTeJIbHbIM pa3BlITIIeM ero
1I0JIOmeHIIH 0 IIpOllcxom.n;eHlIlI xeMopeQenTopHblX KJIeTOI. l.apOTII]:(Ho-
ro «rJIOMyca» y MJIeKOnIITalOII:\nx, y KOTOpbIX, H8 ero B3rJIH,1, :JTOT
Bonpoc .n;o CIIX nop OCTaeTCH OTKpblTblM (Adams, 1958).
B nOCJIe]:(HlIe rO]:(bl ony6JIlIKOBaH PH.D: IIHTepeCHblX paoOT, B KO-
TOpblX K8poTn.n;Hb1H JIa6nplIHT aMcplI6IIH n3yqeH HOBeHmmm IIIeTo]:(a-
Mil QIITOXlIMlIqeCKorO II IITOJIOrlIqeCKOro aHaJIlI3a, n03BO;IIIBIlIIIX
YBlI.n;eTb pJl]:( .n;o Toro He lI3BeCTHblX .n;eTaJIeH, IIMelOII:\IIX npIIIIIIlI-
nlIaJIbHOe 3IIaqeHIIe .n;JIH OQeHKII nplIpo.n;bl cne H(I)JIqeclmx K:leTOK
II npIIHa.n;JIemHOCTIi nx JIII60 K 3H]:(OKpIlHHbIM, JIII60 xeMopeQeIITOp-
HblM :JJIeMeHT8M. Gouder n Desai (1966) nOKa3aJIII n03IITIIBHYlO
peaKQlI10 ImeTOK Ha 06pa60TKY 6lIxpoMaToM K8JIliH no Hillarp II
Hokfelt II <I)JIyopec eH 1I1O no Eranko ( 1955c) . EIIO.'lorII'feCKIIe
np06bl 3KCTpaKTOB 1I3 KapOTlI.n;Horo JIa6IIplIHT8 II 113 cynpapeHa;IO-
BblX KJIeTOI{ Ha.n;nOqeqHlIKOB JIJlrymeK nOKa3aJIli B 060lIX CJIyq8J1X
yqaII:\eHlIe cep.n; e6IIeHIIJI II nOBblllIeHlIe KpOBJlHOrO .n;aBJIeHIIJI (KOHT-
pOJIeM CJIymlIJIa IIepCPY311J1 cep.n; a paCTBopOM PIIHrepa). ABTOpbl
nplImJIII K BbIBO.n;y, qTO KJIeTKn KapOTII.n;Horo JIaOUpIIHTa .'lJIrylllel\
JlBJIHIOTCJI xpOMacpCPUHHbIMII, ceKpeTnpYIOII:\I1MlI 8.n;peH8JIIIH II Hopa.n;-
peHaJIlIH, II He pa3.n;eJIJlIOT TOqKY 3peHIIJI 0 ceH30pHoil IIpnpo.n;e I1X.
HeHcHO TOJIbKO O]:(HO: 38qeM aBTop8M nOHa.n;0611JIOCb .n;JIJI 3Toro OT-
plIIJ;aTb IIHHepBaIJ;IIIO K8poTII.n;Horo JI8611pIIHT8 II ero n8paraHrJIIIOHap-
HblX KJIeTOK Jl3bIK0-rJIOTOqHbIM HepBOM? HaK JII060H opraH, KaK JIIO-
6aJl .n;pyraH 3H.n;0I{pIIHHaH meJIe38, B TOIlI qnCJIe II M03rOBOH CJlOil
Ha.n;nOqeqHIIKOB, XpOM8cpcpIlHHbie KJIeTKII JIa6UpIlHTa 06JIa;J;a1OT a.p-
cpepeHTHblMII npIl60p8MII, CJlrHaJIII3IIpYIOII:\IIMJI B eHTp8JIbHYIO HepB-
HYIO CIICTeMY 0 CPYHI\IJ;IIOH8JIbHOM COCTOHHIIII nepIIcpepUII. MHHepBa-
IJ;HJI JIa6IIpIIHTa K8pOTU.n;HOH BeTBblO H3b1K0-rJIOToqHOro HepBa bblJI8
nOKa3aHa Palme (1934) II Fedele (1943), a Laurent II Krieger
(1965) OnIICaJIII B KanIIJIJIJlpHon 30He RapOTlI]:(HOro JI86IIplIHTa JIJI-
rymKII KJIeTKII C KpynHbIM OBaJIbHblM H.n;pOl\1 II 'IeTKIIMII rpaHHU8111I1.
KOTOpbIe OII.'IeTeHbl lIepBHbIMn BOJIOKH8111I1, 06p8aylOII:\I1MII Ha nx no-
BepXIIOCTn YTOJIII:\eHIIJI II cnnpaJIII. 8T11 OKOIlQaHIIH, npIIHa.n;JIem8II:\IIe
qyBCTBIITeJIbHblM MJlKOTHblM BOJIORlIaM IX n8pbl, 06p83YlOT CnJIeTe-
Hne Ha 1I0BepXlIOCTII opr8Ha, I1pOHlIKaIOT BHYTPb, TepHIOT MHKOTb,
.n;IIXOTOMnqeCKn .n;eJIHTCJI II, He npepbIB8HCb Ha coce,u:Hnx r8Hr.'11I03-
IIblX KJIeTK8X, 1I0.n;XO,u:JlT K KJIeTKaM II cpIl6p03HbiM TJlmaM JIa6J1-
pImT8.
. B OqeHb 06CTOJlTeJIbHoii: pa60Te Rogers (1966) IICCJIBp;OBaHbl aHa-
TOMIIH, rIICTOJIOrIIH, IITOJIOrIIH II IITOXnMnJl KapOTIIp;Horo Jla6nplIH-
r8 Tpex BII]:(OB 6eCXBOCTblX aMcpn6IIH (Bufo marinus, Hyla aurea II
Neobatrachus pictus). Ha OCHOB8HIIn 60JIbmoro KOJllIQeCTB8 rliCTO-
XIIMlIqeCKnX pe8K IIii:, nOCT8BJIeHHbiX C eJlblO onpep;eJIeHIIJI cpepMeH-
TOB, cpeHOJIbHblX aMlIHOB, pn60HyRJlelIHOBOH l\IICJIOTbl, rJllIKOnpOTelI-
96
flpocdem COC!lOO
o
fJ-K/lem
"
.'
ZUlJMIf
----..
.'
PII c. 728. CxelIIa. IIaparaHrJIllOllapHlde RJIeTRJ] JI3 RapoTIip;HOH 06JIaCTH JIHrymRII
(Rogers, 1966)
HOB, cpOCCPOJUIIIII.n;OB II .n;pyrIlx coe.n;IIHeHIIH. a TalnRe Ha OCHOBe IJ;!Y-
TOJlOrlIqeCRIIX oco6eHHocTeH Rogers xapaRTeplI3yeT RJleTOqHblII co-
CTaB CIIHycOII.n;aJlbHoro CnJleTeHlm 8McplI6IIn .n;BYMJI BII.n;aMlI RJleTOK --
a II 13. nepBbIe (a-KJleTKII) co.n;epmaT K8TeXOJlaMIIHbl, IIM€IGl'
OB8J1bHYIO IIJlII HenpaBIIJlbHYIO CPOPMY, KpynHoe JI.n;po C MeJlKIIM
CKy.n;HblM xpoMaTIIHoM II 3epHlicTYIO II.JIII rOMoreHHYIO I.{IITOITJla3MY.
H8 cepnn:Hblx cpe38x YCTaHoB.'leHo, qTO a-KJleTKII npoHII3bIBalOT
BCIO TOJl y CTeHKB H8pymHoii COHHOn: apTeplIII: B a.n;BeHTIIIJ;IIII UX
1a:IO, HO IlX KOJlIflleCTBO YBeJllIqIlBaeTCJI B Memcocy.n;IICTblX TWI\8
CIIHYCOII.n;HOro CnJleTeHIIJI. HeCKOJlbKO RJleTOR Han:;J:eHO Bp;OJlb 60KO,
BOll CTeHKIl rJlaBHOn K8Mepbl :la6IlpIIHTa. B 8.n;BeHTIIIJ;BIO 06 eH COlI.
HOII apTeplI1I OHII He pacnpOCTpaHJlIOTCJI, a B 8;J;BeHTIIIJ;I1I1 BHYTpeUr
Hen COHHOll BCTpeqalOTCJI pe.n;I\O, a-KJleTKlI KOHTaKTHpYIOT C HepB-
HblMII BOJlOKH8MII, 06pa3YIO IIMII Ha HIIX 6YJlaBOBII.n;HbIe CIIHanCLI.
I3-KJleTKII mleHb CJlOmHOH cpOpMbl (pIIC. 28), OHB 06p83YIOT ceTe-
BII.n;HbIe CTPYKTYPbl II CBJl3aHbi OTpOCTKaMlI C 3H.n;OTeJlUeM CIIHYCOIl-
.n;OB, C a-KJleT.RaMII II IIHOf]:(a rpaHII'IaT Henocpe]:(CTBeHHO C npo--
CBeTOM COCy.n;8.
Rogers cpaBHIIB8eT a- II I3-KJleTKIl C I{.lIeTKaMII liapOTIl.n;Hom
TeJla MJleKOnIITalO IIX II Ha OCIIOBaHlI1I 1I3MeHeHIIJI fIICTOXIIMIflle.-
CKIIX pe8KIJ;IIH B HIIX no]:( BJUmHlIeM aHOKCIIII II flInepKalIHIIII, oqer ,;..,
BB.n;HO, CqIlTaeT IIX XeMOpeIJ;enTOpHbIMII. B TO me BpeMJI OH npo-':
BO]:(IIT cpaBHeHIle IIX C a]:(peHaJiIIH- II Hopa.n;peHaJlIIHco.n;epma MII
KJleTKaMII «fJlOMYCHblX» 06pa30BaHIIH MJleKOnIITalO IIX. J];BOHCTBeHr
HOCTb 3aKJlIOqeHIIll aBTopa II TPy.n;HOCTIl, KOTopble, no efO CJlOBaM..
B03HIIKalOT B MopcpOJlOrlI'IeCKOH .n;lIcpcpepeHIJ;lIpOBKe Mem.n;y .n;BYMJI TII.
4 H. A. CMHTTeH
97
naMll RJIeTOIi, p;enaIOT nOHHTIILIMII II Te pa3HOrnaCIIH, IWTOpLle ey-
eCTBOBaJIH B xapaKTepHCTlIKe :JTIIX KJJeTOK B IIpemHHX pa60Tax.
MLI He pHcKyeM p;aBaTb IIpOIl3ll0JILHbIe TOJIIWBaHHH pa60Te Rogers,
op;HaKO, 1I03BOJIHM BLlCKa3aTb HeI\OTopLle co06pameHlIJI 0 npHpop;e
OIIHcaHHblX 1111I a- 11 -KJIeToK. Ooa TlIlIa KJIeTOK MorYT npnHa;:J:-
JIemaTb K naparaHrJIIIOHapHbIM 3JIeMeHTaM, HO a-J\.'IeTKH eo-
p;epmaT .peHo.'lbHblC al\lIlHbl H OOJlap;aIOT eelipeTopHoH aJ\TUDHOCTbIO
(ux I1TOmla3Ma 1\IOmeT 6L1Tb JIuGo rOMoreHHoll, JIII60 3epHIIcToii),
a -KJIeTJ\n no DeeM IIp1I3HaJ\aM 1\IOrYT TpaJ\TOBaTbCH liaK PYP;II-
MeHTapHbIe, 1I0TepJ1BIlIUe xpOMa.p.pnHHoeTb :mel\IeHTbl. MomHO e;J;e-
JIaTb II p;pyroe npep;lIo."!omeHIIe II IlpIl3HaTb -lweTJ\I1 eycTeHTaKY-
}JJlPHbIMH, KaK Ross (19:59), J\oTopbJii onnea."! y.lbTpanpYIiTYPY Kapo-
Tup;Horo TeJIa MJleKOnIITaIO nx. :::ITO 1I0CJlep;Hee IIpep;nO:IOmeHue IIOA-
J\peIIJiHeTCJI pa60TaMII Ishii H Oosaki (1969), J\OTopLle Tome 1I0J\a-
3aJlU p;Ba KJleTO'lHbIX TIIlIa B KapOTII;:J:1I0M JlaGnpuHTe ma6 (Bufo
vulgaris), OAHU 113 J\OTOpbIX eop;epmaT OCMUO.pIlJlbHbIe rpaHYJlbl,
a p;pyrlle JIIIllIeHbl IIX II paCCMaTpUBaIOTCH aBTopaMII B RaqeCTBe
CYCTeHTaKYJlHpHbIX.
CepIDI I1CCJle]:(oBaHlIll Ishii C coaBTopaMH (Ishii, Ishii, Honda,
1966a Ishii, Oosaki, 1966b; Ishii, Honda, Ishii, 1966c; Ishii,
Ishii, 1967; 1970; Ishii, Oosaki, 1969) eJleycTpeMJleHHO HanpaB-
JleHa Ha BblHClleHIIe HHlIepBal(IlOHHbIX oTHollieHIIH B KapoTIIp;HOM
Jla6IIpIIHTe ma6 H MexaHH3Ma B03I1HJ\HOBeHHJI xeMoceH30pHbiX HIII-
IIYJlbCOB. CaMoll 1I0pamaIOru;elI, IWH BblpamaIOTCJI aBTopbl, oc06eH-
HOCTbIO KJleTOI{ HBJlHIOTCJI rp8UY':lbI, CXOP;HbIe C OCMIIO.pHJlbHbIMlI
rpaHYJlaMII M03rOBoro CJlOJl Hap;IIOqeqHHKOB H l{8pOTII]:(Horo TeJla
MJleKOlIlITaIOru;nx. Ha :JTIIX KJleTHaX, IWTOpbIe aBTopbI Ha3LIBaIOT xe-
MOpel(eIITOpHLIMU, I1MeIOTCJI p;na TlllIa HepBllblX OIwHQalluii. OP;HII
113 HHX co]:(epmaT CHHalITIIqeCKne IIy3LlpblHI H MllTOXOH]:(pIIII, JlB-
JIJIJICb IIpeCHHalITIIqeCIDIM 3BeHOM (11. pel(elITOpHaH KJleTKa - IIOCT-
CIIHaIITIIqeCKIIM) - 3.p4>epeHTHblii CIIIIane; P;PyrIle He IIMeIOT CII-
HalITIIqeCKIIX IIY3blPbKOB II CJIymaT IIOCTCliHalITIIqeCKOH MeM6paHOll,
IIpep;Ha3HaqeHHOH P;JlJI pe elITopHOH KJleTKII - a.p.pepeHTHbIH CIIHalIC
(K.Ishii, Oosaki, 1966, 1969). B 1I0IICKaX .pII3HOJIOrHlIeCKOrO 3Ha-
lIeHHJI 3.p.pepeHTHoll IIHHepBa IIII xeMope elITOpHoH KJleTKH 6bIJI IIO-
CTaBJleH pJlP; 3KCIIepIIMeHTOB (K.Ishii, K.Ishii, Honda, 1966a; Ishii,
Honda, Ishii, 1966c; Ishii, K. Ishii, 1967, 1970); aBTopLl YCTaHO-
BIIJIII pOJIb JIa6IIpHHTa B perYJIJI III1 p;LlxaHIIH II yqacTIIe CHMlIa-
TlIlIeCKOll II BaraJlbHOH IIHHepBa I111 B lIHrII6II IIH xeMoceH30pHofi
aKTIIBHOCTII. MHorIIe HeomIIp;aHHLle pe3YJIbTaTbI o6cym]:(aIOTcJI aBTO-
PI;IMII C TOlIJ\n 3peHHJI HX B03MomHoll PO,lIH B MexaHH3Me xel\lOpe-
elI IIII. He KaeaHCb Beero IWMIIJleJ\ca IIO]:(IIHTbIX aBTOpaMtI BOlIpOCOB,
XOlIeTCJI 1I0p;qepIiHYTb, qTO HX pa60TbI, yp;aqHO COqeTaIOru;IIe MOP.pO-
nOrIIqeeKlIe II 4>u3I10JIOrIIQeCKlIe I1ee.'le;J;OBaHliJI Ha O]:(HOM II TOM me
06'beKTe, y6e]:(HTe.'lbHO IIOK83bIBaIOT ]:(BOllnyIO IlHHepBa HIO TIIlIIlq-
HblX ap;peHaJIOBbIX JmeTOK C J\aTexo.TJCo;J;epmaru;IIMH rpaHYJIaMII. C:IY-
qaiiHoe IIPIICYTCTBlIe B HaII60.'lee ariTIIBHLlX xeMoceH30pHbiX 30Hax
kapoTII)J;Horo Jla6HpIIHTa 3TIIX KJIeTOJ\ BOBce He JlBJlJleTCH aprYMeH-
98
TllpOBaHHblM CmI]:(eTeJIbCTBOM JlX Y'laCTIIH B YIia3alIHOii <PYHKl{lm.
XOTJI HaJIUqlIe B l{IITOIIJIa3Me OCIIlIIO<pIIJIbHbIX rpaHYJI B HaCTOHrn;ee
BpeMJI yme He MomeT JlBJIJlTbCH a6COJIlOTHbiM apryllIeHToM B nOJIb-
3Y co]:(epmaHIIH B HIIX aMIIHOB (.n;:m :'JToro Tpe6YIOTCJI ]:(onOJIHIITeJIb-
Hble rIlCTOXIIMlIqeCKlle II 6UOXII:\III'ICClme aHaJIII3bl), TeM He Menee
3TOT IIpll3HaK HBJIJleTCJI O.n;HIIIII II; IIOKa3aTeJIeii :m]:(OKpIIHHoro xa-
paKTepa KJIeTOK. RpOMe Toro, IIl\1('I0lI.\lIeCH rIlCTOXIUnlqeCKUe .n;OKa-
3,\Te:IbCTBa IIpllCYTCTBIIJI IIIOHOH:\1IIII OB, 6110:WrII'IeCI..aJl IIpo6a 3KCT-
pa1..TOB II3 KapOTII.n;Horo JIa611plIIITtl If lIa.n;IIO Ie IHIIKOB JIHrymeK, Ko-
TopaH BbI3bIBaeT O HOTIIlIlIbIii 3CP<PeI,T Ha I,pOBHHoe .n;aBJIeHlIe
(Gouder, Desai, 1966), II, HaIWneu;, YJIbTpaCTpYliTypHoe CXO]:(CTBO KJIe-
TOR, IIIIIeIOrn;IIX ]:(Ba TllIIa I1CpnIILIX OROHqaHIIII B KapOTII.n;HOM JIa611
pllHTe (Laurent, Krieger, 1965) II Ha XpOllIa<P<PIIHHbIX KJIeTKaX Ha]:(-
IIOqe'lHIIKOB (Piezzi, '1966), He BLI3bIBaIOT COIIlHeHIIH B 3HP;OKpIlH-
Hoii npllpo.n;e KJIeTO'lHbIX :J.'Iel\leIITOB KapOTII.!I:HOro JIa6UpUHTa 3eMHO-
BO.n;HbIX. HeCMOTpH Ha 3TII <paKTLI, Ishii II Oosaki (1969) npHnlICbI
B8IOT 3TIIM KJIeTKaM peu;elIToprryIO <PYHKII,IIIO, HO IIpli aTOM np1l3HaIOT,
'ITO <p1I3I10JIOrIlqeCKOe 3I1a'leIIIIC ::J<p<peKTOpHOH IIHHepBal{IIH xeMO
pel elITopnblX KJIeTOK He IIO ]:(aeTCJI OlJ7,JlCHeHIlIO.
B IIpOTIIBOnOJIOlliHOCTL 1\apOTII]:(HOii 30He, KOTOpaJl XOpOIiiO 06-
CJle.n;OBaHa, Mbl He lIalII.TI1I n JIUTCpaType CBe.n;eHuII 0 xpoMaclJcllJlH-
IIblX 3JIeMeHTax B 06JI8CTU .n;pyrIIx ma6epHbiX .n;yr aMclJ1I611H. B CBO-
eii co6CTBeHHOH pa60Te, IWTOpaJl npono.n;IIJIaCb Ha JIllqllHKaX JIJlry-
IIICK, HaM yp;aJIOCb HaHTII rpYIIIILl, a IIHor.n;a OT]:(eJIbHO KJIeTKII TOJlblW
B 06JIaCTII .n;yrll aopTbl (pIlC. 22, B). RaK npaBIIJIO, OBII paclIoJIa-
ra.'1UCb B a.n;BeHTUl{IIII aopTbl II, qTO OqeHb BamHO, p;aBaJIJI OqeHb
CJIa6yIO nOJIOmlITeJIbHYIO xpoMaclJ<pIIHHYIO II IIOp;aTHYro peaKn;IIH.
lomHo .n;YMaTb, qTO UMeHIIO OHn HBJIHIOTCH OCTaTKaMII Tex MaCCHB-
HbIX lIaparaHrJIlleB, KOTOpLIC y ce.'1axIIii o6pa3YIOT MyclJTbl Ha 110]1.-
R.'1IO'III'lHLIX (aKCIIJIHpHbIX) apTepllHx II IIplIJIemarn;IIX K HIIM yqa-
CTKax aOpTbl. II TO II .n;pyroe - IwpeHb no.n;KJIIOqlIqHOH apTeplI1I I.J
.n;yra aopTbi - pa3BIIJIIlCb 113 IV II VI ma6epHbix p;yr.
II3JIOmeHHOe B HacTOHrn;en rJIaBe nOKa3blBaeT, qTO C nepexo.n;OM
1\ Ha3eMHOMY o6pa3Y llill3HII npOll30lIIJIII KOpcHHbie H3MeHeHHJI B
MopclJOJIOrll'reCKoM o6.'1IIKe a.n;penaJIOBOH CHCTeMbT: OCHOBHOH clJYH1\-
unoHa.'1LHLlii oqar ee Imnl{CIITpllpyeTcJI B OKO.'1oIIOqeqHOn o6.'IaCTII,
a OCTaJIbHble ee yqaCTKII IIO]:(BepraIOTCJI pe.n;YKU;IIII. O.n:HoBpeMeHHo
I1aCTyrraeT IIpOrpeCCIIBHLIII :ITaII pa3BIITIIH CIIMIIa-TUqeCKOH HepBHou:
CIICTeMbI. BIIlCcTe C HepBHLl:\1II :1.'1e:\ICHTaMU IIOJIyqaIOT IlIIIpOTWe pac-
npOCTpaHeHue a.n;peHaJIOBbiC 1,.'1eTIm, KOTOpble BCTpeqaIOTCJI B CIIM-
lIaTllqeCRnX Y3JIaX II COIIYTCTBYIOT HepBHbIM Bo.'1OTmalll, BIIJIOTb .n;o
I1X pa3BeTBJIeHIIH B napeHxIIMe BIIYTpeHHIIX opTaHoB.
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XpaHJlIOTCH cocy.n;IICTO-TRaHeBbIe OTHOllieHIIJI C xpOMacpcpIlHHLlMH . )JIe-
MeHT8MII, CBoii:cTBeHHbIe aMcpII6I1J1M; C ]:(pyroii: CTOpOHbl, y HIIX H8-
'1l1HaIOT cpOpMlIpOBaTbCJI Ha.n;nOqeqHLIe meJle3bl, OqeHb CXO]:(IILIe no
rIlCTOJlorIIqeCRIIM II U;IITOJlOrlIqeCRIIM npIl3HaR8M II 3M6pIIOHaJILHOMY
pa3BIITIIIO C meJle3aMII BblmeCTOJlII:\IIX n03BOHOqHbIX.
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IIX B RaqeCTn caMOCTOJlTeJlbHLlX opr8nOll n03BOJIJleT yme y npeCMbl-
RaIOII:\IIXCJI Bbl]:(eJlIlTb II pa3.n;eJlbHO paCCM8TpIIBaTb .n;Ba ROMnOHeHTa
xpOMacpcplIHHoii: TRaHIi IIJlII, TO'IHee, ]:(Be 06J1aCTIi ee p8cnpOCTpaHe-
HIIJI, ROTOpLIe, o.n;HaRO, B U;eJlOM npo.n;OJlmaIOT COCTaBJlJlTb e.n;IIHYIO
CIICTeMY. O.n;Ha 113 HllX - a.n;pen8J10B8H - cocpe]:(OTOqeHa B H8.n;no-
'IeqHIIRaX, .n;pyr8J1, ROTOPYIO y MJleRonIIT8IOII:\llX nplIHJlTO Ha3L1B8Tb
3RCTpaa.n;peHaJlOBoii:, COXp8HJleTCH B CTeHR8X RpynHblx RpOBeHOCHb1X
COCy.n;OB. J];eJleHIIe Bceii: CIICTeMbI Ha Ba pa3.n;eJla y.n;06HO .n;JlH ll3J10-
meHllJI cpaRTIIqCCROrO MaTepIIaJla. O.n;HaRO CJle.n;yeT 3apaHee CR83aTb,
qTO Bbl.n;eJleHIIe B OT]:(eJlbHYIO py6pIlRY BneHa.n;IIOqeqHIIROBOll (3I,CTp8-
a]:(peH8J10Boii:) TRaHn B ll3BecTHoii: CTeneHII npllHIIMaeTCJI HaMIl YC-
JlOBHO. B 3TOT pa3.n;eJl MLI OTHOCIIM TOJIbRO Te «OCTaTRII» BHeHa.n;no-
qeqHIIROBblX xpOl\lacpcpllHHLlX OCTpOBROB TRaHll, ROTopLle «YHaCJle.n;o-
BaHbl» OT pbl6 II RpyrJlopOTblX II COOTBeTCTBeHHO pacnpOCTpaHeHllIO
y 8TIIX npe.n;ROB pacnOJlOmeHbl B RpynHblx RpOBeHOCHblX cocy.n;ax
meiino-rpY]:(Hoii:, 6pIOmHoii: II 386pIOmllHHOH 06J1aCTII. Bblllle yme ro-
BOpllJlOCb 0 TOM, qTO HapJl.n;y C YCJlOil\HeHlleM 06II:\eii: opr8Hll3aU;IIII
U03BOHOqHbiX mIIBOTHLlX llHTeHCIIBHO pa3BIIBaeTCJI II YCJlomHJleTCJI
CllMnaTlflleCRaH llHHepBaU;llH opr8HOB. B neplIo.n; 3apo.n;blmeBoro pa3-
BllTIIJI CllMnaTIIqeCROH HepBHOll CHCTeMbl BLlCIlIIIX n03BOHOqHbiX BMe-
CTe C paCCeJleHneM ee RJleTOR II 06pa30BanlleM nepIIcpepllqeCRllX
Y3J10B II CnJleTeHIIII, Ha nepIIcpepllIO non8.n;8IOT II xpOMacpcpllHHLIe 3J1e-
MeHTLI, JlBJlJlIOII:\lIeCJI npOll3Bo.n;HbIIIIII Toro me Heiip03RTo.n;epM8.'ILHO-
ro 3aqaTRa. Bo BpeMH MIIrpaU;IIII OHII IIJ1II MorYT 3a.n;epmlIB8TbCH B
aaqaTRaX nepncpepnqeCRIIX CIIMnaTUqeCRIIX Y3J10B II Tor]:(a y B3pOC-
100
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l\1II KJleTKaMII (nl!-paraHrJlUII B C06CTBeHHOM CMblCJle CJlOBa), IIJlII npo-
llUKaTb BO BHYTp.mHUe opraHbI BMeCTe C HepBHblMII BOJlOKHaMII. TIa-
paraHrmlOHapHbl8 Y3eJlKII, paCCeJlIIBllIuecH BMeCTe C CIIMnaTlIqeCKOii
nepBHOii CIICTeMOn II XapaKTeplI3YIOII{IIe BbiCOKIIH 3BOJlIOIJ;IIOHHblH
JTan OpraHU3aIJ;UII CIIMnaTO-aApeHaJlOBOn CIICTeMbl, HBJlHIOTCH CTOJIb
me nporpeCCIIBHOH COCTaBHon qaCTbIO xpoMa4>4>IIHHOH CIICTeMbI, KaK
II HaAnOqeqHIIKII, II n03TOMY B 4>YHKIJ;IIOHaJlbHOM CMblCJle He MorYT
OTHOCIITbCH I, :.mCTpaaApeHaJlOBOn CIICTeMe. .
IJ;eJleC006pa3HOCTb HaqUHaTb YCJlOBHOe AeJleHlIe xpoMa4>4>IIHHOH
CIICTCMbI H"8 aApeHaJlOBYIO U 3KCTpaaApeHaJlOBYIO yme C RJlaCCa npec-
:MblKaIOII{IIXCH AIIKTyeTcH TeM 06CTOHTeJlbCTBOM, 'ITO B xOAe AaJIbHen-
llIeii 3BOJlIOIJ;IIII n03BOHOqHblX 3TII ABe TeHAeHIJ;IIII npOAOJImaIOT pa3-
BIIBaTbCH AaJlbllIe, HO, KaK Mbi YBIIAIIM HIDKe, BO B3aUMHO npOTIIBO-
nO.'IOmHbIX IIanpaBJleHIIHX. CTpoeHlIe II 4>YHKIJ;UH HaAnOqeqHblX
me.'1e3 YC.:'.IOiKHHIOTCH, II OHU IIOCTeneHHO npIIo6peTaIOT nplI3HaKII,
CBOllCTBeHHbIe opraHHblM CTpYKTypaM. B TO me BpeMH OCTaTKII BHe-
HaAnOqe'IHIJKOBOn xpoMa4>4>IIHHOH CIICTCMbI B COCYAIICTon II OKOJlOCo-
CYAIICTOll coeAIIHIITeJlbHOH TKaHII, Ha060pOT, npOAOJImaIOT TepHTb
clJYHKIJ;IIIO 3HAOKplIHHblX meJle3. STOT npOIJ;eCC, 06HapymlIBaeMblH B
3a'IaTO'IHoii <popMe elI{e Y pbl6 II no.l"yqllBllIlIll HalI60Jlee IIHTeHCIIB-
Hoe pa3BIITIIe Y 3eMHOBOAHblX, npoAO.::JmaeTCH Y npeCMblKaIOII{IIXCH.
B rrepBYIO OqepeAb, 8TO OTHOCIITCH K ma6epHblM COCYAaM, B KOTO-
pblX na6JIIOAaCTCH CHlImeHue npoAYIJ;IIpyeMblx KJleTKaMII ropMOHOB
1I.'I1I nO.'IHOe npeKpalI{eHlIe ero CIIHTe3a II HaKOnJleHIIH.
B COOTBeTCTBlI1I co CKa3aHHblM B HaCTOHlI{eH rJlaBe paCCMaTplI-
BaIOTCH ABe rpynnbl xpOMa<p<PIIHHblX opraHOB - HaAnOqeqHIIKOBbie II
BHeHaAnO'leqHIIKOBble. B nepBOH rpynne nOKa3aHbi Mop<P0JlOruqe-
n-me nplI3naKII, KOTopble CJle}J;yeT paCCMaTplIBaTb B KaqeCTBe IIpO-
rpeCCIIBHbIX, n03BOJIHIOII{ue CqlITaTb HaAnOqeqHIIKII penTlIJIlIn opraH-
IIblMII CTpYI\TypaMII, II AaHa rlICTOXlIMIIqeCKaH xapaKTeplICTIIKa KJle-
TOK, OTpamaIOlI{aH IIX meJle3IICTYIO <PYHKIJ;IIIO. B CTeHKax KpynHblx
I>pOBeHOCHbiX (II JlIIM<paTllqeCKIIX) COCYAOB onlIcaHa Mop4>OJlOrllH CO-
CYAIICTO-TKaHeBblX OTHOllieHIIH 3KCTpaaApeHaJlOBOH XpOMa<p4>IIHHOn
TKaHII II ee rIICTOXIIMII'IeCKlIe oc06eHHOCTII, KOTOpl>Ie B 06II{eM-TO
MaJlO OTJIUqaIOTCH OT TaKOBon y aM<pu6uH.
HaAnO'le'IHble meJle3b1 npeCMblKaIOlI{lIXCH, IIX aHaTOMU'IeCKOe
paCIIOJlOmeHlIe II MIIKpOCKonU'IeCKaH CTpYKTYpa onlIcaHbI B MHOro-
'lIlCJleHHblX 0pUrHHaJlbHblX pa60Tax: (Reese, 1931; Jungueira, 1944;
Retzlaff, 1949; Miller, 1952; Wright, Chester Jones, 1955,
1957, Liu, Maneely, 1959; Gabe, Martoja, 1961, 1962; Gabe, Ran-
curel, 1964; Gabe, Saint-Girons, 1964) II 0630pax (Vincent, 1897c;-
1922; Berkelbach, 1934; Hartman, Brownell, 1949; Bachmann,
1954; . Hebard, Charipper, 1955; Gabe, Martoja, Saint-Girons,
1964 I). HecMoTpH Ha AOBOJlbHO 60JlbIDOe pa3H006pa3IIe <POpMbI, pac-
I B BTOM o6sope Gabe C coaBTopaMII (1964) BMeeTCH nop;p06HaH CBop;Ka JIIITe-
paTYpld no IllOpcJ>OJIOfBlI II BM6plIOJIOfBII HaAnO'le'lHlIROB y pasHIdX OTpHP;OB
penTIIJIHll.
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a - aI<YJIa; - pbICa (HOCTlICTan); c - aMqlllfillfI (Ichthyo»his); Ii - ..lHrYIlIlia; e - J\IlI .
plfI:(a: ,- I'jJOHOgllJI; g - rOJIYOb; It - CXl1gHa; i - I<pbICa; du - aOp33.J'hHaH 30pTa:
v _ HlIIHHHH nOJIaH BCHa; ica - MCllip('(iCPlIbiC apTcpllH: cltt - xpoMadJdJ"IIuaH TH3Hb;
cl - HOpTI'II<aJIhHaH THaHh; h - nO'lHH (Chestcr Jones, 1963)
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Mil MOp«pOJIOrII'leClUIMII llpIlanaliaMII. H. IIX 'lllCJIY OTHOCIITCR COJIIII-I..l'-
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(xpOMacpcpllIIHoii, IIJIlI «Mmrolloii») II IIlITl'ppeHaJIOBOii «(IWPIWBOii»).
n:OTOpOe Ha6,1J1OJJ:aeTeR B UOJILIIICii lI,TlIl Ml'lIbIlIeH CTCnCIITI Y necx ;1..:Jl-
BOTHblX :noro IWaCCa II ROTOpOC ]-\e.rIaCT I1X CXOJJ:I1bIMIl 1I0 CTPYliTYpC
C HaJJ:IIOqeqHlmaMlI BbllllCCTORlIJ;lIX cpOpM. D MeIIblllCH CTeneHlI ;JTO
MO}KHO CKa3aTb 0 HaJJ:IIO'lC'IIJIlKaX 'IeIllY ii'IaTblX (aMeli, IIIIJ;CpllI-\"')'
Y ROTOpblX a]-\penaJIOBLIl' IiJll'Tlin ;rcnWT n OCHOllllOM na JJ:opaa.1JbllOii
CTopOHe HaJJ:110'Ie'IHlIKOB (pIle. 29, e), a n 60JIbllleii CTeneUH oTO 01'-
HOCHTCR K npCJJ:CTaBHTCJJRI\I JJ:llYX JJ:pyrnx 1I0JJ:KJIaCeOB (lIepClIaXII, I_pO-
IWJJ:IlJIbl), Y KOTOpblX IWCTO'HILll' ;JJ1e:\leIITLI aJJ:pcnaJIOBOH TI,an II
102
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HUX Kjll'TOH JlO:IJI:1JI I\OP:I<IJI!.-
HOrl (Bl'jJXlIl'ii) IIOBl'pXlIOl"TJ[
Haj\nO'II"lIlIlIia }\ IIl1' pllllhi (Lacl'r-
ta agilis). 1I0;lilTiIaH IH'ali:IUlH
no XII'.I.rIapIlY II .'\C"BI'.'I!.TY
OIL X7, ou. 10
P.. c. 31. Ila;(lIn'\("IIJJII\ HII C-
JHII\LI (Lacerta agilis), IlO, Be-
weHHwii Ha Op!.DHrKdiJw. 110-
aTHaH IIl'aK1lIIH nu XIUIJHll-'UY
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CTlIraIOT B 8TOM OTHOllieBIIII CTpoeHIIH HaAnOqeqHIII\OB nTlIq (Hart-
man, Brownell, 1949; Hcbard, Charipper, 1955; Gabe et aI.,
1964). KpoMe Toro, HaAnOqCqHble llieJIC3b1 Bcex npCCMblRaIOIqIIXCH
06JIaAaIOT cBoe06pa3Boll CIiCTeMOH BaCRYJIHplI3aqllII nOpTaJIbHoro
TlIna, a AJIH MHorlIX Xapal\TCpeH 60JIee TeCHblH 1\0HTaRT C nOJIOBbiMIi
meJIe3aMII, qeM C nOql\aMII. MCI\JIIOqCHlIe COCTaBJIHIOT Ha3eMBbIe 1JCpC-
lIaXII (Hartman, Browncll, 1949; Hcbard, Charippcr, 1955; Gabe,
Martoja, 1962; Gabe, Martoja, Saint-Cirons, 1964), Y I\OTOpblX
IIapHble HaAn01JeqHIIIill 06pa3YIOT 60JIbilllle 1\0Mnal\TBbiC MaCCbl, CIIM-
MeTpuqHO JIClliaIqIIC Ha BeHTpaJIbHOH IIOBepXHOCTII noqCI\. IIo Hallln 1
]:(aHHbIM, TeCBblll I\OBTaRT C nOqRaMIi IIMCIOT II HCROTOpblC BII]:(bl HIl.\C-
plIq. Y MOpCRIIX 1JepClIax B 60JIbllllIHCTBe CJIyqaeB HaAII01JeqHIIRII
CJIIIBaIOTCH II 06pa3YIOT OAIiB HcnapBblll opraH. Y MeCTa BbIXO,'l;a OT-
BOAJlIqIIX nOqCqBblX BCB II BAOJIb 8TIIX BeB Ba BCHTpaJIbBoii nOBCpx-
HOCTII pa36pocaHbI Y3JIbi HaAnOqeqHlIl\OBOH Tl\aHH, COCTOHIqlle 113 RY-
'Iel\ aApeHaJIOBbiX II HHTeppeHaJIOBbiX I\JICTOI\. CoeAIIHIITeJIbBOTl\aHBaH
l\ancYJIa, OTAeJIHIOIqaH HaAnoqCqHIII\H OT IIOqCI\, He BC3AC O]:(IIBaRono
BbipallieHa: B MCCTax B3alIMBoro IIpOmll\HOBeHIIH nO'ICqBOll II aApeHa-
JIOBOn Tl\aHen, OC06CBBO no XOAY IIOqCqBblX BCB, RanCYJIa OTCYTCT-
ByeT, rpaHlIqbl MClli,'l;Y OpraBaMII CTIIpaIOTCH.
Y I\pOI\OAIIJIOBblX (Rccsc, 1931; Hcbard, Charippcr, 1955; Ga-
be, Rancurel, 1964; Gabe, Martoja, Saint-Girons, 19(4) JIaAIIO-
'IeqHHI\ 01JCHb 6JII13IW IIpllMbmaeT 1\ nOJIon BeBC II nO.'IOBbIM iliC."'Ie3a 1
II TOJIbl\O B XBOCTOBOll qaCTII COnplIl\aCaCTCH C I10qRaMII IIJIII OT;J;C:ICB
OT HIIX TOBI\IIM CJIOCM PblXJIOR cocAlInlITCJIbBOn Tl\anH. IIHTCppeBaJIO-
Bble THllilI Ha]:(nOqCqHIII\OB I\pOI\OAIIJIOB paCIIOJlOlliCnbl B n;CHTpC llie.W-
3b1, a CBapYllill Ol\pYlliCBbI npcpblBlICTblM CJIOCM xpoMa<j>qnlJ!lIoii TRa-
HII, 1\0TOpaH MCCTaMII I1pOBIIl\aeT BBYTPb II IICpCnJIeTaCTCH C IIBT(,p-
peHaJIOBblMII 8JIeMeBTaMII.
Y 1JcllIyifoIaTbIX (Bcrkelbach, 1934; Rctzlaff, 1949; Bach-
mann, 1954; Wright ct aI., 1955, 1957; Gabe, l\Iartoja, 1961;
Gabe et aI., 1964; CMIITTeB, 1963, 1964) xpoMa<j><j>IIBIIaH Tl\aBb
60JIee IIJIII MeHee TOJICTblM CJIOCM paCnOJIOlliCHa Ha AOp3aJlbBOR CTO-
pOHe HaAlIoqCqHlIl\a II, I\pOMC Toro, nnJICT('lIa B cro qeHTpa."'II,HLle
yqaCTI\II, rAe o6pa3ycT cCTco6pa3BYIO CTPYI\TYPY. Y 3Mell cc 60,lhlTIC
B CcpCAIIHe, Y HIqeplIq - Ba 3aABCM I\pac mCJIe3bl (pIlC. 29, 30). Co-
CAIIHIITeJIbBOTl\aBHaH l\ancYJIa, CTpYI\Typa I\OTOpOR nOAP06no onnca-
Ha Retzlaff (1949), BbipallicBa He Be3Ae 0AlIHaI\OBO 1JCTI\O, a B
pHAe Y1JaCTI\OB COBCCM OTCYTCTByeT. IIpOCJIOHI\II COCAIIHIITCJIbBOH TR3-
BII, BXOAHIqlle B llieJIe3Y, IIHorAa AeJIHT ee Ha ]:(OJIbRII, l\alli;J;aH 113
I\OTOpbIX COAeplliIlT xpOMa<j><j>IIBHYIO II IIHTeppeHaJIOBYIO Tl\aHII.
Y 60JIbllllIHCTBa HIqepnq, no onlIcaBIIIO MBornx aBTopOB, BB.J:IIOqeq-
HIIRII He IIMeIOT HenocpeACTBeHHoro 1\0HTal\Ta C nOql\aMH, JIelliaT
aCIIMMeTplIqBO Ha nOJIOBbiX llieJIe3aX IIJIII 3al\JIIOqeBbi B mesovarium
(y raTTepIIR), npllMblRaIOT 1\ OTBOAHIqIIM nOqeqHblM BeHaM, a Y MC-
CTa IIX CJIIIJIHIIH - n R BlIlliHeR nOJIOR BeHe. OqenIiABO, Tal\Oe pac-
IIOJIOllieHlIe BCTpeqaeTCH He Y Bcex HIqepliq. Kal\ nOKa3LIBaIOT HalllII
f04
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Ma4>lfJlIIlIIan peaIuJ,JIH no XlInJlapny II XeKneJlbTY
a - HagllO'Ie'IJUIH; 6 - nO'II<a; 6 - 6plOlliHaR aopTa. OH. X 3, 06. X 10
aHHbIe, y Lacerta agilis II Agama caucasia Ha noqeqHHlm B
CEoeH rO;IOBllOii 'IaCTII naI\ 6bl nO]:(DeIliCHbl (BKJIIO'IeHbl) B 6pbnI{eii-
hY nOJIOBblX ;'1,eJle3 (pIIC. 31), a B XBoeTOBoii 06JIaCTII CMblKlJ.IOTCH
e neHTpa.'lbHOii (y L. agilis) IIJUI Me]:(lIaJIbHOll: (y Agama) nOBepx-
HOCTbIO nO'leK II OT eJIellbl OT nec TOJIbRO TOHRIIM CJIOCM RJIeTqaTRII
(pIlC. 3:!). Y L. agilis JleBblii IIa nO'IC'IHlm HCCKOJILIW aCIIMMCT-
PIl'IHO paCllo.'101IH'n Ha OTBO]:(flIl\IIX IIO'IeqHbIX BeHax, npaBblll: JIellilIT
Ha HlIlliH(,H 1I0JlOll: DCHe. fI3Y'IeHIIC ecplIiinLIx cpe30B nOKa3bIB3eT,
'ITO Y ;JTOrO Im a HIl\epIII\ 3 peH3JIOB3H 'IaCTL 113].!,IlOqeqHIIKOB 06-
P33YCT IW lIIaI{TIIYIO Macey, IWTopaH B HeeROJILIW CJIOeB paenOJIO-
a,eH3 H3 ]:(op3aJIbHOii CTOpOHC 1I{eJIe3bl II 06pa3yeT OCTpOBKII I1JIII
1I3.'IbqeBII]:(HLIe DLlpOCTbl, IIpOlII1K3I0lqlle mIYTpb llHTeppeHaJIOBOll:
n.;aHIi (pIle. 30, 35). Y Agama Op3aJIbHblll: CJIoii BblpamCH cJIa6ee,
OCTpomm xpoMacpqJlIHIIOH TRanIl pa36pocaHbi B TOJIlU,e Beell: llieJIe3bl
110 Xo].!,y OTBO HIl\IlX 1I0'IC'lnblX BCH II qaCTO npllMLlKalOT K MOqeBbIM
haHaJIbqalll. HaIIOOJIee I'PYIIIIbIC CKOII;rwnIIfl ce .TIemaT Ha Me lIaJIb-
Hoii 1I0nepXnOCTII Ha nOqe'IBIIIWB no].!, RancYJIoii II TeCHO IIpIIMblKa-
lOT h hpynnbIM cocy aM TYJIOBIIIIJ.a (pIlC. 32, 33). .lI.OJIb'IaTOe CTpoe-
HIlI' Ha nO'IC'IHbIX llieJIe3 Y 060Ilx BlI OB HIIJ.epllq lIe BbIpallieHO COB-
celli lIJIn BblpallieHO cJIa60: MClIUWCTOqHbIC npOMemYTKil npoHII3aHbi
105
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. 5... '_ . ..<) iI'" . ......
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J<pal' UlI.J,IIO'll"I1I1IJ<a. UIW IO aUJlTbi. XJlU Ia«l>qJIIIlIIiHl pl'aJ<I\JlH IIn XII:l;IiIPUY
II Xi'J<Bl'JIbTY
a - H<,gnO'le'lHHH; 6 - aO]1Ta. 01<' X 3, 06. X 10
coeAIIIIIITe"ILIIOTKaHII"' m BO.'10 1-\ lIa Mil, liOTOpble GOJIee IIJIII MeHeC paB-
HOMClHIO OnJICTaIOT OUC TI;:aun I13AUO'WQIlIII;:OB. IIMeeTcH rYCTaH CeTL
KaIIIIJIJlHpUB, pa3BeTB.'IeHIILIX II nIlAC Y3Ii:IIX JIeHT B TOJI C aW.'lC3bI.
C.lICf.\yeT UTMeTiITL CIIrTPMY opranIloro l,pOBoo6pall (,I1I1H I13AIIO-
qeQIlIIIWB y )[perMLlI..:aIO 1I X 01. A pn'!HIa,'1 "liaR I,POBL lIocTynacT J
Ha]:(IIO'Ie'lIlIII\1I 113 HeCIW!lbl>IlX IIrTO'IIIII1..:on: 1I0'IC'IIILIX apTCpllii
(a. rellalis), IIO:IOBLIX (3. Spl'I'IIHllira 11,'11I a. o\"arica), IIPTBeii
AOp3aJIbIloii aopTLI II 3HaqIITeJIbIlO pea..:e 1I3 opbmm,eeqIloii apTeplIn
(Retzlaff, 1949). llrTynuB B I1a 1I0'IC'IIIII1,. IC:II>I1C apTC)l1I1I oupa-
3YIOT 6oraTYIO ceTb TOHHIIX 1"IIII1.'lJIHpOB, t..:UTOphll' pa:!IIl'TII.'lHIOTl'H B
lIapCIIXIIMC opralla II OII.lIeT3IOT rpYlIlIbI meJIe3I1CTLIX I"ICTOK no 1I3-
IDIIM Ha6JIIO)J:cIIIIHM, CIIHYCOII]:(HLIO KalIHJIJIHpbl paCIlIHpCHIIoro Tima,
KOTOpLIC xapal\TepIlbl AJIfl IW]:(1I0'I('QIlLIX llieJIC3 l\lJIeIWllIITaIO.IIJ;IIX.
y pelITlI:mii BCTpe'IaIOTCH pl';U":O.
ITYTII IIpIlTOI,a BCII03HOii t..:pOIJII (IIOPT3.1hlIaH CIlCTCMa KpOBOOO-
pa eHJIH) C lIeKOTOpLll\1II lIapllal IIf1MII IIOBTOpHIOTCH y paalIblX OTPH-
)J:OB IIpCCMLII;:aIO I1XrH, J10 n OrJlOnliOM OrpalIll'ICliLI ABYMH UrTOqHII-
KaMII - BClIaMII TYJIOBH a II nO'I("IIILIMII. llnepnJ.le IIOpTa,TlbHaH CII-
CTeMa Gbl.'la onIlcaIla Y 3 Il'ii II 'H'pl'lIax Ecker (!S4(j) II 1I0:1i-i>l' Pet-
tit (1896) II B ]:(a.'lhlleiiUlllx I1rl'."Il' OnaIIlIfIX 1I0 TBepi-I,.'\l'lIa y nrex
nOAKJlaCCOB n OTpHAoB npecMblRaIO I1XCH (01. 0630pLI Spanner.
1929; Hartmann, UruwllclI, 1949; Hchard, Charippl'J', 19;)3;
106
Gabe et at, 1964; II MOHorpacimlO Chester Jones, 1957). Y JIII e-
pHI (Ray, 1936; Wright, Chester Jones, 1957) BeH03HaH KpOBb
nocTynaeT B Ha nOqeqHIIKII II3 Mempe6epHbix (naplIeTaJIbHblx) II cnlI-
lIaJIbHblX .BeH, npoxO HIl\IIX B6JIII3II n03BOHROB. Y raTTeplIll (O'Do-
noghue, 1921) nplIHOCHlI(aH BeHa Ha nOqeqHIIKOB npOIICXO IIT II3
KpaHlIaJIbHOll qaCTII nOqeK II, KpoMe Toro, nOJIyqaeT KpOBb II3 TYJIO-
BIIll\HbIX BeH. BeHbI BCTynalOT B Ha nOqeqHbIe meJIe3b1 II B HIIX pa3-
BeTBJIHIOTCH Ha KanlIJIJIHpbl. MCTeKalOll\aH II3 Ha nOqeqHIIROB KpOBb
lIocTynaeT B 3<p<pepeHTHbie BeHbI nOqeK IIJIII HeIIOCpep;CTBeHHO B HlIm-
HIOIO nOJIYIO BeHY.
Y npeCMbIKalOlI(IIXCH nOpTaJIbHYIO CIICTeMY BaCKYJIHpII3aI{1I1I IIMe-
lOT TPII opraHa: Ha nOqeqHIIKII, IIOqKII II neqeHb (Spanner 1929).
Y MJIeKOnIITalOll\IIX mlIBOTHbiX II qeJIOBeKa OHa coxpaHHeTCH TOJIbKO
B neqeHII, Tor a KaK JIH Ha nOqeqHIIKOB TaKoe cBoe06pa3IIe KpO-
Bo06palI(eHIIH HBJIHeTCH oc06eHHOCTblO npeCMbIKaIOII(IIXCH. B Ha no-
IeqHIIKaX nTIII{ OHa BCTpeqaeTCJI B pe YI{IIpOBaHHOM B e (Hart-
man, Brownell, 1949).
Mbi He BCTpe'laJIII pa60T, CIIeI{IIaJIbHO nocBHlI(eHHblx BblJlCHeHHIO
<p1I3IIOJIOrlIqeCKOrO 3HaqeHIIJI nOpTaJIbHOll CIICTeMbl Ha nOqeqHIIKOB
npecMblKalOlI(lIXCJI. O HaKO caM <paKT CTOJIb YCJIOmHeHHoll CIICTeMbl
BacKYJIHpII3aI{1I1I KOMnJIeKCa meJIe3IICTblX cynpapeHaJIOBbiX KJIeTOK,
KOTopble B npoI{ecce IICTOplI'IeCKOrO pa3BIITIIH cocpe OTOqllJIlICb B
XBOCTOBOD: 06JIaCTII TYJIOBlIlI(a, crpYIInlIpOBaJIIICb BMeCTe C IIHTeppe-
HaJIOBOll TKaHblO II BO MHornx CJIyqaHX IIHKancYJIlIpoBaJIIICb II oT]:(e-
JIIIJIIICb OT coce HlIx opraHoB II TKaHell, CB eTeJIbCTByeT 0 nplIcno-
C06IITeJIbHOM 3Ha'leHIIII TaKoll CIICTeMbi KpOBo06palI(eHIIH. CHa6iKeHIIe
JII060ro opraHa KpOBblO, a BMeCTe CHell IIIITaTeJIbHblMII BelI(eCTBaMII
II KIICJIOpO OM, KOTopble 06eCIIeqlIBalOT HOpMaJIbHblll 06MeH B KJIeT-
Rax, .'le1l\IIT B oeHOBe ero ClIeI{II<plIqeCKOll <PYHKI{IIII. OpraH1I3an:IIH
ClleTeMbl RpOBoo6pall\eHIIH, RaK npaBII.'IO, yeJIOmHfleTefl napaJIJIC.TIbHO
YBeJIlPIPHlllO TOM pOJIlI, KOTOpylO opraH nplIo6peTaCT B npoqecee pa3-
BIITIIH 1I KOTopafl He06xo]:(IIMa JIH no.rIHoqeHHoro <PYHRqlIoHlIpoBa-
HIIH Beex PyrlIx elICTeM opraHlI3Ma. IIHblMII e.'10Ba1\1II, CTPYKTypHaH
()praHlI3aqllH CIICTeMbi KpOBOCHa6meHIIH HBJIHeTCH O]:(HIIM 1I3 nplI-
cnOC06IITeJIbHblX, Bblpa60TaHHblX B 3BOJIIOI{IIII MexaHII3MOB, 06ecne-
'UlBaIOIl\IIX HOpMaJIbHYIO mII3He eHTeJIbHOCTb mlIBOTHoro. ECJIII
Y'IeCTb, 'ITO KOMnJIeKC cynpapeHa.lIOBblX 3H OKplInHbIX MeTOK, KOTO-
pbIe cocpe OTOqlIJIIICb B6JIII3II nOqeK n nplI06peJIII opraHHblll TIIII
CTpoeHIIH, HBJIHeTCH e]:(IIHCTBeHHblM TaKoro po a 06pa30BaHIIeM B
TCJIe npeCMblKaIOIl\IIXCH, TO CJIe yeT yMaTb, qTO IIMeHHO eMY IIplI-
Ha JIellilIT BbinOJIHeHlIe HalI60JIee aKTIIBHoll <PYHKI{IIII 3H OKpHHHOll
llie.'Ie3bl II nplI TOM <PYHKqlIlI, 60JIee CJIOlliHOii no CBOIIM 3a aqaM,
'1eM Ta, KOTOPYIO OHII BblllOJIHHJIII Ha 60JIee nplIMIITIIBHOM ypoBHe
CBoeII TKaHeBOll opraHII3aI{lIlI.
l\aK 1I3BeCTHO, IIHHepBaqlIH JII060ro opraHa II ero COCY IICToro
PYCJIa HBJIHeTCH PyrlIM BamHeHmIIM MexaHII3MOM, 06eCneqlIBalOII(IIM
npoqecebl IIX cneqlI<pnqeCKOrO <PYHKqnoHlIpoBaHIIH. MHHepBaqlIoH-
HbIe nplI60pbl Ha CTeHKax KpoBeHocHLIX COCY OB HBJIHIOTCH neplI<pe-
f07
8, 82 V, 83
A
L
SA
(J (JIJ IJA
a;y 85 84 8,
8;;m.C'
8A
IJA
(JIJ
(J Ad A In v.C'
PH C. 34. CxeIIIId nHHepBal\lIII Hal{nO'le'lHJIROB RpOROAIIJIa
8, - 8. - c nepBoro no nHThlt1: CHMnaTH'lecKHe HepBU, 8ym. C - CHHanTH'leCKalI I1CnO'lK8;
Vi - V. - BeHhl KapAHHaJIbHhle, LV, - nepBut1: JIyM(jaJIbHhlt1: n03BoHoK; 8A - CeI'MeH-
T8pHaH apTepHII; A - apTepllH; In. V. C.- vena cava inferio; G - rOHaJJ;a; Ad - HaA-
nO'le'lHHH; GD - l'eHHTaJIbHhlt1: TpaHT; DA - AOpaaJIbH81I aopT8 (L8wton, 1937)
plIqeCKHMII 1JaCTHMII TeX HepBHblX qeneii, KOTOpble OCYIqeCTB.!IHIOT
pe<j>.!IeKTOpHble aKTbI II perYJIlIpYIOT BCIO CJIO}ImylO p;IIHaMIIKY KpO-
BOTOKa B OpraHaX. HepBHble annapaTbi Hap;II01JeqHIIKOB Y npeCMbl-
KaIOIqIIXCH II IIX qeHTpaJIbHble CBH3II MaJIO II3yqeHbI. CHMlIaTlIqeCKIIe
RJIeTKII H MeJIKHe raHrJIII03Hble CKOnJIeHIIH onncaHbI no cocep;CTBY C
Hap;n01JeqHIIKaMH, nop; KancYJIoii II BHYTPH meJIe3b1 (ROJIOCOB, 1930;
Swinjard, 1937).
B pa60Te Lawton (1937), B KOTOpOH H3yqeHbI MaKpo- II MHl\pO-
CKOnlI1JeCKlIe qeHTpaJIbHble CBH3II HepBHblX lJIeMeHTOB Hap;nOqCqHlI-
Ka KpOKO]:(IIJIOB, nOKa3aHa cerMeHTapHaH HHHepBaqlIH ll3 CIIMnaTII-
qeCKIIX Y3JIOB, OCYIqeCTBJIHeMaH HeCKOJIbKHMII HepBHblMII CTBOJIal\1II
(pHC. 34). Ha MeCTe 3TIIX CTBOJIOB Y MJIeKOnlITaIOIqIIX pa3BIIBaeTCH
06IqHH CIIMIIaTlIqeCKIIH CTBOJI B BHp;e 1JpeBHOro HepBa. IIop;06HaH
Mop<j>OJIOrlf'leCKaH opraHII3aqlIH HepBHoro cHa6meHIIH Hap;nOqeqHII-
KOB peIITlIJIlIH OTpamaeT, BIIP;HMO, npoMemYTOqHblii 3Tall 1f>0PMHPO-
fOB
BaHHH IIHHepBar(HOHHblX OTHOllleHHii B PH Y n03BOHOqHbIX. Pa6oT8
Lawton IIpe CTaBJIHeT IIHTepeC eII:\e II B TOM OTHOllleHHH, '1TO an-
TOp onlIcaJI norpyiKeHlIe OTpOCTROB HepBHblX RJIeTOR CIIMnaTII'IeCRIIX
raHrJIlIeB B M03rOBYIO qaCTb Ha]:(nO'leqHIIIWB II c eJIaJI 3aRJIIOqeHHe
06 IIHHepBar(IIH xpoMa«jl«jlHHHOll: TRaHII nOCTraHrJIHOHapHblMH BOJIOR-
HaMH (He JIHIlIIIB ee, R COlliaJIeHHlO, cHa6llieHHH II nperaHrJIIIOHapa-
MH). HanOMHIIM, '1TO aHaJIoruqHbIe RapTHHbI Ha6JIIO aJI Young (1933)
y aRYJIbi (CM. plIC. 13). OTII 06a Ha6JIIO eHIIH roBOpHT IIpOTUB TOH
TeoplIU, ROTopaH paCCMaTplIBaeT xpoMa«jl«jluHHble RJIeTRII B RaqeCTBe
BToporo neplI«jIeplIqeCROrO HeHpoHa CHMnaTlIqeCROH HepBHoH CH-
CTeMbi.
OnlIcaHHble MaRpo- II MHRpOCROnH'leCRlIe oc06eHHocTII BaCRYJIH.
pH3aqIIII II HHHepBar(HII Ha nOqeqHIIROB penTIIJIIIll: xapaRTeplI3YIOT IIX
opraHHYIO CTPYRTYPY. HalI60JIee nOJIHOe npe CTaBJIeHlIe 06 3H 0-
RplIHHOH iKeJIe3IICTOH «jIYHRr(IIII 3TIIX opraHoB aeT H3Y'IeHlIe r(IITOJIO
rim II qlIToXIIMlIqeCRHX CBOHCTB llieJIe3IICTblX napeHxHMaT03HbI:1.
RJIeTOR.
no Mop«jlOJIOrlIqeCRHM IIplI3HaRaM xpoMa«jl«jllIHHble RJIeTRII HaAno-
'1eqHIIROB penTIIJIIIH He OTJIlIqalOTCH OT RJIeTOR M03rOBoro CJIOJl Ha
IIOqeqHIIROB MJIeROIIIITalOII:\IIX. OHII HMelOT 1I0JIIIrOHaJIbHYIO, Ry6H'1e-
CRYIO, BblTHHYTYIO IIJIII HenpaBIIJIbHylO «jIOPMY (pIIC. 35), JIlImeHbI,
RaR npaBIIJIO, OTpOCTROB, COAepiKaT RpyrJIOe HJIII CJIerRa OBaJIbHOe
HAPO C He pe3RO BbipallieHHblMH, cJIa60 oRpallleHHblMH rJIbl6RaMII xpo-
MaTIIHa II 1-5 HAPblIllRaMII. IJ;HTOnJIa3Ma 6e Ha PHR-3epHaMH (cJIa-
60 nHpoHIIHo«jlIIJIbHa) II OTqeTJIIIBO 3epHHcTa.
XpoMa«jl«jllIHHaH peaRr(HH HBJIHeTCH Mop«jlOJIOrlIqeCRHM Bblpallie-
HlIeM npOAYRTIIBHblX II ceRpeTopHblx npor(eccoB, npOTeRalOII:\IlX B qH..
TOnJIa3Me RJIeTOR, II, CJIeAOBaTeJIbHO, MOllieT CJIYlliHTb RplITeplIeM .AJIJI
Or(eHRII «jIYHRqlIoHaJIbHDrO COCTOHHHH RJIeTOR: B neplIoA IIHTeHCIIBHO-
ro ropMoHoo6pa30BaHHH OHII nepenOJIHHIOTCH HpRO oRpallleHHblMH n
llieJITblH qBeT xpoMa«jl«jllIHHblMII 3epHaMII II onYCTOllialOTCH OT HIIX B
nepHoA BbiAeJIeHHH HHRpeTa. CaMbiM xapaRTepHblM nplI3HaRoM JIIO-
60ll: llieJIe3IICTOll: RJIeTRII HBJIHeTCH D;IIRJIlIqeCRaH ceRpeTopHaH Aeh-
TeJIbHOCTb. ]];JIH Toro qT06b1 y6eAHTbcH B ee HaJIHqHH B xpoMa«jl«jllIll-
HbiX RJIeTRaX HaAIIOqeqHHROB penTIIJIIIH II BblHBIITb IIHaMHRY 3HAO-
RpIIHHOll: aRTHBHOCTII, HaMil 6b1JIH 06CJIe OBaHbI HII:\eplIn:bI (Lacerta
agilis H Agama caucasis) B BeCeHHe-JIeTHIIll: neplIoA (Mall:, IIIOHb) ,
RorAa llilIBOTHbie HaXOAHTCH B aRTIIBHOM COCTOHHIIII, II B 3HMHHH
(AeRa6pb) - BO BpeMH CnHqRII.
Mop«jlOJIOrIIqeCRIIMII TeCTaMII JIH onpeAeJIeHHH «jIYHRD;IIOHaJIbHO-
ro COCTOHHIIH RJIeTOR CJIYlliIlJIII: CTeneHb HanOJIHeHIIH D;IITOnJIa3Mbl
3epHaMII ceRpeTa II CTpyRTypa ROMnJIeRCa rOJIbAllilI, yqaCTlIe ROTO-
poro B «jIopMIIpoBaHIIII ceRpeTa xpOMacp«jlIIHHOll: RJIeTROH Y MJIeROnlI-
ralOII:\IIX nORa3aHbI de Robertis, Sabatini (1960), H. A. CMIITTen
(1965 a, 6; 1966 6, B, 1967), H. Jr. IOIlIReBHq (1966 a, 6) II
B. B. .HrJIOBbiM (1966 a).
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Ha CTa]:(UII BLI]:(eJleHIIH, CUHTe3a UJiU HaROIIJleHUJI ropMOHOB (pUC. 37,
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peTOpHoii aKTIlBHOCTU Iw:eTOK (puc. 38). BLlHCHUTb RapTIIHY KOMII-
JIeKCa fOJlb]:(mu B xpoMa4>«lJIIHHblx KJleTRaX BO BpeMH 3UMHen CIIHqRII
J{OCTYIIHbIMU MeTop;aMH CBeTOBon MUKPOCKOIIUU HaM He y]:(aJlOCb, TaK
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lIIeHTanbHOII pa60Te Biscardi, Wassermann ( 1966) Ha Ha]:(nOqeq-
HllRaX 3Meii: (Xenodon merremii). Ha OCHOBaHHH O]:(HOTHnHoro xa-
paKTepa peaK HII KneTOK Ha BBe]:(eHHe HHKOTHHa, CTHMynHpylOIII;erO'
BLI]:(eneHHe KaTexonaMIIHOB, aBTopbl rrpe]:(1I0naralOT CXO]:(CTBO H B Me-
xaHH3Max Bbl)J;eneHIIH 060HX ropMOHOB.
ITO]:(BO)J;H HTor Bonpocy 0 KneTOqHOM ]:(HMop H3Me B Ha]:(nO'leqHII-
I.ax peIlTlInHII, MomHO CKa3aTb, qTO KneTRII B 06LI lHbIX ycnoBHHX
cO)J;epmaT Hopa]:(peHanHH II, 110 MHeHHIO 60nblllllHCTBa aBTopOB, HMe-
lOT ]:(Ba xapaRTepHblX, TonbKO IIM CBoii:cTBeHHbIX Mop OnOrll IeCKIIX
npH3HaKa (BenH'lHHa KneTOK II BenHqlIHa rpaHyn), He OTnH IaIOTCH no
CBoeMY XIIMHqeCKOMY COCTaBY OT a]:(peHanHHCO)J;epmaIII;HX II, B03Mom-
HO, MoryT B onpe]:(eneHHblX ycnoBIIHX nepeXO]:(IITb B MeTlInHpOBaHHble
OpMbI.
ITo cpaBHeHHIO C ]:(aHHbIMII no 1I3Y IeHIIIO Ha]:(nOqeqHHKOB peIITH-
nHii:, B nnTepaType MeHbllle lIMe eTCH CBe;o;eHHii: 0 TOii: xpOMa IIHHoii:
TKaHII, ROTopaH nH60 06pa3yeT c06CTBeHHo naparaHrnHII II pacnono-
meHa B Henocpe]:(cTBeHHoM coce]:(cTBe C Y3naMH cHMnaTHqeCKoii: HepB-
HOii: CHCTeMbi (IIorpaHlIqHoii: eIIOqRII H 6plOlliHoro CIIneTeHIIH), nH60
OTHOCHTCH K TeM xpoMa «lJHHHblM KOMlIneKcaM, KOTopble 3anomeHbI B
RpoBeHoCHblX cocy]:(ax II BXO]:(HT B CTPYRTypHblii: COCTaB IIX CTeHOK.
Hall60nee CTapble CBe]:(eHHH 0 BHYTPHCOCy)J;HCTblX CKOlIneHIIHX
xpOMa IIHHbIX aneMeHTOB OTHOCHTCH K KpOBeHocHoii: CHCTeMe Tyno-
BIIIII;a, r]:(e, no Ha6nlO]:(eHHHM, c]:(enaHHbIM enw Wiesel (1902a) H 1103-
me Hartman, Brownel (1949), rpYIInbi xpoMa «lJHHHblx RneTOK
nplICYTCTBYIOT B MarllCTpanbHblX KpOBeHOCHbiX cocy;o;ax 6plOllIHoii: 06-
naCTH II 3a6plOwHHHoro IIpOCTpaHCTBa (cnlIHHaH aopTa, 3a]:(WIH 110-
naH BeHa), B HX pa3BeTBneHIIHX, a TaRme B cocY)J;ax 6n:asnemaIII;HX
opraHOB (IIOqRaX, rOHa]:(ax) . 113yqeHHIO xpOMa IIHHoii: TKaHH B
rpY]:(HbiX II weii:Hblx cocy]:(ax pellTlInHii:, HBnHIOIII;HXCH IIpOII3BO)J;HbIMJI.
ma6epHbiX )J;yr, 1I0CBHIII;eHO MHOrO pa60T, B TOM qHCJIe C IICIIOnb30-
BaHlIeM COBpeMeHHblX MeTO]:(OB rHCTOnOruqeCKOro aHanlI3a (Trinci.
1909, 1912; Reese, 1914; Terni, 1931; Palme, 1934; Ask-Upmark.
1935; Adams, 1939, 1952, 1953, 1957a, 1958; Adams, Pickerscill.
Underwood, 1957; Rogers, 1967; Sidky, 1967).
Y 1I3yqeHHblX HaMIl ]:(BYX DH]:(OB HIII;epH (Lacerta agilis II
Agallia caucasia) YCTaHOBneHO Ha ceplIii:Hblx cpe3ax paCIIpOCTpaHe-
HHe xpoMa HHHbIX aneMeHTOB n CTeHKax KpOBeHOCHbIX COCY]:(OB TY-
nOBIIm;a. B aToii: pa60Te BblHBneHbl nOKanHaa IIH a]:(peHanHH- II HOp-
a]:(peHanHHco]:(epmaIII;HX KJIeTOK H HanH'llle B HIIX ceKpeTopHoii: aKTIIB-
HOCTII.
B OTpeaKax ]:(op3anbHoii: aopTbl, KOTOpbIe conpHKacalOTCH C Ha]:(IIO-
qeqHHKaMII HnII IIPOXO]:(HT pH]:(OM C HHMH, xpoMa HHHbIe KneTKH 06-
paaYIOT IIpepbIBHCTYIO, a IIHor]:(a cnnOIliHYIO neHTY B cpe]:(Heii: II Ha-
pymHOii: 060nOqKaX cocy]:(a (CM. plIC. 41, B). ITo Mepe y]:(aneHHH OT
Ha]:(IIOqe IHIIKOB H npH6nlImeHIIH K KpaHlIanbHOMY KOH y TynoBHIII;a,
r)J;e paClIonomeHbl aopTanbHbIe ]:(yrll, KOnH IeCTBO xpoMa HHHbIX
::MCl\leHTOB CHlImaeTCH, H OHH 06paaYlOT a]:(eCb MenKlIe OCTpOBKII. Co-
119
OCTaBneHile IIpenapaTOB, 06pa60-
TaHHblX 6HXpOMaTOM, C TeMlI, ROTO-
pble nOABepraJIIICb HO]:(HpOBaHIIIO,
II03BOJIJleT CR8saTb, qTO B aopTe
IIMeIOTCJI 06a BIIAa RJIeTOR H qTO
60JIbmee ROJIH'IeCTBO Hopa]:(peHaJIHH-
COAepm81IJ;IIX HaxOAHTCH B I'pYAHOM
.VTpe3Re AOp3aJIbHOii aopTbI, a TaR-
me B HIICXOAHIIWX qaCTHX ee ]:(yr
(pIlC. 41, B). B aTOM OTHOmeJIHH
Hallill ;!I;aHHble COBna]:(aIOT C Ha6JIIO-
J];eHIIHMII Muratori C COTpY;lJ;HilKaMH
( 1965a, b) . ABTOpbl npOBeJIH cpaBHil-
TeJIbHOe IICCne;lJ;OBaHHe xpOMa.p.pIIH-
HOD TRaHII, paCIIOJIOmeHHOii B pas-
HbiX yqacTKax TeJIa Lacerta muralis,
HCnOJlbSYJI PHA cne HaJIbHblX rHCTO-
XilMlIqeCRIIX MeTO;lJ;OB: cTapblii MeTO;!!;
X:pOMlIpOBaHHH Henle, IIO;lJ;HpOBaHHe
110 Hillarp II Hokfelt, cepe6peHIIe
no Bodian II .pJIyopec eH HIO no
Eranko II Falck. OHil nplIIIIJIH R
BblBOAY 1.. qTO HaH60JIbmee ROJIHqe-
CTBO Hopa]:(peHaJIIIHco]:(epmaIIJ;HX
RJIeTOR HaXO]:(IITCH B nerOqHO-aOp-
TanbHoii 06naCTH.
B 8pTeplIHX pa3Horo RaJIH6pa-
OT RPYIIHblX ]:(0 MeJIRHX apTepHOJI,-
06a BII]:(a RJIeTOR, RaR npaBlIno, TO
me HaXO]:(HTCJI B HapymHoM II cpe]:(-
HeM CJIOHX COCY]:(IICTOii CTeHRH HJIil
JIemaT B oRpymaIOlIJ;eii cocy]:( coe]:(H-
HIITeJIbHOD TKaHH (pHC. 42, 44) II
3HaqHTeJIbHO peme saXO]:(HT BO
BHYTpeHHHD CJIOD (pilC. 43).
B COCTaBe CTeHOR BeHo3Horo
pyCJIa Mbi TaKme Ha6JIIO]:(aJIH a]:(pe-
HaJIHM- II Hopa]:(peHaJIHHco]:(epma-
IIJ;lIe KJIeTRH, KOTopble paclIpe]:(eJIe- .
Hbl BO Bcex Tpex CJIOHX COCY;lJ;D.
O]:(HaKo HaH60JIee xapaRTepHoD IIX
nOKaJIHSaJweD HBJIHeTCH tunica in-
tima II 6JIH30CTb C aH]:(OTenHeM. B
HHmHeD nOJIOD BeHe 06HapymeHbi
.f50JlbllllIe ROJIHqeCTBa EJIeTOK, co-
}l;epmalIJ;ilx Hopa]:(peHaJIHH, ROTOpbIe
CTenHTCH B;lJ;onb cocy]:(a II MeCTaMH
BblcTynaIOT B ero IIpOCBeT (pHC. 45) .
120
P u c. 41. CxeMa (A) cTpoeHlIH
cepp;e'lHo-cocYP;IICTOH CIICTeMLI pe-
nTIIJIIIH. HIIH\HUH IIpHMoyrOJIb-
linK - yqaCTOK aopTId Ha ypOBHe
IIO'leK, BepXHIIH IIpHMoyrOJIbHIIK--
Y'laCTOK HlIcxop;H eH 'laCTII p;yrll
aOpTld
B - aopTa Ha ypoBHe nO'iHH (8) H Hag-
nO'le'lHHHa (6) (CM. HllHUIHiI: npHMoy-
1'0JIbHHK Ha cxeMe A): 11 - xpoMadJljJHH-
HbIe KJIeTHH B tunica media B nepHoA
allMHet1: CllH'lHH HIIIepllIU>! (Agama cauea-
sis). XpoMaljJljJHHHaH peaKIIHH no XuJl-
JIapny H XeKBeJIbTY. OK. X 3.5, oi'i.X8;
B - H" xogHlI\aH 'laCTb AYI'U aopTH (a)
(CM. BepxHut1: npHMOYI'OJIbHHK Ha cxeMe
A): 6----:HOpagpeHaJIHHcogepmalI\He KJIeT-
HH B tunica media II B tunica intima.
IIoAaTHaH peaHI\HH no XIIJIJfapny H
XeKBeJIbTY. OK X 3,5. oi'i. X 8
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a - D tunica adventicia MCJIIiIlX aI1Tcl1J1ii; (j - D COCgllllJlTCJIbHOfi 1'1;8111I 3aUl11OmJlHlIOm
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(Lacl'rta agilis) B Ul'pJlOP; 311111lIeii cnwJ.Rl'I. Hopap;peHaJlIIHCO;u;epmaU{IlC IUICTKII
B CTenKe cocyp;a. IlO,'WTlIaH peam.\IUI no XIIJlJIapny II XeKBeJILTY
a - npOCBcT IIammJIfIpa; 6 - npocDcT OTBOnRII\ct1: nO'le'IHofi ncnbI; 6 - HopanpCHaJIIfH-
cOAcpmall\IIC I<JIL'THIf. UH. X 5, 06. X 20.
B BCUaX oonee IIIl'JlI'-OrO I-;aJIH6pa, }J.peHIlpYlOIIJ,IIX IipOBb 113 opraHOB,-
nO'Il'I, (pIlC. 4G), IW}J.IJO'Il"IIUUWB (pIlC. 47), II pa3BeTBJIemIflX napne-
Ta:JbllblX cocyp;on II }J.pyrnx ou.rracTeii, 8TOT nplIHn;Hn pacnpl',o:enemIfl
xpmW(PwnUHbIX 8JIeMeHTOn BUJIH311 npOCBeTa COC.Y}J.OB coxpaHHl'TCH.
Do;:\ 8H}J.OTeJIIIeM RanllJIJIHpOn IIa6JIlO}J.al'TCH BbIcTynalOIIJ,ee B llX npo-
CBOT coe}J.IInHTeJIbHOTKaHHoe YTOJIIII.enlIe, n IWTOpOM JIemaT l{pynHhle
3}J.peHaJIlIII- II Hopap;pl'IIamIllCOi l'piI\i1I1 IIe l.;JIl'TKII (pIIC. 48, 49).
1I3Y'Il'HIre ceKpl'Topnoii aHTIIBIIOCTH xpOMaWWHHHbIX KJIeTOK, HO-
TOpblC JIORaJIH30BanbI B CTClmax KponeHOCHblX COCY}J.OB, npOBe}J.eHO
Ha M:'lTl'pnaJIe, B3HTOM B pa3nbIe Cl'30nhl rO}J.a, H nOJIY'lCnbl pe3YJIbTa-
TbI, anaJIOfIl'Inble Tel\l, KOTopble onIIcaHbI n Ha}J.no're'IIIIIKax. Y HIIJ,e-
pun; n 3Hl\IHIIii uepuo}J., C.oupOBom}J.alOIIJ,IIHCH OTHOCUTl'JIbllbII\I nOKoeM
II na}J.eHIIel\l ml'.rre3IlCToii aKTIIBnOCTII, na6JIIO}J.aeTCH CHlImeHlIe BbI}J.e-
.rremIfl ropMOHOll II nepenOJIlICUlle IIMH n;IITOnJIa3Mbl ImeTOK (pHC. 41,
E, B, 45, 47, 48). B BeCeHHe-JIeTHHH lIepHO}J. RapTHHa Ml'UHeTCH II
B liJIeTI,aX M01IiIlO BII}J.eTb ncl' CTa}J.IlII OCllOOO1K}J.l'nIlH cel,pl'Ta (pnc. 42,
44,50,51).
HalllIl }J.alIlibIC, nOJlY'ICllIIbIe Ha HIIJ,l'plIn;aX, IJOIia3bIBalOT , 'ITO n
HID-KHell II Cpl'}J.IIeii: 'IaCTHX TYJIOBHI:u;a CKOIlJIemlH XpOMaWWUHHbIX
K.iIeTOR nCTpe'IaIOTCH B cocY}J.ax BCeX KaJIH6pOB. B cocY}J.ax apTepH-
a:IbHOrO Tllna OIIII, RaIi npammo, paCnOJIOmenbl n HapymHoM II rpe}J.-
125
HOI\I CJlOHX COCYJ-\IICTOll CTeru>U II 11 Of\pYII,afOllWll COCJ-\lllillTCJlbliOii
Tl,aIlIf; II COeT3Ul' ;JTUX C1\OII.'Il'1I1I1l IIaXOJ-\HTOl IVICTI.ar, l'O;\l'pn.;all\I11'
I,m, aJ-\pc WI.'II1 II , Tar.; If llopaJ-\pPlla:lIlII. I JIfl COl'y.[\on nCII03110ro Tlllla
XapaKTl'pllO HOnOl'pl'.[\CTllellIlOp 1I(H1!1PJRfllllll' 1\:11'1'01'; OOOllX IIII OIl I,
'IPOCBOTY cocy.[\a II TOCllafl CIIJI.II, IlX l" :HI]l,OTCJlIII'M. no lCIJl' lIpllu:m-
;{WHIfH COCyp;OB I'; llopep)wii 't:H'1'1I TY.'lonmll,a !,O;I1I'Il'CTIIO X pO !\I <HfJ-
«pIIIIIlLIX OCTpOlll,OIl IIap,aeT II l'iI:mop I1X Y:lll'll('IIIal'1'CH.
3al\<'III'IIIBaH jl<ICCI\IOTIJCIlI\(' COCY;\lfCTO-Tl-<allpIILIX OTIIOHICIIII ii C
xpmralj)(IIllIlIILIIIIII J:ICIIfCHTa:l1H II OGJI3l'TII Ty.-IOIIIIII\a, Y:lICCTIlO Yllo:\lH-
'"
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IiI' MI'JlliOH' cucyp;a JJUleplll\hI
(Lacl'rtil ilgilis). I10Jla!l.Pl'lIiI-
JIIIIICO;ll'p,liaJlllle liJleTl,1I B CTl'II-
lil' \"a a \"(1:-:0("U111. lIoAaTIHul
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enH3aII C 1I0ii COCJUIUIITO.TII,II01"l-<aIlill,11II1I 1I0jl"'II,IIII"HIII) 'Il'(Jl'lIilX X[10-
M3<plI1I1I1Il1010 I,JU'TIm. OOllapymPJlIIMc II. 1\. J\o.1pl'I1m.;olloii rpYIIIILI
KJIOTOI, IIpnuajVIPIl.;aT HOCOI\1Hl'llIlO I-< CllCTl'''p aOfJTa:lbllblX IIlIl'lIa:l,IIO-
'IO'[llIU,OIlLIX IWl\fII.'J.l'I';Con xpol\la(IJlIIIIIlIII.I\: ;);II'IIIl'II1'OII.
COCY.D;LI maoPpIIoii oi'J.-raCTII IIl)('Cl\Ibll.;aIOIl\IIXCH OOCJW;IOBalllol li:l
HOIIIIlOrnX nppp,l'TaBIIH'."'IHX ;JToro I';.TIaCCH, r:la 1111 (,1:11 oopa o 1 n oii.-w-
CTII I{apOTnp'HbIX ;.\yr. .\opTa.'ILllhJ(' II ;ICrO'IIII.U' .QTII II;IY'll'lIbl 'IellL-
llIe. HalluQ.TIee no,o:poUHO onnCaIIbJ GIHlly!H,al\IIH ('oulIoii a)Hl')HIII II
KapOTIlp;nbIll CIIIIYC Y H eplJI (Palmo, t!I::.'t; A(lams, He):\: Sidky,
1967; Hogol's, 19(7). B TOM IIlenC, ]WTOPOl' COOTEI'TCTIIYCT fimIJypl,a-
n;IIII 06Ill,eii cOllIIOii apTepulI IIIJIeIWIIIITaIOIll,IIX II r;:(o Y UIIX lIaXO;!,IIT-
CH KapOTIr uoo TOJIO, Y Hlll,epIl paCIIOJIOil';PIIH l.;apOTIl;tIl'UI ;!,yra
(puc. 52). HOCIW.'ILIW OTBOpcTlIii, IWTopLle COP;!,IIIIHlUT n!lYTpClIIIfOlO
COIIHYIO apTl'pIlIO C l.;apoTII.[\HOii ,1 ...oii, II IIX pa311hlO pa:mop),[ IIPII-
.[\alOT ;noii 00.1aCTII ('hapoTII;:J.IIOIlIY ('IIllYCY'») JlaOllplIllTOIIO;:J.oollI,llr
nu.[\. Y napanOB II XallreJIeOHOB (mapoTn,o:IIblii ('IIlIYC» COCTOIlT 113 p'BYX
II)WTOI';OIl, COP,o:IHII'IlIILIX ll'iI,,o:y couoii II cjJ(IJOjJ II pOl\a!lll LUIII OTllCpCTU-
H:l1II (Adams, J 95 , 1957b). Y :ml'ii l\Il'CTO )liI;I;tB()('I1lUI ('oUIIoii apTO-
purr uo a:l,u rO."'IOBLI C'lIlTaCTrJI 1'011I0:101'11'111(,1 I omIlypl.;a I IIII 11I:101>0-
IIUTalOll\IIX (Ask-U)lmark. 1935; Boyd, 19q ). Y 'll'ppnaXOULLX (Ter-
ni, 193J) II I,POIW:I,nJIOBLIX (Rcpso, 1914; Torni, H131) ;na OUJIaCTL,
126
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Hoii IIPllhl TY.'IOIJIIII(a nIl epIIl(hl
(Lacerta agili,:) B JIeTlIIIii ne-
puo/(. ACIIlIxIHIIIHan aliTIIJlUOCTh
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lI.un no XIIJIJHlPUY II XCIWl'JIhTY
a - IIHTCIIt_-"UDIIO onp:un IIHbJ(' H. r(>T-
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(Agama cauca"is) II .'It'Tllllli IIl'-
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lp
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peHHell COHHOll apTepIIII OT 6e3b1MHHHOll.
Ha CTeHKe BHYTpeHHell COHHOll apTepll1l II IIpllJIema ero Y'laCT-
Ra P,yrII Y H eplID; OIIlIcaHO Me30p,epMaJIbHOe YTOJI eHlIe, IIpOHII3aH-
HOe RalIlIJIJIHpaMII (Adallis et aI., 1957). CIIeD;lIaJIbHblX IIHKaIICYJIH-
pOBaHHblX 06pa30BaHIIll B aTOM MeCTe, KOTopble MomHO 6bl 6blJI0 BbI-
]:(eJIIITb B KaqeCTBe caMOCTOHTeJIbHblX CTPYRTYP, Ha1I0]:(0611e RapoTIIp,-
Horo TeJIa MJIeKOIIIITalO lIx II IITIID;, 3p,ecb HeT _(Adams, 1958). Ho OT-
]:(eJIbHbIe KJIeTOqHble rHe3p,a, rOMOJIOrllqHbIe TeM, KOTopble IIMelOTCH
Y MJIeKOIIIITalO IIX, OTMeqeHbl MHorllMII IICCJIep,OBaTeJIHMII. Y pa3HblX
ceMellCTB H eplln; Trinci (1909, 1912), Palme (1934) II Adams
(1942, 1953) HaXOp,IIJIII IIX B a]:(BeHTIID;III1 RapoTII]:(HOll p,yrlI, B 06-
JIaCTII BHYTpeHHell COHHOll apTeplIlI, y BapaHOB OHII IIMelOTCH B lIepe-
ropop,Re Memp,y RaHaJIaMII (mapOTIIJI,HOrO cIIHyca» (Adams, 1952).
B 0lIy6mmoBaHHblx 1I0p,p06HblX pa60Tax IIO MopifloJIOrll1I II rIICTOXII-
MIIII KapOTIIp,HblX COCyp,OB y Chalcides ocellatus II Scincus scin-
cus (Sidky, 1967) II Y Trachysaurus II Tiliqua (Rogers, 1967)
OlIlIcaHbl RJIeTOqHbIe rpYlIlIbl B ap,BeHTIIIJ;IIII KopHell BHYTpeHHell COH-
HOll apTepIIII II B oRpymalO ell KapOTIlp,HYIO p,yry coe]:(IIHIITeJIbHOll
TKaHB - cpep,1I KOJIJIareHOBblX, 8JIaCTuqeCRIIX II peTIIRYJIlIHOBblX BO-
JIOKOH. no MHeHll10 Rogers, MeJIKlIe RJIeTOqHbIe rpYIIIIbl (B 2, 4,
12 KJIeToK) B COBOKynHOCTII cpaBHIIMbl C RapOTIIJI,Hbl1\l «TeJIOM» MJIe-
ROIIIITalO IIX. IIMelOTcH p,Be pa60Tbi C YKa3aHlleM Ha cy ecTBoBaHlIe
aHaJIOfII1ffibiX KJIeTOK y qepelIax (pIlC. 53), ROTopbIe B Bllp,e He60JIb-
mllx Y3eJIROB IIplIJIemaT CHapY>KII R BHYTpeHHell COHHOi'1 apTepIIII
(Terni, 1931) IIJIII paCIIOJIaralOTCH B ee MbImeQHOll 060JIOqRe (Pal-
128
Ille, H);14). Y I';pOIW}J,IJ.TIOBLIX II :ll\Ipii MLI )1(' I1HW:1I1 YIlOJllllllHIIUll 06
3Toii TImmI.
Pahne (1934) onlleaJ! y Hll.(Cplll II 'wpl'mIX n}J,0.'1L aOpT3JlbHblX
II JlerO'lHLIX ]:(yr II MemJ\Y pa3ueTn:ICIllJ}IMII truncus arteriosus pac-
CCHHHLle naparaIlr.1IIOliapIlL!l' I';:JeTI\II. AllTop nO]:('ICpl-\lIYJI, '1'1'0 OHII
BCTpeqaIOTOI r:lallIlhlM 06pa30M B I uniea media II tunica adventicia.
Bee ::ITII RJICTOqllLIC rpYllllbl, pareeHHnblC Y B3pOC.'ILIX pl'IITHJIlIii
B CTeImaX mauepHblx COCY.L\OB II IIX npOIl3BO}J,HbIX (aopTaJILHbIX, JIe-
rOqHbIX II KapOTIIjJ:I1LIX ]:(yrax), y OWIhllllIHcTBa JlnIBOTIlLIX He 06JIa-
]:(aIOT xpollIaiPiPIIHHOl'TbIO. TIO:IOiI\IITe:lhllaH xpollIac}JiPlIHnaH peaK HJI
ounapymcHa Trinci (19 t ) TO:lhI\O y o}J,lIoro BII}J,il Hll.(epII (Gongy-
Ius ocellatus) II nO.l0ilnITl':lhWIH aprl'IITo<lm.1bHaH nORa:1aIla Pal-
me (1934) y Smaragdeider hse. II OTHOllieHlI1I cO}J,epmaHIIH IIopa]:(-
peHaJIIIHa y Lac('rta mUl alis Jl(J:I 'lleIlLI H('raTIIBHhIl' pC:1y.-lhTilTLI R
I..apOTII}J,HLIX coeY iJX II I10;mTIIBIlLII' II :1l'HJ'lHo-aopTa.ILIlLIX (Mul'a-
tori et. aI., 1 %.Ia) .
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.1lIii il.;a6('plloro npollex01R}J,ellIlH, OI_PYIH(' II hI I..anlI,1JJlHpaMII, IlHOf}J,a
clIHyrOIl}J,Horo TlIna, II TOIIIHIMII IlPpllHLlllIH 1I0JI01..lIaMII. HJIeTl..u He-
npallll.'lhIlOii IlJIII OROII}J,HOii {IJOpMLI, IllIIeIOT ABa copTa H}J,ep (Adams,
19,")2, H),18; Sidky, 19(7). 1I]:(pa O}J,III1X I;:JIl'TOI.. l'PYIlIIhIC (6-10 MK),
nY;lhIphl;OIlIl.L\lIhll' co eJI1\OO O"paIIIl'I1l1hIM XpOllIiJTIIHOJII, }J,pYflle-
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Pile. 53. l'pyulla J\lcmmx xpOJ\laIPIPHHHhlX I\JIeTOK (a) B CTCHKe BHYTpeHHeH
COHHOii apTl'pl1ll 'lepcuaxlI (l'alme, 1934)
5 II. A. CMHTTCH
129 -
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1d:eJIKlle (3-4 Jltl£), nJIOTHbIe, llHTeHCIIBHO OKpallleHHble. YRa3aHllH
Adams (1952, 1958) 0 nplICYTCTBlIll oc06b1x «JIHMljJoQlITOnO;JJ;06HblX»
KJIeTOR, «CBH3aHHblX C HepBaMll» II, B03MomHO, COOTBeTCTBYIOII{llX
IIpoHenp06JIaCTaM Ochoterena (1936) II Boyd (1937), no HameMY
MHeHlIIO, IIpe]:(cTaBJIHIOT IIHTepeC B TOM OTHomeHlIll, qTO, 110 onlIca-
HIIIO Adams, HanOMIIHalOT rJIlIIO, BCTpeqalOII{YIOCH B naparaHrJIIIHX Y
..MOJIO;JJ;bIX mlIBOTHblX II Y 3apo;JJ;blmefl nTII II MJIeKOnIITaIOII{IIX. TIo-
;JJ;06Hble RJIeTKll paCCMaTplIBalOTCH HaMll ;JJ;aJIbme (rJI. VII) RaK
cIIOHrlI06JIaCTlIqeCKaH BeTBb HeMp08KTo]:(epMaJIbHOrO 3aqaTKa, ROTO-
paJl HBJIHeTCH llCTOqHllROM xpoMaljJljJllHHblX 8JIeMeHTOB.
lITaR, B COCY;JJ;llCTOM CIICTeMe pa3HblX OTPH;JJ;OB npeCMblRalOII{llXCH
cocpe;JJ;OTOqeHO eIl{e ]:(OCTaTOqHO 60JIbmOe ROJIIIqeCTBO xpoMaiPiPIIHHoM
TRaHII, XOTH II l.IeHbme, qeM Y pbl6 II RPyrJIOpOTbIX, HO 3HaqllTeJIb-
HO 60JIl,me, qeM Y BblllleCTOHII{llX n03BOHOqHbIX. AJIH Hee xapaKTep-
Ha BHYTpllCTeHOqHaH JIOKaJIII3a IIH: B cocY;JJ;ax apTepllaJIbHOrO TIIIIa
ee IIJIaCTbl II rHe3]:(a HaXO;JJ;HTCH B cpe;JJ;HeM II lIapymHoM 060JIOqRaX
II B oRpymalOII{eM coe;JJ;llHllTeJIbHOM TRaHII, B TO BpeMH KaR B BeH03-
HblX cocY;JJ;ax - BO Bcex Tpex 060JIOqKax. O eIIlInaH 8TOT ljJaKT, Bll;JJ;II-
MO, CJIe;JJ;yeT npll3HaTb ero 8BOJIIO IIOHHOe 3Ha'leHlIe. Y RpyrJIOpOTblX
Bce xpoMaljJljJlIHHble 8JIeMeHTbl, r]:(e 6bl OHII Hll Haxo;JJ;llJIllCb,-B Be-
Hax, apTepllHx llJIII B cep;JJ; e, rpaHllqaT C 8H]:(oTeJIlIeM; y pbl6 ::ITa
JIORaJIlIaa IIH IIOJIHOCTblO coxpaHHeTcH TOJIbRO B BeHax, a B apTeplI-
3JIbHJ.lX cocY;JJ;ax ROJIlIqeCTBO ee pe3Ro CHllmaeTCH II KOHTaRT C aH]:(o-
TeJIlIaJIbHOM cTeHRoM Ha6JIIO]:(aeTcH TOJIbRO Ha OrpaHllqeHHblX yqaCT-
Rax, Y npeCMbrnalOII{IIXCH, TaR me RaR II y aM1jJ1I6llM, STOT IIpOn;ecc
npO;JJ;BIIHYJICH ]:(aJIbllle: BO BHYTpeHHeM 060JIOqKe apTeplIM npaRTlIqe-
CM HHr;JJ;e yme HeJIb3H BCTpeTllTb xpoMaljJljJlIHHblX RJIeTOR, OHII Bce
-«OTO;JJ;BIIHYTbl R lIepllljJeplIll» II pacnOJIOmeHbl B;JJ;aJIII OT apTepllaJIb-
Horo RpoBoToRa, B BeHax me OHII eIl{e rpaHHqaT C npocBeTOM, lfHbI-
MIl CJIOBaMII a]:(peHaJIOBaH CIICTeMa B CBoeM llCTOpIIqeCROM pa3BllTlI1I
CBaqaJIa IIORll;JJ;aeT apTeplIaJIbHOe PYCJIO, a nOTOM - BeH03Hoe. qTO
iRe KaCaeTCJI aH]:(ORpllHHOM aRTllBHOCTll RJIeTOR, TO B cocy]:(ax TYJIO-
BllIl{a OHa nOJIHOCTblO coxpaHHeTCH, a B meMHo-rpY;JJ;HblX pe3RO pe]:(y-
II;llpOBaH3 (rllCTOXllMllqeCRII aTO BbTpamaeTCH B (<HeXpoMaljJljJlIHHo-
CTII» naparaHrJIlleB aTOM 06JI8CTII) II paCCMaTpllBaeTCH HaMIl RaE
CJIe;JJ;CTBHe Tex pe]:(Y lIpyroIl{IIX IIpo eCCOB B ma6epHOM aIIlIapaTe, RO-
Topble OIIlIcaHbI B IIpe]:(bl;JJ;YII{eM rJIaBe.
." I1MelOII{IIeCH B JIIITepaType ]:(aHHble 06 IIHHepBan;IIII llIeMHo-rpY;JJ;-
HblX COCY;JJ;OB penTlIJIllM OqeHb CRY]:(Hbl, 110 cpaBHeHlI10 C xopomlIM
II3yqeHlleM aTOM 06JIaCTII Y MJIeROIIIITaIOII{IIx. O;JJ;HaRO Bce me IIMeIOT-
CJI OCHOBHble cBe;JJ;eHIIH, no ROTOpbIM MomHO CY]:(IITb 0 rOMOJIOrIIII IIX
IIHHepBaU;IIOHHblX OTHollleHllM. I1HHepBan;IIJI RapOTII;JJ;HblX COCY]:(OB Y
penTHJIIIM OCYII{eCTBJIHeTCH BeTBHMII 6JIym;JJ;alOII{ero HepBa, H3bIKOrJIO-
TOqHOro II meMHoro cIIMnaTllRyca (Couvreur, Duculty, 1924; Ter-
ni, 1931; Ask-Upmark, 1935; Adams, 1939, 1942, 1952; Kaushi-
va, 1940; Boyd, 1942), T. e. ll3 Tex }I,e lICT01JHIIKOn, ROTopble H3-
BeCTHbi y MJIeROnHTaIOII{llX. Y HeROTopblX OTPH;JJ;OB (RpORO;JJ;IIJIOBbIe-
Couvreur, Duculty II qepenaXoBble - Fedele) aBTOpbl npe]:(nOJIa-
13Q:
raIOT cy:rn;eCTBOBaHlIe B CTeHl\aX COCYP;OB p;enpeCCOpHOrO HepBa, roMO-
JIOrlIqHOrO HepBY MJIel\OIIIITaIOIT(IIX. OT BepXHeH ropTaHHOH BeTBII
Jl3bII\OrJIOTOqHOrO II CIIMnaTlIqeCI\OrO HepBOB TOHl\lIe BOJIOI\Ha npOHII-.
l\aIOT B I\JIeTOqHble rHe3p;a lIaparaHrJIIIOHapHOH Tl\aHII.
MeJIl\He naparalIrJIHOHapHbIe Y3eJIl\H II OTp;eJIblIbIe xpoMa<I><I>HH-'
HbIe I\JIeTI\H conpOBomp;aIOT HepBHbIe CTBOJIIII\H II BCTpeqaIOTCJI B pa3-
HbIX opraHax HIT(epnD;. OHII Tal\me nonap;aIOTCH BHYTPII IIJIII PHP;OM C
CIIMnaTlIqeCI\HMII raHrJIIIJlMII. ::1Ty IICTIIHHO naparaHrJIIIOHapHyro
Tl\aHb Y peIITIIJInH onlIcaJI eIT(e Wiesel (1902a). OHa OTHOCHTCH R
TOMY TlIny BHeHap;nOqeqHIIROBOH TRaHII, pa3BIITIIe II paClIpep;eJIeHlIe
ROTOpOH 06YCJIOBJIeHbi nporpeCCIIBHblM pa3BHTIIeM CIIMlIaTlIqeCROH
HepBHOH CIICTeMbl.
nOp;bITomHBaH Bce H3JIOmeHHOe B 3TOH rJIaBe, MO>RHO CI\a3aTb, -- )
QTO l\JIaCC npeCMbIRaIOIT(HXCH HBJIHeTCJI cTyrreHbIO B 3BOJIIOD;HOHHOM
pa3BHTHII xpoMa<I><I>IIHHOH CIICTeMbI n03BOHOQHbIX, Ha ROTOpOH yme
P;OBOJIbHO qeTRO BbIHBHJIaCb nporpeCCIIBHaH JIIIHHH ee pa3BIITIIH B
Bllp;e 60JIee IIJIH MeHee o<I>opMJIeHHbIX Hap;nO'leqHbIX meJIe3 H perpec-
elIBHaH - B BIIp;e (<HeXpoMa<I><I>IIHHbIX» y3eJIl\OB B RpOBeHOCHbiX cocy-
p;ax ma6epHOrO IIpOHcxomp;eHHH. B cocyp;ax TYJIOBIIIT(a, IIplIJIemaIT(1IX
1\ Hap;IIOqeqHbiM meJIe3aM, TO me eIT(e coxpaHHeTCH al\TlIBHO <I>Y:UR-
D;IIOHlIpyIOIT(aH meJIe3IICTaH xpoMa<I><I>IIHHaH Tl\aHb, ROTopaH, OqeBIIp;-
HO, Ha p;aJIbHeHllIeM 3Tane <I>IIJIOreHe3a - y nTIID; II MJIel\OIIIITaIO-
IT(IIX - «YHp;eT» Ha nOCTpoeHHe Hap;IIOqe'ffiIIROB.
<1>opMlIpoBaHlIe Hap;nOqeqHIII\OB II Mop<I>OJIOrlIqeCl\oe 060c06JIeHIIe
HX B l\aqeCTBe caMOCTOJlTeJIbHbIX OpraHOB, BnepBbIe HalI60JIee qeT-
RO HaMeqaIOIT(eeCH y npeCMbmaIOIT(IIXCH, CJIep;yeT paCCMaTplIBaTb RaR
IIplIcnoc06IITeJIbHblH MexaHH3M, 06eCneqlIBaIOIT(IIH y BbiCOROOpraHII30-
BaHHbIX mIIBOTHbIX, B qaCTIIOCTII MJIeI\OnIITaIOIT(HX, MaRCHMaJIbHO 6JIa-
rOnplIHTHbIe YCJIOBIIH <I>YHRD;HOHaJIbHOH al\THBHOCTII CIIMnaTo-ap;peHa-
JIOBOH CIICTeMbI. Mop<I>OJIOflIqeCRaJl n;eHTpaJIII3aD;IIH conpOBomp;aeTCJI
y pelITIIJIHH C6JIHmeHHeM II B3alIMHbIM npOHIIlmOBeHlIeM xpoMa<I><I>IIH-
HOH «<M03rOBOH») II IIHTeppeHaJIOBOH «<I\Opl\OBOH») Tl\aHeH, qTO xa- I
paRTeplI3yeT 06:ru;yIO TeHp;eHD;IIIO 3BOJIIOD;IIII II HBJIHeTCH HeOT'beMJIe-
MOH MoP<I>OJIOflIqeCI\OH 'WPTOH Hap;nOqeqHbIX meJIe3 BbICilllIX II03BO-
HOqHbIX. <1>YHI\D;IIOHaJIbHOe 3HaqeHIIe TaROrO c6mDl\eHHJI P;BYX rna- )
HeH OCTaeTCH P;O CIIX 1I0p HeHCHbIM. npep;IIOJIOmeHlIe 0 BJIIIHHIIII 1\0p-
I\OBOrO CJIOJl Hap;nOqeqHIII\OB Ha MeTIIJIHD;HIO Hopap;peHaJIIIHa (Ches-
ter Jones, 1957) He nop;TBepp;IIJIOCb (Coupland, 1953; Olivereau, 1959).
OIIlIcaHHoe y penTIIJIIIH paClIpep;eJIeHlIe RJIeTOI\ B I\pOBeHOClIblX co-
cYfJ;ax TO me He COrJIaCyeTCH C 3TOH ROHD;enD;HeH, Tal\ l\aR 01I1I He
CBH3aHbI C IIHTeppeHaJIOBOH Tl\aHbIO, IIO cop;epmaT MeTIIJIlIpOBaHHLIH
ropMOH.
--!
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S.
L' .aa8a V
DTII:QbI
qTO IIpep;CTaBJIHeT C060H xpoMaw<InIHHaJl CIICTeMa IITlIU;? Ral\IIe
tTpyI\TypUbIe OC06eHHOCTII ee Xapal\TepHbI P;JIH }TOrO RJIaCCa 1I03BO-
HO'lHbIX II IIMeIOTCJI JIll OTJIHqlITeJIbHbIe IIp1I3Hal\lI, CBOHCTBeHHbIe
TOJIbRO nTIID;aM?
CBe)WHIIH, IIpIIBe)WHHbIe B IIpep;bI)J;yrr(IIX rJIaBaX 0 paCnOJIOme-
RIIII xpoMa<I><I>IIHHOH TRaHII B TeJIe P;PyrlIx 1I03BOHO'lHbIX mIIBOTHbIX,
YRa3bIBaIOT Ha P;IIWWY3HOe ee paClIpep;eJIeHlIe l\aR CaMbIH xapal\-
TepHbIH Mop<I>OJIOrlIqeCRIIH IIplI3Hal\. Y IITIID; CTelIeHb pa3p;p06JIeHHO-
CTII meJIe3IICTbIX XpOMaWWIIHHbIX 3JIeMeHTOB IIpeBbIIlIaeT BCe, qTO II3-
BeCTHO P;JIJI p;pyrlIX RJIaCCOB, II TpYP;HO 1I0MaeTCH aHaJIII3Y. B 3TOM
fQTHOmeHlI1I nTIID;bI CTOHT OC06HHROM B pJlP;Y 1I03BOHOqHbIX. ReB03-
"MOlliHO Ha nepBbIH B3rJIJlp; YCMOTpeTb RaRyIO-JIII60 3aROHOMepHOCTb
B paClIpep;eJIeHIIII OTp;eJIbHbIX KOMlIJIeRCOB RJIeTOI\, 1\0TOpbIe B BlIp;e
:MeJIRIIX me3p; pa36poCaHbI B pa3HbIX 06JIaCTHX TeJIa IITIID;,- B CTeR-
,Rax: COCYP;OB, HepBHbIX BOJIORHaX, raHrJIIIJlX, npOCJIoiiRax COep;IIHII-
'TeJIbHOH TRaHII II BHYTpII OpraHOB. ECJIII, Op;HaRO, paCCMaTplIBaTb
-xpoMa<I>Il>IIHHYIO TKaHb IITIID; He II30JIHpOBaHHO, a RaR OP;IIH II3 3Ta-
:nOB Il>IDIOreHeTlI'IeCROrO pa3BIITIIJI II RaR CIICTeMY, Hepa3pbIBHO CBH-
.3aHHYlO C 3BOJIIOD;IIeH CIIMlIaTlI'IeCl\1IX HepBHbIX 3JIeMeHTOB, TO 3ap;aqa
UeCROJIbRO o6.rrerqaeTCH II MHoroe CTaHOBIITCH B03MomIlbIM 060CHO-
naTb.
. OqelIb nop;p06HbIe II BamHbIe CBep;eHIIH P;JIH Xapal\TeplICTIII\H XpO-
:MaIl>Il>IIHHoii: CIICTeMbI IITIID; p;aeT CTapoe IICCJIep;OBaHlIe Kose (1907),
:KOTOpbIH rrpoBeJI HaMIOp;eHIIH Ha 60JIbmOM quCJIe BII]:(OB - OT TOJIbRO
"qTO BbIn;ynlIBilllIxCH rHe3p;OBbIX II p;o B3pOCJIbIX. IJ;eHHocTb ero Ha6-
JlIOp;eHIIH aaKJIIOqaeTCH B TOM, qTO, IIOMHMO p;eTaJIbHeHllIero II CRPY-
nYJIe3HOrO OlIlIcaHIUI MOPWOJIOrlI1I lIaparaHrJIlIeB, OR IIOBCIOP;Y OTMe-
aJI qaCTOTY, C ROTOpOH eMY IIpIIXOP;IIJIOCb BCTpeqaTb TY IIJIII IIHYIO
CTPYRTYPY, B TOH IIJIII IIHOn: CBH3II C P;PyrHMII Tl\aHHMII II opraHaMII.
TiIIaTeJIbHOCTb IIpOBep;eHHblx Kose Ha6JIIOp;eHIIH, orpOMIlbIH MaTeplIa-
aJI, :MHomecTBo 06CJle]:(OBaHHbIX BIIP;OB IITIID; 110M ora lOT Cp;eJlaTb 3aRJIIO-
qeHIIH06 .aHaTOMlIqeCRIIX oc06eHHOCTHX lIaparaHrJlIIOHapHOH CII-
CTeMbI IITIII{ II CqlITaTb IIX BIIP;OBOH IIpIIHap;JlemHOCTbIO. IIMeHHo 110-
BTOMY MbI BOCIIOJIb30BaJilICb waRTlI'IeCI\OH CTOpOHOH CBep;eHIIH, ROTO-
pbIe IIpIIBOP;IIU' 3TOT aBTOp. CaM Kose 1I0qTII He 3aHIIMaJlCJI P;PyrlIMII
1I03BOHOqIlbIMII II He CTaBIIJI lIepe]:( C060H HIIRaRIIX cpaBHIITeJIbHO-
:Mopll>OJIorll'Iec.RIIX. 3aAaq. OH p;ame He 3ap;aBaJICH D;eJlbIO RaR-TO CII-
132
TeMaTII3IIpOBaTb CBOII naoRroAeHllH BHyTpII IIpep;CTaBIITeReH RRaCCa
nTII , II BllP;OBOH 1I0p;60p Y HerO B 113BeCTHOH Mepe ORa3aRCJI CRY-
qaHHbIM (qum, BopOHa, RyplI a, CHerllpb, BJl6RIIR, rORy6b, )W03P;
II p;p.). Op;HaRO ero Ha6Rrop;eHIIH II 1I0P;P06HO OIIlIcaHHbIe MOp«I>OROrH-
qeCRlIe «paRTbI xopomo YIWap;blBaroTcH B TY cxeMY, ROTOpaJl BbITe-
KaeT lIB aHaRIIBa llCTopuqeCRoro paBBllTIIJI CIIMllaTo-aAPeHaROBOH CII-
CTeMbI 1I03BOHOqHbIX. C 8BORro IIOHHblX 1I03II IIH Ha6Rrop;eHHH Kose
nORyqaroT, C OP;HOH CTOpOHbI, y6ep;IITeRbHOe 06'bHCHeHlle, a c P;PyroH,
cRymaT OCHOBaTeRbHbIM 1I0p;TBepmp;eHlIeM IIpaBIIRbHOCTII TOro HallpaB-
JIeHIIH, ROTOpOe IIpOBOP;IITCH HaMil B 113yqeHIIII xpoMa«I>«I>IIHHOH
CIICTeMbI B PHP;Y 1I0BBOHOqHbIX mlIBOTHbIX, II II03BORHroT p;eTaRII3II-
pOBaTb II yrRy6IITb HeROTopble BOllpOCbI.
B RpaTROM 113RomeHIIII Ha6JIIOAeHlIH Kose (1902, 1904, 1907)
CBOP;JlTCH R cRep;yrorqeMY. IIpemp;e Bcero Kose Bce xpoMa«I>«I>IlHHbIe
IWeTOqHbIe CROIIReHIIH Ha3b1BaeT lIaparaHrRIIHMII. lIClloRb30BaHlie aB-
'TOp OM <!TOro TepMIIHa, BIIP;IIMO, He cRyqaHHo, a HBRHeTCH CRep;CTBII-
eM oc060 TeCHoro II paCllpOCTpaHeHHOrO ROHTaKTa xpoMa«I>«I>IIHHbIX
' meMeHTOB y IITII C 1I0rpaHllqHbIM CIiMllaTuqeCRIIM CTBOROM, C lIe-
pll«I>eplIqeCRIIMIi CIiMllaTlIqeCRIIMIi raHrRIIHMll II HepBaMII. 8TIIM Mom-
EO 06'bHCHIITb TO 06CTOHTeRbCTBO, qTO aBTOp paCCMaTplIBaeT Bce
IIaparaHrRIIOHapHble CTpyI\TYPbI IIOP; yrROM 3peHIIH llX BBaIIMOOTHO-
meHIIH C 8ReMeHTaMII CIIMIIaTuqeCROH HepBHOH CIICTeMbI. B OTp;eJIb-
HOH rRaBe OH OlIllCbIBaeT IIOrpaHll'IIlbIH CIIMllaTII1JeCRIIH CTBOJI, )WRIIT
ero Ha TPIi pa3p;eRa - meihIblH, rpyp;HOH II 6promHoH, B Ramp;OM II3
ROTOpbIX 113Y1JaeT aHaTOMlIqeCROe 1I0RomeHlle, rlICTORoruqeCRoe
CTpoeHlIe II IlToRorlIro lIaparaHrRlleB. Hap;IIOqeqHIIROBbIe II Rapo-
'T1Ip;HbIe lIaparaHrRlIlI, a TaRme Te, ROTopbIe 3aRomeHbI B CTeHRax
.a6AoMIIHaRbHbIX RpOBeHOCHblX COCYP;OB II B opraHax, OH TO me pac-
CMaTpllBaeT BMeCTe C CIiMllaTlIqeCROH HepBHOH CIICTeMOH RaR ee CO-
CIaBHYro qaCTb.
B paCllpep;eReHIIII lIaparaHrRlIeB 110 XOP;Y 1I0rpaHIIqHbIX CTBOROB
HMeroTcH Ollpep;eReHHbIe 3aROHOMepHOCTII: Y MOROp;bIX IITII IIX 3Haqu-
'TeRbHO 60Rbme 110 cocep;CTBY C CIiMIIaTllqeCRIIMII raHrRllHMII, qeM Y
"'BBpOCRblX; B TO me BpeMJI, He3aBllCliMO OT B03paCTa, 60RbmllHCTBO
lIaparaHrRlIeB RemlIT BHYTpII IIJIII B6RII3II Y3ROB a6p;oMIIHaRbHoH 06-
JIaCTIi II MeHbme B rpYP;HoH II meihIoH. B 1I0rpaHuqHblx HepBHbIx
CTBORax, COep;IlHJlrorqllX CIIMllaTUqeCRlle Y3RbI MemAY C060H, lIaparaH-
£RlIoHapHbIe RReTRII BCTpeqaroTcH TORbRO B a6p;oMIiHaRbHoH 06RacTII
II TO B 3HaqlITeRbHO MeHbmeM RORlIqeCTBe, qeM B CaMliX Y3Rax.
OllllcaHlie MaTepliaRa Y IITII Kose pa3p;eRIIJI Ha ABe qaCTIi. IIep-
BaH 1I0CBHrqeHa TeM ROMIIReKcaM BHeHap;II01JeqHIIROBbIX xpoMa«I>«I>IlH-
HblX 8ReMeHTOB, ROTopbIe MbI paCCMaTpliBaeM RaR BTopuqHbIe, «I>II-
JIOreHeTllqeCRII HOBbIe, pa3BllBarorqlIeCJI lIapaRReJIbHO C 8BoRro lleH
HepBHOH CIICTeMbI, BTOpaJl - xapaRTeplICTIiRe HaAIIOqe1JHIIROB IITII ,
RaR meRe3IICTbIX opraHOB, ROTOpbIe yme Y IIpeCMbIRarorqlIXCJI BbI)J;e-
ReHbI HaMil B caMOCTOHTeRbHYro CTPYRTYPY opraHHoro TlIlla.
IIaparaHrRIIOHapHbIe CROIIReHHH B 0611RbHOM RORlIqeCTBe Haxo-
J,J;HTCJI PHAoM C Hap;II01JeqHIIRaMII, C a6p;OMlIHaRbHb1M II 3aAHliM OT-
'133
pe3KaMM 1I0rpaHMqHblX CTBOJIOB II BAaJIII OT HIIX: B COJIHeqHOM CIIJIe-
TeHIIII, B COeAMHMTeJIbHOll TKaHII 6plOlliHOll II 3a6plOlllIHHOll 06JIaCTM.
3TII MeJIKlIe lIaparaHrJIliH, IIHOrAa COCTOHmlIe Bcero 113 2-3 KJIeTOK,
KOTOpbIe Mbl BO MHomeCTBe Ha6JIIOAaJIII B IIeplIOA aM6pMOHaJIbHOrO
pa3BIITMH KypMHblX 3apoAbilliell, Kose (1907) OlllICaJI II Y APyrlIx
llTH:U;. OHM JIemaT IIJIM n30JIllpOBaHHO - BHe CBH3II CO CTeHKaMM Ma-
rlICTpaJIbHblX KpOBeHOCHblX COCYAOB AaHHon 06JIaCTII, IlJIn KOHTaRTlI-
pyroT C xpoMa«I>«I>MIIHbIMII CKOIIJIeHIIHMM, KOTopble MH«I>IIJIbTPMPYIOT
COCYAIICTble CTeHKII. Ho qTO OqeHb BamHO B OIIlIcaHIIU aTIIX CTPYK-
TYP Y Kose - TO CBH3b IIX C aJIeMeHTaMII CIIMlIaTuqeCKon HepBHon
CIICTeMbi. ABTOp AaJI OqeHb rrOAP06HYIO xapaKTeplICTlmy B3aUMOOT-
HOlleHIIH rraparaHrJIIIOHapIlblX OCTpOBKOB aTOll 06JIaCTII C nepBllblMU
aJIeMeHTaMII: xpoMa«I>«I>IIHHbIe ImeTim mloo rrO.TJ.HOCTblO 1I0rpYi-I,ellbl B
HepBHble CTBOJIIIKII lIepn«I>epuqeCKMX pa3BeTBJIeHIIB HepBOB. I! Y IIX
K BHYTpeHHIIM OpraHaM, JIlI60 rrpMJIeralOT R HIIM C IIepn«I>epIlII II JIe-
}liaT IIOP; coe]:(IIHIITeJIbIIOTl.aHHOH 060JIOqKOB. Kose OTMeqaeT, qTO Aa-
}I,e Te Memme lIaparaHrJIIIOHapHble Y3eJIKII, ROTopbIe RaR 6YP;TO 6b1
He CBH3aHbI C HepBHblMII CTBOJIIIRaMlI, npII p;eTaJIbHOM paCCMOTpe-
HIIII II COOTBeTCTByroIT(IIX MeTop;ax 06pa60Tl\II ORa3bIBalOTCH B J>OH-
TaRTe C caMblMII MeJIbqaHIlIMMM pa3BeTBJIeHIIHMII HepBHblX nOJIOI>OH.
KpOMe Toro, B lIaparaHrJIIIHX 'I aCTO BCTpe'IaIOTCH raHrJIII03Hbie CIIM-
lIaTlIqeCRlIe RJIeTIm, Ha ROTOpblX xpoMa«I>«I>MHHbIe RJIeTlm CIlP;HT B
BII]J;e llalIOqeR. II Te II P;PyrMe yqacTBYIOT B 06pa30BaHIIII HepBHblX
CIIJIeTeHIIll: 6plOllHOll: 06JIaCTII.
IIpe06JIa]:(alOmee 60JIbllllIHCTBO Bcex CROIIJIeHlIll lIaparallr.TJ.1I0Hap-
HOB TRaHM II oTAeJIbHbIX xpoMa«I>«I>IIHHblX RJIeTOR, pacUOJIOmeHHblx B
raHrJIIIHX M HepBax, JIemaT HelIocpep;cTBeHHo Ha I\alIlI.TJ.JJHpaX MJIII
B6JIII3II HIIX.
C Hallell: TOqRII 3peHIIH, IIpep;CTaBJIHIOT HecoMHeHHbIll IIHTepeC
Ha6JIIOAeHMH Kose, xapaRTepM3YIOmlIe rraparaHr.TJ.IlOlIapnylO TKaHb B
1I0QKaX II cocyp;ax. B OCTaTKax lIepBlIqHOll 1I0qKII, ROTopble BCerP;3
paCIIOJIOmeHbl B6JIII3II Hap;IIOqeqHMROB II rrOJIOBbIX meJH'3, neT raHrJIU-
03HblX CIIMlIaTllqeCRIIX RJIeTOK, a lIaparaHr.iIIIOHapHble CKOIIJIeHIIR
ORpymalOT 1I0'IeqHble RaHaJIbn;bl II OCYMROBaHHbIe MaJIblIlIrMeBbl Te:Ib-
n;a. EOJIbllllHCTBO aTIIX lIaparaHrJIlIeB, 110 CBII]:(eTeJIbCTBY aBTopa,
He IIMeeT BII]J;llMOll: CBH3II C ClIMlIaTlIqeCI\Oll: HepBHOH CIICTeMOll. BTO-
plIquaH IIOqRa cop;epmlIT CIIMlIaTlIqeCKIIe raHrJIIIII, ROHTaKTllpYlOmlle
C lIaparaHrJIlUIMII. ROTopble MOrYT BXOP;IITb B CTpyRTYPHblii COCTaB
CTeHOK IIOqeqHblX apTeplIH II BeH, a MorYT, He3aBIIClIMO OT KpoBe-
HOCHbiX COCYP;OB, IIpOHIIRaTb BAOJIb 1I0qeqHblX RaHaJIbn;eB fJIy60RO B
lIapeHXIIMY 1I0qeK. Cpep;1I Bcex COCYP;IICTblX 06pa30BaHIIH 3a6plOlllln-
HOB 06JIaCTII Kose BbiAeJIHeT 1I0qeqHble BeHbI RaR COCYP;bl, 60JIblle
APyrnx HacbIIT(eHHble xpoMa«I>«I>IIHHoH TRaHblO. B a6p;OMIIHaJIbHOll:
aopTe II B RpynHblX apTeplIJlX 6plOllHOH 1I0JIOCTII, B TOM qnCJIe Hap;-
nOqeqHlIROBOll:, Kose OlIlICblBaeT cROIIJIeHIIH lIaparaHrJIIIOHapHOll TRa-
HII, ROTopble pacrrpocTpaHHIOTcH B tunica media II tunica adventi-
cia IIJIII 3aHIIMalOT OP;lIH lI3 aTHX CJIOeB. qacTb rraparanrmleB B
tunica media He IIMeeT coep;IIHlITeJIbHOTRaHHOll 060JIoqRII II B BII]J;e
134
'Tl\aHeBbIX 3JIeMeHTOB IIpOHII3bIBaeT Cpep;HlOIO 060JIo'lliY aOpTbI II P;PY-
I'IIX apTepllii:. MeCTaMII lIaparaHrmlOHapHaH rnaHb B apTeplIaJIbHbIX
'Cocyp;ax IIpOHlIl\aeT IIO'ITII p;o tunica intima, HO HIII\Orp;a He corrplI-
RaCaeTCJI C 3Hp;OTeJIlleM. BOJIbillOe 1\0JIIIqeCTBO lIaparaHrJIlleB B a6-
AOMIIHaJIbHbIX apTepllHX II BeHax aBTOp OTMeqaeT RaR MOpq,OJIOrlI-
qeCI\YIO qepTY HaCTOJIbRO xapaRTepHyIO, 'ITO OHa Bcerp;a MOHWT CJIY-
H\HTb 01I03HaBaTeJIbHbIM IIplI3HaROM P;JIH BblHBJIeHIIH COCYP;OB Ha IIpe-
IIapaTax.
llplI O1I1IcaH1I1I RpOBeHOCHblX COCYP;OB a6p;OMIIHaJIbHoii: 06JIaCTII
Kose BbIp;eJIIIJI p;Ba xapaRTepHbIX IIplI3HaRa, ROTOpbIe BamHbI C TOq-
RII 3peHIIH cpaBHIITeJIbHO-Mopq,OJIOrlf'IeCRoii: OIJ;eHRII ero q,aRTII'IeCRO-
ro MaTeplIaJIa. llepBbIii: OTHOCIITCJI R TOJIbRO qTO CRa3aHHOMY: B aop-
Te II apTeplIaJIbHbIX RpOBeHOCHbIX cocyp;ax lIaparaHI'JIIIII IIpeIIMy:ru;e-
CTBeHHO paCIIOJIOmeHbl B cpep;Heii: II HapYlliHoii: 060JIoql\ax II HIIRorp;a
He ]:(OCTllraIOT 3HAOTeJIIIH, B TO BpeMH RaR B BeHax OHII JIemaT Y
npOCBeTa COCYP;OB. BTOpoii: RacaeTCH clIMlIaTlIqeCRoii: IIHHepBaIJ;IIII lIa-
paraHrmIeB: Te II3 HIIX, ROTopble cocpep;OTOqeHbI B HapY'l\Hoii: ap;-
BeHTIIIJ;lIaJIbHoii: 060JIOqRe apTeplIii:, CBH3aHbi C TOHI\IIMII CIIMlIaTII-
qeCI\IIMII HepBHbIMII BOJIORHaMII, qerO Kose HII pa3Y He BlI)];eJI B-
lIaparaHrJIIIHX, JIema:ru;IIx II30JIIIpOBaHHO B tunica media apTepllii: II
aopTbl.
llpII 1I0lIbiTRe aHaJIII3a Bcero 06ll1IIpHOrO q,aRTHqeCROrO M3Te-
plIaJIa Kose yp;aJIOCb YCTaHoBIITb OP;IIY qepTY B Mopq,OJIOrII'IeCROM
o6JIIII\e lIaparaHrJIlIeB 3TOii: 06JIaCTII, 1\0TOpaJl HelI3MeHHO 1I0BTOpJl-
eTCJI Y Bcex 06CJIep;OBaHHbIX BII]:(OB nTm:t; IIJIII CBoii:cTBeHHa 60JIb-
illIIHCTBY: cpep;II MaCCbI naparaHl'JIIIOHapHOD: TR8HII, ROTopaH BCIOAY
ROHTal\TIIpyeT C CIIMlIaTlf'IeCROD: HepBHOD: ClIcTeMoii:, IIMeIOTCH Y'IaCT-
RII, JIllilleHHbIe IIJIll 1I0qTII JIIIilleHHbIe 3TIIX ROHTaRTOB. R Ha1I60-
JIee I\pynHbIM ll3 HllX OTHOCIITCJI xpOMaq,q,IIHHaJl TRaHb, IIpOHll3bIBaIO-
:ru;aH tunica media aopTbI II HeKoTopbIX 8pTeplIii:. BTOpbIM TaRHM
yqaCTROM HBJImoTCJI IIepBlIqHbIe 1I0qRII, B ROTOpbIX naparaHrJIllOH8p-
HaH TRaHb 1I0rpymeHa B RaHaJIbIJ;b1 IIJIll oRpymaeT B BIIp;e noJIY-
JIYHIIii: OCTaTRII MaJIblIlIrlIeBblX TeJIeIJ;. 06a 3TH yqacTRa B q,llJIOreHe-
TII'IeCROM OTHOilleHIIII - HalI60JIee ApeBHlIe 06pa30BaHIIH: lIepBII'UI8JI
1I0qRa, ROTopaH HBJIHeTCJI pa60TaIO:ru;IIM opraHOM Y B3pOCJIbIX OCO-
6eii: HII3illlIX 1I03BOHO'IHbIX, Y IITIIIJ; q,YHRIJ;IIOHllPyeT TOJIbRO B BM-
6plIoreHe3e II CMeHHeTCH BTOPlIqHOii:; CTeHRII MarlICTpaJIbHbIX COCY-
]:(OB IIPOXOP;JlT CJIOmHbIii: IIYTb nepecTpoii:RII B CMbICJIe cop;epmaHIIH
n HIIX xpOMaq,q,IIHHoii: TKaHII. HalI60JIee ApeBHHMII B BTOM CMbICJIe
HBJIHIOTCH BHYTpeHHHH II cpe]:(HJlH 060JIOqR8 COCYP;OB. IIHHepBau;HS
3TIIX CTPyRTYP CO CTOpOHbI CIIMlIaTlIqeCROD: HepBHoii: CIIC'feMbl OT-
CYTCTByeT Y RpymopoTbIX II cJIa60 pa3BIITa Y PbI6, HO MomeT IIpO-
JIBIITbCH Ha 60JIee BbICOROii: CTYneHII BBOJIIOIJ;IIOHHOro pa3BIITIIJI.
Y IITIIIJ; - eCJIII BTII q,aRTbI CIIpaBeMIIBbI - cJI860e npep;CTaBlITeJIL-
CTBO CIIMlIaTllqeCRIIX 3JIeMeHTOB B6JIII3II lIepBlIqHOD: IIO'IRII II B
tunica media 6pIOIDHOii: apTeplIaJIbHoii: CIICTeMbI COrJI8cyeTCJI C BTHM
II MomeT CJIymlITb IIORa38TeJIeM Toro, qTO 3TII lIaparaHI'JIIIII OTHo-
CHTCJI R H8II60JIee p;peBHen 'I8CTII 3RcTpaap;peH8nOBOD: CIICTeJlb1.
t35
CJIe]:(yeT TYT me orOBoplITbCH, qTO OTCYTCTBlIe CIIMnaTII'IeCKOH
IIHHepBaD;IIII BOBce He JlBJIJleTCJI a6COJIIOTHbiM nplI3HaKOM «p;peBHO-
CTII» IIaparaHrJIIIH: «p;peBHlIe» naparaHrJIlI1I MorYT 1I0JIy<IIITb ee B
3BOJIIOD;IIII BTOpll'lHO, qTO, HanpliMep, lIMeeT MeCTO B .rpYP;HbIX II
wellHblX (KapOTIIp;HOM) CTpyKTypax. Ho B Tex CJIyqaHX, Korp;a CHM-
naTII eCKaH IIHHepBaD;IIH lIaparaHrJIIIJI OTCYTCTByeT (a YKa3aHIIH H8
HIIX BCTpeqaIOTCH II Y MJIeKOnIITaIOIII;lIx), TaKoe 06'bHCHeHlIe Ha1I60-
JIee npaBOMO'IHO II 060cHoBaHo nOCJIep;OBaTeJIbHOCTbIO IICTOpll eCRO-
ro pa3BIITliH CIICTeMbl. A Te lIaparaHrJIIIII, KOTopble B03HIIRJIII II pa3-
BIIJIIiCb nonYTHO C pa3BIITIIeM B 3BOJIIOD;IIII CHMnaTlIqeCROll HepBHoH
CIICTeMbI, TeCHO CBH3aHbI C Y3JIaMII II npOBop;HIIKaMIi 3TOH CIICTe-
Mbl. 0 HIIX, KaK BTOPllqHO B03HIIKWnx II nporpecclIpyIOIqIIX CTPYR-
TYPax, Mbi yme yrroMIIHaJIII B rJIaBe 06 aMiPII611J1X, y ROTOpblX OHn
BlIepBble nOHBJIHIOTCH. Y nTIID; lIaparaHrJIIIOHapHaJl CIiCTeMa 3HaqlI-
TeJIbHO 60JIee pa3BIITa, 'ITO BbipamaeTCH B 0611JIIIII xpoMaiPiPlIHHbIX
3JIeMeHTOB BHYTPII CIiMnaTuqeCKOrO norpaHII'IHoro CTBOJIa, B nepll-
iPeplIqeCKIiX CIiMnaTllqeCI\IIX raHrJIIIHX II 110 XOP;Y HepBOB, IIP;YIII;IIX
K Hap;IIOqeqmrnaM, nOqI\aM II P;PyrlIM opraHaM. 3TIIM MomHO 06'b-
JlCHIITb 60JIbillOe I\OJIII'IeCTBO IIaparaRl'JIIiOHapHblX rHe3p;, BCTpeqaIO-
III;IIXCJI B COep;IIHIITeJIbHOll TKaHIi no xop;y qpeBHblX HepBoB. B TO
me BpeMH TaI\IIe I\apTIIHbI, Ha6JIIOp;aeMbie y IITIID;, HBJImoTCJI, BII-
P;IIMO, IIpOMemYTOqHblM 3TanOM B 3BOJIIOD;IIII ap;peHaJIOBOll CIICTeMbI,
]:(JIH I\OTOpOrO XapaI\TepeH MaI\CIIMaJIbHblH pa36poc naparaHrJIlIeB.
TaI\oe MHeHlIe 0 npOMemYTOqHOM 3TalIe 3BOJIIOD;IIOHHOrO pa3BIITIIR
CI\JIap;blBaeTCH nOCJIe 3HaI\OMCTBa C pacnpep;eJIeHlIeM xpoMaiPiPlIHHoH
TKaHII B a6p;OMIIHaJIbHOH 06JIaCTII MJIeI\OIIIIT8IOIqIIX, y KOTOpblX ee
paClIpOCTpaHeHlIe SHaqlITeJIbHO cymlIBaeTCJI; eCJIII pa36poc naparaH-
rJIlIeB II BCTpeqaeTCH y MJIeI\OIIIITaIOIII;IIX, TO TOJIbI\O B 3M6plIoreHe-
se, p;a II TO He B TaI\Oll CTeneHII, I\aK y J1TIIIi;. TaI\ I\aK Mbi 3Ha-
eM, TO y BblCWnx 1I03BOHOqHblX rJIaBHblM iPYHI\D;IIOHlIpYIOmIIM
IIaparaHrJIlIeM HBJIHeTCH H8]J;nOqeqHIIK, 06JIap;aIOIII;IIH CJIOmHOll peiP-
JIeRTOpHOH perYJIHD;IIell, TO MomHO IIpep;nOJIOmIITb, qTO y IITIID; MId
3aCTaeM TOT 3Tan iPlIJIoreHe3a, I\Orp;a 60JIbWIIe MaCCbI xpoMaiPiPlIH-
HbiX I\JIeTOK HaXOp;HTCH eIII;e no nYTII I\ CBoeMY rJIaBHOMY MeCTY
Ha3Ha emrn - Hap;lIoqeqHIIRy II BMeCTe C pa3BIIBaIOIqellCJI CIIMnaTII-
qeCI\OH HepBHOH CIICTeMOH p;OCTlIraIOT ero Ha CJIeP;YIOIII;eH CTap;IIII
IICTOpll eCI\oro pa3BIITIIH. B IIOJIb3Y 3TOfO rOBopHT II Te HenpepblB-
Hble lIepeXOp;bI, I\OTOpble CYIII;eCTBYIOT Memp;y HHMII: lIaparaHrJIJIOHap-
HaH TKaUb IIMeeT TeHp;eHD;IIIO OTp;eJIIITbCH OT raHrJIII03HOro CIIJIeTe-
HIIH 6pIOillHOll nOJIOCTII II CBH3aHa THmaMII C caMbiM KpynHblM na-
p8raHrJIIIeM - H8]J;nO'IeqHIIKOM (Kose, 1907; Muller, 1929; c06cT-
BeHHble Ha6JIIOp;eHIIH). B TO me BpeMJI Y B3pOCJIbiX oc06eH
MJIeI\OnIIT8IOIII;IIX rJIaBHaH Macca xpoMaiPiPlIHHblX I\JIeTOI\ y»\e p;ocTlIr-
118. H8p;nOqeq lHI\a; B 3M6plIoreHe3e me MJIeI\OnIlTaIOIqIIX I\apTIIHa
HaIIOMIIHaeT TaI\OBYIO Y B3pOCJIbiX IITIID;.
H Kose 1I0p; epI\IIBaeT, '{TO lIaparaHrJIlI1I Y Bcex IITIID; IIMeIOT CXOA-
Hae cTpoeHlIe II qTO Bce OHII IIoxomu Ha cYIIpapeHaJIOBbIll (HaA-
1I0 eqIU!ROBl>Iii) lIaparaHrJIlill. XpoMaiPiPIIHH8H peaI\D;IIH - eAIIHcT-
136
BeHH8R mlIpOlW paClIpOCTpaHeHHaH B Te rop;bI P;JIH rIICTOXHMH1JeClWrO
BblHBJIeHHH ap;peHaJIIIH01I0P;06HblX COep;IIHeHIIii: - 06HapymlIBaeT
60JIbmyIO BaplIa6eJIbJlOCTb ORpaClm RJIeTOH BO Bcex lIaparaHrJIHJIX,
rp;e 6b1 OHII HII paCIIOJIaraJIIICb, II OTpamaeT, IIO MHeHIIIO aBTopa,
pa3Hble q,a3bl ceHpeTopHoii: aRTIIBHOCTIi RJIeTOH.
ITo Mepe yp;aJIeHHH OT a6p;OMIiHaJIbHoii: 06JIaCTII B RpaHlIaJIbHOM
H Ray aJIbHOM HaIIpaBJIeHHHX ROJIuqeCTBO lIaparaHrJIHOHapHblX RJIe-
'TOR B raHfJIIIHX CHMlIaTHqeCHIIX CTBOJIOB y6b1BaeT, oc06eHHO y
B3pOC.'IbiX IITIIIJ;. CTelIeHb paClIpOCTpaHeHIIJI lIaparaHrJIlIeB B meii:Hoii:
II rpYP;Hoii: 06JIaCTHX. xapaRTep IIX B3aIIMOOTHomeHIIii: C oHpymaIO-
IQHMII TRaHHMII, I1X pa3BIITlIe, IIHHepBa IIJI II BaCKYJIHpII3a HH 01I1I-
caHbl MHorlIMH aBTopaMII (Kose, 1902, 1904, 1907; Terni, 1931;
Muratori, 1932a, b, c, d, 1933a, b, c, d; Watzka, 1930, 1931;
1943; Palme, 1934; Nonidez, 1935b; Chowdhary, 1953; ITplIii:MaR,
1960, 1962; Wirt.z, 1968; MOp030B, 1969, 1970, II P;p.). Kose (1907)
OlIlICaJI HX eJIYIO BepeHH y, ROTopaH HaqlIHaeTCH OT IIpep;cepp;IIii:
II 1I0]:(HHMaeTCH BP;OJIb COCYP;OB p;o RapoTIIp;HOro lIaparaHI'JIIIH,
a Muratori (1932c) OTMeTIiJI HX TecnyIO CBH3b C aHp;OHpIIHHblMII
opraHaMII ma6epHoro IIpOlIcxomp;eHlIJI II OHOJIO CIICTeMbi Baryca.
Kose YRa3aJI Ha BaplIa IIH COHHoro lIaparaRI'JIIIJI, ROTOpblii: B BHp;e
Op;IIHapHOro HJIH p;Boii:Horo oprana paCIIOJIOmeH 110 06e CTOpOHbl mell
II 06Hapy»mJI rpYlIlIbI lIaparaHrJIlIOHapHblX RJIeTOR tunica adven-
ticia 06IT(eii: cOHHOii: apTeplIlI,- HlIme 6IIq,YPRa HII ee Ha Hapym-
HyIO II BHYTpeHHIOIO, HO Bblme OTxomp;eHIIH 1I0]:(RJIIOquqHOii: apTe-
pHH (pIIC. 54). RpoMe HIIX, MeJIRHe lIapafaHrJIIIOHapHble Y3eJIRII
paClIpOCTpaHHIOTCJI B RpaHlIaJIbHOM Ha1IpaBJIeH1I1I 110 xop;y He60JIb-
mlIx COCYP;IICTbIX II HepBHblX CTBOJIIIROB. HapJlJ];y C rIICTOJIOruqe-
CHUM CXOP;CTBOM IIepeqlICJIeHHblX IIaparaHrJIIiOHapHblX Y3eJIROB aBTOp
IIOp;qepHHYJI CYIQeCTBeHHYIO pa3HII Y: Bce lIaparaHrJIIIII, CBH3aHHble
.c RapOTIlp;Hoii: apTeplIeii: ]:(pyrlIMII RPYIIHblMII cocyp;aMII mell II IIpep;-
Cepp;IIeM, COCTOHJIH H3 6eC BeTHbiX RJIeTOH. B TO me BpeMH B na-
paraHfJIIIOHapHblX RJIeTHaX, ROTOpble HaXOp;HJIIICb B CHMIIaTuqeCRlIX
CTBOJInHaX II raHrJIlIJIX meii:Horo II rpY]:(HOrO OT]:(eJIOB (y MOJIOp;OrO
p;p03p;a), OH Ha6JIIOp;aJI HeraTIIBHYIO peaR IIIO RaR IICKJIIOqeHlIe, TaR
me HaR B a6p;OMIIHaJIbHblX CIIMlIaTlIqeCRIIX HepBax (y MOJIOP;Oii: B
pOHbl II CTaporo RJIeCTa).
TIQaTeJIbHOCTb HCCJIep;OBaHHii: 06mHpHoro MaTeplIaJIa, ROTOpbIM
paCIIOJIaraJI Kose, CJIymlIT CBIIp;eTeJIbCTBOM 6eCCIIopHOCTII ero Ha6-
JIIOp;eHIIii:. A aTII Ha6JIIOp;eHHH HBJIHIOTCJI xopomliMII q,aRTUqeCRIIMII
.p;aHHblMII, 1I0p;TBepmp;aIOIT(IIMII BblCRa3aHHble HaMil 06IT(IIe co06pa-
meHnH B aToii: II IIpep;blAYIT(llX rJIaBaX 06 aBOJIIO 1IH CIIMTIaTOap;pe-
'HaJIOBoii: CHCTeMbi 1I03BOHO'IHblX. OHII CBO]:(HTCH R CJIep;YIOIT(eMY. ITpo-
IICXO]:(flIQaJI B IIpo ecce q,IIJIOreHe3a pep;yR IIH II «aTpoq,IIH»
aHCTpaa]:(peHaJIOBbiX yqaCTROB xpoMaq,q,IIHHoii: THaHH, ROTopaH HaCTY-
naeT B pe3YJIbTaTe q,YHR IIOHaJIbHoii: (<HeHymHoCTII», IIpeiKp;e Bcero
KacaeTCH TOii: ee qaCTII, ROTOpaJI cOCpep;OTOqeHa B IIpOII3BOP;HbIX iKa-
6epHbIx HpOBeROCHbiX COCY]:(OB - B RapOTIIAHblX, JIerOqHblX II aOpTaJIb-
HblX Ayrax. OP;HHM II3 IIplI3HaROB pep;YH HH HBJIHeTCH CHllmeHlIe
137
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JIaCTII nTlIl\
ao - aopTa; Cb - RapOTHgHoo Teno;
de - ROpOTHgHI>ItI npOTOR; dn-
genpeccopHHtI HepB; ic - BHYTpeH-
HHH RapOTBAHaH apTepnH; eC - Ha-
pymHaH RapOTBgHaH aPTepnH; X-
IJJIymJ\alOll{ut1: HepB; gn - g. nodo-
sum IJJIymAolOllIel'o HPpBa; IX-
f!abIRo-rJIOTO'lHHt1: HepB; gI X - l'aHI'-
mlt1: HabiRo-rnOTO'lHOI'O HepBo; go-
BepxHHt1: met1:HHt1: CnMnOTB"'leCHnit
yseJI; Icc - JIeBaH oIJll{aH COHHaf! ap-
TepuH; lig. a. - 60TaJInOBa CBf!Sl<a;
Ira - CBHaRa ROpHf! aopTbI; pa-
nel'O'lHaH opTepnH; pet - npeBepTefi-
paJIbHbIt1: npeRapOTHgHbIt1: HepBHbIt1:
CTBOmlR; pth - III H IV napaTH-
peoHJ\HbIe menesbI; rec - npaBaH oI'i_
ll{8H COHHaH opTepnH; 8a - nOAl<n
'D'I'lHaH opTepnH, thyr - ll]HTOBHA-
HOH meneso; t'a - nOaBOHOQH8fI
apTepHH (Adams, 195H. BHgOHaMe.
HrHo)
iBblpaOOTRlI clleqlIllmqecRIIX rOpMOHOB, ROHqeHTpaqlIJl ROTOphlX B
RJIeTRaX OKa3b1BaeTCH HmRe 1I0pora qYBCTBIITeJIbHOCTII xpoMa<I><I>IIH-
HOH peaRqlIlI. Y IITlIq B )TOH 06JIaCTn OHa Bcerp;a oTpnqaTeJILHa.
B TO me BpeMH HalI60JIee aRTIIBHaJl ap;peHaJIOBaJl <I>YHRqIIJI B 8BO-
JIloqnlI II03BOHO'lHblX IIOCTeIIeHHO cocpep;OTOqlIBaeTCJI B 3ap;HeM OT-
p;eJIe TYJIOBlI a, B ROTOpOM pa3BIIBaeTCJI clleqllaJIbHblH 8HP;ORpIIH-
HWH opraH - Ha;:J;IIOqeqHIIR H R ROTOPOMY CTHrlIBaIOTCH xpoMa<I><I>IIH-
HbIe 8JIeMeHTbI. B oRpymemlH 81'OrO opraHa Y IITlIq coxpaHHeTCH erqe
aRTIIBHOCTb 8RCTpaap;peHaJIOBbiX xpoMa<I><I>IIHHblX 8JIeMeHTOB B CTeH-
Rax MarlICTpaJIbHblX COCYP;OB. Op;HaRO IIpOTJDKeHHOCTb COCYP;IICTblX
yqaCTROB, B ROTOpblX BCTpeqaIOTCH BHYTplICTeHOqHble rpyrrllbl n
mraCTbI xpoMa<I><I>lIHHblX RJIeTOR, Y IITlIq MeHbme, qeM Y IIpeCMbI-
RaIOrqIIXCH: RallplIMep, Y HrqeplIq aRTIIBIIO <I>YHRqllOHHpyIOrqlIe RJIeT-
]{1I BCTpeqaIOTCJI He TOJIbRO B CTeHRax a6p;OMliHaJIbHOH aopTbi B6JIH-
311 Ha;:J;1I0qeQHIIROB, HO TamRe - XOTJI II B MeHbmeM ROJIlIqeCTBe-
B rpyp;HOH ee qaCTII (CM. plIC. 41, B). 3aTo Y IITlIq pa3BllBaIOTCJI
HOBbie xpoMa<I><I>lIHHbIe OCTpOBRII BP;OJIb qpeBHblX HepBOB II IIX MeJl-
.RlIX pa3BeTBJIeHIIH, II OHII, RaR IIpaBIIJIO, pe3RO xpoMa<I><I>llHHbI. Ro-
Boe HaCJIOeHlIe xpoMa<I><I>lIHHblX lIaparaHrmleB, IIpOHBJIHIOrqeecH B
Hau60JIee OTqeTJIIIBOM II 6YPHOM pOCTe lIX Y IITlIq, COOTBCTCTByeT.
OqeBII]:(HO, Ha1l60JIee IIHTeHCIIBHOMY Mop<I>OJIOrll'IeCROMY pa3BIITBIO
-cliMIIaTlI'IeCROH HepBHOH CIICTeMbi (Rirt, 1934). ECTecTBeHHo, qTO
B a6p;OMIIHaJIbHOH 06JIaCTII, rp;e coxpaHHIOTCH erqe 8RCTpaap;peHaJlO-
Bble lIaparaHrJIlI1I II rp;e HaCJIalIBaIOTCJI 110 XOAY CIICTeMbi qpeBHblX
flepBOB <I>llJIOreHeTllqeCRll HOBbie xpoMa<I><I>nHHble 8JIeMeHTbI, OHII Ii"
Bcerp;a pa3JIIIqlIMbi P;Pyr OT P;PYJ'a.
B pa60Tax IIpamCRIIX IICCJlep;OBaTeJleH Watzka (1931, 1933) B
Palme (1934), 1I0cBHrqeHHblx cpaBHIITeJIbHOMY II3yqeHIIIO meHHO-
rpYP;Horo OTp;eJIa HeROTopblX 1I03BOHOqHblX, B TOM qllCJIe IITlIq, 110
cy eCTBY 1I0p;TBepmp;eHbI OCHOBHbIe <I>aRTbI, 1I0JIyqeHHble Kose. Op;Ha-
RO 8TII nCCJIep;OBaHlIH IIpep;lIplIHIIMaJIIICb yme C P;PyrHMII qeJIHMII B
6b1.'I1I cp;e.TIaHbl P;PyrlIe BbIBOp;bI.1 TIocJIe BbI]J;BIIHYTOH de Castro
ROHqelIqllll 0 rJIOMycax-perYJIHTOpaX RpOBHHoro p;aBJIeHBH Watzka
(1943), 1I0p;TBepAHB pa60Tbi cBoero yqlITeJIH Kohn 0 cyrqecTBoBaHHB
xpoMa<I><I>lIHHblX II HexpoMa<I><I>IIHHblX lIaparaHrJIlIeB, BblCRa3aJICJI, op;-
HaRO, B 1I0JIb3Y llX HepaBH03HaqHOH <I>YHRqlIlI: xpoMa<I><I>IIHHble pac-
CMaTpnBaJIIICb 1I0-lIpemHeMY B RaqeCTBe BbI]J;eJIllTeJIbHblX 8HP;ORpIIH-
HblX opraHOB, HexpoMa<I><I>IIHHble - B RaqeCTBe peqellTopHblX. Wat-
2ka nCCJIep;OBaJI C 8TOll 1I03IIqlln RapoTllp;HblH IIaparaHrJInH,
Palme - IIpnCepp;e'IHble.
Watzka II Palme 1I0p;p06HO IICCJIep;OBaJIII Y IITlIq RPYIIHbIe co-
CYP;bl melI, IIplICepp;eqHYIO 06JIaCTb II IIplIJIemarqlIe yqacTRl1. ABTO-
pbl, CCblJIaHCb Ha oc06eHHocTn pa3BeTBJIeHIIH COCYAIICTblX CTBOJIOB
Y IITnq II MJIeROlIlITaIOrqnx II Ha HepaBH03HaqHOe paCllpep;eJIeHlIe B
HIIX lIaparaHrJIlIeB, OTMeTIIJIlI, qTO TOqIIYIO IIpeeMcTBeHHocTb YCTa-
HOBIITb TPYP;HO. lIMn OlllIcaHLI HexpoMa<I><I>IIHHble lIaparaRrJIlllI B o
U\eH COHHOH .apTeplIlI, JIerOqHOH apTeplIll, Ayre aopTbi II ee HIICXo-
J{m:u;eii 'laCI'II II Macca MeJIRliX RJIeTOqHblX rpYIIII B oRpymaIOrqnx
139
TKaWIX - OKOJlO IIpaBOrO 6poHXa, Memp;y JlerKlIM II aopToii:, BIIYTPB
HepBHblx CTBOJlOB, B BeHeqHoii: 60p03p;e cepp; a II 3mmapAe. 11H-
TepecHbl co06rqeHHble HMII p;eTamI, xapaKTeplI3YlOrqHe B3alIMOOTHO
weHlIH lIaparaHrJlIIOHapHoii: TI aHII CO CTeHKaMII KpoBeHocHblX cocy-
P;OB II C HepBHblMII CIIJleTeHHJDlII. B 06J1aCTII 06rqeii: COHHOii: apTepIUJ
BCTpeqalOTCH lIaparaHrJIlIOHapHble My«I>TbI Ha IIpeKamWJlHpHblX cocy-
p;ax, cocTOHrqlIe II3 HexpoMa«I>«I>HHHblx KJleTOK. B JlerO'IHbIX II aop-
TaJlbHblX p;yrax My«I>TbI OqeHb pep;KII II IIpe06J1ap;aIOT CBH3H C HepB8-
Mil: IIaparaHrJlHOHapHbIe rpYIIlIbI 3aKJlIO'IeHbI B rYCTYIO ceTb raHrJlH-
03HblX RJleTOK H HepBHblX BOJlOKOH, B 60JlbWlIHCTBe CBoeM MHKOTHblX,
(Palme, 1934). qeM 6J1IIme K cepP;JUT, TeM HepBHoe CIIJleTeHlIe CTa-
HOBIITCJI ryrqe II TeM 60JlbWe B HeM pa36pocaHO MeJlKIIX lIaparaH-
rJlHOHapHblX yseJiKOB, KOTopble qarqe JlemaT BIIYTPII HepBHbIx CTBO-
JlIIKOB. B HapymHoii: 060JlOqKe HlIcxop;Hrqeii: qaCTII aopTbl IIMelOTCJI
lIaparaarJlIIOHapHbIe KJleTKII II Heo6:WIaii:ao rYCToe HepBHoe CIIJleTe-
HlIe, lIS KOTOporo BOJlOKHa IIpOHlIKalOT B tunica media. CTpYKTypa
tunica media, cocToHrqaH 1I3 perYJlHpHo cMeHHlOrqlIx p;pyr p;pyra
MblWeQHblX CJlOeB II 3J1aCTlIqeCKIIX MeM6paH, HapYlliaeTcH MeJlKHMH
HexpoMa«I>«I>IIHHblMH KJleTOqHblMII rpYlIIIaMH II COeP;IIHHTeJlbHOB
TKaHblO. He JlHllIeHO lIHTepeca co06pameHIIe Palme, BblCKa3aHHoe
IIM 110 IIOBOP;Y Tex IIaparaHrJlIIOHapHblX rHe3p;, KOTopble OH BlIp;eJl B-
tunica media: 1I0p;06HaH JlOKaJlII3a H CBH]J;eTeJlbCTByeT, no aBTopy,
npOTHB IIHKpeTopHoii: «I>YHK HII lIaparaHrJlIIOHapHoii: TKaHII, TaK KaR
He HBJlJIeTCH IIOP;XOP;HrqIIM MeCTOM AJlJI CeKpe IIII cop;epmlIMoro KJle-
TOK B KpOBb IIJlII JlHM«I>y. B TO me BpeMH Palme p;OIIYCKaeT B Ka-
qeCTBe rlIlIOTe3b1 BamHYIO pOJIb npHCepp;eqHblX II KapOTIIp;HblX lIa-
paraHrJllIeB BMeCTe C HepBHblMH OKOHqaHHHMII Ha cocyp;ax B pe-
enn;lIH. HaM KameTcH, TaKoe AOlIyrqeHlIe JlIIWeHO BCHKoro
OCHoB.aHIIH, nOToMY qTO JlOKaJlII3a IIH lIaparaHrJlIIOHapHblx RJIeTOK B
tunica media JlBJlHeTCH erqe 60JlbWHM IIpelIHTcTBlIeM P;JlH BOClIpllJl-
TIIH XIIMHqeCKIIX CP;BlIrOB, qeM ]:(JlH BbI]:(eJleHHH ropMoHoB, TaK KaR
rJlaBHOe PYCJlO KpOBII JlemlIT BeCbMa p;aJleKO OT KJleTOK.
ABTOpbl pHp;a p;pyrlIx pa60T 30-x roP;OB BCTanH Ha erqe 60Jlee
KpaiiHlOlO 1I03II HIO II BCJlep; 3a 1I0HCKaMII pe«I>JleKCOreHHblx 30H y
MJIeKOlIIITalOrqlIx CTpeMHJlIICb Haii:TII y IITIII!; aHaJlOrH'IHbIe 06J1aCTII.
BOClIpHHIIMalOrqHe HapHp;y C 1I3MeHeHIIJIMII XHMHq CKOrO eOCTaBa
KpOBII TaIDKe II KOJle6aHIIH p;aBJleHIIH B COCYAHCTblX CTeaKax. nOHCKD
CBOP;IIJlIICb K BblJIBJleHlI10 yqacTKa 06rqeii: COHHOii: apTepHD MOp«I>O-
JlorlI'IeCKII rOMOJlOrlI'IHOrO CIIHYCY MJleKOlIlITalOrqlIx II K H3YQeHIIIO
ero IIHHepBa IIII BMeCTe C IIHHepBa IIeii: KapoTHp;Horo TeJla (Terni,
1931; Muratori, 1932a, b, c; 1933a, b, c, d; Nonidez, 1935,
II 1I03me Chowdhary, 1953). 8TH aBTOpbl 1I0ATBepp;HJlH p;aHHbl&
npep;blp;yrqIIX DCCJlep;OBaTeJleii: (Watzka, Palme) 0 TOM, qTO xapaKTep
pa3BeTBJleHIIii: KPYIIHblX KpOBeHOCHblX COCYP;OB Y IITIII{ He COOTBeTCT-
ByeT TaKOBOMY Y MJleKOIIHTalOrqlIx II, CJlep;OBaTeJlbHO, YCTaHOBHTb Y
HIIX rOMOJlorlI1O KapOTlIP;HOrO clIHyca TPYP;HO. RpOMe TOro, YKa3aHHbIe
aBTOpbl HaWJlII 1I06J1II30CTII OT COHHOii: apTepIIII p;pyrlIe MaCCbl IIa-
paraHrJlIIOHapHoii: TKaHlI, IIp;eHTII'IHble IIO :qDTOJlOrlIqeCKIIM npn3Ha-
140
KaM II pa36poCaHHLle B pa3HLlX YTJaCTKaX II B pa3HLlX KOJIHqeCTBaX.
MHLlMII CJIOBaMlI, OHII npmuJIII K TeM me pe3YJIbTaTaM, TJTO II Upa>R
CKlIe IICCJIeJ];OBaTeJIII. ITaparaHrJIlI1I MOrJIII 6L1Tb xpoMa4>4>IIHHLlMB II
HexpoMa4>4>IIHHLlMII (apreHT04>IIJIbHLlMII II HeapreHTo4>IIJIbHLlMII) II
OTJeHb MHOrOqlICJIeHHLlMH Y HeKOTopLlX BIIJ];OB nTII (Muratori,
1932a, 1933a). M3-3a KOHTaKTOB IIX C pa3HLIMII TKaHJlMII II
TJaCTOrO HaxmKJ];eHlIJI B TaKHX MeCTax, rJ];e Y UTII He YJ];aBaJIOCL
IIOJIyqlITb IIpeccopHLle 34>4>eKTbI (Linden van der, 1934; Jung, 1934),
OTJeBIIJ];HO, B03HHKJIa TeHJ];eH HH HOBOR KJIaCclI4>IIKa BII uaparaHrJIII-
eB. Te II3 HlIX, KOTOpbIe 6b1JIII HaiiJ];eHLI B CTBOJIaX 6JIymJ];aIOm;ero
HepBa, aBTOpLl Ha3BaJIII «BnyTpIIBaraJILHLIMII» (Terni, 1931; Murato-
ri, 1932a, b, c), HJIH <meCIIMDaTHTJeCKHMII», a, CJIeJ];OBaTeJILHO, «He-
HCTIIHHbIMH). K TOMY me OHII 6L1JIII (meXpoMa4>4>IIHHLIMII». TOJIbKO
TOT yqacToK COHHOR apTepIIII IITII , KOTOpblR paCIIOJIOmeH HaJ]; OT-
xomJ];eHlIeM IIOJ];KJIIOTJuqHOR apTepIIII II IIplIJIeraeT K ganglion nodo-
sum 6JIymJ];aIOm;ero HepBa, 6L1JI IIplI3HaH roMOJIOroM KapOTIIJ];Horo
cHHyca MJIeRODIITaIOIltHX Ha OCHOBaHlI1I CKOIIJIeHHH «3DIITeJIII01IJ];HLlX
KJIeTOK)) (IIoJ];06HbIX KJIeTKaM KapOTHJ];HOro TeJIa MJIeROIIIITaIOm;IIX)
H IICTOTJHIIKOB IIHHepBa IIII, KOTOpaJl OKa3aJIaCb HJ];eHTII'lHOR MJIeKO-
DIITaIOm;HM. B pe3YJIbTaTe II3yqeHIIJI HepBHoro cHa6meHHH 3TOii 06.:.
aaCTII Muratori (1932c, 1933a,b,c,d) 3KCDepHMeHTaJIbHO IIOKa-
3aJI, TJTO MJlKOTHbIe BOJIOKHa tIYBCTBIITeJIbHOro xapaKTepa II3 KJIeTOK
ganglion nodosum (DpeHMym;ecTBeHHO), a TaKme BeTBII H3L1RO-rJIO-
TOtIHOro HepBa (Terni, 1931; Ask-Upmark, 1935), IIPIICYTCTBlIe KO-
TOpblX Y IITH OCDapHBaJIB Nonidez (1935b) II Chowdhary (1953),
HBJIHIOTCJI a4>4>epeHTHbIMII IICTOTJHIIKaMlI ee IIHHepBa IIII, a CIIMDa-
TuqeCRHe HepBLI - 34>4>epeHTHLlMII. B HeJ];aBHIIX pa60Tax E. K. Mo-
p030Ba (1969, 1970) 1I0J];TBepmJ];eHO yqaCTHe Jl3L1KO-rJIOTO'lHoro Hep-
Ba B IIHHepBa HII RapoTIIJ];HOR 06JIaCTII nTII . JJ;aHHLlR Y1J:aCTOR
06m;eR COHHOR apTepliH IITII OTJIlItIaeTCJI OT RapOTIIJ];Horo clInyca
MJIeRODHTaIOm;IIX TOJIbRO CTpoeHHeM caMOR CTeHRII cocYJ];a, B ROTO-
pOM He Ha6JIIOJ];aeTCH IICTOHTJeHIIJI MblmeqHOrO CJIOH, xapaRTepHoro
J];JIH MJIeRODIITaIOm;HX.
AopTaJIbHoMY DaparaHrJIIIIO 6b1JIa IIpmmcaHa Ta me 4>YHR HH
pe4>JIeRCOreHHoro opraHa, RaR II RapOTIIJ];HOMY Teny, Ha OCHOBaHIIB
rlICTOJIOrllqeCROrO CXOJ];CTBa II IIHHepBa IIII (Muratori, 1937).
HeJ];ocTaTOqHaH apryMeHTa IIH BLlBOJ];OB 0 4>YUR IIOHaJIbHOR pOJIH
RapOTIIJ];HLlX II aOpTaJIbHO-JIerOqHLlX DaparaHrJIlIeB y IITII HaXOJ];HT
OTpameHlle II B ('OBpeMeHHbIX pa60Tax. JJ;ame Te IICCJIe!{OBaTeJIII..
ROTopble paccMaTpHBaIOT RapOTII]:(HYIO o6JIacTb MJIeRODIITaIOm;1IX RaK
peq}JIeRCoreHHYIO 30HY, COMHeBaIOTCJI B aHaJIOrlIqHOR 4>YHR IIII ee-
Y IITII . TaRoe 3aRJIIO'leIllIe MLI HaXOJ];HM B MOHorpa4>IIII pbHHoro
CTOpOHHHRa 3TOR TeopUIi Adams (1958). BO-IIepBLlx, aBTOp IIp II-
JJ:epmHBaeTCH MHeHHH, qTO Y IITII HeT RapOTIIJ];HorO cHHyca, IIOJ];06-
HO TaROBOMY Y MJIeRODIITaIOIqIIX. Bo-BTOpblX, pa3BIITIIe HapymHoH'
COHHon apTepHII DTIII.\, OTJIuqHoe OT pa3BIITIIJI ee Y MJIeKOIIIITaIO-
m;IIX, 3aTpYJ];HHeT TOqHYIO JIORaJIH3a IIIO rOMOJIOrB'lHbIX o6.'1acTeii y
Tex II J];pyrIIX. M, HaROHe , B-TpeTbIlX, J];JIJI IIplI3HaHIIJI rOMOJIOrlIU
f41
Adams cqnTaeT 06HaaTeJIbHblM IIPIICYTCTBne KJIeTOK, KOTOpbIe, Bbl-
nOJIHHH KaK II B RapoTII HOH 30He MJIeKOnnTaro nx pOJIb cIIe n n-
TJeCKnX pe enTopHLlX aJIeMeHTOB, OJImHLI 06HaaTeJIbHO JIemaTb n
.a BeHTII nn apTeplIn. IIplIcYTcTBne B llIeHHOH 06JIaCTn nTnI He O -
Horo, a HeCROJIbKIIX lIaparaHrJIIIOHapHLlX Y3eJIKOB aaCTaBJIHeT nCKaTb
cpe n HIIX KaROH-JIII60 O IIH, KOTOpLlH COOTBeTCTBOBaJI 6L1 KapOTII -
HOMY Teny MJIeKOIIIITaro IIX II 6L1JI 6bl OTBeTCTBeH 3a pel.\enTopHYro
YHK IIro aTOH 06JIaCTII. O HaRo Adams, OCHOBblBaHCb Ha aHaJIlIae
JIIITepaTypbl II Ha OqeHb no p06HblX IICCJIe OBaHIIHX I.\nTnpyeMoH
JdM RCCepTa IIII Chowdhary (1953), IIplIaHaeT, TJTO 110 I.\HTOJIOrn-
'leCKnM II ame nTOXIIMnqeCKIIM nplI3HaRaM Bce naparaHrmllI O II-
HaKOBbl. OH, Bn IIMO, C COmaJIeHlIeM OTMeqaeT, qTO JIemaT OHII rpoa-
HMn He TOJIbKO B tunica adventicia, HO II B tunica media, n qTO
B pH e YTJaCTKOB KapoTII HOH 06JIaCTn nTnl.\ nCTpeqaroTcH KaK ap-
rlIpO IIJIbHbIe, TaR II He aprnpO HJIbHble RJIeTRH, qTO, HaKOHel.\,
e IIHCTBeHHbIM OTJIlIqlIeM naparaHrJIHeB Pyr OT Pyra MorYT CJIY-
mlITb paaMepbl aTHX yaemmB II qTO, BepOHTHO, HalI60JIee RpynHoe
IIa HIIX eCTb «RapOTH Hoe TeJIO». He060CHOBaHHOCTb TaKoro OBo]:(a
KaK BeJIIITJIIHa Y3JIa, HR06bl CJIyma aH B nOJIb3Y nplI3H8HnH 3a
(<KapOTII HbIM TeJIOM) nTH oco60ro, He CBOHCTBeIlHoro pyrlll\l
naparaHrJIIIHM xapaKTepa, OTJeBII Ha. A Bbl3BaHa OHa Bce TOH me
He06xo IIMOCTbro JIH aBTopa BLI eJIIITb cpe II Bcex HexpoMaifJ<I>HH-
HLIX naparaHrJIlIeB IITII CTPYRTYPY, KOTopaH 6L1JIa 6bl OTBeTCTBeH-
Ha 3a xeMope elITopHYro YHK IIro.
B pa60Te Wirtz (1968) o6CJIe]:(oBaHo cep]:(l.\e II npncep]:(eqHbIe
COCY LI roJIy6eH II RYP Ha ceplIHHLlx cpeaax. OH onlicaJI MHoro-
TJIICJIeHHLle rpynnLl a peHaJIOBbiX KJIeTOR Y nepe HeH CTeHRn JIe-
BOH JIeroqHOH apTeplIlI, Ha aa HeH CTeHKe JIero'lHoro CTBo.la, B yr-
JIY Mem y aopToH II BeTBbro JIeBOH KopoHapHoll apTeplIlI, a TaKme
Mem y ]:(yroll aopTLI, JIerOTJHOH apTepHII II OCHOBaHlIeM cep l.\a.
ABTop 1I0]:('IepRHYJI, qTO, HCCJIe YH cep e IITn , OH HallleJI IIapa-
faHrJIlIlI TOJIbRO y MeCTa OTXOm,iJ;eHHn KopOHapHblX COCY]:(OB OT
KpyrrHLlX COCY OB rpY Holl nOJIOCTIi II He BCTpeTnJI nx Ha nYTII
aJIbHelllllero CJIe]:(OBaHIIH II B pa3BeTBJIeHHHX BeHe'lHLlX COCY OB.
OH OTMeTIIJI TaK}l{e OTCYTCTBHe xpoMaifJ<I>nHHoH TKaHH B caMOH
Mblllll.\e cep l.\a. B COOTBeTCTBlI1I C 3a)];aTJeH, nOCTaBJIeHHOll Wirtz,
qT06bl BLlHCHHTb xeMope eIITopHYro pOJIb rJIOMycHOH TRaHII JIH pe-
rYJIHl.\nlI KopoHapHoro KpoBo06pa eHIIH, aBTOp cTpeMIITcH o eHlITb
IIOJIyqeHIIbIe peaYJIbTaTLI. OH nOJIaraeT, TJTO rJIOMYCLI He cnoc06Hbi
YJIaBJIlIBaTb na eHlIe HanpHmeHIIH RnCJIOpO a B paaBeTBJIelll1HX KO-
pOHapHbIX COCY OB, TaR KaR B TOJIl e MIiORap a OH IIX He HameJI;
IIX pacnpOCTpaHeHne OrpaHlJ'IlIBaeTCH KPYIIHblMn cocy aMIi rpY HOll
IIOJIOCTII II YCTbHMIl BeHeqHblX apTepliH. OqeHb cnpaBe JIIIBO, Ha
Ham B3rJIHJ];, II ero IIplIaHaHlIe, KOTopoe OH eJIaeT Ha OCHOBaHlI1I
DTepaTypHblX nCTOqHIIKOB 0 TOM, TJTO xpOMa nHHble rpaHYJILI B
pe enTopHblX (rJIOMYCHbIX) RJIeTKaX IIMeroT HeHCHoe aHaqeHlIe.
IIpIIBe eHHLle Wirtz aKTLI IIO TBepm aroT IIpaBIIJIbHOCTb cpaB-
HIITeJIbHO-Mopil>OJIOrlIqeCKoro IIoAxo a, ROTOpblH IIpOBO IITCH B Ha-
142
CTOJuD;ell pa60Te, )];JIH il>YHRD;IIOHaJIbHOH oD;eHlm: lIaparaHrJIHeB BLlC-
IlIIIX 1I03BOHO'lJILlX: IIX OCTaTRII coxpaHHIOTCH elD;e B cocy)];ax ma-
6epHoro IIpOHCXOm)];eHIIJI, HO IIX HeT B pa3BeTBJIeHHJIX RopOHapHblX
COCY)];OB cep)];D;a, ROTOpbIe He CBOHCTBeHHbl 6eccOCY)];IICTOMY cep)]; y
KpyrJIOpOTbIX II HBJIHIOTCH CTpYRTYpaMII II03)];Hemuero IIpOlIcxom-
)];eHlIH.
M3y eHlIe IIaparaHrJIHeB IITIID; II IIX pa3BIITHH C IICIIOJIb30BaHlIeM
COBpeMeHHUX MeTO)];OB rlICTOXIIMHqeCRoro aHaJIII3a Ha aJIOCb B 60-x
op;ax (ilplIHMaR H CMIITTeH, 1959; IIpIIHMaR, 1960, 1962; CMHTTeH
H MaJIIIHOBCRaH, 1960; Muratori, 1962; Muratori, Battaglia, Mo-
donesi, 1965a, b; CMIITTeH II Ey)];aHD;eB, 1968; CMIITTeH, 1970,
1971) .
3a)];aqa HaIDHX IICCJIe)];OBaHlIll 3aRJlIO aJIaCb B cpaBHHTeJIbHOM
II3yqeHIIII 3M6plIoreHe3a II rlICTOXIIMlIqeCRIIX CBOHCTB llIeHHo-rpy]:(-
HLIX lIaparaHrJIlIeB IITIID; (ilplIHMaR II CMIITTeH, 1959; IIpHiiMaR,
1960, 1962) H lIaparaHrJIlIeB 6pIOlliHOH IIOJIOCTII - M03roBOro CJIOJl
Ha)];IIOqeqHIIROB II IIapaaOpTaJIbHLlX CROIIJIeHIIH (CMIITTeH II MaJIH-
HOBCRaH, 1960).
EOJIbIDOii: cpaBHIITeJIbHblH MaTepHaJI, IIMeBIDIIHCH B paClIopHme-
HIIII 8. X. IIpHiiMaR, 1l03BOJIIIJI eH YCTaHOBIITb, TO IIaparaHrJIH-
OHapHble Y3JILI meHHo-rpY)];HOH 06JIaCTII IITII , RaR RPYIIHble (Ra-
POTH)];HOe TeJIO II OROJIOBaraJIbHblH lIaparaHrJIHH), TaR II MeJIKlIe
(CBH3aHHble TOlIorpail>n ecRII C lIapaCHMlIaTuqeCRIIM OT]:(eJIOM HepB-
HOll CIICTeMbI IIJlIf paCIIOJIOmeHHLle B COCTaBe CIIMlIaTII eCRIIX HepB-
HblX CTBOJIIIROB), CXO]:(Hbl Mem)];y C060H II C RapOTII)];HLlMH lIapa-
l'aHrJIIIHMH B3pOCJIbIX MJIeROlIlITaIOlD;lIX - RpOJIIIROB, RpblC, MLlllIeH,
I\lOpClmX CBIIHOR, ROllleR, lIeC OB (l1pnHMaR, 1962). 06a KPYIIHblX
Y3JIa HMeIOT )];OJIbqaTYIO CTPYRTYPY, Ram)];aH II3 )];OJIeR IIpe)];CTaBJIe-
Ha rpYlIlIoH clIeD;nil>II t IeCKIIX lIaparaHrJIHOHapHblX KJIeTOR C RaIlHJI-
JIHpaMII CIIHYCOII)];HOro TlIlIa. B Ram)];OM lIaparaHrJIIIH BCTpe aIOTCJI
KJIeTI\:1I ]:(BYX TIIIIOB. O)];HII II3 HIIX OBaJIbHoii:, oKpyrJIOH IIJIH UOJIII-
rOHaJIbHoii: il>OPMbI, C RPYIIHblMH CBeTJIblMII IIY3bIPbROBUP;HLlMII H]:(-
paMH, C MaJIblM CO)];eplKaHlIeM xpOMaTIIHa II 1-3 H)];pLlmRaMlI. B H -
pax 3TIIX KJIeTOK Ha1l60JIee CHJIbHO pa3BIIT OROJIOH)];pblmRoBbIH
xpOMaTllH, IIpe)];CTaBJIeHHLlii: RpYIIHblMII rJIbl6RaMII, IIHOr)];a IIpeBbl-
maIOID;HMll pa3MepbI H)];pLlmRa; OCTaJIbHOH BHeH)];pLlmRoBblll xpOMa-
TIIH IIpe)];CTaBJIeH IIeIKHbIMII rpaHYJIaMII. IITOIIJIa3Ma B TaRllX KJIeT-
Rax CBeTJIaH, CJIaOO IIllpoHIIHoil>llJIbHaH, HHOr)];a BaRYOJIII3IIpOBaHHaH.
RpOllw 3THX RJIeTOR B COCTaB )];OJIeR BXO)];HT RJIeTRII C Menee pa3-
BIITblM 060)];ROM IITOIIJIa3Mbl, TeMHOORpameHHblM H)];POM C ROMlIaRTHO
paCIIOJIOIKeHHblM xpOMaTHHOM II 60JIbmllM llCJIOM H)];pLlllleR (3-
7). IITOIIJIa3Ma IIX IIllpoHllHoil>IIJIbHa (pllC. 55).
Ha6JIIO)];eHHJI Ha)]; IIpOD;eccaMII pa3BIITIIH RJIeTOR B 6PIOIIIHLlX lIa-
paraHrJIHHX IITII (CMllTTeH II MaJIIIHOBCRaH, 1960) II cpaBHeHII9
HX C meHHo-rpY)];HLlMll (ilplIHMaR II CMIITTeH, 1959; IIplIHMaR, 1960,
1962) IIpHBeJIH R y6em)];eHIIIO 0 D;IITOJIOrnqeCROM e)];IIHCTBe II O]:(H.
HaKOBLlX IICTOqHllRaX HX pa3BIITIIH. Pe3YJIbTaTLI 3TIIX pa60T, IIO)];-
RpelIJIeHHLle aBTOpa)];IIOrpail>II eCRIIM II3yqeHIIeM pa3BIITIIH 6pIOmHLlx
143
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IIICTII"JlOBbIM 3eJlCIIJ.\JII-nll pOllllllOI\l
OK. X 7. 06. X 1,0 (npenapaT 3. X. IIpHIlMaH)
naparaHrmWB IITII:Q (CM. rJl. VIII), ]:(aJlJl HO;IMOilillOCTL IIpU;lHaTb,
qTO xpoMaiPqJJlllo6J1aCTbi pa3.BIIDaIOTCJI H3 Toii nCTll1l I\JIPTO'llILIX
nOIIYJIH:QlIll, KOTopwe IIO BbIXO)];e 113 Heiipo:.JI..TO; Cplll<\JlhIlOii ,13 "':Ia)];KJI
.)];IIiPiPepeH:QHpYIOTCH B CIIOHrHOO.'1aCTUqeCI\01I1 lIanpan;tellllil. O)];I10ii
II3 CaMblX Xapal,TepHblX qepT 3TIIX CIIOHrUofi.TIaCTlI'IeCI,IIX I..;ICTOI, HB-
JlHeTCH MCJlIWe, 60raTOe XpOMaTIIIIOM H]:(PO C Y31UlM OfiO,L(IWM :QIITO-
IIJla3MLI. ITO)];06HLIC R.TIeTKII OIlHCaHLI MHOrO'IIIC;ICII"hllllll allTOpalll1l Y
pa3HblX il,IIBOTIIbIX, B pa3HbiX lIaparaHrJlIIHX II 13 pa;lIIhlX yy.aCniaX
TeJla. B Bllp,e 60JlblllllX 1IJ111 MeJlKIIX CKOn:lCnilii IIX "aO;IID;J:aJlII B
3M6pHOreHC3e, Y MOJlO)];LIX n B3pOCJlbiX oeofieii (Kose, lUU7; Smith,
1924; Hclt, 1925; de Castro, 1928; Watzka, 194R; Hollinshcad,
1943; CMllpHoBa, 1948; Langer, 1952; Ross, 1957. 19,")9, II )];p.)
II TpaKTOBa:m I10-pa3HOIIIY nan )];ereHCpaTilBlILIC qJOplllLl, .).Jym..:QlIo-
HaJibIlbIe CTa]:(UII, PY]:(III1ICIITapHLIC :J.'1CMeIlTLI If pcmc - mil.. pC3epll-
HLrii MaTepnaJi ]:(JlH nOlloo6pa30llallllH xpoMml)( nIlHLlx :JJleMeHTOB.
lICIIOJlb30mmlle aBTopa]:(lIorpaqJIlIl, ]:(olIycKalOuwfi lIanUOJlee 06'b-
enTHBIIblii aIlaJlH3 IICCJle]:(YCMoro MaTCplIaJla, TIm..a3aJl, 'ITO MeJlIW-
H)];epIlLIe KJlenHI coxpmIHIOT CIlOCOOHoeTb I.. MIITOTlltICCnO IY ]:(eJle-
HHIO B TeqCIIlle ]:(JlIITCJILHOro nepno)];a 3Mfipllorencaa n lIBJlHIOTCH
MaTepllHclwii n:JleTlwii, 113 KOTOpOll iPOpMllpYIOn"H xpoMaiPcPlIIIO-
6JIaCTbI. B IIpOTIIBonOJlOinIlOCTb 11M CIIMIIaT06J1aCTbI, 3a I.;oTOpLI 1ll
144
no 0611\eMY IIplI3HaHIIIO 3aKpelIIIJIaCb pOJIb IIpOIl3BO)];IITeJIeH II, cne-
)];OBaTeJIbHO, IICTOTJHIIRa XPO)Ia4>il>HHHbIX )JIeMeHTOB, TepHIOT ClIoc06
HOCTb K MlIT03Y II IIpeBpam;aIOTCH B aM6plIOHaJIbHLle CIIMlIaT0I.\IITbl
3a)];OJIro )];0 Toro, KaR KOHqaeTCH B03HlIKHOBeHlle II C03peBaHIIe XpO-
Ma4>QmH06JIaCTOB (CM. rnaBY VIII).
HapH)];Y C 06m;IIMII TJepTaMII, xapaKTeplI3YIOm;IIMII Mopil>onorlI-
qeCKOe C03peBaHIIe lIaparaHrJIlIeB 6pIOlliHOH II llIeHHo-rpY)];HOH 06
JIaCTeH IIT II CXO)];CTBa IIX C TaKOBblMII Y MJIeKOlIlITaIOlI\lIX, lIMe-
IOTCJI II MemBII)];OBble PI13JIIITJIIH. 110 na6JIIO)];eHIIHM IIpIIHMaK, aTO
B lIepBYIO Oqepe)];b 60JIbmaH IIJIII MeHblllaJl BbipameHHOCTb P;BYX KJIe-
TOqHblX TIIIIOB, a TaKme CTelIeHb IIlIpOHIIHOil>IIJIIIII I:(IITOIIJIa3MU.
Y IITIII.\ KJIeTKII lIepBOrO Tllna C KPylIHblMII H)];paMII IIMeIOT 60JIee
DLlpameHHYIO IIlIpOHIIHOil>IIJIbHYIO 3epHIICTOCTb, qeM B aHaJIOrlITJJILlX
KJIeTKaX MJIeKOIIIITaIOm;HX (lIpooMaK, 1962), a Y aMil>II6IIH IIHpOHII-
HOil>IIJIIIH em;e 60JIee OTTJeTJIIIBa, oc06eHHo Y MOJIO)];LlX oc06eii (CMIIT-
TeH, 1963) . OTJeBII)];Ho, aTOT IIplI3HaK OTpamaeT pa3HbIe crylIeHII
4>IIJIOreHeTllqeCKoro C03peBaHIIH KJIeTOK.
06pa60TKa MaTepllaJIa IIo]:(aTOM KaJIIIH, IIO Hillarp, H6kfeIt
(1955), )];JIH 1I0JIyqeHIIH peaKl.\lI1I C HOpa,iJ;peHaJIIIHOM B lIaparaH-
r;rIHHX llIeHHo-rpY)];HOH 06JIaCTII IITlIl.\ )];aJIa 01'pIII:(aTeJIbHLle pe3yJIb-
TaTbl (lIpIIHMaK, 1960). Muratori (1962), IICIIOJIb30BaB aTY me
peaKI.\IIIO Ha HOpa)];peHaJlIIH, a TaKme MeTO)]; EO)];lIaHa Ha apreHTO-
il>IIJIHIO aOpTaJIbHO-JIerOqHbIX II )];PyrlIx MeJIKIIX lIaparaHrJIlIeD llIeHHO-
rpY)];Hoii: 06JIaCTII Y KaHapeHRII II )];OMalliHero rOJIy6H (paCIIOJIOmeH-
HbIX Ha BLlIIYKJIOii: CTopOHe )];yrII aopTbl, BRJIIOqeHHhlX B CTBo;r
6JIym]:(aIOm;ero HepBa H D YJIbTIIM06pOHXllaJIbHOe TeJIbl.\e), 1I0JIyqllJI
TaKI-1e me OTpHI:(aTeJIbHble pe3YJIbTaTbl. B 1I0CJIe)];YIOm;eH pa60Te Mura-
tori C coaBTopaMu (Muratori et aI., 1965a, b) npHMeHlIJI )];JIH Tex me
rteJIell: 4>JIyopeCI:(eHTHhIH MeTO]:( H 4>uRCaI.\HIO r.TIYTapaJIb)];erH,iJ;OM
(IIO Tramezzani et aI., 1964) )];JIH BblHBJIeHIIH Hopa]:(peHaJIlIHa B Kapo-
T1I)];HOM lIaparaHrJIlI1I I.\LlIIJIHT II Tome He 1I0JIyqllJI II0311TIIBHbiX pe-
3YJIbTaTOD. B HameH pa60Te, IIpOBe)];eHHOH IIO MeTo]:(y Falck II Owman,
Mbl IIOJIYTJlIJIII OTTJeT.rIHBYIO il>JIyopeCl.\eHI:(IIIO 3eJIeHOBaToro TOHa B 06-
JIaCTII COHHOH apTepHII (KapOTII)];HbIH lIaparaHrJIIIH), D BOCXO)];Hm;en
TJaCTH )];yrII aopTbl II B alIlIRap)];e npe)];Cep)];IIll:, TlIlIHqHYIO )];JIH HOp-
a)];peHaJIIIHa (pIIC. 56, A, B).
B pe3YJIbTaTe IICCJIe,iJ;oBaHIIH, OTRpLlDaIOm;IIX IIPIICYTCTBlIe CIIe-
I:(1Iil>IITJeCRIIX ropMOHOB B llIeHHo-rpY)];HLlX lIaparaHrJIIIHX, IIX MomHO
Ha3BaTb «xpoMail>il>IIHHbIMlI» II, CJIe)];OBaTeJIbHO, CXO)];HbIMII D aTOM
OTHOllleHlIlI C a6)];OMIIHaJIbHLlMII II JII06L1MII )];pyrllMII, 06JIa)];aIOm;IIMII
a)];peHaJIODOH aH)];OKpIlHHOH il>YHKI:(IIeH.
PaClIpe)];eJIeHlle HOpa)];peHaJlllHCO)];epmam;IIX KJleTOK B naparaH-
rJIIIHX a6)];OMIIHaJlbHOH II 3a6pIOllIIIHHOH 06J1aCTeH II3yqeHO HaMII Ha
B3pOCJlLlX Kypax II )];OMaIliHeM rOJly6e. ROHl.\eHTpaI:(IIJI HOpa)];peHa-
JlIIHa B lIaparaHrJlIIOHapHblX KJleTKaX aTIIX yqaCTKOB TYJlOBIIm;a 3Ha-
.TJIITeJlbHO Dblllle, qeM B meHHo-rpY]:(HbIX, II 6e30TKa3Ho BblHBJlHeTCJl
MeTO)];OM 1I0)];lIpOBaHIIH, MeHee qYBCTBIITeJlbHbIM, KaK II3BeCTHO, TJeM
4>nyopeCI:(eHTHbIH. PaClIpOCTpaHeHlIe Hopa]:(peHaJlIIHCOn;epmaIIJ;HX KJle-
145
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Pile. 56. <l.>JIyopeel\I'IIUIiH IIOpa,T{pl'IW;Il1IlcoJWP;-Jmu II"\: HJIl'TOJ;: II IIHparanrJIIlHX
A - B RapoTHgHo!I'I UbIllnellHa. I] - B 06JlaCTH gyrll aOp:rbl 'HIm:! (110 ([>anbHY II OBMI'IIY)
OH. X 3, 00. X 10
TOI>, no HalllIlIl1 II a U.TIIOlJ,CliIlH 111 , 11(' III11PCT ],a"llx-:I1Tfiu 3al;OIlUIl1CP-
HOCTen. Hapngy c agpcIIaJTIIIIrOJ cpll,all nll1n olin pa;!i'Jpoeallhi II 0;.\/1-
1I0'IhY IIJTn ofipa3YIOT II1C.1IaIC lIaparaIlr:IIIOnapUhlC Y3C:II>U no XO;Iy
nOrpaHIl'lHLIX CTBO:IOB II H'.n;JlX COl'YlJ,Jll'TO-HCpllllLIX IIY'lhOH: npn-
MblhalOT I> agBeHTIlI /lU apTcpnwI, I, llepBIIM 1 cTBommaM, /I:III :Ie-
maT 13 coe]:(IIllUTl':Il,IlOTl,aliIlLIX llpoe.TlOiil,ax opranOB, B ORO:IOIIO'J.e'l-
HOII I>JTeT'laTI.e, n CTCT/I,C nO.TIoii BCllM II B 3aupIOllIIIIIHOM IIpOCTpaH-
CTne (pIlC. 57). IlOrTOHIIHO Ollll BCTpeqaIOTCH B l,anCY:le
Ha]:(IIOqe'lHIIIWB. OAlIaI,O llce ;JT/I OKO:lOcocYAnCTLIC II Olw:lOllepBHLle
naparaHrJllII1 npeIJMYlIJ,CCTlleHlIO eocpc]:(oTO'ICllLI B GplOlliHOM IJ 3a6-
pIOllIlIHHOM npOCTpaHCTllax; B l,ay; a.TJI.IIOl\1 II l,pallna:ILIIOI\[ lIanpaB-
JIeHIIHX KO.TIlI'ICCTBO I1X YUhIBacT.
CaMLIM l\pYlIllJ,II\[ llaparaur:mCM UTilI..(, I\al\ II BeCX BLICIIIIIX n03-
BOHO'lHbIX, HII.1HeTCfl l1agIl(l'IC'IIIIII-;. 3TO 1III1,anry.1J1pOllaHHLlii, 'Wll.le
napHblii opraH, paClloilOilWll Ha I>pal1lla.1JIoIlOM KOHQe nO'lel>
(CM. pIlC. 14), B TCeHOI\[ I,OIlTaI>Te C UO.1JOBl,I m meJTe3aMIl. OnlI-
canlle r/lCTOJIO/'II'ICCI\Oro CTPOPllJUI lIa; 1I0'JC'J.Il JIl\OII nTIIQ BCTpC'IaeTCH
elIJ,e B pa60Tax ncplloii 1I0JIOnJIHLI npOIll.1Joro CTOJICTIIH (Nagel, 183G).
BOJTee n03AlIl1e lIer;le]:(OBaJl/lH IlpIlHa :ll'il,aT Hahl II. (1891), \"in-
cent (1897c), Kosc (1907) II B HalliC BpCI\[H Hartman, Kn'Jff,
McNutt, Carver (1947), Hartman, Browllcll (1949).
146
!{aK n:meCTIIO, neT.;WUIIJ],IIOl' paCnO.'lOi!;eIlIlP Ua,;LOI, xpoMal./Jl./HIHIIOll
'I'Ranll cpeJ],n OCTpOBIWB II Tpauer')'JI «({WPI\OBOl'O» BCII.I,eCTBa upH]:(aeT
HaJ],1I0qe1.fHHKaM I1TIIQ TaI\YIO XapaKTl'prr)'lO l\apTHUY KpymeBa, 1{8-
..yro HCJlbBH ECTpeTIIT(' JIll Y o;UlOro II;I IlpPJI,CTaUllTeJIPii. APyrIlx
K'IaCCOB n03BOIIOqHLIX 'i-\;1I130TII(,rx (l'IIl. pill'. 2H, G, 58). PRALl XPO-
r.ra!J>qmHHb1x KJlCTOl\ II pOll1I3aJl "I oun:ILII "1M I,O.TIII'ICCTBOM l,aUH:IJIH-
pOB, HepCAKO cuaBIlI11M III'H, lIe lIMelOII.I,I1MII npucneTa, a T3KIKe cn-
HYCOllA:1MII pa3HbIX pa;u.repon. I\Hiller (1920) OlTnea.'I xpoMa!J>!J>IIu-
r:"
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Pile. 57. rpymlbj XpO H\(I)4)jllllIhIX I,JIl'T/JK l[h'IIJIl'Hlia
A - B aABCHTH HH apTCpnOJIbI (llonaTlIan pcaml'''1 no '(II.II,I] pny II XCHBCJIbTY); B - \J ncpB-
HO I CTBOJIHHC (xpoMadJdJmman peaH Hn no XII"" 'puy II XCI<BCJlbTY). UM. X 5. 06. X 40
147
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Pile. 58. H3l\IIOql''JHIIK KYP1IlI.bl. Ap;pellaJIIIH- 11 1I0pa;J.p(,lla.'lIlHCOp,epilm lIe
I\JIl'TRIl. IIOP;3TH3H pe31;:1\IIH 110 XIIJIJI3pny II XeRneJILTY
OH. x 5, oiL X 40
nbre 3eplla Iial{ B 1I.I1TOIIJla3Me aApeHa!IOBblX 1\.lJ(,TOIi. Tal' II B
IIpOCBeTe KaIIIIJl.TIHpOB, II cqeJl 3TII lIaXO lm IIIOP«PO:lOrIl'ICl'IH1:\1 ;1,0-
Ha3aTe.TIbCTBOM CeHpeI IIII aJ\pCllamlH3 B HpOllh I.
IIopTaJibIIaH CIICTeMa lI.npHYJlHlI.lIII I{POBII "pIICYll\a, 1I11;:t,IIMO, }Jce:\1
BII]:(aM IITIIlI.. Ee BIIepBblC OIIIICaJlII ell\e n npOIllJlOIII CTO:IeTlIII. Hart-
mann II Brownell {1949) OTMeTllml, qTO MopqJOJlOrIl'll'CI>Oe YCTpoii-
CTDO IIOpTaJlbHOn CIICTeMbl IITIIlI. IIpIlHlI.IIIIlla:lbHO He OT.1II'I; eTCH OT
CTpOeUlIfI Mempe6cplloii 1I0pTaJlbllOii CIICTeMbl y penTlI:JIlii.
06 IIHHepllaQl1II HaAIlO'Ie'IIIIfIWB nTIII I13BeCTIlU Cp3B1IIITeJlbIl0
Ma.'lO (Giacomini. 1898b; Miiller, 1929). Be3MHHOTHJ.le CmrrraTlI-
qeCI{Ue BO!IOIma 1l0J],XO;:t,HT I> U3;1,IIO're'llillIiY n rTIlO.1C n. splan-
chnicus minOl', B Mellhllleil: CTcnCIIIl B 11. splilllrhniclIs major,
a TaHmc 113 IIImmIlX rpYJ],HblX CIIlI1II3TlPICClmx raII rJJJICB. 13HYTPII
lIaAIIO'IC'IIIIII\a OlIll o6pa:!YIOT eIIJleTCIIIIH II:! BOJlOlWH II raIIrJlIl0:!-
lIb IX CIIMnaTlI'ICCRIIX I>.TICTOIi, on:ICTalOll.{I1X XpO?tIa«p«pIIIIHble 3J1ellleHTbl
(Bachmann, 1934).
HopaApeHaJlIIHCOAeplIiall\IIe Ime TIm , no 11 a III 11 III Ha6mo)];eHIIHIII, He
, nOp,06I1bIC "a"THllbl ('I'I,,,t'I\III1, OIlIlI"11l1l1ldt' II ;lJ'ynBHI aBTopaMH, B q,lCTlIOI"TlI
A. C. AOreJIeM (1S!I't), Ill' nop,TBt'pmp,11IOTCH cOIIllCMelIIIbIMU 3JIelnpOHIIO}III-
RpOCROIIH'IeCI,IIMIl IICCJIeP;OllaIIIIHMII, lIa OCIIOB31I1I11 IWTOPIdX MOllil\O rOBopllTb
o BId;:{t'JIeHIIII II fipOBeliOCIIOl' PYCJIO CO;tepiKIBlOro rp3I1Y.'1, 110 He ca}IllX rpa-
HYJI.
148
IIMelOT RaROn-JIII6o 3aROHOMepHOH JIORaJIH3a:QHII BHYTPII M03rOBOrO
CJIOJl Ha)];IIOqeqHHROB, a 6eCIIOpH)];OqHO paClIpe)];eJIeHLI Cpe)];H a)];pe-
HaJIIIHCO)];epmaIIJ;IIX (pIIC. 58). O)];HaRO IIX ROJIlP:leCTBeHHble co-
OTHomeHIIH Y pa3HbiX OTPH)];OB IITIII:( IIpe)];CTaBJIHIOT HHTepec. B ce-
PIIII pa60T Ghosh C coaBTopaMII (Ray, Ghosh, 1961; Ghosh,1962;.
Ghosh J., Ghosh A., 1962, 1964; Datta, Ghosh, 1965; Ghosh, Datta..
1969), IIOCBHlIJ;eHHLlX I:(IITOXIIMIItIeCROH xapaRTepIlCTHRe Ha)];IIO'Ie'lJIlI-
ROB IITIII:(, IIORa3aHO COOTHomeHlIe a)];peHaJIllHa II HOpa)];peHaJIIIHa B-
4>HJIOreHeTII'IeCROM II OHToreHeTIIqeCROM pa3BIITHH. C IIOMO.m.blO PII-
COBaJIbHOrO alIlIapaTa B M03rOBOM Be.m.eCTBe Ha)];IIO'Ie'lJIHROB 6L1JIII
O'Iep'IeHLI ROHTYPbl )];BYX RJIeTO'lHLlX rpYIIII: IIO)];aTII03IITIIBHbIX HOp-
a)];peHaJIIIHCO)];epmaIIJ;HX II Tex, ROTopLle TOTaJIbHO ORpamlIBalOTCH
npII B3aHMo]:(eHCTBHII C COJIHMlI xpOMa (a)];peHaJIIIHCO)];epmaIIJ;HX) .
ApeaJIbl Tex II )];pyrHx OlIpe)];eJIeHbl IIJIaHHMeTplIqeCRII (Ghosh A.,
1962; Datta, Ghosh, 1965). Y IITIII:(, CTOHIIJ;IIX Ha pa3HLlX CTY-
lIeHHX 8BOJIIO:QIIOHHOH JIeCTHH:Qbl, COOTHOllieHlIe a)];peHaJIIIH- If Hopa)];"-
peHaJIIIHCO)];epmaIIJ;IIX RJIeTOR 3aROHOMepHO H3MeHHeTCH (pHC. 59).
,[(IIarpaMMa )];eMOHCTpnpyeT IIO)];'beM IIpOl ecca MeTHJIHI:(IIII Hopa)];pena-
JIIIHa B a)];peHaJIIIH, 110 Mepe 8BOJIIOI:(IIOHHOrO pa3BlITHH, II ]:(OCTII-.
meHlIe MaRCIIMYMa B 4>IIJIOreHeTlIqeCRII 60JIee 1I03)];HIIX rpYlIlIax IITlIll
(Passeriformes) 1. ABTOp IIOJIaraeT, qTO HaH]:(eHHaH 3aROHOMepHOCTb
IIplIJIOmHMa K )];pyrHM mIIBOTHbIM, r)];e IIpHMIITIIBHOCTb opraHlI3a:Q1I1t
]:(OJImHa COOTBeTCTBOBaTb CJIa60MY pa3BIITIIIO CIIoco6HOCTII R MeTII-.
JIHpOBaHlI1O ropMOHa. R COmaJIeHIIIO, TaRIIX pa60T HeT Ha )];PyrlIx
1I03BOHO'IHLlX, HO 6L1JIO 6L1 OqeHb IIHTepeCHO IIpOBeCTII IIX, HalIpH-
Mep, Ha pbl6ax. Ho TeXHlIqeCRII OCYlIJ;eCTBJIeHlIe TaRon 3a)];aqII 3a-
TpY)];HeHO RpanHIIM Mop4>OJIOrH'IeCRIIM pa36poCOM a)];peHaJIOBOn TRaHa
Y pLl6 II OTCYTCTBlIeM (<Ha)];IIOqeqHIIRa) RaR rJIaBHOrO opraHa, B RO-.
TOpOM COCpe)];OTOqeHa a)];peHaJIOBaH 4>YHRI:(IIH B opraHlI3Me 8THX mlI-
BOTHbIX. OqeHb B03MomHO, qTO He TOJIbRO BHYTPH OT)];eJIbHbIX RJIaC-
COB II03BOHOqHblX, HO H Mem)];y HIIMH - OT HlI3ll1HX R BbICIlIlIM.
CYlIJ;eCTBylOT aHaJIOrH'IHble COOTHomeHIIH. TaRoe IIpe]:(IIOJIOmeHHe alI-
plIOpHO MomHO 060CHOBaTb BLICOKIIM MoP4>o4>YHRI:(IIOHaJIbHbIM pa3-
BIITHeM CIIMlIaTlIqeCROn HepHHon CIICTeMbI, ROTopaH 06JIa)];aeT TOH-.
ROOTperYJIlIpOBaHHblM MexaHII3MOM )];enCTBIIJI HOpa)];peHaJIIIHa, Hr-
palOlIJ;ero Y BblCIlIHX 1I03BOHO'IHbiX pOJIb Me)];lIaTopa. Ha BbICORon.
CTYlIeHII 8BOJIIOI:(IIOHHOrO pa3BIITIIH HOpa)];peHaJIIIH RaK Me]:(HaTOp 6e-
peT Ha ce6H 4>YHRI:(1I1O xpoMa4>4>HHHoH TRaHII, ROTopaH, TePHH B HeM
Heo6xo)];IIMOCTb (B TaRon CTelIeHH, KaR 8TO CBonCTBeHHO HII3mlIM
II03BOHOqHbIM), MeTIIJIlIpyeT ero B a]:(peHaJIIIH.
Pa60Tbi A. Ghosh (1962), J. Ghosh H A. Ghosh (1962), Datta
II A. Ghosh ( 1965) YRa3blBalOT elIJ;e II Ha TO, qTO Mop4>OJIOrlI-
1 TIpop;nOJIOllieHHe Etkin, BldCRa3aHHoe 11M B AHCRYCCHII no nOBOP;Y pa60TId
Ghosh Ha III MelliP;YHapop;HoM cIIMn03HYMe no Cp8BHHTeJIbHOH BHp;OKpHHOJIO-
rHH B HnoHIIH (1961) 0 TOM, 'lTO Y MeJIRIIX lliIIBOTHIdX, He 'lYBCTBYIO IIX ce6H
3all\ll eHHIdMH OT BHemHHX BparoB (R ROTOpldM OTHOCHTCH II MaJIeHbRHe
neB'llIe nTII'lRII), npeBaJIHpyeT cop;eplli8HHe Bp;peHaJIIIHa, a Y TaRIIX, RaR JIeB,-
Hop8p;peHaJIHH8, He JIlIllieHO OCHOB8HIIH II HYllip;aeTCH B caMOCTOHTeJIbHOM 1I3Y-
'leHHII.
149'
PELECANIFORtfES
.PhoZacrocora.:c ni'ler
CICONIIFORNES
.Bu.buZcus ibis
GALL/FORNES
.6allus domesticus
- loo u Adr.NOr.
50
lOO U
50
lOOf .
50W
eOLUNDIFORNES l; f_
CoZum.ba livia LIJ
CUCUUFORNE8 loof _
EUd!Jnamis scolopaceus 5ot..J.J
PASSER/FORNES lOO l
Passer domesticus (I:Ploceidae) L
Acridotl1eres tristis (I: 8turnidaeJ 50
Holpastes cafer (I: Pucnonotidae)
Corvus splendens (I: Corvidae)
P u c. 59. fllCTorpaMMa OTHOCIITeJIbHblX nponopl\lIH 8p;peH8JII1H8 II Hopap;peHa-
J1JIHa y p;eBHTlI BIlP;OB nTlIl\ (Chosh, 1962)
"IeCRlIe B3alIMOOTHOllieHIIH Mem]:(y ROpOH II M03rOM Ha]:(IIO'leqHIIRa
.(IIJIH Mem)];y cYlIpapeHaJIOBOH H HHTeppeHaJIOBOH TKaHHMII) IIpO-
:HBJIHIOTCH Y pa3Hbix mHBOTHblX 1I0-pa3HoMY H Hym)];aIOTCH B 06'b-
lICHeHHH. B pa60Tax Wright H Chester Jones (1955) II Chester
Jones (1957) MeTIIJIlIpyIOIqlIe CBOHCTBa M03rOBOH TKaHII CBH3L1BaIOT-
oCH C KOpTHRaJIbHOH TOliorpail>lIeii: H BJIIIHHlIeM ROpKOBOH TRaHII Ha
8TOT IIpoI:(ecc. IIo )];aHHbIM Ghosh C coaBTOpaMII (1962, 1965), co-
epmaHHe HOpa)];peHaJIIIHa y IIpe)];CTaBIITeJIeH pa3HbiX OTpH)];OB II BH-
)];OB IITIII:( He 3aBIICIIT OT Mopil>OJIOrH'leCKIIX COOTHomeHIIH II 06'b-
-eMOB KopKOBoro H M03rOBoro BerqecTBa. B CBH3II C 8TIIM KOHI:(eIII:(IIH
Chester Jones 0 3HaqeHIIH CTPYRTYPHLIX KOHTaKTOB ROpLl II M03ra
11 Ha)];IIOqeqHHRaX Ha MeTIIJIHI:(IIIO, 1I0JIyqlIBllIaH B HaCTOHIqee BpeMH
paClIpOCTpaHeHlIe, Hym)];aeTCH, 110 ClIpaBe)];JIIIBoMy MHeHIIIO Ghosh,
B lIepeCMOTpe. Toro me MHeHIIH IIplI)];epmlIBaeTcH Euler (HJ63) .
OTMeqaH, qTO B il>HJIOl'eHe3e 06e TKaHH He BCer)];a BCTpeqaIOTCH
BMeCTe, OH CqlITaeT, qTO 6IIoJIOrlIqeCKOe 3HaqeHHe TeCHLlx KOHTaK-
""TOB xpoMail>il>HHHbIx KJIeTOK C KOpROBOH TRaHbIO Ha]:(IIOqeTJHIIROB He
1I0JIyqlIJIO eIqe y)];OBJIeTBOpIITeJIbHOrO 06'bHCHeHIIH.
IIHTepeCHbl pe3YJIbTaTbl pa60Tbi J. Ghosh H A. Ghosh (1964),
1I0JIyqeHHLle IIPH 1I3yqeHlI1I OHToreHe3a na Ha,JJ;IIOqeqHHRaX MOJIO-
LlX (OT 2 )];0 10 )];Hen: IIOCJIe BblJIyn JIenlIH) II B3pOCJIbiX IITHI:(.
"Y Tex II )];pyrlIX 06'beM I:(lITOXIIMlItIeCKII BLlHBJIHeMLlX KaTeXOJIOBLlX
ropMOHoB O)];IIHaROD, B OTJIllqHe OT MJIeROlIlITaIOIqIIX mlIBOTHLlX II
°TJeJIOBeKa, y KOTOpblX 6HOXlIMHqeCKHMH MeTO)];aMII (Shepherd, West,
1953; West et aI., 1953) IIOKa3aHO YMeHbmeHlIe HopaApeHaJIIIHa II
llOBblmeHlIe a)];peHaJIIIHa IIpOIIOpl.\HOHaJIbHO D03paCTY. B HamCH pa6o-
Te Ha KyplIHLlX 3apO)];bllllaX H I:(bIIIJIHTaX (CMIITTeH II MaJIHHOB-
-CKaH, 1960) Mbl OTMeTH.TIII Te me oc06eHHocTII, OTJIIITJaIOrqlIe n;blII-
.JIHT OT KpLlC, MblllleH II RpommOB (CMIITTeH, 1958). }lme B 1I03)];HlIe
-CpOKH 8MoplIoreHe3a Y I:(bIIIJIHT Mbl He MOrJIH OTMeTHTb IIpeHMy-
lI:(eCTBeHHoro HaJIH'lIIH RaROrO-JIII6o O)];HOro copTa KJIeTOK. RpOMe
150
Toro, ]:(JIJI pacnpe]:(eJIeHIIJI IIX B «ceTl\e» M03rOBOrO Be eCTBa Han
1I0qeqHIIKOB xapaKTepHO He OqarOBOe, a paBHOMepHoe paCIIOJIome-
nue II, CJIe]:(OBaTeJIbHO, paBHoqeHHoe B OTHomeHUU TOlIorpacIJllu K
KOPKOBOMY CJIOIO. 3peJIOpom]:(aIOIIJ;UeCJI nTeHqbl 06JIa]:(aIOT, BII]:(UMO
TaKIIM 3anaCOM KaTeXOJIOBblX ropMoHoB, KOTOpbIe 06eCneqlIBaIOT
clJYHRqlIIO B3pOCJIbIX. MeTUJIIIPYIOIqaJl CUCTeMa Y HIIX K 3TOMY BpeMe-
HII ]:(OCTuraeT, CJIe]:(OBaTeJIbHO, yme BblCOKOll cTyneHu pa3BIITIIH.
He3aBucIIM0 OT Toro, KaK B ]:(aJIbHenmIlx UCCJIe]:(OBaHUJlX 6Y;J;YT-
pemeHbI Bonpocbl 0 3H81JeHUII B3auMo ellCTBIIJI (<KOPbl» II «M03ra».
H8]:(nOqeqHUKa U l:aKOBO 3HaqeHUe KOJIUqeCTBeHHblX nOKa3RTeJIen B.
co]:(epmaHuu Toro nJIU ]:(pyroro ropMoHa B Ha]:(IIOqeqBHKaX n03BO-
HOqHbIX, BamHO, 1JTO OHU UMeIOT KaKon-To 60JIbmOn clJYHKqlloHaJIb-
Hbln CMblCJI, oTpamaIOT UH]:(IIBu]:(yaJIbHbIe lIoTpe6HocTII pa3Hblx BII]:(OB.
II no]:(qepKUBaIOT 3HaqeHIIe Ha]:(nOqeqHOll meJIe3b1 KaK caMOCTOJlTeJIb-
Horo opraHa. no OTHomeHUIO K nTuqaM BHTepeCHbIe CBe]:(eHUH 06
aKTIIBHon pOJIU Ha]:(nOqeqHUKOB npu nepeJleTaX ]:(aHbI B pa60Te
Naik II George (1964). )J;aHHble 3TUX aBTopOB, KOTOpbIe IICCJIe-
]:(OBaJIU MurpupYIOIqUX CKBOpqOB B pa3Hble BpeMeHa ro]:(a - OT OKTH6-
pJl ]:(0 anpeJIJI, nOKa3blBaIOT, qTO Henocpe]:(CTBeHHO nepe]:( nepeJIe-
TOM B qUTOlIJIa3Me KJIeTOK M03rOBoro CJIOJl Ha]:(nOqeqHUKOB Ha6JIIO-
]:(aeTCJI YCUJIeHHan IIpO]:(yKqUH a]:(peHaJIIIHa II Hopa]:(peHaJIIIHa,
B03paCTaeT aKTUBHOCTb KIICJIOH clJoccJlaTa3bl, YBeJIUqUBaeTCH KOJIU-
qeCTBO MUTOXOH]:(pun II co]:(epmaHue PHI\.
B IITOre aHaJIU3a MopcJIOJIOrUII a]:(peHaJIOBOn CUCTeMbl nTuq Mom-
HO OTMeTUTb CJIe]:(YIOIque oc06eHHocTu, xapaKTepu3yIOIqlIe 3Tan ea.
clJIIJIOreHeTlf'IeCKOrO pa3BUTUJI.
B pH]:(Y 1I03BOIIOqHblX Ha]:(nOqeqHUKII nTuq ]:(OCTuraIOT BbicOKon.
MopclJOJIOrUqeCKOn II clJYHKqlIoHaJIbHOn CJIOmHOCTII II HBJIHIOTCJI cne-
quaJIII3UpOBaHHbiMII 3H]:(OKpUHHbIMJI opraHaMU.
3KcTpaa]:(peHaJIOBbie opraHbl B cocy]:(ax ma6epHoro npOUCXOlli]:(e-
HUJI OTJIBqaIOTCJI OT Ha]:(nOqeqHIIKOB ]:(OJIbqaTbiM CTpoeHueM I! pe3Ko.
CHlImeHHblM co]:(epmaHueM ropMoHoB. Pa3pacTaHlle coe]:(uHIITeJIbHOn
TKaHU, npoHU3bIBaIOIqen naparaHrJIUOHapHble Y3eJIKII, ]:(eJIUT ux Ha.
]:(OJIbKU II CJIymuT O]:(HUM U3 BblpameHlln pY]:(UMeHTaquII. )J;pyruM
npU3HaKOM, xapaKTeplI3YIOIqlIM yracaHue clJYHKqlIu, HBJIJleTCJI na]:(e-
Hue IIpO]:(YKTUBHOCTII cneqlIclJU'leCRUX RJIeTOK lIaparaHrJIlIeB.
Y nTuq, HapHJJ:Y C PY]:(UMeHTapHblMII Y3eJIKaMU 3KCTpaa]:(peHa-
JIOBOll CUCTeMbI, OTpamaIOIqIIMU perpeCCUBHoe HBJIeHlIe B IBOJIIO-
qIDI, 1I0JIY'IaIOT mlIpOKoe pacnpOCTpaHeHlIe MeJIKUe a]:(peHaJIOBble-
OCTpOBKII II OT]:(eJIbHble KJIeTKII, COIIYTCTBYIOrn;ne IIOrpaHII'lHblM CUM--
naTlI'leCKIIM CTBOJIaM II Bcen neplIclJepll'leCKOH HepBHon CIICTeMe._
OHII paCCMaTpllBaIOTCH HaMIl KaK nporpeCCIIBHbIH nplI3HaK B pa3BU'CUB
a]:(peHaJIOBOn CIICTeMbl, OOYCJIOBJIeHHblii CJIe yIOIqIIM 3TalIOM IBOJIIO-
quu HepBHOll CIICTeMbI, KOTOpaJl ]:(OCTuraeT mupOKoro apeaJIa Y nTlIq_
r.aa8a VI
1.JIEKOnIITAIO IIE
B IIpe]:(bl]:(YIIJ;I1X rJIaBaX Mbl CTpeMIIJIIICb IIOKa3aTb, qTO CTPYKTYP-
Hble KOHTaKTbI naparaHrmleB C cocY]:(I1cToii II HepBHOii CIICTeMaMII B
PHJJ:Y II03BOHOqHblX mIlBOTHblX B n;eJIOM oTpamaIOT CPYHKn;I10HaJIbHOe
Ha3HaqeHIIe xpoMacpcplIHHOH CIICTeMbJ Ha pa3Hblx ypOBHHX aBOJIIO-
.D;I10HHOrO pa3BIITIIJI lliIIBOTHbIX. C]:(eJIaHO aTO 6b1JIO C Toii l.{eJIbIO,
'ITo6b1 nO]:(OHTII K OCHOBHOH 3a]:(aqe Hamero IICCJIe OBaHHJI, KOTopaH
3aKJIIOqaeTCJI B On;eHKe CPYHKn;I10HaJIbHOH pOJIII Bcex xpOMacpcpIlH-
HbiX opraHoB MJIeKOIIIITaIOlI.{lIx II paccMoTpeHlIIl IIX C TOTIKII 3peHIIH
npIlHa]:(JIemHOCTII K e]:(I1HOH aH]:(OKpIIHHOH a peHaJIOBOii CIICTeMe.
B HaCTOJlID;eii II lIoCJIe]:(yIOID;I1x rJIaBaX Mbi IIOlIbiTaeMCJI oCYID;ecT-
BIITb aTY 3a]:(aQY.
Kor]:(a 3axo]:(IIT peqb 0 naparaHrJIIIOHapHOH CIICTeMe MJIeKOnIl-
TaIOID;IIX, TO caMoii pacnpOCTpaHeHHOH ee xapaKTepIlCTIIKOH CJIymIlT
..cJIe]:(yIOID;ee YTBepm]:(eHIIe: 60JIbmIlHCTBO lIaparaHrJIlIeB perpecclIpyeT
cpaay nOCJIe pom]:(eHIIH, coxpaHHeTCJI TOJIbKO M03roBOH CJIoii Ha]:(-
JIOqeqHIIKOB II qaCTIIqHO napaaOpTaJIbHblH. Ha]:(o nOJIaraTb, 'ITO Ta-
ROe cym]:(eHlIe MOrJIO B03HIIKHYTb rJIaBHblM 06pa30M B pe3YJIbTaTe
60JIbmoro 3HaqeHIIJI Ha]:(nOqeqHblX meJIe3, KOTopble ]:(eiiCTBIITeJIbHO
OCTlIraIOT Y B3pOCJIbiX oco6eii MJIeKOIIIITaIOID;IIX BbiCOKOii CTPYKTYP-
Hoii II CPYHKn;IIOHaJIbHOii CJIOmHOCTII II IIrpaIOT Y HIIX rJIaBeHCTBYIO-
ID;YIO pOJIb cpe]:(1I ]:(PyrIlx aJIeMeHTOB aTOH CIICTeMbi. MeJIKlIe me
naparaHrJIIIII, BO MHomeCTBe onIlcaHHble B pa3HblX opraHax pa3-
.JIIIqHbIMII aBTopaMII B pa3Hble rO]:(bl, OqeBII]:(HO, He nOJIb3YIOTCJI mIl-
pOKoii II3BeCTHocTbIO, a Te 113 HIIX, KOTopble pacnOJIOmeHbI B Ka-
pOTII]:(HOii II npIlcep]:(eqHOH 06JIaCTHX II 3aCJIymIlJIII lIonYJIJlpHOCTb,
no 06ID;eMY np1I3HaHIIIO, 6e3Ha]:(emHO nOTepJIJIII CBJl3b C aH]:(OKpIlH-
Hoii CPYHKn;I1eii II npIlOOpeJIII He CBOHCTBeHHYIO 11M pen;enTopHyIO.
HapR]:(Y C aTIIM HeT HIIRaKoro COMHeHIIJI B TOM, qTO YTBepm]:(eHlIe
06 «06paTHOM» pa3BIITIIII naparaHrJIlIeB nOCJIe pom]:(eHIIR He JIII-
meHO OCHOBaHIIJI: B aM6plIoreHe3e MJIeKOnIlTaIOID;I1X OHII BCTpeqaIOT-
oCR B 60JIbmeM KO.TIIIqeCTBe II 60JIbmlIX pa3MepOB, qeM Y B3pOCJIbiX
oco6eii. nOaTOMy- perpecc naparaHrJIIIeB II 3HaqeHIIe MeJIKIIX napa-
raHrJIIIOHapHblX OCTpOBROB Y B3pOCJIbiX IIOJIOB03peJIbiX mlIBOTHbiX
IIO]:(JIemaT pa360py B HaCTORII.{eH II CJIe]:(YIOID;eii rJIaBax. Perpecc na-
paraHrJIlIeB Mbl paCCMaTplIBaeM B KaqeCTBe IIplI3HaKa peKaIIIITYJIJI-
n;IIII, IIOBTOpJlIOID;erO TO, qTO 6bIJIO cBoiicTBeHHo xpOMacpcpIlHHoii CII-
<\TeMe HII3mlIX n03BOHOqHbiX (KPyrJIOpOTble, pbI6hl), a 3HaqeHIIe MeJI-
.152
KHX IIaparaHrJIHeB C TOqKH 3peHHJI HX npHHa]:(JIemHOCTH K BKCTpaa]:(-
peHaJIOBOn maHII, OCTaBmefICJI B HaCJIep;CTBO OT HII3R00praHI<I30BaH-
HblX mlIBOTHbiX II cocpep;OT01JeHIlOn B6JIll31I IIJIII BHYTPII ap;BeHTlIqllll
HpynHblx apTepHH, JIll60 pa3BllBmeHCJI BHOBb Ha 60JIee II03]:(HeH CTY-
neHII BBOJIIOqIlOHHOH JIeCTHIII.J;bI - BMeCTe c pa3BIITIIeM CllMnaTIIqe-
CROll HepBHOH c,I1CTeMbI - II OKa3aBmeHCH pa36pocaHHOR BMeCTe C
HeH 110 Bc,eMY TYJIOBIII:u;Y.
ECJIII 06paTIITbCJI K JIIITepaTypHbIM IICT01JHIIKaM, TO JIerKO y6e--
]:(IlTbCJI, qTO cpe]:(11 onIlcaHIIH xpOMacpcpIlHHOH CIICTeMbi y II03BOHOq-
HblX mIlBOTHbiX MJIeKOIIIITaIOIII;IIM nOCBJlIII;eHO HalI60JIbmee 1JIICJIO IIC--
CJIep;OBaHIIH. nOMHMO Hap;IIOqeqHbiX meJIe3 y HIIX onIlcaHO orpOMHoe'
qlICJIO naparaHrJIIIeB. B pa60Tax, OTHOCHIII;I1XCJI K BTOH TeMe, lIa-
paraHrJIlI1I HaHp;eHbI B caMblX pa3HLIX MeCTax TeJIa B3pOCJIblX mH-
BOTHblX B BII]:(e MeJIKIIX rHe3]:( B napeHXllMe opraHOB, B HX Kan-
cYJIax llJIII B oKpymaIOIII;en coe]:(IlHIITeJIbHOn TKaHII, 1JaCTO PMOM C
raHfJIll03HblMII KJIeTKaMII IIJIII HepBHblMII CTBOJIIIKaMII: B CJIIOHHOH
meJIe3e (IfBaHoB, 1930); B napeHXllMe nOqeK, nOqeqHblX JIOXaH-
Kax (IfBaHoB, 1930; Watzka, 1943); B IIpe]:(cTaTeJIbHOH meJIese
(Bucura, 1907; Ciardi - Dupre, 1936); B HllqHIIKaX (Bucura, 1907;
Winiwarter, 1910; IIBaHoB, 1930; Watzka, 1943); B meiiKe.
MaTKll; B TOJICTOM KllmeqHllKe (I1BaHoB, 1930, 1945); B JIenUlX
(l\ynpllJlHOB, 1959); B Tpaxe06pOHXllaJIbHOH 06JIaCTll - Ha nepep;-
HeH CTeHKe Tpaxell B MeCTe ee 611cpypl<aI.J;I111 II B HaqaJIbHblX
OT]:(eJIaX 6POHXOB (Muratori, 1964, 1965); B ropTaHll (Watzka, 1966;.
Schonberger, 1967); Mem]:(y JlpeMHOH BeHOH II BeTBHMll Jl3bIKO-fJIOT01J-
HOro HepBa - B6JIll3II canaliculus timpanicus BllCOqHOH KOCTII (Zett.er-
gren, Lindstrom, 1951); B fJIa3HOH Op6I1Te (Botar, Pribek, 1935); B
CllMnaTllqeCKllX Y3JIaX nOrpaHllqHOrO CTBOJIa II B ]:(pyrlIx neplIcpepllqe-
CKllX CnJIeTeHIIJlX II raHfJIllHX (3aJIKIIH]:(, 1937; npllHMaK, 1959;
RaplIeHKo, 1964; de Ribaupierre et at, 1966; Siegrist et at, 1968);
B ]:(yre aopTbi (Busacchi, 1912-1913), B TOM me MecTe, r]:(e II03me
Penitschka (1931) OlIllCaJI Hap;cep]:(eqHbIH naparaHfJIllH, Palme
(1934) - paraganglion aorticum supracardiale II rp;e Muratori
(1935) II Truex (1950) HalliJIll OCTpOBKll; B oKpymaIOIII;eii aopTY
coe]:(IIHIlTeJIbHOH TKaHll, B npllJIemaIII;IlX cocy]:(ax II HepBHblX CTBo-
JIax 6JIymp;aIOrn;ero II ]:(enpeccopHoro HepBOB (Muratori, 1933b, c,
1935; Nonidez, 1935a, b; Celestino da Costa, 1935a, b); B YCTbe
BepxHen nOJIon BeHbI - Memp;y oKpymaIOIII;eH ee coe]:(IIHIlTeJIbHOU-
TKaHbIO II a]:(BeHTllqlleH, 60raToH B 3TOM MeCTe BOJlOKHaMll MllOKap]:(a
(Muratori, 1964); B BIIllKap]:(I1aJIbHOH mIlpoBoH KJIeTqaTKe, B6JIII3II
JIeBOH II npaBoH BeHeqHblX apTepllH (Wirtz, 1968); B_ cep ,aeqHblJL
ymKax (Pannier, 1935); B cTeHKax_npe]:(cepp;IlH (True , I95'O) ,
B HapymHoH 060JIOqKe JIerOqHOH apTepllii (rpllropbeBa, 1954; Ryn-
pllHHOB, 1959); Memp;y JIerOqHOH apTeplleH II aopTOH (Seto, 1935);-.
B 1I0]:(KJIIOqllqHOH (Hollinshead, 1940); Mem]:(y COHHOH II npaBon'
1I0]:(KJIIOqHqHOH II BORpyl' 60TaJIJIOBOH CBH3KII (Goormaghntigh, Pan-
nier, 1939); B BeTBU JIerOqHOH apTepllU, nllTaIOIII;eH aOpTa.:IbHO-JIe-
rOqHbIH naparaHfJIIIH (Hughes, 1967); 110 xo,ay 06rn;eH COHHon'
f53:
llpTepml (White, 1935; IIplIiiMal\, 1959) II 6eCqlICJIeHHOe qlICJIO pa60T
C OIIlIcaHlIeM (<I\apOTII]:(Horo TeJIa» B 6IIclJYPl\aqIIII COHHOll apTeplIlI.
Ha pnc. 119 ]:(aHa cxeMa, I1JIJIIOCTplIpYIOIqaJl IIplIMepHoe paCIIpe]:(e-
JIeHlIe naparaHrJIIIOHapHblX Y3eJII\OB Y MJIeI\OnIITaIOIqIIX.
Cpe 1I lIepeqlICJIeHHblX pa60T Mbl cneqlIaJIbHO ynOMJlHYJIII CTapble
ya60TbI, OTHOCJlIqlIeCH 1\ HaqaJIY Tel\YIqero CTOJIeTIIJI (uaIIpIIMep,
Wiesel), II CaMbIe cOBpeMeHHbIe, IIpoBe]:(eHHble HOBefunHMII MeTo]:(a-
_Mil :meI\TpOnHoii: MlIl\pOCI\OlIlI1I (Siegrist C coaBTopaMlI) IIJIII rIICTO-
XIIMIIII (Muratori). l\IbI Tal\me C03HaTeJIbHO 06'be]:(IIHIIJIII B :nOl\f
cnlICl\e lIaparaHrJIIIII, 1\0TOpble, 110 I\JIaCcIIclJnl\aqlI1I pa3HblX aBTopoB,
OTHeceHbI 1\ pa3HbiM rpYIIII3M: ]:(aIOIlUIX n He aIOIqIIX 1I03IITIIBHYIO
.x.pOMaclJclJIIHHYIO peal\qlIIO B RJIaCClIqeCI\Oll ee MO lIclJlIl\aqlIlI, IIo-pa3-
HOMY IIHHepBlIpOBaHHbIe, pa3BIIBaIOIqneCJI II3 pa3HblX IICT01JHIII\OB II
JlI\06bI He 0]:(H03HaqHble 110 clJYHl\qIlIl. HalIOMHIIM OCHOBY, Ha 1\0-
. TOpOii: c03]:(aJIIICb Bce 3TII npe]:(CTaBJIeHIIJI: OTCYTCTBIIe rIICTOXIIMII-
1JeCI\II 06HapymlIMbix rOpl\lOHOB B 3TIfX opraHax; paCIIOJIOmeHlIe IIX
B6JIII3Il I\pynHblx COCY]:(OB Tal\Oll 06JIaCTII, r e perIlcTpaqlIH reMO-
_ IIHaMlIqeCl\lIx II XIIMlIqeCI\IIX c BlIrOB I\pOBII OqeHb BamHa JIJI aHa-
JIII3a YCJIOBlIll M03roBoro I\poBo06paIqeHIIJI (]:(yra aopTbi II 06JIaCTb
. COHHbIX apTeplIll); 06IIJIbHaJl qYBCTBlITeJIbHaJl IIHHepBaqIIH, OTBeqaIO-
IqaJl Tal\lIM Tpe60BaHIlHM reMO]:(IIHaMIII\II, II, Hal\OHeq, pa3BIITlte
II3 lIapaCIIMnaTlIqeCI\IIX nCTOqHlIl\OB - B IIpOTIfBOnOJIOmHOCTb BceM
]:(PyrlIM lIaparaHrJIIIHM, rlICToreHe3 1\0TOpblX CBJl3aH C CIIMnaTlIqe-
CI\Oll HepBHOll CIICTeMOll.
He IICqepnblBaIOIqaH Bcex ]:(eTaJIeii:, HO ]:(OBOJIbHO nOJIHaJl CBO]:(l\a
cBe]:(eHlIll II IIpII3HaI\OB, 110 1\0TOpblM 6b1JIa ]:(aHa I\JIacclIqml\aqIIJI
IIaparaHrJIlIeB, BOmJIa B PYI\OBO]:(CTBa (Watzka, 1943), r]:(e Bce na-
paraHrJIlI1I ]:(eJIJlTCJI Ha «, xpoMaclJclJIIHHble» IIJIII CIIMlIaTlIqeCl\lIe, If
(meXpoMaclJclJIIHHbIe», IIJIII IIapaClIMnaTII1JeCl\lIe, KOTOpbIe pa3BIIBaIOT-
CH II3 IX II X nap M03rOBblX HepBoB. K qlICJIy nepBblx OTHOCJlTCH,
RpoMe M03rOBoro CJIOJl Ha]:(noqeqHIII\OB, «cB060]:(Hble» naparaHrJIlI1I
(rpynlIbi I\JIeTOK B opraHax, B nOrpaHlIqHOM CIIMlIaTlIqeCI\OM CTBOJIe,
IIO XO]:(Y CIIMnaTllqeCRIIX HepBOB). K qlIC-JIY BTOpblX - l\apOTII]:(Hblll
naparaHrJIlIll, Ha]:(cep]:(eqHblll II MeJIKlIe rpYIIIIbI, pacceHHHble B CTIIO-
JIax 6JIym aIOIqerO II H3bIR0-rJIOTOqHOrO HepBOB.
OTcy1'CTBlIe IIJIII HelIOCTOJlHCTBO xpoMaclJclJlIHHoll peal\qlIll B pH]:(e
.lIaparaHrJIlIeB 6b1JIII O]:(HIIM II3 IIOBO OB, Kal\ yme rOBoplIJIOCb Bblme,
K C03]:(aHIIIO II ]:(Pyroll 1\0HqelIqlIlI, He CBJl3aHHOll C pa3BIITIIeM 11
npOIfcxom]:(eHlIeM naparaHrJIlIeB. 3a (<HeXpoMaclJclJIIHHblMII» CTPYI\TY-
paMII 6b1JIa nplI3HaHa clJYHl\qlIJI BOCIIplIJlTllJl pa3 pameHlIii, nOCTY-
lIaIOIqIIX co CTOpOHbI I\pOBII. IIMeuHo Ha MJIel\OIIIITaIOIqIIX 6b1JIII BlIep-
Bble npOBe]:(eHbI MopclJOJIOrlI1JeCl\lIe IICCJIe]:(OBaHIIJI, Ha OCHOBaHlI1I KO-
TOpblX Driiner ( 1925) II de Castro ( 1928) BblCKa3aJID BTY
mnOTe3Y. OTcYTCTBlIe cneqlIclJlI1JeCKOll oKpacKII XpOl.laclJclJIIHHblX KJIe-
TOK II 06IIJIbHaH, 31\CneplIMeHTaJIbHO OKa3aHHaJl, 1JYBCTBIITeJIbUaJl
HHHepBaqlIJI aJIII IIOBO de Castro IICKJII01JIITL BTII I\JIeTKII II3
CIICTeMbi IIaparaHrJIIIOHapHblX, C1JIITaTb IIX peI(elITopHblMII II AaTb
TeM caMbiM MopclJOJIOrlIqecl\oe 060cHoBaHIIe clJII31I0JIOrlI1JeCKHM aH-
:154
HblM Hering (1924, 1927) 0 pecIJJIeKcoreHHoH pOJUI KapoTII]:(HOR
30HbI. TI03me, no TeM me nplI3HaKaM K KapOTII]:(HbiM «rJIOMycaM»
6b1JIII nplIcoe]:(IIHeHbI OKOJIOCep]:(eqHble - Kapp;lIo-aOpTaJIbHble (Penit-
schka, 1931; Muratori, 1935, 1937; Nonidez, 1935a, b, 1936.
1937 a, b), HaJIlIqlIe KOTOpblX xopomo rapMoHlIpOnaJIO C «p1I31I0JIOrlI-
qeCKIIMII ]:(aHHblMII 0 pe«pJIeKCOreHHblx 30Hax B 06JIaCTII p;y;rII aop-
Tbl (Heymans et a!., 1933; Heymans, 1934).
TIpe]:(cTaBJIeHIIJI 0 CTpYKTYpax, OTBeTCTBeHHblX 3a peqeIlTOpHYIO
«PYHKqIIIO Kap]:(IIO-aOpTaJIbHblX II KapOTIIp;HbIX 30H, KOTopbIe 06pa-
30BaJIIICb B pe3YJIbTaTe Mop«p0JIOrlIqeCKIIX IICCJIep;OBaHlIii 20-30-x ro-
P;OB TeKyrn;ero CTOJIeTIIJI, npO]:(OJImaIOT, BonpeKII HOBbIM «paKTaM, roc-
nO]:(CTBOBaTb B HayqHOH JIIITepaTYpe. Memp;y TeM HOBbie ]:(aHHbIe He-
YKJIa]:(blBaIOTCJI B rrpOKpYCTOBO JIome STOii KOHqenqlIlI, KOTOpR.JI YCTa-
peJIa II ]:(aBHO Tpe6yeT nepeCMOTpa. HeCMOTpJl Ha TaKoe DOJIOme-
HlIe, B no]:(aBJIHIOrn;eM 60JIbllllIHCTBe SRcneplIMeHTaJIbHblX pa60T aB-
TOpbl, nOJIyqaH SJIeKTpOHHOMIIKpOCKOnUqeCKlIe II rlICTOXUMuqeCKlIe
cBe]:(eHuH, C HeCOMHeHHOCTbIO CBII]:(eTeJIbCTByIOrn;lIe 06 SH]:(ORpIIHHOD
nplIpo]:(e naparaHrJIlIeB. He nblTaIOTCH nepeCMOTpeTb Mop«p0JIOrlIqe-
CKlIe OCHOBbI cTapoii TeoplllI, a P;IICKYTIIPYIOT 110 BOIIPOCY 0 TOM,
KaKUM 06pa30M STII HOBbIe «paKTbI IIplIJIOmlITb K cyrn;eC'fBYIOrn;eii
TOqKe 3peHlIH, T. e. Kan: paCIlIH«PPOBaTb IIX 3HaqeHIIe B pe«pJIeKCO-
reHHblX peaKqlIJIX YKa3aHHblX 30H.
MbI 06'be]:(IIHHeM Bce naparaHrJIlI1I MJIeKOnIITaIOrn;IIX, paHee pa3-
6HTbIe Ha OCHOBaHIIII IIX CTPYKTypbI, «PYHKqIUI II npOlIcxom]:(eHIIH Ha
caMOCTOJlTeJIbHble rpynnbl II OTHOCIIMbIe B pJl]:(e CJIY'IaeB K pa3HbiM
CIICTeMaM, B e]:(IIHYIO SHJJ;OKPIIHHYIO. 06'beJJ;IIHeHlIe IIX 060CHOBaHO
TpeMH 06CTOJlTeJIbCTBaMII. TIepBoe - STO MHOrOqlICJIeHHble II pa3HO-
CTopOHHlIe «paKTbl, nOJIyqeHHble K HaCTOHrn;eMY BpeMeHII nYTeM qlI-
TOJIOrlIqeCKOro H I'HCTOXHMlIqeCKOro aHaJIH30B, KOTopble n03BOJIHIOT
C 60JIbmoii y6e]:(HTeJIbHOCTbIO KOHCTaTHpOBaTb e]:(HHCTBO STOH CIICTe-
Mbl II OTHeCTII ee K a]:(peHaJIOBbIM CTpyKTypaM SHP;OKpIIHHoro XapaI{-
Tepa. BTopoe - BTO cpaBHlITeJIbHO-MOp«pOJIOI'HqeCKlIe CBe]:(eHHJI, 110-
JIyqeHHble HaMH Ha c06cTBeHHoM MaTeplIaJIe II II3 aHaJIII3a JIIITe-
paTypHblX IICTO'lHIIKOB, cBII]:(eTeJIbCTBYIOrn;He 0 TOM, qTO IIaparaHrJIlIlI,
pacnOJIOmeHHbIe y MJIeKOnHTaIOrn;HX B a]:(BeHTHqlIlI COCy]:(OB meiiHO-
rpY]:(HOH 06JIaCTII (110 XO]:(y COHHblX, JIerOqHblX apTepHH II ]:(yrII
aopTbI, B snlIKap]:(e IIpe]:(cep]:(uii II B cep]:(eqHblX ymKax), HBJIHIOTCJI
He qeM HHbIM, RaK MeJIKHMII OCTpOBKaMH, OCTanmlIMIICJI OT HeKor]:(a
pacnpOCTpaHeHHblX IIJIaCTOB xpOMa«p«PIIHHOH TKaHII y KpyrJIOpOTbIX
II pbI6. TpeTbe - 1I3yqeHIIe sM6plIoreHe3a, pe3YJIbTaTbi KOTOpOro 110-
l\a3blBaIOT KaK e]:(HHI.1H rJIlIaJIbHblii IICTOqHIlK pa3BIITIIH KJIeTOK M03-
rOB oro CJIOJl Ha]:(nOqe'lHIIKa II KapOTU]:(Horo TeJIa - TlInlIqHOrO npep;-
CTaBIITeJIH (<HeXpOMa«p«PHHHbIX» naparaHrJIHeB, TaK II npH3IIaRII pe-
RannTYJIHqHII B pacnpe]:(eJIeHHII lIaparaHrJIHeB.
B npe]:(b yrn;HX rJIaBaX OblJIO nOKa33Ho, qTO MHOriOJIeTHlie CIIOpbI
o HenOCTOJIHCTBe IIJIII OTCYTCTBHII xpOMa«p«PHHHOH pea HII B pHp;e
naparaHrJIlIeB II, CJIep;OBRTeJIbHO, OTCYTCTBUe 'B 1:IIIX IT'OpMOHOB B H3-
CTORrn;ee BpeMJI nOTepHJIII 3HaqeHHe. BHe]:(peHlIe III 61I0JIorlIIO HOBblX
155
--
A
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2
5
l' B C. 60. XpOMaclJclJlIHHaH peaKl\IIH (A) B KJIeTKaX KapOTHp;HOrO naparaHrJIIIH
"1\pOJIHKa; lIop;aTHaH peaKl\HH (B) B KJIeTKaX KapOTHp;Horo naparanrJIHH KOillKII
{no XIlJIJIapny II XeKBeJIbTY)
J - JJ;OJIbI<B C JlB'l'eHCHBHOil: peaI<qHet1:; 2 - JJ;OJIbI<B co Mallott HJIH OTCyTCTBYIOIIIet1: peaI<nuett
.(lIpHilMaI<, 1959J. PUCaB. 8.11II8paT, On. X 10, oil. X 90
bleTO OB IICCJIe OBaHIIJI n03BOJIIIJIO BLlJlBJIHTb ;n RJIemaX :3TIIX OpraHOB
MlIHlIMaJIbHOO CO epmaHlle rOpMOHOB (KO.1luqeCTBa .KOTOpLiX 6bIJIII,
BII]:(IIMO, HIDKe 1I0pora qYBCTBlITeJIbROCTH XpoMaclJclJHHHoii peaKD;III1)
.11 HaXO IITIt IIX ame Y TeX mlfBOl'HLlX, Y KOTOpLiX HIII\TO IIX IIpem
fte He OIIIICLlBaJI. B HeKOTopblX naparaHrmfJIX 06HapymeHa IIpOA)TR-
D;HJ1 MOHoaMIIHOB, cBoiicTBeHHaJl £CeM TeM opraHaM - B nepBYIO oqe-
pe b MoarOBOMY C,JIOIO Ha]:(IIOqe'IHIIKOB, KOTopLie BceMII nplI3HaIOTCH
:3I1)];OKpUHHbIMII.
B 1959 r. no HameMY npe]:(JIOmeHIlIO llpniiMaR npOBeJIa pa60TY
_110 HsyqeHIIIO paClIpe eJIeHIIH a peHaJIIIBa II Hopa peHaJlHHa B Ha-
pOTII HOM opraHe MJleROIIHT8IOm;HX B p8sHLIe lIeplIO LI nOCTHaTaJIb-
Horo OHToreHe3a (KpOJIIIK, Mblmb, Kphlca, MOpciKaH CBHHKa, KOIllKa,
c06oJIb) C uplIMeneHlleM XpoM8clJclJUJ:IHOii peaKD;1I11 B MO]:(IIclJIIKaD;IIH
Hillarp H Hokfelt (1953, 1955) II 1I0 aTHoro MeTo]:(a, IIpe JIO-
meHHoro TeMII me aBTopaMII JIH 06H8pymeHIIJI Hopa peHaJIIIHa. Xa-
paKTep II03HTlIBHOH peaKD;1I1I MeTOK KapOTII1{HO'l'O naparaHrJIIIJI Ha
8 penaml.H II Hopa]:(peHaJlIIH, nOJIY1JeHHblH llpllHMaR (pIlC. 60, A, B),
8HaJIOrH1JeH peaRD;IIII B M03rOBOM CJIoe H8 n01Je1JHHROB (CMHTTeH.
1958), BCJle S8 ony6JIIIKOnaHIIeM pa60TLI llplIHMaR (1959), Mu-
156
scholl et a1. (1960) IIOKa3aJIU npUCYTCTBue Hopa,npeHamlHa Ii 3KCT-
paKTax II3 KapOTH HbIX TeJI TeJIJlT C nocJIe yroIII;eH 61I0JIornqe-
CKOii: IIp060ii: Ha II30JIH'pOBaHHoii: MaTRe KpblCbI. I\OJIUqOCTBO Hopa]:(pe-
HaJIlIHa B 3KCTpaKTaX COOTBeTCTBOBaJIO KO H e TBY KJIeTOK, onpe]:(e-
JIHeMbiX Ha rHCTOJIOrHqeCKHX Cpe3aX nOCJIe 06pa60TKH IIO aTOM KaJIUH
IIJIH q)JIyopeCI(IIPYIOIII;HX Ha clJopMoa-KaJIbI(HYMXJIOp HblX upenapa-
Tax (Eranko, 1955c). B CJIe yIOIII;eM ro y Rahn (1961) IIo]:(p06HO
oxapaKTepH30BaJI I(HTOJI9rHqeCKHe H rHCTOXH1.{HqeCKHe cBOii:cTBa IDle-
TOK KapoTH]:(HOTO naparaHrJIliH II Bb CKa3aJI COMHeHHe B IIX YBCTBII-
'feJIbHoii: TIplIpO e, npHMKHYB TeM caMbIM K qlICJIY Tex HeMHorllX
clBTOpOB, KOTopble Ha OCHOBaHHH CTPYKTypbl, paCUOJIOmeHHH II IIplI-
CYTCTBHJI KaTeXOJIaMUHOB, Ha nam B3rJIH]:(, IIpaBHJIbHO OI(eHHJIH 110-
JIyqeHHble ]:(aHHbIe. Muratori et a1. (1965a) nO]:(TBep]:(illIH Ha6JIIO-
.'(eHHJI 3. X. npHii:MaK, 1I3yqHB MeTO]:(OM Hillarp H Hokfelt
(1955) KapOTII HOe TeJIO KpOJIIIKOB H KomeK. CpaBHHBaH IIPH 3TOM
Hopa]:(peHaJIlffioBYIO ,peaKI(IIIO KJIeTOI, RapOTllJ];HOrO 'feJIa C ]:(PynIMH
naparaHrJIHHMII (no]:(KJIIOq1f1lHbIMH, aopTaabHO-JIerOqHblMII II Ha]:(no-
qeqHllKoM), aBTopbl OTMeTHJIH, qTO KOJIHqeCTBO xpoMaclJclJlIHHblx KJIC-
TOR, co epmaIII;HX HopaJ1:peHaJIIIH, y6b1BaeT B KpaHHaJIbHOM HanpaB-
JIeHHH.
PeaKI(UH Ha Hopa]:(peHaJIlIH. no ]:(aHHbIM npHil:MaR (1959, 1962),
XapaKTepH3yeTcH pH OM oc06eHHocTeii:. Bo-nepBblx, RHTeHCHBHOCTb
ee HapaCTaeT no Mepe YBeJIUqeHHH 'B03paCT8 mHBOTHbIX; BO-'BTOpblX,
no Mepe pa3BHTHH 60JIbmee qHCJIO RJICTOK BOBJIeKaeTCJI B IIpoI(CCC
CIlHTe38 ropMOHOB; B-TpeTbHX, ropMOH o6HapymnBae'I'CH IIpeHMYIII;e-
CTBeHHO B KpynHblx ilmeTI<ax KapOTIIp;HOrO TeJIa C KpynHblMII CBeT-
JIblMH JI]:(paMH H peme B MeJIKlIX, MaJIO HclJclJepCHI(lIpOBaBHblx; B-qeT-
BepTblX, He y 1!cex 06CJIe OBaHHblx mlIBOTHbIX KJIeTKH KapOTH Horo
TeJIa ].I:aIOT n03IITH'IJHYIO peaKI(HIO - y HeKoTopblX OHa B TeqeHHe
Bceii: mH3HH OCTaeTCJI HeraTIIB'Hoii: (Kpblca, Mblmb, MOpCKaJl CBIIHKa).
nOJIy'IeHHble naHHble rOBOpJlT 0 TOM, TO 1I03HTH1B'HaJl peaKI(HJI,
BbIHBJIHIOrn;aJl B KJIeTKaX MOHoaMHHbl, 3aBHCHT OT CTeneHH 3peJIOCTH
KJIeTOqHblX 3JIeMeHTOB II, COOTBeTCTBeHHO, pa3HOH ClIoc06HOCTH CIIH-
Te3HpOBaTb H HaKanJIHBaTb nHKpeT. RpOMe Toro, Y PMa MJIeKOnH-
TaIOIII;HX B KapoTII]:(HOM naparaHrJIHn, TaK me KaK B iHeKOTopblX
IIpIlcep]:(eqHbIX, peaKD;HJI BOBce ne o6nap.YiRHBaeTCJI. J];e HTKII pa60T
IIO]:(TBep].l:IlJIH ee nOCTOJIHHoe OTCYTCTBHC B KapOTIIp;HOM IIaparaHrJIHH
onpe eJIeHHblX BII OB mHBOTHblX, He 06HapymHB ee CTapblMII MeTO-
aMH H He UOJIyqHB n03IITlIBBblX pe3YJIbTaTOB IIPII IICnOJIb30BaHHH
COBpeMeHHblX Mo nclJHKaD;Hii xpoMaclJclJlIHHoii: peaRD;HH II 1I0]:(aTHoii
no XIIJIJIapny II XeRBeJIbTy.
O HaKO clJJIYOpeCI(eHTHblii: MeTO]:( <DaJIbKa II OBMeHa n03BOJIHJI
IIOJIyqlITb y MJIeKOIIHTaIOIII;HX 1I0JIO)KHTeJIbnyIO peaKI(IIIO Ha MOHoaMII-
Hbi B KapOTH HOM naparaHrJIlI1I RpOJIHKa, RomKH, TeJIeHKa, C1!IIHLII
II B JIerOqHO-aOpTaJIbHOM naparaHrJIHH (Fillenz, Wood, 1966; Mu-
ratori et at, 1966; Dolezel et at, 1968; Chiarini, 1970). HaMH OHa
06HapymeHa y KpblCbI, MOpCKOil: CBIIHKII B Tex lIaparaHrJIIIJlX, K01'O-
pble paHee CqHTaJIIICb (<HeXpoMaclJclJHHHbIMII» (CMHTIeB, 1970, 1971).
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B ap;B!'HTllIl)ljI BOCX01\HlI\l'ii 'Ianu I\)TI-I a0l'Tbl KpblCbI (no <1>aJIbRY II OmICHY)
A - OH. X 3, 00. X 10; B - 01<. X . lIO X 40
IlOl,a;mIlO npllCYTCTBIIC 1l0pa;_\l-l'Ua:llllla 1\ aOpTiI.'lhllhlX naparaUr IlUIX
(pnc. (ii, A, R), n Jlaparaur.'IJlHX .'Il'ro'IIlOii apT!'pUlI (pue. 62) II n
l,apOTII.o.BOM (pne. 63, A, B).
Y cncx B IIOJIY'IeIlIIII nO:IU.I,UTC,TlhUbIX CEC;l.CIIlIll 0 r.opMOB3JIbHOM
CO;l.epmllMOM lIapar3IIr.'1110Hi1pULIX 1,.1IeTOl, cnll.o.CTe:1J,eTBOB3JI He 06
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TOlIJI33Me HJICTOR l,apOTII;l.llllro 11 fJ,pymx ('HCXpmI3«P«PIIHHbIX) 1I3pa-
r3BrJIlleB MJIeIWIIUTalOIl\IIX II OOH3aTC:lhBO JIll OH Bo06r:u;e ]:(OJImelI
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n3pHbie CTpYlnypbl, B liOTOphiX IIC YAaCTCH oOHapymuTb MOH03MHIIhI
'IJ:lyopeCQeIITliLIM 11.'111 :llOubnl AP TlnI, I\IOi-liCT OLITb, er:u;e 60JIee qyn-
cTBIITeJIbHbIM I\le 1'O;l.OM. TO n :JTOM ne iiY;l.CT HllqerO Y]:(IIBHTeJIbHoro.
P3CCl\laTpIIllaH lIap3ranrJIII01H1IHl 'JO Tl,anh (' :mo.llO[{IIOHHOll TOqI>II
.lpeHIIJI, He TPYWIO yoefJ,IITbCfl, 'fTO. Tl'pHH II IIpOI(ec.ce IICTOPWlC-
cl.;oro p33BIITllfl IPYHl\QIIOlla.ilLHOe ;l1Ia'Il'III1J 3H,J,0l';PIlHIILIX ilie:Ie3, IIO-
oGHLle naparanrJIlllI Tl'pHIOT Cnol"OoIIOCTh npo YQUPOB3Tb IIHKpCT,
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TOI\ 11<1 paranrJIlUl B 6u4>Yl1KanIlIl
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B PYAUMeHT3pnbIU OCTaTHII.
lIo ne 'fOJIbIW npUMCnelIIIfJ nOBbIX rIICTOXIIl\lll'IeCIHIX l\IeTO]:(OH If3-
MCHIIJIO 1l0JIOmeH1Ie HCIIJ.cii n o6napYJIamo lIIOn03l\IIIHbl Ta l, rJ;c I1X
ne HaXO]:(Ilml. He MCIIec ]:(ellIonCrpaTIumo npIlCYTCTBIIC ropllIOHOB B
I>JIeTI>3X HapOTII]:(HOro naparaHrJIIIH nOlma3HO B aJICI\TpOHHOl\lm\po-
CKOlHPIeCI\IIX pa60T3x (Level', Boyd, .1957; Level', Lewis, Boyd,
1959: Garner, DUllcan, 1958; Hoffman, Birrell, 1958; Ross,
195 ; Kock de, Dunn, HW4; Palkama, H)6:>; MUl'atori, Battag-
lia, Modonesi, 19li5(b); Battaglia, IH68, 1970; Battagli'1. Ma-
riani, 1970; Biscoe, Stelrbens, 1965, H)6G, 19G7; Chiocchio
et aI., 1966; Fillenz, Woods, 1966; Kraus, 1966; Kraus, Mar-
tinek, 1967; Hoglund, 1967; Bliimcke, Rode, Niedorf, 1967;
AI-Lami, Murray, 1968a,b; AI-Lami, 1969; Grimley, Glennel',
1968; Chen, Yates, 1969; Chen, Yates, Duncan, 1969; Yates,
et aI., 1970; Zapata ct aI., 1969; Morita et aI., H)69, 1970;
Abraham, 19G8, 1970).
Bee oe-a nCKJIlO'ICIIITJI 1Ic--eJIe]:(on3TeJIII CYOllIIIRpocKonn'IecKoro
cTpoeHHH KapoTIl]:(1l0rO n3p:uanr.'lIIH naXO,I:\HJIU B HTDnJIa3l1Ie K-IeTOK
rpaHYJJbl C aJIeKTpOHHOJ1JIOTHhIlII eo]:(epmIlllIbIlII, no]:(06HbIe no CTl)YKT)'-
pc rpaHYJIallI M03fOBoro CIOH HajJ,nO'le'lHIIFon. TIpIlHa]:(JIemHOCTh co-
]:(epmllMoro rpaHYJI H MOHOal\UJHaM (a]:(pen3.'1IIIIY II Hopa]:(pena.TJIIIIY)
6hI.fla y6e]:(HTeJIbHO 1l0lH13RHa B HaAnO'le'IHIIKe :1BTopa]:(lIorpaiPn'IeCIUlM
MeTO]:(OM B :JJIeKTpOniiOllI l\lImpOCIWne (Axelrod ct aI., 1961; Hempel,
J\ra[t, 1968) II 6nOXIIMII'IeCIUlM RnR.flHaOM rpaHYJTHpHbIX' ())paK-
nii (Blaschko et aI., 1957), a B HapOTII]:(1I0M naparaHr.llIlI-
nCTOIIJ.eHlleM rpanYJI lIo("JIe pe:JepnIllln:!a IIII ,IHlBOTIIOro (Lever et
aI.. 195\"); Chen ('t aI., 19li9) II anTopa,I:\norpaiPncii B :JJIeHTpOH-
HOM lIIlIKpOCIwne (Chen, Yates, 1 UGU).
BTopoii qepToii YJIbTpacTpYI TYPHOii opranH;Ja lIn l\apOTlI I10rO
uRparaHrJIIIJI JIBJIH€TCJI cBoiicTBcHHoe BeeM ooeJle,I:\OnaHHbIM BII;:J;aM
160
MJIeKOnHTaIOIII;HX HaJIH'IHe Heo]:(uop-op;Hora KJIeTaqHara COCTa'Ba.
B IIpemHlIX pa6aTax 'cO' CBeTOBbiM MIIKpDCKOIIaM (CMHpHDBa, 1948;
de Kack, 1951, 1954, 1959; Willis, Tange, 1959; Rass, 1957;
Pessacq, 1961; IIpHHMaE, 1960, 1962) (pIIC. 64, 65) H lID YJIbTpa-
CTpyKTYPHbIM npH3HaiKaM Hau6D.lIee aTqeTJIHBa pa3J1If'IaIOTCJI p;Ba THiia
RJIeTaE (Lever et aI., 1959; Hoffman, Birrell, 1958; Rass,.
1959; De Kock, Dunn, 1964; Biscae, Stehbens, 1965, 1966; Chi-
occhio et aI., 1966; Kraus, 1966; Kraus, Martinek, 1967; Fil-
lenz, Waads, 1966; Hoglund, 1967; AI-Lami, Murray, 1968a, b;
Chen, Yates, 1969; Marita et aI., 1969, 1970) - 6aJIee 3JIeR-
TpOHHaIIJIDTHbie II pa3pemeHHble. B aCTaJlbHaM DnHcaHHe Map.pa-
JIarJf'IeCEHX IIpU:3HaKaB :JTIIX EJIeTaE p;aeT'CJI pa3HblMH a'BTOpaMH 110-
pa:3HOMY.
TpeTbllM BonpocaM, KOTapOMY 'B caBpeMeHHaH JIHTepaType yp;e-
JIHeT.cH 6aJIblllae BHHMaHHe npH a6CJIe,p;aEaHHH EapaTHgHDrD IIapa-
raHrJIHH, HBJIJleTCH 1I3Y'IeHHe IffiHepBa HH H YJIbTpaCTpyRTypbl CH-
HaUTIf'IOOE'IiX CBJl3eH.
Bce HaEOIIIffiIllHOCJI E HaocTOHlI\eMY HpeMeHH .paRTHqeCEHe CBe-
p;eHHJI a cy6MHRpaCROnHqOCEall apraHH3a HH ImpDTH,IJ;Horo IIaparaHr-
JIHJI IIMeIOT 3'HaqeHHe p;JIJI IIaHIIMa.HIIH .pYHI,wmHaJIbHaH paJIII KJIeTOR
EapaT HaM a6JIaCTH H pa36npaIOTcH HaMH !B CffiJl3II C TeMII TpaE-
TaBEaMH, EaTopble CYIII;eCTBYIOT B Ha'CTaJlIII;ee 'BpeMH no 3TOMY 1I0II-
pocy.
Lever c c'OaBTopaMH (1959) p;aJIII 6aJIblllaH .paETHqOOEHM Ma-
TepHaJI, xapaETepH3YIOIII;HH TaHEYIO CTPYRTYPY RapaTIIp;Horo napa-
raHrJIIIJI II npaBeJIII rHCTDJIarHqe'CEOe H l'H'CTOXIIMUqeCEae era 06cJIe-
p;aBaHHe. OHH nalliJIH, qTO IIaIIYJIJll(IIJI rpaHYJI B TeMHblX EJIeTEaX
BbllIIe H aHH 6aJIee oCMlla.pIlJIbHbI, qeM B CBeTJIbIX, ] qTa p;JIJI na-
CJIep;HHX xapaETepHa JIe60JIblllae EOJIllqeC'rBa HJIH aTCYTCTBHe q>aHYJI.
8TII RJIeTOq'Hbie Bapllal(HI:1 H IIepexap;Hble .papMbl Memp;y HHMH aBTD-
pbl CqHTaIOT BblpameHlleM l(JIEJIHqeCEOM p;eHTeJIbHOCTII RJIeTOE, Ha !BOB-
p;epmlliBaIOTCJI aT CROJIbEa-HH6YP;b aIIpep;eJIeHHblX BblCRa3b1BaHIiH 0
.pH3HOJIOrHqeCHOH CYIII;HaCTH 1II\JIOB; OHH TaJIbEO npep;naJIamHTeJIbHO
raBOpJlT 0'6 HX ceEpeTapHaH IIpllpap;e. OCHCBHaJl MblCJIb aBTop-on Ha-
IIpaBJIeHa Ha ToO, qTo6bI nOHJlTb MexaHH3M JIaEaJIbHarO .pU:3HOJIOrn-
qeCKOra ]:(eHCTBlIJI EaEara-Ta rapMOHaJIbHarO areHTa (B03MamHa, Ra-
TeXaJIaMHHOB), HaXO]:(HIII;erDCJI B rpaHYJIaX B 3anaCHaM .popMe. 8TOT
areHT p;aJImeH JlBJIJlThCJI MeCTHbiM aRTlIBaTapOM qYBCTBUTeJIbHbIX
aEaHqaHHH 'HepBa XepHHraH TeM CJIaCa6CT'BOBaTb HX pe enTopHaH
<PYHE lIH B IIaHlIMaHHH ue Castro. BnpaqeM, CTpeMJIeHHe Hamil
.pH3HOJIarnqe.CEHM pa3p;pamlITeJIb P;JIJI qYBCTBHTeJIbHoro HepBa IIp'll-
Hap;JIemliT iRe TaJIbEa Level', Lewis, Boyd, Ha CBaHCTBeHHO :r,1H0rJ'1M:
aBTapaM, a6'bJlCHHIOI Hl\1 npIlcyTcTBHe !f>eHaJIbHblX aMHHaB B Eapa-
TlI]:(HaM naparaHrJIlIlI HX 3HaqeHlIeM B aCYIII;eCTBJIeHlIH pe.pJIeETOp-
Haro aKTa.
ECJIII IIplI3HaTb TaqEY 3peHHH Lever C caaBTapaMII a JIaEaJIbHaM:
p;eHCTBJlII Bblp;eJIJleMbiX RJIeTEaMH rapMaHaB, TO aCTaeTCH HeIIaHJlT-
HblM, naqeMY Bblp;eJIeHlIe EaTeXaJIaMIIHaB B EpaBb II3 RapaTIIp;Horo
6 H. A. CMHTTeH
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rraparaHrJIlUJ I!eCl\a (A) II Rom-
1m (B) «TpJ\\Iloflp;epm,j('» 11 «cneT-
JIOHp;epHb\(,» KJIeTKIf
ORp. MPTnJ10BhIM 3e.lIeHbIM-nnpOHn-
HOM. 0,<. X 7 06. x 40 (IIpnfiMaR,
1962)
P II c. 65. MoaroBoH CJIOH nap;-
nO'le'lHIIRa upoJIlIRa. «TeMHo-
flp;epHlde» II «CBeTJIOJIp;epHlde»
RJIeTUII
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HnIlOM. OH. X 7, 06 X 40 (npena-
pal' IIpllilMaR)
napara.arJlHJI OrpaHUqllBaCTCH l\ICCTHb1M p;eHCTBlIeM, a Bb1p;e.JIeHlle Tex
me KaTeXOJlaMIiHOB 113 APyrHx naparaHrJlHeB, HanpHMep, HaAnoqeq-
HHROB, HBJlHeTCH 06IJJ;IIM. HenOHHTHO II iIlPyroe 06cToJlT,eJlbCTBo. AB-
TOpbl, XOTJI II Haxop;HT HeKOTopble cxop;Hble qepTbi B CTpoeHIIH Ime-
TOR KapOTH,D,HOro naparaHrJlHJI n Hap;nOqeqHHKa, HO He IIplf3HalOT,
O)J;HaKO, HX IIOJlHOH aHaJl.orHII Ha TOM OCHOBaHHH, qTO ClIMnaT3RTO-
MIiJI He conpOBomp;aeTCH p;ereHepa HeH OKOHqaHHH Ha nx IIOBepx-
HOCTII.
Memp;y TeM :nOT .paKT KaK pa3 H Hap;O paCCMaTplIBaTb RaK IIOP;-
TBepmp;eHHe op;HOTlinHOCTII HHHepBaQIUl M03roBoro CJlOH Hap;IIOqeq-
HIIKOB II KapOTIIp;Horo lIaparaHrJlHJI If, CJlep;OBaTeJlbHO, RaK JIlIillIlllH
P;OBOP; B IIOJlb3Y aHaJlOr1f1f 3TlfX OpraHOB. TIOqeMY 3TO IIMeHHO TaK, CTa-
HeT JlCHO nocJle paCCMOTpeHIfH IIHHepnHQII.OHHblX OTHOllieHIIH B Ha.n;no-
qeqHIIKe, KOTopble 1f3J10meHbI Hlfme.
Ross OIIHCblBaeT B p;OJlbl\aX rJIOMyca p;Ba KjleTOqHb1X TIIIIa, ORPy-
meHHblX CIiHycolIp;aMu: «XeMOpe enTOpHble) II oKpymalO IIe llX «cy-
C'I'eHTaKYJIHpHble), KOTopble Ire OKOJlORJIeTOqHOH JIOKaJIH3a HH nepe-
XOAHT E HHTepcTH Hll, rp;e conpoBomp;alOT aKeOHbI. CycT,eHTaKYJIHp-
Hble RJIeTKH 110 CBOeiMY 'pa-onOJIOmeHIIIO BOKpyr xeMopeIJ;euTopHblx
COO'11BeTCTBYIOT II THrry JIeBepa c COaB'I'OpaMH (Lever et a!., 1959),
HO [1.0 CTpyRTYPHo--4JYHK HOHaJIbHbll\1 OBOHCTBaM TpaKTYIOTCJI POOCOM
EaR 1I0]:(p;epmHBalOIJJ;1Ie 3J1eMeHTbI. Ross Tome He HallleJi B KapOTlIp;-
nOM lIaparaHrJIlI1I ROHTaKTOll HepllHblX OKOHlIaHlIH, xapaKTepHblX P;JlJI
3.p.pepeHTHbiX CIIHancoB. Ho allTOp, npHp;aBaJl 60JlblllOe 3HaqeHHe
nplICYTCTBlI10 3JIeRTpOHiWIIJIOTHbiX rpaHYJI C .peHOJIbHblM oop;spmll.,
Mb1M, KOTOpble CXOP;HbI C TaKOBbiMII B MoafOBOM CJloe Hap;nOqe1fHlI-
KOB, nOJIaraeT, qTO IIX HaJlUqlle T'pe6yeT OCTopomHOCTIl B TOJIKOBa-
HHII IIpllp08bI «rJIOMYOOblX) KJleTOR II uepeCMOTpa Y'CTaHOBImillIlXCH
B3TJIJlP;OB Ha HIiX RaK Ha pe eIITOpHLIX (de Castro, 1926; Boyd,
1937; Hollinshead, 1943). OH cOBepllleHHo npaBUJIbHO nHllIeT, qTO
cJIa6aJ1 xpoMa.p.pHHHaH peaK IIH B 3TIIX RJIeTRax 06'bHCHHeTcH CKY"
P;OCTblO H paa6pocaHHocTblO rpaHYJI 110 HTOnJIa3Me.
E e 60JIee npaBIIJIbHO, C Halliell TOqKH3peHIIH, K o eHKe 1/I;aH-
HbiX 3JIeRTpOHHOH MHKpOCKonlIU IIOP;XOP;HT de Kock II Dunn (1964).
lfx rJIaBHOe B03pameHue npOTliB ceHcopHoH .pYHK HII KapOT -
Horo naparaHrJIIiH 3aKJIIOqaeTCH ;B TOM, qTO BPHA JIH XHMH"'leCKlIMlI
pe eIITopaMII KpOBOTORa MOryT 6LITb KJIeTKH, KOTopble II30JIUpOBaHbI
OT npOCBeTa KanHJlJlJlpOB (nOMHMO nJlOTHoro 3Hp;OTeJIHJI II ero 6a-
3aJIbHOH MeM6paHbI) Mo:r.qnblMH KOJlJlareHOBblMH iBOJlORHaMH, IIP;y H-
MH B IIpOP;OJlbHOM H IIOIIepe'IIlOM HanpaBJIeHIIHXII 06pa3YIO m.1H
P;O illecTII CJIOeB. R qH JlY TKaHell, OTp;eJIHIO HX ClIe H.pK'leCRHe
KJIeTKH RapOTHp;Horo 'feJla OT KpOBII, aB'IOpbl OTHOCJlT H 1I0Mep-
mnBalO IIe KJleTim. 06 II30JIH llII KJleTOK OT npOCBeTOB COCYAOB 00-
06 a1OT II I1J;pyrHe aBTopbl (Lever, et a!., 1959; Ross, 1959; Bis-
coe, Stehbens, 1966; AI-Lami, Murray, 1968a). TIo Ross (1959),
06a RJIeTOQHblX TlIna OTp;eJleHbI OT CIIHYCOHP;OB IIHTepCTII IIaJIbHLlMII
IIpOCJIoimaMH KOJlJlareHa, .pH6p06JIaCTOB H 003MHKOT"HbiX HepBllbiX
ROJIOROH C IIX 060JIOqKaMII II3 illBaHHOBCKlIX KJIeTOK.
6" 163
13 pa60Te BiscDe II Stehbens (1966) nDKa3aUbi [lJIDTHbie TDJI-
-'CTble 'CJIDH RDJIJIarOOa, IWTDpble OTp;eJIHIOT [lpOCBeT Ra'llHJIJIHpO'B DT
cne HlIJHqe'cRnx RJIeTDK RapDTH)];HDrD TeJIa, a TaRme 3Hp;OTeJIHaJIbHbIe
""CTeHRH KalIHJIJIHpoB C liIeHecTpan:nHMu, qTD, ImR H3BeCTHD, CaMO
nO' ce6e JlBJIHeTCH xapaRTepHblM nplI3HaRDM 3HP;DRpIIHHblX meJIe3.
lID Ha6JIIOp;eHHHM AI-Lami II Murray (1968a), liIeHeCTpa HH 3a-
TJlHYTbI TDHRHMH nepenDHRaMII, qTO, Ha Ham 13'3rJIHp;, MomHD pac-
1J;eHHBaTb, RaR IIplI3HaR CHlImeHHDH :mp;DRpIIHHDH IiIYHIO::\IIII RJIeTDR.
AiBTDp OnHCaJI eIU;e DP;HO' BamHDe Ha6JIIOp;e'HHe: THmeJIaH aHDRCHJI
Bbl3bIBaJIa He HCqe3HOBeHHe OCMUDIiIHJ[bHblX rpaJIyJI B RJIeTRaX Ra-
pDTI:J;AHOro TeJIa, RaR 3TD D6b1qHD Ha6JIIOp;aeTCH, a Hapa,cTaHHe nx.
CDp;epmaHHe Tex HJIH HHblX aMHHOB B RJI,eT-RaX RapDTHp;HDro lIa-
paraHrJIHH B HacTOHln:ee BpeMH P;H'CRYTHpyeTcH. Battaglia (1968)
BblHBHJI Y KpbllC 'l'Dn:bRD ap;peHaJlHH, 'B TO BpeMJI RaR B M03rOBDM
cJloe HaMO"IeqHHRa Tex me mHBDTHbiX - 06a aMHHa. B pa6DTe MD-
rita C CDaB'l'OpaMII (1969) B RapDTHgHOM lIaparaHrJIHH KDIlIRU nD-
Ra3a o TJeTbipe TH.IJa RJIeTDR II D6cymp;aeT,cH 'cDAepmaHHe B HHX
ap;peHaJIIIHa, HDpa,n;peHaJlHHa, p;DIIaMIIHa II cepDTDHHHa (RJleTRH p;HIiI-
liIepeH Ii:pYIOTCH Ha OCHOBRHHII BeJIH"'lUHbI, RDJlHqeCTBa, «pDpMbI H
3JIeRTp01IHDH nJIDTHOCTH rpanYJI). B 1970 r. Te me {I!BTOPbl coot)-
IIJlH, qTO 4>apMaROJIDrlt'tleCRne Boop;eHC'TBlIJI tIJln C!fnMYJ1J:I,n:na: I{Q-
pDTI:I!p;HDrO HepBa MeHJlIOT DTHDCHTeJlbHDe RDJIH"'lOCTBD pa3HblX TlIllOB
RJleTDR. KpoMe Toro, 3J1eRTpDHHDMHRpOCROIIH IeOI\'lIe IICCJlep;DBaHHH
IIOCJIe)J;HHX JIeT IIORa3a.lIH, qTD MOplilDJlDrIm 'II RDJlH"'leC:TBO OCMIro-
IiIIIJIbHblX rpaHYJI 3aBHCHT DT nplIMeHHeMbiX IiIHRcaTDpDB (Duncan,
Yates, 1967). Bce ::ITD 3aTpY)J;HHeT 'II'H'l'epnpeTa HIO 'lIOJlYTJaeMbIX
aHHblX.
He'CMO'l1pJl Ha pHP; HeJlCHOCTeH, HaM RameTCH npa'BHJIbHblM CTJII-
"TaTb, qTD B RapDT HDM IIaparaHrJIIIH MJIeRDIIHTHIOIU;UX IIMee'l'CH p;Ba
RJIeTDqHblX TIIIIa (He Cql:lTaH 3Hp;OTeJlIIH II CDep;IIHI:ITeJIbHDTRaHHbIX
.3JI-8MeHTOB), DP;IIH 1:13 RO'TOpblX DTHDCHTCH R CIIIe II«pHqeCRHM napa-
raHrJlI:IDHapHblM H .cop;'epmI:lT rpaHYJlbI RaTeXOJla!MIfHOB, a APyroH R
BCIIDMDraTeJlbHblM - «, CYCTeHTaRYJlHpHblM» (<IIOp;p;epmIIBaIOIU;HM),
«rJlHaJIbHblM). BHYTpH nOIIYJlH IUI cnen:MIiIHTJecRHX RJleTOR He DP;HD-
pDP;HblH COCTaB D6Y'CJlDBJleH HX pa3HblM IiIYHRn:HDHaJIbHblM COCTDH-
DHeM. qTD me RacaeTCH ap;peHaJiIIH- II Hopa:p;peHaJIHHCOp;epmaIU;HX
RJleTDR B COCTa'Be RapDTHp;HDro lIaparaHrJlIIJI, TO' nORa IIX MomHD
Bb eJIHTb TOJIbHD rHCTDXHMHqeCKIIMH MeTop;aMIi. Cne HaJIbHblX YJIb-
Tpa'CTpyRTYpHblX xapaKTepn.CTHR P;JlH Tex H p;pyrnx HeT.
HalI60JIee IIHTepeCHbiM II BamHbIM BDlIpDCDM B YJIbTpaCTpYRTyp-
HDM DpraHII3an:HH RapDTH)];HDrD naparaHrJIIIH MJleRDIIIITaIOID;IIX HBJlJleT-
CJI HCCJlep;DBaHHe ero CHHanTlIqeCRIIX CBH3eH, RDTDPblM B HaCTDHID;ee
BpeMJI npHAae'l'CH 60JIbmOe, eCJIH He rJlaBIIDe, i,'maqeHHe npM Dn:eHKe
liIymtD;HDHaJIbHOH pDJIH ero RJIeTDR. PaCCMOTpeHHe HHHepBa HDHHblX
D'IIHOmeHHH 'HaM RameTCH BamHblM He nDTDMY, qTO apXHTeRTDHHKa
II IiIYHR UOHaJlbHaJl CYIU;HOCTb HepBHblX CBH3eH (HX nplIHa)];JlemHOCTb
R TJyncTBHTeJIbHblM HJIII p;BHraTeJIbHbIM HeiipDHaM) MDmeT "lJTD-JIH6D
II3MeHHTb B Dn:eHRe IiIH3HDJIDrIf'IeCRDH pDJIH RapDTHAHDro oprana RaR
t
3HP;ORpIIHHOH meJIe3b1, Ii nOTOMY, 'ITO B 3THX BOnpOCax HMeeTCJt
.oC{)6eHHO MHOro HeOOOCHOBaHHblX IIpep;nOJIOmOOIIH II 3aRJIIOqeHIIH.
XapaRTep 1I01lCROB B :COBpeMenHhiX IICCJIep;OBaHlIHX CJIOmIIJICH
"Ha 4>oHe P;IfCRYCCIIH, a B IIOCJIeARee BpeMH yme I'ocnOJJ;CTBYIOIII;IIX
YTBepmp;eHIIH 0 .pYHR HOHaJIbHOH pOJIII RapOTIIp;Horo TeJIa RaR xe-
1dOpen:eIITOpa. QT06b1 pa306paThCH B HOBbiX cP.aRTax II nx TOJIKOBa-
mm:, Heo6xop;IIMO BOIIOMHIITb TO, qTO 6b1JIO IIo;rryqeHO B npemUIIX
"IlCCJIep;OBaHIIHX - p;o HaqaJIa cy6MIIRpOCROlIuqeORoro II3yqeHIIH CTpyK-
TYp.
lf3yqeHHe Mall'pO- II MIIRpOCROnHqeCROH Mop.pOJIOrIIII 'lIepBHOfO
Ha6meHIIH RapOTlt]l:HOH 06JIaCTH IIloIeeT p;JIHTeJIhHYIO 'lICTopIIIO,
Ha IIpOTHmemI1I ROTOpOH 06cymp;aJIOCb II pelllaJIOCb MHoro P;IICRYC-
ClIOlI'HblX .BOIIpoCOB. U3JIOmeHIIe OIIIICaTeJIbHblX II oomnepIIM8HTaJIbHblX
MaTepHaJIOB 110 IIHuepBan:IIII l\apOTIIp;HOH 06JIaCTII y pa3Hblx :r.meRO-
IIIITaIO:rn:IIx II IfC"IepnLIBaIO:rn:YIO OBOP;RY JIIiTepaTypbl MOlli'HO HaHTII
n CTaTbJlX II MOHorpa.pIIHx Hering (1924, 1927), Driiner (1925),
de Castro (1926, 1928, 1929, 1951)t Boyd (1937a, b), A. A. CMlIp-
HOBa (1945), B. H. IIoHoMapeBoH (1946), Abraham (1953),
T. A. rpnropbeBOH (1954). Adams (1958), Comroe (1964) II p;p.
Ha OCHOBaHIfII aTUX HCCJIep;OBaHIiH IIHHepBa IIH lRapOTI oH 06JIaCTH
Y MJIeROIIIITaIO:rn:IIX IIpeACTaBJIHe'l'CH CJIep;YIOIII;IIM 06pa30M.
TpII HepBa BXOP;HT B COCTaB RapOTIIp;Horo CIIJIeTeHIIH - H3b1RO
fJIOTOqHbIU, 6JIymp;aIOIII;IfH II CHMlIaTIIqeCRIIH, ROTOpble HHHepBII'pYIOT
06JIaCTb p;CJIemm 06:rn:eH COHHOH apT.epIIR. Y HeRoTopblX BIIJJ;OB mII-
BOTHhIX R HUM [l,pIICOep;lIHHeTCH 1I0P;'b1I'3b1"IHblH. MJlROTHbie H 6e3MH-
ROTHbie BeTBlI aTIIX HepHOB pa3BeTBJIJlIOTCH m CTeHRax RapOTIIAROfO
clIHyea II B RapOTIIp;HOM «rJIOMyce», 06paaYH a.p4>epBHTHbie II a4>-
.pepeHTHble HepBHhIe OROH'IaIIIIH (Wilson, Billingsley, 1923; de
Castro. 1926, 1928, 1951; Hiegele, 1928; CMlIpHoBa; 1948; Vaida,
Duca, Lazar, 1966). IIo OlIlI'CaHIIHM H. f. CMHpHOBOH (1948),
OqeHb 'CXO;!);HblM C TeM, qTO p;aeT de Castro, a.p.pepeHTHaH IIHHep-
Ba lIJ1 TRaHeH RapoTIIp;HOro naparaHrJIIIH ocYIII;eCTBJIHeTCH TOJlCTblMII
II cpep;Hero KaJI1I6pa MHROTHbiMII nepBHbfMII 'BOJIORHaMlI, ROTopble,
TepHH MIIcmlH, IIpli Bxo]:(e B P;OJIbRlI meJIe:abl pa3BeTBJIHIOTCH B HlIX
II 06paaYIOT CJIOmHble HepBHble alIlIapaTbi Ha [lOBepXHOCTU «'I'JIOMYC-
HLIX)} RJIeTOR. Mop.pOJIOrIIqeCRIIMII ()C{)6eHHocTHMII, xapaRTepII3YIO-
III;IIMll qync-TBlITeJILHbIH TIIn oRouqaHIIij Ha «fJIOMYCHblX)) RJIeTRaX,
JlBJIHIOTCH TepMMHaJIbHble peTIIRYJIHpHble nJIaCTIIHRlI II CeTeBIIJJ;Hble
CTPyRTypbl. OT Tex me a.p.pepeHTHbIX BOJIOROH, ROTOpbIe IIHHepBII-
PyIOT «rJIOMY;CHble)} RJIeTRII, nOJIyqaIOT qyBCTBUT,em:'Hbie OROH"IaHIIH
II COCY]:(bI RapoTIIp;Horo «rJIOMYCa)}. H. r. CMlIpHOBOH OIIIIcaHbI pa3-
BeTBJIeHIIH op;Horo n TOro me a.p.pepeHTHOfO BOJIORHa Ha «rJIOMYC-
HblX)} RJIeTRaX II Ha CTeHRax COCYP;OB. 8.p.pepeH'l1Hble O1WHqaHlIJI B
TRaHHX RapOTII)]:Horo lIaparaHrJIIIH 06pa30BaHbI TOHRllMII 6e3MHROT-
HbiMII BOJIORHaMII, ROTOpble ORaH"IIIBaIOTCJI Ha «rJIOMYCHblX)} RJIeTRaX
CIIUpaJIeBIIp;HblMll HaMOTRaMII C TOHRlIMII ROHn:eBbiMII ROJIeqRaMII II
HlIlwrp;a He- 06pa3YIOT ceTeBlIp;HblX CTPYRTYP, CBOHCTBeHHblX qyocTBII-
Te.J1bHblM HepBHblM OROHqaHIIHM.
1R.
TIpIIBe}];eHHble I1CCJIe}];DnaIIHH KDHl1;enblX DT}];eJlDn HepnHblX CTPYK-
TYP RapDTII}];HDH D6J1aCTH DTJlII"'laIO'fCH Tll{aTeJlbHDCTblO II qpe3BblqaH-
HDH TDqHDCTbIO DIIHCaHHH, CnDHCTBeHHblMII TaKIiM aBTDpaM, KaK
de CastrD H BceM pa6DTaM IDKDJlbl B. 11. JIaBpeHTbeBa, R KDTDpbIM
IIpIIHa,rJ;JlemHT lIy6J1HRal1;I1H H. r. CMHpHDBDH ( 1948). <1>aKTH'IeCnaa
cropoHa 3THX HCCJIe}];OBaHHH IllITe-peCHa C JI.BYX l'DqeK 3peHHH. Bo-
IIepBblX, -cone.pllleHHD HCHO, qTD HeT If30JlHpOBaHHDH IIHHepnal1;IIH
«TJlDMYCHblX)} KJleTDR, KpDBeHOCHbiX c.ocY}];OB «rJlDMyca)} (n CDCTaB
CTeHDR RDTOpblX, HapH}];Y C APyrllMH TKaHHMII, BXD}];HT II caMII «rJlo-
MYCHble)} KJle'IKH) H RapDT HDrD cIIHyca, a HMeIO'ICH HepBHble C'IBO-
JlbI, KOTOpbIe pa3BeTBJlHIOTCH BO Bcex TKaHHX 6II.pypKa11;1lH 06ll{eH
COHHDH apTepHH H B 6J1H3aemall{eM paoclIIlIpeHIIDM yqacTRe. 06 0'6-
ll{HX 'BeTJJJIX, HHlIepBHpYIOll{nx <:ITII C'IPYKl'ypbl, IIMeIOTCH }];aHHble II
B APyrIlx pa60Tax (Meiyling, 1936; JIbice, HKO'B 1939; CMHpHDn.
1945; MuratDri, 1937).
Bropoe, qTD JIBCTnyeT If3 IIpIIne}];eHHblX IIiCCJle}];DBaHIIH If qTD 'Ie-
IIepb IIHOI' a .aa6L1BaeTCH, :nD }];anHD YCTaHonJleHHaH ill 3K<mepIIMeH-
TaJll/HbiX II o:r:nu;aTeJlbHblX pa6DTax RJlaCCII"'leCKIIMII HeiipDrIICTOJlDrII-
qeCKIIMII MeTop,aMII }];BOHHaH IIHHepn3.I1;HH KapDTII}];HOH meJIe3bI - 3.p-
c}lepeHTHaH II a.p.pepeHTBaJI. AD 20-x rD}];DB TeRYll{erO CTDJleTHJI
npeIIMYll{eCTBeHHDe BHIIMaHl1e y}];eJlHJlOCb CHMlIal'II"'leCJ\on HHHepna-
J:\IIII. Eo rJlaBe aToro HalIpanJleHIIH CTOHJI KDhn II erD IlIKOJIa. Ha-
'IlIHaH C repe}];HHbI 20-x rof!;DB 1I }];D BaCTDJllI{ero BpeMeBII rJlaBHblH
mITepeC anTopDn COCpe}];DTDqHJlCH Ha If3YQeHHII cHa6meHHH DpraB8
qYBCTBIITeJlbHbl'MH BepBaMIi. C1.I8Ha lIHTepeCOB H Banpa'BJleHHH nc-
CJle DBa'HlIH 6LIJIa Bbl3BaHa IIDHCKaMH Mop.pDJlOrHqeCKDrO cy6cTpaTa.
ocym;eCTBJlHIOm;ero cllHyc-pe.pJleKc Hering II xeMDpeJ:\elI IIIO, IIPUIIII-
CblBaeMYIO «rJlDMYCHblM» KJleTRaM de CastrD.
YqaCTlie CIIMIIaTlIqeCKHX nDJlDKDH B IfllHepnal1;HII KapDTII;J;HDH
06J1aCTII IIplI3HaeTCH, D}];HaRD, He TDJlbKD CTapblMII anTDpaMII II IlIKD-
JlDH KDhn, HD II B 6DJlee IID3}];HHX pa60Tax (Ask-Upmark, 1935;
.JIbloeHKoB, 1939; Palumbi, 1940; Abrahalll, 1953, 1964). MHTepec-
BD OTMeTHTb, qTO -caM de CastrD - D}];IIB 113 DCBDBOnOJlOmHIIRDB
MDp DJlDrllqeCKHX HOCJl8}];DnaHHH 'Iync'I'BIlTeJlbHDii mmepnaJ:\I1H pa3-
BIIJlKH 06111;eH CDHHDH apTepIDI II KapDTIf,lI;BDrD TeJla - OIIHCaJi B CBDeH
nepBOH pa6DTe 1926 I'O}];a ABOHHYIO .nIlHepnal1;lIIO KJleTDK KapD'III}];-
HorD lIaparaHrJlHJI H IIpH3BanaJi erD meJle:3IICTYIO .pYHKJ:\IIIO. Peq,-
JleRTOpHoe BJlHHBHe IIpn ocym;eCTBJlBHHII meJle3lIiCTDH .pYHKI1;IIH 3a-
RJlIOqaJlOCb, lID anTOpy, B CJle}];YIOll{eM: H3MeHeHHe KponHHDro }];aB-
JleHHH H COCT3'na KponlI B036y ae'I qYBCTBIITeJlbHble HepBHble
DKDHqaHIIH, DT KDropblX lID J:\eHTpOCTpe m.TeJlbHblM nyTHM H3b1KD-
rJlOTOqHDro Hepna pa3]:(pameHHe IIepe}];aeTCH B IIpD}];DJlrDnaTblH M03r II
oTTy}];a nD a4>4>eKl'OpHblM nOJlDKHaM TDro me HepBa J:\eHTpaJlbHblU
HMIlYJlbC }];OCTHraeT RJleTrOK RapOTH}];HOrD TeJla H Bbl3b1BaeT I1X ce-
KpeJ:\HIO. OqeHb BamHO DTMeTH'Ib, qTD B cnoux cym}];eBHHX de Cast-
rD (1926) nbl<CKa3b1BaJlCH He TOJlbRD '0 .pYHKI1;HH caMHX KJleTDK Ra-
poTHJ{HOrD lIaparaHrJlHH, HD DTMeqaJl H .pH3HDJlDrlIqeCRDe 3HaqeHHe
CeRpeTa, nbl}];eJlHeMOrD IIMII B KpDnb. OH He npIIllIICblBaJi eMY TOJlb-
166
1\0 MCCTBYIO pOJIb pa3;ZJ;pamlITeJlH pert;elITOpOB aToro me Opra'H8,
.a nOJIaraJl, qTO ceEpe-r, IIona;ZJ;aH .B Rp<>BeHocHoe PYCJlO, OEa3bIBaeT
06III;ee ;ZJ;eHCTBlIe Ha opraHII3M. TaRHM 06pa3OM, de Castro, XOTJI
.II rOBOpIIJI TOJIbRO 0 lIa-paCHMlIaTIIqeCEOM acpcpepOOTHOM cHaomemm
RapOTIIAHoro IIaparaHrJlHH II eqIITaJi CIIMnaTuqecElIe BOJlOKHa HHlIIb
:nPOXOAHIII;HMII qepe3 Hero R CORHbIM apTeplIJIM, H'O p8J3;ZJ;eJlHJI MHeHJIe
CBOIIX npe;ZJ;IlIHCTBeHHlIKOB 0 meJleaIICTOM XapalJ\Tepe EapO'l1H;IJ;Horo
IIaparaHrJlIIH. B 1928 r., Ha OCHOBaHlI1I HOBblX aRcnepIIMeHTOB,
de Castro OTRa3aJlCH OT CBOIIX npemHIIX nOJlOmeHIID:. nepepe3aJl H3b1-
Ro-rJlOl'Oq}lblH nepn BUYTPII qepelIa II, CJle;ZJ;OBaTeJlbHO, Y;ZJ;aJl1IJ1 acp-
. .cpepeHTHbie nOJlOKHa, OH He HallleJi ;ZJ;ereHepIIpyrom;IIX OROHqaHIIH H8
-«rJlOMYCHblX» RJleTEaX H Ha aTOM OCHOBaHHH YTBepm;ZJ;aJl, 'ITO CHHYC-
nepB He co]:(epmlIT rt;eHTp06emHblX ceRpeTOpHblX BOJlOROH, a TOJIbRO
{Jert;eITTOpHble II, CJleAOBaTeJlbHO, EapOTIIJ{1IblH «rJlOMYC) He MomeT
6bITb meJleaOH, a HBJlHeTCH qYOCTBIITeJlbHbIM opraHOM, ROTOpblH CJIy-
JIHIT ;ZJ;JlH nOCnplIHTHJI RaqeCTBeHHblX II3MeHeHIIH EpOBII. R JK :mme-
CTHO, aTO UOCJle;O;Hee YTBepm;ZJ;eHIIe de Castro He nO;ZJ;TBep;ZJ;IIJlOCb.
Biscoe C coaBTOpaMII (1969) IIOBTOpIIJlII aROnepIDIeHT de Castro,
:nepepeaan H3b1EO-rJlOTO'lHblH HepB BHYTPII 'IepMa; B pe3ym.TaTe
aJleE'l1pOIiHOMHRpOCROnH'IeCEOrO I13Y'IeHMJI EapOTII;ZJ;Horo IIapaTaHrnRJI
B 1I03;ZJ;HlIe CpOEII IIOCJle OlIepart;1I1I 6bIJia 06HapymeHa ;ZJ;ereHepaII;HH
acpcpepeHTHblx CIIHanOOB Ha «fJlOMYCHblX) EJleTEax.
C EaTeropIIqeCROH TO'IROH 3peHIIH de Castro 0 KPYHRrt;III1 Ea-
pOTII;ZJ;Horo TeJla em;e II B Te rO;ZJ;bI He CorJlaCIIJlCJI Sunder-Plassmann
(1930), ROTOpblH YTBepm;ZJ;aJl, qTO RapoTH,U;Hoe TeJlO He MomeT
QblTb aHaTOMIPIeCIWH OC'l!OBOi'1 CIIHY'CHbiX peKPJleRCOB Hering 110 TOH
IIpuqlIHe, "'ITO cnen;RcpIlqeC {IIe pen;enTopbl !H8XO;ZJ;HTCH He B HHM, a B
CTelllie EapoTlIJtHOI'O cllHyca. HOOEOJlbEO II03me Palumbi (1940)
mICaJl, qTO pert;eIIrt;IlH ocyro;eCTBJlHeTCH EOHl1;eBblMll 1Ip1I6opaMII H3bIRO-
rJlOToqHOrO II 6J1yiK;ZJ;arom;ero HepBoB, EOTopble 06pa3yroT CJlomHble
pa3BeTBJleHnJl B apTepllaJlbHblX CTem\BX, B nepIIapTepllaJlbHOH 06J1a-
CTII II BOEpyr ;ZJ;OJleE «rJloMy.ca) II 'ITO pecpJleEC Bbl3b1Ba-eTCJI He
pa3;ZJ;pameHlleM n;eH'11pOCTpeMIITeJlbHOH CllCT.eMbl EapOTH;IJ;HOro TeJla,
a 06YCJlonJleH pen;ellTOpHbfMlI aIInapaTaMII, pacnOJlOmeHHblMlI BO BceH
RapOTH,U;HOH 06J1aCTll. .
TaEllM 06pa30M, OOBORYIIHOCTb KPaRTH"IeCEIIX ;ZJ;a'HHbIX, nOJlyqeH-
HblX B 30-40-e rO;ZJ;bI, roBOplIT 0 TOM, 'ITO EapOTII;ZJ;HaH 06J1aCTb
06J1a.u;aeT ;ZJ;BOifHOH llHHepBal1;Hei'1 II qTO HeTBII O;ZJ;HOro HepBHoro CTBO-
JIa MOryT JlHHepnlIponaTb pa3Hble TImHn aTOH 06J180CTII.
RaE me paCn;eHllnaro1'CH IIHHepnan;noHHble OTHomeHIIH B RapoTII]:(-
HOM lIaparaHrJllI1I Tenepb, IIOCJle BblHCHeHIIH cy6MHKpocEonuqecElix
CTPYRTYP aTOll 06J1aCTll?
fJlanHOe BHIIMaHlIe :ureRTpoHHblX MllRpOCROIIHCTOB y;ZJ;eJlHeTCH
Ol1;eHEe HepnHblX OROHqaHlIll Ha RJleTEax. BblHcHHeTcH oonpoc, 1Ip1I-
na;ZJ;JlemaT JIll OHII al}>«jJepeHTHblM IIPOBOD;HlIKaM, HBJlHIO'TCJI JlII qyB-
CTBIITeJlbHblMll (II, cJle;ZJ;onaTeJlbHO, OTBeqarom;IlMII rJlanHOH, no cOBpe-
MeHHblM B033peHHHM, KPYHRI1;IIII EJleTOE - pen;eITTOpHOH) IIHH EJleTRII
RapoTlJ,Il;Horo naparaHI'JlIIH o6J1a;ZJ;aroT ;ZJ;BOllHOH IIHHe}ffian;IIeH. Ross
167
(1959) o6HapY1Ium TOJIbEO TeCHblM EOHTaET Mem}];y xeMopen;enTOp-
HOH EJleTROM H aECOJleMMOII, HO He nu}];eJl nY3bIpbRon, xapaRTepHbIX
}];JlH HepnHblX OEOHqaHIIU, II CTPYETYPHOU Cnen;IIaJlII:3an;nII MeM6paH
F MeCTe EOHTaETa. B pa60Tax Biscoe, Stehbens (1966), Hoglund
(1967), Reis, Fuxe (1968), Abraham (1968, 1970) o6HapymeHa
}];nOMHaH IIHHepnan;IIH I\apOTU}];HOrO IIaparaHrJlIIH If onIIcaHbI EOHTaET-
HbIe CTPYRTypbl, EOTopbIe Y}];OBJleTBOpHIOT conpeMeHHbIM EplITeplIHM
IIpeCHHanTlfqeCEnX :meMeIlTOB, npIlHa]J}1811{aIIJ;IIX 3.p.pepeHTHbIM BO-
JlOEHaM (Palay, 1956, 1958; de Robertis, 1958; Taxi, 1965; Gray,
Guillery, 1966). Ha 3J1eETpoHHorpaMMax y Biscoe, Stehbens (1966)
IIOEa3aHO, qTO E MeM6paHe EJleTOE I TIIIIa nplIMbIRaIOT HepnHbIe OEOH-
qaHIIJI, EOTopbIe CO}];epmaT MeJllme IIY3bIPblUl, MIITOXOH}];pmi II He-
60JlblllOe EOJlIIqeCTnO Be3IIEYJl C 3J1eETpOHHOnJlOTHOH Cep}];u;eBIIHoM.
I1J1a3MaTll'IeCEUe MeJ\I6paHbI HepBHbIx OEOHqaHIIM II EJleTOE I Tuna
pa3}];eJleHbl lIJ;eJlbIO B 100-200 A II IIMeIOT Cnen;HaJIII3IIpOBaHHbIe
3J1eRTpOHHOnJlOTHbIe 30HbI. Ee3MHEoTHbIe HepBHbIe BOJIOEHa BHYTPII
}];OJIeE Tome qaCTO CO}];epmaT MIIl\pOBe3IIEYJIbI, MIITOXOH}];PIIII II HeMHO-
ro OCMIIO.pIIJlbHblX IIY3b1pbEOB.
RaE Biscoe II Stehbens (1966), TaE II Hoglund (1967) y}];e-
JIHIOT 60JlblllOe :BHII-MaHUe cTpYETypaoM, EOTopbIe Ha OCHOnaH1IU nbl-
llIenpl:liBe}];eHHbIX pa60T Palay II Gray C coanTopaMII He MOryT lIC-
TOJIEOBaTb JIHaqe, qeM 3.p.pepeHTHbIe CIiHanCbI. Mop.pOJlOrIIqeCliOe
YCTPOMCTBO IIX ronopIIT 0 TOM, 1JTO XHMIIqOCEaH nepe}];a1Ja ]:(OJlIHIIa
II}];TII OT Hepnnoro OEOH1JaHIIJI E Cnen;I1.pII1Je'C1\I1M EJIernaM RapOTII]:(-
Horo TlIna, II qTO, CJIe}];OBaTeJIbHO, RepBHble OEOHqaHIIH HBJlJlIOTCH
IIpeCIIHaIITlIqe I CEHMII, 3w.pepeHTHblMli [10 OTHOllleHIIIO E EJleTE3M.
MHTepec anTOpOB E 3TIIM CTpYKTypaM HBJIHeTCH BnOJIHe eCTOOTneH-
HbIM, TaE EaE 'Bce OHIf CTOHT Ha II03nn:IIHX xeMopen;elITOpHOM .pYHE-
u;IIH EapOTH}];HOro TeJla II ]:(JlH HIIX fJlanHbIM HnJlHe'l'CH BbIHCHeHIIe
a.p.pepeHTHOM IIHHepnan;Im opraHa. B EaqeCTSe }];OEaaaTeJlbCTBa npn-
CY'I1CTBIIH nOCJle HeM B iKapOTII}];HOM TeJle Ha EJlernaX I TIIna Bis-
coe II Stehbcns CCbIJIaIOTCH Ha pa60TY Gray II Guillery (196£),
B EOTOpOM e-CTb yRa.aaHIIH Ha CHHaIITIIqeCEne nY3bIpbEJI II B a.p-
.pepeHTHblX HepBHblX OEOH1JaHIIHX, a Ross (1959) II Hoglund (1967)
npe}];nOJIaraIOT }];BoiiHyIO IIHHepBan;IIIO HX HaIIO}];o6IIe EJIeTOE nnY-
COBbiX coco'IKon. PaccMaTplIBaH EJleTEII HapOTII}];HOrO TeJIa EaE pe-
n;enTopHbIe, Ross cpaBHIInaeT }];eTaJUl 'cTpoeHIIH HepBHblX OEOH1JaHIIH
Ha HIIX C TaEonbiMII B }];pyrHX ceHcopHblX opraHax (Engstrom, 1958),
f}];e aBTopbI TaE me He HaXO}];IlJlII n EapOTII}];IIOM TeJle cnen;II.pII1Je-
CEIIX CTPYETYP n YTIaCTEaX EOHTaRTon aECOHa C n;lITOnJla3MOM pe-
n;enTopHbIX EJleTOE. RpoMe TOro, OCblJIaHCb Ha TO, 1JTO B xeMope-
u;enTopHblX EJleTKaX HaH}];eHa HeCIIen;II.plIqeCEaH ac.Tepa3a II an;eTIIJI-
XOJIJlH3CTepaaa (Koelle, 1950), Ross npe}];nOJIaraeT, qTO IIMeIOIIJ;I1e-
CH n rJlOMYCHbIX EJleTEaX nyabIpbRn CO}];epmaT an;eTIIJlXOJlIIH II HB-
JlHIOTCH CIIHanTIITIeCEIIMII IIY3bIpbEaMlI, EOTopbIe, nbI}];eJlHHCb I1il
EJleTEII, Boa6ym}];aIOT npOXO}];HlIJ;lIe MIiMO aECOHbI.
B nOCJle}];YIOIIJ;IIX 3EcneplIMeHTaJlbHblX JIC'CJle}];OnaHIIHX Biscoe c
coanTopaMII (1967, 1969) np06J1eMa }];BOHHOii lIHHepnan;I111 RJleTOK
168
RapO'TII;IJ;HOrO' naparaHrJIlfR O'KO'H1JaTeJIbHO' ,pellieHa, Ha 'Halll B!3rm:r;n;,
B IIO'JIO'mllTeJIbHO'M CMblCJIe. DocJIe lIepepe3KII Cll'H}'1CHOrO' HepBa a'B-
TOpbl nO';rry1JIWII gereHepaTIIBHble 1I3MeHeHIIH 'B 'HepBHblX OKO'H<IaHlIJIX
Ha «rJIO'MYCHblX)} RJIeTKaX, AO'Ka3a l B lIX 1JYBiCTBIITeJIbHYIO !II'PHp'OJJ;Y.
)J;aJIee O'HII nO'BTO'pIIJIII O'nbiT de CastrO' C HHYTplI1JepenHO'M lIepe-
pe3KO'M H3LTKO'-rJIQTQqHOrQ HepBa (BiscQe, Lall, SampsQn, 1969)
II TO'me Q6Ha.pym1IJIII ;n;ereHepan;1I1O CIfHaneO'B Ha RJIeTKaX I TllIlIa,
IIQ;n;Q6HYIO TaKOBQM nOCJIe lIepepe3KII ClrnYCHO'rQ HepBa. 3TIIMII OlIbI-
TaMII ]:(QKa3aHQ, <ITO' nep-BHble QROH'IaHIIH HBJIHIOTCH 3<p epeHTHbIMII,
a HeMpoHbI, KOTQPbIM QHII .npIIHap;JIemaT, HaXQ;n;H'TCH B CTBQJIe M03ra
II H'BJIHIOTCH napaCIIMlIaT:H"'IeCI<IIMII. B 3 CTIepIIMeHTe iC y;n;aJIeHIIe'M
BepXHerQ mei'mQrQ Y3JIa BiscO'e II Stehbens (1967) TO'me !IIQJIy<IIIJIII
p;ereHepan;IIIO CIIHaIICOB, HQ IIQCJIep;HlIe 6b1JIII pacnQJIQmeHbI He Ha ImeT-
Kax I TIIIIa, a B IlIBaHHQBCKIIX 3JIeMeHTax, qaCTQ QKQJIO 8H;n;QTe-
JIlfR, II IIQKa3aJIII 3TIIMII OlIbiTaMIf a<PrPepeHTHYIO IIHHep'Ba:1I;1f1O opra-
na nQCTraHrJIHO'napHblMII CIIMlIaTIIqeORIfMII BOJIOKHaMII. nO'MlIMQ AO-
Ra3aTe.lIbeTBa HaJIIf1JIIH 3<p<pepeHTHO'MII a<p<pepeHTHOM IIHHepBa!l1;Im
RapQTII)J;H'Oro lIaparanrJIIIH anI IICCJIe}];O'BaHlfR YC'TanOBIIJIII pa3'HYIO
JIQKaJIII'3an;IIIO 3«p-<pepeHTHbiX HepBHblX ORQnqaHIIM: rrapaCII1M.na:TIIqe-
CKIiX - Ha IIQBepXHOCTII «rJlQMY'CHQM)} (.nO' HallieMY \3HAQRpmrnQM IIa-
paraHrJIHQHapHQM) KJIeTlm, a 'CIIMII8''I'H"'Ie'CRIIX - 'B Q PymalOIII;IIx ee
rJIIIaJIbHblX II coeAIIHIITeJIbHQTKaHHblX 8JIeMeHTax. 3Haqe1Ilfe \3TOrQ
<paKTa pa3QUpaHO HaMII HIIme, I'p;e AeJIaIOT'CH ,BbrBOP;bI 0 3aKQHOMep-
HOCTHX IIllHepBan;un 'CIIMlIaTQ-ap;peHaJlO'BQM o(JHCTeMbl C n03IIn;IIM ee
lICTOpnqeCKO'rO' pa3BIITIIH (CM. CTp-, 179-181).
B <p1I3I1O'JIOrIIqeCKIIX pa6QTaX IIQCJIep;HerQ BpeMeHH, OTHQCHIII;IIX-
CH K IIpO'6JIeMe <PYHKII;IIO'naJIbHO'M pOJIII KapQTIIp;HO'rQ «rJIO'Myca»
(Eyz.aguirre, KO'yanO', 1965 a, b, c), aBTQPbl npep;nQJIQmIIJIII, qTO
13 xeMO'pen;eIIII;IIII KapQTIIp;BO'rO' TeJIa «rJIO'MYCHble)} KJIeTKII nrpalO T
PQJIb IIpeCIIHaIITH"'IeCKQrO' 3JIeMeHTa, a qyncTBIITeJIbHble HepBHble
'QKO'HqaHIIH - IIO'CTCIIHaIITIIqeCKO'rO', II qTO' BeIlI;eCTBa, BbiAeJIHeMbIe II3
«rJIO'MYCHblX)} KJIeTO'K BQ BpeMH pa3p;pameHIIH, MQryT Bbl3b1BaTb xeMO-
pen;eIITO'pHble IIMIIYJIbC:bI. IIpII pa3HblX BIIp;ax pa3p;pameHIIH Bblp;eJIJI-
IOTCH pa3Hble BeIlI;eCTBa, B qaCTHO'CTII, an;eTIIJIXO'JIlfH, KO'TO'PblM QHII
HaIlIJIII B TKaHHX KapOTIIp;HO'rQ TeJIa (Eyzaguirre, KO'yanQ, TaylQr,
1965) .
OTHO'CIITeJIbHO' QrpaHlI1JeHIIH <pII3I1O'JIQrlI1JeCKO'rQ p;eMcTBIIH Be-
TII;eCTB, .Bblp;eJIHeMbIX 1I3 KapO'TIIp;HbIX KJIeTO'K, MeCTHbiM 3HaqeHIIeM
B Ka1JeCTBe <paKTQPO'B, pa3p;pamalOIII;IIX HepBHble OKO'uqaHIIH TO'JIbKO
3TO'ro O'praHa, Mbl yme BblCKa3aJIII CBQe O'TplIlI;aTeJIbHO'e O'THQrneHlIe
K 3TO'MY MHeHIIIO. B pa6O'Te Bliilllcke C cO'aBTO'paMII (1967) 3Ta TQq-
Ka 3peHIIJI HaxO'p;IIT XQPQllIee MO'p<pO'JIO'rlI1JeCKO'e II rIICTO'JIO'rlI1JeCKOe
IIO']:(KpeIIJIeHlIe. 8KcIIepIIMeHTaJIbHO' Bbl3BaHHO'e KIICJIO'PQp;HO'e rO'JIO']:(a-
HlIe y 6eJIbiX KpblC CQIIPQBQmp;aeTCH Bblp;eJIeHIIeM KaTeXO'JIaMIIHQB II3
KapO'TIIAHO'rQ TeJIa: C IIO'MO'III;blO <pJIyO'peCn;eHn;IIII QUHapymIIBaeTcH p;e-
.<pIIn;IIT, a 3aTeM IIO'qTII IIO'JIHO'e IICqe3HO'BeHIIe KaTeXO'JIaMIIHO'B II3
KJIeTO'K, B TO' BpeMH KaK B KPO'BII YBeJIIIqIIBaeTCH KO'J1lI1JeCTBQ ap;pe-
HaJIIIHa II HO'pap;peHaJIIIHa: 3JIeKTpOHHaJl MIIRpO'CKQnIIH IIO'Ka3b1BaeT
. 169
npII aTOM Bce CTa,D;IIH IIX BbI]:(eJIeHIIJI - p;BlImeHIIe rpaHYJI E neplI-
tjleplIlI EJIeTElI, CJIIIJIHlIe MeM6paHbl rpaHYJI C MeM6paHOM EJIeTEII.
npopblB MeM6paHbI II Bblp;eJIeHlIe :meETpOHHOIIJIOTHOrO cop;epmIIMoro
rpaHYJI B pacmIIpeHHoe MemEJIeTOqHOe npOCTpaHCTBO. HapHP;Y C 3TIIM
B EJIeTEax Ha6JIIO]:(aeTcH Ha6yxaHIIe MIITOXOHP;pIIM, paCillIIpeHIIe 1I0p B
Jlp;epHOM MeM6paHe II paclIap; xpOMaTIIHa. 113BecTHo, qTO Bce 3TH
EapTIIHbl xapaETepHbI II P;JIH EJIeTOE M03rOBoro CJIOJl Hap;rrOqeqHII-
EOB B lIeplIOp; aETIIBHoro Bblp;eJIeHIIH ropMOHOB (IIpIIMMaE, 1965a, 6,
1966).
Hap;o OTMeTHTb IIpaBIIJIbHYIO, Ha Ham B3rJIJlp;, o eHEY (Bliim-
cke C coaBTopaMII, 1967) EaE c06cTBeHHblx, TaE II JIIITeparypHblx
p;aHHblX. OHII IIOJIaraIOT, qTO EaTeXOJIaMIIHbI B YCJIOBIIJIX OlIbiTa BbI-
p;eJIHIOTCH He TOJIbEO II3 «pel\eIITOpHblX) EJIeTOE EapOTIIp;Horo TeJIa,
a II3 BceX naparaHrJIIIeB, B TOM qlICJIe II Hap;cep]:(eqHblX II C 0p;06-
peHlIeM OTHOCHTCH E pe3YJIbTaTaM YIIOMHHYTblX Bblme IJ!II3IIOJIOrlf1Ie-
CEIIX OlIbiTOB Eyzaguirre, Koyano, ROTopble II3MeHHJIII 3JIeETpIIqe-
Clrn:e pa3pJlAbi CIIHYCHoro HepBa IIOCJIe aIIIIJIIIEal\IIII ap;peHaJIIIHa, HOp-
ap;peHaJIIIHa II cepOTOHIiHa II He 06HapymlIJIH yqam;eHIIH pa3pHp;OB.
Bliimcke C coaBTopaMII H8. 3TOM OCHOBaHlI1I cnpaBep;JIIIBO YTBepm-
p;aIOT, qTO pelJ!JIeETOpHaH P;yra, COep;IIHHIOm;aH «pel\enTopHYIO) EJIeT-
EY qepe3 p;bIXaTeJIbHblM II Ba30MoTopHblM l\eHTpbl II lIepn:lJ!epH"'le-
CKHe HepBbI C p;blXaTeJIbHon: MYCEYJIaTYPOH, He nymp;aIOTcH B EaTe-
XOJIaMlIHaX B EaqeCTBe nocpep;HimOB Memp;y pel\enTOpHblMII RJIeT-
EaMII II OEOHQaHlIHMII ClIlIYCHoro HepBa.
OOBOEyilIHOCTb Bcex OBep;eHIIi'I, fiOTOpblMH Mbi palOIIOJIaraeM B Ha-
CTOHm;ee BpeMH 0 CTpYEType, JIemaIII;eH B pa3BIIJIEe COHHOn: apTepnll
MJIeEOIIIITaIOm;IIX, (pIle. 66) 1I03BOJIHeT 'CEa3aTb CJIep;YIOIII;ee. B co-
C'TaB 3TOro ysemm BXOp;JIT RJIeTKII, EOTopbIe CIIHTe3IIpYIOT II BbI]:(e-
JIHIOT aTeXOJIaMIIHbI, IIHHepBIIpYI01'CH qYBC11BHTeJIbHbIMH II alJ!lJ!e-
pOOTHbiMII npII6opaifolH, OCYIII;eCTBJIHIOIII;lIMII pelJ!mmTOpHoe Bblp;eJIeHII8"
EaTeXOJIaMlIHOB, T. e. IJ!YHEqIIOHIIpYIOT TaE me, EaE MO'3rOBOM CJIOR
Ha,D;nOqeqRIIKOB. MHblMII CJIOBaMII, B'ce crmpeMeH'Hbie IICCJIe.n;onaHIiH
,n;alOT HaM B pyRII lJ!aETbI, Ha OCHOBaHlIII EOTOpblX p;OJImHO IIplI3HaTh
3a a'l10M CTpYETYPOM 3H]J;OEpIIHHble CBoikl'Ba. Op;HaEO HapMY CO CXO]:(-
CTBOM C H OqeqHIIEOM RapOTIIp;HblH lIaparaHrJIIIM 06JIap;aeT II He-
E01'OpblMIi oco6eHHOCTJlMII. OH IIpep;CTaBJIHeT C060M pa3p;eJIeHHbln: H3
P;OJIbEII Y3eJIOE MeJIEIIX EJIeTOE, EOTopbIe EaE Obi 3aMypOBaHbI B
EPYIIHble BOJIOlOInCTble CTPYETypbl eoe]:(HHIITeJIbHOH TEaHII. 06'beM
BOJIOEHIICThlX ilIpOCJIOeE II IWJIlf1IeCTBO OIIJIeTaeMblX HMII MeTOE nop;-
lIepmeHbI pe3EHM BII,D;OBbiM EOJIe6aHIIHM (pIIC. 67, A, B, B, F). KJIeT-
Ell cop;epmaT 'MaJIO ropMOHOB II :B 60JIee MeJIRllX IIO cpaBHeHIIIO C
M03rOBMM CJIOeM Hap;IIOq,eqHIIEOB OCMIIOIJ!IIJIbHblX rpaHYJIaX (IIO p;aH-
HbiM Lever et a!., 1959, pa3Mep HX B EapOTIIp;HOM TeJIe paBeH
0,05-0,15, oR M03roBOM CJIoe Hafl/lIOqeqHItEOB - 0,05-0,4 Jltn). aTO
cBoiiicTOO HBJIJleTCJI IIOC'l1OJlHHblM II xapaETepHblM jI];JIH EapoTlIp;HbIX
lIaparaHrJIIIHB Bcex MJIeEOIIIITaIOIII;llX. Pa3HIIu;a Ha6JIIOp;aeTCH TOJIbEO
B CTeIIeHII IJ!JIyopeCl\eHI\IIII, CBOHCT,BeHHoi'1 pa3HblM mll'BOTHbiM II OT-
pamaIOm;eH MHHblllee IIJIII OOJIblllee ROJIlf1IeCTBO 6noreHHblx aMIIHOB
170
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a - HapOTHAHhIt1: napal'aHI'JIHt1:; 6 - HapYIHlIaH COHHaH apTepHH; 8 - BHYTpeHHHR COHHaR
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B 1,."1eTK3X. )].pyrlIl\I CTOJIb me xapaI\TCpHbIM II CTOJlb me nOCTOHH-
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BCIITIII.(IID.JlbHOii 060JIO'me 3pTepIlll, T. e. Bp;amI OT npOCBeTa c-oey]:(a
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c IlCHrp3Jlbnbll\l KpOBOTOKml IJ COHIlOii apTepIIII, no, "3K IIOHa33JIH
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CBCTOB K3nUJIJIHpnOrO PYC.'13 TOJlCTbIMII npOCJIOllKaMII OOep;I1HIITeJlb-
BOTI-\iUIHblX :JJIeIlIC!lTOB, ua3a.'1LIlL1MII MeMOp31Ialllll, IIIB3HIIOBClmMU
R."1CTI,aMII II MHHOTHbIMII II Ue3l1UIIWTHbIMII HepHHblMII BOJIOI,H8?1111.
3TII CTPYKTYPbl He TOJILKO lIe cnoc06CTHYIOT BbIllO::lHeHIIIO «rJIOlI!ye-
HbDIIn) H.'IeTI\aMII .pYHKI.(1II1 pC elI I1I1 XII nI'reCKnX cp;BnrOB, npone-
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HoiL B aTOM OTIIOIliCHIIII na;.w nplI3HaTb CllpaBe]:(:IIIBOCTb COlllHeHUH,
BbICJ-\a:J8HTII,IX de Kock II Dunn (1964). TaKaH CTpYKTypa, 0P;H3KO,
Ma.l.O Y;:I.OBJIeTBOpHeT II Tell! TpcuonaHIUIM, IWTOpbIe nYllilIbl ]:(JlH oey-
IllCl'" II:ICHIIH :mp;OI,pIIIIIIoii (PYHI{ IIII. II\C.'IC3bl. :IlIllIeHlIbIe npOTOKOB
II nI,I;J.e.'IHIOw.ne npO;:J.Y"Tbl JKII:mcp;CfiTC.'IbHOCTII Henocpe]:(CTBeHHO B
1'1)0111>, I1ya';;:J.3IOTCH B Tom,OCn'lIHbIX CTPYI';Typax, OTp;eJlHIOIllllX II-
'TOn."111:J1II)" i(';C.'Ie::!lICTbIX WleTOK OT I,POBOTOK3.
n plI Jlo):lxop;e 1\: II:!Y'ICIIIIIO CIICTCMbl n<1p8raHrJlIleB MJIClWIlIIT3IO-
I II,( C I1CTOpIPIeCInIX n03lUlllii, npn y'IeTe UX B33IIMOOTHOllleHlIll e
17f
KpoBeHOCHbI'M PYCJlOM Ha pa:3HblX ::nanax 3BO'JlIOn;HH 'Bce CBO'HCTBeH-
HbIe MaKPO'- II MllKPOCKOIIHqe<:KHe qepTbi CTPO'eHIIH KapO'TH]:(HO'ro 118-
paraHrJlHH Hera rnCTaXHMHqeCKHe C'BO'llCTBa JlerKO HaXO HT 06'bJlC-
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TO' 'KapO'TII,IJ;Hblll naparaHrJIIIH xapan:TeplI3yeTCJI aTpoiPuen II 8a-
nYCTeBaHIIeM .ceKpeTOpHblX ,meMeHTOB, IIPO'paCTaHueM CO'ep;IIHIITeJlb-
Hon TI\aHblO (CM. pIIC. 67) H paCIIO'JlO'mCmIeM 13 HapymHO'H 060'-
JlO'qKe TO'n apTepUH, KO'TOpaH KO'rAa-TO' iPYHKI1;IIoHIIPO'BaJla B Kaqe-
CT'Be Tpe'Tben ma6epnO'H yrH.
A qTO MOmHO' CKa:aaTb 0' ]:(PyrlIx T8KHX me o6pa3orIJaHIIHX, KD-
TO'Pble mnpOKO' paCIIpOCTpaHeHbl B O'Jlb JlerO'1JHDH apT{ pUII, yrH aDp-
TbI, III YIliKax cep]:(ua, YCTbe KopDHapHblX apTepIIH II KO'TDpble-
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TIIJJ;HYIO II Kap]:(Ho-aOpTaJlbHYIO peiPJleKCareBHble 30HbI? .
IIpII paCCMO'TpeHlI1I pa6O'T 110' XPO'MaiPiPlIHHO'll CIICTeMe MJleKO'IIII-
TalOm;IIX 'B XPO'HDJlOrlI1JCCKOM IIDpHJJ;Ke 6pOCae'l'CH B rJla3a cJlep;ylO-
m;ee D6cTaJlTeJlb'CTBO. B Han6DJlee CTapblX pa60Tax naparaHrJlIIII
oc06eHHo IIpIIcep]:(e'IfHOH O'6J1aCTII, paOCMaTpHBalOTCH aBTopaMII KaK
aH]:(OKplIHHble meJle3b1 (KO'hn, 1903; Wiesel, 1906; Busacchi
1912-1913, II JUl.). HaqlIHaH C 30-x rO'fJ:O'B, HHTepec R HRM pe3Ko
BO'apa.cTaeT II nOHBJlHeTCH MHDro ny6J1UKa.:Qun, 13 KO'TDPblX aBTopb]
C nplICTpaCTIIeM u3yqalOT MopiPDJlOrHqeCKUe oc06eHHocTII aTIIX me.
CTpYKTYP II, cpaBHHBaH C KapDTHAHblM rraparanrJlHeM, TpaKTYIOT IIX
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m;en, a He BbI]:(eJlHTeJlbHDn iPYHKI1;lIll naparaHrJIlIOHapHblX KJleTDK.
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paraHrJlHO'HapHblX KJleTOK MelliP;Y ]:(YI'OH aopTbI II JleI'OqHDH apTepllen,
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O'6pa30BaHIIH: paragangliO'n aO'rticum supracardiale ,11 paragang-
liO'n aO'rticum infracardiale II Memp;y HIIMH O']:(IIH HJlII HeCKO'Jlb-
KO MeJlKIIX.
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(B), Ilec,-\a (B), l\pOJIlma (F)
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apTepIIII TJ,a ee pa3BeTBJleHIIH Ha npaBYK) II JleBYIO. MHaraqlICJleHHble
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apTeplIaJIbHblX 'cacy]:(aB I1PY]:(HoOII 'lJaCTII TYJfOBiHrn;a. Penitschka, Pal-
me, Seto HalllJlII cxa]:(CTBO IIX C RapaTIITJ,HblM IIaparaHrJIlIeM, II Ram-
TJ,bIH II3 aBTapaB IIa cBaeMy BbICRa3an:CJI 00 HX WYRKQIIH. Ha OCHOBa-
HMH IcpaameHIIJI ca oC,TPYIRTypaH RapaTlI\!I;HOroO TeJla, C HexpaMa4>WHH-
HOCTbIO era RJleTOR, tOaCY]:(IICTblM C'Ha6meHHeM II paCtIIoOJlameHlIeM B
06J1a!C'I1H OROH'IaHIIH AeIIpec,capHoOra HepBa, a'HaJlar1'!lfH'O ]:(enpe,ccopy
RaipaTH]:(HoOra TeJla (de Castra), Palma npeJ{TIOJlamIIJI, 1fTa cyrrrpa-
Rap IIaJlbHhIe ilIapal'aHrJIHII MoOryT IIrpaTb paJib perYJlJlTapoB RpOBJI-
Haro TJ,a'BJleHHJI. Op;HaRa aH naTJ,'IepRHYJI npH 3TOM, qTO TaRae cym-
]:(6HHe TI.oOJlmHa pa'CCMaTplInaTb'CJI He 6aJIbIDe, RaR nmIlaTe:3a. B TO me
BpeMH Palme C'OB6pllleHHa npaBHJlbHO paCCMaTplrnaJi JlORaJlH3aQIIIO
lIa,paraHrJlHeB '13TJ,aJIIi aT rrpOCHeTa cocy]:(a (aH Haxa]:(HJI ilIX He TaJlbRO
B tunica adveniicia, Hoo IIHorTJ,a H B tunica llIedia), RaR IIPH-
3HaR, raBapmD;IIH npOTIIIn IIX HHRpe'TapHaH WYHKQHH. OH IIpeAlloJla-
I'a.rr !MeCTHae BJlIIHHiHe BbI]:(eJlJleMbIX RJleTRaH Bern;e'CTB Ha 6J1II3J1ema-
rn;IIe HepBHble BO,lIORHa, a He 'Iepe3 RpaBb. ::ha IIOCJle;:J;HJIJI Ta'IRa
3peHIIJI, pacnpocTpaHeHHaJl B HaCTaJlrn;ee BpeMH, He UMeJla, aJ{HaRO,
HmmROH waRTH'I6CRaH OCHOBbi n el'a nOCJlep;OBaHIIJlX.
B 30-e II 40-e ra]:(bl II n03me TY me a6J1aCTb HOCJle]:(OIBaJlII II
IIaJlY'IHJlII RaBble ]:(aHHbIe na pacnpep;eJleHHIO na.paraHrJlHeB, IIX TOH-
RaH QIITaJlarIIII IIHHepBaQHII II BacRYJlJlplI3aQHH Muratori (1933b, c,
1935, 1937, 1959), Nanidez (1935a, b, 1936, 1937b), Pannier
(1935), Gaarmaghtigh. Pannier (1936, 1939), Hallinshead
(1939, 1940a, c), B. B. KynpHHHaB (1954, 1959). EbIJIa alIIIcaHa
M'HameCTBO naparaHrJIHeB B CTBaJle 6J1YiRTJ,aIOrn;era HepBa (Muratari,
1932a, b, c; 1933b, c; de Castra, 1951), Mem]:(y lIa]:(KJlIO'IIIlfHaH ap-
TepneH II 06rn;eH caHHaH cnpaBa H Ha aapTaJlbHooH yre 1B6JI1'!3H
na KJlIO'IWfHaH apTepHH CJleBa (Nanidez, 1935a, b), y OCHOBaHIUI
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RapaHapHaH apTepIIII y 3apa]:(blllIa JleTY'IeH MblIDII (Celestina
da Casta, 1936), Ha 3a]:(HeH CTeHRe IIpe]:(cep]:(HH H B CepTJ,e'IHbiX Ylll-
flax (Pannier, 1935; Gaarmaghtigh, Pannier, 1939), B meHHaM
CIIMnaTHRyce (de Castra, 1951). Celestina da Casta, RaTapblH
II3Y'IaJl pa.cnpe]:(eJIeHlIe naparaHrJlHeB Y JleTy'IIIX MbillIeH, HallleJi
arpOM'Hble HX mWnJ1eHHH B pa3HbiX Y'IaCTRaX llIeII. ABTapbl IIO-pa3-
HaMY paCQeHll!BaJiM WYHH.QHIO Bcex 311HX a6pa30naHHH. B OCROBe pa3-
HaH aQeHRH JlemaJlII Te me CnapHbIe BanpOCbl, 1\aTOpbIe Ba3HIIRaJlII
B OTHallleHHM RapaT1'!jD;Haro naparaHrJlHJI - xpaMa4>WHHHoCTb II ne-
xpaMaW<pHHuaCTb, xa,paRTep Ba01\YJlHpH3aQ:HH H IIHHepB8QH1M, CBH3II
C 6J1ym]:(aIOrn;IIM HepBOM. I'hfeHHa B 'STaT nepH ,HHT6'HCll!BHOro '06-
CJle]:(aBaH:HH llIeHHa-rpYP;HaH a6J1aCTH B03H1mJlII Te RJla,c.cHWHRaQHM
naparaHrJlIIeB, a IWTapblX raBaplIJlOCb Bblllle.
gCJlH B ilIepBblX pa6aTax aBTapbl 6bIJ1I1 aCTapamHbl B ClBOIIX Bbl-
Bap;ax II HHTepnpeTaQIIM naJlY'IeHHbIX waRTaB II 3aRJlIOqeUHJI IIX He
aTJlM'IaJlI'!'Cb e]:(HHa06:Qa3HeM, TO B IICCJleTJ,aBaHMJlX nOCJleTJ,YIOrn;ero Bpe-
t74
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BOBbi cep]:(e'IHO-COCY IICTOll 30HbI C TOqKII 3peRIIH ee pe.n;eIITOpHOU
!j>YILKn;IIII (Knoche, Schmitt, 1963). 0 B03MomHOCTli 3H,I1;OKpIIH-
HOU tPYHKn;lHI qaCTO aiI(e He yrroMIIHaeTCJI II OCblJlOK Ha COOTBeT-
CTBYIO YIO JlIITepaTYPY He p;eJlaeTCJI. 06cym]:(eHIIH B OCHOBHOM CBO-
JlTCH 'K O]:(HOMY - rroxom JIll TOT 1IJ11l IIHOU rraparaHrJllIll Ha iKa-
pOTII]:(HbIU IIJlII Ha]:(cepp;e'IHbIU II, CJle OBaTeJlbHO, HlBJlJleTCH JIll OH
pen;eIITOpHblM opraHOM IIJIII HeT. A caMII RapOTII;l];Hble II Ha]:(ce,p-
p;eqHbIe naparaHrJIlI'II B xo]:(e ::JTnx IICCJle]:(oBaHlIu rrpeBpaTIIJlIICb KaK
6bI B 600c.llopHble pen;ellTopHble CTPYKTypbl. Y 60JlbmlIBCTBa aBTopoB
aa 3TOT 'Cqe'l HllKaKIiX 'COMHeHIIU yme He B03HlIKaeT.
B HaCT()JI ee BpeMll B -cynpaKap]:(IIaJlbHOM npOCTpaHC"l1Be H3Be-
CTHO 6e,CqIliCJleHHOe MHomeCTBO Mem\HX lIaparaHrJIlieB (]:(o 56 rro
Blessing, 1963), IIHor a OOCTOH IIX II3 HeCKOJlbKIIX KJleTOR II qa-
cro H8'3b1BaeMbiX MIIKporraparaHrJIlIHMH (RYIIPIIHHOB, 1959; Murato-
plI, 1964, 1968) (pilC. 68, A, B, B). Y pa3HbIX mlIiBOTBblX OHII
pa3Ho06pa3HbI no 'pacrrOJlOmCHlIIO, KOJlllqeCTBY II CBH3HM C oKpymaIO-
IIMII TKaHHMII. By]:(yqu 06'beAIIHeHbl 'C 60Jlee KpyrrHbIMII 'lIaparaHr-
JlIIJIMII, OHII yJIn!j>nn;IIpYIOT'CH B !j>YHKn;noHaJlbHoM OTHOllleHlI1I KaK
opraHbl clIHo-KapoTII]:(HbiX II Kap]:(lIo-aOpTaJlbHblX 3GH, yqaCTBYIOII:\IIX
B ocy ecTBJleHIIII peWJlel(COreHHblx !j>YHKn;IIU (Heymans, Neil,
1958; KYIIpnJlHOB, 1959). Mop!j>oJlOrn'IeCKlle IIOCJle]:(OBaHIIH Nonidez
(1935a, 1936, 1937b), K0110pblll OITIICaJl lIIaparaHrJlIIII IB 6m}>ypKa-
.IJ;IIII 6e3b1MJlHHOll apTep:m, y OCHOBaUIIJI rrOp;KJlIOqlIqHOll, Ha]:( YCTbeM
60TaJlJlOBa rrpoTOKa II B JlerOqHOU apTepIIII (opraH Penitschka),
crrocOOcTBOBaJlII TaKOMY 06'be IfHBHMIO. Nonidez HallleJl '8 aOp'laJlb-
HOM II JlerO'IHOM «rJloMyce» i[l;ua TlIIIa KJleTOK, II 06HblX 'reM, 1\0-
TOpble orrlIcaHbl B KapOTII]:(HOM 'leJle: OAHII KpYUHble, 06J1a]:(aIOlIJ;lIe
xpOMa !j>HlI'HOll 3epHIIC'fOC'lbIO, ]:(pYTlIe MeJlKMe - HexpoMa!jKpIlHHble.
llHTepeooo OTMBTIITb, qTO CIDdrraTuqeCKYK> nHHepBan;IIIO «rJlOMYCOB»
Nonidez paCCMaTpHBaeT KaK IIHHepBan;IIIO COCYP;IICTOll CTeHKlI,
a cHa6meHIIe qYBCTBIITeJlbHLlMII OROHqaHIIJIMII II3 p;errpeccopHoro Hep-
Ba pa3J1l1qaeT Ha TOM TOJlbKO OCHOBaHIIII, qTO O]:(HII US HUX (xe:uo-
pen;ellI'lopbl) IIMeIOT B CBoeM COCTaBe «rJloMycHble» KJleTfftIl, a p;py-
rile (npeccopen;errTOpbl) «rJliOMYCHbIX» KJleTOK He IIMeIOT II OTJlII'IaIOT-
CJI no CBoell CTpY!\Type. B OTBeT 'Ha IIHTeprrpeTan;IIIO Nonidez Mb]
MomeM CKa3aTb, 'ITO rro]:(o6noe ]:(eJleHne Ha xeMO- II IIIpoocopen;err-
TOpbl MOmHiQ 6bIJIO 6bl IIpnHJlTb JllIllIb B TOM cJlyqae, ecJlII 6bI Ta-
KaJl Mop!j>OJlOrIIH (namI'IlIe «rJlOMY-COB» B COOTBeTCTBYIOIIJ;IIX M€-C'lax}
Bcer a ICOBilIa aJIa 6bl C WII3I10JlOrlI'IOCRH o6HapymeHHblMH xeMope-
n;elITopHblMn 30HaMD, qe1'O, O]:(HaKO, Ha CaM()M ;n:eJle HeT.
B OqeHb no]:(po6HOM CpaBHIITeJlbHOM I1c,cJleAOBaHIIII y qeJlOBeKa
II 1I03BOHO'IHbiX Muratori (1959) paCCMaTplIBaeT I'IICT()tPII3IIOJlOrIIIO
IIpeooope enTopOB II, RaK OH Ha3blBaeT, «IIaparaHrJlIIOIIo]:(06HYIO xe-
Mopen;errTOpnyIO TRaHb» KapOTlI)J;HbIX, ITO]:(RJlIOqnqHbIX II aOpTaJlbHO-
JlerOqHblX «npn a'l1KOB». OH rrOBTOpJle'I', qTO "lYBCTB1ITeJlb'Hble OKOH-
qaHIIH HepBa Hering - de Castro (BeTBII Jl3b1KO-fJIOTOqHOro)
B CTeHKax KapoTII]:(HOii apTepIIII II aHaJlOrlIqHbIe qYBCTBBTeJlbHlde
175
5
p u c. 68. CxeMId (A, B, B)
pacnpep,eJIeBJIH nepu6pOHXnaJIb-
HbIX MnRponaparaHrJIneB (Mu-
ratori, 1968)
A
B
aKQHqaHIIJI HepBa Ludwig-CYDn (BeTBII 6JIym]:(aIOm;erQ) CTHMYJIlIpy-
IOTCJI ]:(aBJIeHlIeM KPQBH (6apo- IIJIH IIpeCCDpen;enTDpbl) H BJIlIHIOT Ha
cepp;e'fHo-cocY]:(l1'CTble pCljmeiKcbI. Te me HepBbI BeTBJI'l'CJI II ]:(aIOT
pen;enTOIpbl B Tex me IIpeoCapen;elITapHblx 3OHax, nO' Ha CTpyKTY-
pax, ROTapbIe IIa]:(Q6HbI naparaHrJIIIJlM II qynCTBIITCJIbHbI K XIIMIIqe-
cKOMy ;cocTaBY KPQBM. Pa30IIpaH B cBaen iCTaTbe Ij>II3IIOJIarIIqeCKIIe
31j>Ij>eKTbI C KapQTHJ];HbIX II aapTaJIbHblX HepBDB, Muratari aqeHb
Rep'HO an;eRlIIBaeT IlIIIPQTY IIX ]:(eifCTBMH: aH Ila]:(QepKlfBaeT, QTO DHR He
DrpaH'IIQII'BaIOTCH Cepp;eqHa-COCY]:(IICTblMII II pecnnpaTapHblMII. 'II, OCbI-
JIaaCb na pa60TY Heymans, Bauckaert, Reguiers (1933), YKa3b1-
BaeT Ha CBJIIIHHlIe KapQTII;IJ;HblX pelj>JIeKCOB Ra ceKpen;IIIO ap;penaJIIlHa
Ma31'0IBblM CJIaeM H8AIIDQeqHIIKOB. Mem]:(y TeM IIQ3me, Kar]:(a 6b1J1
pa3pa6aTaH Ij>JIyapecn;eHTHbln MeTa]:( «I>aJIbKa II KQrp;a B KapaTIIp;-
HbIX II JIerDQHD-aapTaJlbHblx naparaHrJIIIHX 6b1JIII o6HapymeHbI ManO'-
aMIIHbI, '1101 HX pelj>JleRTO'PHQe BbI]:(eJIeHIIe IIpM pa3]:(pameHIIII YKa'3aH-
HbiX 3an CTaJIII TpaKTOBaTb B RaqecTBe Bem;eCTB, ]:(eiicTRYIOm;IIX JIa-
KaJIbHO Ha HepB'HbIe aKOH'IaHIIJI 3TIIX me CTPYKTYp. CJIep;OBaTeJlbHO',
p;JIJI 'CTOpoHHIIHaB xeMapen;euTapHaii Ij>YHKn;IIM lIaparaHrJIIIeB aTKpbl-
THe !B nux MQHO'aMIIHaB CJIymlIT He ]:(aBO']:(QM B lIaJlb3Y HHp;aKpIIH-
Bon era WYHKn;IIII, a Tpa.KTyeTcH B KaqeCTBe lj>aKTapa, Y'IaCTBYIO-
m;el'O B MexaHII3Me Me'CTHaro pen;elITO'PHarO' pelj>JleKca. 3a6bIBaIOTCH
p;ame Te lj>aKTbl, KQTQPble 6blJIlI OTKpblTbi OCHQBOIIO'JIamHIIRaM'II xe-
Mopen;enTOpHQra 3HaQeHIIH KapOTJI]:(HblX II Kap]:(IIQaOpTaJIbHblX 3an.
R QHCJlY TaKIIX 3a6bITbiX lj>aKTO'B, aQeBII)J;HO, Ha]:(a aTneCTII II pelj>-
JIeKTOpHO'e BbI]:(eJIeHIIe a]:(peHaJIIlHa Ha]:(IIDTIeqHIIKaMII IIOCJIe pa3)J;pa-
)KeHIIJI KapQTIIp;HblX 3QH, anlI'CaHHOe nIIDHepaMH 'H3yqeHIIJI BTO'ii
176
IIpOOJleMbi (Heymans c coaBTOpat.UI, 1933; Heymans, 1934; Hey-
mans, Neil, 1958).
TaRHM 06pa30M, :mCIIepllMeHTaJlbHbIe ,naHHble y6e HTeJlbHo rro]:(-
TBepm]:(aIOT, qTO pe.pJleRTOpHoe BbI]:(eJleHlIe a]:(pCHamma HacTyrraeT
He 'J10JlbRO ,B iRapOTH]:(HOM rraparaHrJlMII, HO II B H8WIoqe'IHIIRe II,
OqeBH;l];HO, B APyrHx naparaHrJlHHX, HllJlHIOrn;nXCH 3H]:(ORpIIHHbIMM op-
raHaMII, II qTO 06ycJloBJleHo OHO .pn:3IIOJJOrJlqeCHOn rroTpe6HOCTbIO
Bcero opraHII3Ma, a He Hym]:(aMII Tex qYBCTBIITeJlbHbIX ORouqa-
Hnn, nOTOpbIe IIMeIOTCH B RapOTIIAHon 06J1acTII. IIpII 3TOM 3Rcrre-
plIMeHTaJlbHO YCTaHOBJleHO (Eyzaguirre, Koyano, 1965b), qTO alI-
n.'IIIRan;IIJI RaTeXOJlaMIiHOB ITa 06J1aCTb 6IIcJlYPRan;IIII COHHOll apTepnII
He BJlllHeT Ha pe.pJleRTOpHoe yqarn;eHIIe pa3pH]:(a. CJle]:(OBaTeJlbHO,
IIpe]:(nOJlOmeHlIe, qTO Bbl]:(eJlHeMblH RJleTRaMII (aH]:(oreHHbln) HOp-
a]:(peHaJlIIH ]:(eHcTByeT MeCTHO II pa3]:(pamaeT HepBHble OROHqaHIIH 3TO.
ro opraHa, aRcnepllMeHTaJlbHO He IIO]:(TBepm,l];aeTCJI. TeM He MeHee
1\Iuratori (1959), YIIOMHHYB 0 .pe.pJleRTOpHoM BJlIIJIHMII 'IYBC1'BII-
TeJlbHOH RapoTII]:(Hoii: 30Hbl Ha ceRpeTopHYIO .pYHRn;IIIO HaAnOqeqnII-
ROB, He AeJlaeT 113 3Toro HHI,aIHIX IIbl'BO]:(OB. OH, Hao60pOT, npII-
cOeJJ;IIHHeTCH R Teopun Koeh (1931), no I\OTopon RapOTIIAHble II
JleroqnO-aOpTaJlbHble 301lbl, IIOMIIMO 06rn;ei'1 ,D;JlH HIIX XHMopen;enTOp-
HOR .p1I3IIoJlOrlIqeOKOH .pYHl,n;UII, MorYT 6bITb If Mop.pOJlOrlf'IeCRll
06'bep;IIHeHbI MeiR]:(y C060ll Ha TOM OCHOBaHIIII, qTO Bce OHII IIpO-
1I30lllJlII II3 CIICTeMbi ma6epHbiX apTepnaJlbHLIX ]:(yr II, TaR me RaR
3TII IIOCJle]:(Hne, nOJlyqaIOT lIHHepBan;IIIO OT Jl3bIRO-rJlOTOqHOrO IrepBa
(RapOTII]:(HbiR CIIHYC, pa3BIIBllIuiicH II3 III ma6epHoR p;yrlI) II OT
6J1ym]:(aIOrn;ero (RopeHb nO]:(RJlIOqHqHOR apTepHH, ]:(yra aopTbl II 60-
Ta.rrJlOB rrpOTOR, rrpOHCllIe]:(IlIIfe WOTBCTCTBeRHO II3 IV II VI ma6ep-
HbIX AYr). OHU ]:(encTBIITeJlbHO CBH3aHbi 110 npoIlcxom]:(cHIIIO C ma-
6epHblMII AYraMU, HO He Ran xeMopen;errropbl, a RaR 3H]:(OKpIIHHblC
meJle3b1.
B MOHorpa.pHII B. B. HynpIIHHoBa (1959), IIOCBHrn;eHHoll HepB-
HOMY anrrapaTY COCY]:(OB M8J10rO Rpyra RpoBo06parn;eHIIJI, o6lIJlbHaH
«rJlOMYCHaJl» T.RaHb, ROTOPYIO OH OnIICaJl MBeHTIID;IIlI JIerOqHOH
apTepIIII II KOTOpaJl BeCbMa cxo]:(rra .c RapOTn HbIM «rJIOMYWM»,
Tome paccMaTIHlBaeTcH RaR cy6cTpaT xeMopen;eIIn;IIlI. IIop;p06Hble
I\lOp.poJlOrlf'I'OCRIIe p;aHHble HyrrpIIJlHoBa rrORa3bIBaIOT, qTO pJlP;OM C
«rJlOMYCHOll» TRaHbIO .rremaT HepBHbIe RJleTRU II qTO «rJlOMYCbI»
Bc.er]:(a CBH3aHbi C HepBHblMII -ceTHMII, OIIJleTaIOrn;IIMH IIX II pa'cnpo-
CTpaHJlIOrn;HMIICH B c.TeHRax COC.y]:(OB. COCTOJIT OHII TI'3 MHROTHbiX
n 6e3MJlROTHbiX BOJlOROH. ABTOP, OTMeqaJl HepaBHoMepHocTb IIHHep-
Ban;IIII cOCY]:(IICTblX CTeHOR MaJlOrO Rpyra RpoBo06parn;eHIIJI II oc060e
o6IIJlIIe ee y YCTbH JlerOqHOR apTepIIII II y HaqaJla 60TaJlJlOBOH
CBJl3RII, HaXO]:(IIT 06'bHCHeHlIe 3TOMY (TaR me RaR II Muratori,
1959), CCblJlaJlCb Ha Koch (1931), CqnTaIOrn;ero, qTO MeCTa BeTB-
JIeHIIR apTeplIaJlbTIblX ma6epHblx ]:(yr JlBJlHIOTCH Ha1I60Jlee BblpameH-
HblMlI pecI>JleRCOreHHbIMH 30HaMII. O]:(HaRo HynplIHHoB crrpaBe]:(JlIIBO
OTMeqaeT rJlaBHbIe cJla6b1e Me'CTa 3TOR ROHQenI(IIII - OrpaHIIqeHHOCTb
pacrrpocTpaHeHIIJI TnlIllqHblX UJlOMyCOll), IIJlII IIaparaHrJllIeB, B op-
f77
ramI3Me II o61I\ue TPYl-\HOCTII, I-\OTOpble BCTpeqaeT MOPWOJlOr np"
COIlOCTaBJleHIIII WU3UOJlOrnqeCKIIX II MopWOJlOrllqeORIIX ]:(aHHblX: MOp-
4K>JlorUqeCRII 06HapymeHHbIe qYBCTBIITeJlbBbIe OROHqaHUH BCTpeqa-
lOTCJI peme, qeM wn3UOJlOrUqec,rme.
Mbl yme rOBOpUJlU, qTO n COBpeMeBBbIX pa60Tax no ltlOpq.OJlO-
rHU rpyp;BblX naparaBrJlUeB aBTopLI Be Y'r lIX 1I3yqeHUC B cpaBBU-
TeJlbHOM acneRTe C RapOTU]:(Bblltl II H8,ACep;:J;eqHbIMli naparaHrJlUHMn,
ROTOpbl{J CqUTalOT peqenTOpBbIMlI opraBaMII. B pa60Tax, B ROTOpblX
JI.erO'lHbIH «rJlOMYC» u3yqeB ;meKTpOHBOMlIKpOCRonuqeCKll, ero RJleT-
RII Tome pacqemrnalOTCJI RaR xeMopeqenTopBble (Knoche, Schmitt,
1963; Battaglia, Mariani, 1970).
TaRMM 06pa30M, BamBeHllIue BonpOCLI 0 peqJJleRTopBoM BJlUHHUII
Ha mlI3HeHHO BamHLIe WYHRqllll opraHn3Ma, eCJIlI OHII UCXOP;HT II3
COCY]:(HCTblX yqaCTROB meHHO-l'pYP;HOII 06.;IaCTH, CBH'3b1BalOTCJI C
«rJlOMycaMII) RaR IICXO]:(HLlltl nYHRTOM peW.;IeRca. BnpOqeM, «fJlOMY-
CbIl> IIOnOJlb3YlOTCH qaCTO ]:(JlH IIo]:(06HblX nocTpoeHuH, eCJlII OHn Jle-
maT U lHe RpOBeHOCHoro PycJla U BHe rpY]:(HOM nOJlOCTII. B GJleRTpOH-
HOMIIRpOCROnUqeCROH pa60Te Elfvin (1968) HallleJl B HHmHe6pbl-
meeqHOM raHrJlIIII KpOJllma «JleTKU C OCMIIOWBJlbHblMII rpaHYJlar.ru,
KOTopble no wopMe, pa3MepaM II IIJlOTHOCTII TlIImqHbI, HaK OH nOJla-
raeT, ]:(JlH RJleTOK M03rOBoro CJlOH Ba]:(nOqe'IHUKOB. Elfvin CqllTaeT
3TOT THin 'RJleTOK BHOBb OTKpblTblM II 3aTpY]:(HHeTCJI orr.pe]:(eJllITb IIX
WYHKquoHaJlbHoe 3UaqeHlle B peryJlHTopHOII aKTU'BHOCTU raHrJlUJI.
HecMoTpJl Ha CXO]:(CTBO HaiI]:(eHHblx UM rpaHYJI C xpoMaWWIffiHblMII
M03rOBoro CJlOH Ba]:(IIOqeqHIIKa, aBTop HII CJlOBa He rOBoplIT 0 TOM.
qTO ero «HOBbIe) KJleTKU II MOPWOJlOruqeCRU H WYHRn;UOHaJlbHO CO-
OTBeTCTBYlOT naparaHrJlUHM 6plOlliHon nOJlOCTU, KOTopble BClO]:(Y CO-
npOBOlli)J;alOT CUMnaTuqeCRue CnJleTeHUJI II Yllie MHoroRpaTHo OIIU-
caBbI paHee. AHaJlOflIqHbIe KJleTKII 06HapymeHbl B BepXHeM meHHOM
Y3J1e RpblCbI Siegrist C coaBTopaMu (1968), ROTOpbIe Tome IIOJla-
ralOT, qTO OrnpbIJIB HOBbIH BU]:( }\.;IeToR. J13yqeHue yJlbTpaCTpYK-
Typbl II WYHRqIIII RJleTOqHblX 3J1eMeHTOB a]:(peHaJlOBOH CIICTe bI, pac-
IIOJlOmeHHblX B Y3J1ax CIIMnaTuqeCROH HepBHon CUCTeMbI, ROHeqHO,
He JlUllieHO UHTepeca, HO pe3YJlbTaTbi pa60T 6bIJIll 6b1 60Jlee 3qxpeR-
TIIBHblMII, eCJlu aBTopbl 6bIJlU 6bl 3HaROMbi C CUMIIaTo-aApaHaJlOBOH CII-
CTeMOH B qeJlOM, C IICTOpUqeCRHM npOIllJlblM ee II3yqeHHH II BbITe-
RalO uMu oTclOp;a cOBpeMeHHbIMu IIp06J1eMaMu.
HapJlP;Y C pa60TaMII, ROTOpbIe no cylI\eCTBY HeRplITHqeCRII BOC-
IIpRHIIMalOT TeoplIlO 0 peqJJleKcoreHHoM 3HaqeHlIII Rap]:(IIOaOpTaJlb-
HblX IIaparaHrJllIeB, oT]:(eJlbHble MOPWOJlOrII BblCRa3b1BalOT Ha 3TOT CqeT
II HeRoTopbIe COMHeHIIH. IIoBo]:(oM K 3TOMY CJlymaT, C O]:(HOH CTO-
pOHbI, HOBbIe HaXO]:(KH (<Hexpor.IaWWUHHbIX) naparaHrJllIeB B MeCTax,
paCIIOJlOmeHHblX 3a npe]:(eJlaMB Tex 30H, r]:(e OHII BnepBbIe 6blJlII
OIIlIcaBbI, II JlBJlHlOlI\lIeCH peWJleRCOreBHbIMII (Watzka, 1966, Schon-
berger. 1966, 1967, 1969). C ]:(pyroH CTOpOHbl, 06IIJlbHaH qyBCTBII-
TeJlbHaH IIHHepBaqlIJI (rpBropbeBa, 1954; AOJlro-Ca6YPoB, 1958) II
WII3IIOJlOrlIqeCRII 06HapymBBaeMbIe peWJleRCOreHHble 30HbI (JlHTepaTY-
PY 110 peWJleRCOreHHblM 30HaM CM. y B. H. qepHlIroBcRoro, 1960
178
II B. A. JIe6e eBoll, 1965) ORa3aJIHCb He TOJIbRO B YRa3aHHblX CO-
CY IICTbIX yqaCTRaX, a paClIpOCTpaHeHbI no Bcell CIICTeMe COCY]:(HCTO-
ro pYCJIa. II B Moplj>oIj>YHlU HOHaJIbHOM OTHOllieHIIH CBJlBlUlbI He C
naparaHrJIIIOHapHblMM Y3eJIRaMH, a C caMOll COCY]:(HCTOll CTemWll.
B aTOM CMblCJIe 3aCJIymlIBaIOT ynoMIIHaHIIH COMHeHIIJI B O:QeHRe xe-
Mope:QeIITOpHOll pOJIII (<HeXpoMaIj>Ij>IIHHbIX) lIaparaHrJIlIeB, BblCRa3aH-
Hble He]:(aBHO Schonberger (1967). Hall]:(JI TaRlIe Y3eJIRII B CRJIa]:(-
Re CJIII3IICTOll ropTaHII B TeCHOM ROHTaRTe C raHrJIH03HbiMII RJIeT-
RaMH, aTOT aBTOp rOBoplIT, qTO MeCTO TaRHX naparaHrJIlIeB He CBJI-
3aHO C TeMII cocy]:(aMII, ROTopble npe]:(CTaBJIHIOT BamHoe 3BeHO B
reMO]:(IIHaMIIRe RpOBII, II nOTOMY BpJl]:( JIll IIX MomHO paC:QeHIIBaTb
B RaqeCTBe pe:QeIITopOB B nOHIIMaHIIII Heymans II de Castro. K co-
maJIeHIIIO, Schonberger He ]:(eJIaeT II3 CBoero 3aRJIIOqeHIIJI 60JIee
IlIIIpOROrO 0606m;eHIIJI. 3HaqlITeJIbHO paHbme, qeM Schonberger, HO
110 cym;eCTBY TY me MblCJIb BblCRa3aJIa T. A. rplIropbeBa (1954)
B OTHOllieHlI1I RapOTII]:(Horo IIaparaHrJIIIH. OHa BH]:(HT O]:(HO II3 CJIa-
6b1X MeCT . TeopIIII de Castro B TOM, qTO RapOTII]:(Hoe TeJIO, BCTaB-
JIeHHOe no xo y nIITaIOm;IIX M03r COCy]:(OB II npe Ha3HaqeHHOe no
de Castro ROHTpOJIIIpOnaTL XHMlI3M RpOBlI, t.wmeT OCTaBaTLCH H BHe
3Toro pyCJIa. BaCRYJIHpII3a:QIIH RapOTM]:(Horo lIaparaHrJIHH B pMe
CJIyqaeB IIpOIICXO]:(IIT H II3 p;pyrHx HCTOquIIROB, HanpHMep, Hapym-
HOll COHHOll apTeplIlI, 3aTbIJIOqHOll, rJIOToquOll H ]:(pyrlIx, He IIMeIO-
m;HX OTHomeHIIH R cHa6meHHIO M03ra RpOBbIO. A C pa36IIpaeMblx
HaMH aBOJIIO:QIIOHHbIX II03II:QIIM cOBepllleHHo HCHO, qTO «COHHOMY)
IIaparaHrJIIIIO COBceM He 06H3aTeJIbHO 6b1Tb B a]:(BeHTH:QHII BHYTpeH-
HeM COHHOll apTeplIlI: JIORaJIII3a:QHH pY]:(HMeHTapHoro OCTpOBRa MomeT
cp;BlIraTbCH II Ha HapymnyIO COHHYIO apTepmo H Ha RaROll-JIII60
]:(pyroll yqaCTOR, nplIHa]:(JIemaBIlIIIII Ror]:(a-TO III ]:(yre ma6epHblx
apTeplIll.
l1.JIJI Toro qT06b1 BblJlBIIJIaCb HalI60JIee nOJIHaJl RapTHHa Bcell CII-
CTeMbi BHeHa,WIOqeqHIIROBOll xpoMalj>lj>HHHon TRaHM, IIpe]:(cTaBJIJleT
IIHTepeC paccMoTpeTb C aBOJIIO:QIIOHHOll TOqRII 3peHIIJI IIHHepBa:QlIoH-
Hble IIplI60pbl IIaparanrmleB, oco6eHHo alj>lj>epeHTHble. Bblllle yme ro-
BOpHJIOCb 0 TOM, qTO HapH]:(y C nOJIOmlITeJIbHblMII ]:(aHHbIMII 06 IIH-
HepBa:QIIH BHeHaJJ;IIOqeqHIIROBbIX lIaparaHrJIHeB (nHHec, 1924; Hol-
linshead, 1937) IIMeIOTCJI pa60Tbl, HX OTplIIWIO, m;IIe qTO nO]:(TBep-
m]:(aIOT II HeROTopble Ij>H3HOJIOrlIqeCRlIe 3RcneplIr.leHTbl. Peqb 3]:(eCb
II]:(eT He 0 Bcen 3RCTpaap;peHaJIOBOn xpoMalj>Ij>IIHHOn TRaHII, a 06
OTp;eJIbHblX ee yqaCTRax. MI,l IIOJIaraeM, qTO cnopbl 0 nepBHblX OROH-
qaHIIJIX :QeHTp06emHblx HepBOB Ha naparaHrJIIIOHapHblX RJIeTRaX He
HMeIOT CTOJIb IIpIIHn;IIIIHaJIbHOrO 3HaqeHHJI, RaROe nplI]:(aeTCJI IIM MHO-
rliMII IICCJIe OBaTeJIHt.m. OCHOBaHHeM R TOMY CJIymaT CJIe]:(YIOm;IIe CO-
06pameHIIJI.
B IIpO:Qecce aBOJIIOn;IIOHHOro pa3BlITIIJI MHorlie opraHbI xpoMalj>-
Ij>IIHHOn CIICTeMbi B pe3yJIbTaTe H3MeHeHIIn YCJIOBlIii cym;ecTBoBaHHJI
mlIBOTHoro TepHIOT, RaR MI,l BH]:(eJIII, 3HaqeHHe JIH opraHII3Ma II
peAY:QlIpYIOTCJI. B pe3yJIbTaTe pe YRn;1I1I B03HlIRaIOT JIII60 pY]:(HMen-
TapHble opraHbI, COXpaHJlIOm;HeCJI B 3aqaTOqHOM COCTOJIHIIH B Teqe-
179
HlIe BCerO lIepHO]:(a mlI3HH OpraHlI3Ma, JlII60 npOBH30pHbIe, Cym;ecT-
BYIOm;IIe II q,ymm;IIOHlIpYIOm;He TOJlbKO y 3apO]:(blllIeR.
HaM 113BecTHO, qTO y HII3ll1IIX n03BOHOqHbIX (MHHor) ClIMnaTII-
qeCKaH HepBllaH CIICTet.m HaxO]:(HTcH B TaKOR 3aqaTOqllOR cTa]:(HH
pa3BHTIIH, qTO HII B MOPWOJlOrlIqeCKOM, HII B QJlI3HOJlOrHqeCKOM OT-
HOllleHHHX He 06eCneqIlBaeT HHHepBaqmo xpOMaWWIIHHOR TKaHII 8TIIX
mHBOTllblX. CIUnIaTlIqeCKaH IIJlHepBaqHH pa3BIIBaeTCH II03]:(lIee H8
1I0CJle]:(yIOm;HX 8Tanax q,HJlOrelle3a rr03BOHOQHblX II 6epeT Ha ce6H
q,YHKqIIIO BHella nOqeqHHKOBblX xpOMaWWIIHHblX ::meMeHTOB. HeJlb3H
JlH IIpe]:(IIOJlOmIITb, qTO KaKHe-TO He60JlblllHe YQaCTKH xpOMaq,q,IIH-
HOR TKaHH OCTaIOTCH He 3aTpOHYTblMH 8TlIM rrpoqeccoM II npe6b1-
BaIOT B COCTOHHIIII KaK Ubi eCTeCTBeHHOR 3q,q,epeHTHoil «]:(eHepBa-
qHIII). ::ho npe]:(nOJlOmeIlHe, eCJlII 0110 cnpaBe]:(JlIIBO, OJlmIlO HaRTH
OTpameHlIe II B IIH HBH yaJlbHoM pa3BlITIIH mHBOTHbIX - B IIaparaH-
rJlIIOHapHblX opraHax, peKanIITYJlHpYIOm;IIX HCTOplIqeClmR npoqecc.
C 3TOR TOqKH 3peHHJI npe CTaBJlHIOT IIHTepeC pa60Tbl Muscholl,
Vogt (1963, 1964), Brundin (1966), JI. B. CaKOBlIq (1964, 1969),
Coupland (1965), Coupland, Weakley (1970) H Winckler (1969).
Brundin, COqeTaIOm;HR B CBoeR pa60Te q,H3HoJlOrHqeCKIIR 3KCIIe-
pIlMeHT C ::)JIeI\TpOHHOMIlKpOCKOnHQeCI HM HaOJlIO,lWHHeM, nOKa3aJl, qTO
B yCJlOBIIHX CTpecca HacTynaIOm;aJl y HOBopom]:(eHHblX KpOJlIIKOB rJlII-
KoneHHH CHHmaeT co epmaHlIe KaTeXOJlaMHIIOB B KJleTKaX M03rOBO-
ro CJlOJl lIa]:(nOQeqIlIlKOB, HO He BJlHHeT Ha KJleTKII IIpeaOpTaJlbHOrO
naparaHrJlHH. ABTOp CqHTaeT, qTO rJlIIKOneIIIIH, Bbl3bIBaIOm;aJl pe«iJ-
JleKTOpHoe BbI]:(eJleHHe KaTeXOJlaMHHOB II3 M03rOBoro CJlOJl Ha nOqeq-
HHKOB qepe3 qpeBllblR HepB, He OKa3b1BaeT allaJlOrHqHOrO ]:(eiicTBIIH
Ha BHeHa]:(nOQeqHIIKOBblii IIaparaHrJlIIR IIOTO!\lY, QTO OH JlHllIell I1H-
HepBaqlIlI. OqeHb npaBHJlbHO, lIa Halll B3rJlH , II pyroe 3aKJlIOqe-
RHe Brundin, OCHOBaHHoe TO »m Ha OTCYTCTIIHII IIl1l1epBaqll1l npeaop-
TaJlbHOrO IIaparaHrJlHH: 1I0CJle ]:(aqII CI -Tllp03HHa IIJlII ]:(IIrIl pOK-
CHtPeHIIJlaJlaHlIHa BKJIJOqeHIIC II nOCJIC YIOm;ee cnOHTaHHoe IICqe3HO-
BeHlIe pa]:(HoaKTHBHoro Hopa]:(penaJIIIHa npOIlCXO HT B naparaHrJlIlHX
},fe]:(JleHHee, qeM B M03rOBOM cJloe Ha]:(nOQeqIlIlKOB.
JI. B. CaKOBlIq, IIccJlep;YJl IIHHepBaqlIIO 6pIOIliHo-aOpTaJlbHblX IIa-
paraHrJllIeB (IeJlOBeKa, p;eJlHT ero pa3BlITIIe lIa ]:(Ba neplIo]:(a. B lIep-
BOM Ha6J1IOp;aeTCH HeIIpepbIBHblR pOCT naparaHrJlHeB II BblCOKaJl WYHK-
qlIoHaJlbllaH aKTIIBHOCTb xpOl\mq,WIIHHblX IweTOK (K COmaJleHIlIO.
aBTOp He nIlllleT, KaI\IlM TeCTOM OHa nOJIb30Ba."laCb npH onpen;e Ie-
HHII aKTHBHOCTH). Bo BTOpOM - npeKpam;ClIHe pOCTa H nOCTeneH-
HaH pe]:(YKqIlH naparaHrJlIlH, KOTopaH npo]:(OJImaeTCJI lIeCKOJIbKO neT.
Ha npOTHmeHlUl IWTOPbIX coxpaHHIOTCJI OT]:(eJlbHble rpynnbl q,YHKqIlO-
lIaJlbHO aKTIIBllbIX XPOMaWq,lIHHbIX KJleTOK. ABTop no]:(qepKIIBaeT, qTO
B lIepBOM IIepIlo]:(e 6pIOlliHOll aOpTaJlbHbllI na paraHrJlIlR OQeHb 6ep;eH
IIRHepBaqHoHHblMII CTpYKTypaMII. Bo BTopml neplIo]:(e KOJlIIQeCTBO
IIX 3allieTHO YBeJlIIQHBaeTCH, oc06eHHO B Tex yqaCTKaX naparallrJlIIH,
r]:(e, no Ha6J1IO eHIIHM J1. B. CaKOBlIq, npOHCXO]:(HT 3aMem;eHHe IIa-
peHXHMaT03HblX ::meMeHTOB coe]:(HHIITeJlbHOll TKaHbIO, a rHe3 a em;e
coxpaHIIBllIlIXCH xpOMaWWllHHblX KJleTOK OCTaIOTCH nO-IIpemHeMY co
180
CRY,IJ;Ht.IM ROJIH'IeCTBOM HepBHblX aJIeMeHTOB. HapH]:\Y C BOJIOKHaMII
II OKOHtJaHIIJlMn 6e3MJ1KOTHbiX HepBOB B COe]:\nHnTeJIbHOTKaHHOH CTpO-.
Me pe,IJ;yn;IIpylO erocH lIaparaHrJInH JIOKaJIn3YIOTCJI HepBHbIe OKOH-
qaHIIJI MHKOTHblX qYBCTBnTeJIbHbIX BOJIOKOH B <popMe CJIOmHblX IIH-.
RalICYJInpOBaHHblX RJIy60qKOB, ROTOpbIe, no MHeHnlO aBTopa, OTHQ-
CJlTCH He K xpOMa<p<pnHHbIM KJIeTKaM, a TOJIbKO R COe]:\HHnTeJIbHO-
TKaHHblM npOCJIOHKaM. MHKOTHbIe n 6e3MHROTHbie HepBHble BOJIOKHa
opneHTnpOBaHbl no XO]:\Y KpOBeHOCHblX COCy]:\OB. TIo Mepe yrJIy6JIe-
HnH IIpOn;eCCOB pe]:(YKn;nn ROJInqeCTBO HepBHblX OROHqaHnH YBeJIlIqlI-
BaeTCH. ABTOp 06HapymnJIa nx Ha TOM me MeCTe n 1I0CJIe 1I0JIHOro.
nCqe3HOBeHnH opraHa.
I1HTepecbl CaKOBlIq CBO]:\HTCH K BbIHCHeHnlO nHHepBan;IIn 6plOm'-
Ho-aOpTaJIbHOrO lIaparaHrJInJl qeJIOBeKa. Ha OCHOBaHnu 6JIn30CTII Hep-
BHblX OROHqaHnH K KpOBeHOCHbIM cocy]:\aM OHa ]:\eJIaeT BbIBO]:\ 06 IIX
xeMopen;elITopHoH <pYHKn;un. l1.JIH Hac l\IaTepUaJI CTaTbn CaROBnq IIpe]:\-
CTaBJIJleT ]:\pyroH nHTepec. OnucaHHblH elO npeaOpTaJIbHblH 6pIOIliHOH:
lIaparaHrJInH HBJIHeTCJI peKalInTYJIlIpYIO IIM opraHOM. TIepBblH nepnog.
ero pOCTa n aKTIIBHOCTn xapaKTepu3yeTcH CRY]:\HOH nHHepBan;IIeit
II 1I0BTOpHeT TeM caMblM <pnJIOreHeTnqeCKlIe CBOHCTBa IIpeAKOB. BTO-
pOH nepnop; pe]:(yKn;un conpOBom]:\aeTCH pa3BlITUeM coe]:\nHIITeJIbHOR
TKaHII n KpOBeHOCHblX COCY]:\OB n lIapaJIJIeJIbHO UX nHHepBan;IIeH.
I1HHepBnpYIOTCH He napeHxuMaT03HbIe aJIeMeHTbI, KOTOpbIe TaK It
OCTalOTCJI co CKY]:\Ht.IM KOJIlIqeCTBOM HepBHblX aJIeMeHTOB, a 60JIee 1103]:\-.
HlIe coe]:\nHnTeJIbHOTKaHHble 06pa30BaHnH II COCY]:\bl, KOTopble IIOJIY-
qalOT cnMlIaTnqeCKne HepBbl BTOpnqHO II He CBH3aHbi HelIocpe]:\CT-
HeHHO C rrapeHxnMaT03HbiMn aJIel\IeHTaMu peKamlTyJInpYIO ero,
opraHa.
Winckler (1969) OTMeTIIJI, qTO BO BHeHa]:\nOqeqHnKOBOH xpoMa<p
<pnHHoH TKaHn 6plOlliHOH 06JIaCTn OqeHb l\IaJIO <pJIyopecn;npYIO IIx
HepBHbIX BOJIOKOH, a aJIeKTpOHHOMUKpOCROnlIqeCROe n3yqeHne ee He
06HapymUJIo a<p<pepeHTHbIX HepBHblx OROHqaHlIH Ha rraparaHrJInOHap-
HblX RJIeTKaX Hn BO BHYTpnYTp06HOM pa3BuTuII, HII B 1I0CTHaTaJIh-
HOM (Coupland, Weakley, 1968, 1970).
TIpnBe]:\eHHble ]:\aHHbIe MorYT paCC IaTpnBaTbCH KaK IIpII3HaKn,.
peKalInTYJInpylO ne a<p<pepeHTHYIO «]:(eHepBan;IIIO» xpoMa<p<PIIHHoH
TKaHn Hn3ll1nX 1I03BOHOqHbIX. C aTllX me rr03nn;nn Mbl paCn;eHllBa-
eM paClIpe]:\eJIeHlIe a<p<pepeHTHbIX ClIMnaTnqeCKnx IIplI60pOB He H
cnen;n<plIqeCRlIX KJIeTKaX KapOTn]:\Horo lIaparaHrJIIIJI, a B oKpymalO eii.
TRaHII (CM. BbIllie OlIlIcaHne onbITOB Biscoe, Stehbens, 1967). TaKo-
BO B HaCTOJl ee BpeMJI COCTOHHlIe Borrpoca 0 naparaHrJInHX MJIe-
KonIlTalO nx.
TIpem]:\e qeM IIepeXo]:\nTb K xapaKTepllCTnKe caMoro KpYIIHOro.
IIaparaHrJInH - M03rOBoro CJIOJl Ha]:\IIOqeqHnKa, 1I0]:\Be]:\eM HeKOTOpbIe'
nTorn CKa3aHHOMY B aToH rJIaBe n 1I0nMTaeMCH KOpOTRO IIpOaHaJIII-
3npOBaTb MOP<P0JIOrlIqeCKne <paKTbl 0 naparaHrJIIIHX MJIeROlInTalO nx..
ROTOpbIe xapaRTepn3YlOT IIX KaK cTylIeHb B aBOJIIOn;nOHHOM pa3Bn-
Tnn xpOMa<p<pnHHOH aH]:\oKpIIHHOH cnCTeMbI n03BOHOqHbiX mnBOTHbIX.
IIo]:\qepKHeM e e pa3, qTO B «HeXpOMaq,<pIIHHbIX» lIaparaHrJInHX B.
181
'IIaCTOHIIJ;ee BpeMH ]:\OKaSaHO npDCYTCTBlIe ropMOHOB 9JIeKTpOHHOMDKpO-
.cKOnD'IeCRDM aHaJIDSOM, iPJIyopec eHTHOH rDCTOXDMlIeH D 6IIOXDMD-
qeCKHM DCCJIe]:\OBaHHeM 9KCTpaKTOB. O]:\HaKO a,TlInlISM CTpoeHHH na-
araHrJIDOHapHWX KJIeTOR II naparaHrJIDOHapHOH TKaHII OTJIlIqaeT DX
OT MosrOBoro CJIOH Ha]:\nO'Ie'IHnKOB.
B I'JIaBe 0 MnHorax 6bIJIO nOIiaSaHO, qTO BCJI TOJIIIJ;a CTeHRII
cep]:\ a DJID, KaK ero HaSWBaIOT, paCIlIHpeHHblH nYJIbCHpylOIIJ;1lli yqa-
CTOR apTepnaJIbHOH Tpy6KII npOHD3aHa xpoMaiPiPuHHblMIi 8JIer.IeHTa-
MD, TaK me RaK D CDCTeMa ]:\op3a;rbHoii aopTbl, HaqllHaH OT II ma-
06epHoro cerMeHTa. B rJIaBe 0 pbl6ax rOBOpDJIOCb, 'ITO, XOTJI OCHOBHbl8
lIJIaCTbl Xpor.laiPiPllHHOH TKaHD 3aJIeralOT B CTeHKax MarlICTpaJIbHblX
TyJIOBIIIIJ;HbIX BeH, O]:\HaRO DMeeTCfI eIIJ;e MHoro yqaCTKOB apTepDaJIb-
Ho-aOpTaJIbHOH CDCTeMbl, r]:\e OHa coxpaHHeTCJI (HanpDMep, «aRCII-
JIHpHOe cep]:\ e» CeJIaXDH}. XpoMaqJiPuHHaH CDCTeMa KpyrJIOpOTblX D
IJbl6, iPYHK DOHHpYIOIIJ;aH y HHX K31> 3H.IJ,OKpDHHaH, y HaseMHblX mu-
BOTHblX npeTepneBaeT IISMeHeHIIH, CBHsaHHbIe C pe]:(YR HeH ma6epHWX
COCy]:\OB. H anor.fH 11M, 'ITO ]:\ame BHYTPIi "JIaCca aMiPn6HH MomHO npo-
CJIe]:\DTb 1I0]:\06Hble lIepecTpoiiKD, HalIpDMep B VI nape ]:\yr: y XBO-
CTaTbIX 60TaJIJIOB npoToK HBJIJleTCH iPYHK IIOHDPYIOIIJ;DM COCY]:\OM, a y
.oecxBocTWX - HeiPYHR DOHHpYIOw;e H OOTaJI::IOBOll CBHSROH. BeHosHblH
CIIHYC, HacwIIJ;eHHblH ceKpeTDpYIOIIJ;DMD xpoMaiPiPlIHHwMII 9JIeMeHTa-
MD, y HDSIlIIIX nOSBOHOqHblX nCqeSaeT B IIpon;ecce <);JHJIOreHe3a KaK
caMOCTOJlTeJIbHblH Y'IaCTOK nplIeep]:\eqHblX COCY]:\OB, a y IITD yme
n;eJIIIKOM BXO]:\DT B COCTaB npe]:\cep]:\uH.
A]:\peHaJIOBaH TKaHb MJIeKOnnTalOIIJ;lIX BCTpeqaeTCJI B Tex OT]:\e-
JIax cepJ:\IJ;a (lIpe]:\cep]:(De, cep]:\e'IHble YIllKD), Ky]:\a onycTDJICJI B xo]:\e
-3BOJIIO DII BeH03HblH CIIHYC, H pa3MeIIJ;eHa Ha Tex KpoBeHocHwx co-
<cy]:\ax, B iPopMlIpoBaHIID ROTOpblX npDHHJID yqacTDe ma6epHwe (B YCTbe
.JIerOqHOH apTepDD II 110 ee xO]:\y ]:\0 paSBeTBJIeHDJI Ha HapymHyro
:II BHYTpeHHIOIO, B ]:\yre aOpTbI y MeCTa OTxom]:\eHIIH 60TaJIJIOBOH
-eBJlSKD, B 6eSbIMJIHHOH, nO]:(KJIlOqH'IHOii apTepDHx D B 6DiPypKa DII
'COHHOH). 8TII yqaCTKH xpoMaiPiPlIHHoH cnCTeMW MJIeKOIIIITalOIIJ;DX,
xapaKTepHsYIOIIJ;IIe nos]:\HHH 8Tan 8BOJIIO IIOHHOrO pasBDTIIH xpor.miP-
iPlIHHOH CIICTeMbl, DMelOT cJIe]:\ylOIIJ;De CTpyKTypHbIe oc06eHHocTH.
TIaparaHrJIDII y MJIeKOmITalOIIJ;DX acnOJIOmeHbI B tunica adven-
ticia RpoBeHocHblX COCY]:\OB DJID B oKpymalOIIJ;eH ee mlIpoBoH KJIeT-
-qaTKe llIeHHo-rpY]:\HoH 06JIaCTD. B cooTBeTcTBylOIIJ;DX rJIaBaX MW'IIPD-
BMII IIo]:\p06Hb1H iPaKTDqeCRIIH r.IaTepDaJI, 1I0KaSblBalOIIJ;DH rpa]:\IIeH'r
lIepe:\feIIJ;eHIIJI xpoMaiPiPDHHbIx 8JIeMeHTOB BHYTPII CTeHOK KpoBeHoc-
HblX COCY]:\OH - OT tunica intima (KpyrJIOpOTWe pbl6bl) K tunica
media (seMHoBo]:\Hble, npecMwKalOIIJ;DecH) II K tunica adventicia
(IITD bl). TIpII BceH paSHOpe'IlIBOCTII BO BSrJIH]:\ax Ha RapoTD]:\HbIH na-
paraHrJIDH y MJIeKOIIIITalOIIJ;nx (Ha ero npoDcxom]:\eHlIe, DTOJIOrnlO IT
iPYHRn;IIIO) e]:\DHCTBO BSrJIH]:\OB DMeeTCH TOJIbRO B OTHOllieHlI1I ero JIO-
KaJIDSa IIH Ha HapymHoH 060JIOqKe COHHOH apTepDD (IIJIII oKpymalO-
IIJ;eH coe]:\DHDTeJIbHOll TKaHD) (Kohn, 1900, 1903; Smith, 1924,
ode Castro, 1926, 1928; CMHpHoBa, 1948; TIpnllMaR, 1962; Ha]:\eeB,
Y JIYM6eRoB, 1969, n ]:\p.). B Ha pymHoH 060JIOqKe apTeplIll. II B 91I1I-
.182
KapJ];e npeJ];CepJ];UH paCIIOJIOmeHbl U BCe J];Pyrue IIaparanrJ1Jrn: KPYII-
HblX COCYJ];OB rpYJ];HOH qaCTII TyJIOBUlI\a (Kohn, 1903; Wiesel, 1906;
Penitschka, 1931; Muratori, 1932-1937; Palme, 1934; Seto, 1935;
Nonidez, 1935-1937; KYIIPIIHHOB, 1959, II ]:\p.). B nepBblX rJIaBaX
Mbl ]:\OCTaTOqHO 1I0J];p06HO 060CHOBaJIU iPYHKIJ;DOHaJIbHYIO eJIeco06-
paSHOCTb HaxomJ];eHUH xpoMaiPiPuHHoH TKaHU B6JIUSII MarUCTpaJIbHO-
1'0 RpOBOTOKa Y Husmux n03BOHOqHblX II lIepeJ];BumeHHe ee B rJIy6L.
CTeHOK RpOBeROCHblX COCY]:\OB Y Bblcmux, no Mepe 1I0TepII aTIIMII op-
raHIISMaMII lIoTpe6HocTu B TaKOH iPYRK UU. Hau60JIee nOJIHOe OTpame-
Hue 8Toro IIpo ecca Mbl BCTpeqaeM Y MJIeKOnIITaIqIl\UX.
llOTepH iPYHK IIU, eCTeCTBeHHO, BJIeqeT sa C060H II pH]:\ J];Pyrux
CTPyKTypHblX IISMeHeHuH, B eJIOM xapaKTepusYlOlI\IIX 6es]:\eHcTBu8
U aTpoiPulO lIaparalIrJIlIeB: OHU rYCTO IIpopaCTalOT coe]:\UHUTeJIbHOH
TKaHblO, KOTOpaJl ]:\eJIUT meJIesy Ha J];OJIbKII II lIepeCJIalIBaeT OT]:\eJIb-
Hble RJleTKU, caMII meJIeSUCTbIe 8JIeMeHTbi sanYCTeBalOT, MeJIbqalOT,
Bblpa6aTbIBalOT HlIqTOmHbIe ROJIlIqeCTBa ropMOHa UJIU COBceM TepHIOT
8Ty ClIoc06HOCTb. He60JIblllaH BeJIUqUHa OCMUOiPUJIbHblX rpaHyn, KO-
TOpaJl HapJl]:(y C J];pyruMII IIplISHaKaMII npnHIIMaeTCH B paCqeT IIPII
o eHKe 8TIIX opraHOB, Tome MomeT OTpamaTb npo ecc HeJ];opaSBUTUJI
IIHKpeTOpHOH iPYHK UII B flUX.
llpu 1I0]:\xo]:\e K UCCJIe]:\OBaHlIm IIaparaHrJIUeB C IICTopUqeCKIIX
IIOSII UH CTaHOBJlTCH 1I0HJlTHblMU II MHorue J];pyrue iPaKTbl. llapa-
raHrJIUOHapHbIe OCTpOBKlI, JlBJIHHCb pY]:\UMeHTapHbIMU OCTaTKaMU Kor-
J];a-TO 6b1BIlIUX CIIJIOmHblX IIJIaCTOB U THmeH xpoMaiPiPlIHHoH TKaHlI,
MorYT 6blTb KpynHee, MorYT 6blTb MeJIbqe, a MorYT COCTOHTb Bcen>
JIUllib lIS HeCKOJIbKUX RJleTOK. OCTpOBKII U RJleTKII BHYTpn OCTpOB-
KOB MorYT OTJIlIqaTbCJI J];pyr OT J];pyra U 110 cIIoc06HOCTII K Bblpa-
60TKe IIHRpeTa - B OJ];HIIX OHa coxpaHHeTCH B 60JIbllleH CTelIeHlI,
B ]:\PYI'IIX B MeHbllleH UJIII COBceM UCqeSaeT B npo ecce aBOJIIO UU.
B aTOM CMblCJIe MomHO 6b1JIO 6bl IIpUSHaTb cyll\eCTBOBaHue «XPO-
MaiPiPIIHHbIX», (meXpoMaiPiPIIHHbIX» U «CMelllaHHbIX» lIaparaHrJIUeB,
HO IIpII aTOM HII B KOOM CJIyqae He KJIaCTb B OCHOBY TaKoro ]:\eJIe-
HUH IIX (<Bal'aJIbHOe», IIJIn «CUMlIaTlIqeCKOe», rrpoucxomJ];eHue, a TOJIb-
KO CTeIIeHb peJ];yK IIU. OJ];HaKO BO IIS6emaHue nYTaHHI.J;bI U3-sa He-
COCTOHTeJIbHOCTII yqeHIIH 0 UX paSJIUqHOM IIpoucxomJ];eHIIII, a rJIaB-
Hoe us-sa Toro, qTO B HaCTOHlI\ee BpeMJI Te IIaparaHrJIUII, KOTOpbl8
CqUTaJIIICb (meXpoMaiPiPuHHbIMU», 6JIaro]:\apJl HOBOR TeXHIIKe CTaJIR
xpOMatPiPuHHbIMU», - OT 8TUX TepMHHOB CJleJ];yeT OTKaSaTbCJI.
BIIOJIHe IIOHJlTHbl II Bapua uu B paCIIOJIOmeHUII Tex IIJIU UHblX
IIaparaHrJIUOHapHblX rHeSJ]; y paSHblX mlIBOTHbIX, oc06eHHO UMelOlIJ;IIe-
MeCTO 110 OTHOllieHUIO K MeJIKIIM yseJIKaM. OqeHb BOSMomHO, qTO
60JIblllOe KOJIUqeCTBO naparaHrJIUOHapHblX OCTpOBROB y mUBOTHblX,
110 cpaBHeHlI10 C TaKOBblMII y qeJIOBeKa, 06'bJlCHHeTCH paSHOH CTe-
lIeHblO UX pe]:\YK UII: Y qeJIOBeKa, KaK y Hau60JIee BbiCOKoopraHU-
<lOBaHHOro CYlI\eCTBa, OHa 3alllJIa J];aJIbme, qeM Y J];pymx opraHU3-
MOB. <1>opMa KJIeTOR, eCJIII rrpOCJIe]:\UTb sa HeH B npo ecce mOJIIO-
IIOHHOro paSBUTUH, MeHHeTCH, IIpeBpall\aeTCH us MHorOOTpOCTqaTOR
(cep]:\D;e MIIHor, «aKCUJIHpHOe cepJ]; e» CeJIaXUH) B 1I0JIlIrOHaJIbHYIO.
183".
IIJIII OBaJIbHYIO (nTII bl, MJIeROnIlTalO lIe); pe]:(YR IIH OTPOCTI{QB II]:\eT
IIOCTeneHHO, II eCJIII OHII BCTpeqalOTCH IIHOr]:\a Y MJIeKOIIIITalO IIX,
'.1'0, HeCOMHeHHO, IIpe]:\CTaBJIHIOT c060ii CTPyKTypbl, YHaCJIe]:\OBaHHble
OT CBOIIX MHOrOOTpOCTqaTblX IIpe]:(KOB.
B IIpOTIIBOIIOJIOmHOCTb aRCTpaa]:\peHaJIOBbiM lIaparaHrJIIIHM M03-
rOBOH CJIOH Ha,lVIOqeqHImOB MJIeKOnIlTalO IIX ]:\OCTllraeT BblCOKOii MOp-
tP0JIOrIlqecKoii II ciJYHK IIOHaJIbHOH CJIomHOCTII. B IIpO eCCe 8BOJIIO-
"QIIOHHOrO paaBIITIIH OH IIpll06peTaeT opraHHYIO CTpYKTYPY II KaR
caMOCTOHTeJIbHblH aH]:\OKpIIHHbIH meJIeallCTbIH OpraH 06JIa]:(aeT CJIom-
HOH CIICTeMOH KpOBo06pa eHIIH II IIHHepBa IIII. CJIOmHOCTb CBII]:\e-
'.I'eJIbCTByeT 0 BblCOKO ClIe I1aJIllallpOBaHHoii ciJYHKD;IIII, ]:\JIJI BbIlIOJIHe-
HIIH KOTOpOH MoarOBOH CJIOH Ha]:\IIOqeqHIIKa Hym]:\aeTCJI B CTporOH
II TOqHOii perYJIH IIII HalIpaBJIeHIIH TpaHClIopTa ropMOHOB, ocy e-
CTBJIJleMOrO opraHHbIM KpOBo06pa eHlleM.
ApTepIlaJIbHbIH nplITOR HpOBlI R Ha]:\IIOqeqHIIKaM, ClICTeMa pac-
IIpe]:\eJIeHIIH opraHHbIX apTepIlH, apTeplIOJI II KanlIJIJIHpOB B KOpKO-
BOM II M031'OBOM CJIOHX, MopiPOJIOrIIqeCROe CTpoeHlIe BeHoaHoro
OTTORa, COCY]:(I1CTbIe CBH3U H3]:(nOqeqHOII meJIeabl co CMemHblMn
opraHaMII, HaJIIIqlle apTepilOBeH03HblX 3HaCTOM030B II npu.CTeHOqHblX
CiPIIHRTOPOB, perYJIIIpylO I1X MeCTHbIH KPOBOTOR,- Bce aTO OTpam3eT
;CJIOmHble OHTO- II iPIIJIOreHeTIIqeeKIIe ocofieHHOCTII opraHa.
OCHOBHblMII IICTOqHImaMII KpOBOCHa6meHIIH Ha]:\nOqe'lHlIROB MJIe-
KOIIIITalO I1X mIlBOTHblX II qeJIOBeKa HBJIJlIOTCJI HIImHIIe ]:\naiPpar-
MaJIbHbIe, nOqeqHbIe apTepIlIl II aopT3 (Landau, 1907; I1BaHoB, 1930;
30JIOTYXIIH, 1923; AypIl blH, 1945; raJIb OBa, 1958; qIlqIlHa]:\ae,
19626), a T3Kme ]:\OnOJIHIITeJIbHbIe - qpeBHaH II 1I0HCHllqHaH (30JIo-
'TYXIIH, 1923; raJIb OBa, 1958; QlIqIlHa]:\3e, 19626), apTepIlIl mIlpo-
BOH KanCYJIbl IIO'lKII (30JIOTYXIIH, 1923), BHYTpeHHHH ceMeHHaH, JIe-
BaH meJIY]:\OqKOBaH (r3JIb OBa, 1958) II BepxHe6pbDlGKe'lH3H (qllqIl-
Ha]:\ae, 19626). CIIcTeMa paaBeTBJIeHlIH apTeplIH Ha KanlIJIJIHpHYIO
CeTb, 110 ]:\aHHbIM CT3pbIX aBTopOB (Nagel, 1836; Henle, 1865) n
60JIblllIlHCTBa COBpeMeHHblX (Bennett II Kilham, 1940; Bennett, 1941;
AypIl blH, 1945; CaIIIIH, 1964, II ]:\p.) , IIpOXO]:\IIT pa3]:\eJIbHO B KOp-
KOBOM II MoarOBOM CJIOHX Ha]:\IIOqe'lHIIRa. HecKoJIbKO IIHOe MHeHIIe
IIO aTOM)' Bonpocy BblCR3aaHO TOJIbKO B pa60Te H. A. qlIqlIHa]:\ae
{1962a), KOTopaH, Ha OCHOBaHlI1I CBOIlX aKClIepIlMeHTaJIbHblX ]:\aH-.
HbIX, CqIlTaeT, qTO apTepIl3JIbHaH ceTb MOal'OBOI'O Ha]:\nOqe'IHIIKa IIpe)];-
CTaBJIHeT C060H IIpO]:\OJImeHIIe RpOBeHOCHOH ceTII KOpbl. AJIJI IIHTe-
pecYIO I1x Hac BonpOCOB 0 IIYTJlX TpaJICIIOpTa ropMOHOB Ha]:\nOqeq-
HIIKa yKaaaHHble ]:\eT3JIlI 3pTepIlaJIbHOrO CH36meHIIH opraH3 He CTOJIh
cy ecTBeRHbl. UHII I1HTepeCHbl TOJIbKO C ToqKII 3peHIIJI B03MomHofi
.aHaTOMlI'leCKOH CBH3I1, a OTCIO]:\a II iPYHK IIOHaJIbHOrO BJIIIHHIIH IIpIl-
.JIeralO eii K M03rOBOM)' CJIOIO KOpbl Ha]:\nOqeqHIIKa. BOJIee BamHoe
3Ha'leHlIe IIMeeT paCCMOTpeHlIe CIICTeM BeH03Horo OTTOKa KpOBII OT
Ha]:\IIOqeqHIIKOBOii meJIeabl II IIX CBJlaeH C cocy]:\aMn 6JIlIaJIema nx
.opl'aHOB.
ApTepIIII MoarOBoro CJIOH Ha]:\nOqeqHIIKOB, paCIIaBilllICb Ha KaIIIIJI-
J1J1pbl B ero IIapeHXIIMe, OIIJIeTalOT xpoMaciJciJnHHble KJIeTKII II co611-
184
palOTCH B BeH03HYIO CeTb, rrpOHII3b1BalO.l.QYIO BCIO TOJIII\Y M03rOBOH
qacTII: Ha]:\nOqeqHlma (Bennett, Kilham, 1940; Bennett, 1941). 110
HanpaBJIeHIIIO K :qeHTpaJIbHbiM OT]:\eJIa 1 Hail;nOqeqHlma BeHbl YKPYII-
HHIOTCH, CTeHKII IIX HCTOHqalOTCH H oGpa3YIOT IlIHpOKlie BeH03Hble'
ClIHYCOH]:(bl.
HaR 1I3BeCTHO lI3 PYROBO]:\CTB H I\IHOrO IHCJIeHHbIX ;mClIepHMeHTaJIb-
HbIX pa60T, H3 CJIHHHHH BCH03HbiX CHHYCOlI]:(OB o6pa3yeTcH :qeHTpaJIb-
HaH BeHa Ha]:\nOqeqHIIRa, KOTopaH BlIa]:\aeT B HHmHlO1O nOJIYIO H no-
qeqHYIO BeHbI. t.lacTb apTepHaJIbHOii KpOBH RopRoBoro CJIOH Tome-
]:(peHHpyeTCJI :qeHTpaJIbHOH DeHoii. Ho, RpOMe ;noii OCHOBHOii MarlI-
CTpaJIII, MHOrOqllCJIeHHble BeH03Hbie BeTOqRII 06pa3YIOTCH B6JIII311 KaII-
CYJIbi Ha]:\rrOqeqHIIKa II iPOpMIIPYIOT 60JIee KpYIIHble 8KcTpaoprallHble
CTBOJIbI, KOTopble coe]:\IIHHIOTCH C BeHaMIi oKpymalOlI\lIx opraHOB. CJIIIH-
Hlle BeH03HbiX COCY]:\OB Ha]:(nOqe llIIiKa C ]:\yroo6pa3HblMIi BeHaMII no-
qeK, OlIlIcaHHoe ell\e B rrpOIllJIOM BeRe, 6bIJIO rro]:\TBepm]:(eHo n03]:\HeH-
IlIIiMIi pa60TaMII (HIICeJIb, 1928; llIeHiPaHH, 1949; faJIb:qoBa, 1958).
Kpol\le TOro, MHOrOqllCJIeHHbIe BeHbI, OTXOJ:\HlI\lIe OT IIoBepXHOCTI'I
Ha]:(IIOqeqHIIKOB, coe]:\IIHHIOTCH C CIiCTeMofl BOpOTHOii BeHbl lIeqeHII
(CanIlH, 1961, 1964), Bna]:(alOT B BeHbI AliaiPparMbI (faJIb:qoBa, 1958;
CaIIIIH, 1964), rJIIiCCOHOBOii RanCYJIbl II rrapeHXIIMbi neqeHII(llIeH-
iPaHH, 1949; CanllH, 1964), o60JIOqeR CeJIe3eHRII, no]:(meJry]:\OqHOH
meJIe3bl, nlIlI\eBo]:\a II Kap]:\lIaJIbHOH '1aCTII meJIY]:\Ka (CamlH, 1961,.
1964), a TaKme B BeHbI rrOJIyJIyRHbIX Y3JIOB COJIHeqHOrO CIIJIeTeHIIH
(CanIlH, 1964). Harrison (1957) II Harrison II Holy (1960) 110-
Ka3aJIII, qTO IIpli CIIa3Me apTepllii M03rOBoro CJIOJl Ha]:\nOqeqHIIKOB-
B03MomeH lIepexo]:\ RpOBIi B KOpTIIKaJIbHble CIiHyCOII]:\bI.
OC06bIH HHTepec B MopiPOJIOrllqeCKOii OpraHlI3a:qlllI CIiCTeMbi KpO-
BOTORa Ha]:\nOqeqHblX meJIe3 npe]:\CTaBJIHIOT Te CTpyKTypbl, KOTOpblft
IIrpalOT pOJIb npllcnoco6JIeHIIH, perYJIllpylOlI\lIX MeCTHoe KpoBoo6pa-
lI\eHlle B opranax. fJIaBHbII\f11 113 HIIX HBJIHIOTCH ciPlIHKTepoo6paa-
Hble 06pa30BaHliH (pacrrOJIOmeHHLle B CTeHKax MCJIKIIX apTepliH n
BeHYJI, T. e. B npe- II rrOCTRalIlIJIJIHpHOM pYCJIe), apTepHo-BeH03Hble
aHaCTOM03b1 B napeHxnMe opraHa IlJIIi BHeopraHHO II CIiHycOII]:\aJIb-
Hble RanIlJIJIJlpbI.
fJIa]:\Klle MbIIlI:qbl cocy]:\nCTblX ciPIIHKTepOB, nOCTpoeHllbie II3 :qlIp-
KYJIHpHO IlJIII IIpO]:\OJIbHO lI]:\y.I.QIIX BOJIOIWH, cna6meHbi IIHHepBa:qlIOH-
HblMIi IIpll6opaMII, oueCneqllDalO.l.QHMII IIX 6blCTpoe COKpall\eHlle IlJI1£
paCCJIa6JIeHlle II COOTBeTCTBeHHO 3aKpblTlie IlJIIi OTKpblTlie npOCBeTa
cocy]:\a.
OCHOBHblM iPII3110JIOrlIqeCKliM na3HaqeHlleM aTIiX CTPYRTYP, KaK
II3BeCTHO, HBJIJleTCH 06eCneqeHIIC TOH IlJIIi HHOii HanpaBJIeHHOCTIi KpO-
BOTOKa B opraHe, OTDeqalOlI\ee B Kam]:\blH ]:\aHHbIii MOMeHT nOTpe6-
HOCTHM opraHlI3I\Ia. B 3aBIICliMOCTli OT l\IeCTOHaxom]:\eHIIH ciPIIHKTe.
pa B npeKanllJIJIHpHblX apTepllOJIaX IlJIIi B CTeHRax nOCTKanIlJIJIHp-
HblX BeH OH UJIII 3a]:\epmllBaeT npliTOR RpOBIi K RanlIJIJIHpHOii CeTJI
opraHa, mIll, 3aTpY]:\HHH OTTOK, MomeT ]:\errOHlipOBaTb ee B opraHe..
$YHK:qIlOHaJIbHaJl ]:\IIHaMIiKa opraHa, HenpepblBHO MeHHlOlI\aH ero cne-
:qniPllqeclmfl MeTa6oJIII3M, Tpe6yeT COOTBeTCTBYlOlQeH ClIe:qllaJIII3a:q1I1f
185.
RpOBOTOKa, BKJIlOqeHDe IIJID BbllilllOqeHDe RanDJIJIHpHoro PYCJIa. ElI\e
Claude Bernard B JIeSIJ;HHX, qDTaUHblX B I oJIJIem ]:\e (J>paHc B 1869 r.,
IOBOplIJI 0 60JIblllOM 3HaqeHDII RanDJIJIHpHOrO KpOBOo6pall\eHDJI B op-
raHax, IIplICIIOC06JIeHHOrO ]:\JIJI BblIIOJIHeHIIJI OlIpe]:\eJIeHHblX 3a]:\R'I B
pa3HbIe B03paCTbi 11 B pa3Hble MOMeHTbl mII3He]:\eJlTeJThHOCTH opra-
HD3Ma (Bernard, 1871). OH me YKa3bIBaJI Ha TO, qTO 3HaqeHlIe
-CDCTeMbI KaIIDJIJIJlpOB, KOTOpaJl 06eCIIeqlIBaeT DnTIIMHblB: KOHTaKT TKa-
neBblX, napeHXDMaT03HbIX 8JIeMeHTOB opraH a C KpOBLlO, pa3BDBaeT-
.CH B 8BOJIIOIJ;DD nOCTelIeHHO D ]:\OCTlIraeT BbiCOKoro ypOBHJI TaM, r]:\e
]:\JIH 06eCIIeqeHDH iPII3DOJIOrlIQeCKDX nOTpe6HOCTeH Opl'aHII3Ma Tpe-
oyeTCJI MeCTHoe opraRHoe ]:\BDmeHDe KPOBIl, B KaKOH-TO Mepe He3a-
BDCDMoe OT coce]:\HDX opraHOB II OT 0611\ero MarDCTpaJIbHOrO KpOBO-
'TOKa. II03]:\Hee 8TII me BOIIpOCbl 8BOJIIOnIIII cOCy]:\IICTOH CIICTeMbI 110]:\-
P06HO pa3pa6oTaJI B cBoeH KHllre 11. 1\1. AOreJIb (1903-1904), r]:\e,
B qaCTHOCTD, ]:\aJI rJIy60KIIH aHaJID3 iPYHKIJ;IIOHaJIbHOH 06YCJIOBJ'IeH-
HOCTII CTpyKTypHblX B3aDMOOTHomeHDH COCy]:\OB B cpaBHDTeJIbHO-aHa-
TOMDqeCKOM IIJIaHe.
(J>YHKIJ;IIJI ciPlIHKTepoB Hepa3pblBHO CBH3aHa C iPYHKIJ;IIeii ]:\pyrlIx
perYJIJlTOpOB opraHHoro KpOBOTOKa - C apTepDO-BeH03HblMII aHaCTO-
M03aMD IIJID, IIHaqe, CTpYKTypaMD, 06eCIIeqlIBalOII\IIMII IIpJlMYIO CBH3L
apTepDH II BeH. B YCJIOBDHX 3aKpblTIIH IIpOCBeTa OTBO]:\HII\IIX BeH II,
CJIe]:\OBaTeJIbHO, 3aTpY]:\HeHHoro OTTOKa, apTeplIaJIbHaH KpOBb HeIIO-
cpe]:\CTBeHHO IIOIIa]:\aeT B KpynHble BeHbI II ]:\aJIee B 0611\IIii KpOBo-
"TOK, MHHYH MeCTHYIO KanDJIJIHpHylO ceTb opraHa. ApTepDo-BeH03Hble
.aHaCTOM03b1 pacnOJIaralOTCH IIJID BHeopraHHO, B oKpymalOlI\IIX cep03-
.HblX 060JIOqKaX, D Tor]:\a IIpII COOTBeTCTBYlOlI\DX YCJIOBDJlX MorYT
MeHHTb KpOBOTOK BO Bcer.1 opraHe, nepeBO]:\JI HymHoe KOJIlIqeCTBO
RpOBD B coce]:\HlIe opraHLI, IIJID HaXO]:\HTCJI BHYTPD Opl'aHa, pery-
.JIDPYH KanlIJIJIJlpHOe KpOB006pall\eHIle B RaKOii-JID60 ero qaCTD. Pe-
rYJIHIJ;DH OCYlI\eCTBJIHeTCH HepBHblMII IIpD60paMD. I1x CTpoeHIIIO, KaK
B paBHoii CTeneHD D CTpoeHDIO caMIIX apTepDO-BeH03HbIX aHaCTO-
M030B II ]:\pyrDx COCY]:\DCTblX CTPYKTYP MeCTHoro KpOB006pall\eHDJI B
'pa3HblX opraHax, 1I0CBHII\eHa 60JIblllaJl JIDTepaTypa.
B qaCTHOCTll, YCTaHOBJIeHO (Nonidez, 1942; rpDrOpbeBa, 1954;
lliy6uH, 1952), qTO apTepDo-BeH03HbIe aHaCTOM03b1 B II306IIJIIIII cHa6-
.meHbl MHKOTHblMD HepBHblMD BOJIOKHaMD qYBcTBIITeJIbHOii IIplIpO]:\bl,
KOTOpbIe o6pa3YlOT Ha HDX CJIOmHO IIOCTpOeHHbIe peIJ;eIITOpbl, D QTO
-6e3MHKOTHbIe BOJIOKHa JlBJIHIOTCJI IIOCTraHrJIDOHapHblMII OTpOCTKaMII
CIIMnaTHqeCKIIX KJleTOK, OCYlI\eCTBJIHlOlI\lIMH ]:\BDraTeJIbHbIe peaKIJ;1I1I
aHaCTOM030B.
o COCY]:\DCTblX CTpYKTypax Ha]:\nOqeqHDKa II oKpymalOlI\HX ero
opraHOB, IIMelOlI\HX OTHOllieHlIe K perYJIHIJ;IID MeCTHoro KpOBOTORa
]:\aHHoii 06JIaCTH, B JIIITepaType DMeeTCJI cpaBHDTeJILHO MaJIO CBe]:\e-
HDH. OARaKo apTeplI0-BeHo3Hble aHaCTOM03b1 onDcaHbl B IIapeHXIIMe
M03rOBoro CJIOH Ha]:(nOqeqHHKa (RpacyccKaH, 1923) D B PblXJIOH coe-
JJ;DHDTeJIbHOii TKaHD, oKpymalOlI\eH Ha]:\IIOqeqHDRH (Bennett, Kilham,
1940). A. A. RpacyccKaJl YKa3aJIa Ha Me]:\yJIJIHpHble apTepIIII Ha]:\-
nOQe"'IHIIK8., RHor,D;a IIJ:\YlI\lIe B COIIpOBOm,D;eHIIII BeH, KOTOpbIe BbIXO-
186
HT D3 CDHYCOD OB D BJIIiBalOTCH B :QeHTpaJIbHYIO BeHY I1JID aHa-.
CTOM03DPYIOT C Be.HaMD oRPymalOlI.\DX OpraHOB. IIMelOTcJI TaKme IW-
pOTKDe COCY DCTbIe CBJl3D Ha IIOqeqHDKa C PyrDMII opraHaMII
B qaCTHOCTH C 1I0qKOH (HDCeJIb, 1928; llIeH<paHH, 1949). HapH Y
C 8TDM D3BeCTHbi HelIocpe CTBeHHble aHaCTOM03bI, CBH3blBalOlI.\De IIpH-
MblMH KaHaJIaMH apTepDD C BeHaMD. 8TD KaHaJIbI BMeCTe C MHoro-
KpaTHO OIIDcaHHbIMII B Ha IIOqeqHHKaX MJIeKOIIDTalOlI.\DX rJIa KOMbl-
meqJJ:blMD c<pDHKTepaMD, paCIIOJIOmeHHbIMD B CTeHKe :QeHTpaJIbHOK
BeHbl (qnqDHa 3e, 1962B; CaIIDH, 1961, 1964) II IIPDHOCHlI.\DX co-
cy ax Ha]:(IIOQeqHDROB (llIeH<paHH, 1949), CJIymaT MexaH1I3MaMII, pe-
rYJIlIpylOlI.\IIMD HalIpaBJIeHDe KpOBOTOKa qepe3 Ha IIOqeqHHKOBbie me-
JIe3bl (CaIIDH, 1961, 1964). HaK IIpaBIiIJIO, xpoMaciJ<PDHHble KJIeTKH
He KOHTaKTDpylOT C apTepDaJIbHOH CTeHKOH, a OMblBalOTCH KpOBblO.
npOXO JllI.\eH 110 KalIDJIJIHpHOMy pyCJIy. B COOTBeTCTBDH C 8TDM, B
06b1QHbIX WD3DOJlOrDqeCIUlX YCJlOBHHX rOpMOHbI, Bbl eJlHeMble xpO-
MaW<PDHHOH RJIeTROH, IIOlIa alOT R IIpOCBeT RamlJlJIHpa II BblHOCHTCJI
n;eHTpaJlbHOH BeHOH B 06lI.\nH I{POBOTOR. <P1I3DOJIOrDqeCKDH CMblCJI
CTPYKTypHOrO KOHTaKTa (<aH oKpDHHaH KJIeTKa - KaIIDJIJIHp» pa306paH
Bblme; OH XapaKTepeH JlH BbiCOROOpraHD30BaHHbiX mHBOTHbIX, y KO-
TOpblX 8H OKpIIHHbIe ciJYHK:QDH a peHaJIOBbiX opraHOB B 60Jlbmeii Mepe,
qeM y HD3ll1DX mDBOTHbIX, 1I0AqDHeHbi HepBHblM perYJlH:QDHM. ACIIH-
XpOHHO <PYHKD;IIOHDpYIOlI.\IIe meJle3IICTble RJleTRD IIOCTOJlHHO n, BIIJJ;II-
MO, He60JIbmHMD IIOP:QDHMD IIOCbIJIalOT qepe3 KalIDJlJlHpHylO CDCTe-
MY meJIe3bl B 06lI.\DH KpOBOTOK HeKOTOpOe KOJIDQeCTBO IIPO Y:QII-
pyeMbIx elO KaTeXOJIaMIIHOB. OAHaKO IIPII CTpeCCOBbIX peaK:QlIJIX
(WII3HOJIOrDqeCKDX IIJlD 8KClIepIiIMeHTaJlbHbIX), Bb13blBalOlI.\HX 1I0BbI-
llIeHHYIO lIoTpe6HOCTb OpraHD3Ma B 8TlIX ropMOHaX, meJle3a CDHXPOH-
HO Bbl6paCblBaeT BeCb IIHKpeT, Bblpa60TaHHblH B xpOMaW<PHHHblX KJleT-
KaX. MomHo alIpHopHO IIOJIaraTb, qTO MaCCOBOe KOJIDQeCTBO Bbl]:(e-
JIHeMOrO DHKpeTa, BO-lIepBbIX, D Heo6xo lIMOCTb 6bICTpoii ero peaJlH3a-
:QDD, BO-BTOPbIX, Hym alOTcJI JlH OCYlI.\eCTBJIeHIIH <p1I3IIOJlOrlIqeCKOrO
8<p<peKTa B DHblX IIYTJlX BDmeHDH KpOBII (a BMeCTe C Heii Drop-
MOHOB), 1I0MlIMO 3aMe JIeHROrO TOKa 110 KaIIIIJlJlHpHOii CIICTeMe. 110
aHaCTOM03aM II, KOneqHO, He 6e3 yqaCTIIJI c<pIIHKTepOB, ropMOHbI MO-
rYT B 60JlbmHX ROJIDqeCTBaX IIOCTYlIaTb DJIIi B o6ID;IIii KpOBOTOK, IIJlII
B coce]:(HlIe opraHbI B 3aBIICDMOCTII OT ciJD3IIoJlOrlIqeCRHX IIOTpe6-
HOCTeii.
MHoro06pa3HbIH OTTOR KpOBD Ha IIOqeqHHROB, B qaCTHOCTD, OT
M03rOBoro CJIOH, HalIpaBJIeHHbIH, RaR MbI rOBOpHJlII, B pa3Hbie opra-
Hbl (IIOqRII, lIeqeHb - Qepe3 BOPOTHYIO II IIeqeHOQHYIO BeHbI, ceJle-
3eHRy, IIO meJIY OqJJ:ylO meJle3y, IIDID;eBO , Kap Dii meJlY Ka, IIOJlY-
JIYHHbIe Y3JIM, KOpy Ha IIOqeqHDKa), B COqeTaHDD C BHYTPHCOCY H-
CTbIMn CTpyKTypaMD, peryJlDpYIOlI.\DMH BHYTpDopraHHblii KpOBOTOK,
06eCIIeqDBalOT BlImeHDe ropMoHoB B HalIpaBJleHDD, HymHoM B Kam-
blH aHHbIH MOMeHT JlH opraHlI3Ma. ABDmOODe 8TO no CBODM <p1I-
3HOJlOrHqeCKlIM B03MomHOCTHM IIpD6JlDmaeTCJI B TaKlIX CJIyqaHX R
TI)MY, KOTopoe IIOCTOHHHO CYlI.\eCTByeT y HD3ll1DX II03BOHOqHblX (rJIaB-
HbIM 06pa30M y KpyrJIOpOTbIX D pbI6) D BbllIOJlHHeTCH y HDX Ma-
187
rDCTpaJIbHblMll COcy]:(aMD. TaKDM OOpaSOM, apTepUO-BeH03Hble aHa-
'CTOMOSbl B xpoMaiPiPDHHblX opraHax MomBO paCCMaTpDBaTb KaK aBO-
.JIIO IIOHHO BbIpa60TaHHbie CTPyKTypbI, OpDcooc06JIeHHbie AJIH HOBblX
YCJIOBuii BblBeAeHDH DHKpeTa 113 KJIeTOK, KOTOpble B opoD;ecce pas-
BIITDJI OTO]:(BDHYJIDCb OT r.mrDCTpaJIbHbiX COCYAOB D ORa3aJIIICb B KOH-
TaKTe TOJIbKO C KaODJIJIHpaMD.
C aTOH TOqKD speHDH SaCJIymDBalOT BHDr.laHIIJI HaJIllqDe apTeplIO-
BeHOSHblX aHaCTOMOSOB B Apyrux IIaparaHrJIDHX, OlIlICaHIII>IX paSHbl-
Mil aBTopaMII. Goormaghtigh II Pannier (1939) HaIllJID DX B HaA-
cep]:(eqHbIX lIaparaHrJIIIHX, B TO I MeCTe, rAe KaOIIJIJIHpHblH KpOBOTOR
..npeHDpyeTcH OTBOAH Dr.1II BeHaMD. Celestino da Costa (1944) 0011-
caJI IIX B coeADHDTeJIbHOii TKaHlI, oKpymalOII:(eH RapOTIIAHblH oapa-
raHI'JIIIH, de Castro (1951) - B Mem\HX apTeplIHX, KOTopble JlBJIJlIOT-
'CJI KOJIJIaTepaJIJlMH 60JIee KPYIIHOH apTeplID, IIIITalOlI{eH KpOBblO Ka-
IIDJIJIJlpHYIO ceTb KapOTDAHoro oaparaHrJIDJI.
l1.JIJI Goormaghtigh D Pannier (1936, 1939) 13TH IIpIICIIOC061I-
"l'eJlbHbIe MexaHDSMbI, CYlI{eCTBYIOII:(IIe HapHAY C KalIDJIJlJlpHOH ceTblO,
CJIymDJlD OCHOBaHDeM ]:(JIH ciJYHKD;JIOHaJIbHOrO paSAeJIeHDH DMD cep-
AeqHbIX IIaparaHrJIDeB Ha ABa TDIIa: Te II3 HHX, lIaparaHrJIDOHap-
Hble aJIeMeHTbI ROTOpbIX cooplIKacalOTCJI TOJIbKO C KaIIDJIJIJlpaMH (OIID-
.caHbl Pannier B aODRapAe CepAeqHblX YllIeK) , HecYT DBKpeTopHYIO
4>Y,HK IIIO, T. e. JlBJIJlIOTCH HCTDHHblMD lIaparaHrJIHHMD, aTe, KOTO-
.pble paCIIOJIOmeHbl OOA Ayroii aOpTbI (olIDcaHbl Penitschka KaK pa-
raganglion aorticum supracardiale), IIpDJIemaT K apTepDo-BeHosHblM
;8.HaCTOMosaM II HBJIJlIOTCH, KpoMe Toro, xeMope eoTopaMII. C TaKDX
:me IIOSDD;IIH paccMaTplIBaJIII 3HaqeHIIe apTepllo-BeHosHbix aHaCTOMO-
:80B de Castro (1951) II OT'laCTD Boissezon (1944).
Mbl IIOJIaraeM, qTO opnCYTCTBDe B APyrDx lIaparaHrJIIIHX, KpOMe
HaAIIOqe'IllIIKOB, apTepHO-BeBOSHbIX aHaCTOMOSOB HBJIJleTCJI He pas'b-
.eAIIHJlIOII{HM, KaR CqDTaJID Goormaghtigh II Pannier (1936, 1939),
. a 06'beAIIHJlIOID;HM IIpDSHaKOr.l AJIH iPYHKn;IIOHaJIbHOrO eADHCTB3 B
. KaqeCTBe IIHKpeTopHblx opraHoB. B IIOJIbSY aTOro me rOBopDT D rycTaH
KaIIlIJIJIJlpHaJl cen, OTMeqeHHaJl BceMII 6e8 IICKJIlOqeHDJI aBTopaMD H
_AaBmaJl IIOBOA nepBblM DCCJIeAOBaTeJIJlM K 060SHaqeHIIIO KapOTIIAHoro
TeJIa COCYAIICTbIM KJIy6KOM (Arnold, 1865). B IIpOD;ecce IICTOplI'le-
CKOI'O paSBIITHH IIaparaHI'JIDll, 0 KOTOpblX DAeT peqb, aBOJIIODpOBaJIII
B CMblCJIe ciJYHK DOHaJIbHOH 3HaqDMOCTD cBoeii HHKpeTopHoii ciJYH-
K DII, OAHaKO OHII BcerAa OCTaBaJiIICb aHAoKpDHHblMII OpraHaMH. Ha
'KaKOM-TO, OqeBIIAHO, (mpOMemYTOqHOM», aTalIe aBOJIIO IIII II03BOHO'l-
HbiX OHII IIpD06peJID apTepDOBeHOSHbie aHaCTOMOSbi AJIJI YCOBepmeH-
.CTBOBaHDJI opraHHoro KpOBOTOKa II perYJIHD;DII TpaHCIIopTa ropMOHOB.
He MeBee CJIOmHO YCTpoeHa CIICTeMa HHHepBa 1I1I HaAIIOqeqHIIKOB.
B HaCTOJl ee BpeMJI Mbl paCIIOJIaraeM OI'pOMHblM qlICJIOM pa60T,
nOCBH eHHbIX MopciJOJIOrDD HepBHblX aJIeMeHTOB HaAIIOqe'IllIIKa D aK-
.cTpaopraHHblM HepBHblM CBJl3J1M Y MJIeKOOIITalO DX I. 8KclIeplIMeH-
I JIIITepaTYPY no IIHHepBaqHH Hap;nOqe'iHIIROB CM. Y Bachmann (1954), ArapRo-
Ba (1964) II Coupland (1965a).
;188
TaJIbHble HCCJIep;OBaHlIH, HanpaBJIeHHble Ha Bt.IHeHeHlle 4>YHK IIOHaJIb-
HOrO 3HR'IeHIIJI BHYTpIlOpraHHblX HepBHbIX IIpll60pOB, CBII]:\eTeJIbCTBY-
lOT 06 06lt1IbHOii qyBcTBIITeJIbHOii II p;BllraTeJIbHOii IIHHepBa 1I1I aTO-
ro opraHa.
B a6]:\oMRHaJIbHb1X naparaHrJIlIHX Hollinshead (1936) II B M03ro-
BOM CJIoe Ha]:\IIOqe'IllIIKOB (Hollinshead, 1936; IIJIbIlHa, 1939) 60raTo
.npe]:(CTaBJIeHt.I MJlKOTHble BOJIOKHa II :ux ROH eBble OT]:\eJIt.I, nepe-
pom]:\alOI:I\lIeCJI npII aKCTlIpna 1I1I cnllHHOM03rOBbiX Y3JIOB (IiJIbIlHa,
1948) II ganglion nodosum 6JIym]:\alOI:I\ero HepBa (ArapKoB, 1964).
BoraTcTBO qYBCTBIITeJIbHblX IIplI60pOB CBII]:\eTeJIbCTByeT 0 BblCOKOM
pa3BIITIIH CHrHaJIbHOii CIICTeM.bI B aTOM opraHe.
HapHJ:\Y C qYBCTBIITeJIbHblMII BOJIOKHaMII B COCTaBe qpeBHOro Hep-
Ba II]:\YT ceKpeTopHble BOJIOKHa, IIOKa3aHHbie eI:l\e M. H. Qe60Kca-
pOB.bIM (1909, 1910); npll pa3]:\pameHIIII OHII cnoc06HbI MeHHTb xa-
paRTep ceKpe 1I1I ropMOHOB. PacnpocTpaHeHHoe MHeHlIe 0 CIIMlIaTII-
qecKoii HHHepBa II11 Ha]:\IIOQeqHIIROB nperaHrJIIIOHapaMlI, OCHOBaHHoe
rJIaBHblM 06pa30M Ha TOM, qTO He II3BeCTHa JIOKaJllf3a JI BToporo
neplI4>eplIqeCROrO HeiipOHa, B HaCTOllIII;ee BpeMR aap;o IIplI3HaTb He-
COCTOJlTeJIbHblM. PemalOI:I\YIO pont II aToM noUPOM CLIrpaJIU 9Kcne-
pllr.reHTaJIbHO-rIlCTOJIOrllqeCKlIe IICCJIe]:\OBaHHH, IIpOBe]:\eHHble B JIao()-
paTopll1l B. II. JIaBpeHTbeBa B. II. IIJIbllHOii (1939) II 1I0]:\TBep
»\]:\eHHble C HeKOTopblMII BaplIa IIHMH cD. A. AYPIl blHblM (1942) II
f. B. ArapKoBbiM (1964). B. II. IIJIbllHa B HeCKOJIbKIIX ceplIHX OlIbI-
TOB Ha B3pOCJIbiX co6aKax II HOlliKax C nepepe3Koii qpeBHblX HCp-
BOB, aKCTllplIa lIeii 1I0JIYJIYHHbiX Y3JIOB II lIepepe3Koii CIIIIHHOM03ro-
BblX KopemKOB nOKa3aJIa, qTO OCHOBHaH Macca ceKpeTopHblX Hep-
BOB Ha CBoeM IIYTII OT cnllHHoro M03ra K M03rOBOMY CJIOIO Ha]:\IIO-
qeqHIIKOB npepblBaeTCJI II qTO aKCOHbI BToporu HeiipOHa OKaHqlIBalOTCH
B meJIe311CTOii napeHXIIMe opraHa. Snl OnblTbi y6em]:\alOT B TOM,
qTO MeCTOnOJIOmeHlle BToporo HeiipOHa OqeHb Baplla611JIbHO, qTO OH
l\WmeT HaXO]:\IITbCJI B caMOM 1\103rOBOM CJIoe Ha]:\nOqeqHIIRa, B ero
KanCYJIe II B COJIHeqHOM CIIJIeTeHlI1I (pIlC. 69), a, 110 ]:\aHHblM cD. A.lly-
pll blHa II f. B. ArapKoBa, eI:l\e II B KOpKOBOM CJIoe. OH MomeT
2IemaTb B JII060M MeCTe Ha npOTHmeHIIII Bcero CIIMlIaTllqeCKOrO
CTBOJIa - OT CII:UHHOM03rOBbiX KopemKOB ]:\0 M03rOBoro CJIOH Ha]:(no-
qeqHIIKOB BKJIlOqIlTeJIbHO. HarJIJI]:\HbIM nO]:\TBepm]:\eHlleM B03MomHO-
CTII TaKHX Baplla lIii B paCCeJIeHIIH HeiipoHoB BToporo 1I0pH]:\Ka Y
B3pOCJIbiX JlBJIHIOTCJI ar.r6plIoJIorllQecKJle IICCJIe]:\OBaHIIH A. M. fOJIy6a
( 1936) , a TaKme Hallill cpaBHIITeJIbHO-aM6plloJIorllqeCKlle ]:\aHHbIC
(CMIITTeH, 1958, 1962, 1963), nOKa3aBmlle oce]:\aHHe MllrpllpYIOII:(HX
CIIMnaTllqeCKIIX aJIeMeHTOB B JII060M 113 YKa3aHHblX yqaCTKOB CIIMna-
TllqeCROrO HepBa.
HapH]:\Y C BOJIOKHaMII, KOTopble IIpepblBalOTCH Ha HeiipOHax BTO-
poro 1I0pJl]:\Ka, B. II. IIJIblIHa IIpOCJIe]:\HJIa TaKme ]:\ereHepa 1I1O He-
60JIblllOrO qllCJIa HepBHblX OTpOCTKOB BnJIOTb ]:\0 TepMIIHaJIeii M03ro-
Boro CJIOH Ha]:\nOqeqHIIKOB, HO HII pa3Y He BII]:\eJIa IIepepom]:\eHIIJI
nx OROHqaHlIii Ha xpoMa4>4>IIHHblX KJIeTKax. ITo MHeHlI10 aBTopa, ]:\0
Tex 1I0p nOKa TaKlle OKOHqaHIIH He 6Y]:(YT Haii]:\eHbI, IIpem]:\eBpe-
189
P II c. 69. Cxe!lla pacnoJIome-
RlfH CIIMnaTlI'IeCKlJX Heii:poHoB
BTOpOro nOpHlJ,Ka, IIHHepBnpylO-
II{IIX M03rOBoe BeTI{eCTBO Hap;-
nO'Ie'lHlfKa MJIeKOnmalOTI{IIX
1 - npel'aHI'JIHOHapHoe BOJIOI<HO II
Het1:pOHY, aa.nel'alOIl\eMY B MOal'OBOM
BeIl\CCTBe HagnOqeqmma; 2 - TO me,
aa.nel'alOII1cMY B I<ancyJIe H8AnO'lC'l-
HHI<a; 3 - npel'aHI'JIHOHapHoe BOJIOR-
HO R Het1:poH)', S8JIel'alOIl\eMY u 110-
JIYJIYHHOM l'aBI'JIHH (I1JIbHHa, 1939).
MeHHO rOBOpIlTb 0 HaJIIIqIlIl npHMOii I1HHepBa IIII xpoMaciJqJHHHblX
KJIeTOK nperaHrJIIIOHapaMII. B IIOCJIe]:(YIOl I1e rOAbI HII caMoii B. II. I1JIb-
I1HOii, HII .J];pyrIlMII aBTOpaMII nOAo6HbIe OKOHqaHIIH He 6blJIII 06Ha-
pymeHbI. Pa60Tbi Coupland ( 1965), nORa3aBmero TlInlIqHbIe aciJciJe-
peHTHble HepBHbIe OKOHqaRIIH Ha KJIeTRaX M03rOBoro CJIOH Ha.J];no-
qe'lHIIKOB RpblC, Coupland II Weakley (1970) Ha KpOJIIIKaX HOCJlT
OIIIICaTeJIbHblH xapaKTep II He pellialOT BOlIpoca 0 IIplIpO.J];e BOJIOKOH.
C KOTOpblMII OHII CBH3aHbl.
IIpII II3yqeHIIII I1HHepBa I10HHbIX OTHOllieHIIH B KapOTII]:(HOM na-
paraHrJIIIII CJIe.J];yeT Y'IHTblBaTb onbiT mCIIepIlMeHTaJIbHOrO II3yqeHIIH
HepBHblX CBH3eii B M03rOBOM CJIoe Ha.J];IIOqeqHIIKa. TaK me KaK II
B nOCJIe.J];HeM, nperaHrJIIIOHapHbIe BOJIOKHa, I1.J];Y.I.qIle K KapOTIIAHOMY
IIaparaHrJIIIIO, MorYT, He AOCTIIrHYB ero, npepblBaTbCJI B m060M MeCTe
110 XOAY CIIMIIaT eCHoro rrepBa. I1x coxpaHHocTb IIJIII, Ha060poT, ne-
pepom.J];eHlIe IIpli 8RcTlIplIa I1H BepxHero llIeiiHOrO Y3JIa 1I0ABepme-
Hbl 60JIbllllIM IIH.J];I1BII.J];yaJIbHbIM KOJIe6aHIIJlM. B pa60Te Lever' et a1.
(1959) IIOCJIe IIpOlI3BeAeHHoH aBTopaMII CIIMIIaT8KTOMHII MOrJIlI .J];e-
reHeplIpOBaTb TOJIbRO IIperaHrJIIIOHapHbIe BOJIOKHa II OCTaBaJIMCb I1H-
TaKTHblMH OKOHqaHIIJI Ha RJIeTKaX KapoTII.J];lIoro lIaparaHrJIIIJI. B 8K-
clIepIlMeHTe Biscoe, Stehbens ( 1967) , OqeBII.J];1I0, 60JIbmlIHCTBo BO-
JIOKOH OKa3aJIOCb 1I0CTraHrJIIIOHapHbiMII (TeJIa IIX HeiipoHoB paCIIOJIa-
raJIIICb B ganglion cervicalis superior) , II aBTopbl Ha'6JIIO.J];aJIII IIX
.J];ereHepa I11O B KapOTJ1.J];HOM lIaparaHrJIJ1II.
Pe3YJIbTaTbi 8KcneplIMeHTaJIbHOrO IICCJIe.J];OBaHIIJI B. II. I1JIbIlHOii.
YTBepm.J];alO.I.qlle nOCTraHrmWHapHYIO CIiMnaTIIQeCI{YIO IIHHepBa 1I1O
xpoMaciJciJIIHHblx RJIeTOK Ha.J];IIOqeqHIIKa, BamHbI AJIJI Hac e.I.qe II B
APyroM CMblCJIe. OHII IIMelOT npHMoe OTHOllieHIIe R TeM IIOJIOme-
HIIHM 0 rJIHaJIbHOH IIpllpO.J];e xpoMaciJWIIHHblX aJIeMeHTOB, KOTopble
pa36IIpalOTCJI BO BTOpOH qaCTII HallieH pa60TbI II KOTopbIe HeO]:(HO-
KpaTHO HaMIl BblCKa3bIBaJIIICb (CMHTTeH, 1958, 1962, 1963, 1964).
XpoMaciJciJHHHaJl KJIeTRa IIHHepBIIpyeTCJI nOCTraHI'JIIIOHapHblMII BOJIOK-
HaMil CIIMnaTHqeCKOH aepBHOH CIICTeMbl. OHa, CJIe.J];OBaTeJIbHO, TaK
190
me RaK Bce OCTaJlbHble TKaHeBbie :meMeHTbI BHYTpeHHUX opraHOB,
nOJlyqaeT BOJlOKHa OT ]:(BYXqJleHHOro s«lJ«lJepeHTHoro 3BeHa BereTaTIIB-
HOll HepBHOll CIICTeMbi. TaKaJI lIwp«lJoJlOrlIqeCKaJI opraHlI3an;IIJI HepB-
HbiX CBJI3ell JIBJlJIeTCH O]:(HIIM 113 apryMeHToB B nOJlb3y Toro, qTO
xpoMa«lJ«lJIIHHaH MeTKa IIpOU30lllJla B npon;ecce IIH]:(IIBII.n;YaJIbHOro II
IICTOplIqeCKOro pa3BIITUJI He II3 HenpOHa (Hellp06J1aCTa). ECJlU me
npU3HaTb ee HellpOHHoe npOlIcxom]:(eHue, Ha]:(o ]:(onYCTUTb cy ecTBo-
BaHlIe TpeXqJleHHOro s«lJ«lJepeHTHoro 3BeHa CUMnaTlIqeCKOll HepBHon
CIICTeMbl. TaKoe .n;ony eHue npoTUBOpeqlIJlO 6b1 OCHOBHblM IIpUHn;II-
naM CTpOeHIIJI BereTaTUBHOll HepBHOll CHCTeMbi II COCTaBIIJlO 6b1 IIC-
KJlIOqeHlIe, JlOrlIqeCRII He YKJla]:(blBaIO eeCJI B 06 yIO cxeMY BHHep-
Ban;IIOHHblX OTHomeHUll BHYTpeHHIIX opraHOB. AJlJI naparaHrJllIeB,
II B qaCTHOCTU ]:(JlJI Ha]:(IIOqeqHIIKOB, CJle]:(OBaJlO 6b1 Tor]:(a AonYCTIITb
cy ecTBOBaHlIe Oc06b1x, TOJlbKO IIM OAHIIM CBonCTBeHHLlX HepBHblX
CBJI3ell, He OIIpaB]:(aRHOe HII «lJYHKD;IIOHaJlbHblMU nOTpe6HOCTHMH amx
()praHOB U HUKaRHMII ]:(pyrIIMu co06pameHUJlMU.
IIepequcJleHHble «lJaKTbI II BblTeKaIO ue II3 HIIX BblBO]:(bl n03BO-
JlHIOT HaM c.n;eJlaTb 3aKJlIOqeHlIe, qTO BCJI naparaHrJLIIOHapHaJI TKaHb,
pa36poCaHHaJI B TeJle MJleKOnIITaIO IIX B BH]:(e MeJlKBX B KpYIIHblX -
KJleTOqHblX CROIIJleHlIll, npllHap;JlemlIT R e]:(RHOll xpoMa«lJ«lJuHHoll cn-
-eTeMe, «lJYHKD;IIOHaJIbHO npe]:(Ha3HaqeHHOll ]:(JlJI Bblpa60TRH B Bbl]:(eJle-
HWI KaTeXOJlOBblX aMUHOB. Cpe]:(II IIaparaHrJLIleB OCHOBHaJI IIHRpe-
TOpHaJI «lJYHKD;IIJI IIplIHa]:(JlemlIT M03rOBOMY CJlOIO Ha.n;nOqe1J.HIIKOB RaK
BblCORO CIIen;UaJlII3UpOBaHHoMy opraHY C BHYTpeHHell CeKpen;IIell. Oc-
TaJlbHble MeJlKlIe BHeHa]:(IIOqeqHUKOBble naparaHrJlIIU, KOTopble pacno-
JIOmeHbI B RpynHblx pa3BeTBJleHIIJIX aopTbI U nplIcep]:(eqHOll 06J1aCTH,
.n;OCTlIrJlII Ha ]:(aHHOM STaIIe 8BOJlIOn;III1 MaKCIIMaJlbHOll Mop«lJOJlOrlI-
qeCKOll PY]:(UMeHnUl;IIII U «lJu3IIoJlOrlIqeCKOll pe]:(YKn;IIII (KapOTB]:(Hblll
naparaHrJllIll, JleroqHO-aOpTaJlbHble U HeKOTopble p;pyrlIe). fpynnbl
naparaHrJlUOHapHblX KJleTOK, KOTopble BCTpeqaIOTCJI B COCTaBe HepB-
HblX CTBOJlOB, raHrJllIeB II npeTepMlIHaJlen, JIBJlJIIOTCJI conYTCTBYIO-
ID;BMII aJleMeHTaMlI BereTaTIIBHOll HepBHon CIICTeMbI. OHII paCCeJlJIIOT-
CJI BP;OJlb HepBOB B neplIo]:( UX pOCTa II BMeCTe C HUMB MorYT P;O-
CTuraTb IIeplI«lJeplIqeCKIIX opraHOB. 8TII rpynnbl He npUHap;JlemaT K
pY]:(BMeHTapHblM meJle3IICTblM 06pa30BaHIIJIM, a nOJIBUJlIICb II paSBII-
JIIICb B 8BOJlIOn;UB nOCTeneHHO,- IIapaJlJleJlbHO COBepmeHCTBOBaHUIO
CIIMnaTlIqeCKOll HepBHOll CIICTeMbi. CpaBHUTeJlbHO-MOIXPOJlOrlIqeCKlIe
AaHHble II 8KcneplIMeHTbI Ha l.meKOnlITaIO X nORa3b1BaIOT, qTO IIH-
HepBan;IIJI IIaparaHrJlIIOHapHOll TlmHII ocy ecTBJlJIeTCJI nOCTraHrJlIIO-
HapHblMII BOJlOKHaMII CIIMIIaTII'lf!CliOrO HeiipOHa.
qACTb BTOPAR
OHTOfEHE3
CMMTIA TO-A,1],PEHAJIO Bon CMCTEMbI
r.rtU6U VII
PA3BHTHE ITAPArAHrRHEB
H ITPOHBREHHH PERAITHTYRHJJ;HH
B IIepBOH qaCTU 6b1J1H onucaHbl Te Mop«lJo«lJu3IIoJlOrUqeCRUe «lJaK-
Tbl, KOTOpbIe xapaKTepu3YIOT Opl'aHbl IIaparaHrJlUOHapHOH CUCTeMbl Y
B3pOCJlbiX oco6eH pa3Hbix rpynn II03BOHOqHbIX. BblJlO nORa3aHO 3Ha-
qeHHe cpaBHUTeJlbHo-Mop«lJoJlOrUqeCKOrO no]:(xo]:(a K u3yqeHUIO :3Toii
CUCTeMbI ]:(JlH OCMbICJlUBaHUH II On;eHKu ee y MJleKOnHTalO ux mIIBOT-
HblX II qeJlOBeKa. PaCCMOTpeHue CTPYKTYPHblX oco6eHHoCTeH U «lJYHR-
n;UOHaJlbHOH aKTUBHOCTU IIaparaHI'JlUeB C II03IIn;UH UX UCTOpUqeCKO-
ro pa3BuTuH IIoKa3blBaeT, qTO II3MeHeHUH rlIcTOJlOrlIqeCKOH CTPyKTYPbl
II B3auMooTHomeHuH C oRpymalO uMII TKaHHMU II opraHaMII HBJlH-
IOTCH OTpameHueM IIplIcnoco6uTeJlbHOH :3BOJlIOn;UIl BHYTPU rpynnbl
n03BOHOqHbIX mUBOTHbIX.
ITYTb Mop«lJoJlOrHqeCKOH :3BOJlIOn;IIH xpoMa«lJ«lJHHHoH CHCTeMbI Ha-
meJl OTpameHue H B IIpOn;ecce OHToreHe3a naparaHrJllIeB. Xopomo
U3BeCTHbi II3MeHeHUH xpoMa«lJ«lJIIHHoH TKaHII no KOJlUqeCTBY, CTPYK-
'type II rlICTOXUMUqeCRUM CBOHCTBaM B IIeplIO]:(bl 3apoAbimeBoro II
nOCTHaTaJlbHOrO pa3BUTUH opraHlI3Ma. ECJlII nO]:(OHTII R UX paCCMOT-
peHlI10 C :3BOJlIOn;UOHHbIX n03IIn;UH, TO OHU HaXO]:(HT y6e]:(IITeJlbHOe
06'bHCHeHlIe KaK npU3HaRU peKanIITYJlHn;IIII. H3MeHeHIIH, ROTophle B03-
HURalOT B «lJUJlOreHe3e, HBJlHIOTCH pe3YJlbTaTOM 1I3MeHeHIIH B OHTO-
l'eHe3e, npOIICXO]:(H UX Ha pa3HbiX CTa]:(IIJlX pa3BIITIIH II Be]:(y 1IX
R YKJlOHeHUIO OT Toro IIYTII, KOTOpblM XapaRTeplI30BaJlCH OHToreHe3,
II 3aKpeIIJlHIOTCH B qJHJlOreHe3e. MH]:(UBII]:(yaJlbHOe pa3BIITIIe oc06n
(OHToreHe3) CKJIa,D;bIBaeTCH 1I3 ]:(BYX OCHOBHbIX npOn;eCCOB, Hepa3pbIn-
HO CBH3aHHbIX ]:(pyr C ]:(pyroM U Uffi' UX IIapaJlJleJlbHO ]:(pyr APyry:
nOBTopeHUH (peKaIIUTYJlHn;UH) CTpoeHUH npe]:(KOB, rJlaBHbIM 06pa30M
3TanOB IIX pa3BUTUH, II II3MeHeHUH npucnOC06UTeJlbHOrO xapaKTepa
(a]:(anTan;IIH) R YCJlOBHHM cy ecTBOBaHUH. Bce U3MeHeHIIH, KOTopble
B03HURaIOT B OHToreHe3e U 3aKpenJiHIOTCH B 3BOJlIOn;IIII, HBJlHIOTCH
a]:(anTUBHhlMII (illMaJibraY3eH, 1940). CJle]:(oBaTeJlbHO, nOHBJleHUe HO-
BblX CBOHCTB IIJlU Mop«lJoJlOrUqeCKIIX CTpyRTYP y B3pOCJlhlX oco6ei'1
HBJlHeTCH CJleACTBlIeM n;eJloro pH]:(a OHToreHeTlIqeCKIIX nepecTpoeK.
192
3HaKOMCTBO C UUMU ]:(OnOJlHHeT Te npe]:(CTaBJleHlUl, KOTOpbIe CKJla-
p;bIBaIOTCH npH u3yqeHIIII qmJloreHUII MeTOp;OM CpaBHHTeJlbHOll MOp«lJO-
JlOrlIlI.
]J,JlH n03HaHUH 3aKOHOMepHocTeii: aBOJlIOn;IIII IIaparaHrJlIIOHapHoii:
CIICTeMbl 06a npon;ecca, COBeplllaIO IIeCH B OHTOreHe3e - peKanUTY-
JlHn;IIH U a]:(aIITan;IIH, IIHTepeCHhI U BamHhI B O]:(UUaRoBoii: CTeIIeHII.
OUU IIO]:(KpeIIJlHIOT U B II3BeCTHoii: Mepe IIpOBepHIOT Hamu 3HaHIIH 0
IIYTHX «lJIIJlOreHe3a a]:(peHaJlOBOll CIICTeMhI II cnoc06CTBYIOT yrJly6J1eH,.
HOMY auaJlU3Y 3aKOHOMepHocTeii: aToro npon;ecca.
IIccJlep;omwuH OHToreHe3a ap;peHaJIOBOll CIICTeMhI Y n03BOHO'IHLlX
mlIBOTHhlX Ha'laJIIICL no c eCTBY Torp;a me, Ror]:(a nOJlBIIJlIICL CBe-
]:(eHIIH 06 OTp;eJIbHbIX opraIlax aTOll CIICTeMhI. ABTophl, KOTophle p;aJlH
IIepBbIe KpaTKlIe II IIOCJlep;yIO ue 60Jlee no]:(p06Hble OIIlIcaHIIH napa;-
raHrJlIIOHapHblX opraHOB, ux cBH3eii: C CUMnaTlIqeCROll II cOCYP;IICTOii:
CUCTeMaMU y B3pOCJlhlX oc06ell n03BOHOqHhlX llillBOTHblX, 6h1J1II U
nepBblMII IICCJlep;OBaTeJIHMII aM6puOHaJiLHOrO pa3BUTUH. OP;HU II3 HIIX
TOJIbKO IIOIIYTHO KaCaJllICb aM6plIoreHe3a, p;pyrue II3yqaJlII ero cne-
n;IIaJlbHO, HO npelIMy ecTBeHHO y BhlcmlIX II03BOHOqHhlX II MJleKOnlI-
TaIO UX, TpeTbII - Y HU3ll1HX UJlII B cpaBUHTeJIbHO-aM6puoJlOrlI'IeCKOM
IIJIaHe y pa3HbiX rpyrrn II KJlaCCOB n03BOH01JHbIX. R HaCTOH eMY
BpeMeHH HaKOnUJlOCb orpOMHoe qUCJlO pa60T, p;06h1Thle «lJaKTbl II KOH-
KpeTHhle pe3yJIbTaThI KOTOphlX BO MHorOM nOBTOpJlIOT ]:(pyr p;pyra.
TIpe]:(MeTOM aTUX pa60T cJlymaT n;UToreHe3, rUCToreHe3, opraHoreHea,
a B 0606 eHIIHX II CBO]:(Kax II BOIIpOChI CIICTeMoreHe3a. IIOCJIep;HlIll,
IIpaB]:(a, 60Jlbme paCCMaTplIBaeTCH B npemHUX IICCJle]:(OBaHIIHX CTaphlX
aBTopOB II OTHOCIITCH R KOHn;y npomJloro IIJIU Ha'laJlY TeKy ero
CTOJleTRH.
OCHOBHbIe BOIIpOChI, IWTophle CTaBHJlHCb UCCJle]:(OBaTeJlHMH B pa3-
Hble rO]:(LI, H Te OCHOBHhle UTorn, «lJaKTH'IeCKaH CTopOHa KOTOphlX
]:(aeT YBepcuuOCTb B ee ]:(OCTOBepHOCTU, paCCMOTpeUhI B HacToH eii:
pa60Te. Becb al\mUpnqeCKuii: l\IaTepnaJi HcnoJlb3yeTcH npUMeHUTeJlbHO
R TeM 3a]:(aqaM, ROTophle IIOCTaBJleHhI B uaCTOH eM IICCJle]:(OBaHIIII,
II TpaKTyeTcH C Y'leTOl\1 Tex MaTepHaJlOB no q,1IJ10reHUU, KOTophle
1I3J10menhl Bblllle, a TaKme c06CTBeHHhlX ]:(aUHhlX no OHToreHe3y,
nOJlyqeHHblX ua pa3HblX II03BOHO'lHhlX,- npeHMy ecTBeHHO Bhlcmux.
Xopomo II3yqeHHOe pa3nUTHe Ha]:(nO'le'lHhlX meJIe3 y Bhlcmnx
n03BOHOqHbIX, MJleKOnnTaIOIIJ;HX II qeJloneKa (Mitsukuri, 1882;
H. Rabl, 1891; Zuckerkandl, 1901, 1912; Wiesel, 1902b; Soulie,
1903; Kohn, 1903, 1930; Poll, 1906; Hays, 1914; Goormaghtigh,
1914, 1921; Celestino da Costa, 1917, 1926; Weyman, 1922; Rau,
Johnson, 1923; Hammar, 1925; Hett, 1925; Kohno, 1925; E. Miller,
1926; Wrete, 1927; Keene, Hewer, 1927; IIBaHoB, 1930; Brauer,
1932; rOJIy6, 1936; Coupland, 1952, 1956, 1965; Crowder, 1957,
CMRTTeH, 1958, 1962; CMHTTeH, MaJiIIHOBCKaH, 1960; Adams, 1958;
KpYMUHH, 1960; CeMamKeBII'I, 1960; KOBpnmRo, 1962a, 6, 1970;
Kan;HeJlbCOH, 1965, 1966a, 6; 1967a, 6, 1969; Giordano-Lanza, 1965)
npe]:(CTaBJIHeT C060ll pH]:( IIOCJlep;OBaTeJlbHbIX aTanOB rUCToreHe3a KOp-
ROBoro II M03rOBoro CJlOeB, B TeqeHlIe KOTOphlX R n;eJlOMHqeCKoii: aaK-
7 H. A. CMH'rTeH
193
JIa]:(KC 6Y]:(YIII;Cii ROpbl nO]:(XOp;HT 6pIOffiHblC OT]:(CJIbl CIIMIIaTUqCCKOH
CIIO'IKU II MlIrpUpYIOIII;UC U3 HIIX napaOpTaJIbHblC KJICTOqHblC KOM-
IIJICRCbI. BHaqan:c, nplI6JIIImaJlCb R KOPROBOMY 3aqaTKy, OHII TOJIb-
KO npUJICmaT R cro MCP;IIOp;Op3aJIbHOH nOBcpXHOCTlI, a IIOTOM BHCP;-
pHIOTCH B ero CHTpaJIbHblC o TP;CJIbi. IIp0]:(oJImuTcJIbHOCTb U MOp«lJO-
JIOruqeCKHe oc06eHHOCTII :3Toro IIpOn;ecca BapLIIPyIOT Y pa3HblX IIpC]:(-
CTaBIITCJICn MJIeKOnUTaIOIII;ux II COOTBCTCTByIOT JIII60 npO]:(OJImUTCJIb-
BOCTU sM6plIOHaJIbHOrO IIepuo]:(a, JIII60 - qTO qaIII;c - OXBaTblBaIOT
pa3HbIe OTpC3KU nOCTSM6puOHan:bHoro OHTorCHC3a.
COOTBOCIITCJILHOC «lJopMlIpoBaHUC ROpKOBOH II M03rOBon TKaHCH
Hap;nOqC1JHblX mCJIC3 B OHTorCHC3C MJICKOnIITaIOIII;IIX (u IITU ) B II3-
BCCTHOH CTCIICBU nOBTopHCT STOT npo ccc B «lJUJIOrCHC3C TCX n03-
BOBOqHblX, Y KOTOpblX XapaKTCpBO IICpBOHaqaJIbHOC pa3]:(CJICBIIC ROpbl
II M03ra II IIOCJICp;yIOlI1ce IIX CJIIIHHUC: Y pbl6 OHU II30JIHpOBaHbI
]:(Pyr OT p;pyra, y aM«lJu6IIH II pcnTllJIuii cOIIpuKacaIOTcH, a y MJIC-
ROnUTaIOIII;ux M03roBoH CJIOH IIOJIBOCTbIO oRpymcH KOpOH. Hclt
(1925) U HCKOTOpblC p;pyrlIc IICCJICp;OBaTCJIII BIIp;CJIII B STOM npH3Ha-
Ke pa3BIITIIH Ha]:(IIOqCqBblX mCJIC3 Y MJICKOIIUTaIOIII;IIX mlIBOTHbiX 0
qCJIOBCKa xopomcc nO]:(TBcpmp;CHIIC 6IIorCBCTUqCCKOrO 3aKOHa.
rOpa3p;0 60JIee IIHTepeCHO U BamHO 6b1JIO 6b1 p;JIH II3yqCBIIH co-
OTHomeBIIH «lJIIJIOrCHC3a II OHTorCHC3a cpaBHIITL sM6plIOHaJIbHOC pa3-
BIITUC p;pyrlIx opraHOB IIaparaBrJIIIOHapHOH CUCTCMbI II COIIOCTaBUTb
UX C cpaBHIITCJILHO-aHaTOMlIqCCKHMU p;aBHblMII. TaKoc cpaBHCHIIC Mor-
JIO 6b1 ]:(aTb 60JICC qCTlWC npCp;CTaBJICHIIC 0 «lJIIJIOrCHCTlIqCCRUX npc-
06pa30BaHIIHX CIICTCMbl, 06JICrqUTb II nOMOqb YCTaHOBJICHIIIO rOMO-
JIOruqHOCTII CTPYRTYp. R COmaJICHIIIO, HC]:(OCTaTOqHOCTb CBC]:(CHIIH 0
pa3BIITIIII opraHOB naparaHrJIIIOBapHOH CUCTCMbI HC ]:(aCT B03Mom-
HOCTII npOBCCTII ]:(eTaJIbHOrO aHaJIU3a II Cp;CJIaTL 0606III;CHUC B STOY
SaIIpaBJICHUU. ,
HCCMOTPH Ba 60JIbmOH HHTCpCC, npOHBJICHHblH K II3Y1JCHUIO sM6-
plIorCHe3a IIaparaHrJIIICB, II 60JIbmOC qlICJIO paooT Ha STY TCMY, IIX
06'bCKTaMU CJIymIlJIO OrpaHUqCHHOC qUCJIO IIaparaBrJIIIOHapHblX CTPYR-
TYp. RaK IIpaBUJlO, STO 6bIJIU KapOTU]:(HbIH U 6pIOmHOH napaaOpTaJlb-
HblH IIaparaBrJlIIH. Bblro]:(HO OTJIUqaIOTCH B STOM OTHomCBlI1I pa60Tbi
H. M. ROBplImKo (1962a, 6, 1970), ROTopaH II3Y1JaJIa pa3BIITUC M03-
rOBOro CJlOH H&p;n01Je1JHIIKOB, 6pIOmHoro aOpTaJlbHOrO, Hap;ccp]:(CqHhIX
II RapOTup;HblX naparaBrJlUCB y 'ICJlOBCRa U JJa60paTopHbiX mOBOT-
HblX B cpaBnUTCJlbHOM acnCKTC.
RpOMC TOro, BCT CKOJlbKO-HII6Y]:(L CHCTCMaTHqCClmX HCCJlC]:(OBa-
Bun OHTorenC3a y pa3HblX IIpcp;CTaBIITCJIcii II03BOBOqHbiX mUBOTBhIX.
T. C. BCT cpaBHIITCJlbHOH SM6plIOJlOrUH CIICTCMbI, KOTopaH OXBaTblBR-
JIa 6b1 BCC ypOBBU pa3BIITUH n03BOBOqHbiX II BCC CTPYI{TYPbl, BXO-
P;HIII;I1C B UX COCTaB U nOCTCIICHBO SBOJlIOUpYIOIII;uc. Mcmp;y TCM U3-
BeCTHO, HaCKOJIbRO BamBO cpaBHCHuc 3apop;blmCH Bau60Jlcc OT]:(aJlCn-
HbiX p;pyr OT p;pyra «lJ0PM, y KOTOphIX Ha paHHlIX STanax lIH]:(IIBII-
aJlbHOro pa3BIITIIH MomBO BaHTII 60JlbmC qCPT Cxop;CTBa U, CJlC]:(O-
BaTCJlbBO, JIcrqc YJlOBIITb rOMOJlOrlIIO CTpyKTYP, qCM Y B3pOCJlblX,
p;aJlCKO pa30mC]:(IlIUXCH B «lJuJlorCHCTlIqCCKOM pa3BHToH.
194
He MeHbmee 3Ha1JeHlIe UMeJIO 6b1 II3yqeHUe MJIeKOIIUTalO UX, ux
1I0]:(KJIaCCOB, OTpJIP;OB, ceMellcTB II BIIp;OB. 3HaHIIe 8M6puoreHe3a MJIe-
KonIITalO UX, KaK Hau60JIee pa3BIIToll «lJOPMbI II Hau60JIee YP;aJIeH-
HOll OT CBOIIX IIpe]:(KOB, Tome 3HaqlITeJIhHO 06J1erqlIJIO 6bI 3 aqy.
IIocJIe]:(oBaTeJlbHoe II3yqeHUe pa3BIITIUI OT paHHIIX P;O n03]:(HUX CTa-
]:(Ull II03BOJIUJIO 6b1 BbiJIBIITb IIPU3HaKlI, Han:60Jlee XapaKTepHble ]:(JIJI
OT]:(eJlbHblX rpYIIn BHYTPU 8Toro KJIaCca.
CKY]:(OCTb Hamux 3HaHUll no 8M6plIoreHe3Y IIaparaHrJIUeB He JIU-
maeT, O]:(HaKO, npaBa npOaHaJlU3IIpOBaTL II3BeCTHbie «lJaRTbi II cp;e
JIaTb HeKOTopble BbIBO]:(bI 06 OHToreHeTUqeCKHX 3aKOHOMepHOCTJIX,
HMelO UX OTHomeHue K «lJIIJIOreHe3Y. 06 IIM IIpU3HaKOM, xapaRTep-
HLiM P;JlJI Bcex 6e3 IICRJIlOqeHUJI 3apoAblmeii: MJIeKOIIIITalO IIX (II IITHn;) ,
JIBJlJIeTCJI pa3BIITue Ha BeHTpaJIbHOll IIOBepXHOCTII 6plOmHoll aopTbi II
ee BeTBJIX 06HJIbHOrO CKOIIJIeHUH xpoMa«lJ«lJHHHoii: TKaHII, Ha3b1BaeMO-
ro (<opraHOM QYRepRaHAJlJI», a TaRme npeaOpTaJIbHblM, IIJIU napa-
aOpTaJlbHbIM 6plOmHblM naparaHrJIlIeM (Zuckerkandl, 1901; Kohn,
1903; Vincent, 1910; .HXOHTOB, 1913a, 6; Celestino da Costa, 1926;
l1BaHoB, 1930, 1945; Coupland, 1952, 1956, 1960, 1965; CMHTTeH,
1958, 1962; ROBpumKo, 1962a, 6, 1970; Lampinen, 1964, U MH. p;p.).
Pa3BIITIIe 8Toro opraHa HalI60J1ee nOAP06HO II3yqeHO Y Jla60paTop-
HbiX H p;OMamHUX mUBOTHblX, y 06e3bJlH U qeJIOBeKa. fpaHUn;bI ero
paCIIpOCTpaHeHUH, KOJIlIqeCTBO TRaHeBbiX KOMnJIeRCOB, BXOp;JI UX B em
COCTaB, «lJoPMa OCTpOBKOB II IIpOTHmeHHOCTb BP;OJlb aopTbi pe3KO pa3-
JlUqalOTCJI y pa3HbiX rpynn mIIBOTHbiX. Y KOllleK, HaIIpUMep, 8TO
IIpO]:(OJIrOBaTbIe, HUTeBII]:(Hbie TeJlbn;a, KOTopble npOCTUpaIOTCJI OT Me-
CTa ]:(eJIeHIUI aopTbI, Y c06aKII - 8TO HenapHblll opraH, KOTOpblll ne-
peXOp;IIT Ha art. mesenterica inferior, Y RpOJIIIROB - BepeTeHo06pa3-
Hble TeJla, RpaHlIaJIbHblll KOHen; KOTOpblX Bpe3aeTCJI B KOPKOBYIO
qaCTb Ha]:(nOqeqmlKa U IIOCTeneHHO UM oKpymaeTCJI. B IIpOTIIBOno-
JlOmHOCTb KpOJIUKaM y KpbIC U Mblmell 6plOmHble IIaparaHrJIIIU JIemaT
TOJIbRO pJI]:(OM C 3aqaTROM KOpbl, He B]:(aBaJICL BHYTPb ero, a 06pa-
30BaHIIe M03rOBoro CJIOJI Ha]:(nOqeqHUROB Y HIIX npOIICXOAHT nYTeM
Murpan;UII KJleTOqHblX 8J1eMeHTOB B 3aqaTOR KOpTIIRaJIbHOrO CJlOJI.
Y qeJlOBeRa npeaOpTaJIbHble IIaparaHrJIUII JIemaT 110 60KaM aopTbi
B Bu]:(e CIIMMeTpUqUblX AJlUHHblX TeJI U CIIYCKalOTCJI ]:(0 MeCTa ]:(e-
JIeHUH aopTbI. HaK npaBIIJlO, y Bcex 3apo]:(blmell MJIeKOIIIITalO IIX
Bblme II HlIme OCHOBHbIX IIaparaHrJIUOHapHblX TJImell, KOTopble COCTaB-
JIHIOT c06CTBeHHO opraH QYKepKaH]:(JlJI, BCTpe1JaIOTCJI MHOr01JUCJIeH-
Hble MeJIKUe naparaHfJIHOHapHble rHe3Aa, paCCeJIHHble B BIIp;e MeJI-
KHX rpYIIn UJIH OTAeJlhHblX RJIeTOK (pIIC. 70).
Y 60JIbmUHCTBa MJleKOIIUTaIO UX B TOT neplIO]:(, KorAa Macca
xpoMa«lJ«lJIIHHblX rHe3A pa36pocaHa B 6plOmHOll 06J1aCTII,- B rPY]:(HOll
IIOJIOCTII HX OqeHb MaJlO. MCKJllOqeHlIe COCTaBJIJIIOT HII3mue nJlan;eH-
TapHble - HaceKOMOHAHble II PYKOKpblJIble, 0 ROTOpblX CKa3aHO HlIme.
M3BeCTHO, 1JTO opraH QYRepKaHAJIJI ,He IIOCTOJIHHOe 06pa30BaHH9.
Y 60JIbmlIHCTBa MJIeROnUTalO IIX OH xopomo pa3BUT TOJILKO B 8M6p1I0-
HaJlbHOM IIeplIOAe, a 3aTeM nOCTeneHHO perpeccupyeT. CpOKH, B Te-
qeHlIe KOTOpblX npOIICXO]:(IIT ero 06paTHoe pa3BIITIIe, nOABepmeHLI
7* iDS
D!2J .5
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MN NN
P H C. 70. Pacnpep;eJIeHue aop;OMllHaJIbHOll xpoMaCPCPuHHoH TKaml Y 3apop;bIIIIeii
lJeJIOBeKa (peKoHcTpYK IH) (Coupland, 1965)
60JIbIDIIM He TOJIbKO BII]:(OBblM, HO U HH]:(UBII]:(yaJIbHblM KOJIOOaHIIHM.
Y HeKOTopblX mlIBOTHblX OH perpecclIpyeT KO BpeMeHU pom]:(eHUJI,
y ApyrIIX - B HaqaJIe IIOCTHaTaJIbHOrO OHToreHe3a, y TpeTbUX, na-
nplIMep y qeJIOBeKa, 6proIDHOii IIaparaHrJIuii HaqUHaeT nOCTeneHHO
aTpoqmpoBaTbCH co 2-ro ro]:(a BHeYTp06Hon mU3HH U CTaHOBUTCJI
MaRpOCKOIIlIqeCKU Hepa3JIIIqlIMbiM R neplIo]:(y nOJIOBOn 3peJIOCTII (.HXOH-
TOB, 1913a, 6). 110 AaHHblM Wiesel (1902b),OH HCQe3aeT R 10 ro-
AaM, no r. <I>. MBaHoBY - K 15 ro]:(aM, a Zuckerkandl 06uapymuJI
eFO OCTaTRH y 39-JIeTHen meHl UHbI. H. M. ROBpuml\O (19626) OIIH-
c JIa Ha :MeCTe 6proIDHoro aOpTaJILHOro naparaHrJIHJI CROnJIeHIIJI KJIe-
196
-rOK, ]J;alO IIX nOJlOmlITeJlbHYIO xpoMa«lJ«lJIIHHYIO peaKQIIIO Y B3poCJlbiX
JlIO]:(eH ]:(0 75-JleTHerO B03paCTa.
ApyraH KapTIIHa Ha6J1IO]:(aeTCJI Y HaceKOMOH]:(HLIX II pyKORpblJlblX.
"Y emeH U JleTyqUX MbimeH UX OIIIICaJi Celestino da Costa (1917,
1920, 1935a, 1936a, b), y Ma]:(araCKapCRUX HaceKOMOH]:(HbIX (Cente-
tes ecaudatus, Hemicentetes semispinosus, Ericulus setosus) Adams
(1958a). B sM6pllOHaJlbHOM pa3BIITIIII emeH opraH U:YKepKaH]:(JlH XOTJI
.II CY.I.U;eCTByeT, HO HalI60Jlee nOJIHOrO pa3BIITIIJI ]:(OCTllraeT TOJIbRO
IIOCJle pOiR]:(eHIIJI. Ero KJleTKII )J;oJlro COXpaHJIIOTCJI B He)J;u«lJ(pCpeHQU-
pOBaHHml COCTOJIHIIU II, BU]J;UMO, HaqlIHalOT C03peBaTb TOJIbliO nocJle
pomJ];eHlIJI; «lJYHRQUOHlIpyeT STOT opraH B TeqeHlIe BceH mll3HII (Ce-
lestino da Costa, 1936a). Y HeKOTopblX HaceKOMOH]:(HbIX (Adams,
1958a) IIpeaOpTaJlbHblH naparaHrJlIIH O'IeHb pacnpocTpaHeH II 3aHlI-
l.meT y 3apo]:(blmeH 06mlIpHYlO TepplIToplIlO. OH IIpe]:(CTaBJleH CKOII-
:reHlIeM KJleTO'IHLlX rHe3]:( Ha BeHTpaJlbHOH IIOBepXHOCTn 6plOmHoii
.aOpTbi II ]:(BYMH ]:(op3aJlbHblMII IIapaaOpTaJlbHbIMII cToJl61IRaMil IIapa-
rallrJIJIOUapnOH TKaHU, KOTOpaJI THneTCH B]:(OJIb 6plOmHoH, rpY]:(HoH
n meiinOM 06J1acTeii 3apo]J;blma. Cnepe]:(1I OHII ]:(OXO]:(HT ]:(0 OCHOBa-
HnH qepena II ]:(OCTllraIOT ypOBHH IipaHlIaJlbHOrO nOJilOca BepxHcro
meUHoro Y3J1a, a C3a]:(11 CIIYCKalOTCJI Hume 6U«lJYPKa:u;UU aopTbi. CTOJI-
OHKII IIpOXO]:(HT Jlu60 B BH]:(e CIIJlOIliHOll JlCHTLI (y Ericulus), JlII60
06pa3YIOT IIpepLIBUcTLIll TJI;H (y Centetes) II TJIHYTCH no npO]:(OJlb-
Hoii OCII TeJla pH]:(OM C CHMIIaTlIqeCIaIMII CTBOJIallIH. B TOH 06J1aCTU,
r]:(e pa3BIIBalOTCJI Ha]:(IIOqeqHmm, napaaOpTaJlbHble CTOJl6uKII pacno-
JIaralOTCJI ]:(op3aJlbHee meJle3 U CBOOO]:(HO C HUMII co06 a1OTCJI, a B
TeqeHlle nnYTpHYTp06Horo pa3nUTIIJI OHII perpeccllpYIOT OqeHb He3Ha-
'IUTe.lIbHO.
OcooeHHocTLIO HaceKOMOH]:(HbIX HBJlHeTCH II crroc06 06pa30BaHIIH
M03rOBoro CJlOH HaAITOqe'lHlIROB, ROTOpblll «lJopMupyeTCJI He IIYTeM
MllrpaQlI1I IweTOK, a cJlymuT Herrocpe]J;CTBeHHLIM rrpO]:(OJlmeHlleM IIpe-
aOpTaJlbHOrO naparaHrJIJIJI, ROTOpLIM, BHe]:(pJIHCb B ROpTUKaJlbHblH
CJlOH, rrOKpblBaeTCJI 11M KaH manKOll. TaKoii cnoc06 He HBJlHeTCJI,
BIIpOqe l, IICKJlIOqeHIIeM )J;JlJI HaCeKOI\IOH]:(HblX, OH onucaH He TOJlbKO
y emell (Celestino da Costa, HJ:36a), HO II Y KpOJIIIKOB (Kohn,
1903; Coupland, 1956, 1965; CI\HlTTeH, 1962).
TaKlIl\1 06pa30M, pa3BIITue Ha]:(nOqeqHUKOBOH U BHeHa]:(nOqeWII-
KOBOU naparaHrJlIIOHapHOH TKaHII y HaCeKOMOJI]:(HbiX II PYKOKpblJlblX
CJle]:(yeT npOTUBonOCTaBJlJITb, Ral{ 3TO ]:(eJiaeT II Adams (1958a), APY-
rlIM MJleKOIIHTaIOIIJ;IIM. .
113 3TIIX, XOTH U CRY]J;IIb1X CBe]:(eHUU 0 pa3BUTIIU naparaHrJllIeB
Y HeKOTopblX MJleKOrrUTalOlQlIX (B CMblCJI{) OXBaTa pa3HblX rpYlln MJle-
KorrIITaIO IIX), MomHO c]:(eJlaTb HeKOTOpbIe BblBO]:(bI.
TIpeaOpTaJlbHblH naparaHrJluii 3apO]:(blmeH MJleKOnUTalO IIX pacno-
JIOmeH, J\ali IIOKa3b1BaeT caMO Ha3naHlIe, OIiOJlO aopTbi U I\IeCTal\1H
COIIpllKaCaeTCJI C ee a]J;BeHTIIQlleii. AopTa II conYTCTBYIO lIe en Ma-
rllCTpaJlbHble BeHbI y KpyrJlopoTbiX II pbl6 HaCbI eHbI, KaK I\lbl 3HaeM,
xpoMa«lJ«lJIIHHon TKaHblO II JIBJlJIIOTCJI rJlaBHblM MeCTOM ee pacnpo-
CTpaHeHIIJI. JIerKo IIpe]:(CTaBIITL, 'ITO rrpeaOpTaJlbHbln naparaHrJllIn
197
JIBJlJIeTCJI 3M6plIOHaJlbHblM IIpII3HaROM, ROTOpblii: peRaIIIITYJllIpyeT CO-
OTBeTCTBYlOlIJ;lIe CTPYRTypbl, xopomo pa3BIITbie B 3apOAbillieBOM ne-
plIo]:(e II Y B3pOCJlblX oc06eii: IIpe]:(ROBbiX «lJOPM. Y HaceROMOJIAHblX
IIpeaOpTaJlbHblH IIaparaHrJlIIH AOCTlIraeT HalI60Jlee MOIIJ;Horo pa3BH-
TIIH. OH nO]:(XO]:(IIT, ImR yme CRa3aHO, CBOIIM nepe]:(HIIM ROHD;OM R
qepeIIY, a 3a)J;HIIM - R pa3BIIJIRe aopTbi. Y ]:(PyrlIx MJIeROnIITaIOIIJ;IIX.
60JIee BbiCORO opraHII30BaHHLIX, OH cocpeAOTOqlIBaeTCJI B 06JIaCTII
6plOmHoii: aopTbi. CJIe]:(OBaTeJIbHO, paCTHHYTOCTb IIaparaHrJIliOHapHOH
TRaHII 110 ]:(JlIIHHOH OCII TYJIOBlIlIJ;a - IIplI3HaR HII3ROH «lJIIJIOreHeTn-
qeClWH opraHlI3an;IIII mlIBoTlIoro, it ROHD;eHTpan;IIJI ee B6JIII31I HaA-
nOqeqHIIROB - CBII]:(eTeJIbCTBO HanOOJIee BbiCOROH opraHII3aD;lIlI. ECJIJI
yqeCTb, qTO HaceROMOfl]:(Rble HIJJIJIIOTCJI HalI60.rree IIplIMIITIIBHLIMH MJIe-
ROnIITaIOIIJ;IIMU, H3 ROTOpblX pa3llHJIlICb Bce COBpeMeHHble IIJIal{eHTap-
Hble, TO IIpOn;ecc OHToreHeTlIqeCIWH peI\anIITYJIJIl\IUI «lJIIJIOrOHe3a IIa-
paraHrJIlIeB IIpe]:(CTaBJIeH ]:(OBOJIbHO HarJIJI]:(HO. OG 3Tml ronopn r II
CpORII pa3BIITIIJI IIpeaOpTaJIbHblX IIaparaHrJIHeB, ROTOpblC y HaceRO-
MOJI]:(HbIX npO]:(OJImalOT ern;o «lJYHKQIIOlInpOBaTb Y B3pOCJlbIX II>lIBOT-
HbIX, npU6JIHmaHCb II no 3TIIM nORa3aTeJIJIM K CBOIIM npe]:(lmM, B TO
BpeMH KaK COBpeMeHRbIe B3pOCJIbie «lJOPMbl P;PyrlIx IIlJIeKOlIIlTalOrn;IIX
«lJYHKD;1I1O YTpaTHJIII.
Bo MHorlIX npe]:(bI]:(yrn;IIX rJlaBaX IIepBoH qaCTII HaCTOJIrn;eH paoo-
Tbi cpaBHIITeJIbHO-Mop«lJOJIOrlIqeCKII 6b1JIO IIOKa3aHO, qTO IIIH0rlIe lIa-
paranrJlulI TepJIIOT CBoe «lJYHKQIIOHaJIbHOe 3HaqeHIIe P;JlJI opraHH3Ma
B pe3yJlbTaTe II3MeHeHIIH YCJIOBIIH cyrn;ecTBOBaRIIJI II nOCTeIIeHHO pe-
]:(YD;IIpYIOTCJI. M3BeCTHO, qTO CYlIJ;eCTByeT HeCKOJIbKO TIIIIOB pe]:(yK-
D;IIII opraHOB. OAIIH II3 HHX npOJIBJIHeTCJI B TOM, qTO opraH cyrn;e-
cTByeT JIlImb B 3M6plIoreHe3e BbICOKopa3BIITbiX B «lJIIJIOreHeTlIqeCROM
OTHomeHlI1I «lJOPM II IICqe3aeT R IIIOMeHTY pOlli]:(eHIIH IIJIII Ha KaKOlII-
TO 3Tane OHToreHe3a - BpeMeHHbIe, IlJIII npOBH30pHble opraHbl. OHU
06eCneqlIBillOT «lJYHKD;1I1I opraHII3Ma ]:(0 Tex nop, nOKa He C«lJ0PMH-
pOBaJlCJI II He HaqaJI «lJYHKD;IIOHlIpOBaTb aHaJIOrlIqHblH opraH, npH-
CYIIJ;IIH B3pOCJIbiM mIIBOTHbIM.
TaKOMY onpe]:(eJIeHHIO BnOJIHe OTBeqae1 opra.H QYKepKaH]:(JIJI.
A paCIIpe]:(eJIeHlIe B HeM HepBHblX CTPYKTYP (CaKOBlIq, 1964, 1969)
)J;aeT gOIIOJIHIITeJlbHblH MaTepHaJI B IIOJlb3Y TaKoro onpe]:(eJIeHIIJI. Pe-
l\aIIlITYJlUpyIOrn;ee 3HaqeHIIe xapaKTepa IIHnepBan;HH 6pIOIliHo-aOpTa.iIb-
HblX naparaHrJIlIeB pa306paHo B rJIaBe VI.
M3BecTHo, qTO IIpeaOpTaJIbHblH IIaparaHrJIIIii: CJlymlIT «MaTplID;eH»,
IIOCTaBJlJIlOlIJ;ell KJIeTOqHblll MaTepHaJI ]:(JIH M03rOBoro CJIOJI Ha]:(IIO-
qeqHIIROB. CIIoc06, IIPII IIOMOIIJ;II KOToporo OCYlIJ;eCTBJlHeTCJI 3TOT npo-
n;ecc II «lJopMlIpyeTCJI HaAIIOqeqHaJI meJIe3a MJIeKOIIIITaIOIIJ;IIX, 'fome
3aCJIymlIBaeT BHIIMaHIIJI. Bo BpeMJI 3M6plIoreHe3a M03rOBOH CJloii: Ha]:(-
nOqeqHIIRa JIII60 COCTaBJIJIeT qaCTb npeaOpTaJIbHOrO naparaHrJlHJI R
IIpHMOM CMblCJle CJIOBa, T. e. ero «lJopMHpOBaHlIe II]:(eT nYTeM U-06-
pa3Horo BpaCTaHIIJI B KOpTIIKaJILHble 3aQaTRII, KOTopble B BII]:(e ma-
IIOQeR IIJlII RaIIlOmOHOB ca]:(JITCJI Ha KpaHlIaJlbHble OTpe3KII IIpeaOpTaJIb-
Horo naparaHrJIIIJI (pIIC. 71), JlII60 IIYTeM MlIrpa:u;IIII II3 Hero RJIe-
TOQHOrO IIOTORa BHYTPb ROpbl. B nepBOM CJIYQae B3aHMOOTHOmeHBJI
198
a
p )I c. 71. Cnocoo oopa30BaHuH
xpuMa4>4>IIHHoii TRaHII HaAIIO'le'l'
HIIKa B 1M6pllOreHe3e KpUJUlKa
(peKoHcTpY K l\IIf1)
a - aopT3 (Coupland, 19(5)
,., ""
(J,IMM
'-----'
IMN
A\
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1103rOBoro U KOpKOBOro CJlOeB B 3M6pHOreHe3e MOmHO CKOpee Ha3-
BaTb IIpUJlemaHlIeM ]:(pyr K ]:(pyry, a BO BTOpOM cJlyqae B3aUMHLlM
nepeMemlIBaHlIeM. "Y HeKOTopLlX MJle1{OIIUTaIOIII;IIX, R KOTOpLlM IIpU-
Ha]:(JlemaT npUMHTIIBHble IIJlan;eHTapHble, M03rOBon CJIon Ha,n;IIOqeqHII
KOB 06pa3yeTCJI nocpeACTBOM nepBoro cnoc06a, y APyrlIx nocpeA-
CTBOM BToporo. B IIpon.ecce ]:(aJlLHenlllero pa3BIITIIJI ROpTIIKaJlbHblO
maIIOqKU IIOJlHOCTbIO oKpymaIOT M03rOBon CJIon.
HaM II3BeCTHo, qTO AJIJI HeKoTopblX B3pOCJlblX «lJOPM HUmeCTOJI-
III;UX rpYIIII mUBOTHbiX HaIIpIIMep, aM«lJu6un) IIplIJIemaHUe «M03ra»
]I «ROPbl» 6e3 B3aUMHoro IIpopacTaHHH JIBJlJIeTCJI xapaKTepHblM MOp-
cP0JIOrUqeCRIIM cBoncTBoM II COXpaHJIeTCJI B TeqeHlIe Bcen m1I3HII.
3aTeM HaqUHaeTCJI B3aUMHoe npoHuKHoBeHlIe, KOTopoe IIpUBOAIIT K
(<IIorJIOIII;enUIO» M03ra Kopon II «lJopMllpoBaHIIIO HaAIIOqeqHblX meJle3
MJIeKOnIITaIOIII;ux C IIX xapaKTepHon CTPYKTypon.
Ha OCHOBaHIIU cRa3aHHoro, Mbi IIOJIal'aeM, qTO U-06pa3Hoe Bpa-
CTaHHe M03rOBoro CJIOJI Ha]:(IIOqeqHIIROB B 3M6puoreHe3e MJIeKOnlI-
TaIOIII;HX JIBJlJIeTCJI 60JIee «]:(peBHIIM» cnoc060M «lJopMlIpOBaHUJI HaAITO-
'IeqHblX meJIe3, a MlIrpan;HJI 60JIee (<HOBblM». OqeBIIAHO, MomHO npeA-
nOJIOmUTb, qTO cIIoc06 opraHoreHe3a uaAIToqeqnblx meJle3 B KaKon-To
Mepe oTpamaeT ypoBeHb <pIIJloreHeTHqeCKOrO pa3BIITIIJI mIIBOTHbiX.
J];JIJI 60JIee YBepeHHoro cymAeHIIJI no 3TOMY BOIIPOCY HymHbI, oAHa-
KO, )J;OIIOJIHUTeJIbHble IICCJIeAOBaHUJI MHorlIX rpynn MJIeKOnIITaIOIII;lIx
no BOcxo]:(JIIII;eii: JleCTHUn;e «lJUJIOreHe3a.
BTopon TIIII pe]:(YKn;UII opraHoB - py]:(uMeHTan;IIJI. PYAHM6HTap-
HbIe opraHbI COXpaHJIIOTCJI B TeqeHlIe Bcen mlI3HII opraHII3Ma, HO
B BII]:(e CTPYKTYP, YTpaTIIBmux UJIII CHH3IIBmUX CBoe OCHOBHoe «lJYHR-
n;UOHaJIbHOe 3HaqeHUe B npon;ecce HCTOplIqeCKOrO pa3BIITUJI.
o pY]:(IIMeHTapHblX opraH ax IIaparanrJIIIOHapHon cnCTeMbi BbIcmlIX
1I03BOH01JHblX yme IlIJIa peqb B npe)J;bI)J;yIII;IIX rJIaBax. CpaBHHTeJlb-
nO-Mop«lJOJIOrUqeCROe II3yqeHUe IIOKa3aJIO, qTO B CTPYKTypHOM U «lJYHR-
IJ;UOHaJIbHOM OTHOmeHIIJIX, a TaKme no IIOJIOmeHUIO B6JIU3U CTeHOK
RpynHblx RpOBeHOCHbiX COCYAOB mell II rpYAII (COHHblX, JlerOqHblX,
iUS.
aOpTbI}, OHII JIBJlJIIOTCH OCTaTl\aMII TOH TKaHII, KOTOpaJI UblJla XOpO-
mo pa3BIITa Y npep;Iwn STIIX opraHII3MOB. BblJlo IIOl\a3ano, qTO cne-
n;mlm'lecKHe KJleTRII KapOTII]:(Horo H Ha]:(cep]:(e'lHbIX naparaHrJlHeB
B3pOCJlbiX }I\HBOTHbIX IIMelOT aTlInHqHOe CTpoeHlIe II pe3IW IIOIlIDKeH-
HYIO SIIP;OI{PIIHHYIO <PYHKI{IIIO. Xo]:( rlICToreHe3a STIIX napcmmrmIeB
II KJleTO'lHbIe IICTO'lHIIIUI IIX pa3BHTIIJI, KOTOpbIe OKaaamlCb C]:(I1HbllIlIl
P;JlH naparaHrJIIIeB UPIOIlIHOII (1.103rOBOH CJlOH Ha]:(IIOqe'lHIIKon) II meH-
Hoii (RapOTII]:(Hblii naparaHrmlii) 06J1aCTen II, CJle]:(OBaTeJlbHO, nocJly-
HmJlII B nOJlb3Y I1X e]:(I1Hoii a]:(peHaJlOBOii WYHRI{IIII - OIIlIcaHbI Hume
(CMIITTeH, 1958, 19G2; IIPUHIIIaK II CIIIIlTTeH, 1959). 3]:(ecb 3aCJlY-
»>llBaeT 6bITb onre'leHHbUI SMupIlOJlOrIlQeCROe Ha6J1IO]:(ellIlc Adams
(1958a) Ha HaceKOlIIOH]:(HbIX, IWTopoe nO]:(KpeIIJlHeT, Ha lIaIli B3rJlJI]:(,
PY]:(lIlIIellTapHbIII xapaKTep ImpOTlI IIOrO naparaHrJlUH.
IIplI OlIlicaHIIII naparaHrJlIlCB Y 3apo]:(bIllIeii lIIa;:{araCIwpCImx na-
CCROl\IOJI]:(HbIX Adams IIO]:('lCpIWYJI, qTO BCH (<BHYTpIlII<I]:(IIO'letIIIIIKO-
BaJII) naparaHrJlIIOHapHaJI TKalIb I{I1TOJlOrIl'leCKII II rIICTOJlOrlI\IeCKU
CXO]:(Ha C «BHeHa]:(nO'le'IIIIIIWBOih), KOTopaH pacnpOCTpaHella B]:(OJIL
Bceii ]:(JIlInbl aopTbI, BnJlOT}, ]:(0 IWpOTIIP;HOH pa3BIIJlKII. Ha OCIIOBa-
HIIII MOPWOJlOrH'leCKOrO CXO]:(CTBa BHe- II BHYTpHHa]:(nO'leqHIIKOBOii
TKaHII Adallis paCCMaTplIBaeT TY II ]:(PyrylO RaK e]:(IIHYIO CIICTeMY.
B TO me BpeMJI OH 06par:u;aeT BHUMaHlIe Ha TO, 'lTO 3TH IIaparaH-
fJIIIOHapHbIe MaCCbI OTJllIqnbI 110 CTPyKTYpC OT ImpOTII]:(Horo IIapa-
raHrJlIIH, ROTOpbIH JlCmlIT Y MCCTa RapOTII]:(Hon 6HwypKaI{IIJI '. 0 qeM
MomeT CBII]:(eTeJlbCTBOBaTb 3TOT waKT? Adams nOJIaraeT, 'lTO pa3J1U-
'lIIJI IIX B CTpYKType BCRJlIO'laIOT B03MomHOCTb npOIlCXOil\]:(eIlIIH Ka-
pOTB]:(Horo IIaparaHrmIJI B3 Toro }Re HCTOqHIIKa, 'lTO II napaaOpTaJlb-
HaJI TRaHb, KOTOpaJI B TaKOM 1I306HJlIIB lIMe eTCH Y 3apO]:(blmeii Ha-
CeKOMOJI]:(HblX, 'lTO p;OCTBraeT caMbIX BepXHIIX pa3BeTBJleHIIii aopTbI,
BIIJlOTb ]:(0 COHHblX apTepBH. TOQRa 3pCHBJI Adams He OTpamaeT
06'bel{TIIBHOro nOJlOmeHIIJI BelI{oii. BbIme MbI BII]:(eJlH, 'lTO no CTPyK-
Type, CBJI3JIM C ]:(pyrBMII THaHHIIIH II rHCTOXBIIHI'IeCImlll cBoiicTBa}f,
KOTOpbIe rOBopHT 0 pe3KOlli CIIIIn,eHIIH rOplllOHaJlbHOH <PYHlm;HII, Ka-
pOTII]:(HblH IIaparaHrJIIIii JIBJlHeTCJI 3HP;OKpIlHHOii, HO PY]:(B}feHTIIpO-
BaHHon meJle30n, IIpIIHa]:(Jlemar:u;eii K e]:(IIHoii naparaHrJlIlOHapIIoii CII-
CTeMe II BblIIOJlHJIBllIen B <pIIJlOreHeTII'leCKOM IIpOmJlOllf nOJlHO eH'
HYIO, ap;peHaJlOBYIO WYHKn;IIIO.
06'bJICHeHlIe Ton pa3HIII{bI B CTpYRType, KOTOpylO Ha6J1IO]:(aJi Adams
B 3M6plIOHaJlbHOM KapOTIIp;l101II naparaHrJlIIII, 110 cpaBHelllI10 C ]:(pyrII-
MB, nplIMbiRalOlI{lIlII1I K HeMY naparaHrJlIIJIMII, CJle]:(yeT, C Hamen TOq-
KII 3peHIIH, IICKaTb B pa3HbIX TlInax pe]:(YKQIIII: C o]:(Hon CTOpOHbI,
06pa30BalllIe npoBu30pHoro opraH a (npeaOpTaJlbHbln naparaHrJllIn),
C P;Pyron,- PYP;IIIIIeHTapHOro (KapOTII]:(Hbin IIaparaHrJllIn). IIpOBII30p-
I 8TO me MTIeElIe BIdCIi:a3IdBaeT Celestino da Costa (1935b), C'lIlTaH, 'lTO KJleTKB
RapoTlip;rroro Daparanrmm HBJIHIOTCH Pyp;IIMeHTaprrIdMIi BJIeMeHTaMIi YUJIOT-
HeHrrOH cOCYP;liCTOH CTellRII. a TepMlirre «PYP;IIMeHTapHIdH» B DOlllIMaDIIlI Ce-
lestino da Costa rOBopliTCH B rJIaBe VIII. ABTop BRJIap;bIRaeT B nero IIrro
cOAepmaHlIe, OTJIII'lHOe OT Hamero.
200
HLIB - 3TO BpeMeHHaH, HO IIOJIHO eHHo4>YHRIJ;llOHHpYIOm;aH meJIe3a,
PY IIMeHTapHLIll - 3TO nOCTOHHHaH, HO IIOqTlI He 4>YHKD;HOHHPYIO-
:m;aH. IIx 4>YHI{ I1II COOTBeTCTllYIOT nx CTpYKType.
y IIJIO OB qeJIOBeRa HMeIOTCH Ba RpynHLIx Ha cepJ];e'lHbIX IIa-
paraHrJIlIR (IlllmHHH II BepXIIHH) II HeCKOJIbKO MCJIKHX (Palme, 1934;
HOBpllmKo, 1962a, -1970). HHmHlIll, B 3MupHoreIle3e KOToporo npe-
06JIa aIOT xpoMacpcpHIlHbIe 3JIeMeHTLI, He YBeJIII'lIlBaRCb K pomJ];eHIlIO,
aaqlIHaeT perpeCCIIpOBaTb H y B3pOCJILIX COCTOIIT 113 He60JIblllOrO qllC-
JIa HexpOMacpcpIlHHbIX 3,'1eMeIiTOB. BepXIIIIll, Y ROTOpOro IIpe06JIa aIOT
Hexpmmcp<jJHIIHLIe IweTKII, 1I0CTeIIeHIIO YBeJIH'lIlBaeTCH H Y B3pOC-
JIblX CO]];epmIlT HeCKOJIbKO HexpOMacpcpHHHLIX KJIeTOqHblX OCTpOBKOB.
HexpOMacpcplIHHLIe KJIeTlU1 IIMeIOT IIOJIIlrOHaJIbllYIO CPOPMY, Tor a KaK
xpOMacpcpllHHLIe MorYT 06JIa]];aTb OTpOCTKaMH (Palme, 1934; Boyd,
1960), T. e. coxpaHHIOT e e 4>OPMY, CBoBcTBeIlHYIO MIIHoraM. TaKIIM
o6pa30M, HeROTopble MOPCPOJIOrIl1JeCKlle qepTLI II rHCTOXHMH1JeCKHe
CBOBCTBa KJIeTOIi B OHToreHe3e 1l0BTOpHIOT 3Tallbl cpIl.'10reHe3a. PeJU'I<-
U;HH BepXHf:r0 IIaparaHrJIHH H eT CKopee no Tlmy PY HMeHTaD;JIII, aHa
IlllmHero 06paayeTcH IIpOBH30pHLIll opraH. Ha6JIIO]:(eHIIH Palme HJI-
JIIOCTpnpyIOT perp<;ccHIO IIaparaHrJIHeB Y qCJIOBCRa.
}' mlIllOTHblX HaJ];cepJ];eqHLIX IIaparaHrmleB 60JIbllle, qeM y qeJIO-
BeKa, II IIX perpeccHH HJ];eT Me JIeHHee (Palme, 1934). OC06CHHO
MHoro xpoMacpcpllHHoii TKaHII y aapoJ];LIlliell JIeTY1JIIX Mblllleii, CROIIJIe-
HIIH ROTOpOll 06paaYIOT :n;eJIYIO eIIb IIaparaHrJIIIeB B OJIb llIeH I]
rPY Hoii qaCTII TYJIOIJII a (Celestino da Costa, 1935a, 1936b). 3a-
'laTOK Ha]];H,JIJOqllqHOro IIaparaHrJIlIH Y KpOJIIIRa (D'Agostini, 1959),
pa3BHBaIO HJICH 113 MeaeHxHMHoro YTOJI CmIH IIpaBOJI HaJ];KJIIOqHq-
HOll apTepUH, IIOCJIe pom eHIlH OTJ];eJIHeTCH OT apTepHH, ero coeJ];HHII-
TeJIbHOTKamJaH CTpOMa pa3paCTaeTCR II y BapOCJILIX KpOJIHROB
:rrpeBpa aeTCR B Py6 OBYIO TKaHI>, JIHIlICIlIlYJO cne Hcpll'leCKHX K.JIe-
TOK. AOpTaJIbIlO-JIeroqHblH IIaparaHrJIHll, IICCJIeJ];OBaHHblH BO BHYTPH-
YTp06HOM paaBIIT1I1I Y HOBopomJ];eHHblX II B3pOCJIbiX KOllieK (Hug-
hes, 1967), IIHBOJIIOlipyeT lIapaJIJIeJIbHO IIIIBOJIIO IIII cocYJ];a H 3apa-
:m;eHIIH ero IIpOCBeTa. BOTaJIJIOB IIpOTOK B 3M6pHoreHeae poraToro
CKOTa (Harms, 1966) oRpymeIl KOJIb OM lIaparanrJIlIOHapHOH TKa-
HH B tunica adventicia II t. media IIpOTOKa II CJIHBaeTCH C JIe-
rOqHbIM naparaHrJIIIeM. TIocJIe pomJ];eHHH, HeCMOTpH Ha IIHBOJIIOD;H-
OHHbIe npo eCCbI B car-10M IIpOTOKe, 3TH yaJIbI HeKOTopoe BpeMH CO-
xpaHHIOTCH, no C rOJ];aMH aapaCTaIOT coeJ];HIIIITeJIbHOII TKaHbIO.
HarJIflJ];IIYIO KapTlIHY nOCTeneIlHoro oopaTHoro pa3BHTIIH 06Hapy-
mIIBaIOT B OHToreHeae a6J];OMIIHaJILUbie H peTponepIITOHHaJIhHl>le IIa-
paraur:ulII. Em;e B 1901 r. Zuckerkandl OIIIICa,'I B 06JIaCTII 6pIOIlI-
HOB aopTbi 1Ie.'10BCqeClwro 3apo]:(blllIa naparanrJIlIII, IWTOpblll HaaBaH
ero nMeHeM II KOTOpbIll 1I0J];BepraJICR 06CJIeJ];oBaIllIIO r-morlIMIi aBTO-
par.1II (Kohn, 1903; Poll, 1906; Vincent, 1910; HXOHTOB, 1913, a, 6;
Ce]estino da Costa, 1926; I1naIIoB, t 930; Goormaghtigh, 1936,
II J];p.). OpraH I(YKepKaH]:(JIH, KaK IIpaBIIJIO, IICqeaaeT 1I0CJIe pom e-
HHH, TOJIbKO Y HeKOTopblX mITBOTHbiX (c06aKII) OH MomeT COXpaHHTL-
CH B TeqeHne mII3H11 (Celestino da Costa, 1926). TI pOD;ecc HC1Je3-
201
HOBeHlIJI, no YTBepm]:(eHIIIO aBTopa, oco6eHHO ]:(eMOHCTpaTHBeH Y Jle
Ty<IUX MblllleH, Y ROTOpblX OT 06HJlbHOrO CROIIJleHIIH xpOMacpqJlIHHoH
TRaHII Y 3apO]:(blllla OCTalOTCJI Y B3pOCJlbiX oc06eH JlHllIb OT]:(eJlbHbI&
rHe3]:(a. Y 6eJlbiX Mbllllen (Goormaghtigh, 1936) Ha MeCTe IICqe3HYB-
llIero xpoMacpcplIHHoro IIaparaHrJlJIJI, OIIlIcaHHoro Kohn (1903), B peT-
pOIIepliTOHlIaJlbHOM IIpOCTpaHCTBe OCTaeTCH IIpepblBlicTLIn mHYP IIapa-
raHrJlHOHapHblX KJleTOR B KOHTaRTe C IIOqeqHOn II HlImHeH IIOJlOH:
BeHaMH, C ganglion solaris H HHme ero. CoxpaHIIBllIlIncH IlIHyp
KJleTOR COJ];epmlIT RaR xpOMacpcplIHHbIe, TaR 11 HexpOMacpcplIHHble :,)JIe-
MeHTbI. Y qeJlOBeRa OCT3TKII 6b1BllIeH 06JIJIbHOH xpoMa4>CPIIHHon TRa-
HH IIJlO]:(OB OIIIICaJi B 1930 r. r. <D. MBaHOB B a}:l;BeHTIII.\IIII 6plOllI-
HOIi: aopTbl.
B Ha6nlO]:(eHHHx Ha]:( paCIIpOCTpaHeHHeM naparaHrJlHOHapHon TRa-
HH Y {3apo]:(blllleH 60JlbmOe MeCTO 33HIIMaeT HX OROJlOr3HrJl1l0Hap-
Hoe paCIIOJlOmeHlIe, 6J1aro]:(apH KOTOPOMY OHH II 1I0Jly<IIIJIH CBoe Ha3-
BaHHe. TIocTOJIHHaJl 'I'onorpa4>lIqeCRaJl CBJl3b IIaparaHrJlIIOHapHblX
RJle'l'OR C Y3J1aMII Cl<IMIIaTlIqe,CROn HepBHon CIiCTeMbl OTMeqeHa B 60Jlb-
llIeH IIJlH MeHbllleH CTeIIeHH Hce 1H HCCJle}:lOBaTemIMH xpOMacp4>HHHoii
CHCTeMbl II OCHOBaHa Ha rII'CTOreHern'IeCKOM pO]:(CTBe. Mx IIpIICYTCT-
BHe Y 3apO]:(blllleH B pa3HbiX Y'IaCTKax CIiMnaTlIqeCKO:ll CHCTeMbl OIIH-
caJlII MHOI'He aBTopbl, HO HaH60Jlee l.\eHHblMIi IICCJle.n:OBaHHHMH, coxpa-
HHBIlIIIMH 3HaqeHHe ]:(0 HamlIX }:IHeH, Ha]:(o IIplI3HaTb pa60Tbi Kohn..
a T3Kme Celestino da Costa (1917, 1936a, b) H 60Jlee n03]:(HIOIO CTa-
TblO Adams (1958a).
Kohn HamrCaJi pH):{ CPYH]:(aMeHTaJlbHbIX pa60T, 0606 eHHblx B MO-
Horpa4>IIH 1903 r., r]:(e IIplIBeJi 061l1IIpHblH !fJaKTlIqeCRIIH MaTeplIaJl 110-
naparaHrJlHOHapHoH CHCTeMe H c]:(eJlaJi Maccy Ha6J1IO]:(eHIIH Ha]:( pa3-
BHTHeM OT]:(eJlbHbIX ee opraHOB Y pa3HhlX BII]:(OB mIIBOTHbIX. C ero.
HMeHeM CBJI3aH B3rJlH}:I Ha IIpOHcxom]:(eHlIe xpoMacp4>IIHHOH TKaHIi 1I3'
CIIMIIaTH'leC'IWH II TeopHH HepBHoH 4>YHKIJ;lIH xpoMa4>tPlIHHblX KJleTOR.
TIoCJle]:(HHe Kohn OiIIlICaJi Y 3apO]:(blllleH KpOJlJma, KpblCbl, MOpCROD:
CBIIHKII, MbTIlIII, CBHHblI, co6aKII, KOIllRll, qeJlOBeI a B IIOrpaHlIqHOM
CTBOJle H BO MHorliX HepBHblX CIIJleTeHIIHX TYJlOBII a:. p1. aorticus
abdomin3lis, pI. mesentericus inferior, pI. coelicus, pI. hipogastricus.
RaR npaBJIJIo, B 6plOlliHOH 06J1aCTII II nplIJleraroID;Hx yqaCTRaX
IIorpaHlIqHOrO xpoMa4>CPlIHHoro CTBOJIa xpoMacpcplIHHblX RJleTOR 60Jlb-
llIe, a B Kay]:(aJlbHOM H RpaHHaJlbHOM HanpaBJleHIIJ'IX R0J1lIqeCTBO ilK
B CTBOJIaX HCTOID;aeTCH. TOJlbKO Y -3apO]:(blllleH HaceKOMOH]:(HbIX II PY-
RORpbIJIbiX MomHO rOBoplITb 06 06HJlbHOM II nOCTOHHHOM IIPIICYTCTBJIlI
IIaparaHrJlIIOHapHblX 3J1eMeHTOB 110 ]:(JlHHe Bcero CIiMIIarnqeCROro.
CTBOJIa (Celestino da Costa, 1917, 1920, 1926, 1936; Adams, 1958a).
OqeBH]:(HO, "[{TO pacnpOCTpaHeHHaH B;J;OJlb Bcero TYJlOBIIID;a 3aRJIa]:(-
Ka xpOMacp4>IIHHOH TRaHH Y HH3mlIX nJIan:eHTapHblX OTpamaeT peRa-
IIlIryJIIIPYIOID;IIH IIpOl.\eCC B OHToreHe3e II03BOHOqHbIX, a KOHl.\eHTpa-
I.\HH ee Y BblCllIlIX n03BOHO'IHbIX Ha OrpaHH'IeHHOM Y'IaCTKe aopTbI-
3]:(aIITHBHbIH, H3IIpaBJIeHHblH Ha 4>opMHpoBaHlie H3J{IIMe'IHHR3, ROTO-
pblH B 3BOJIIOD;HOHHOM OTHOllieHIIH HBJIHeTCH 60JIee cOBepmeHHoH 3H-
}:IORpHHHOH meJIe30ii II xapaRTeplI3yeT agpeHaJIOBYIO CHCTeMY BbIC.
202
mHX II03BOHOqHblX. ::ho IIO]:(TBepm]:(aIOT H IIpOU;eOCbl RJIeTOqHblX ]:(H4>-
4>epeHU;1IpOBOR, H,D;ym;IIe 6b1CTpee B 6plOllIHblX IIaparaHrJI1IHX, 'leM B
meil:HblX. 8TO 6b1JIO O'leHb XOpOIliO nORa3aHO e e B pa60Tax Celesti-
no da Costa Ha em ax (1920, 1926) II Ha ]:(pyrHx MJIHKOIIHTaIOII{1IX-
sail:u;ax, RpoJI1IKaX, KOIliKax, JIeTyqHX Mbllllax II CYMqaTblX (1936a).
B HacTOHm;eil: rJIaBe 1I3JIOmeHbI 4>aRTbI H HeKoTopble co06pame-
HHH 0 peRanHryJIHU;IIHX xpoMa4>4>HHHoil: (a]:(peHaJIOBoil:) CHCTeMbI 110-
3BOHOqHl>1X. O]:(HaKo MbI 06pa aeM BHHMaHHe HaTO, qTO He Bce
-cBOMCTBa 3TOiI: CUCTeMbl peKaIIHTYJI1IPYIOT pa3BIITHe ee B 4>IIJIOreHe-
se. CpaBHIITeJIbHO-Mop4>OJIOrHqecRoe II3y<IeHlIe CHMnaTO-a]:(peHaJIOBoil:
-CliCTeMbI nOKa3aJIO 3HaqlITeJIbHOe 3ana3]:(blBaHlie B pa3BIITII1I CHMIIa-
TH'loc,KIIX ee 3JIeMeHTOB 110 cpaBHeHlI10 c aApeHaJIOBblMH (xpoMa4>-
tlJ1IHHbl III). B IIpOI1;ecce rliCToreHe3a C1IMlIaTO-xpoMa4>4>1IHHbIX ROA{II-
neRCOB BblHBIiJIaCb APyraH 3aROHOMepHOCTb - 60JIee II03]{Hee MOp4>O-
J10rIPIeCKOe C03peBaHHe xpoMa4>4>IIHHbIX 3JIeMeHTOB II COOTBeTCTBeH-
HO 60JIee II03AHee HaqaJIO IIpO]:(YRU;IIH MOHoaMIiHOB B H1IX no cpaB-
HeHlI10 C CIiMIIaTII'leCKHM1I 3JIeMeHTaMli (CM1ITTeH, By]:(aHu;eB, 1968).
Pa60Ta Enemar c coaBTOpaMIi (1965), ecJI1I y<IIITblBaTb B03paCT II3Y-
"IeHHbIX HMII 3apo]:(blllleil:, rOBopliT B nOJIb3Y TaROM 3aKOHOMepHOCTD-.
TIpou;eccaM rHCTOreHe3a II ]:(II4>4>epeHu;IIpOBKII KJIeTOK B CIiMnaTO-
xpoMa4>4>IIHHbIX ROMnJIeKCaX IIOCBHm;eHa CJIe]:(YIOm;aH rJIaBa.
P,n,aaa VIII
P A3BIITIIE AAPEHAJlOBJ>IX RJlETOR
II IIX IIOJlOiKEHIIE B rEHETlIqECKOII CIICTEME
HEPBHOII TKAHII
rlICTOreHea xpOMacpqJlIHHon CIiCTeMbi Ha Bcex :nanax IICTOpliH
ero 1I3yqeHIIJI Y n03BOHOqHbiX 6b1JI CBH3aH C IICCJIe]:(oBaHIIHMD paa-
BIITIIH Bel'eTaTllBHOn HepBHon CIiCTeMbi. 8TO Bbl3BaHO IIOCTOHHHblM
coce]:(cTBoM BereTaTDBHblX HenpoHoB lIepncpepnqeCROn HepBHon TRam!
C xpOMacpcplIHHblMIi :,meMeHTaMIi II HaJID1JlIeM 60JIbllllIX CIiMIIaTO-Xpo-
MacpcplIHHblX KOMllJIeKCOB, BCTpeqaIOlI{lIXCH Ha Bcex CTa,IJ;IIHX OHTore-
HeTlPIeCROrO pa3BIITliH opraH1I3MOB.
MCT01JHDR IIpOIICXOiR}WHIIH RJIeT01JHbIX :meMeHTOB xpm.mcpcpHH-
HOH II BereTaTIIBHOH HepBHOH CIICTeM - O]:(IIH 1I3 caMblX CTapblX BO-
npOCOB, p;e6aTHpOBaBIlIIIXCJI B TeqeHlIe MHorliX p;eCHTRJIeTlill. B pe-
3YJIbTaTe MHOroqRCJIeHHblX OIIlIcaHnll II :mCIIeplfMeHTaJIbHO-3M6pII0-
JIOrlf1JeCRIIX pa60T BblHBIIJIIiCb pa3HOrJIaCIIJI, KOTOpbIe IIPlfB€JIIf K CO-
3]:(aHIiIO ]:(BYX TeOpliJI - 0 Me3eHXIIMHOM II neHpmmTOAepMaJIbHOM IIX
pa3BIITHIi. ABTOpOM IIepBOH 6bIJI Remak (1847), BTOpOH Balfour
(1877). B03HnKaeT JIll BereTaTIIBHblH HeHpoH Ha MeCTe 1f3 Me3eHXII-
Mbl IIJIII HBJIHeTCH IIplilliJIblM 3JIeMeHTOM 1I3 HeJlp03KTo]:(epMbI - BOT
BOIIpOC, KOTOpblH BblHCHHJID IICCJIe]:(OBaTeJIIi. MecTHoe Bo.3HlfKHOBe-
HDe 3JIeMeHTOB BereTaTIIBHOll HepBHoH CHCTeMbl I 060cHOBbIBaJIOcb.
. rJIaBHblM o6pa30M TeM cpaKToM, qTO KJIeTKII 06HapYiRliBaJIliCb Ha ne-
plIcpeplIlI paHbllle Toro Bpe IeHD, Kor]:(a B HIIX MOiRHO 6bIJIO 06Hapy-
iRlITb OTpOoCTKII II HepBHbIe BOJIOKHa, CBH3bIBaIOII{IIe IIX C BblllleJIeiRa-
II{IIMII OTp;eJIaMII. O]:(IIaKO elI{e CTapbIe aBTopbl nplillIJIIi K npaBIiJIb-
HOMY IIpe]:(IIOJIOiReHIIIO II nOIm3aJIli Mlirpan;nIO 3aQaT01JHbIX KJIeTOK Ha
neplIcpepIIIO ern;e p;o BpaCTamIH TYp;a HepBHblX BOJIOROH (Poll, 1906;
Kohn, 1907; Kuntz, 1920). HallllI Ha6JIIO]:(emm, KOTOpbIe Mbi lIMeJIII
B03MOiRHOCTb npOBeCTIi Ha 60JIblllOM IaTepnaJIe y pa3HblX rpynu
II03BOHOqHbIX, IIOJIHOCTbIO C 3TllM COrJIacyIOTcH (CMIITTeH, 1962), 0]:(-
HaKO OHll paCXO]:(JlTCH C Ha6JIIO]:(eHIIHMR Muller If Ingvar (1923), KO-
TOpbIe CqIITaJIII, qTO Mnrpan;IIH npOIICXO]:(IIT TOJIbKO B]:(OJIb HepBHblx
nYQKOB, CPOpMIIPYIOllJ,llX 6YP;YIn:lIe rami communicantes. no HalllHM
p;aHHbJM, KJIeTRII nepecemnOTCJI B MeCTa 6YP;YlI{nx 3aRJIap;OK nepncpe-
pll'IeCRRX Y3JIOB Ha TaKIIX paHHIIX CTap;IIHX, Kor]:(a OHII He IIMeIOT
p;aiRe HaMeKa Ha pOCT OTpOCTROB.
I 0630p JIIITepaTypld II cOBpeMeHHYIO Ol\eHRY Me3eHXIIMHOll: TeopillI eM. y
A. r. RHoppe (1948, 1949a, 6); A. r. RHoppe II JI. n. CYBOPOBOH (1961) H
Kuntz (1954).
204
HeHp03RTo epMa bHwe CDMllaTOO aCTHqeCRHe R eTRH B ya ax CDM-
naTH'ieCROii :qeIIOqRD MOmHO HaiiTD: yme TOr a, KOr)];a HeT em;e npD3-
HaROB ]:(HBepreHD;HD ]:(ame B CnHHa bHblX Y3 aX. nOCRO bRy BWCe e-
HDe K eTOK II IIpO]:(BDmeHDe IIX R MeCTY 6Y]:(YID;IIX BereTaTIIBHblX Y3-
OB Ha1JHHaeTCH He B]:(O b HepBHWX BO OROH, a 110 CDHD;HTHa bHOH
ceTHe Me3eHXlIMHblX K eTOR, IICC e OBaTe H, BH HMO, IIpDHHMa II HX
3a MeCTHble IIpOII3BO]:(HbIe CRJIepOTOMOB. Ha paHHDX CTa]:(IIJIX B yqa- -t;
CTR8X (mepeCe eHDH» Heiip03KTo]:(epMa bHbIe 3 eMeHTW II01JTII COB-
ceM He OT HqaIOTCH OT oKpymalOm;ux RJIeTOR Me3eHXIIMW. Mop<pOJIO-
rnqeCKD HX MomHO BWHBJITb TO bKO B TOM c Y'lae, ecml OHn paclIo-
omeHW He B O]:(DHOqKY, a rpYIIllaMH D 06pa3YlOT IIpII 3TOM 60 ee II H
MeHee II OTHYIO « opomRY». B 3TDX yqacTKax R eTKH HeiipoaRTo ep-
Mbl HMelOT BbITHHYTYIO, DHor]:(a BepeTeH006pa3HYIO <poPMY. B (<KOHe'l-
HblX CTaHD;HHX» lIepeCe eHDJI, KaROBblMII HB HIOTCJI IIepH<pepDqeCKDe
BereTaTDBHbIe Y3J1W, HeiipoaRTo]:(epMa bHble R eTRH 06pa3YIOT, XOTH
H pa3pblx eHIIYIO CTPYRTYPY, HO 110 cpaBHeHDIO C K eTKaMII oKpymalO-
m;eR Me3eHXHMbl, 60 ee ROMIIaKTHYIO. ITo 3TOMY IIplI3HaRY ame Ha
paHHDX CTa]:(JIHX He D<p<pepeHD;DpoBaHHoro 3a1Jarua CIIMlIaTII1JeCRDe
raHr DD MomHO erKO OIIpe]:(e HTb, qTO 1I03BO JleT BeCTH Ha6 IO)];e-
HHH Ha]:( ]:(a bHeRIlIDM XO]:(OM DX pOCTa D D<p<pepeHD;lIpOBKD.
B OCHOBe TeopHH Balfour (1877) 0 HeRp03KTO epMa bHoM pa3-
BHTDH BereTaTHBHOR D xpoMa<p<pIIHHoR CIICTeM, DMeBllIeR MHOro1JDC-
eHHWX CTOpOHHHROB II IIoMe oBaTe eR (Mitsukuri, 1882; Babl H.,
1891; Kohn, 1900, 1903, 1907; Zuckerkandl, 1901, 1912; Wiesel,
1901; Soulie, 1903; Poll, 1906; Celestino da Costa, 1917, 1926,
1936a, b; Hett, 1925; Keene, Hewer, 1927; MBaHoB, 1930; ro y6,
1936; Coupland, 1952, 1956, 1965, D p.), emDT IIpe]:(CTaB eHDe,
BblCRa3aHHoe BIIepBwe Schenk (1876) \ 0 MHrpaD;HH K eTOqHWX 3 e-
MeHTOB 113BHe D a bHeRllIee IIX C03peBaHIIe Ha MecTax 6Y DX
nereTaTHBHblX Y3 OB.
B HaCTOHm;ee BpeMJI BOIIpOC 0 HeRp03RTO epMa bRoM IIpODCXOm-
eHDII CJIMIIaTUqecRoH II xpoMa<p<plIHHoH TKaHD MomHO CqDTaTb B
IIpHHD;JIne pemeHRblM, HO ]:(eTa H 3Toro IIpoD;ecca - TaKDe, RaR yqa-
CTHe B HHX HeRpa bHoro rpe6HH IIJTII HepBHoR Tpy6KH, OCTalOTCJI OT-
RPWTI>IMI'I. Pa3BHTDe CHMIIaTHqecRoH HepBHoH CDCTeMbl 110 OIIlIcaTe L-
HWM pa60TBM His (1897) Ha RYpDHWX 3apO WllIax D Froriep
(1907) Ha 3BpO)];bllllaX CRaTOB H RpomrnOB IIpODCXO DT D60 D3 Heii-
pa bHoro rpe6HJI D CIIDHa bHWX raHr DeB (His), H60 D3 HepBHoH
Tpy6KH (Froriep) . ITo Froriep K eTKll MDrpJIPylOT B O b lIepe -
HUX KopelliKoB, 110 Detwiller (1937) - BAO b 3a]:(HIIX, a 110 Kohn
(1907) - B]:(OJ1b Tex H ]:(pyrHX. Pacxom]:(eHDH B MHeHDHX 06 yqBCTDII
HeRpa bHoro rpe6HH II H nepBHoH Tpy6RD B pa3BHTHH CHMIIaTH'ie-
CROH HepBHOH CHCTeMbl He 61>1 II pellleHW H 3RCIIep1IMeHTa bHoii aM-
6pDo orDejj. 8KCIIepJlMeHTB bHble IIpDeMbl, ROTopble IIpmnJTII Ha CMe-
RY OIIHcaTe bHWX Dcc e]:(OBBHDH B IIepBoii qeTBepTH TeRym;ero CTO-
eTHH, OKa3a HCb II O)];OTBOpHbIMll B BblHcHeHHD MHorDX 3aROHOMep-
t Schenk, 1876 -l\IIT. no A. r. KHoppe, 19498.
205
HDCTen lIH IIBH]:(yaJIIJHOro. pa3BHTHH, HD B BDIIpooaX pa3BHTHJI
IIepHcpepHTfeCRDH HepBHDD CHCTeMbl He BHeCJlH rrDJlHDll HCHDCTH H He
YCTpaHlmH CYlI{eCTBYIOm:nx pa3HDrJlaCHn. BD3HIIRJIH HDBbIe npDTIIBD-
pe1JHBbIe cpaRTbl, RDTDpLle IIDCTaBIIJlH 110.]:( CDMHeHHe BD3MOiRHOCTb
OROHqaTeJlbHorD aKcneplDleHTaJlbHDro pellleHHJI BDnpDca 0.6 HCT01JHH-
Rax D6pa3DBaHlIH 1I0rpaHlI1JHb1X CHMnaTHqeCRHX CTBDJlOB. B DIIblTax
C y aJleHHeM y <JapolILlme:A JlJlrymRH raHrJlHD3HDn nJiaCTHHRII BMeCTe
C ADp3aJlbHDn qaCT:bIO nepBHDn Tpy6RH (Muller, Ingvar, 1921) HJlH
RaYTepH3a:qIIH HX Y RypHHblX 3apD]:(LlllIen (Muller, Ingvar, 1923,
CampenhDut, 1930), P63YJlbTaTbi RDTDpblX nD.n;TBepiR.n;eHLI B aHaJlD-
rlIqHLlX :mcnepHMeHTax (Yntema, HammDnd, 1945; HammDnd, Yn-
tema,1947), CIIMnaTnqeCKIIe3aqaTRH He pa3BHBaJlHCb. Ha aTDM ooHD-
BaHIIH 6L1J1 c]:(eJiaH Bbmo.n; 0. MHrpa:qHn RJleTDqHblX aJleMeHTDB CHMna-
TIIRyca 1Jepe3 .n;op3aJlbHLle RDpeIliRH. B TO. iRe BpeMH B pa3HbiX MD.n;II-
cplIRa:qnJlX Tex iRe :mClIepHMeHTOB, IIpD]:(eJlaHHbIX D.n;HDBpeMeHHD, HD
HeaaBIICHMo. DT aTHX aBTDpOB (Kuntz, 1926; Uchida, 1927a, b, c) II
n03]:(Hee JDnes (1937. 1939, 1941), 6b1J1D IIDRa3aHD, 1JTD pa3BHTIIe
CHMIIaTII1JeCKHX HepBHLlX 3a1JaTRDB ne HapymaeTCH II 1JTo., CJle]:(DBa-
TeJlbHO, pa3BIITHe 3J1eMeHTo.B CHMIIaTlJ<leCRo.n HepBHDn CIICTeMbl IIpD-
IICXDAIIT II3 HepBHon TPy6KH IIYTeM MlIrpa:qHH 1Jepe3 HeIIOBpeiRAeH-
HLle BeHTpaJlbHLle RDpemRII.
HeCDCTDHTeJlbHDCTL 3RcnepHMeHTaJlbHDrO pemeHIIJI IIpD6J1eMbl 0.
npDHCXOiR]:(eHHn BereTaTII-BHDH HepBHDH CIICTeMLI CTaJia D1JeBH,D;HDH.
IID1JTII D]:(HDBpeMeHHD, HD He3aBIICIIMD DT :mCIIepHMeHTaJlbHbIX pa6oT,
nOJIBIIJlIICb HDBbIe HCCJle.n;OBaH1IH OIIHCaTeJlbHDro XapaRTepa, RDTDpLle,
B DTJlII'lHe DT_ IIpe]:(LI]:(y X, 6L1J111 BbIlIOJIHeHLI He 06:r:qerIICTDJlDrn1Je-
CRHMH MeTo.n;aMH, a CIIe:qHaJlbHbIMII HenpDrHCTDJlDrn:qCCRIIMII.
]J.. It fDJly6 (1936) B pa6DTe 110 HHHepBan;IIII HaWI01Je1JHHRa nD]:(-
pD6Ho. DCTaHaBJlIIBaeTCH Ha RJleTOqHblX ]:(HcpcpepeHn;IIpOBRaX BereTa-
THBHDH CIICTeMLI B aM6pHDreHe3e RYP, TeJlHT II 1JeJlOBeRa; Juba
(1937) IIDCBJllI{aeT Crren;HaJlbHDe IICCJle]:(DBaHlIe pa3BHTHIO CHMIIaTlPIe-
CRDro IIDrpaHHQHDro CTBDJla B 3M6pHDreHe3e RYP, a Mihalik
(1940) - B 3M6pHDreHe3e RpDmrna. TDJlbRD Kuntz (1945), HaYQIIB-
IlIHH B 3TDM HaIIpaBJleHHII IIpe]:(cTaBIITeJleH Bcex RJlaCCDB nDaBDHD1J-
HbIX, npHMeHJleT (npeHMym;eCTBeHHD) rnCTDJlDm1JeCRHe RpaCHTeJlll.
HecMDTpH Ha HCIIDJlb3DBaHIIe pa3HLlX cnDc06DB llMnperHan;HII (roJly6
pa60TaJi no. BYKe-BHJlbIllDBCRDM)', Juba - no. KaxaJlIo, Mihalik-
no KaxaJlIO Ii ]J.aBeHIIDpTY), Juba H Mihalik BCJle]:( aa caMHM Ra-
mon y Cajal (1908) IIplI3HaIOT, 1JTo. MeTD]]; cepoopeHHH MaJID IIpH-
roAeH ]:(JlH pellleHHH BDUpOOOB paHHHX RJlero1JHLlX .n;HcpcpepeH:qHpDBDR.
HalI6DJlee IIJlO]:(DTBDpHbIM B aTo.M DTHDmeHlUl DKa3aJlCJI KapHDMeTpH-
1JeCRHH MeTDA, npHMeHeHHLlH A. f. KHDppe (1948), ]:(JlJI H3YQeHHH
Wlc}lcpepeH:qHpoBDR RJleTo.1JHDro MaTeplIaJla pa3HLlX 3a1JaTKDB, B To.M
qllCJle H HeHpaJlbHDro..
Pa36npaeMblH Bo.rrpDC 0 IIYTJlX MHrpan;HH RJleTo.qHDro MaTepHaJia
II3 :qeHTpaJlbHhlX yqf)CTRo.B HeBpaJlbHblX 3aKJla.n;OR He 6L1J1 rrpe.n;Me-
TDM Harnero Cnen;HaJlbHDro R3yqeHIIJI. TeM He MeHee 6oJlbmoH MaTe-
pHaJi cepHHHLlX cpeaOB aapDALlmeH Jla6DpaTDpHLlX iRlIBOTHLlX D
206
IITH:q, RO'fOpbIe HaXO]:(IIJIHCb B HameM paCIIopHmeHHH, II03BOJIHeT HaM
npHcoeAHHHTbCH R Ton rpynlIe HCCJIeAOBaIDIn (OImCaTeJIl>HLIX II 9R-
CIIepHMeHTaJIbHbIX), ROTopLle nplI3HaIOT HepBHLle BamllW II CnHHaJIb-
Hble raHrJIHII IICT01JHHRaMH nepH4>epH'leCRHX CIIMIIaTH1JeCRlIX HenpoHoB
(Balfour, 1877; Onodi, 1886; van Wyhe, 1889; His, 1890, His,
1897; Rabl, 1896; Hoffman, 1900; Held, 1909; Miiller, 1920;
Miiller, Ingvar, 1921, 1923; Gless, 1940; KHoppe, 1948, 1949a, 6,
HAP.). KJIeTRH IIOlIa]:(aIOT B 3aRJIa]:(RY CIIHHaJIbHblX raHrmIeB 110 3aA-
HHM RopelliRaM H CJIeAYIOT AaJIee B 1I0rpaHH1JHble CTBOJIbI CHMnaTll-
qecRoii CHCTeMLI. AORa3aTeJIbCTBaMH B IIOJIb3Y 3TOro IIOJIOmemm cny-
maT CJIeAYIOID;IIe Ha6JIIOAeHHH: 1) pa3BHTHe 3a1JaTROB IIepH4>epH1Je-
CIlliX CHMnaTH'leCRHX Y3JIOB HacryIIaeT paHbme, qeM MomHO ROHCTa-
THpOBaTb 06pa30BaHHe nepeAHHx RopemROB HepBHon TIJy6RH; B TO
me BpeMH yme B 3TOT paHHHll: IIepHO]:( 3M6pHOHaJIbHoro pa3BHTIIJI
110 xO]:(y 3aAHHX RopelliROB H BIIJIOTb AO 6pIOIliHo-aOpTaJIbHbIX CHM-
IIaTHQeCIUIX Y3JIOB BblRBJIHIOTCJI HeIIpepbIBHbIe «AOpOmRH» MHrpHpyIO-
ID;HX RJIeTOR, 2) Ha 60JIee II03JJ;HHX CTaAIIHX, RorAa pa3BHBaIOTCJI
nepe]:(HHe RopelliKII, B HX COC,TaBe pe]:(RO BCTpe1JaIOTCH RJIeTRH, a ecJllI
l! BCTpeqaIOTCH, TO IIpHHaAJIemaT OHH R rJIHaJIbHblM 3JIeMeHTaM.
B 3TOM OTHOllleHHH Ham MaTep1IaJI COBIIa]:(aeT C AaHHLlM1I Miiller 1I
Ingvar (1923) H f\Hoppe (1948), nORa3aBIlIHX, qTO BeJIII1JHHa HAep,
ROTOpbIe BCTpe1JaIOTCJI B lIepe]:(HHX RopelliRax, IIpH6JIHmaeTcH R pa3-
MepHOMY RJIaccy rJIHaJIbHLlX HAep TOll: me CTa]:(HH pa3BHTIIJI.
B 0630pax KYHl a (Kuntz, 1945, 1954), A. r. KHoppe (1948,
1949); JI. B. CYBOPOBOll: H A. r. l\Hoppe (1960); A. r. KHoppe
II JI. B. CYBOPOBOll: (1961) IIo]:(p06HO pa306paHa 4>aRTH'leCRaJl CTO-
pOHa 1ICTOpH1JeCIUI CnOiRlIBmHXCH HanpaBJIeHHll: 1ICCJIe]:(oBaHlIn 0 IIpO-
HCXOiR]:(eHJllI norpaH:K'IHOrO CTBOJIa CHMlIaTlIRyca 1I IIOABe]:(eHbl HTOr1I
no OHToreHOOY BereTaTHBHOll: HepBHoii CHCTeMLI y II03BOHOqHbiX iRlI-
BOTHbIX. B cOBoKynHOCT1I 3TH 0630pHLIe CTaTbH ]:(aIOT 1I0qTH 1Icqep-
IIblBaIOID;ee npeACTaBJIeHlle 06 ypOBHe HallilIX 3HaHlin B ]:(aHHOll: 06-
JIacTII.
ECJI1I 3RClIep1IMeHTaJIbHaH 3M6pHOJIOrHH He pa3pelllHJIa CIIOpHbIX
BOIIpOCOB 06 Y1JaCTHH TOro 1IJIH HHoro OT]:(eJIa :qeHTpaJIbHbiX 3aqaT-
KOB B I'HCrOreHe'3e lIep1I4>epIPlecROll: HepBHoii C1ICTeMbl, TO B OTHO-
meHH1I pa3BHTHH ee H3 Hell:p03RTo]:(epMLI pemaIOID;YIO pOJIb CblrpanH
HMeHHO 3RCIIep1IMeHTaJIbHbIe pa60TLI. Campenhout (1930), Ham-
mond II Yntema (1947) IIORa3aJllI Ha RYP1IHbIX 3apOALlmax, qTO
ABYXCTopOHHee y]:(aJIeH1Ie HepBHoro rpe6HJI BMeCTe C IIp1IJIemaID;1IM1I
YQaCTRaM1I HepBHoii: Tpy6RH BneqeT 3a c060ll: 1I0JIHOe 1ICqe3HOBeHH6
CHMlIaTlI1JeCRIIX raHrJI1I03Hb1X RJIeTOR, a TaRiRe BHYTPII- II BHeHa]:(no-
'1e1JH1IROBOll: xpoMa4>tIJHHHOH TKaHII. OARaRO, HMO CRa3aTb, 1JTO He
TOJIbRO 3TII OIIbITLI, HO II BeCb IICTOplIqeCR1IH IIYTb IIccneAoBaH1Ill:
)Toii IIp06JIeMbl ]:(ORa3blBaeT eWIHLlH IICT01JHHR pa3BHTHJI xpoMa4>4>1IH-
Hoii TRaH1I BMecTe C C1IMnaTlI1JeCROll: HepBHOH C1ICTeMOH 1I3 Heiipo-
3RTOAepMaJIbHOn 3aRJIa]:(R1I 1I OTBepraeT Y1JaCTHe Me3eHXllMLI.
B npo:qecce Bcex 3T1IX IIccneAoBaHIIH, npOBe]:(eHHLlX Ha pa3HLlX
rpYIInax n03BOH01JHLlX - OT pLl6 O 1JeJIOBeRa, YTBepA1IJIOCL npeA-
207
CTaBJIeHlIe, lITO IIOCJIe MllrpaD;DD KJIeTOK 113 D;eHTpaJIbHblX HepBHblA
aatJaTKOB Ha IIepmpepDD 06pa3YIOTCH CDMlIaTO-xpoMalfJlfJ1IHHble IWMII-
JIeKCbl, D3 KOTOpblX .n;lIlfJlfJepeHD;npyroTCJI, C O]:(HOil: CTOpOHbI, IIMIIa-
THtJeCKlIe raHrmm, C ]:(pyroil: - xpoMalfJlfJlIHHaH TKaHb. IIOqTII BCe HC-
CJIe]:(OBaTeJID, 3aHHMaBillHeCH MoplfJoreHe30M IIaparaHrJIHeB, BbICKa3b1-
Bamr KaKOe-JIH60 MHeHHe 0 pO]:(OHatJaJIbHoil: KJIeme, II3 KOTO}lOil: pa3-
BDBaIOTCJI CHMIIaTlI'IeCKDe II XpoMalfJlfJHHHbie 3JIeMeHTbl, .n;aBaJIH eil:
pa3Hble HallMeHOBaHHH - «py.n;HMeHTapHaH», «1IH]:(DlfJlfJepeHTHaJl»,
<mapaClIMlIa THKo06pa3yrom;aH», «CIIMIIaTo06pa3YIOm;aH», « CDMIIa Toro-
HDJI», (mepBII'IHaJl CDMlIaTlI'leCRaH KJIeTKa» II JW. D IIpe.n;JIaraJIH cxe-
Mbi ee .n;HlfJlfJepeHD;HpOBlm:
CIIMIl8ToroHlIH
I
!
CIIMnaT06JIaCT
!
CIIMnaTII'lecKaH
raHrJIII08H8H KJIeTKa
t
<DeoxpoM06JIacT
!
CDeOXpOMOl\IIT
(no Poll, 1906)
CIIMnaTIIQecKaH 6JIaCTeMa (blastema simpatico)
I
t
CIIMITaTOrOHlIH
!
CIIMnaT06JIaCT
!
CIIMnaTIIQecKaH
raBrJIII08HaH KJIeTKa
t
Cocep;HHH ( 06aBOqHaH)
KJIeTKa
t
CDeoXpOM06JIaCT
!
<DeoxpOMOl\HT
(no Watzka, 1943)
HeHp06JIacT
CIIMnaToroHHH
I
t
CIIMITaT06JIaCT
!
CIIMnaTIIQecKaH KJIeTRa
t
<DeoxpoM06JIacT
L
CDeOXpOMOl\IIT
(Pettinari, 1953 l\HT. no
Giordano L3llza, 1935)
IhaK, JIH60 ]:(Be (Watzka), JIH60 o]:(Ha (Poll, Pettinari) Ma-
TepDHCKaJl K,lIeTKa.
«CHMIIaTntJecKaH 6JIaCTeMa» Watzka, 113 KOTOpOil: pa3BHBaIOTCJI
06e TKaHD, aHaJIOrntJHa, BH]:(DMO, 110 CBOIIM IIOTeHD;HJIM «CHMIIaTO-
xpoMalfJlfJ1IHHoil: TKaHD» Zuckerkandl (1901, 1912), 3JIeMeHTbI KOTO-
poil: Tome IIpO.n;yU;UpyroT «CHMIIaT06JIaCTbI» H (mapaCD lIIaTo06pa3YIO-
m;De» KJIeTKD,
Em;e B 1913 r. It M. HXOHTOB OTMeTHJI, tJTO XO]:( ]:(DlfJlfJepeHII;H-
pOBKD, IIpe]:(JIOmeHHblil: Poll, HHRTO He Bu.n;eJI ad oculus, D OH
JIMeeT TOJIbKO TeOpeTlI'IeCKOe 3HaqeHHe. Ho, KaK 6y.n;eT BD.n;HO II;]
208
H1nReCJIe YIOm;ero, OH He IIMeeT If TeOpeTH'IeCRoro 3Ha1JeHlrH, TaR RaR
He COOTBeTcTByeT ]:(eHcTBIITeJIbHOCTJI. A B 1965 r. Coupland B CBO-
eH MOHorpa.p1I1i OTMeTIIJI, qTO xpoMa.p.pIIHHbIe KJIe'N\II If CIiMIIaTH'le-
CRlIe HeHpoHbl B03HIIRalOT 1I3 MeJIRIiX RHTeHCIIBHO 6a30.pIIJIbHbIX RJIe-
TOR. llo Tpa mJ;1I1I OH Ha3bIBaeT 3TH RJIeTl\lf «IIepBHqlIblMIi CHMna-
TH'leCRIIMID), HO IIPII 3TOM 01JeHb IIpaBIIJIbHO IIO,wIepRIIBaeT, qTO ]:(0
CHX 1I0p HeII3BecTHO, IIpe]:(CTaBJIJIlOT JIH OHII C060H 06m;ylO CTBOJIOBYIO
RJIeTRY, 113 ROTOpOH pa3BHBalOT'CJI 3JIeMeHTbi ]:(BYX TRaHeH (CIiMnaTII-
qeCROH H xpOMa.pqJlIHHOH), IIJIH CYlI(eCTByeT ]:(na Tlfna IIepBH'lHbIX 6a-
30qJlIJIbHbJX RJIeTO!K. llpH3HaBaJl, '1 TO HeJIb3J1 H3yqaTb Mop.poreHe3 6e3
H3YQeHlfH IICTOqIJIIROB 06pa30BaHHH 6a30qJlIJIbHblX IIepBlIqHblX cm,rna-
TlIqeCRIIX 3JIeMeHTOB, Coupland IIOJJaraeT, o HaRo, qTO B HaCTOHm;ee
BpeMJI pa3]];eJICHlie IIX 110 rIICTOJIOrlf'leCRIIM IIplI3HaRaM HeB03MomHO.
HeeMoTpH Ra RplITlIqeCRlIe 3aMe'laHIDI 110 IIOBO]:(Y reHeGa xpoMacp-
«pIIHHbIX RJIeTOR, BblCRa3aHHbIe elI(e B Ha'laJIe TeRYlI(ero CTo.rreTllJl
.HXOHTOBbIM (1913), ]:(0 CIIX IIOp HeT pa60T, B ROTOpblX cneqlIaJIbHO
o6cym]:(aJICH 6bl 3TOT BamHblH B TeOpeTllqeCROM OTHOllleHlIli BOIIpOC
(Coupland, 1965). Bo Bcex COBpeMeHHblX PYROBO CTBax II ony6JJH-
ROBaHHblX 0630pax nOBTOpJllOTCJI Bce Te me CBe]:(eHIIJI 0 rIInOTeTH'le-
CROH «CIIMIIaTOrOHHlI». I1CRJJlOqeHlIe COCTaBJIHeT mIllib pa60Ta
B. H. lloHoMapeBoH (1946), B ROTOpOH paCOMaTpliBaeTCJI TOJIbRO Ra-
pOTII HbIH IIaparaHrmIH. IIcxo]];H H3 IIpe JIOmeHHOH H. r. XJlOnlIHblM
(1946) eCTeCTBeHHOH CIICTeMbi TRaHeH, ROTOpaJl IIORa3bIBaeT Tom]:(e-
CTBO npeBpam;eHIIH 3RCnJJaHTlIpOBaHHblX TRaHeH, HMelOm;lIX e]:(IIHblll
IICTO'lHHR npOI'ICXom]:(eHHH, B. H. lloHoMapeBa IIpOBeJIa 3RCIIepIiMeH-
TaJIblJbIH aHaml3 IIPIIPO]];bl RapOTII]];HOrO IIaparaHrJlIIJI RYPIIHblX 3apo-
]:(blllIeH B YCJJOBIIHX 3RCIIJJaHTaqlfli. OHa IIplilliJIa R 3aRJIlOqeHHIO, 1JTO
RJleTRII, 06pa3YlOII(IIe lIapeHXHMY opraHa II Tom]:(eCTBeHHble no xa-
paRTepy pOCTa II npeBpam;eHlIll C T'lIIIOM pOCTa 3IIeH]:(IIMbl If rJJHII, HB-
JlJlIOTCJI IIpOII3BO]:(HbiMlI raHrJIlWHapHOii IIJlaCTIIHRII If B RaqeCTBeHHOM
OTHOllieHIIH OJJmHbI 6b1Tb OTHeceHbI R rJJlf03neH]:(HMHOMY TIIIIY rlI-
CTOJJOrH'JeCROH CIiCTeMbl H. r. XJlOIIlIHa.
HalliH c06CTBeHHble IICCJIe]:(OBaHIiJI rlICToreHe3a IIaparaHrmIOHap-
HblX opraHOB 6plOlliHOll II mellHo-rpY]];HOll 06J1aCTell 6b1JIIi IIpOBe]];eHbl
Ha IIpe]:(CTaBIITeJIHX Tpex rpYIIII n03ll0HOqHb1X: a M.p1I61IJIX , IITlIqax II
MJleROIIIITalOlI(lfX (CMIITTeH, 1958, 1961, 1962, 1963; CMIITTeH II BJIa-
OC-llpIIHMaR, 1959; C I1ITTeH II MaJiIIHOBCRaH, 1960; llplIHMaR, 1960,
1962; llplIHMaR II CMIITTeH, 1959).
B HaCTOJllI(ell rJlaBe H3J10meHbl pe3YJIbT'aTLI 3T1IX pa60T, B ROTO-
pblX Mbl IIblT3JIliCb BblHCHIITb IIplIpO]:(Y pO OHaqaJIbHOll RJIeTRII xpo-
Ma.pcpIIH06JJaCTa. B ::JTOM IICCJle]];OBaHHII Hac IIHTepeCOBaJIII, O]:(HaRO,
He TOJIbRO TeOpeTH'leCRlIe BOIIpOCbl 0 reHeTH'leCIWll CIICTeMe HepB-
HOll T'RaHH II 0 TOM MeCTe, ROTopoe 3aHliMalOT B Hell xpOMacpcplIHHble
3JJeMeHTbI. 1I3YQeHIIe rlICToreHe3a - O HO 1I3 BamHeHllIliX 3BeHbeB B
qemr Tex Mop.pOJlOrlIqecRlIx IICCJIe]:(OBaHIiH If Tex 3a]];a 1 1, ROTOpbIO
CTOHT rrplf pellieHlI1f cnopHblX BOIIpOCOB 0 «xpOMaCPCPHHHblX» If (<He-
xpOMacpcpIIHHbIX» naparaHrJlIIJlX, 0 IIX «CIIMIIaTII'leCROll» II <<IIapaCIIM-
lIaTlI'IeCROH» IIplIpo]:(e, 0 IIX «a]:(peHeprlIqeCROll» H «Hea]:(peHeprH'le-
209
CROM» 4>YHRqIIHX. B 3T1IX CnOpHblX BO'npO'CaX, RO'TO'PLle IIO'.n;PO'6HO' II3-
nO'meHbl B IIepBblX rnaBaX, ]:(aHHbIe 0'6 3M6plIoreHe3e IIaparaHrnlieB II
o pa3BIITHII CneD;1I4>H1JecRlIx KJIeTO'R Bcer.n;a IICIIO'JIb3O'BaJIIICb aBTO'pa-
Mil .n;mI ]:(O'Ra3aTeJIbCTBa TO'M IIJIII IIHO'M RO'HqeIIqlIlI. Oc06eHHO' O'CTPO
3TO OIqYID;ae'.I1CH 110 OTHO'llieHlI1O R RapOTII]:(HO'MY IIaparaHrmllO. qacTo
aBTopLl 3M6pIIO'JIO'I'II1JeCRIIX pa6O'T CTpeMIIJIIiCb HaMTII O'6'bHCHeHlIe
UiMelOID;IIMCJI pa3HO'rJIaCIIHM 0 CTPYRTYpHO'M II qIITO'XU,MlIqecROM CBO'e-
O'6pa3IIM RapOTII]:(HO'ro IIaparaHrJIIIJI, 0'6 O'CO'6eHHO'CTHX erO' pa3BIITliH
(TaR me RaR MO'P4>O'JIO'rli IJLlTaJIIiCb HaMTII cy6cTpaT .n;JIJI Tex 4>1I31I0
nOrlIqeCRIIX pe4>JIeRCO'B, RO'TO'PbIe B RapO'TII]:(HO'M O'6JIacTli O'6HapymlIJIII
4>II3IIO'JIO'rlI). EID;e Watzka (1943) O'TMeTIIJI, qTO pa3JIII1JHbIe B3rnH]:(bI
Ha CTPO'eHlIe aTO'ro O'praHa CBH3aHbi C IIpe]:(CTaBJIeHHJlMII a era pa3-
BIITlIII. B Rpyr a6cym]:(aeMLlx BO''lIPO'OO'B 6bIJIO' BO'BJIe1JeHO' npO'IICxO'm-
.n;eHlIa RapO'TH,D;HOro IIaparaHrJIHJI, IIplIqeM RO'HqeIIqlIH 0 M83eHXIIMHO'H
rrplIPO'.n;e ero RJleTO'R (BenO'it, 1928; BO'yd, 1937b II ]:(p.) O'Ra3anaCb
HaII6O'JIee CTO'MRO'M UO' cpaBHeHlI10 C MHeHlIeM 0 Me3eHXIIMHO'M IIPO'IIC-
XO'm]:(eHIIII ]:(pyrlIx IIaparaHrJIlIeB II ]:(0' CIIX IIO'P He C1JIITaeTCH pemeH-
HO'M (Adams, 1958). B IIO'.n;.n;epmRY me XPO'Ma4>4>IIHHbIM (mapaCIIMna-
TlIqeCRIIM» naparaHrnlIHM Bbl.n;BHraJIO'Cb IIO'JIO'meHlIe 0 .n;BO'MCTBeHHO'M
IIX IIPO'IICXO'm]:(eHHII Ha TO'M O'CHO'BaHIIH, 1JTO' MeJIRlIe lIaparaHrJIIIO'Hap-
IIbIe Y3eJIRII BCTpe1JaJIIiCb JIH6O' 110 XO']:(y IIapaCIiMIIaTlIqeCRHX HepBO'B,
BpaCTalOID;IIX B RapO'TII.n;HLlM IIaparaHI'JIlin (H3bIRO'-rJIO'TO'qHbIM II 6nym-
.n;alOID;IIM), JIH6O' 110' XO']:(y CIu.maTHqeCRIIX (Watzka, 1943). CJIO'BOM.
lleHTp THmeCTH H OTBeTCTBeHHOCTb 3a TY IIYTaHHqy, RO'TO'paH ]:(0' cero
BpeMeHII rocnO)];CTByeT B O'THOllieHIIH MO'p4>O'JIO'rIPIeCRO'M H 4>YHRqIIO'-
HanbHOM O'qeHRH 3TOM CTPYRTYpbl, JIemliT He TOJIbRO' Ha 3RcnepliMeH-
TaJIbHblX II O'IIIICaTenbHLlX IICCJIe]:(O'BaHIIHX OOPO'CJIblX O'Ca6eM, HO' II Ha
HCCJIe]:(O'BaHIIHX rnCTO'reHe3a.
Y1JHTblBaH, q'.I1O' 1I3yqeHIIe pa3BHTHH HepBHblx 3a1JaTRO'B Hym]:(aeT-
CH B O'6O'3Ha1JeHIIII O'T.n;eJIbHbIX cTa]:(IIM CO'3peBaHliJI RJIeTO'qHO'ro MaTe-
plIaJIa, 1I3JIOmeHIIlO 4>aRTII1JecRO'rO MaTeplIaJIa MLI npe]:(nO'CbIJIaeM ne-
pe1JeHb CyIqecTBYIOID;IIX RJIaCclI4>IIRaqIIM.
.c ]:(aBHHx IIO'P IIPH 1I3Y1JeHIIII I'HCTO'reHe3a CO'Marn1JeCRO'M II Be-
reTaTIIBHO'n HepBHO'M CHCTeMbl IICCJIe]:(oBaTenH IILlTaJIIICb Ha O'CHOBa-
HIIH MO'P4>O'JIO'rJllIeCRIIX IIplI3HaRO'B pa3.n;eJIIITb CO'3peBaHIIe HepBHblx
RJIeTOR Ha 3TaIILI. OCHO'BaTeJIII CO'BpeMeHHLlx npe]:(cTaBneHIIM 0' pa3-
BIITHII RJIeTO'qHblX 3JIeMeHTOB HepBHO'rO' 3a1JaTRa (His, 1889, 1890;
Schaper, 1897) ]:(aJIH xapaRTepliCTIIRY caMblM paHHI'IM 3TaIIaM 3TO'rO
IIPO'qecca B qeHTpanbHO'n HepBHO'M CIiCTeMe, 06O'3HaqIIB PO']:(O'Ha1JaJIb-
HbIe RJIeTRII HepBHO'M Tpy6KII «IIH]:(II4>4>epeHTHbIMII» RJIeTRaMII
(Schaper) IIJIII ,«3aqaTRO'BbIMH» (His). lIO' Schaper, «"lIH.n;n4>4>e-
peHTHbIe» RJleTRII HBJIHIOTCJI 6IIIIO'TeHTHLlMII II IIPO'H3BO']:(JlT HeMpo-
HanbHble H BCIIO'MO'raTeJIbHLle 3JIeMeHTLI HepBHO'M CIiCTeMbl, IIO' His
aHM CJIymaT PO']:(O'Ha1JanbHLlMII TO'nbRO' ]:(JIJI HeMPO'6JIaCTO'B II O'TJIH1JaIOT-
CH OT 3nIITeJIHO'IIo.n;O'6HLlX RJIeTOK, ]:(alOID;IIx rJIHIO.
YnO'MHHeM nO'IIYTHO', 1JTO' IIO'MIIMO' TID;aTeJIbHO' O'Tpa6aTaHHO'ro 4>aK-
TK'IeCRO'ro MaTeplIana, RO'TO'PblM IIO'nyqlIJIII B HaCJIe]:(CTBO aT His no-
cne.n;YIOInlIe nORO'neHIIJI IICCJIe.n;OBaTeJIen, 3Ha1JeHlIe ero pa6O'T HOCHT
I
210
nplIH HIIlIaJIbHbln xapaKTep. OCHOBHble ero IICCJleJJ;OBaHHJI IIO TCOpIIII
pa3BIITIIJI HepBHon CIICTeMbi (1889, 1890), 060cHoBaBffilIe MHeHIIC 0
rIICTOI'CHCTII'ICCKOH ('aMOCTOHTeJlbHOCTl1 HCHpoHa, Herrocpe]:(CTBCHHO
IIpe]:(IlICCTBOBaJlII cfJoPMYJIlIpOBKe Waldcyer (1891) HenpoHHoH TCO-
plIH II OKaaaJlII HCOOMHCHHOC BJlHHHIIC Ha ce 1I0CJlC]:(YIOlIte,C paaBHTHC.
B ]:(aJlbHCnIIllIX OlIlICaTCJlbHblX (Ramon y Cajal, 1906, 1908) II 8KC-
IICplIMCHTaJlbHblX (Harrison, 1904, 1907) pa60Tax TOqRa apCHHH
His nOJlY'lIIJia ffilIpOKOC paclIpocTpaHeHHc. YTBCp]:(HJlOCb rrpeJJ;cTaB-
JleHHe, qTO Ha OIIPCJJ;CJlCHHOM 8TalIC I'JI.CTOrCHe3a 6HIIoTeHTHbic HC-
roa:cfJcfJcPCHD;lIpOBaHHblC KJlCTKII nOCTCIICHHO IIpeBpalItaIOTCH B ]:(cTep-
MllHHpoBaHHble 8J1CMCHTbI II nplI06pcTaIOT CIICD;lIcfJlI'ICCKIIC MOpcfJOJlO-
rlIqCCKlIC IIplIaHaKII, IIpc]:(ffiecTBYIOIItIIC 06pa30BaHIIIO apeJlblX HCHpO-
HOB II COIIpOBO>K]:(aIOIItCH HX rJlHH. His, Ramon y Cajal, Harrison
HMeJlH CpC]:(H CBOIIX COBpeMCHHIIKOB IIpOTHBHIIKOB, KOTOpblC aalD;lIlIta-
nil CllHD;HTlIaJlbHOC CTpocHHe HcpBHoro aaqaTKa (Sedgwick, 1895;
Hardcsty, 1904) IIJlII ]:(OKa3blBaJlII HaJllIqHC OC06blX IIJlaaMO]:(CCM, co-
e]:(HHHIOlItllX KJlCTKII ]:(pyr C ]:(pyroM (Held, 1909). HOHD;elID;IIH aHTII-
HCHpOHIICTOB paa]:(CJlHJlaCb RaK 3apy6emHbIMH IICCJlC]:(OBaTCJlHMII (Bok,
1928; Bakcr, Gravcs, ,1939), TaK II OTCqCCTBCHHblMII (.HCBOHH, 1939;
1948; rOJlb]:(IIH, 1940, 1947, 1949, H ]:(p.). O]:(HaKo B HaCTOHlItec BpC-
MH paCCMOTpCHIIC 8TOH IIp06J1CMbI C HeHpoHHblX 1I0alID;Hll (Saucr,
1935a; Angulo y Gonzales, 1939; XJlOIIHH. 1946; BHHHlIKoB, 1946;
RHoppe, 1948) ]:(OJlmHO cquTaTbCH C]:(IIHCTBeHHO npaBHJlbHblM, IIMCIO-
lqRM CpC]:(II apCeHaJia apryMeHToB B CBOIO nOJibSY H 8J1CRTpOHHOMIIK-
jpOCKOlIlIqCCKIIC JJ;OKaaaTeJlbCTBa (Duncan, 1957; Panncsc, 1960;
II plIllMaK, 1963, II ]:(p.). Y JlbTpacTpyKTypHaH opraHlIaaD;IIH HepBHoro
aaqaTRa HC OCTaBJlHCT COMHeHRH B cro KJleTOqHOM CTpOCHlI1I Ha JlIO-
6blX CTa]:(IIHX paaBIITHH, B TOM qlICJle caMblX paHHIIX.
CPC]:(H OlIpc]:(eJlCHllH, KOTOpblC ]:(aBaJlHCb KJlCTKaM HcpBHoro 3aqaT-
'Ka II KOTOpblC IICIIOJlb3YIOTCH B COBpCMCHHblX pa60Tax, HaH60Jlec
Y]:(06Hbl TC, KOTOpblC oTpamaIOT 8TalIbi cfJYHRD;HOHaJlbHOrO COCTOH-
HIIH - IIJlIOpIIIIOTCHTHOCTb, ]:(cTepl\IHHaD;IIIO, cIIoc06HOCTb K IIPOJllIcfJe-
paD;HH II MHrpaD;IIII. Bcc IIpe]:(JlOmCHHblC KJlaCclIcfJIIKaD;JIH HBJlHIOTCH,
KOHCqHO, IIpR6J1R3RTCJlbHblM oTpamcHlIeM HClIpepblBHoro IIpoD;ccca
rlICTorCHe3a HepBHoH TKaHII. O]:(HaKO, XOTH OHH II HOCHT qepTbI CXC-
MarnaaD;HII, HO y]:(06Hbl ]:(JlH KpaTKoro 060aHaqCHIIH oT]:(eJlbHbiX 8Ta-
IIOB, eCJlH 8TH 8TalIbi COOTBCTCTBYIOT CTeIICHH MoPcfJocfJYHRD;IIOHaJlb-
HOll 3pCJlOCTII KJlCTKH.
Ha IIplIMCpC D;eHTpaJlbHOH HCpBHOH CIICTCMbl IIpe]:(JlaraJlOCb BbI]:(C-
JIHTb HCCKOJlbKO cfJa3, KOTOpblC B COBOKYIIHOCTII OlIpC]:(CJlHIOT RCTO-
pRIO Bccro IICpHo]:(a 8M6plIOHaJlbHOH p;mpcfJcpCHD;IIPOBKII KJleTOK B
HepBHblX aaqaTKax. Hcss (1957) Ha3bIBaCT TplI. TaKIIX CTa]:(IIH:
1) IIPOJllIcfJcpaD;RII, 2) ]:(IIcfJcfJcpcHD;IIaD;IIII, 3) pOCTa R coapCBaHIIH.
Hamburgcr (1957) aHaquTCJlbHO 60Jlcc no]:(p06HO paccMaTplIBacT 110-
CJlC]:(OBaTCJlbHbIe CTYIICHII. pa3BIITIIH CHaqaJla IIJlIOpmIOTCHTHblX 8J1C-
MeHTOB, aaTCM KJlCTOK, ]:(cTepMIIHlIpoBaHHblx B HCllpOHaJlbHOM Ha-
npaBJICHIIR, II p;aJICC CTaJ{IIII IIPOJllIcfJcpaD;lIlI, MlITpaD;IIII, 1I0HBJlCHIIH
nplIaHaKoB P;IIcfJcfJcPCHD;IIPOBKII, IIpopaCTaHIIJI aKCOHOB, pOCT, 3pCJlOCTb
II CTapeHIle HenpOHa. CTa]:(1I1I 4>YHKD;IIOHaJlbHOn aKTIIBHOCTII Ham-
burger CBH3bIBaeT C pH]:(OM 4>epMeHTHblx CIICTeM, KOTopble rrOHBJlHIOT-
CH B pacTY lIx RJleTKax. CTaAHII rrpoJlII4>epaD;1I1I II HaqaJIbHOll II4>-
4>epeHD;IIaD;1I1I (Hess, 1957) II CTa]:(IIII II.'llOpHrrOTeHTHOCTII II Heiipo-
HaJlbHOii ]:(eTepMIIIIaD;1I1I (Hamburger, 1957) xopomo YKJla blBaIOTCJI
B xapaRTeplIcTIIKy A. r. HHoppe (1948, 1949) IIpe3YMrrTIIBHoii HepB-
Hoii Tpy6KH II CTa]:(IDI raCTpYJlHD;HII. A. r. HHoppe (1948), A. r. HHOp-
pe II A. B. CYBopoBa (1959) Ha IIplIMepe pa3BIITIIH HepBHoii Tpy6KII
II raHrJllI03HOn rrJlaCTIIHKII ]:(alOT CJleJ{YIO YIO IwacclI4>IIKaD;IIIO. TIep-
Bbln IIepIIO]:( (<IIpe]:(BapIITeJlbHOii) AH4>4>epeHD;IIpODKII: 1) 4>a3a, IIpe]:(-
meCTBYIO aH ]:(eTepMIIHaD;IIII: He]:(eTepMIIHlIpOBaHHble B HeiipaJlbHOM
HaIIpaBJleHIIII KJleTRII IIpe3YMlITIIBHOii HepBHoii IIJlII raHrJllI03HOH
IIJlaCTIIHKII, 2) 4>a3a HeiipaJlbHoft ]:(eTepMIIHaD;IIII: eTepMIIHlIpOBaH-
HbIe D HeiipaJlbHOM HarrpaBJleHlI1I RJleTKII HepBHon rrJlaCTIIHKII II Tpy6-
KII (Me]:(YJlJl06J1acTbI) II raHI'JlII03HOn IIJlaCTIIHKII (l'aHrJlII06J1aCTbl).
3) 4>a3a Heiip06J1aCTlIqeCKOii II 1'JI1I06J1aCTlIqeCKOii eTepMIIHaD;lIlI:
rrpoHeiip06J1acTbI, KOTopble 110 pHJ{Y HeCIIeD;II4>II'IeCKIIX rrpII3HaROB Ha-
qlIHalOT OTJllIqaTbCH OT 8J1eMeHTOB, pa3BnBalO IIXCH B HenpOI'JllIaJIb-
HOM HarrpaBJleHIIII. BTOpoii rreplIo]:( crreD;n4>II'IeCKOn ]:(1I4>4>epeHD;IIpOB-
KII: 4) 4>a3a Heiip06J1acTa (06pa30BaHlIe HeiiplITa II Henpo4>nopIIJlJI).
5) 4>a3a pOCTa II C03peBaHIIH HenpOHa (MOJlo]:(on ]:(1I4>4>epeHD;I1PYIO-
£qlIncH HenpOH YCIIJleHHO paCTeT, IIplI06peTaeT eH]:(pIITbl, cy6cTaa-
D.IIIO HIICCJlH II rrepBbIe CIIHarrTlIqeCKlIe CBH3II), 6) 4>a3a 3peJlOCTII:
3peJlbln, 1I4>4>epeHD;lIpODaHHbln HenpoH 06J1a]:(aeT Bcen rroJiHoToii
xapaKTepHblx ]:(JlH HeiipoHa CTpyKTYP II 4>YHKD;IIii, HO IIpO OJl)IWeT
Me]:(JleHHO pacTII II lOmeT 06pa30BbIBaTb HOBbIe CBH31I.
]J;Ba IIepIIo a HBJlHIOTCH callIblll1Il BamHblMII B pa3BIITIIII RJleTOq-
HblX 8J1eMeHTOB B JlI060M 1I3 HepBHblX 3aqaTKOB - 6Y]:(b TO D;eHTpaJlb-
Hble (crrIIHlIon II 1'0JlOBHoii M03r) IIJlII IIeplI4>epnqeCKlIe (qYBCTBII-
TeJlbHbIe H DereTaTIIBHbie Y3J1bl). O]:(IIH II3 HIIX xapaKTepII3yeT co-
60ft crrocooHoCTb K HeCrreD;II4>II'IeCKOii, ]:(PYl'oii K crreD;II4>II'IeCKOH
4>YHKD;IIOHaJlbHOn aKTIIBHOCTII HeiipoHoB. H OCHOBHblM MOp4>04>YHK-
D;IIOHaJlbHblM rrplI3HaKaM rrepBOI'O IIepIIo]:(a OTHOCHTCH cIIoc06HOCTb K
MIITOTlIqeCROii aKTIIBHOCTII II MlIl'paD;IIII RJIeTOqHblX 8J1eMeHTOB (He-
CIIen;lI4>n'IeCKlIe IIOKa3aTeJlII paHHIIX 8TarrOB I'IICTOl'eHeaa), BTOpOI'O
rreplIo a - CTPYKTypHoe 4>opMIIpoBaHlIe pe4>JleKTOpHbIX J{Yr II paaBII-
Tile JlORaJlbHblX II l'eHepaJlII30BaHHblX pe4>JleKTOpHbIX aKTOB ]:(BlIra-
TeJlbHOn aKTIIBHOCTII (crreD;II4>IIqeCKlIe IIOKaaaTeJlII IIoa]:(HIIX 8TaIIOB
I'IICTOl'eHeaa) .
MIITOTIIqeCKaH aKTIIBHOCTb, a TaKata MlIrpaD;IIH KJleTOK TeCHO CBH-
aaHbl C rrpOD;eccaMII ]:(eTepMIIHaD;1I1I B HepBHOM aaqaTKe. PO]:(OHaqaJlb-
HaH Henp08KTo]:(epMaJlbHaH RJleTKa He]:(eTepMIIHIIpOBaHa B KaKOM-JlII60
Ita ]:(BYX BoaMomHblX HarrpaBJleHlIn 1I4>4>epeHD;IIpOBKII - HenpaJIbHOM
IIJIII rJllIaJlbHOM. 3a CqeT ee ]:(eJleHIIH ocy eCTBJlHIOTCH rrpOJlII4>epa-
TIIBHbln IIpOD;ecc, HapaCTaHlIe MaCCbI KJleTO'IHOI'O MaTeplIaJIa B aaqaT-
Ke, BblCeJleHlIe el'o II IIepeMe eHlIe B ]:(PYl'lIe TKaHII II opraHbI aapo-
blma. BII IIMaH ]:(II4>4>epeHIwpOBKa Ha Be KaTel'Op:n1I KJleTOK Ha'IlI-
HaeTCH e e B IIeplIO]:( rrpOJlII4>epaD;IIII, BblpamaHCb B 6HMo]:(aJlbHOCTII
212
};PIIBbIX H):\epHbiX pa3MepOB B HepBHOH TPy6Re II CIIIIHaJIbHblX l'aHr-
JIIIHX (RHoppe, 1948, 1949) II B CTpYRTypHO- I1TOXIIMlIqeCROH AHcfJefJe-
peH IIpOBRe H):\ep B CIIIIHaJIbHblX II rreplIcfJeplI ecRIIx CIIMTIaTII eCKlIx
Y3JIaX (CMIITTeH, 1958, 1962). XapaRTepHblMII CBOHCTBaMII Bcex RJIe-
TOqHblX IIOIIYJIHD;IIH 8Toro rreplIo]:(a HBJIHIOTCH MIITOTlIqeCRaH aRTIIB-
HOCTb, rrpoJIlIcfJepaD;IIH II MlIrpaD;IIH, COBIIa):\alOm;IIe C TaROBblMII cpe):\R
He]:(lIcfJcfJepeHD;IIpOBaHHblx 8JIeMeHTOB.
TaRIIM 06pa30M, MopcfJOJIOrlIH H]:(pa OIIpe]:(eJIHeT rrYTb pa3BIITIIH.
)J;JIH HeHpaJIbHOH reHepaD;IIII RJIeTOR, ROTOpbIe BCTYIIIIJIII Ha IIYTb ):\e-
TepMIfHlIpOBaHHoro pa3BIITIIH II IIplI06peJIH BII):\IIMYIO MOpcfJOJIO-
rlI eCRYIO ):\IIcfJcfJepeHD;IIpOBRY, no IIpO]:(OJImaIOT pa3MHomaTbCH, IIpe):\-
JIOmeH TepMIIH IIpoHeHp06JIaCT (Mihalik). ;:hOT TepMIIH y]:(06eH ]:(JIJI
0603HaqeHIIH RJIeTOR 8TOH CTa):\IIII, AeTepMIIHlIpoBaHHblx B HeHpaJIb-
HOM HaIIpaBJIeHIIII. 110 MopcfJOJIOrlIQeCRIIM IIplI3HaRaM His CqIITaJI
qTO e]:(I1HCTBeHHbIM RplITeplIeM, He OCTaBJIHIOm;IIM COMHeHIIH B HeHpo-
6JIaCTIIqeCROH IIplIpo]:(e RJIeTRII, MomeT CJIymnTb OTpOCTOR, BbipacTaro-
m;nii n3 rreplIRaplIOHa. B TOM MeCTe, r]:(e TeJIO RJIeTRII IIepeXOJ{IIT B
HepBHblH OTpOCTOR, RaR IIpaBlIJIO, Ha6JIIO]:(aeTCH I1HTeHCIIBHoe ORpa-
IlIIIBaHlIe IWTOIIJIa3MbI. no HallilIM Ha6JIIO]:(eHIIHM, 8TOT rrplI3HaR o .
HOCIITCH R cpaBHIITeJIbHO rr03]:(HIIM 8TarraM C03peBaHIIH rrpOHeHp06JIa-
CTa. 3HaqlITeJIbHO 60JIee paHHIIH BblpamaeTCH B CTPYRTypHOM II D;n-
TOXlIlIlHQeCROM I13111eHeHIIII H):Iep, H]:(pblllleR II xpOMoD;eHTpOB. Ho B JIIO-
6011I CJIyqae, Aame TOI'):Ia, Iwr]:(a ROHyco06pa3Hoe Y):\JIIIHeHlIe TeJIa
rrpoHeiip06JIacTa, 06YCJIOBJIeHHOe pOCTOM OTpocTRa, MeHHeT ero cfJop-
MY, MIITOTlIqeCRaH aRTIIBHOCTb coxpaHHeTCH. MIIT03bl 1II0mHO BII]:(eTb
II Tor]:(a, Ror]:(a HaJIIID;O pOCT H]:(ep IIpOHeHp06JIaCTOB II, CJIe]:(OBa-
TeJIbHO, MeHHIOTCH I1X pa3MepHbIe COOTHOllieHIIH C H]:(paMII RJIeTOt(
rJIIIaJIbHblX reHepaD;IIH. Ho 8TO IIOCJIe]:(Hee MomHO OTMeTIITb TOJIbRO
B caMOM HaqaJIe. IIpoHeiip06JIaCTbi crroc06Hbi R IIIHTOTIIqeCROIIIY
pa3MHomeHIIIO ):\0 Toro IIeplIo]:(a, rrORa B HIIX He pa3BIIT HeiipocfJlI6plIJI-
JIHpHLlii arrnapaT II HeT HCHO BblpameHBblX OTpOCTROB. C rrOHBJIeHlIeM
IIMrrperHllpyrom;IIXCJI cepe6pOM OTpOCTROB, HemHon HeHpocfJ1I6puJIJIHP-
HOD ceTim BORpyr H):\pa, ROTopoe R 8TOMY BpeMeHH CTaHOBIITCH rry-
3blpbROBII]:(HbIM II CBeTJIbIM, MIIT03bl rrpeRpam;aIOTCH. RJIeTRa IIepexo-
):lilT B CTa]:(IIIO HeHp06JIaCTa II HacTyrraeT IIeplIO]:( crreD;IIcfJlIqeCROH
]:(IIcfJcfJepeHD;IIpOBRII RJIeTRII.
Henp06JIaCT He ]:(eJIIITCH 1, HO rro OIIlIcaHHIO A. f. Rnoppe (1948,
1949) coxpaHHeT Ha HeRoTopbln IIeplIO]:( BpeMeHII crroc06HoCTb R
MllrpaD;IIII. Y Hac HeT 06'beRTIIBHbIX AaHHblX, ROTopble 6b1 rr03BOJIIIJIII
orrpe]:(eJIeHHO BblCRa3aTbCH B OTHOlIIeHlI1I crroc06HOCTII HeHp06JIaCTa R
rrepe]:(BlImeHIIIO. B TO BpeMH RaR Mbi y6e]:(IIJIIICb B IIOTepe MIITOTIIqe-
CRon aRTIIBHOCTII Henp06JIaCTOB (IIpII II3yqeHIIII ceplIHHblX cpe30B 3a-
pO]:(blllleii MHorlIX BII]:(OB mlIBOTHblX, II, rJIaBHOe, IIpII aBTopa):\IIOrpa-
cfJlI'IeCROM IICCJIe):\OBaHIIII RYPIIHblX 3apO]:(bllllen), TPY):\HO CYAHTb 0 CO-
xpaHeHlI1I IIJIII rrOTepe Henp06JIaCTaMII MID'pa IIOHHOn crroc06HOCTII.
1 B pa60Te He paCCMaTpHBaIOTC-H Te ep;Hnll'lHlde HCCJlep;OBanHH, B KOTOpldX BId-
Cli:a3IdBaeTC-H B03MOlliHOCTb MIlTOTUqeCKOrO p;eJleHHH, HR061d IIpHc aH p;alli&
3peJlldM HeHpOHaM B3pOCJlOro opranll3Ma.
213.
MomHo TOJILKO IIpe]:(nOJIaraTb, QTO 11.0 Toro MOMeHTa, IIOKa ueiIpo-
CJIaCTbl OTHOCIITeJIbHO eJIJ;e «cB060]:(Hbl» OT CBH3eiI C oKpymalOJIJ;llMll
TKaHHMll II ]:(pyr C ]:(pyroM, OHll MOryT nepeCeJIHTbCH. B TeqeHlle 3TO-
1'0 IIeplIo]:(a MeHHeTCH BeCb CTPYKTYPHbliI 06JIIIK HeiIp06JIaCTa: pa3Bll-
1IalOTCH HeiIpo!jJ1I6pllJIJIbI, IIOHBJIHeTCH TllrpOll]:(, paCTYT II AH!jJ!jJepeu-
n;HpYIOTCJI OTpOCTKll, !jJOpMlIpYIOTCH MemHeiIpOHUbIe II TKaHeBbie CH-
HanTUqeCKII€ npll60pbl - HeiIp06JIaCT IIpeBpaJIJ;aeTCH B HeiIpOH.
BTOpoiI IIepllO]:( ]:(II!jJ!jJepeHIJ;llpOBKII HeiIpoHOB HaMIl He paCCMaT-
pIIBaeTCH, TaK KaR He IIMeeT npHMoro OTHomeHIIH I\ pa36IIpaeMoii:
np06JIeMe. J(JIH nepBoro neplIoAa B ]:(aHHoiI pa60Te IIpllHHTbI cJIel1;yro-
JIJ;IIe 0603HaqeHHH oT]:(eJIbHbiX STanOB l1;ll!jJ!jJepeHIJ;llpOBKII HeiIpoHa:
1) HeAll!jJ!jJepeUIJ;llpOBaHHaH KJIeTKa, 2) npoHeiIp06JIaCT, 3) Heii:po-
06JIaCT, 4) 8M6plIOHaJIbHbliI neiipoH. YqllTblBaH IIIHor006pa31le OIJ;eHOl{,
.a TaKme ]:(PyrlIx TepMllHOB, Mbi orOBaplIBaeM IIpll3HaKII, KOTOpblMII
xapaRTeplI3yeM Ram]:(blii II3 STIIX DTanOR.
H e ]:( II !jJ !jJ e p e H IJ; II P 0 B a H H bl e K JI e T K II HepBHoro 3aQaT-
Ra-STO MeJIKlIe O]:(HOpO]:(Hble KJIeTRII, KOTopble xapaKTepll3ylOTCH
Y3KOii 30HOD: IJ;IITOnJIa3Mbl II paBHOMepHblM pacnpeAeJIeHlleM xpOMaTII-
Ha B H]:(pe. Mx rJIaBHbIe !jJYHKIJ;IIOHaJIbHble IIplI3HaKII - 6llnOTeHT-
HOCTb, CIIoc06HOCTb K MIITOrnqeCKOMY ]:(eJIeHIIIO II MlIrpaIJ;lllI.
II p 0 H e iI p 0 6 JI aCT bI - 8TO KJIeTKll, ]:(JIH KOTOpblX xapaKTepHO
nepepaCIIpeAeJIeHlle xpOMaTllHa B H]:(pe, ROHIJ;eHTpaIJ;llH ero BOKpyr
.HApblmeK, IIOCTeneHHoe YBeJIlIqeHlle pa3MepOB H]:(ep, pOCT H]:(pblmeK II
-o6pa30BaHlie KOHYCOBll]:(HOD: !jJOpMbl KJIeTOK, 06YCJIOBJIeHHOH pOCTOM
.aKCOHa. Mx rJIaBHbIe !jJYHKIJ;llOJIaJIbHble IIpll3HaKII- l1;eTepMllHllpO-
BaHHoe pa3BIITIIe B HeiipaJIbHOM HaIIpaBJIeHlI1I II coxpaHeHlIe CIIoc06-
HOCTII K MllT03Y II MlirpaIJ;IIII.
H e iI p 0 6 JI aCT bl xapaKTeplI3YlOTCH KpyrrHblM, CBeTJIblM, IIY-
3blpbKOBllAHbiM HAPOM, pOCTOM aKCOHa II nOHBJIeHlIeM B IIeplIKaplIOHe
HemHoii ceTKll neiipo!jJll6pIIJIJI. I1x maBHbIe !jJYHKIJ;IIOHaJIbHbIe npll-
.3HaRII - IIOTepH CIIoc06HOCTII K MllTOTllqeCKOMY AeJIeHIIIO, HO, B03-
.MOmHO, em;e II coxpaHeHlIe MlIrpaIJ;IIOHHbIX CBoiicTB.
;3 M 6 plIO H a JI b H bl e Hen p 0 H bl xapaRTepll3YlOTCH pOCTOM OT-
pOCTKOB, 06pa30BaHlleM HepBHblX BOJIOKOH II CIIHaIITllq€CKIIX CTPyR-
"TYP (CJIe]:(OBaTeJIbHO, 3aMbiKaHlleM pe!jJJIeKTOpHbIX Ayr) II IIOHBJIeHII-
.eM cy6cTaHIJ;IIII HllCCJIH. I1x rJIaBHbIe !jJYHKIJ;HOHaJIbHble IIpII3HaKII-
CIIoc06HOCTb K npOBe]:(eHIIIO HepBHoro B036ym]:(eHllJI II ocym;eCTBJIe-
Hmo pe!jJJIeKTOpHbIX aKTOB.
Mop!jJOJIOrlI'leCKaH xapaKTepllCTIIKa rJIllaJIbHOn reHepalJ;lllI 3Ha-
'llIITeJIbHO CJIOmHee II3-3a He3HaqHTeJIbHblX npe06pa30BaHIIii 8TllX KJIe-
"TOK. no aHaJIOrlIJI C HenpaJIbHblMH SJIeMeHTaMll HX MomHO Ha3BaTb
npOCIIOHrH06JIaCTaMll, B03HHKalOJIJ;llMJI B pe3YJIbTaTe AII!jJ!jJepeHlJ;ll-
PYIOJIJ;IIX MllT030B 6IInOTeUTHon He]:(II!jJ!jJepeHIJ;IIpOBaHHoii KJIeTRH,
oCIIOHrlI06JIaCTaMII II 3M6pnOHaJIbHOii rJIneii. O]:(HaKO, eCJlH IIpocnOHrJlO-
6JIaCTbI OTJIllqalOTCH OT IIponenp06JIaCTOB CTPYKTYpoii xpOMaTlIHO-
BbiX I'JIbl60K II paBHOMepHblM paCIIOJIOmeHUeM lIX B HApe, TO OTJIlfqlITb
HX OT HeAII!jJ!jJepeHIJ;IIpOBaHHbIX KJIeTOK Ha paHHHX cTaAIIHX rlICTore-
ne3a HepBHblX 3aqaTKOB HeB03MomHO. Ha 60JIee II03AHHX cTaAHHx
.214
IIepH!jJeplIqeCKaH I'JIIIJI, B OTJlHqHe OT D;eHTpaJlbHOH (HaIIplIMep, aCT-
pOD;IITOB), TOme He HMeeT xapaKTepHblX CTPyKTYPBblX qepT, CBOHCT-
BeHHLlx TOJlbKO eH Op;HOH. Ho IIOCTeneHHO nOHBJlHeTCJI Hap;emHblH
KplITeplIH ]:(JlH ee onpe]:(eJleHlIH B raHrJlHHX II HepBHblX CTBOJIaX B BII]:(e'
xapaRTepHoro paCIIOJlOmeHIIH OKOJlO HeHpOHOB, B KaqeCTBe CaTeJlJlII-
TOB II llIBaHHOBCKIIX :meMeHTOB. llOJlBJleHlIe 8Toro IIpll3HaKa B aaqaT-
Kax HepBHblX Y3J10B COBIIa]:(aeT 110 BpeMeHH CO CTap;IIeH 06paaoBaHIIJI"
HeHp06J1aCTOB II IIOTOMY MomeT C IIOJlHLlM IIpaBOM 6blTb oxapaRTepll-
aOBaHO CTa]:(HeH CIIOHrII06J1aCTOB.
Pap;H 06'beKTIIBHOCTII cymp;eHIIH, a TaKme B03MomHOCTII YHII!jJK-
D;lIpOBaTb IIpOD;eCCLI paaBIITIIH KJleTOK II IIpllaHaKH IIX Mop!jJOJlOrH'le--
CROH ]:(1I!jJ!jJepeHD;IIpOBKII B 3BOJlIOD;IIOHHOM pHJU' II03BOHO'IHLlX, 6b1JJ'
IIpOBe]:(eH D;IITOJlOrllqeCRIIH aHaJllI3 Y npe]:(CTaBIITeJleH Tpex rpynll
II03BOHOqHbiX - MJleKOIIIITalOID;IIX, IITIID; II aM!jJ1I6nH. Y aapOp;LImeii:
MJleKOnlITalOID;nx (KpbICbl, Mblmll, KpOJllma) II IITIID; (Kypbl) II3yqeH
rIICTOreHea 6plOmHbiX II meHHblX naparaHrJllIeB, y aM!jJ1I6IIH (JlHrym-
KII) - 6plOmHbIX.
Bbl60p mlIBOTHblX 6blJl 06YCJlOBJleH paaHLIM xapaKTepOM !jJ0PMII--
pOBaHHH naparaHrJlIIOHapHOH TRaHII B neplIop; 8M6plIOHaJlhHOrO paa-
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rJlbl6KII !jJeJlbreH-IIOJlOmIITeJlbHbIX xpOMOD;eHTpOB MorYT JlemaTb-
3J(eCb B6J1lI31I H]:(epHOR 060JlOqKII IIJlII B JIl060M ]:(pyroM Y'laCTKe XPO-
MaTIIHOBOR ceTII. qlICJlO IIIIPOHIIHO!jJIIJlbHbIX H.n;pblmeR 3]:(eCb 60Jlbme
H pa3MepbI IIX MeHbllle. RIITH xpOMaTHHa MeCTaMII IIMelOT rpy6b1c
CqepTaHHH II 1I0]:('1epKIIBaIOT TeM caMblM PIICYHOK Bcero ROMIIJleKCa
xpOMaTIIHOBblX CTPYKTYP H]:(pa-rJlbI60K, aepeH, BOJlOKHIICTLIX 06paao-
BaHIIR, paccpe.n;OTO'leHHblX 110 nceR IIJlOCKOCTII H.n;pa, BII]:(IIMOi'I Ha cpe-
ae (pIIC. 72, A).
reHeTH IeCRaH CBH3b H]:(pbllllel{ C XpOMOCOMaJlbHLIM alIlIapaTOM Y
napOCJlblX oc06eii (Brachet, 1957) H B 8M6plIonrne3e (Caspersson.
1950) ]:(aBlIO n;meCTHa. no HalllIIM Ha6J1IO]:(eHIIHM, IIO Mepe pa3BIITHJI
Hei'Ip06J1aCTII'IeCKIIX H cIIoHrlI06J1aCTIIqeCKHX 1I0IIYJlHD;IIR lIapauyK--
JleOHapHblH xpOMaTIIH pacxo.n;yeTcH, B TO BpeMH KaK HAPbllllROBblH
alIlIapaT YBeJllIqlIBaeTCH B pa3Mepax. O.n;HaKO B pa3HbiX KJleTOqHbIX
IIOIIYJlHD;HHX ero YBeJlHqeHlIe IIpOHBJlHeTCH no-pa3HOMy: B HeRp06J1a-
CTax pacryT H]:(PbIIllRJI, B CIIOHrlI06J1aCTaX YBeJlII'IHBaeTCH IIX '1IICJlO.
B IIOCJle]];HIIX MHomeCTBO XPOMOD;eHTpOB II paaHo06pa3IIe pa3MepOB
JI.n;pLlIIIeK, Hepe.n;KO BblHBJlHeMblX B BII.n;e TOqeqHOH 6aao!jJIIJlIIII BIIYTpn:
crYlIteHIIH xpOMaTIIHa, ROTOpbIe, BCpOHTHO, MomHO paCCMaTpHBaTb R
KaqeCTBe CTa]:(JIH coapeBaHIIH H]:(pblmeK.
B peaYJlbTaTe lIepepaClIpe]:(eJleHHH H.n;epHoro xpOMaTIIHa B COCTa-
Be ClIlIHaJlbHblX yaJlOB Ha paHHIIX CTa]];IIHX pa3BIITIIH OlIpe]:(emIIOTCH
]:(Be Mop!jJOJlOrlIqeCKII paaHbIe rpYIIIIbl KJleTOK, OTJllIqalOlItlIeCH ]:(Pyr
OT .n;Pyra IIO IIHTeHCIIBHOCTH OKpaCRII H]:(ep. B KJleTRaX, B ROTOpblX
H.n;epHblH xpOMaTIIH ROHD;eHTpHpyeTcH B II36paHHbiX yqaCTKaX, B6J1H-
3II KPYIIHblX H]:(pblmeR, IIHTeHCIIBHOCTb OKpaCKII H.n;ep CJla6aH, B TO,
BpeMH R.aK B ]];pyrlIx, y ROTOpLIX xapaKTepHblM CTPYKTypHblM IIplI-
aHaROM HBJlHeTCH paBHOMepHoe paClIpe]:(eJleHlIe xpOMaTIIHa B HApe II
r]:(e XpOMOn;eHTpbl CBHaaHbl Mem.n;y C060R 60Jlee rpy6b1MII II TOJlCTblMIl
HIITHMH, peaRn;HH <DeJlbrHHa 60Jlee IIHTeHCIIBHa - H]:(pa OKpamllBalOT-
CH TeMHee.
Mbl HeO]:(HOKpaTHO OTMeqaJlII (CMIITTeH, 1958, 1962; llplIRMaK II"
CMIITTeH, 1959), qTO II3MeHeHHe IIHTeHCIIBHOCTII !jJeJlbreHOBcKoll
,mpaCKII H]:(ep B neplIo.n; paHHero rlICToreHeaa HepBHblX yaJloB He CO-
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qlIHe H]:(pa nplI06peTalOT pa3HYlO IIHTeHCIIBHOCTb ORpaCRII ( CM.
plIC. 72, A). O.n;HaKo, yqlITblBaH OTBeTCTBeHHOCTb TaRoro BblBo.n;a,
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HTeJlbCTBO, '1TO II3MepeHHH KJleTO'lHblX H]:(ep He IIpolIaBo.n;IIJlOCb, MO-
meM TOJlbRO IIpe.n;IIOJlOmIITeJlbHO CY]:(IITb 0 IIplIqlIHaX BII]:(IIMOR ]:(1I!jJ-
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pacIIpe.n;eJleHIIH xpoMaTHHa II KOHD;eHTpaD;IIII ero B O'leHb MaJlblX 110.
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HlleM BeJllIqllHbI JI ep II COOTBeTCTBeHHO 113MeHeHlIeM KOHn;eHTpa IIII
B HIIX xpoMaTIIHa.
TaR RaK IIORa3aTeJlII pa3MepoB H pa He IIfpaJIH pellialOlI\eii: pOJIH
JlH eJleii: HaCTOH efO IICCJle OBaHIIH, TO IIpOCIIOHflI06J1aCTbi Mbl Ha-
3b1BaeM «TeI\IHOH epHbIMII», a IIpoHeii:p06J1aCTbl «CBeTJlOH epHbIMH»
RJleTKal\1II B RaqeCTBe MopqJOJlorllqeCKIIX OlIpe eJleHllii, ROTopble CJlY-
maT IIpll3HaRaMII HaqaBllIeiicH lHpqJepelI lIpOBKII RJleTOqHblX CTPYK-
TYP B 3aqaTl{e HepBHoro Y3J1a. 113 Ha6J1IO eHllii: aJlbHeii:IlII1X cTa lIii
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XpOMaTilIla (1)
2 - Hnepll3H 060JIO'lH8. 3 - RnpbIlliRO (no Hertl, 1957
pa3BIITliH CTanOBIITCH HCHO, lTO 113 KJleTOK CO CBeTJlbll\1II H paMll
cPUpMnpyeTCJI HeiipaJlbHaJl nOnYJlH IIH RJleTOK, II3 RJleTOK C TeMHbl-
MIl H IJaMII - fJlIIaJlbHaH nOnYJlH I1H.
Y UC:mqeHlle pa3Mepa H pblllleR B paCTY IIX Heiip06J1aCTllqeCKIIX
IIOnYJIHU;IIHX II BblCOKaH IIlIpOnllHOcPlIJlIIH IIX 6b1J1II YCTaHOBJleHbl II
PYrIIl\m aBTOpaMH (llilipHona, 1955; JIenllHcoH C COTp., 1956; Ro-
POqRIIH, 1965). 8TH IIpll3HaRII CnlI eTeJlbCTBYIOT RaR 0 HaROnJleHIIH
XlIl\lIIqeCKH aRTIIBHblX Be eCTB (RnCJlbIX 6eJlKOB II PHR JI pbIIllKa),
Tall: II 0 paCTY I1X ROHTaKTax IIonepXHOCTII H pblllleR C napaHYRJleO-
JlHpnbIM xpOMaTIIHOM. YBeJlIl IeHIIe qllCJla H pbllIleR B H pe, Ha6J11O-
p;aeMoe B fJlII06J1aCTaX, Tome CJlymIIT nblpameHlIeM HaROIIJleHIIH Be-
eCTn BHYTPII H pblllleR II B e e 60JlblIleii CTeIIeHII YBeJlIlqlIBaeT
IIonepXHOCTII conpIiROCHOBeHIIH, cnoc06CTBYH TeM caMblM paCllIllpe-
HIIIO ROHTaRTOB C HAepHblM COAepmllMblM (Hyden, 1943; Caspers-
son 1941, 1950; Brachet, 1957; ReAPoBcRIIii:, 1959). B opnrllHaJlb-
HOii: pa60Te Heltl (1957) O Ienb AeMOHCTpaTnBHO nOl\a3aHO
(pIlC. 73), qTO xpoMaTIIH, Henocpe CTBellHO IIpmlbnmlO I1ll R H -
PbllllRY II 06b1qHO 0603HaqaeMblii IIapaHyRJleapHblM, TaR II XpmIaTIIH,
KOHTaRTlIpYIO lIii C 060JlOqRoii: H pa, I\IOPcPOJlOfllqeCRII CBH3aHbi Mem-
;J;Y c060ii: U;e.iIbIM PHAOM IIepexo HblX cPOpM II no CYTII eJla HB.iIHIOTCH
e1J;HHOll clIcTeMoii.
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CTBO rcoii: Cnc.TeMbl, YCTaHOBJleHHOe B CBoe BpeMJI Caspersson (1941)
II pa3BIITOe B IIOCJle YIO IIX pa60Tax. npllQeM CJleAyeT OTMeTIITb, qTO
YHRU;IIOHaJlbllaH CBH3b 3TIIX CTPYKTYP IIMeeTCH HaJlIIU;O B JlI060M
cnyqae - ocy eCTnJIHeTCH Jill I\lem y IUlMIi Henocpe cTBeHHblll ROH-
221
TaKT IIJlII 3TII CTPYRTYPbl IIpOCTpanCTBeHBO pa3p;eJleHhI (Rep;poBcKHH
1959).
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pa3Hoo6pa:me CTPYKTYPHblX KOHTaKTOB Jlp;pblllleK II xpOMaTIIBa, Ba-
plIalUIII B pa3Mepax II qIICJle Jlp;pblllleK CJlep;yeT, OqeBHp;BO, pac-
CMaTpHBaTb RaK Mop4>OJlOrlI'IeCROe BblpameBlIe MeTa60JllI'IeCRIIX ROp-
peJlJln;IIH 3TIIX CTpYKTYp. B p;HBepreBTBO pa3BIIBalOII{HXCJI RJleTKaX
YKpYIIHeHHe Jlp;pLlllleK B OP;BIIX CJIYqaHX II YBeJllI'IeHHe HX qlICJla B
P;Pymx Be MomeT, BIIP;IIMO, CJIYmlITb TecTOM P;JlH OIIpep;eJleBIIH 4JYBR-
n;IIOHaJibBoro Ba3BaqeHHH 3TIIX CTPYRTYP B TOM II P;PYI'OM cnyqaHX.
If Te II APyrlIe RJleTKlI, 6yp;yqII 3M6pIIOHaJlbBbIMH, CIIBTe3IIpyIOT 6e J
JlOK. TaM II TYT peqb IIp;eT 06 IIHTeHcH4JIIRan;HH CIIBTe3a 6eJlRa
(Brachet, 1957) B paCTY eH BepBBoH TKaBII. Ho yqacrne PHR
Hp;pblllleK B IIpOn;eccax CIIBTe3a BblpamaeTCH, B03MomBO, B OP;BOM
cJlyqae B pOCTe pa3MepOB IIX, B P;PyroM B YBeJllI'Iemm qHCJla. Halll
MaTeplIaJl Be rr03BOJIHeT CY]:(HTb n TOM, qeM OIIpe]:(eJiHeTCH IIX pa3Boe
Mop4JoJlOrHqeCROe BblpameBHe R KJleTOqBLlX IIOIIYJlHn;HHX HepBBoro
3aqaTRa. B BaCTOHlI\eH pa60TA ::!TOT IIplI3HaK IIplIBHMaeTCH B Ka-
qecrne Mop4>OJlOrlI'IeCKOll xapaKTepHCTIlIm p;II4>4>epeBn;upOBKII KJle-
TOqBLlX CTpYKTYP, 3aKOBOMepBO IIOBTOpHlOlI(HllCJI Y pa3HbiX BHp;OB
mlIBOTHblX II, KaR 6yp;eT BIf]:(1I0 naJlbllle, IIMelO lIll MeCTO B 3aqaTKaX
Bcex BepBHbJX Y3J10B, B TOM qUCJTf'! U CHMrrarnqeCKIIX.
,D;aJlbHeHllIee pa3BIIrne II pOCT RJleTOR B qYBCTBIITeJlbBhlX Y3J1aX
BblpamaeTCH B IIOHBJleBlI1I pHp;a Mop4>oJlOrlIqeCKIIX IIplI3HaKOB, KOTO-
pble rrplIBop;HT R 4>opMlIpoBaBIiID xapaKTepmlX H MBorORpaTHO OIIII-
caBBLlX KJleTOK - HeHp06J1aCTOB H CIIOBm06J1aCTOB, BXOP;H HX B CO-
CTaB cnlIHHOM03rOBbiX Y3J10B II03BOBOqBbiX mIIBOTHbIX, B qaCTHOCTII
MJleKOIIHTaIOII¥Ix. B BellpaJlbBbJX 3J1eMeHTax YBeJlUqlIBalOTCH pa3Me-
phi JI]:(ep, IIap;aeT KOHl eBTpan;IIJI ,D;HR, paCTeT rreplIHYImeapHaH 30Ba
n;IITOIIJla3Mbl (CM. plIC. 72, B, B), 4JOPMUPYIOTCJI OTpOCTKH II HeH-
po4J1I6plIJlJlHpBblH arrrrapaT (pIIC. 74). B Kamp;OM Jlp;pe OP;BO IlJI11
ABa Jlp;pbJIlIKa, KaK IIpaBHJlO, pe3KO rrupoHIIHo4JIIJlbBble, IIpO]:(OJImaIOT
YKPYIIBHTbCJI. CJIa60 OKpameHHbJe, BemBble HIITII XpOMaTlIHa 06pa-
3yIOT 60J1ee IIJIII MeBee paBBOMepBO H8THHYTYIO ceTb. XpOMOn;eBT-
phi pep;KII, a OCJlII BCTpeqaIOTCH, TO IIpep;CTaBJleBbl 6J1ep;HbIMII, OqeBb
MeJlKIIMII Y3eJlKaMII. HYKJleoJlHpHblH xpOMaTIIH cJla60 BblpameB.
Mem,u;y BeJIHq}{BOH Hp;pbllllKa II OKOJlOHp;pbIIllKOBLIM xpoMaTImOM B
rrpon;ecce pocTa Ba6J1IOp;aeTCJI 06paTBaH 3aBIICIIMOCTb: qeM 60JThllle
HP;pbIIllKO, TeM MeBbllle Macca XpOMaTlIHa IIpllJleraeT K HeMY. Co3p;a-
eTCH BIIeqaTJleBlle, qTO HYlmeOJlHpHbill xpOMaTIIB «pacxop;yeTCH& Ha
pac 1O Maccy Jlp;phlllleK. 3'1'0 Ba6J1IOp;eBlIe BaXOp;IITCH B rrOJIBOM
COrJlaCHH C 1I3BeCTHbiMII p;aBHbIMU 06 yqacTlI1I Hp;epBoro XpOMaTIIH8
B reHe3e Hp;pblllleK (Caspersson, 1941, 1950; Painter, 1955; Brachet.
1957; Hertl, 1957; Rep;poBcKIiH. 1959).
Pa3Mepbl rJllIaJlbHblX 3J1eMeBTOB YMeBblllalOTCJI, KOBn;eBTpan;HJI
,D;HK B Hp;pax llOBbllllaeTCH, 30Ha n;IITOllJla3MLI y6bJBaeT II K KOHL(Y
9M6pllOHaJibBoro pa3BIITIIH oKpymaeT TeMHbie Hp;pa B BII]:(e Y3Koro
o60AKa (CM. pHC. 72, B, B). Ro BpeMeHH 4JopMHpoBaBIiH rIICTOJlOrlI'Ie-
222
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HOCTII IJ;UT(lIIJla;!MhI, 3a'WTHn 0Y)J,YIlI.nx CUMnaTIPleCIUIX Y3J10B MnmHO
uTlliI'IIITh cpe]:(I1 OI{pymaIOu oll rnHl{IITnaJlhllOii Me3eUXIlMHoii TImHn.
IIpOI\CCC ]:(uncpreHTIIOrO pac'men£'nrm I\JIeTO'lHhIX ;)JJeMCIITon CIlM-
naTlI'I£'CIHIX ya.TIon nOrpalllI'I1IOrO CTBOJJa n npuuIJ;nrre no.n;'1UHneTCH
223
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3M6plIoreHe3a II 3aTpY]:(HIIJICH BblCKa3aTbCJI CROJIbKO-HII6YAb OlIpeAe-
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6JIaCTOB, IIpe]:(IIOJIaraH B03MomHOCTb HX pa3BIITIIH 113 Me3eHXIIMbl.
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BalOm;ero :qIIToreHe3 xpoMacl>IfJIIH06JIaCTOB OqeHb 1I0]:(p06HO, paCIlIHpH-
eT IIpe]:(cTaBJIeHIIH 06 3TlIX IIpo:qeccax II BlIepBble BHOCIIT HOBoe B
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CJIeAOBaJIII. Hett HaqlIHaeT IICCJIe]:(OBaHlIe AIIcl>cl>epeH:qlIpoBKH HepBHO-
ro 3aqaTRa C HepBHoll Tpy6KII II CIIIIHaJIbHblX raHrJIlIeB II 3aKaH'IlI-
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BHIIMaHlIe Ha MHTOTlIqeCRYIO aRTIIBHOCTb. CpeAII Bcex ero Ha6JIIO]:(e-
HlIll 8TO TIOCJIeAHee IIMeeT oc060 cym;ecTBeHHoe 3HaqeHlIe. M3Y'IeHIIB
MlIrplIpYIOI:qIIX 1fJ0PM RJIeTOK 1I03BOJIHeT eMY IIpaBIIJIbHO xapaRTepn-
30BaTb C Mopcl>OJIOrlIqeCKOll TOqKH 3peHIIH KJIeTOqHbie rpYIIIIbl, KOTO-
pble B IIpo:qecce lIepeCeJIeHIIJI 06pa3YlOT CKOIIJIeHHH OKOJIO HaAIIO-
"IeqHIIKOB: 3TO KJIeTRII C Y3KIIM 060AROM :qIITOIIJIa3MbI II rlIIIepxpoM-
HblM HAPOM. MMeHHo Te yqacTKII, rAe cocpeAOTOqeHbI rpYIIlIbI TaKIIX
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1
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Henmnx p;oiPiPepeH:qlIpYIOm;ox MHT030B ]:(aeT p;Be BeTBII pa3BHTIIH-
CHMIIaT06JIaCTHqeCRyIO H iPeoxpoM06JIaCTlIqeCRYIO. BamHblM llIarOM
BlIepe]:( B pa60Te 3Toro aBTopa HBJIHeTCH TO 06CTOHTeJIbCTBO, qTO OH
CBH3aJI 06pa30BaHHe xpor.miPiPHH06JIacTa (iPeoxpoM6JIaCTa) He C Ton
I\JIeTROn, 1\0TOpaJl yme Bbl6paJIa IIYTb CBoero pa3BHTIIJI 1\ Henpo-
6JIacTY, a C IIpe]:(llIeCTBYIOm;en CTa]:(Hen, em;e He]:(uiPiPepen:qnpoBaH-
Hoii B l\aI\OM-JIII60 H3 HarrpaBJIenHII pa3BHTHH. Hett lie rrpo]:(aeT
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XOTH H ROHcTaTHpyeT ee HMeHHO B 3TH X I\JIeTl\ax. Ho Mbi 3TOT
iPaKT paCD;eHHBaeM Ral\ Han60JIee cym;ecTBeHHbln. Har.l p;yr.meTCH,
qTO TaRoe cymp;eHHe aBTopa ropa3p;0 6JIHme IIOP;BOP;HT R rrpaBHJIb-
HOMY IIpep;CTaBJIeIlHIO 06 HCTOqlllIRaX pa3BHTHH xpoMaiPiPHHHblx KJIe-
TOn, qeM IIpep;llIeCTBYIOm;He co06pameHHJI 06 3M6plIOHaJIbHOn CHMIIa-
Tn'leCIWU ImeTI\e I\al\ rrpep;TeqH xpoMaiPiPHH06JIaCTa. <1JaKTbI 06 3M-
6pHOHaJIbHOM pa3BIITHH HenpoHa CBII]:(eTeJIbCTBYIOT, qTO Heupoll TepJl-
eT crroc06HoCTb R MHTOTH'IecRoMy p;eJIeHHIO B paHHlIe CpOKH CBoero
06pa30Ba1lHH. 113 OIIHCaHHJI Hett. TPYP;1I0 CKa3aTb, p;o RaKHX CpOROB
3M6pHOHaJIbHOrO pa3BIITHH 3apop;blllIa OH Ha6JIIOp;aJI MHT03b1. Ho ]:(ame
OP;HO TO, qTO OH IIOp;MeTHJI HaH60JIbmee 1\0JIHqeCTBO MHT030B B
MeCTax CROrrJIeHHH MeJIRIIX nmepxpOMHblX I\JIeTOK, JlB.'IHeTCH 6mlb-
llIn\l IIJIIOCOM ero pa60TbI.
B rrOCJIep;YIOm;oe rop;bI 60JIbmHHCTBO HCCJIep;OBaTeJIen rnCToreHe3a
M03rOBom CJIOH Hap;II01leqHHKOB y 3apOP;blmen qeJIOBeKa (MBa-
HOB, 1930; Keene, Hewer, 1927; Wrete, 1927; rOJIy6, 1936; Coupland,
1952; Growder, 1957; !to, 1959; Boyd, 1934-1935; 1960) H y p;py-
rnx mHBOTHblX: y 6eJIbiX KpblC (Pancratz, 1931), y Mblmen (Goor-
maghtigh, 1935; Waring, 1936), Y 6b1KOB H KYP (fony6, 1936), y JIo-
IDll]:(en (Meiyling, 1938), y RpOJIHROB (Coupland, 1956, 1965), y
30JIOTbiX XOMHKOB (!to, 1957) 110 cym;eCTBY IIOBTOpJlJIH TO, 'ITO yme
6b1JIO CRa3aHO 0 III!.0Hcxomp;eHHH xpoMaiPiPHH06JIaCTOB IIHOHepaMH B
H3yqeHHH lIaparaHrJIHOHapHon CHCTeMbi IIJIH OCTaBJIHJIH 3TOT BOIIpOC
OTRpblTbiM. 3aTpYP;HeHHH, CTOHm;He Ha IIYTH 1\ pa3pemeHHIO 3Tom BO-
IIpoca, OqeBIIp;HbI. He TOJIbl\O B pa60Tax, B 1\0TOpblX CTaBHJII1Cb H 06-
cymp;aJIHCb ]:(pyrHe BOIIpOCbI rHCToreHe3a H p;HiPiPepeHD;HpoBRH IIa-
paraHr.'meB H B ROTOpblX OCTOqllHK pa3BHTIIJI xpoMaiPiPHH06JIacTa
paCCMaTplIBaJICJI 1I0IIYTHO, HO p;ame crre:qHaJIbHble HCCJIep;OBaHIIH He
Bcerp;a rrpIIBOp;HJIH aBTopOB K orrpep;eJIeHHoMy CY>Kp;eIlHIO. OCHOBHaH
IIpHqlIHa 3Tom - B 60JIbmOM MopiPOJIOrnqeClmM Cxop;CTBe Bcex 3M6pHO-
HaJIbHblX I\JIeTOqHblX 3JIeMeHTOB. IIIar BIIepep; B 3TOM OTHomeHHH cp;e-
JIaJI Hett, KOTopbln HapH]:(y C D;HTOJIOrlIen IIepeXO]:(HbiX lWeTOqHblX
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eJIeHHe B yqaCTKaX, IIpH6JIHmeHHblX K 3aqaTKaM 6plOmHblX IIapa-
raHrJIlIeB, II IIX iPYHKD;HOHaJIbHble oc06eHHOCTH - 06IIJIHe MHTOTII'le-
CRHX iPHryp.
RpOMe 3apop;blllIen KYP, Mblmen H I\pblC, D;HTOJIOrHH xpoMaiPiPlIH-
HbiX rraparaHrJIHeB 6pIOmHon 06JIacTH H lWeTOqHble HCTOqHHKH HX
npOHCXOmp;eHIIJI H3yqeHbI HaMH y aMiPH6IIn.
235
B JIa60paTOpHblX YCJIOBHHX 6b1JIH BbiBeAeHbI JIH'lHHKH H rOJIOB8-
CTHKII ABYX BHAOB JIHrymeK - Rana arvalis H Rana temporaria.
B IIepHOA sapOAbimeBoro pa3BHTHH OAHOBOSpaCTHbie IIOnyJIHI.\IIH HK-
plIHOK paCKJIaAblBaJIHCb 110 paaHblM aKBapHYMaM. aJIbHeiimHD: xOA
aapOAbillieBoro pa3BHTIDI CHCTeMaTlPIeCKH IIpOBepHJICH II 110 Mepa
BaA06BOCTH BHOBb COpTHpOBaJICJI. TaKHM 06pa30M BblBOAHJIHCb JIH-
IJRHKH H rOJIOBaCTHKH, naXOAHII:(HeCH Ha OARHaKOBbiX CTaADHX pa3-
BHTHH. IToCJIeAOBaTeJIbHble CT8.AHH JIHqHHOqHOrO pa3BHTHH OIIpeAe-
JIJlJIB: 110 BHenrneD: clJoPMe TYJIOBH a, CTeIIeHH paSBHTHH Hapym-
BbiX ma6p H ROHeqHOCTeD:.
rOJIOBaCTHKOB clJHKcHpOBaJIH eJIHKOM IIJIII IIOCJIe AeKaIIHTa HH II
qaCTH'lHoro y]:(aJIeHHH BHYTpeHHOCTeii (RHmeqHHKa, meJIYARa, IIe'le-
HH) Ha CJIeAYIO HX CTaAHHX: 23, 24, 25, 26, 27, 28, 29, 31, cera-
JIeTKa (xapaKTepHcTHKa OTAeJIbHblX CTaAHD: B3J1Ta H3 Ta6JIH bI Rugh.
1962). Pa3BHTHe IIaparaHrJIHOHapHblX RJIeTOK H3yqeHo Ha clJpoHTaJIb-
HbiX cepHiiHbIX cpe3ax B 06JIacTH 6plOmHOD: aopTbI, rAe Ha Me]:(H-
aJII>BOM Rpae HHTeppeHaJIOBbiX S8.RJIaAOK paCIIOJIOmeHbI HaH60JIoo
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TOqHbIe TJlmH CHMlIaTHqeCKHX CTBOJIOB (pHC. 81, A), HS MaTeplIa-
JIa ROTOpblX pa3BHBalOTCH OCTpOBKH IIaparaurJIHOHapHblX KJIeTOR Ba
IIepeAHeD: IIOBepXHOCTH aopTbi H ee BeTBHX (art. coeliaca), H3 Me-
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HeD: nOJIOD: BeHbI. OTAeJIbHble IIaparaHrJIHOHapHble yseJIRH IIpOHHRa-
lOT B IIOqeqHYIO IIapeHXHMY, rAe MoryT IIpOXOAHTb BAOJIb 1I0qeqHbiX
KaHaJIb eB.
.n;HlfJlfJepeH HpOBRa KJIeTOqHblX 3JIeMeHTOB B CHMIIaTlIqeCRHX S8-
IJaTKaX Y aMIfJH6HD: IIsyqeHa 110 TeM me MoplfJOJIOrHqeCKl'IM H rHCTO:-
XHMHqeCKHM IIp03HaKaM, ROTopble 6b1JIII HCnOJIb30BaHbI Ha ]:(pyrllx
mIlBOTHbiX.
RJIeTOqHaH ]:(IIIfJcl>epeH HpOBKa aaqaTKOB qyBcTBIITeJIbHblX Y3JIOB
Ha ABe IIOIIYJIJI HH II MlIrpa HJI HX B CHMIIaTHqeCRHe raHrJIHII 110-
rpaHlIqHoro CTBOJIa Ha6JIIOAaeTCH Ha 23-25-D: CTaWIJIX pa3BHTHJI.
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IIpOXOAHT Mem]:(y MarHCTpaJIbBblMII COCYAaMH II MYCRyn:aTYPOH XBO-
CTOBOD: qaCTH TYJIOBH a 110 HaIIpaBJIeHHIO R MeAHaJIbHblM KpaHM
.nOqeR, rAe BIIepBble YAaeTCJI Ha6JIIOAaTb IIOJlOmHTCJIbHYIO peaKl(lIIO
B xpoMacl>cl>IIHHblX RJIeTRaX Ha 26-D: CTaAHH paSBHTHH. CMemaHHwn
RJIeTO'lHbIH COCTaB THmeH, H;:ur HX BAOJIb CTeHOR CIIHHHOH aopThl.
aaTpYAHHeT BhlJlCHeHHe KJIeTOqHblX HCTOqHHKOB xpoMacl>cl>HHHhlX 3JIe-
Me.H'lOB Y rOJIOBaCTHKOB, TaK KaK B MeCTax COIIpHROCHOBeHHH aopTbI
C IIOqKaMH H HaA HHMH AHcl>cl>epeH HpYIOTCH BCe TPH KJIeTOqHble
IIoiIYJIJI IIII - rJIllaJIbH8.H, BeiipaJIbHaJI II xpoMaclJcl>HHHaH. 3TH MeCTa.
no aHaJIOrHH C MJIeROIIHTalO HMH, MomHO HaaBaTb IIapaaOpTaJIbHbl-
MH naparaBrJIIIJlMH. OAHaKO B OTAeJIbHblX yqacTKax, paCIIOJIOmeHHblx
n03aAH OCHOBHOro CROIIJIeHHH xpoMalfJcl>RHHOD: TKaHH, BCTpeqalOTCR
RJIeTOIJHble rpYIIIIbl, MlIrpHpYIOII:(He Ha CHMnaTHIJeCKllX Y3JIOB B CTeH-
RY CIIHHHOD: aopTbi (pHC, 81, A). Oc06eHHOCTH RJIeTOqHblX CTPYRTYp
236
B 3TIIX MeCTaX II03BOJIHIOT rOBopHTb 0 TOM, qTO KJIeTKH, ROTopLle
BHep;pHIOTCH B CTeHRY aopTbl II cnymaT HCTOqHHROM IIaparaHrJIHO-
HapHLlX, CXOP;HLI CO CIIOHrH06JIaCTII'IeCRHM 1'IIIIOM RJIeTOqHLlX reHe-
paquii Y p;pyrHX RJIaCCOB 1I0aBOHOqJIbiX (pHC. 81, B). B 3THX RJIeT-
Rax, IIpOHlIRaIOm;IIX H B ROpy Hap;IIOqeqHIIROB, BIIepBbIe IIOJlBJIJle:rCH
xpoMacIJlIJHHHaJl peaRqHH (pHC. 82).
C H M II a T H q e C Rile Y 3 JI bI H Rap 0 T H ]:( H bl ii II a par a H-
r JI H ii. ¥ 3apop;blmeii RpblC H Mblmeii RapoTIIp;Hblii naparaHrJIHii B
BIIp;e IIapHoro opraH a paCIIOJIOmeH B 06JIaCTH 6HclJypRa:I1;HH 06m;eii
COHHOii apTepHH, B ee HapymHoii 060JIOqRe (IIpHiiMaR, CMHTTeH,
1959). HaqaJIo paaBHTHJI RapOTHp;Horo IIaparaHrJIHH COBIIa]:(aeT C
B03HHRHOBeHHeM 6HclJypRaqHH (y RpblC Ha 14,5, y Mblmeii Ha 13-e
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XID!HOrO THIIa C RpYIIHblMH CBeTJIblMH oRpyrJILlMH Jlp;paMH H C 1-
2 Jlp;pblmRaMH. 3TH RJIeTOqHble rpYIIIIbI paCIIOJIaraIOTCH IIpellKym;ecT-
BeHHO Ha BeHTpOJIaTepaJIbHOii CTeHRe BHYTpeHHeii COHHoii apTepHH.
MopclJoJIorHqeCRH OHH He OTJIlPIaIOTCH OT RJIeTOR ap;BeHTlIquaJIbHOrO
CJIOJl CTeHRH RapOTHp;Hoii apTeplIB B, BBP;BMO, 06pa3YIOTCH B peaYJIb-
TaTe ee paapaCTaHHJI. MLI He 6epeMCH CYP;BTb 0 TOM, B03HBRaeT JIB
3TO 06pa30BaHBe B pe3YJIbTaTe YTOJIrn;eHBH C06CTBeHRO ap;BeHTBI1;B-
aJIbHOii OOOJIOqRII COHHOii apTepBH, RaR 3TO P;YMaJIB ern;e lIepBbIe
BCCJIep;OBaTeJIB pa3BBTBH RapOTHp;HOrO IIaparaHfJIBJI (Arnold, 1865;
H.am;eHRo, 1887; Marchand, 1891; Paltauf, 1892; Groschuff, 1896;
Jacoby, 1896; Verdun, 1898; Heinleth, 19(0) H RaR IIOJIaraIOT MHO-
rBe COBpeMeHHble aBTopbl (Hammar, 1934; Boyd, 1937a; Celestino
da Costa, 1935, 1939; Schwarz-Karsten, 1944), IIJIB JlBJIJleTCH IIpO-
B3BOP;HblM IIpBJIeraIOrn;eii it COcy]:(y Me3eHXBMbI (Rabl H., 1922; Smi-
th, 1924; Agduhr, Hammar, 1932; Pischinger, 1934, 1937). C YBe-
peHHOCTbIO MomHO CRaaaTb TOJIbRO OP;HO, qTO 3JIeMeHTbl, clJ0PMB-
PYIOm;He (<RapOTIIp;Hoe YTOJIrn;eHlIe» Ha 3TOii CTap;BB 3M6pBoreHe3a
RpblCbl B MbimlI, He JlBJIHIOTCH (<IIpIlIllJIblMB», a 06pasyIOTCH B pe-
3YJIbTaTe paaMHomeHBH RJIeTOR MeCTHoro xapaRTepa, Bl'paIOm;1IX pOJIb
CTPYRTYpHoro OCTOBa 3Toro opraHa. HBRaRIIX IIpB3HaROB BCeJIeHBH
B 3TO MeCTO HeiipaJIbHbIX 3JIeMeHTOB ern;e He BMeeTCH (pIIC. 83, A).
Ha CJIeP;YIOm;eM 3TalIe MopclJoreHe3a (15,5-16-e CYTRH Y 3apo-
]:(blllIeii RpblC, 14-e CYTRH Y 3apop;blmeii Mblmeii) B MeaeHXHMHOM
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p;eHBH (pHC. 83, B, B).
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242
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npD]:(yqnpYIOm;ne a]:(penanlIHa (Bo.3Mo.mHD, au;eTlWXo.nnH?). IlepBble
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OTpocmOB RaK eWIHCTBeHHLIX rryTeH MlIrpaqlIlI. OHII cym;ecTBYIOT
B 60Jlee n03]:(HlIe cpORn 8M6plIoreHe3a, Rorp;a MHorne 3aRJla]:(RH yme-
IIMeIOTCH. Ho eCJlH Y'IecTb, qTO MOPcPoreHe3 naparaHrJIHeB HaqlIHaeT-
CH II03me, qeM CHMnaTH1JeCRIIX Y3J10B, Rorp;a HepBHbIe BOJlORHa BO"
MHomx Y'IaCTRaX TYJlOBlIII{a yme pa3BIIJlIICb, TO 8TOT IIplI3HaR H6
MomeT 3p;eCb IIrpaTb pemalOIqeH pOJIII. TI03TOMY npII II3yqeHHH pa3-
BIITHH RapOTH]:(Horo naparaHrJIlm BamHblM HBJlHJlOCb BLIHCHHTb, MIII'-
PIIPYIOT Jill RJleTRII II3 Bcex Tex IICTO'lHIIROB, 113 ROTOpLIX npolIcxo-.
]:(IIT HHHepBaqlIH RapOTII]:(Horo naparaHrJlIIH, IIJlII MlIrpaqllH orpaHII-
'IIIBaeTCH RaRHM-JlH60 O]:(HIIM IICTO'IHHROM II RaRlIM HMeHHO - CIIM-
naTH'IeCKHM HJlII napaC.IIMnaTH1JeCRHM?
PellieHII6 BTOporo Bonpoca 0 xapaRTepe MeTOR, BCeJlHIOr:u;IIXCS
B RapOTII]:(HbIB IIaparaHrJlIIB, npeCJle]:(OBaJlO qeJlb YCTaHOBHTb 110 qlI-
'lOJlOrIPl8CRHM nplI3HaRaM, IIpHHa]:(JlemaT Jill OHII R TOB me IIOlIY-
JlHQIIH, ROTOpLIe 06pa3yIOT Ha]:(nOqeqHbIH naparaHrJIHH, IIJIH HBJlHIOT-
CH 0 T JI II q H 0 ii OT HIIX nonYJlHqlIeH RJleTOO\.
V RpblC B 6IIcPYPRaqIIII 06r:u;eH COHHOii apTepIIII Me3eHXHMHaH
3aRJla]:(Ka 3aRaHqlIBaeT CBoe cPOpMlIpOBaHlIe, RaR yme CRa3aHO.
y. 15,5-16-M cymaM pa3BIITHH, y MblmeH - R 14-M CYTRaM, nocJle-
qero HaqHHaeTCH BCeJl6HII6 HeHpaJlbHLIX 8J1eMeHTOB. Ha 8TIIX CTa]:(IIHX
K o6J1acTH 6IIcPYPRaqIIII 06r:u;eH COHHOH apTepIIII nJlOTHO nplIJIeralOT
BepXHIIH meHHLIH CIIMnaTlI1JeCRHH Y3eJl (ganglion cervicale supe-
rior) H 'IYBCTBIITeJlbHbIH Y3eJl 6JIym]:(alOIqero HepBa (ganglion no.-
dosum). B 3TOT nepllo]:( 06a Y3J1a pe3RO OTJIli'IaIOTCH no qIITOMOp-
cPOJlOJ'H1leCROH xapaKTeplICTIIRe RJIeTOqHblX 8J1eMeHTOB, II B TOM 'IIIC-
Jl6 no pa3MepaM HeHp06J1aCTOB (pile. 86). TaRaH 'IemaH pa3HlIqa..
HapHP;Y C xopomo OIIpe]:(eJiHeMbIMII nYTHMII MlIrpaqIIII (CM. pilC. 83)..
P D c. 85. CxeMa I, A, B, B, r, D - rpaclJn'leCRaH peRoHcTpyKl\IIH apTeplIii
neBoH ma6epHoH p;yrJl JI ux DpOU3BO):\HIdX B cepHli 3apo):\ldIIIeH 'lenOBeKa, DORa-
3IdBaIO aH B3aUMOOTHOmeHJle H3IdKo-rJIOTO'lHOrO HepBa JI CBMnaTB'leCRoro.
CTBOJIa J( OROH'laTeJIbHOH BHYTpeHHei'1 COHROH apTeplIu Ha nOCJIep;OBaTeJIbHIdX
CTap;HHX pa3BUTJlH;
II, A, B, B - rpalJlH'leCRaH peRl'lIcorpYFI1HH o(j.naCTH FsparnnHot1: 6BlJlYPR8I1ID1V Tpex aapo-.
\.\hImeA 'leJIOBeHa, nOHaal>lBalO1I\8H MeCTOnOJIomeHHe MeaOg e pM8JJbHOI'O }'WIOTBeHBH nepBB'!-
HOI'O aa'laTRa R8pom):\H01'0 TeJIa Hero lIepBIINX CBHaeA; Acc - O(i1I\8H COHHaH 8pTepBH;
Act - HapVH<HBH COHHaH 8pTepHH; Ad - BHYT eHHHH COHHaH 8pTepHH; Aph. as - BOCXo-
JlHII\8H I'.JIOTO'lHaH apTepHH; Ap - JIeI'O'lHaH apTepHH; ar. Ao - Jlyra aopTIII; ar. Ap - gyra
nerO'lHot1: apTepHH; Ar. A. - 3-H 8pTepH8JJbHaH gYl'a; ar. A. - {o-H 8pTepB8JIbB8H j.,}'I'&;
cb.- RapOTHgBoe TeJIO; Con - MeaogepM8JJbHOe YllJIOTHeHlfe; Gcs - BepxBBt1: meltm.It1: yaeJII
Gn - G. nodosum (inymA8lOlI\eI'O HepBa; Gp - G. petrosum HahiRO-I'JIOTO'lllOrO BepB8; NgI -
HahiRO-rnOTO'IIIhIt1: HepB; Nh - nog'bH3hI'lHhIt1: HepB; NIs - BepxHHt1: 1'0pT8BBhIit HepB; Ns-
cHMnaTH'lCCRHt1: HepB; Nv - 6JIYlliA8lOlI\Bt1: HepB; re. N l - COHHaH BeTBb H3hIRO-I'JIOTO'lll0I'o-
HepBa; rph. Nv - I'JIOTO'IHaH BeTBb (inymA8lOlI\eI'O HepBa (Boyd, 1937b)
245.
. ,.,.
: \.. :.\".. \.. . .' ) . 't) ..
" ,t, :;:,,,,,. ':", .
'. A ..,', .>, .: ., . .. '>,... . .
'1:.;.t:.. » l' ? '..
-':' , - ../
. ".
"'It'-;
,
P If c. 6. OTHOCIITeJlbHble pa3-
lIIephI RJIeTOH BepXHero meiiHoro
cJnmaTJlQecn:oro Y3JIa If 'lynCT-
IIJITeJIhHOrO Y3JIa 6JIymp;aIOll\ero
HcpBa 3apop;bllUa lIIblllllf (15 cy-
TOH pa3BJITJlH)
a - BepxHHt1: meAHhlt1: yaw; 6-
qYBcTBHTeJIbHbIfi yaeJI. OHp. MeTHJIO-
BbiM aCJIeHbIM-nHpOHHHOM. OH. X 7.
06. X 20 (lIpHI1MaH, CMHTTeH, 1959)
.,
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, ,';f, /:-"-",: ,,' i. ,-."
)1aeT B03MOffiHOCTb 6e:WIlIII60'IHO onpe;:J;emlTb pa3BlITlIe cue JI(IJlIqe-
CKUX KJIeTOK KapOTU Hora naparaHrJIUH KpblC II Mbllllen U3 BepXHero
ruenHoro CUMnamqeCIWro Y3J1a.
HalI60Jlblllee KOJlII'IeCTBO CIiMnaT06J1aCTOB paCUOJIaraeTCR B Kpa-
HlIaJlbHOn qaCTIi BepXHera menHoro Y3JIa, n TO BpeMH KaK B Kay-
J{aJIbHOn CKaUJlIIBaeTCR upelIMy ecTBeHHo CUOHrlI06J1aCTbi II, IIO-BII-
1TIIMOMY, 3]:(eCb me OCTaeTCR HeKOTopoe KOJIUqeCTBO MaJIO]:(lIcPcPepeH-
IIpOBaHHbIX ::meMeHTOB.
3TO paBJIlIqlIe B KJIeTOqHOM COCTaBe KpaHlIaJIbHOn II Kay]:(aJIbHOn
'IaCTen BepxHero llIenllOrO Y3J1a oc06eHHo OTqeTJlIIBO OTMeqaeTCR K
HaqaJIY MUrpa IIII (16-17-e CYTKII y KpblC, 14-e CYTKII Y Mblmen),
KOI']:(a II3 Kay]:(aJlbHOn qaCTU era B Me3eHxIIMHYlO 3aKJla]:(KY Kapo-
"'T1I HOrO uaparaHrJlIIR HaqUHalOT BblCeJlRTbCR KJIeTRII. B OCHOBHOM
OHII upe]:(cTaBJleHbl cuoHru06J1aCTaMII II HeKoTopblM KOJllIqeCTBOM CIIM-
-naT06J1aCTOB (eM. plIC. 83, B).
B ueplIo]:( BblCeJleHIIR H]:(pa cnoHru06J1aCTOB 60Jlee UHTeHCIIBHO
'OKpamIlBaIOTCR uo <DeJibreHY, qeM R pa IIOKORJIJ;IIXCR cUOHrIl06JIa-
TOB BepXHero menHora Y3J1a. no 3TOMY uplI3HaKy OHII Jlerno OT-
.JllIqalOTCR OT M63eHXIlMHbiX KJIeTOK, BMeCTe C KOTOpblMII BCTYnalOT
B 3aKJla]:(KY KapOTII]:(Horo uaparaHrJlHH.
Y 17-19-cYTO"lHbIX 3apo,w,lllIen KpblC II 16-17-cyroqHbIX 3apo-
.J];Llmen MLlmen KapOTH]:(Hblii: uaparaHrJIIIii: upe]:(CTaBJIReT C060H oKpyr-
.JIOe 06pa30BaHlIe, COCTOHJIJ;ee II3 HeCKOJlbKIIX CJlOeB KOH eHTpHqeCKII
paCUOJlOmeHHblX KJleTOK. B 3TO BpeMH B HeM pa3JIHqalOT (oco6eHHO
.xopomo Y KpblC) Be 30Hbl: HapymHaR, COCTORJIJ;aR II3 MlIrplIpylO-
JIJ;IIX 3JH'MeHTOB HeiipaJlbHOra npoRcxom]:(eHHR, H BHYTpeHHRR, co-
roRJIJ;aH II3 KJleroK COeAHHIITeJIbHOTKaHHoro TlIua II KpoBeHOCHblX
-COCY OB. HaJIlIqlIe ]:(BYX 30H B pa3BIIBalO eMCH KapoTII]:(HOM TeJIe
uepBble 6b1J1o OUHcaHO Kohn y 3apO]:(blmeH CBIIHbII II BnOCJle]:(CTBHH
nO]:(TBepm]:(eHO Watzka (1938, 1943), Schwarz-Karsten (1944) II ]:(p.
n03]:(Hee, Y 19-20-cYTOqHbIX 3apo]:(blllleii: KpblC MIITplIpOBaBilllIe
lIoHrn06J1aCTbI rpyuuaMII upO]:(BHraIOTeR B eHTpaJlbHble yqaCTKII
"KapOTlI]:(Horo uaparaHrJlIIH, cPOpMIIPYH BOKpyr eHTpaJlbHOii apTepIIII
.J];OJlbKH, OT]:(eJleHHble ]:(pyr OT pyra UpOCJlOHKaMII coe]:(IIHIITeJlbHOH
TKaHII. qacTb cnoHrn06J1aCTOB upeBpaJIJ;aeTCR 3]:(eCb B cue IIcPlIqe-
.246
CRlle RJIeTOqHble 3JIeMeHTbI RapOTH]:(Horo naparaHrJIIUl II nplI06pe-
TaeT CTpYRTYPY, XapaRTepHYIO ]:(JIH naparaHrJIIIOHapHblX 3JIeMeHTOB,.
OnRCaHHYIO Bblllle (CM. rJIaBY VI II plIC. 75).
qTO RacaeTCH MlIrplIpOBaBm:nx neup06JIaCTOB, TO, Y'IIITbiBaH IIx..
nepml>eplI'IeCROe pacnOJIOmeHlIe II npOABllmeHlIe B]:(OJIb CTeHOIK Rapo-
TIIAHblX apTeplIU, CJIe]:(yeT CqlITaTb, q1'O OHII He 06pa3YIOT 3JIeMeH1'OB
RapoTIIAHOro naparaHrJIIIH, a npJlHIIMalOT B ]:(aJIbHeUmeM yqaCTlIe
B WOpMHpOBaHIIH BHYTpJlCTeHOqHblX HepBHblX Y3JIOB COCY]:(OB.
CpaBHeHne rlICTOreHe3a RapOTH]:(Horo naparaHrmUi Y RpblC II Mbl-
meu ]:(aeT npaBO rOBoplITb 0 npRHQHlllIaJIbHOM cXo]:(CTBe 3TOI'O npo-
u;ecca Y Tex II APyrlIx H B TO me BpeMJI OTMeTHTb HeROTopble BIt-
AOBble pa3JIHq1lH. TaR, Y RpblC pa3B11THe RapoTII]:(Horo TeJIa Ha'IlI-
HaeTCH n03me II JI]:(eT 60JIee Me]:(JIeHHo. BJIaro]:(apH pacTHHYTOCTII
npoQeccoB fiJIeTOqHblX ]:(IIwwepeHQlIpoBOR B 3M6plIOHaJIbHOM pa3BII-
TIIH RpblC Y HIIX OT'leTJllIBee MomHO npOCJIeAIITb oT]:(eJIbHble 3Tanbl
pa3BIITIIH RJIeTOR npII B03HIIRHOBeHHII O]:(HlIX WOPM H3 ]:(PyrlIx_
M3Y'IeHlIe HHHepBaQIIH RapoTIIAHOro naparaHnIIlH (ilplIUMaR r
CMIITTeH, 1959; IIpHuMaR, 1962) nORa3bIBaeT, qTO B Hero BXOl.\HT.
BOJIOIma OT BepXHero meUHoro CIIMnaTlflOC,Roro Y3JIa, OT qYBCTBII-
TeJIbHoro Y3JIa 6JIym]:(aIOm;ero HepBa II, IIO-BII]:(IIMOMY, OT BeTBII H3bl-
IW-rJIOTO'IHOro HepBa. ORaHqlIBalOTCH OHII Ha RJIeTKaX RapOTHAHOrO\
TeJIa B BIIAe neTeJIeR HJlII nyroBOH. B 3TOM HalllII ]:(aHHble nOJI-.
HOCTblO nO]:(TBepm]:(aIOT BblCRa3aHHoe B pH]:(e pa60T (Boyd, 1934-
1935, .1937; nOHOMapeBa, 1946; Watzka, 1938; Schwarz-Karsten,.
1944, II ]:(p.).
O]:(HaRo HeJIb3H COrJIaCHTbCH C YTBepmJ{eHlIeM HeROTopblX aBTO-.
pOB, qTO IIHHepBaQIIH onpeAeJIHeT RJIleTOqHblU COCTaB RapoTII]:(Horo
naparaHrJIIIH, T. e. MlIrpan;IIH RJIeTOR OC eCTBJIHe'f1CH BAOJIb BOJIO-
ROH, II]:(YIIJ,HX II3 Bcex Tpex nCTOqHIIROB HHHepBaI{IIII. no HaIllIIM.
Ha6JIIO]:(eHHHM, MlIrpan;IIH RJIeTOR B RapOT Hoe TeJIO MJIeROIIIITalO-.
m;IIX npOHCXO]:(IIT TOJIbRO II3 Be.pXHero llIeUHOrO CHMnaTIIqecRoro Y3-
JIa; RJIeTRII me II3 qYBCTBIITeJIbHoro Y3JIa 6JIym]:(alOm;ero HepBa He'
BcrynalOT B KapOTII]:(Hoe TeJIO, a npOXO]:(HT MHMO. 81'0 3aRJIlOqeHII8'
BblTeRaeT II3 II3JIOmeHHbiX Ha6JIIO]:(eHHu Ha]:( XO]:(OM II nYTHMII MII-
rpan;1II1, a TaRme cpaBHIITeJIbHbIM II3Y'IeHlIeM n;IITOJIOrlIH 060HX Y3-
JIOB H pa3BIIBalOm;erOCH RapOTH]:(Horo naparaHrJIIIH.
MCTO'IHIIRaMII B03HIIRHOBeHlIH cnen;IIwlIqeCRIIX 3JIeMeHTOB Rapo-
TII]:(HOro .naparaHrJIHH HBJIHIOTCH He Heup06JIaCTbl, RaR YTBepm]:(aJI
Schwarz-Karsten, a CnOHrHOOJIaCTbI, RaK nORa3aJIa B. H. IIoHoMa-
peBa (1946) B RYJIbrype TI\aHII.
Y IITHQ, 110 cym;eCTBY, HeT 06JIaCTII, I'OMOJlOrII'IHOU 611wypRaQIIII"
06m;eu COHHOU apTeplIlI. OT art. subclavia OTXO]:(IIT apTeplIaJIbHblH.
«RapoTII]:(HbIH CTBOJI), KOTOpblH pa3BeTBJIHeTCH Ha art. cervicalis an-
terior, art. vertebralis II art. carotis communis (CM. pHC. 54)..
8Ta 06JIaCTb pacnOJIOmeHa 3Ha'IIITeJIbIliO HHme, qeM c06cTBeHHo 6H-
wypRan;IIH, a IIMeHHO Ha rpaHIIn;e Mem]:(y llIeeu II rPYAHou nOJIOCTblO.
Oc06eHHocTblO nTIIn;, RpOMe Toro, HBJIHeTCH TOIIorpawuqecRaJl:
6JIH30CTb meuHo-rPY]:(HbIX naparaHrJIHeB C napaCHMnaTIIqeCRHM OT-
241
eJlOM HepBHOH elICTeMbl, a IIMeHHO - C qYBCTBIITeJIbHLIM Y3JIOM
6nymAalOI.qerO HepBa. B OTJllfTlHe OT MJleIWnlITalOIqIIX, r]:(e BepXHlIii
meHHblH CIIMnaTIIQeCI\IIll Y3eJI II ganglion nodosum paCUOJlOmeHLI
B 6JIII31\OM eoce TBe ]:(pyr C APyroM II C l\apOTlI]:(HLlM naparaH-
rJlUeM, y nTUQ OHU pa306IqeHLI. BepXHlIll mellHLlll CIIMnaTIPIBCimll
Y3eJI pa3BIIBaeTCH B BepXHell 'IaCTII mellHoii o6JIacTII, a ganglion
nodosum, Hao60poT, HaXO]:(IITCH BHli3Y, Ha rpaHlII B C rpyP;HOll no-
.JIOCTblO, B6JIII3II mellHO-l'pYAHbiX naparaHrJIlIeB.
3TH aHaTOMlIqeCI\IIe oco6eHHOCTII npe]:(CTaBJIHIOT p;OnOJIHIITeJIbHbIe
E03MomHOCTII, I\al\ yme Cl\aaaHO, ]:(JIH paCCMOTpeHIIH IICTO'IHIII\OB npo-
Ifcxom]:(eHliH naparaHrJlIiOHapHLlX I\JIen)l\, nYTell IIX MlIrpaQlI1I (II3
HMnaTII'IeCRIiX IIJIII napaclIMnaTlIqeCI\IIX 3J1eMeHTOB HepBHOll CIICTe-
Mbr) II, 'CJIe]:(OBaTeJIbHO, BblHCHeHlI1I HaJIlIqlIH IIJIII OTCYTCTBIIH l\aRIIX-
'TO 06IqlIX 3al\OHOMepHOCTell ]:(JIH nTlIq II MJIeI\OnIITaIOIqIIX.
YI\a3aHHbIe BonpoCbl, no HameMY npe]:(JIOmeHIIIO, 1I3Y'IeHbI
::i. X. llplIllMaK (1960, 1962) Ha MaTeplIaJIe aapo]:(LImeH RYP nop0]:(LI
c6eJILlH Jlel'ropH» B pa3Hble CpOKH IIHI\y6aqIIII - OT 1 CyTOR ]:(0
BLlJIynJIeHIUI II BblJIynliBmlieCH 1,5, 15, 30-CYTo'lHble qLmJIHTa. llplI-
BOP;IIM ee O'IIRCaHIIH.
Pa3BliTHe llIellHo-rpY]:(HbIX naparallrJIlieB HaqIIHaeT'CH Ha
'6-e CYTI\II 3M6plIoreH63a 3apOp;Llllia qLlnJIeHl\a. Ha]:( pa3BIIJIKOll rpyp;-
:HOH aopTLI II nO]:(RJIIO'IlIqHOll apTepHU y CTeHI\II 06IqeH COHHOll ap-
'TepnH 06pa3yeTcH Me3eHXIIMHaH aaKJlap;Ka, COCT()Hm;aH II3 BepeTeHO-
o6paaHbix KJIeTOI\, pacceHHHLlX cpe]:(II 06HJIbHOro ROJIJf'IeCTBa IIeTeJlb
KanlIJIJIHpHOll ceTII. RapoTII]:(Hblll naparaHrJIlIll pacnOJIaraeTCH He-
CKOJIhGO Bblllle (l\paHlIaJIbHee), 'IeM I\pynHLlll, OI\OJlOBaraJlbHLIll; 06a
oOHII JIemaT Memp;y IqIITOBII]:(HOll II napaIqIIToBII]:(HOll meJIeaaMII, PH-
.p;OM e ganglion nodosum (pIIC. 84, A). B nplIJIemaIqeM 1\ MeaeH-
XIIMHOH aaRJra]:(l\C 'IYBCTBIiTeJlbHOM yaJIe 1\ 3TOMy- BpeMBHII yme OT-
"IeTJlHBO HaMe1JaeTCH MOPWOJIOrli1JeCl\aH ]:(IIwwepeHQlIpoBRa RJIeTOq-
Horo MaTeplIaJIa: I\JIeTl\1I HaXO]:(HTCH Ha CTa]:(IIII npoHeHp06J1aCTOB,
:x.morue nplI06peTalOT IIpli3Hal\II HellpOOJIaCTOB; 'IaCTh cnOHrlI06JIa-
CTOB pacnOJIaraeTCH yme B6JIIIaII HepBHblX 3J1eMeHTOB, ol\pymaH IIX
"Hano]:(06II6 CaTeJIJIIITOB, p;pyrlIe JlemaT cB060]:(HO II 06paaYlOT He60JIb-
mile rpynnbI.
Ha 7-11-e CYTl\II 3M6plIoreHe3a 3apO]:(Llma QLlnJIeHl\a cl\onJle-
HIiJl cnOHrli06J1acTlPlool\IIx KJIeTOqHLlX Mace B conpOBom]:(eHIIII OT-
JJ;eJlbHLIX HeHp06JIaCTOB (CM. plIC. 84, B) BLlCeJIHIOTCH; II3 'IYBCTBII-
TeJIbHOrO Y3JIa 6nym]:(alOIqero HepBa II IIepe]:(BlIralOTCH no HaIIpaB-
JleHlI10 R Me3eHXliMHOll 3al\JIap;l\e l\apOTII]:(Horo naparaHrJIIIJI. qacTb
II3 flHX pacnOJIaraeTCH no ee nepliweplIlI (CM. plIC. 84, B), Ha-
D.0p;06II6 roro, 'ITO Ha6JIIO]:(aeTCH y MJIel\OnlITalOIqlIX (llpUllMal\, CMIIT-
"'IeH, 1959) II WopMlipyeT QeHTpaJlbHYIO qaCTb opraHa. ApyraH qaCTb
ttmrplIpYlOlI{HX I\JIerotJHblX 3J1eMeHTOB npo]:(BlIraeTcH ]:(aJIee B I\ay-
AaJIbHOM HanpaBJIeHUII II B6JIIIaII 3a1\JIa]:(I\II napaIqIIToBII]:(HOll meJle3L1
o6pa3yeT I\pynHbllllllellHo-rpY]:(Holl naparaHrJIlIll (pIIC. 87).
Ha lO-11-e CYTI\II II n03me cpe]:(II MlIrpnpYIOIqIIX cnoHrn06JIa-
CroB (MeJlI\IIX KJIeTOI\ C He60JIbllllIM 060P;I\OM QIITOnJla3MLI, I\OMnaI\T-
248
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Pile. 87. O'l'peaOR nepBHoro CTlIOJIHlm (a) 6JlYllip;aIOn ero HepBa, HP;Yll\erO K
ruei-ino-rpyp;noMY naparanrJlHIO (6) aapop;hlIlla KypHl\1d (9-e CYTKH HHKy6aWIlI).
OHp. MeTHJIOBbIM 3eJICHbIM-llHPOHHHOM, OH. X 7. 06. X 20 (npenapaT IIpHI1Mall)
P II c. 88. Xpmm.pcpnHOOJlaCTLI n CTHO.TIC OJlyit\p;aIoll\ero nepna 3apop;hIwa RyplIl\bl
(12 CyTOK IlHRy6al lIlI)
Ollp. MemJIOBbIM 3CJIrIIbII\I-ITIIPOHlIlIOM. OR. x 7. 06. x 20 (lIpniiI\IaH, 19(0)
(..... ........- ...-,;, -'.'.. .
....... ,,'t, .t.):;.. . ',"
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.."'., '''''''.'' ... 'IiiIL.'!!!IL,;. ...
"'. "- '';h.-", . . ,';" .';. ". .. ....... .......-
-::. ;.: «'.,.' .. ,. ........' __ .AlP
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: _ :; . <t .>. .,.
.; !:* \'-':,f ;. t ;; " ::. ::.;. ..k. «
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HblM JI]:(pOM II B aTJIIPI1I6 aT CDaHrlIa6JIaCTaB MJIelRaDIITalODJ;IIX ca-
epmaDJ;HX TaJIbRO 1-3 H]:(pblmRa) II eAHHH'IHbIX aeHp06JIaCTOB
(RpYDHbIe RJIeTRH C 06mHpHon DHpaHIIHaWIIJIbHaH 3aHaH qIITODJIa3-
.MbI, RpYDHblM CBeTJILIM H]:(paM II 1-2 H]:(pbllliRaMII) DaHBJIHeTCH
TpeTHH TIID RJIeTOR. OHII IIMelOT IIY3blpbRaBII]:(Hae, 6e]:(Hae xpaMa-
TIIHaM HAPa (MeHbllllIx paaMepOB, qeM H]:(pa HeHp06JIacTaB), C 1-
2 H]:(pblllIRaMII II ]:(aBaJIbHa mlIpORHM a6aJJ;RaM qHTaDJIa3Mbl. nO' Be-
.JIlIqlIHe a6a]:(Ra (H BeJIH'IIIHe RJIeTKH B qeJIaM) aHa 3aHIIMaeT cpe]:(-
Ree naJIameHlIe Mem]:(y rJIlIaJlbHblMH II HeHpaJlbHblMII ::meMeHTaMH
Ii pe3Ra aTJlH'IaeTCH aT HIIX aqeHb CJla6aH IIlIpaHIIHaWIIJllIeH. HH B
'yRaaaHHbl6 cpmm, HII B ]:(aJlbHeHllIeM 8TH RJleTRII He a6HapymlIBa-
.lOT B CBoeH qHTaDJla3Me HeiipaWII6plIJIJIJIpHaH ceTII II He IIMelOT
OTpaCTRaB. nO' qlIToJlarnqeCRHM DpH3HaRaM am RJleTKII cxa]:(HLI C
xpaMa4>4>IIHa6J1aCTaMII, pa3BIIBalODJ;IIMIICJI B ]:(Pyrux DaparaHrJlIIHX
(CM. pHC. 75) MJleRaDHTalOlI{HX II DTHq - KapaTII]:(HbIX II 6plOmaa-
aapTaJlbHblX (CMIITTeH, 1958, 1962; CMIITTeH, BJla]:(OC-npIIHMaR,
1959; nplIHMaR, CMIITTeH, 1959; CMIITTeH, MaJIHHaBCRaH, 1960).
B TeqeHII6 12-15 CYTaR aM6plIaHaJlbHoro pa3BIITIIH RypHHblX
sapo]:(LlmeH B DaparaRrJlIDIX II Ha IIym MlIrpaqlIH R HHiM DpalIcXa-
.JJ;IIT MaCCOBae Dpea6pa3aBaHIIe RJleTOR, Da3BaJIHIODJ;ee DpOCJle)J;HTb ]:(II-
HaMIIKY Dpoqecca pa3BIITHH xpoMaWWIIRa6JIaCTaB. Pop;aHaqaJIbHOH
KJleTKaH ]:(JlH RHX HBJlHeTCH aJleMeHT reMHOORpameHRblM H]:(paM
-cDaHrUa6J1aCTH'IeCRaro THDa. B DepeXa]:(HblX wapMax nO' DYTH MHrpa-
qIIII K naparaHrJIIIHM YBeJlH'IlffiaeTCH 3aHa qlITanJlaaMbl DpII a TCYT.
CTBIIII cDeqlIWHqeCKIIX ]:(JlH HepBRaii KJleTRII opraHalI]:(aB. Bce aHII
paCDaJIaralOTCH rpYDnaMII B DeTJIHX RaDIIJIJlHpaB clIHycaH]:(Hara TilDa
II aTJIH'IaIOTCH ]:(Pyr aT ]:(pyra pa3JIH'lIWH CTeneHblO 3peJiaCTII.
TaRUM o6pa3aM, lJalIYJlHqHH RJIeTaK C rlIDepxpaMHblM H]:(paM, Bbl-
.ceJIHHCb II3 ganglian nadasum, p;alOT HaqaJla cDeqHaJIbHbIM RJIeTRaM
DaparaHrJIUeB - xpaMaWWIIHa6J1aCTaM. KpaMe roro, rpYHllbl TaRHX
me RJIeTaR BCTpeqalOTCH B CTHOJIe 6JIym]:(alOIqera HepBa, RpaHllaJlbHee
II Ray]:(aJlbHee ganglian nadosum (pIlC. 88).
qYBcTBIITeJlbHblH Y3eJI 6J1ym]:(alODJ;ero HepBa He HBJIHeTCH, a]:(Ha-
Ra, y RyplIHblX 3apap;blmeH ep;IIHCTBeHHbIM IICTaqHIIRaM a6paaaBaHIIJI
"lmeTaqHblX 8J1eMeHTaB llIeHHa-rpY]:(HbIX DaparaHrJIlIeB. ORaJia llIen-
HblX llaparaHrJIHeB IIMelOTCH MeJIRlle CIIMDaTlIqeCRlIe Y3eJlRlI, paCDa-
JIameHHble lIa xap;y HepBa, Ha3BaHHoro Terni (1927, 1931) DpeBep-
Te6paJIbHblM, DpeKapaTII]:(HLIM CTBaJIHRaM. nO' MHeHlI10 Terni, DpeBep-
Te6paJlbHbIH CIIMDaTH'IeCRIIH HepBHblH CTBaJI HBJIHeTCH aca6eHHOCTblO
CTpaeHIIH CIIMDaTIPIOCRaH HepBHaH CIICTeMLI DTlIq. nO' RJleTO..'IHaMY
caCTaBY II CTeDeHII 3peJIOCTH RJIeTaR B DepHa]:( 8M6pllal'eHe3a 8TII
CIIMDaTH'IeCRlIe yaeJIRII aTJIHqalOTCH aT y3J1aB DarpaHlPIHaro CTBaJla.
,l];IIwwepeHqlIpaBRa B HIIX DO' BpeMeHII aTCTaeT aT ]:(IIwwepeHqllpOB-
KII RJleTaR B IIal'paHlIqHblX raHrJIHJIX. B IIX caCTaBe BCTpeqalOTCH
Bee TPII RJleTOqHLl6 IIaDYJIHqllH - HenpaJIbHaH, rJllIaJlbHaH II xpa-
MaWWIIHHaH, RaTapLl6 MlIrplIpYIOT B RapaTHAHLIH DaparaHrJIlIH. Kpa-
1tf6 TOm, B wapMHpOBaHIIH DapeHXIIMLI RapOTHAHaro DaparaHrJIHH
'KYPIIHbIX 3apa]:(blmeH DplIHIIMalOT yqaCTHe RJleTRII, MlIrplIpYIODJ;H6 B
250
lIlCiillo-rpYl IILJe 1I'lIwrallrJIIIII co l'TOpOIlJ.!. IIpOTUIIOIIOJlOllmoii 'IYR-
CTIIUTl'.'lhnOMY y:I."IY G"IY"'I aIOIl\pro 1Il'l'lIa II COCTalll' 1Il'l'lIl!hlX Hl'TO-
'll'I.., n;tyu ux OT lIepXIIl'1'O lIIl'iillOro CIiMllaTIl'leCIWro Y:\J1iI (pile. 89,
A, /J).
110 l'I'OI,aM lIIrl'allllU II 001,l'MY IIcpcce.'UTIOIU;II'\CH HJIeTOI.., Ita;tO
;(y taTh, '!TO CIlMnaTlI'lCCl\lIl' Y:IJII,I Y IITIIU; IIrpaKJT IIC:JHa'IIITe.'II,IIYIO
pO.ll. n 1I0CTpOCIIIIII napl'IIXII:\It.I lIIl'iillo-rpYAIILIX naparanr:IIIl'II. O;t-
HaW) TaIWrO po;ta <<I>OJIII'IPCTIIPlllliUlI) CTOpOII.1 II ;tallHOl\l CJIY'WC He
JlrpaeT cy ecTRPlllloii PO:III. l'opa3..'1.0 OOJlLIlICe 3lHI'IeIllte - nplllI-
1 lIIl11a.l"lIoro xa 1'.1Iaepa - IIMPl'T (IJaI>T, 'ITO Y nT1l1l IH'TO'lIIlIIWl\I Iwe-
6
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lIIl'iiIlOj"1I Y;J.T\a H lIIl'iillll'I.Py;J.III1 1 Y 1101 pOI rallr.IJIlO 3a I'0;U,IIIHI I';YIIIII(I>I (II CYTOK
III11<YOIIIIIIII)
II - III1'fiilo-T}i'ynllnfi na]lHraHrJmii: (j - I\UUJlaJilUI C'l1I\1I1aTltQee1\J1X 3,,:;If'I\.1PIITCJB; 6 - '1)"8-
CTDIlTC.II>I1bIii Y3CJI GJIYH'AilIOIIII'rn ''''\1":1. 01<11. MeTlIJIODbIM 3PJICilbIM-mII"JlUlilOM. OH X 7,
oiL X 20
n - COCTaD M1trIIHp)'1011lI1X H.lI("rOH UJ],UJlh 1It."IHlHOil BeTO'IJUH, l1JJ,yu\eil liT nl']IXnerO Iuc'fiHoro-
CJIMnaTlI'll'eHoro Y3J,a (I'M. "G» I1a 111I1'. !j!I, A) (1I\1I1,iMm<, 1%0)
251
'I'OqHOrO MaTeplIaJIa lIaparaHI'JIHeB CJIymaT KaK CHMnaTuqeCKlIe, TaK
H lIapaCIIMnaTIPIeCKlIe Y3JIbl. B 1I0JlbSY P;BYX IICTO'IHIIKOB, IIOMHMO
.OIIIICaHHoro, roBOpHT MHOrO'llICJIeHHble MeJIKlIe napal'aHI'JIlIlI, nplI-
-CYTCTBYIOIqHe KaK 110 XOP;Y 6JIYmp;aIO el'O HepBa (Bblme II HlIme
ganglion nodosum), TaK II BHYTpII CIIMnaTlIqeCKIIX lIyqKOB, OTXO-
J.l:JI IIX OT OOpXHel'O meMHoro CIIMnaTlIqeCKoro YSJIa IIJIII IIpeBep-
1'e6paJIbHblX IIpeKapOTH,IJ;HblX CTBOJIIIKOB Term. ITo D;HTOJIOI'IIqOCKHM
nplISHaKaM OHII HuqeM He OTJIlIqaIOTCH P;PYI' OT P;Pyra: oKPymeHbI
COep;IIHIITeJIbHOTKaHHOM KanCYJIOM II cop;epmaT B CBoeM COCTaBe, Ha-
pHP;Y C xpoMa4>4>IIH06JIaCTaMII, nepexop;Hble 4>OpMbl OT KJIeTOK cnOH-
TII06JIaCTII'IeCKOrO THna.
I{ KOH Y 3M6plIGHaJIbHoro nepHop;a npOD;ecc p;II4>4>epeHD;IIpOB:KH
:xpOMa4>4>HHHblX 3JIeMeHTOB Y KYPJIHblX sapop;blmeM He aaKaH1J:lIBa-
-eTCH. B naparaHI'JIIIHX BblJIynHBmlIxcH D;blIIJIHT II B Te'IeHlIe 30 p;HeD:
IIOCJIe BblJIynJIeHIIH coxpaHHIOTCH, HapHp;y C KJIeTKaMII I'JIHaJIbHOI'O
"TlIlIa, MHOmeCTBO nepexop;HblX 4>OpM K xpoMa4>4>IIHHblM. I{JIeTOqHble
.3JIeMeHTbI B HIIX rpynnlIpyroTCH B P;OJIbKII II oKpymaIO'l'CH CIIHyCO-
"HAHblMII KaIIIIJIJIHpaMII. -
B 3M6plI0l'eHe3e II nOCTHaT3JIbHOM neplIop;e paSBIITHH meD:Ho-
TPYP;HblX lIapal'aHI'JIlIeB y KYP He yp;aeTCH YCTaHOBHTb MeTOp;aMII
.xpOMHpOBaHIIH II IIO]J.IIpOBaHHH cneD;II4>IIQOOKOD: peaKD;IIII Ha MOHO-
.aMlIHbI (ap;peHaJIIIH II Hopap;peHaJIIIH).
JaHHepBaD;IIH KapOTIIp;Horo IIaparaHTJIIIH y sapop;blmeM KYP qeTKO
npOCJIemlIBaeTCH yme Ha 7 -9-e CYTKII 3M pII0I'eHeaa. 06IIJIbHOe Ko-
.JIuqeCTBO BOJIOKOH, H,IJ;y IIX II3 ganglion nodosum, BCTYTIaIOT B I\a-
pOTIIP;HblM opraH II B CTeHKY 06 eM COHHOD: apTepIIII II paSBeTB-
.JIHIOTCH Ha TOHKlIe, 'IaCTO IISBIIJIIICTble nperepMIIHaJIbHble BeTOqKII.
ITo xop;y IIX BCTpeqaIOTCH BaplI'KOSHbie B3P;yTIIH C HemHOM HeMpO-
'4>II6pIIJIJIHpHOM ceTKOH. ITpeBepTe6paJIbHblH npeKapoTIIp;Hbli\: CTBOJIHIt
.Ha HMnperHlIpOBaHHblX cepe6poM npenapaTax 06HapymlIBaeTcH n
:B1Ip;e TOHKOI'O IIyqKa HepBHblx BOJIOKOH, KOTopble UP;YT Ha neKOTO-
pOM paCCTOHHlI1I OT CTBOJIa 6JIYmp;aIO ero HepBa, IIpOKClIMaJIbHee
-ganglion nodosum (pIIC. 90). Ha ypOBHe aaKJIap;KII IITOBII HOH
II lIapa IITOBH]:(HOH meJIes 3TOT CTBOJIIIK TepHeTCH, TeCHO nepenJIe-
"TaHCb C BOJIOKHaMlI, npOXOp;H 1IM1I B COCTaBe 6JIYmp;aIO eI'0 HepBa,
'ITO aaTpyp;HHeT IIX paSrpaHlIqeHlIe IIpII BCTYnJIeHHH B KapoTIIp;HblH
napal'aHI'JIIID:. ITpeTepMIIHaJIII SaKaH1J:IIBaIO'l'CH Ha ero KJIeTKaX pa3-
Ho06paaHLlMII 110 4>opMe HepBHblMII OKOHqaHIIHMlI, 06pasYH lIeTeJIb-
.fill, KOJIe1ffill, rryI'OBKII. BamHo 1I0p;'IepKHyTb, 'ITO nop;06Hble OKOHqa-
HIIH HMeIOTCH II Ha CTeHKe 06 eH COHHOH apTepIIII (ITpIIHMaK, 1962).
HepBHoe cHa6meHIIe OKOJIOBaraJIbHoro 1I8paraHI'JIHH 3Ha'IIITeJIbHO 6ep;-
"Bee (BO BpeMH 3M6plIOHaJIbHorO pasBIITIIH) KapoTIIp;Horo II oc eCT-
BJIHeTCH TOJIbKO BOJIOKHaMII 6JIYmp;aIO el'O HepBa.
OCHoBHble BblBOP;bI, KOTOpble CKJIap;blB8IOTCH B pe3YJIbT8Te IISY-
"'leHIIJI. 3M6plIOHaJIbHOl'O rlICTOreHesa meiiHo-rpyp;Hblx naparaHI'JIlIeB
y aapop;blmeH RYP, CBOP;HTCH K CJIeP;YIO eMY. Bo-nepBblx, RJleTKlI,
yqacTByro IIe B IIX 06pa30BaHIIII, IIpIIHap;JIemaT R MeJIKIIM I'llnep-
XpOMHblM nonYJIJID;IIHM cnoHI'II06JIacTuqecKOl'O TlIna II CXOp;HbI 110 D;H-
:252
4 6 7 4 5 I 8
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--
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I' II c. !)O. CxelIIa 3aKJIaJ];KlI KapOTlIP;lloro naparaHrJIlIH II ero lIHHepBaLU1H H3
g. nodosum 6JIYlli}J;aIO ero BepBa 3apO}J;LIDJa KYPJll\LI (8 CYTOK HHRy6aI(Xm)
1 - g. nodosum; 2 - CTBO.'I n. vag1; 3 - npCBcpTeOpaJlhHhIt1: npCHapOTHAHHt1: CTBOJlHH;
-1 - COHH8H apTcpHFI, pa3BHJlHa; oS - HcpBHhIe BO.'IOHFla B HapOTHAHoM napal'aHI'JlH\I!; 6 -
OHOJIOII]\I!TOBHAHhIC lliCJlC3h1; 1 - OHO.'IOBal'a.1IbHhIlt napal'aHI'JIHIt; 8 - Il\HTOBHAHaFi lliCJleaa;
9 - nHIl\CBOA IlIpHi1MaH. 1!J(2)
TOJIOrHqeCKIIM npll3HaKaM C OIIIIcaHHbIMII B 6plOllIHblX naparaHrJIIIHX.
BO-BTOPbIX, 8TII RJIeTIUI I1MeIOT p;BoiicTBeHHoe npOIIcxomn:eHHe 11;1
nap8ClIl\maTIIqeCKIiX (npeIlMy:r:u;eCTBeHHO) II CHMnaTllqeCKHX HepBHblX
Y3:IOB. YqaCTlIe qYBCTBHTeJIbHblX Y3JIOB napaCHMnaTH'leCKHX HepBOB
B 06pa30BaHIIII napeHXIIMbi llIellllo-rpYP;HblX naparaHrJIlIeB nTIIl\
(B TOM qIlC.'IC RapoTIIAlIOro) ;). X. llpnllMaR paCCMaTpXIBaeT C pa3-
HUX TOqeK JpeHIIH. C OP;HOII CTOpOHbl, Ha TOM OCHOBaHlIlI, qTO B
COCTaBe 6.'lymAalOlQerO HepBa IIMelOTCH CIIMnaTHqeCKHe BOJIOKHa II
HepBHb1P K'ICTKII (fOPOAIIHcI.;aH, "1936; fOP;IIHOB, "1939, H p;p.) MomHO
npep;nO:laraTb, 'ITO HapHP;Y C nOCJIep;IIIIMII B CHCTeMY 6JIymAalOlQerO
HepBa npOHIIKalOT H cnOHrII06JIaCTH'leCKIIe KJIeTKII CIIMnaTH'leCROrO
npOlICXOi-K;:J:eHIIH. TaKoe npep;nO:IOmCIIne TeM 60JIee BepOHTHO, qTO
BepxHIIii llIeiiHblii CIIMnaTII'leCImii yac.'I B nepIIOp; 8M6pIIOH8JIbHOrO
pa3BIITIUI l\bllIJIeHRa TeCHO npXl.'leraeT K CTBOJIY oJIymp;alOr:u;ero Hep-
Ba. C pyroii CTOpOHLI, HCJlb:\fl OTInu\aTb II B03 lOmHOCTli IIpenpa-
r:u;eHllH R:IeTOR I'JIII3JlbHOl'O TlIlIa napaCIIMnaTH'leCROro npollcxomAe-
HIIH n xpmwq)(InlHouJl8cTbl. l\IPJlI.;aH. rllllepxpOMHaH RJleTOqHaH no-
nYJIHl\IUI lIa panHlIX CTaAIIHX 8MopnoreHe3a Tomp;eCTBeHHa nep;lItP-
tPepeHl\npOBaHHbIM KJleTl\8M, a n03me cxop;Ha co cnOHrII06JIaCTaMII,
n3 ROTOpblX pa3B1IBaeTCH HCH nepntPepllqeCKaH rJIIIH. fJInaJIbHble
8.'IeMenTbI, nOBclOAY conpOBOH{AalOllI,lIe CIiMnaTHqeCKlIe II napaCHMna-
TIIQeCRIIe HeiiponLI, npnnap;JlemaT I, MaKpOrJllI1l - OJIllrop;eHp;porJIHII
n MorYT 06JIa]:(aTb 0p;HllaR0BLIMII lIOTeHl\uaJIbHblMII B03MmKHOCTHMH
253
B CMblCJIe CBOIIX npeBpa eHllii. B-TpeTbllx, BO BCeX lIaparaHrJIIIHX..
II3 RaROro 6bl IICTOqHIIKa OBII 1111 pa3BIIBaJIIICb, peaKn;IIII XpOMlIpO-
BaHIiH II IIOp;lIpOBaHIiH He p;aIOT nOJIOmlITeJIbHOrO pe3YJIbTaTa lIa nplI-
CYTCTBlIe ap;pellaJIIiHa II Hopap;peHaJIIIHa (T. e. no RJI8Ccu«lmRa:n;IIH
W atzka 011I1 Bce (<HexpOMatPCPIIHHble»). B-qeTBepTbIX, IIHHepBan;IIH
KapOTIIJ\HOro naparaHrJIIiH 06eCneqllBaeTCH BOJIOKHaMlI, lIP;yl.I\IIMIi 113-
ganglion nodosum M, BeCbMa BepOHTHO, qepe3 Hero BP;OJlb 6JIym-
p;aIO eI'O HepB8. H. HIIM nplIcoe]:(IIHHIOTCH CIiMnaTlI'IeCKlie HepBbl..
KOTopble nepenJIeTaIOTCH C qYBCTBIITeJIbHblMIi IIapaCIIMnaTlIqeCRII-
Mil BOJIOKHaMIi 6JIY)H]:(aIOlQerO HepBa nepep; TeM, KaK BCTynlITb B
KapOTII]:(Hblii naj>araHrJIllii. AaHHbIe TIplIiiMaK nOJIHOCTbIO COBnap;aIOT
C JIIiTepaTypHblMIi (Boyd, "1937; Watzka, 1943; Adams, 1958), 110-
Ra3bIBaIO IIMII, qTO HepBHbIe BOJIORHa, npIIHap;JIema lIe pa3HbiM
HepBHblM CTBOJIaM, coep;IIHHIOTCH p;pyr C p;pyroM 3ap;OJIro P;O TOro,
ImR p;OCTMrHYT KapOTHp;Horo naparaHrJIlIH II 06pa3YIOT CJIOmHOe CfiJIe-
TeHlIe. 0 npe06JIap;aHIIII B 8TOM CnJIeTeHlI1I CIiMnaTlIqeCKIIX BOJIOKOII
(no HamliM p;aHHbIM, 8TO IIMeeT MeCTO TOJIbRO y MJIeKOnIlTaIO IIX).
no cpaBHeHIlIO C BeTBHMII 6JIymp;aIO ero, H3bIKO-rJIOTOqHOrO II no]:('b-
H3blqHOro, roBOpliTCH B pa60Tax Danielopolu, Manescu (1928), Cele-
stino da Costa ("1935, 1939, 1940), Boyd (1937b), Watzka (1938).
B pe3YJIbTaTe 113yqeHIIH MOPCPOJIOrllqeCKIiX II rlICToreHeTlIqeChllX-
3aRoHoMepHocTeii RapOTIIp;Horo naparaHrJIIIH y RpblC, Mblmeii II hYpo
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HorO Y3JIa 6JIymp;aIO ero HepBa; MlIrpan;IIH KJIeTOK B KapoTIlp;HbIK
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3TO He TaK: KapOTIIp;Hblii naparaHrJIIiH IIHHepBlIpyeTcH II3 Tpex IIC-
TOqHIIKOB Toro II p;pyroro npOllcxomp;eHIIH, a KJIeTKII B Hero Bce-
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254
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pO OHaqaJIbHblH :meMeHT xpoMa4>4>IIHHOH Tl\aHII npe CTaBJIeH MeJIKOH
TeMHOH epHOB KJIeTROB, KOTOpaH B TeqeHHe JIIITeJIbHOI'O BpeMeHII
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3JIeMeHTOB HepBHoro 3aq8TR8 OHa yqaCTByeT? OTBeT Ha aTOT Bonpoc
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B IIOCJIe]:(yIOlI1;ne cpOTm p83RnTIIH (7 -"1 O-c cYTlm) IIpOIlCXO;'];IIT
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CTaMII II 1I03BOJIHeT OTHeCTII nx K fiJIeTI\aM, npOp;OJIlliaIOII.\HM CIIH-
Te:mpOBaTb J];HK (pnc. 91, A, B).
258
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80pTaJIbHblM CIIMnaTJI1IeCKHM 3aKJIa]J;KaM (cp. plIC. 93, A, B C
pllC. 93, B). HBJIHeTCH JIH Talioe n8]:(eHlIe pe3YJIbTaTOM IIOBbimeHIIH
aCIIHxpOHHOCTII pa3MHomeHIIH, lHacTynaIOr:u;eH 6JIa 1 ro]J;apH MHTpan;IIII
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HO IIpep;IIOJIOmIITb, qTO B Y3JIaX lIepBJI1IHOH CIIMliaTlIqeCKOH l\eIIO'IIUI,
RJIeTO'IHblH MaTeplIaJI KOTOpOH paCXO]J;yeTcH Ha 06pa30B8HHe BceH
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B - yqaCTOK, Oql'P'lCllIIbiii 113 A HBaApaToM. MeTRa onl'eaeJIHCTCH BaA l'UOBrHO-
6J1aCTaMH If xpoMa4>tPlI1I06Jlac raMlI, CIIMllaT06JJaCTId He Ae.1HTCH. OK. X 10,
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EllKa aapU;J.blllIa Kyplll\1d (1 CYTOl;: mmyual\IIII). l\1CTKa onpeAeJIHeTCH Ha;J. cnOH-
rlI06.'laCTa 1II (6), cmmaTOu.lacTbl (a) lie AeJIHTCH
{)H. x 10. 06. X 90
KapOTIUlJoro
'CTa TI,3HII,
'TE3HI'II H.
naparallrJllUI l,ypllHOro 3apO,l.J;blllla rJUIaJIbHblH Tim po-
COOTBeTCTBYIOIl 1I ii no relleTlIqeCKOii KJIaccmlHlRal HII
r. X.lollllua L11I03Uf'I1jJ.IllIlHbJM :meMeHT3M HepBHoH
TE3HII.
IhITepeCH3 T3 Ol\eHEa EJIeTOK, ROTOPYIO cp;eJIaJI Celestino da
Costa (1936, 1955) B pe3YJIbTaTe Ha6JIIOp;eHnii Hap; IIpOn;eccaMII
JJ;wI)q,epf'uIJ;Ilpomm RJJeTO'IllLIX :;meMeHTOB, yqaCTBYIOIqHX B q,OPMH-
pOBa1lI1II naparaHr.'IUI'B, B 'laCTHOCTII l,apOTIIp;Horo. Celestino da
Costa CBblllle 40 1teT (1913-19;)5) pa60TaJI Ha]:( IIp06JIeMOii q,YHK-
1.\1I0Ha:IbHOrO 3Haql'1I1IH l,apOTlI]:(lloro II ]:(pyrIIX lIaparaHrJIlIeB II B
neEOTopblX pa3p;eJIaX 6JIII3KO nOAXO,l.J;lIJI, Ha Halll B3rJIHp;, K peme-
HlIIO Bonpoca 0 ero iIJlI3110.'IOrIPlCCKOii pOJIII. OH nOJIaraJI, q10 co
<:TOpOHbl ::!MupnOJIOrIlII HCT ]:(3HULIX, ROTOpbIe CJIymllJIH 6b1 B rrOJIb3Y
BHOBL n03111maBllIe ii I,J1accIIq,III,al\UH rraparaHrJIIIeB, II OqeHb mHBO
pearllponaJI Ha pemI:mIO TCOpllll 06 3H,I.J;ORplIHHOll CYl.l1;HOCTll Kapo-
nqHoro naparaHrJIlIH (C£'lestino da Costa, 1936) co CTOpOHbl
4>1I3110."'lOroB II 1l1Opq,OJIOron. OlIllcLlBaH lIecTpblii COCTaB RJIeTOK B
3aqaTEaX naparaHr:lllen, Celestino da Costa 60JIblllOe BHIIMaHlIe
yp;e:Ill:I 1<teJIKoii TeMHOH,I.J;epHOii ImeTRe «PY,I.J;IlMeHTapHoro BIIp;a», nplI-
CYTcTBlle KOTOpoii OTMe'laJIll nce IlCCJIep;onaTeJIII rlICTorCHe3a CIIMlIa-
TII'recEoll HepnHoii ClICTeMLI. TaK RaK alnop ynoTpe6JIHeT TepMHH
«PY]:(InleHTapIlLIll», 1,0TOpLlii q,lIrypHpyeT II B HacTOHl.l1;eii pa60Te,
TO BO 1136ea,aHlle IlCnpaBII:lbHLIX TOJIKOBaHnll YTOqHIIM IIOHHTIIH,
265
ROTopLIe B HIIX BRJIa)J;LIBaIOTCJI. PYP;HMeHTa:qlIeH 06I:qeIIplIHHTO Ha3LJ-
BaTb npOHBJIeHlIe peP;YR:q1l1I OpraHa, ROTOPLIii B npo:qecce cPlIJIore-
He3a YTpaTIIJI IIJIII CHII3IIJI OCHOBHOe cPYHR:qIIOHaJIbHOe 3HaqeHIIe II
COOTBeTCTBeHHO II3MeHIIJI CBOe CTpOeHlIe. B MaTeplIaJIaX npep;LIP;Y-
m;IIX rJIaB 6LIJIO nORa3aHO, 'ITO MHOrlIe naparaHrJIlI1I BLICIlIIIX n03-
BOHOqHLIX II RJIeTlill, IIX COCT8BJIHIOI:qlIe, JlBJIHIOTCH Pyp;IIMeHTapHLIMH,
T. e. YTpaTIIBllIlIMII 6LIJIYIO cPYHR:qIIOHaJIbHYIO aRTIIBHOCTb. «PYP;II-
MeHTa.:qIIJI) naparaHfJIlIeB B nOHIIMaHlI1I Celestino da Costa nplIH-
:qllnlIaJIbHO IIHaH. OH paccMaTplIBaeT «Pyp;IIMeHTapHLIe» naparaHrJIlI1I
MJIeROnlITaIOI:qIlX RaR aBOJIIO:qIIOHHO Hep;Ora3BIITLIe opraHLI, T. e. He
p;OCTlIrmlIe eI:qe MOPcPOcPYHR:qIlOHaJIbHOH nOJIHo:qeHHOCTII, a He B R8-
qeCTBe CTPYRTypLI, YTpaTIIBmeH B npo:qecce cPJIJIOreHe3a cPYHR:qllo-
HaJIbHYIO aRTIIBHOCTb.
R COmaJIeHIlIO, OqeHb MaJIO CBep;eHllii DMeeTCJI 06 IICTOqHIIRaX
pa3BIITIIH ap;peHaJIOBOH TRaHII Y HII31l111X n03BOHOqHI>IX. B CBH311 C
pa36HpaeMLIM BonpOCOM HeJIb3H He BcnOMHIITb 0 pa60Te Chevrel
(1889), B ROTOpOH aBTOp, II3yqaH mCToreHe3 «aRCIIJIHpHOrO cepp::qat
pLI6, npe]:(nOJIOmIIJI, qTO naparaHrJIIIOHapHaH RJIeTRa no cBoeii npll-
pop;e npep;CTaBJIHeT OC06LIH BlIP; rJI1I1I CIIMnaTlflIeCRIIX Y3:IOB. J],JIJI
Toro BpeMeHII p;ame TaRoe npep;nOJIOmeHlle aBTopa CBIIp;eTeJIbCTBO-
BaJIO 0 BLICOROM TeXHlIlleCROM ypOBHe ero IICCJIep;OBaHIIH, n03BOJIIIB-
meMy' rOBoplITb He TOJIbRO 0 TEaHeBOM IICTOqHIIRe pa3BIITnH napa-
raHrJIlIeB, HO II 0 RJIeTOqHOM.
3aRaHqlIBaH II3JIOmeHIIe MaTeplIaJIOB 0 RJIeTOllHLIX IICTOllHIIRaX
pa3BIITIIH xpOMacPcPlIHHoii CIICTeMLI, nOCTapaeMCH nop;LITomlITb pe-
3YJIbTaTLI IIX II3YlleHIIH.
MI>l npe]:(nOJIaraeM, qTO nYTb pa3BHTJIH B OHToreHe3e xP01l1acPcPlIH-
HOH RJIeTRH MomHO npep;CTaBIITb CJIeP;YIOI:qIlH cxeMOH:
HeAIIIP<I>epeH HpOBaHHaH RJleTRa HeiipaJIbHOrO 3aqaTRa
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CIIMnaT06JIaCT 3aRaH'IlIBaeT npOJIlIcPepaTIIBHYIO aRTIIBHOCTb II CTa-
HOBIITCH Ha rryn p;eTepMIIHlIpoBaHHoro pa3BIITIIH P;O Toro, EaR Ha-
'IlIHaeTCH HOBo06pa30BaHIIe xpOMacPcPlIHHoii MeTRII. 01leBIIp;HO, po-
JJ;OHaqaJIbHYIO «CTBOJIOBYIO» RJIeTRY CJIep;yeT cqlITaTb nJIIOplInOTeHT-
HOH TOJIbRO B CMLICJIe oupa30BaHIIH II3 Hee P;BYX KJIeTOqHbIX nouy-
JlJI:qIIH - rJIlIaJIbHOH II xpOMacPcPIIHHOH. ECJIII me nplI3HaTb cyr:u;eCTB«;:
BaHlIe ep;IIHOrO IICTOqHIIRa P;JIH Bcex Tpex KJIeTOqHLIX nOIIYJIH:qIIII,
TO Torp;a Ha]:(o paCCMaTplIBaTb xpOMacPcPlIHHble ;)JIeMeHTLI Ral{ caMO-
CTOHTeJIbHYIO BeTBb RJIeTOR, pa3BIIBaIOI:qlIXCH HapHP;Y C TJIIIaJIbHLIMII
II HepBHLIMII aJIeMeHTaMII.
266
B 3aHJIlOqeHIIe OCTaHOBllMCH EpaTHO Ha HeHOTOpLIX TeOpeTlI'Ie-
(;IrnX BOlIpOCaX, CBH3aHHbIX C 8TOH IIp06J1eMOii.
HapHAY C OpraHOreHe30M, HOTOpLIH OTHOCIITCH H qlICJlY BamHLIX
np06J1eM B 8BOJlIOD;IIOHHOii Mop<}>OJlOrlIlI, He MeHblllee 3HaqeHIIe IIMeeT
II3yqellue D;nTOJlOrlIqeCEIIX II rIlCTOJlOrlI'IeCHIIX ]:(II<}><}>epeHD;IIpOBOE.
H. r. XJlomlH (1946) nOJiaraJl, qTO 8M6plIoJlorlIqeCEIIii MeTOA B
H3YQeHlUI ]:(1I<}><}>epeHD;lIpoBHB THaHeii IIMeeT HOJlOCCaJlbHOe 3HaqeHBe
npII COIIOCTaBJleHlIII IIpOI{eCCOB BHAIIBBAyaJlbHOrO pa3BIITIIH C <}>aH-
TaMll lla cpaBHIITeJlbHOH aHaTOMBII. OH cnoc06cTByeT nOHIIMaHIIIO
2aRoHollleplIocTeii fiaH <}>BJlOreHe3a, TaE II OHToreHe3a.
BcnoMoraTeJlbHI>Ie 8J1eMeHTI>I HepBHoii CIICTeMLI OTJlII'IaIOTCH 60Jlb-
IDllM MOp<}>0<}>ll3IIoJlOflIqeCHIIM pa3H006pa3IIeM MeTOqHLIX TlInOB, EO-
Topble He IIOAXOAHT HII nOA HaROH 1I3 qeTLIpeX rpynn THaHeii HJlaC-
CIIQeCfiOll rIICTOJlOrnB. Bce MHoro06pa3IIe rJllIaJlbHLIX 8J1eMeHTOB no
cBoeii MOp<}>OJlOflIqeCHOH B <}>YHHD;IIOHaJlbHOH xapaHTeplICTIIHe BXO):{IIT
B COCTaB Bcex qeTblpex rpynn THaHeii, 6y):{yqII B TO me BpeMH
reHeTIIQeCfill e):{IIHbIM. 1I3yqeHIIe IIX rlICToreHe3a B 8Hcnep1IMeHTaJlL-
HblX YCJlOBIIHX nOMoraeT 06'1.e):{IIHeHIIIO IIX B e):{IIHLIii rJllIaJlbHO-8neH-
):{IIMHblii THII (XJlOIIIIH, 1946). IIx rlIcTo6J1aCTIIqeCEIIe (TKaHe06pa-
3YIOID;IIe) nOTeH:QIIB B 8ECnJlaHTaTax 06HapymlIBaIOT '1epThI CXO):{CTBa,
HeCMOTpH Ha pe3HYIO pa3HIID;y B CTpYEType IICXO):{HLIX TEaHeR. qep-
TLI CXO):{CTBa, OqeHb HpHO BblcTynaIOID;Be B YCJlOBIIHX HYJlbTIIBlIpo-
BaHHJI BHe opraHII3Ma II OTJllI'IaIOID;lIe IIX OT THaHeii ):{pyroro npo-
IICXom]:(eHIIH, B COBOEynHOCTH Ha3BaHLI H. r. XJlOnIIHLIM «TRaHeBOD:
]:(eTepMllHaD;Heih> .
IIpII HYJlbTIIBlIpOBaHIIH pa3HLIX yqacTHoB raHrJlII03HOD: IIJlaCTIIH-
HII, ):{aIOID;eH (<aHTOMe3eHXHMY», Ha6J1IO):{aeTcH pOCT, xapaETepHLIH ]:(JlJI
MHorlIX THaHeD: (xpOMaTo<}>opOB, xpHII\a, HeiipaJlbHLIX 8J1eMeHTOB
II ):{p.). Cpe):{II HBX IIMeIOTCH THaHH, EOTopLIe no rlICTOJlOrlIqeCHIIM
oc06eHHOCTJlM pOCTa B RYJlbType 06HapymlIBaIOT OTqeTJlIIBOe CXO):{-
CTBO C pOCTOM BcnOMOraTeJlbHLIX TEaHeH HepBHOH CIICTeMLI (XJlOnllH,
1946). 8TIIM <}>aHToM ]:(OEa3LIBaeTCH, qTO HJleTOqHLIH MareplIaJl B
8MUplIOHaJlbHOH 3aHJla):{Ee, eID;e ):{O Toro HaH OH npOllleJi Bce CTa):{IIII
]:(1I<}><}>epeHD;lIpOBIrn, 06HapymlIBaeT cBoiicTBeHHLIe eMY ifIICT06J1aCTII-
qeCHlIe nOTeHD;IIII.
rJllIono):{06HLIii pOCT xapaHTepeH, HaH rOBOplIJlOCb Bblllle, ):{JlH Me-
TOH EapOTII):{Horo naparaHrJlllH (IIoHoMapeBa, 1946). RJleTOqHLIe reHe-
paD;1I1I rJlnaJlbHOrO TlIlIa, HOTopLIe HBJlHIOTCH xpoMa<}><}>IIHo06pa30Ba-
TeJlbHLIMII 8J1eMeHTaMB B RapOTH]:(HOM II ):{pyrnx naparaHrJlIIRX,
YCTaHOBJleHHLIe HaMil Ha Tpex rpynnax n03BOHOqHLIX (nTHD;LI, aM-
<}>II61I1I, MJleHOIIHTaIOID;lIe), ne):{YT, CJle):{OBaTeJlbHO, HaqaJlO OT 06pa-
30BaTeJlbHI>IX HJleTOE raHrJllI03HOH nJiaCTIIHIrn. E):{BHcTBeHHoe B03pa-
meHlIe, HOTopoe, Ha Halll B3rJlJlJ{, MomeT 6LITb BLI):{BIIHYTO npOTBB
rJIII06J1aCTllqeCEOH npBpO):{LI xpOMa<}><}>IIHHOii THaHH B IIOJlb3Y Heiipo-
6J1aCTBqeCROH - 8TO B3BeCTHLIe Haxo]:(1rn y 6ecn03BOHOqHLIX. Gaskel
(1914) II Ziller-Perez (1942) nOHa3aJlII nplICYTCTBlIe a):{peHaJiIIHa
(nOJlOmIITeJlbHaH xpoMa<}><}>IIHHaH peaHI{IIH) B HeEOTopLIX II3 MOTOp-
HLIX HeiipoHoB 6pIOllIHOii D;eIIO'lRII aHHeJllI)];, a Ehinger, Falcl(,
267
Myhrberg (1968), Rude (1969), Bianchi (1969) o6HapymBJIH
B IllIX .pJIyopeCI{eHI{IIIO MOH08MIIHOB. rOBOpH 0 .plIJlOreHeTlIqeC;HOM
3HaqeHIIII 3Toro .paHTa, Ziller-Perez BLlCHa3aJlCH B nOJIb3y .pYHH-
I{BOHaJlbHOrO 06"1>ep;IIHeHHH B op;Hon IDle THe P;BYX .pYHRD;IIn - HepB-
Hon (MOTopHon) 11 3HP;OHpIIHHon (xpOMa.p.pIIHHon), HOTopble B p;aJIL-
HeiimeM pacnpep;eJlIIJiIICb II P;BYX CTpYHTypax - B CIIMnaTlIqeCHOn B
xpOMa.p.pIIRHon. CJIe]:(OBaTeJIbHO, B ep;IIHon HepBHon HJIeTHe aBTopLl
BIIp;eJIII .plIJIoreHeTlIqeCKOrO npep;meCTBeHHHHa ]:(BYX 3J1eMeHTOB II HaK
6L1 npep;pelllaJlII ero HenpaJIbHYIO nplIpoP;Y.
B HaCTOHID;ee BpeMH HeCOCTOHTeJIbHOCTL nop;06HbIX coo6pameHIIB
HCHa. Tenepb II3BeCTHO, qTO MOHoaMIIHLI Y BLlcmlIX n03BOHOqHLlX,
B TOM qlICJIe Y MJleHOnIITaIOID;IIX, nplICYTCTBYIOT He TOJlblW B XpO-
Ma.p.pIIHHLlX HJleTRaX, HO o6HapymeHLI .pJlyopeCI{eHTHLlM MeTop;OM B
B D;IITOnJla3Me CIIMnaTlIqeCHIIX (MOTOpHLlX) HenpOHOB (Eranko, Hiir-
konen, 1964; Norberg, Hamberger, 1964). CJIep;OBaTeJIbHO, B CJIY-
qae C aHHeJIIIp;aMII peqb IIp;eT He 0 .pIIJIOreHCTlIqeCHOM npep;llIeC'f-
BeHHIIHe P;JIH IDIeTHII MJIeHOTIIITaIOID;IIX, a 0 HJleTHaX, aHaJlOrlIqHLlX
no .pYHHD;IIII. Ho eCJlII p;ame P;OrryCTIITL, qTO Y aHHeJIIIp; 06e .pYHH-
I{1I1I - HepBHaH II 311p;OHplIHHaH - 06"1>ep;IIHeHLI B O]:(HOM HenpaJlb-
HOM 3J1eMeHTe, TO MomHO TIpep;nOJIOmlITb IIX p;aJIbHeiimee p;IIBepreHT-
Hoe pa3BIITlIe. B 3TOM OTHomeHlI1I MomHO onHTb COCJlaTbCH Ha
H. r. XJIOnlIHa (1946), HOTopLln rOBOplIJI, qTO (mpIIHD;HlIaM pa3p;e-
neHIIH II CMeHLI .pYHHD;IIn B JlBJIeHIIHX p;HBepreHTHoro pa3BIITlIH THa-
Hen HeCOMHeHHO nplIHap;JlemlIT nepBOCTeneHHoe 3HaqeHlIe».
B 3aIDIIOqeHlIe XOqeTCH OTMeTIITb, qTO 1I3yqeHlIe 3M6plIoreHe3a
naparaHrJllIeB HMeeT 3HaqeHIIe P;JIJI pellleHHH TIpIIHD;IITIlIaJlbHLlX Bon-
pOCOB HaCTOHID;en pa60TLI. PeaynbTaTbl ero CHIIMaIOT 3HaqeHIIe P;IIC-
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269
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OCYIqeCTBJleHHII ceHpeTopHoro npo:qecca.
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nOJlHo:qeHHOCTb peillJleETopHoH meJle3HCTOn aHTHBHOCTII.
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mma (15-20 eA. Ha 100 Hera TeJla', IIO]:(liOmHO) II He Bbl3blBaIOT
ceRpe:qllIO' JlOpaApeHaJlHHa (H6kfelt, 1951). AnH IICCJleAOBaHHH
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JIeHHH RJlHHllqeCIUI BblpameHHoro rllnorJlHReMUqeCHOrO mOI\a, ROTO-
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ro 3a611BaJlJI B pa3I1bIe CpORll nocJle BneAeHIIH HHcYJlIIHa: qepe3 9,
11, 24, 48 qaCon, 3 H 6 CYTOK. Bo Bcex onbiTax O]:(HH HaAnoqeq-
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MaR), APYl'Oii - B CBeTOBOM MllHpOCRone (1OIlIHeBHq); B nOCJIeAHeM
CJIyqae npOBO]:(HJlaCb peaH:qHH Ha npllCYTCTBIIe B RJleTKax aApeHa-
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Ha.l{nOqeqHIIROB He.l{aBHO nORaaaHId 6UOXIIMII'leCKIIMII MeTO.l{aMH B cepllll pa-
60T Viveros et al. (1969a, b), Viveros, Arqueros, Kirshner (1969).
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Y ROHTpOJlbHblX II IIop;onblTHblX mHBOTHbIX. PaCIIOJlOmeHHe ROMnJleR-
ca fOJlbp;mH B OCHOBHOH Macce RJleTOR onblTHOH meJle3bl MeHHeTCH.
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OT]:(eJlbHbIX 3epeH IIJlH tl>parMeHTOB HHTH. B TOD me ]:(OJlbRe BH]:(Hbl
fiJleTRII, 3anOJlHeHHbIe rpaHYJlaMH ceRpeTa. ROMIIJleRC fOJlb]:(mII B HHX
coxpaHHeT CeTqaTYIO tl>OPMY H OROJlOHp;epHoe pacnOJlOmeHHe. HaH-
60Jlee OTJlHqHTeJlbHOll qepTOH CTpyRTypbl MHTOXOH]:(plIll B xpoMatl>-
tl>HHHbIX RJleTRaX RpLIC, Haxop;Hm;HXCH B COCTOHHHII mORa, HBJlHeTCH
YHIItl>IIRa HH IIX tl>OpMM H IIpeBpam;eHlIe B ORpYrJIbIe HJlH ctl>epolI]:(-
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CH YBeJlIf'leHHe pa3MepOB MHTOXOH]:(pHH II B03paCTaHHe HX qHCJla
(CM. pHC. 100, B).
B pa3HMe CpORH nocJle BLIBe]:(eHIIH RpLlC H3 mOROBoro COCTOH-
HHH MomHO Ha6J1IOp;aTb IIpo ecc BOCCTaHOBJleHHH ceRpeTa B RJleTRaX
(IOIllReBHq, 19666). qepe3 9 II 12 qaCOB 1I0CJle BBe]:(eHHH IIHCY-
JIIlRa H BLIBe]:(eHHH RpblC H3 mORa xpoMa4JqJHHHaJl peaR HH OCTaeTCH
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HaJlIIH, p;aIOT OTpHD:aTeJlbayIO rHCTOXIIMHqeCRYIO peaRn;HIO. HaqHHaH
C 24 qaCOB, nOJlOmHTeJlbHaH xpoMa<I><I>HHHaH peaR HH 1I0CTeIIeHHO
HapaCTaeT, 3aXBaTMBaH Ha 2-e II 3-II CYTRH Bce HOBLIe n HOBbIe
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IIOqeqHHRa RpMCLl ORpamnBaeTCH CTOJlb me IIHTeHCHBHO, RaR Y ROHT-
pOJlbHMX mnBOTHbIX.
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pe3 9-12 qaCOB ero CTpYRTypa MaJlO oTJlnqaeTCH OT TOD, ROTopaH
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XOp;Hm;HXCH B mORe: Ha6J1IO]:(aeTCH IIporpeCClIpYIOm;aH rlIIIepTpotl>HH,
pa3pMXJleHne n paClIpOCTpaHeHHe «P;IIRTHOCOM& no Bcen HTOnJla3Me
RJleTOR. qepe3 24 qaca 3J1eMeHTbl ROMIIJleRCa HaqHHaIOT CTHrHBaTbCJI
R JlP;PY. 3TOT IIpo ecc nOCTelIeHHO HapaCTaeT, II R 6-M CYTKaM pac-
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MHRpOCROIIIf'IeCROM ypOBHe 06HapymHBaeT eJlblH ROMIIJleRC lIepe-
CTpoeR MeM6paHHblx cnCTeM xpoMatl><I>IIHHLIX RJleTOR B pa3HMe <I>a3bI
nx mn3He]:(eHTeJlbHOCTII. OHn onncaHLl ]:(OCTaTOqHO no]:(p06HO B cepllII
pa60T ITpHiiMaR (1965a, 6, 1966, 1967). Mbl RpaTRO OCTaHOBIIMCJI
Ha Tex pe3YJlbTaTaX, ROTopble IIrpalOT HaH60Jlee cym;ecTBeHHYIO pOJlb
B ceKpeToo6pa30BaTeJlbHOH H Bblp;eJlHTeJlbHOH <I>YHK IIHX RJleTOK: HB
274
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B mOKOBOM COCTOJlHHH KpblC Ha6JIIO]:(aeTCJl Bbl6poc aApenaJIHH3I
H3 KJIeTOR. IIoKa3aHO HeCKOJIbKO IIYTeii, ocy eCTBJIJlIO HX em BLI-
]:(eJIeHHe. AApeHaJIHH COAepmHTCJI B KpynHblx 3anaCHbiX rpaHYJIax..
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HeHHJI BblCOKOii 3JIeRTpOHHOii nJIOTHOCTH (pRC. 98). CTeneHb-
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KaTeXOJIaMHHOB. 3TO CTaHOBHTCH oc06eHHO JlCHO B YCJIOBHHJ( 3Kcne-
plIMeHTa. IIpH HHTeHCHBHOll CTHMyJIJlqHH ceRpeTOpHom npoqecca..
Bbl3BaHHoro mOKOM, Ha6JIIO]:(aeTCJI pe3Koe na]:(eHHe qHCJIa 3anaCHblx..
rpaHYJI, Ha MeCTe KOTOpblX OCTaIOTCH MeM6paHLl, JIHllIeHHLle OCMHo-.
cpHJIbHOrO COAepmHMoro, COXpanJlIO He pa3Mepbl 6LIBmHX rpaHYJL
H nepeXOAHLle cpOpMbl K HHM (pHC. 101). TaKaH ceKpeqHrr nYTeM
paCTBOpeHIIJI co]:(epmHMoro rpaHYJI H Bbl]:(eJIenHH em 6e3 nOBpem-
]:(eHHJI MeM6paHbi HBJIJleTCH HaH60JIee pacnpoCTpaneHHoii (CMHTTeH..
19656) H xapaKTepHa ]:(JIJI MHorHX 3HAOKpHHHLlX meJIe3. CY]:(b6a
nonaBmllX B qHTOIIJIa3MY KaTeXOJlaMHHOB H3BeCTHa em;e H3 nepBLlX
3JIeKTpOHHOMHKpOCKOnHqeCKHX pa60T de Robertis, vaz Ferreira
(1957a, b), de Robertis, Sabatini (1960). ELIJIO IIOKa3aHO,
qTO OIIH B paCTBopeHHOM BH]:(e AJlcpcpYH]:(HPYIOT qepe3 nepnrpa-
HYJIJlpHYIO H KJIeTOqHYIO MeM6paHbl, a 3aTeM nYTeM IlImoqHT03a
nepeHOCHTCJI qepe3 qHTOnJIa3MY 3H]:(oTeJIHaJIbHOll RJIeTKH B KpOBOTOK_
HapHAY C TaKHM MexaHH3MOM Bbl]:(eJIeHIIH KaTeXOJIaMIIHOB Y KPblC..
HaXO]:(JlII{HXCJI B COCTOJlHHH mORa, HMeeT Memo cpparMeHTaqHH 3a-
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II3 OTAeJIbHblX 3epHbimeK, Kam]:(oe H3 ROTOpblX COAepmHT KanJIIO Ra
TeXOJIaMIiHa H oRpymeno MeM6paHoii (pHC. 101). IIpIl pa3pyIIIeHHII
MeM6paHLI 3anaCHbiX rpaHYJI HJIH nYTeM oTmHypoBKH Me;IKHe ny-
3blpbKH nonaAaIOT B qHTOnJIa3MY (pHC. 102). MeJlROBe3HRYJIHpHblm
KOMIIOHeHT qHTOnJIa3Mbi xpoMacpcpHHHoii KJIeTKH, IIOHBJIJlIOm;HllClI B
YCJIOBHJIX 06HJIbHOrO BblAeJleHHH KaTeXOJIaMHHOB, HBJIHeTCH, KaK no-.
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PH]:(Y C ]:(HcpcpY3Heii COAepmHMoro MeJIKHX Be3HKYJI qepe3 KJIeTOq-
HYIO MeM6paHY rryTeM IIpeABapHTeJIbHOm CJIHHHHJI 3THX MeM6paH
B qHTOnJIa3Me 060c06JIHIOTCJI qeJIbie KOMnJIeKCbI TaKDX IIY3LlPbKOB..
GHII BCTpeqaIOTCH KaK no nepHcpepHII ImeTKII, TaR H B Cy63H]:(0-.
TeJIHaJIbHblX H MemKJIeTOqHLlX npoeTpaHCTBax (pHC. 103). 3TII
+--'
PH C. 102. Pa3JIII'lHble 4>OpMId aanaCHbiX rpaHYJI xpmm4>4>IIHHIdX KJIeTOK Hap;no-
'le'lHJIKa KpblCbI B 4>a3c Bblp;eJIeH;JIH ceKpeTa. Pacnap; RaTeXOJICOp;eplli8lI\HX rpa-
HYJI Ha OTp;eJIbHble aepHa. MHOrO'lIlCJIeHHIdC MeJIRJIC Be311KYJIbi B l\lIronJIaaMC.
I1HCYJI;JIHOBbIH roOK
J!&Z - MeM6p8Hh1 I'PaHYJI (P8apymeHHe); 7IIA - rpaHYJIbi B BHAe TYTOBOA Hrollbl; om..
e. - OTIIIIfypoBI<8 Be3MI<TJIbI; nA. e - 8JIeI<TpOHHOnJIOTHbie Be3HI<YJIIi (UpHiblal<. 1965)
277.
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C MHorO'IIICJll'llllLIMH I\aTeXOJlCOp;epmall\lIMI1 IIc;nmYJlaJl\ll n paclIlllpelllIOM cyo;)n.
p;OTeJIJHI.TlbIlOM npoCTpallcTHc (cn)
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(IIpHIIMaH, 19(5)
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(pHC. 104). CpeAn HIIX - <<IlYCTbW», 8J1eRTpoHllonp03paqIlble H 3a-
278
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pacnpOCTpaHHIOTCH no HTOnJla3Me KJleTOR. Qepe3 9, 12 II TeM 60Jlee-
qepe3 24 qaca nocJle BBe,ll;eHIIH HHCYJlHHa MHOlKeCTBO 'l'aRHX nyablpb-
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HbiU Ha plIC. 105. Pa3Mep rpaHYJl nOCTeneHHO YBeJlH'IHBaeTCH H
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JlHpHLle H MelKKJleTOqHbIe npOCTpaHCTBa IIJlH C nOMOlI\bIO nHHO HTO-
3a (IIpHllMaK, 1967).
IIpH aHaJllI3e ceKpeTOpHblX tl>a3 xpoMatl>tl>HHHblx KJleTOK B CBe-
TOBOM MHKpOCKone moKOBoe COCTOHHHe II nepBblH IIeplIO]:( nocJle Her()
xapaRTepH3YIOTCH, KaK Mbl BH]:(eJlH, CTpYKTYPHOH H3MeHqHBOCTbID
KOMnJleKCa fOJlb,ll;IKH: OCMHOtl>HJlbHblU KOMOqeK, xapaKTepHbJll ,lI;JlH
nepHO,ll;a (mOKOJl» KJleTRH, pa3pbIXJlHeTCH, OCMHOtl>HJlbHLIe HIITH ,lI;po-
6HTCH II paCCblTIaIOTCH no HTOnJla3Me.
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BOH MHKpOCKonbl, HeBOJlbHO HarrpamHBaeTCJI aHaJlOrHH B nOBe,ll;eHHH
MeJlKOBe3IIKYJlJlpHOrO KOMnOHeHTa KOMIIJleKCa fOJlb,ll;IKH, ,lI;OCTHraIOlI\e-
ro MaKCHMaJlbHOH rHlIepTpotl>HH K MOMeHTY HaH60Jlee IIHTeHCHBHor()
CHHTe3a ceRpeTa B KJleTKe, H Tex tl>parMeHToB oCMlIotl>HJlbHOrO RJly6-
Ka KOMnJleRCa fOJlb]:(IKH, KOTOpblU paCCblTIaeTCH B 8TO BpeMH no.
Bcell HTOnJla3Me. XapaKTep ,lI;HHaMHKII B pa3Hble tl>a3b1 CeKpe IIII.
Tex H ]:(pyrHX CTpyRTYP - cy6MHKpOCKOnHqeCKHX rry3bJpbKOB H OCMHO-
tl>HJlbHOrO MaTepHaJla «,lI;HKTHOCOM», BblHBJlHeMOrO B CBeTOBOM MHK-
pOCKone, rOBopHT 0 B03MOIKHOCTII IIX HH]:(eHTHtl>HKaJ..t;HH, HeCMOTpJl
Ha pa3HbIe napaMeTpLl BeJlHqIIH. «)J.HRTHOCOMbI» HBJlH10TCJI 4>YHK Ho-
HaJlbHblMH aHaJlOraMH Be3HKYJI, pa3Mepbl ROTOpLlX He nO,ll;]:(aIOTCH
Ha6J1IO]:(eHHIO B CBeTOBOM MHKpOCKone C ero HH3KOH pa3pemaIOII\eH
clloco6HOCTbIO. Bo BCJlKOM cJlyqae H Ta II ]:(PyraJl CTpYRTypa 8KBII-
BaJleHTHLl B TOM CMblCJle, qTO HBJlJlIOTCJI HenOCpe,ll;CTBeHHblMH yqa-
CTHHRaMH HaKOnJleHHH B KJleTKe ceKpeTopHoro IIpO,ll;YKTa. H3BeCTHO,.
'ITO CIIoc06HOCTb KOH,lI;eHCHpOBaTb npO,ll;yKTbl ceRpe HH Ha nOBepx-
HOCTJlX CTpYKTYP KOMnJleKCa fOJlb,ll;IKH 6b1J1a BnepBLIe nOKa3aH3
)J.. H. HacoHoBLlM (1923, 1924) II B HaCTOHID;ee BpeMH npH3HaeT-
CJI KaK HaH60Jlee BepOJlTHaJl (Dalton, 1952, 1961; faTeH6lI, 1957;
Hirsch, 1960; TOKHH, 1963; Herlant, 1964, H ,lI;p.). IIoJlHMop-
tl>H3M CTPYKTYP KOMnJleRCa fOJlb]:(IKH paCCMaTpHBaeTCH B CBJl3H C
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Ta60JllITbi II cl>epMeHTbI (CMIITTeH, 1965 a, 6).
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MeM6paHHblx CTPYRTYP II o:o;eHRII IIX yqaCTBJI B Dpo:o;ecce ceRpeTO-
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.8 B3BeCTHOH Mepe, Hamil JRClIeplIMeHTbl C BBep,eHlIeM pe3epmma
(CMIITTeH, MOCRODRBH - HeoDy6J111RoBaHHble p,aHHble). OlIbiTbi CTa-
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BBOp,IIJlCJI pe3epnBH B p,03e 0,1 M.It Ha 100 a Beca TeJla. MaTeplIa;r
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MaR (1965a, 6, 1966, 19(7) B IOmReBlIq (1966a, 6) IIPB DpO-
Be]:(eHIIII pa60Tbi C BHCYJlIIHOM: Op,BH Hap,IIOqeQHIIR 6paJiB p,JlJI 3J1eR-
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opraHOB]:(OB RJleTRB Ha CBeTOBOM ypOBHe. M3MeHeHBJI, HaCTYnaK.-
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HlIe CpORII 1I0CJIe pe3eplIIIHB3a:O;BII mBBOTHbIX, TaR II B n03p,HlIe.
Pe3YJlbTaTbi onbiTOB, B RpaTROM II3J10meHIIB, 3aRJlIOqaIOTCJI B cJle-.
p,YIOIII;eM.
llpOJlOHrlIpOBaHHoe p,eHCTBBe pe3epnlIHa, ROTopoe BbI3b1BaeTCJI
HapymeHlIeM npo:o;eCCOB CBJl3bIBaHBJI RaTeXOJlaMBHOB, He BJleqeT 3a
C060H op,HoBpeMeHHoro BOCCTaHOBJleHIIJI ropMOHOB (Holzhauer, Vogt..
1956; Eranko, Hopsu, 1958, 1961; Callingham, Mann, 1958..
1962; Klein, Kracht, 1958; Coupland, 1959; Yates, 1963; Cle-
menti, Zocche, 1963, H p,p.), IIMeIOIII;eI'O MeCTO IIpII IIHCYJlIIHOBOM
mORe. B pe3YJlbTaTe xpoMacl>cl>BHHble RJleTRII HeROTopoe BpeMJI IIOCJI&
BBep,eHHH pe3eplIBHa He 06HapymHBaIOT IIplI3HaROB ceRpeTopHOH aR-
THBHOCTII. HMeHHO 1I03TOMY, OqeBIl)J.HO, Ha 3J1eRTpOHHorpaMMax, 110-
JIY'leHHblX nocJle pe3eplIIIHII3a:O;BB mBBOTHblX, HeT HlIRaRBX CTPYR-
TYIJ, ROTopble xapaRTeplI30BaJiB 6bl Bblp,eJleHlIe .ropMOHOB, Ha6J1IO]:(a-
eMoe nocJle BBep,eHHH BHCYJlIIHa. ROMIIJleRC fOJlbp,mB B 60Jlbm1IH-
CTBe RJleTOR Op,HOTHlIHO paCIIOJlaraeTCJl B BII]:(e ROJlnaqRa Hap, Jlp,pOM,
a cy6MBRpOCROIIlf'IeCRB OH npep,CTaBJleH Be3IIRYJlaMB, :O;IICTepHaMII II
RaHaJlb:o;aMII B6J1II3II HApa. llocJle nepBoro BBep,eHIIJI pe3eplIIIHa Ha-
6J1IOp,aeTCJI pa3pemeHIIe 9J1eRTpOHHOIIJlOTHOrO Cop,epmHMoro rpaUYJl
(pIIC. 106) ; 1I0CJle BToporo - YMeHbmeHlIe BX qlICJla. Ha MOOTe
BCqe3aIOIII;BX rpaHYJI qaCTO OCTaIOTCJI IIYCTble 9J1eRTpOHHOlIp03paq-
Hble IIY3b1pbRH, CBll)J.eTeJlbCTBYIOIII;IIe 06 IICTOIII;eHBB ropMOHOB (rlIc-
TOXBMlf'IeCRBe peaR:O;IIB Ha RaTeXOJlaMBHbl B 3TOT neplIop, p,aIOT OT-
+-
P He. 104. XpoMacIJilJ.uHHaH RJIeTRa Hap;nOqeqHJfRa RpbIcbI. TIPOP;YRTbi peClm-
Te3a KaTeXOJIaMHHOB B Be3HRYJIaX 30HId rOJILp;llil!
xT - ROMlIJIeRC 1'0JIhAJRU; ce. e.- CBeTJIh/e IIJIeRTpOHHonp03pa'IHhle Be3MRYJlh/; nA. ._
8:'IeRl'pOHHonJIOTHble HOBoor!pa30BaHHhle BC3HRYJIh/; .At'" - M1'ITOXOHJlPJIH; .II - H):IpO (npuA-
lIIaR. WIT. no CMHTTeH, 1965)
281
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1I0m]:(aeTCJI ncpecTpoiiIiOii 1\Il'1\I6paHlIblx CIICTCIII H:lCTKII, II B TOM qllC.Jle
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B «CTepTOM» Bllp,e, 110 B IIPIlHQlllIl' xapaKTepll3yroT CIIHTeTHqeCKlle
npOQceehl B QIITOn.Jla3}1l' (pIlC. 107). HoeJIe pl'.!CpmnIU3aQlln ,KlmOT-'
UblX UII Ila O]:(HOIll 113 TaIlOH nl' 1I0JIB:JHIOTeJI CTPYI{Typbl, onncalI-
UbIe II ;JliCnl'plillICIITax C lIBC]:(l'lIl1l'lII IIl1eYJIlIIIa II xapalnepU:JYIOIl{IIC
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I{POCKOIIII'ICCI{3JI MCM6p3Ha, IWTOpaJl, 110 conpeMellHbIM n033peHlIJIM,
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282
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.JIJlIOr:u;eH CeKpeTOpHYIO ee ]:(eJlTeJlbHOCTb (CM. Douglas, 1966;
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HaJlbHbIX IIpO]:(YKTOB B 06blqHblX YCJlOBIIHX mII3He]:(eJlTeJlbHOCTII KJle-
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KY OHn ocyr:u;eCTBJlJlIOTCJI HepBHblM rryTeM, JlBJlJlIOTCJI, OqeBII]:{HO,
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HblM aKTOM, B03MomHblM TOJlbKO IIpII COOTBeTCTBYIOr:u;eM BblCOKOM
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eilCTBlle pe3epnllHa H llHCYJIHHa IIpep,CTaBJIHeT llHTepec B ABYX OT-
HOllleHHHX: IIOqeMY B OAHOM cJIY'Iae (pe3epIIllH) HJIeTHa OTBeqaeT
yme B paHHlle CpORll 3apOAblllleBoro pa3BllTHH, B P,PyroM (HHCY-
JIllH) - TOJIbHO B 1I03AHUe CpORH, II nOqeMY B nepBOM CJIyqae IIMe-
eTCH lIepIlOA oCB060mAeHIIH HJIeTHll OT ee cop,epmHMoro, a BO BTO-
pOM CJIyqae OH He HacTynaeT? H3BeCTHO, qTO B OHToreHeTlIqeCIWM
pa3BllTllIf BblCllIlIX 1I03BOHOqHbiX nOJIHOTa rOpMOHaJIbHOH .pYHl\IJ;lllI
06VCJIOBJIeHa CTelIeHbIO 3peJIOCTII RaH nepll.pepHqeCKIIX, TaR II qeHT-
p:lJIbHblX perYJIIlpYIO llX MexaHll3MOB. B D;enlI peaRD;llH «rllIIOTaJIaMII-
TJp.C'.HaH 06JIaCTL - lIaparaHrJIIIOHapHaH HJIeTHa» 06eCneqlIBaeTCJI CIIO-
C06HOCTb opraHll3Ma K npop,yRIl.lIll, Bblp,eJIeHllIO II peD;enD;llH ropMO-
FOB. 3pCJIOCTb I\amAOrO ll3 3BeHbeB D;enH HrpaeT cy ecTBeHHYIO pOJIb
295
B peaJIll3an;HH 06m;erOpMOHaJIbHbIX perymm;Hii. HaCTOHm;aH pa60Ta
orpaHHqHBaeTCJI H3Y'IeHHeM TOJIbRO caMoro uepmpepHqeCROrO 3Be-
Ha - meJIe3HCTOH RJIeTRH. TIepecTpoiiRa MeM6paHHbiX CIICTeM B IIpO-
qecce ropMOHo06pa30BaHIIH H BbI,l];eJIeHHH, JlBJIJlIOm;aHCJI Mop.pOJIOrH-
qeCRHM BblpameHHeM ,l];HHaMHRII ceRpeTopHoro npon;ecca, npe,l];CTaB-
JIHeT C060H nepH.pepHqeCRHii MexaHII3M ceRpeTopHOH aRTHBHOCTII.
STOT MexaHH3M B a,l];peHaJIOBbiX RJIeTRaX C03peBaeT OqeHb paHO-
O,l];HOBpeMeHHO C HaCTYIIJIeHHeM UpO,l];YRTHBHoii CIIoc06HOCTH - II 06ec-
neqHBaeT HCTOm;eHHe H peCIIHTe3 ropMOHOB. Pa3HHn;a B cpORax peaR-
n;HII 3apO,l];blmeBbIX RJIeTOR Ha BBe,l];eHHe pe3epnHHa II IIHCYJIHHa 06yc-
JIOBJIeHa, BH,l];IIMO, pa3HblMH MexaHH3MaMII ,l];eiiCTBIIH 3TIIX Bem;eCTB
Ha opraHH3M II 3aBHCHT OT n;eHTpaJIbHbIX perYJIHn;Hll. L(eHTpaJIbHOe
,l];eHCTBHe HHCYJIHHa (West, 1955, 1961; JIeii6coH, 1962) BJIeqeT 3a
c060ii HapymeHHe yrJIeBO,l];Horo 06MeHa, B TO BpeMJI RaR pe3epUHll
HapYlliaeT npon;ecc CBH3bIBaHlIH RaTeXOJIaMHHOB. S.p.peRT HX ]:(ellCT-
BIIJI Ha 3M6pHOHaJIbHYlO RJIeTRY 3aBIICHT OT CpOROB C03peBaHHH Tex
IIYTeii, 110 ROTOpblM ocym;eCTBJIHIOTCJI HX BJIHHHHH. KaR nORa3bIBalOT
rlIcrOXHMHqeClme peaRIJ;IIU (CMHTTeH, 1967) If ROJIH'leCTBeHHble
onpe,l];eJIeHHH (CTa6poBcRHii, 1964, 1965), 1I0C.:.Ie ,l];eiiCTBIIH HHCY-
JIHHa B Ha,l];nOqeqHHRaX RypHHbIX 3apO,l];bImeii BCer,l];a OCTaeTCH RaRoe-
TO ROJIH'leCTBO ceRpeTa, qTO 06'bHCHHeTCH CJIe IOm;IIM 06pa30M. ECJIH
,l];0 14-15 CYTOR pa3BHTIIJI RYPIIHbIX 3apO,l];bIllieH a,l];peHaJIOBaH CH-
CTeMa HIIRaR He pearHpyeT Ha HHCYJIHHOBYIO rHnOrJIHReMHIO, TO,
OqeBII,l];HO, B 60JIee U03,l];HHe CpORH 3Ta peaRn;HH HapaCTaeT Me,l];JIeH-
HO, II pa3BIIBalOm;HHCH opraHH3M He B COCTOHHHH peaJIH30BaTb Bce
lJanaCbl ropMOHOB.
Pe3YJIbTaTbI H3yqeHIIH ceRpeTopHblX D;IIRJIOB II 3HaROMCTBO C JIH-
TepaTypoii 0 ropMOHo06pa30BaHHH H HaROnJIeHHH ceRpeTa B XpO-
Ma.p.pHHHbIX RJIeTRaX Ha,l];IIOqeqHHROB n03BOJIJlIOT HaM BblCRa3aTb He-
ROTOpbIe co06pameHHH 110 3TOMY 1I0BO,l];Y.
- BonpOCbl .pYHRQHOHaJIbHOH Mop.pOJIOrIIH xpOMa.p.pHHHbIX RJIeTOI{
CBH3ilHbi C np06JIeMOii MOHO-H ,l];IIMop.p1l3Ma RJIeTOqHbIX .pOPM, IIC-
CJIe,l];OBaHHe ROTOpoii B HaCTOHm;ee BpeMH npHHHJIO mHpORHH pa3-
Max. 06pa3YIOTCH JIll ,l];Ba ropMOHa M03rOBoro CJIOH Ha,l];UOqeqHH-
ROB - a]:(peHaJIHH H HOpa,l];peHaJIHH B ,l];BYX THuax R':IeTOR, HJIU Mem-
,l];Y HUMH cym;eCTBYIOT B3aHMHbie nepeXO,l];bI - 3TO TOT Bonpoc, RO-
TOpbIH ,l];OJImeH HHTepeCOBaTb Mop.pOJIOrOB.
HeO,l];HopO,l];HOCTb xpOMa.p.pHHHbIX RJIeTOR Ha,l];nOqeqHHROB OTMeqa-
JIaCb H B IIpemHIIX pa60Tax, OTHOCHm;I1XCH R nepBOll nOJIOBHHe Te-
Rym;ero CTOJIeTHH (HaupHMep, Bennett, 1941), B ROTOpblX yRa3blBa-
JIOCb Ha pa3JIH'IHYIO IfHTeHCHBHOCTb ORpaCRH IJ;HTOnJIa3Mbl, Ha UPH-
CYTCTBHe «TeMHbIX» II «CBeTJIbIX» }meTOR, Ha ORCH.pHJII>HYro
I[ 6a30.pHJIbHYIO peal{n;1I1O 3epeH. B Te rO,l];bl Mop.pOJIOrHqeCROe pa3-
Ho06pa3He xpoMa.p.pHHHbIX RJIeTOI( paCn;eHHBaJIOCb He RaR cym;ecT-
BOBaHUe R.TIeTOqHbIX THIIOB sui generis, a RaR CTPYhTypHoe Bblpa-
meHHe pa3Hbix CTa,l];HH ceRpen;HH.
296
C 50-x rO.o;OB HaqlIHaeTCJI HOBW:D: :qHRJI IICCJIe.o;OBaHII:D:, IIOCBJlIqeU-
HblX Bonpocy COCTaBa RJIeTOR M03rOBOrO CJIOH Ha.o;IIOqeqHHROB, HO'
TOJIROBaHlIe .pH3HOJIOrHqeCROrO 3HaqeHIIJI pa3HblX RJIeTOR npHo6peTa-
eT HHOH xapaRTep. Ha1JaJIOM 3TOrO HRJIa MOmHO CqHTaTb nOHBJIe-
HHe B ne1JaTH pa60T Bander (1950, 1954) H Eranko (1951, 1955a,
b) . Bander onlIcaJI B Ha.o;nO'leqHHRaX Mblmen TeMHble .pYRCHHO-
.pHJIbHble RJIeTRII (F-RJIeTRH) H CBeTJIble, IIHRpHHO.pHJIbHble (P-RJIeT-
RU), OTJIHqalOIqHeCH .o;pyr OT .o;pyra H no BeJIH1JHHe. <DIIRCIIPYJl
. Ha.o;nOqeqHHRH Mblmen B pa3Hoe BpeMH CYTOR, Bander YCTaHOBHJI
ROJIe6aHHJI qHCJIeHHOCTH Tex II ,l];pyrHx THnOB RJIeTOR B HOqHOe H
.o;HeBHoe BpeMH: .o;HeM npeo6JIa.o;aJIH F-RJIeTRII, HO%IO ROJIH1JeCTBO
Tex II .o;pyrHx, oco6eHHo P-RJIeTOR, B03paCTaJIO. Bander paccMaTplI-
BaeT P-RJIeTRll B RaqeCTBe cne H.pHqeCRHX a.o;peHa IIH06pa3YlOlQllX,
a F-RJIeTRH npe]:(nOJIO>KllTeJIbHO CqllTaeT Hopa.o;peHaJIHH06pa3YIO-
IqHMII. ABa THna RJIeTOR OH II03me OnHCaJI y co6aR, RomeR, Rpo-.
JIHROB, MOpCRHX CBHHOR II RpblC. CBOII 3aKJIIOqeHIIJI aBTOp OCHO-
BblBaeT Ha COIIOCTaBJIeHIIH ROJIHq-eCTBa Toro IIJIII .o;pyroro aMHHa-
B Ha.o;IlOqeqHIIRaX mHBOTHoro C IIpO eHTOM o6Iqen nJIoIqa,l];II ceqe--
HHH M03rOBon qaCTII Ha.o;IIOqeqHII'ROB, 3aHHMaeMon P- II F-RJIeT-'
RaMII.
TIoCJIe.o;YIOIqHe, OqeHb MiHOrOqlICJIeHHble HCCJIe.o;OBaHHJI CBonCTB;
xpoMa.p.pIIHHblx KJIeTOR xapaRTepH3YlOTCJI pa3pa6oTRon cne IIaJIb-
HblX rlICTOXIIMlIqeCIrnX MeTo.o;OB Ha CBeTOBOM II 3JIeRTpOHHOMHRpO-.
CROIIH1JeCROM ypOBHJIX, II03BOJIHBmHX pa3.o;eJIbHO BbIHBJIHTb RJIeTRH,_
co.o;epmaIqHe a.o;peHaJIHH II Hopa.o;penaJIIIH (Hillarp, Hokfelt, 1953,
1955; Eranko, 1955a, b, C, 1956, 1957; Eranko, Palkama, 1959;.
Wood, 1962, 1963; Coupland et aI., 1964; Coupland, Hopwood,
1966; Tramezzani et aI., 1964). Hillarp H Hokfelt, a II03iRe Klein
H Kracht (1958) IIpe.o;IIOJIO>KIIJIII, 1JTO Bce RJIeTRII M03rOBoro CJIOJl.
Ha]:(IlOqeqHIIRa o6pa3YlOT Hopa.o;peHaJIHH, HO H3BeCTHbln IIpo eHT H3-
HllX - onpe.o;eJIeHHbln .o;JIJI Ram.o;oro BII.o;a mHBOTHbIX - cIIoco6eH Me-
TIIJIHpOBaTb .ero C o6pa30BaHHeM a.o;peHaJIHHa. KJIeTRH, cnoco6Hble K
MeTIIJIJI HII, HaRanJIHBalOT 3.o;peHanHH, He CIIOc06HbIe R MeTH HIJ;IIII --
Hopa.o;peHaJIIIH.
AJIH aHRJIH3a paClIpe.o;eJIeHHJI a.o;peHaJIHHa H Hopa.o;peH8JIII-
Ha IICnOJIb30BaJIlICb ,mcnepHMeHTaJIbHble MeTO]:(bl. "Y MblllleH co.o;ep-
>KaHne 060HX aMHHOB CHnmaJIOCb IIOCJIe O;:J;HORpaTHoro BBe;:J;eHHH
pe3epnHHa, y RpblC TOTaJIbHaH y6bIJIb a;:J;peHaJIHHa H Hopa.o;peHaJIHHa.
Ha6JIIO.o;aJIaCb nOCJIe IIpO]:(OJImnTeJIbHOn o6pa6oTRH pe3epnHHOM (Era-
nko, Hopsu, 1958, 1961). Ha Ha.o;nOqe'lHHRaX RpblC pe3YJIhTaTbl
ROJIHqeCTBeHHblX XllMHqeCRHX onpe]:(eJIeHIIH a.o;peHaJIHHa II Hopa.o;pe--
HaJIHHa conOCTaBJIHJIHCb C nORa3aTeJIHMH HTo.poToMeTpHH rHCTO-.
XHMHqeCRllX peaR HH - xpoMa.p.pHHHoii, llo.o;aTHoH II .popMaJIHH.pJIYO-
peC eHTHOH (Eranko, Raisanen, 1961; Eranko, Hopsu, 1961).
H3MepeHHH, npOBe.o;eHHbIe B HOpMaJIbHblX YCJIOBHHX H lIocae HCTO-
IqeHHH RaTeXOJIaMHHOB pe3epnHHOM, nORa3aJIH, qTO IIHTeHCHBHOC'fb.
xpoMa.p.pHHHon peaR HH MeHHeTCJI COOTBeTCTBeHHO II3MeHeHlllO CYM-
MapHoro co.o;epmaHnH RaTeXOJIaMHHOB, a HHTeHCHBHOCTb lIo.o;aTHoH
29
II «pJIyopec eHTHOii peaR Hii CBH3aHa C IIORa3aTeJIJlMH ROH eHTpa nH
"HOpa,lJ;peHaJIHHa.
TaRHM 06pa30M, ROJIHQeCTBelIHble H3MepCHlIH 110,lJ;TBep HJIII cne-
H«pnqHOCTb rHCTOXHMHqeCRIIX peaR Hii, a, CJIe OBaTeJIbHO, npHCYT-
CTBHe B M03rOBOM CJIoe Ha]:(nOqeqHHROB ,lJ;BYX THIIOB RJIeTOR, O,lJ;HH H3
ROTOpblX CO,lJ;epmaT Hopa]:(peHaJIHH, B ]:(pyrHx npHCYTCTByeT rpynna Ra-
TeXOJIaMHHOBblX coe,lJ;HHeHHii, CYMMapHO RblHBJIHeMbiX xpOMa«p«pIIHHoii
peaR Heii.
COOTBeTCTBeHHO rHCTOXHMHqeCRIIM peaR IIHM, BbIHBJIHIO JlM a,lJ;-
peHaJIHH II Hopa,lJ;peHaJIHH, B RJIeTRaX 1I0-pa3HOMY pacnpe enHIOTCH
«pepMeHTbI. OCTPOBRH RJIeTOR, 06JIa,lJ;aIOIqHe BblpameHHoii «pJIyopec-
n;eH IIeM H xpOMaTOrpa«pHqeCRIIM MeTO,lJ;OM 06HapymHBaIOIqHe BbICO-
RlIii ypOBeHb CO,lJ;epmaHHH HOpa,lJ;peHaJIHHa, 110 ]:(aHHblM Eranko,
{1951, 1956), He cO,lJ;epmaT RHCJIOM «poc«paTa3bl, HO OTJIHqaIOTCJI BbICO-
Ron aRTHBHOCTbIO Hecne H«pJlQeCROii XOJIHH3CTepa3b1 (Eranko,
Hopsu, 1958). TIplI 3RCnepJlMeHTaJIbHOii rHnepnJIa3J1lI MOarOBOrO
CJIOH Ha,lJ;nOqe'llIHROB, Bbl3BaHHoii npO,lJ;OJImHTeJIbHbIM BBe,lJ;eHIIeIlI Hn-
ROTHH8 (Eranko, 1955), YBeJIHqHBaeTCH pa3Mep «pJIyopec HpYIOlQHX
OCTpOBROB H COOTBeTCTBeHHO paCmJlpHeTCH 30Ha RneTOR, HeraTHB-
HblX B OTHomeHHII KHCJIOH «poc«paTa3b1, HO 06JIa,lJ;aIOIqIIX BblCOROH
ROH eHTpa IIeH Hecne H«plIqeCROH XOJIHH3CTepa3b1 (Eranko et aI.,
1959a, b). BnpOqeM IIpJl o eHRe 3HaqeHHH «pepMeHTHblx CJlCTeM B
pa3Hblx RJIeTOqHblX rpYlIlIax M03fOBoro cnOH Ha,lJ;IIOqeqHIIKOB Ha]:(o
IIMeTb B BH,lJ;Y, qTO ,lJ;aHHble pa3JIHQHblX aBTopOB 0 JIORaJIII3a HH
4>epMeHToB, B qaCTHOCTH RHCJIOH «poc«paTa3bl, He COBlIa,lJ;aIOT II npo-
"THBOpeqaT ,lJ;pyr ,lJ;pyry. TaR, no Picard, Vitry (1959), Hopa pe-
HaJIHHOBble RJIeTKll cO,lJ;epmaT RHCJIYIO «poc«paTa3Y.
B pa60Tax 3. C. Ka HenbcoHa H COTPY,lJ;HHROB ero JIa60paToplili
RneTO'llIbIH COCTaB M03rOBoro CJIOH Ha,lJ;nOqeqHHROB HCCJIe,lJ;OBaH y
JRllBOTHblX pa3Hblx RJIaCCOB, OTpHJJ;OB II ceMeiicTB. ]];HMOp«PH3M RJIeTOK
(a,lJ;peHanHH-H HOpa,lJ;peHaJIHHCO,lJ;epmaIqHx) YCTaHOBneH aBTopaMH Ha
'OC.HOBaHHU rHCTOXHMHQeCRUX, TUHI>:TOpUanblIblX, Mop«ponOrUqeCRUX II
"Tonorpa«pUqeCRHX oc06eHHOCTen.
TIpHBe,lJ;eHHbIH lIepeqeHb pa60T U MHomeCTBO ,lJ;pyrlIx He IIepe-
""1IHCneHHbIX HaMH, a TaIDRe c06CTBeHHble ,lJ;aHHbIe, 1I0nyqeHHbIe Ha
npe,lJ;CTaBHTeJIHX pa3HblX RJIaCCOB n03BOHOqHblX (aM«pu6uHX, penTU-
.JIHHX, IITH aX II MJIeROIIHTaIOlQHX), ,lJ;aIOT OCHOBaHHe fOBOpUTb 0 cy-
eCTBOBaHUH RJIeTOR, O,lJ;HH U3 ROTOpblX B 06blqHbiX YCJIOBIIHX CO-
epmaT a,lJ;peHanIIH, ]:(pyrue HOpa,lJ;peHaJIIIH. JIHTepaTypHble ,lJ;aHHble
II c06CTBeHHblH OlIblT Ha MO,lJ;eJIHX C UHCyJIHHOBblM 1lI0KOM, U3JIO-
-meHHble Bblme, IIORa3bIBaIOT He TonbRO pa3,lJ;eJIbHYIO JIORaJIU3a UIO
'Toro II ,lJ;pyroro aMUHa B RJIeTKaX M03rOBOI'O CJIOH Ha,lJ;nOqeqHUROB,
HO npu HeROTopblX YCJIOBHHX H pa3,lJ;eJIbHOe HX Bbl,lJ;eJIeHHe.
HapH,lJ;y C HeCOMHeHHblM «paKTOM HaJIlIqHJI KJIeTOR, ROTopble, KaR
IIpaBllJIO, CO,lJ;epmaT a,lJ;peHaJIHH II Hopa,lJ;peHaJIHH H I1MeIOT onpe,lJ;e-
.JIeHHYIO 30HaJIbHYIO nORaJIH3a llIO, OCTaIOTCJI HeHCHbiMII MHorne Bon-
pOCbI, Tpe6YIOIqHe ,lJ;anbHeiimHx nOHCROB. flBJIHeTCJI cnopHblM, McmHO
.J1H CqHTaTb Ram,lJ;YIO II3 3THX ,lJ;BYX KJIeTOqHblX rpYlIn caMOCTOJl-
:298
eJIbHblM THOOM, HJIIi OHM MOrYT nepexop:HTb O;:J;MH B P:Pyroii B 3a-
illUCUMOCTII OT YCJIOBuii BHYTpeHHUX (06MeHHbIe peaRIJ;uu), BHelllHUX
.( ce30HHOCTb, 3RcnepUMeHTaJIbHbIe B03p:eiicTBUH), IIJIU UX CJIep:yeT
C'IUTaTb CTa]:(HJlMII CMHTe3a ropMOHOB B ep:UHOM RJIeTOqHOM Tuoe.
He Y Bcex BHp:OB qeTRO pa3JIUqUMbl ]:(Ba Tuna RJIeTOR: Y HeROTO-
'pblX UMeeTCH TOJIbRO Op:UH TUO (JIHrymRa) . OTMeqeHbl ce30HHble
II3MeHeHUH, a TaRme ROJIe6aHUH ROJIUqeCTBeHHblX COOTHOllieHuii ap:-
'peHaJIUH-H Hopap:peHaJIUHcop:epmalI{ux RJIeTOR npu U3MeHeHIIII «PU-
-3UOJIorUqeCROrO cTaTyca opraHU3Ma, HanpUMep, B lIepuop: 6epeMeH-
:HOCTH (RaD;HeJIbCOH, 1967a). B 3RCliepUMeHTe (OOCJIe BBep:eHUH HH-
-CYJIUHa IIJIU pe3eplIUHa) RJIeTRU npu06peTalOT CXOP:HYIO CTpyRTYPY 11
BeP:YT ce6J1 op:IIHaROBO B CMblCJIe cnoc06HOCTU MeTUJIUpOBaTb Hopap:-
;peHaJIIIH B ap:peHaJIIIH (Picard, Vitry, 1964). BBep:eHue HHCYJIHHa
B p:03ax, 06eCneqlIBalOlQUX u36upaTeJIbHOe Bblp:eJIeHHe TOJIbRO ap:peHa-
.JIUHa (IIpuiiMaR, 1965), OOYCTomalOTCH 3JIeRTpOHHOnJIOTHbie RaTe-
XOJIaMllHcop:epmalI{IIe rpaHYJIbl RaR u3 «TeMHblX» RJIeTOR, ROTOpblC
paCCMaTpUBalOTCJI B HaCTOHlI{ee BpeMH B Ra'leCTBe HOCUTeJIeii Hopap:-
peHaJIHHa, TaR II u3 «CBeTJIblX», CqIlTalOlI.\UXCH ap:peHaJIUHcop:epma-
ID;UMH (puc. 117), U qTO, CJIep:OBaTeJIbHO, yRa3aHHaH yJIbpacTpyR-
-rypHaH xapaRTepUCTURa He COOTBeTcTByeT IIplIHHTOMy MHeHUIO ()-
.ropMOHaJIbHOM cop:epmlIMOM 3TIIX RJIeTOR.
Halliu p:aHHbIe, 1I0JIyqeHHbIe opu u3yqeHIIII ceRpeTopHblX D;IIRJIOB
:xpoMa«p«pMHHblX RJIeTOR B 3JIeRTpOHHOM MHRpocRoIIe, n03BOJIHIOT BbI-
.CRa3aTb RplITUqeCRlIe 3aMeqaHUJI II 110 HeRoTopblM p:pyruM BOIIpO-
IcaM, 06cymp:aeMblM B JIIITeparype B CBH3II C IIp06JIeMOii RJIeTO'IHoro
P:HMop«p1I3Ma.
ECTeCTBeHHo, qTO JIORaJIIICTUqeCRaJl TeHp:eHD;IIJI, IIpOHBUBmaHCJI
D Mop«pOJIOrlI'IeCRIIX uCCJIep:oBaHIIHx II HaOpaBJIeHHaH Ha IIHTeHCUB-
Eblll nOIICR MeCT pa3p:eJIbHOrO 06pa30BaHuJI RaTeXOJIaMIIHOBblX coe-
JJ;uHeHlIii - ap:peHaJIIIHa II Hopap:peHaJIUHa, OTpa3UJIaCb U Ha 3JIeRT-
-P0HHOMURpOCROnlIqeCRIIX IICCJIep:oBaHuHx. ¥me B lIepBblX me paOOTax
aBTopbl IIblTaJIIICb HaHTII pa3HuD;y B TOHROH cTpyRType xpoMa«p«puH-
:HblX RJIeTOR, yCTaHoBIITb Ha 3TOM OCHOBaHlI1I cy6MuRpOCROnlIqeCRUe
.oTJIllqUH Memp:y ]:(BYMJI TUlIaMU RJIeTOR.
C nepBblx me maroB 3JIeRTpOHHOMURpOCROlIII'IeCROrO u3yqeHlIJI
naparaHrJIUeB 6bIJIO ROHcTaTupoBaHo, 'ITO ropMoHbl cop:epmaTcH B
D;UTOOJIa3Me RJIeTOR B Bup:e 3JIeRTpOHHoHeOpOapa'IHblX rpaHYJIHpHO
.o«popMJIeHHbIX CTPYRTYp. Cpe]:(u MHOrOqnCJIeHHblX p:oRa3aTeJIbCTB B
nOJIb3Y 3Toro YTBepmp:eHuH cJIymuJIa 3RcoepUMeHTaJIbHaH CTIIMYJIH--
I(IIJI Bblp:eJIIITeJIbHOH «PYHRD;1I11 M03rOBoro CJIOH Hap:nOqeqmiROB, co-
npoBomp:aBmaHcH op:HoBpeMeHHblM Bbl6pacbIBaHueM B RpoBeHoCHoe
. 'pyc."'lO rOpMOHaJIbHOrO cop:epmuMoro RJIeTOR U oCB060mp:eHlleM HX
.IJ;IITon:Ia3Mbl OT rpaHYJIHpHblX CTPYRTYp. CTpYRTYpa 3TIIX rpaHYJI
r."'laBHbIM o6pa30M II 06cymp:aJIaCb IICCJIep:oBaTeJIHMII.
I yJIbTpacTpYRTYPHblM cBollcTBaM RaTeXOJIaMIIHOBblX rpaHYJI OT-
110CIITCH IIX Be."'lllqUHa, «popMa, BHYTplirpaHyJIHpHOe CTpOeHJle II Ha-
.JIllqlle HapymHoii MeM6paHbl. 8TII qepTbi CTpOeHlIJI CJIymlIJIU UCCJIe-
,p;OB3Te.1IHM B K3qeCTBe rJIaBHblX RpUTepHeB p:.1IH cymp;eHHJI 0 TOM,
299
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P If c. 117. Tl'MHhIe (lnl» II CIIl'TJlhII' (CI» X pOJ\laiP II1I1lIhlC KJII'TKIi 1I<1P;1I0'IC'IHlIRa
RpblCbI. IIHCYJIllllolIJ.\ii mOK
Clt - COJ\epmaT I'paHYJIbI; mlt - onYCTorneHbI (lIpHAMaH. 19(5)
BO BCeX Jill IIaparaHrJlIIOHapHblX KJIeTRaX Hap;IIOqeqHIIROB rpa:ayJll.I
op;HGTllnHbI 110 CTpYRType, IIJlII llX MomHO RaR-TO RJlaccmplIqupo-
RaTb.
3aCJlymlIBalOT RplITlIqeCROrO paCCMOTpemIJI, C HallieH TOqRll 3ptJ-
HT<m, Te pa60TbI, ROTOpbIe B OCHOBY p;llIpcpepeHn;llaJlbHOrO IIop;xop;a
K & eTOqHblM TllIIaM IIaparaHrJlIIOHapHOH TRaHII RJlap;YT IIJlOTHOCTb II
BeJIUqIIHY ceRpeTopHblX rpaHYJI. XapaRTepHO, qTO cpep;II aBTopOB, OIIII-
palO nXCH B CBOUX orrIIcallIIHX Ha IIJIOTHOCTb rpaHYJlHpHOrO Co;n;ep-
mliMoro RJIeTOR Kalt Ha rJlaBHblH CTpYRTypHblH IIpn3HaK ;n;IIcpcpepeH-
qnpoBRII, HeROTOpbIe npe;n;JlaralOT RaRYIO-JlIIOO CBOIO MO;n;IIcplIRaqmo
MeTOp;a cpnRCaU;IIn Thallll (Wood, Barruett, 1964: Coupland, Hop-
wood, 1966; 1964). OHn IIORa3aJlII, qTO IIpn npOBep;eHIIn npoU;eP;ypbl
.pnRCaU;IIII TRaHII COqeTaHlie rJIYTapaJlbp;erIIp;a C 6IIXpOMaToM (Wood,
Barrnelt) IIJIn 3a6ycpepeIIHoro rJIYTapaJIbp;erIIp;a C oCMneM, 6IIXpOMa-
TOM KaJIIIH, MOJIIl6p;eHOBOIiIICJIblM HaTplIeM II IIO;n;aTOM RaJIIIJI (Coup-
land, Hopwood, 1966) p;aeT B03MOmHOCTb YCTaHOBllTb qeTRylO pa3-
IIIID;Y B CTeneHII jJIeRTpOHHOH HeIIp03paqHOCTn RaTeXOJIaMIIHOBbIX
rpaHYJI. B aTIIX YCJIOBnHX IIJlOTHOCTb rpaHYJI Hopap;peHaJllmCOp;epma-
m;nx KJIeTOR 3HaqlITeJIbHO Bblllle a;n;peHaJIllHCOp;epmar:u;lIx. B cop;ep-
maTeJIbHOH CTaTbe Elfvin (1965), B ROTOpOH I'JIaBHOe BmmaHlIe
y;n;e.JIeHO IICCJIep;OBaHIIIO YJIbTpaCTpYRTypbl IilleTOqHblX IIoBepxHocTeH
n cneqlIaJIn3au;nn IIX B HanpaBJIeHlI1I ceRpeTopHoH aRTIIBHOCTII RJIe-
TOR, 06par:u;eHO BHllMaHne II Ha CTPYRTYPY qnTOnJIa3MaTlllIeCRIIX
rpaHYJI, HaIIOJIHeHHblX RaTeXOJIaMIiBaMII. B OP;HIIX RJIeTRaX Co;n;ep-
mlIMoe rpa:ayJI IIMeeT pa3HYlO CTeIIeHb pa3pemeHHOCTII (1-H TIIn) ,
B p;pyrllX O;IJ;HOpO;IJ;HbIH HeIIpoHIIqaeMblH ;n;JUI aJIeRTpOHOB MaTepnaJI
HanOJIHHeT 06bIqHO He Bce B:ayTpeHee npOCTpaHCTBO rpa:ayJIbI, a Ra-
RYIO-TO ero qaCTb, TeCHO ROHTaRTlIpYJI IIPII aToM C RaRHM-JIn60 yqa-
CTKOM rpaHYJIJlpHOH IIOBepXHOCTIi. OCTaJIbHOe IIpOCTpaHCTBO rpaHYJIbI
MomeT 6blTb 3aIIOJIHeHO HemH03epHIICTblM IIJIOTHblM MaTeplIaJIOM
(2-H TIIII).
PyraJl rpYIIna IICCJIep;OBaTeJIeH B RaqeCTBe RplITeplIJI pa3HOTlIII-
HOCTH IilleTOR IIaparaHrJInOHapHOH TRaHn Bbl;n;BIIraeT He CTeIIeHb
TIJIOTHOCTII rpaHYJI, a llX pa3Mep (Yates et aI., 1962; Wasserman,
Tramezzani, 1963; Benedeczky, Pup pi, Tigyi, 1965; Benedeczky
ctal.,1965). .
IIoMnMo MaJIOH p;OCTOBepHOCTII aTYlX pa60T n3-3a HII3ROro TeXHII-
qenwro ypOBIIJI HX BbIlIOJIHeHnH, oQoo eHllH, ROTOpbIe ;n;eJIa-
IOTCH B HeROTopblX II3 HllX, Bbl3bIBalOT B03pameHIIJI. ABTOpbI
lIepBOH pa60Tbl B aTOM HarrpaBJIeHHll (Yates et aI., 1962) ncxo-
p;nJIII n3 3RcneplIMeUTaJIbHblX 6nOXnMIIqeCRnX nCCJIe;n;OBaHnH, ROTOpbIe
nORa3aJIII, qTO B pa3HblX cppaRlJ;IIHx n;oHTpmpyraTa M03rOBoro CJIOJl
Eap;rrOqeqmlliOB rrpeBaJlIlpyeT JlIr60 ap;peUaJInH, JIn60 Hopap;peHaJlllH
(SchUmann, 1957; Eade, 1958; Hagen, Barrnett, 1960). SchUmann
(1957) n3yqaJI pacnpep;eJIeHne aMnHOB B rOMoreHaTax, IIOJIyqeHHblX
H3 Ha]:(nOqeqHnROB p;OMalllHnx nTIIU;. OH Bblp;eJIJlJI cppaRqlI10 Hll3Roro
yp;eJIbHOI'O Beca, 60raTYIO a;n;peHaJIIiHOM, II cppaRqlI10 BblCOROro y;n;eJIh-
.Horo Beca, co;n;epma ylO IIpenMyID;eCTBeHHO nopa;n;peHaJIIIH. Eade
301
(1958) U II03me Hagen, Barrnett (1960), pa60TaH Ha rOMoreHaTax
Hap;IIOqeqHIIROB 6bIROB, TalOl\e IIOJIyqUJIII HalI60JIee JIerRUe cppaRquu..
HacblrqeHHbIe a]:(peHaJIUHOM, U HalI60JIee THmeJIbIe - Hopap;peHaJIIIHoM.
TaRaJl 3aROHOMepHOCTb p;aJIa Yates C coaBTopaMII (1962) OCHOBalme
IIpep;IIOJIaraTb, qTO pa3Mep rpanyJI B xpoMaCPCPIIHHblX MeTRax ltlOmeT
CJIymUTb OP;HIIM 1I3 IIpU3HaROB MopcpOJIOrlI'IeCROrO pa3JIIIqlIH Memp;y
P;ByMH TUlIaMU RJIeTOR.
TaRuM IIyTeM B OIIUCaTeJIbHOll aJIeRTpOHHOMURPOCROIIUqeCROH JIU-
TepaType IIOJlBUJIOCb MHeHlIe, qTO cop;epmalIJ;ueCJI B IIaparaHrJIIIO-
HapHblX KJIeTRaX 'I'OpMOHbl UMelOT pa3Hoe CTpYRTypHoe ocpopMJIeHue..
a UMeHHO, qTO OIITUqeCRU IIJIOTHbIe RalIJIU Hopap;peHaJIlIlIa, orpa-
HUqeHHbIe OP;HOROHrypHOll rJIap;ROll MeM6paHoll, IIMelOT 60JIbllllIH p;ua-
MeTp, a RalIJIII ap;peHaJIIIHa - MeHbllllIll. iho cymp;elme 6bIJIO aRCT-
paIIOJII!JJOBaHO U3 pa60T C rOMoreHaTaMII.
Benedeczky, Puppi, Tigyi II Lissak ('1965) UCRaJIlI aRCIIepu-
MeHTaJIbHOe IIO]:(TBepmp;eHlIe CBoeH TOqRJI 3peHUJI 0 pa3Mepax rpa-
nyJI, BBOP;JI RpbIcaM U co6aRaM UHCYJIUH. ABTOpLI CIIpaBep;JIUBO IIpep;-
IIOJIal'aJIU, qTO eCJIn IIpaBUJIbHa ROHqeIIqlIJI 0 COOTBeTCTBUII pa3MepOB.
ceRpeTopHblX rpaHYJI TOMY UJIU UHOMY ropMOHaJIbHOMY cop;epmllMO-
MY, TO IIOCJIe BBep;eHUH mlIBOTHOMY UHCYJIIIHa II36upaTeJIbHOe Bblp;e-
JIeHUe Hap;IIOqeqHIIRaMII ap;peHaJIUHa p;OJImHO 6bIJIO IIpUBeCTU R U3-
6UpaTeJIbHOll aBaRyaqulI U3 RJIeTOR MeJIRUX ap;peHaJIIIHCOp;epmalIJ;ux:
rpanyJI. OHII CqIITaJIU, qTO up;eHTucplIRaqlIJI aTUX P;BYX TIIIIOB rpanyJI
UMeeT 3HaqeHUe P;JIH IIOHUMaHIIJI MexaHII3Ma pa3p;eJIbHOH ceRpequu
ap;peHaJIUHa U Hopap;peHaJIUHa. Ha OCHOBaHIIU U3MepeHUll p;uaMeT-
pOB rpaHYJI, aJIeRTpOHHOMlIRpOCROIIlI'IeCRIIX Ha6JIIOp;eHlIll U ROJIlIqe-
CTBeHHblX OlIpe]:(eJIeHIIH RaTeXOJIaMUHOB, IIpOBep;eHHbIX Ha OP;HUX I[
Tex me mUBOTHbIX, aBTopbl CqUTaJIII, qTO IIop;TBepp;UJIU CEOIO ROH-
qeIIqulO II p;ORa3aJIII, qTO ceRpeTopHble rpanyJIbI pa3Horo «PYHRqUO-
l;[aJIbHOrO Ha3HaqeHIIJI UMelOT pa3JIH"'IHYIO BeJIUquny: Hopa]:(peHaJIU-
HOBbIe RPYIIHbIe, ap;peHaJIUHOBbIe - MeJIRue. Pe3YJIbTaTbl, IIOJIyqeH-
Hble B pa60Te Benedeczky C coaBTopaMlI, pacxop;HTCH C HalliUMU
p;aHHbIMII. Halliu Ha6JIIOp;eHIIH Hap; HaROIIJIeHlIeM RaTeXOJIaMIIHOB B-
ceRpeTOpBblX rpaHYJIaX xpOMacpcplIHHbIX RJIeTOR B Hap;IIOqeqHIIRaX
RpblC IIOCJIe aRClIeplIMeHTaJIbHoro IICTorqeHIIJI UHCyJIIIHOM y6emp;alOT
B TOM, qTO lIepBbIe IIpU3HaRU HaROIIJIeHIIJI ceRpeTa 06J13aTeJIbHO Bbl-
pamalOTCJI B IIOHBJIeHUII MeJIRlIX UJIU MeJIbqaHmlIX RalIeJIb ceRpeTa.
no Mepe pa3BIITIIJI B MeTRe CIIHTeTlIqeCRIIX IIpoqeccoB II ]:(aJIbHell-
llIero HaROIIJIeHlIJI [,OpMOHOB pa3MepbI RalIeJIb YBeJIII'IUBalOl'CJI. OR-
PymeBHble rJIap;RIIMIi MeM6paHaMlI OBII paCTYT, IIpeBparqaJlCb IIO-
CTelIeHHO B 3peJIbie rpaHYJIbl ceRpeTa. MbI lIop;qepRlII!aeM, qTO BCH-
RaJl BHOBb 06pa30BaBHaJl rpaHYJIa Bcerp;a OqeHb MaJIeHbRUX pa3-
MepoB. He TOJIbRO HalllII pa60TbI, HO II peayJIbTaTbl Bcex p;pyrlIx
IICCJIep;OBaTeJIeH, IIplIMeHJlBIlIUX IIHCYJIIIH B RaqeCTBe CTHMYJIJlTOpa
Bblp;eJIIITeJIbHOH CPYHRqUII Ha]:(IIOqeqHUROB, rOBOpJlT 0 TOM, qTO IIOCJIB'
IIHCYJIUHOBOro 1lI0Ra xpOMacpcplIHHbIe RJIeTRU HalIOJIHHIOTCH MaCCOR
MeJIRIIX, IIJIOTHbIX rpanyJI, ROTopbIe TOJIbRO IIO IIpOllleCTBlI1I II3BeCT-
Horo BpeMeHII p;OCTlIraIOT pa3MepOB 3peJIblX rpaHYJI.
302
OIIIll6Ra aBTopOB MOrna OTHOCIITLCJI II K ycnOBIIHM aRcneplIMeH-
Ta. ABTOpbI lie P;OBO]:(IWII mllBOTOblX P;O 1lI0KOBOro COCTOJlHIIH, npe-
]:(OTBpa aH ero 3a6narOBpeMeHOblM BBe]:(eHlleM p;eRCTp03bI. ::ho Morn().
cy ecTBeHIfO 1I3MeHIITb pe3ynbTaTbi onbITa. 113BecToo, qTO p;ame rny-
60Roe ROMaT03Hoe COCTOHHlIe C BblpameHHblMII cyp;oporaMH II no-
Tepcn peqmeRcoB He Bcerp;a Bep;eT R a6coJIloTHO IIonHOMY IICTO e-
Hmo oap;nOqeqHIIKOB OT ap;peHanIlHa. Ecnll me 1'0BOpllTb 0 «npep;-
1lI0Re». ROTOpblii, BII]:(IIMO, liMen MeCTO B pa3611paeMoii pa60Te, TO'
TeM 60aee Henb3J1 P;YMaTb 0 MaCCOBOM Bbl6poce a]:(peHanIiHa. B nplI-
BOP;IIMbIX aneKTpOHHorpaMMax, HapMY C OqeHb MenRIIMII HOBo06pa3o-
BaHOblMII KannJlMII ceRpeTa, MHoro «CTapbIX» 3penbiX rpaHyn, OCTa-
BIlIIIXCH He Bblp;eneHHbIMII 113 KneTKII IIocne IIHcynlIHoBoro B03p;eii-
CTBnH. CqIITaTb IIX (<Hopap;peoanIlHOBbIMH» HeT HHKaKIIX OCHOBa-
HIID, TaR RaR npll I13MepeHlIHX II nOp;CqeTe npoqeHTHoro COCTaBa"
rpaHyn aBTopLI He rOBopHT, B KaKIIX yqaCTJ\ax M03roBoro cnOH Hap;-
nOqelIOnKa RpblC OHII rrpOH3BOp;I1nIlCb: p;enanOCb nIl 3TO Ha (<HOp-
a]:(peHa.TIIIHOBbIX» OCTpOBRax HnH oa OCTanbHOll: Macce xpOMacpqIHo-
ObIX R.'1eTOR - HeIl3BeCTOO.
Mbl nonaraeM, qTO CTeIIeHb nJIOTHOCTII rpaHynbl 3aBIICIIT OT CTe-
neHn ROHqeHTpaqll1I aMHHOB B rpaHyne, ROTOpaJl, cnep;OBaTenbHO,
H o6ycnoBnllBaeT ee CTPYRTypHpOBaOHOCTb. CHIImeHHe ROHn;eOTpa-
qlIn RMIIHOB CHHmaeT IInOTHOCTb co]:(epmHMoro rpaHyn II, Ha060pOT,
HOBo06pa30BaHIIe HX nOBbllllaeT ee. l1I1blMII cnOBaMII, CTeneHb aneRT-
pOHHOii IInOTOOCTII ceRpeTOpHoii rpaHynLI HBnHeTCJI cyry60 CPYHl{-
qIloHanbIIblM IIpH3HaROM H JlBnJleTCH MopcponOrIPIeCRIIM OTpameHHeM:
ceKpeTOpHoii aRTHBHOCTII mo6oii mene3HCTOii RneTRII. TI03TOMY pac-
CMaTpllBaTb CTerreHb rrnOTHOCTH TOll: HJIII HHOii ceRpeTopHoii Tpaoynbl
RaK cneqllcpnlIeCKHii npH3HaR, npIlCY HII RaKoMy-nlI60 onpe]:(eneo-
HOIllY TUllY n\ene3I1CTOii RneTRII, B IIpHHqHne He BepHo. Hallin Ha6-
:I1O]:(eOIlH HaJJ; CTPYRTypoii rpaHyn B pa3HbIe CPYHKqHoHanbHbie IIe-
PIlOJJ;bl KleTOI, - 6YP;b TO 8;J;peHaJIIIHco]:(epma HX IIJIH Hopap;peoamm-
CO:J;eplliall{lIX - He n03BOnJllOT OTMeTHTb RaRHx-nH60 MopcponOrHqeCRHX
oc06eHHOCTell:, npHCY IIX TeM 11.'111 p;pyrIlM. faMMa MopcponOrIPIeCKIIX
113I11eHeHUII, npeTeprrcBaeMbIx a]:(peoa.:rIlHoBoii II.'1H oopap;peoanHooBoii
rpRlly.:ron R nepHop; nepenOnOeHIIJI aMlIoaMH H P;O nonHoro IICTOll{e-
HIIH OT III1X, Bcer;:J;a o;:J;I1HaKoBa.
HaM npeJJ;cTaB:IHeTCJI, 'ITO nepcneKTHBHoCTb cy6MHRpOCKOllIlqe-
CROro I13yqeHlIH XpO Ia.pcpliOObIX KneTOK 3aKnlOqaeTCH He B rrOIICKax
lOpcpo.10rll'H'CI-\IIX pa3,llPHlU Mf'>I, y a;:J;peoanHO- 11 oopa]:(peoanHHCo-
;:J;eplliall{I1MII K.leTI.aMII, a B RblHCHCHIIII nOKan1l3aqHII B RneTKe 6IIO-
CIIOTe3a KaTeXO:IallIIlHOB 11 MopcpOJIOrnqeCKOrO rrpOHBneHIIH aToro npo-
u.ceCR. COBOKYIIHOCTb Becx P;811J1I,TX 0 MOHO- 11 ]:(HMopcpIl3Me IWCTOqOblX
cJ>OPIll. no.'1yqeHHaH B CBeTOBOM MHRpOCKonc, CBH]:(eTcnbcTByeT 0 TOM,
qTO B IIReTOHll{ee BpCMH ]:(OCTnrHYTbl 60nblllHC ycrrexH B orrpcp;ene-
01111 OCIJOfiHOro .lJ;crro a.lJ;pCHanllHa II Hopa.lJ;peHanlIHa, 00 OCTaeTCJI
lIeHCllblM Han60nee Bamooe 3Beoo B 113YlIeOHII 3TOii IIp06neMLI-
MeCTO I1X o6pa30BaHliH. Klein II Kracht (1958) B CBoeM OqeHb
Tll{aTc.'1bIlOM I1CCnep;OBaIlHII He Mornll OTBeTIITb Ha aTOT Borrpoc II
303
'B COOTBeTCTBlIn C MUeHHeM Hillarp, H6kfelt (1953) MmJIII TOJIbJ\O
aRCTpanOJIHpOBaTb cpaKTbl, nOJIyqeHUble 61I0XHMIIKaMII (Kirchner,
Goodall, 1957) 06 06pa30BaHHH ap;peuaJIlIua II3 uopap;peHaJIHHa IIY-
TeM nplIcoep;IIHeUHJI MeTIIJIOBOH rpYIIIII.I. Ha 3TOM OCUOBannH OHn
nOJIaralOT, qTO Bce KJIeTKH M031'0BOrO CJIOH Hap;nOqeqHHRa rrpHcno-
. C06JIeULI R CIIUTe3Y Hopap;peHaJIHHa RaK IIpep;meCTBeHHHRa ap;peHa-
.JIlIua II TOJIbKO qaCTb 1I3 HIIX nplI06peTaeT cIIoc06uoCTb K MeTIIJIJI-
n;IIII Hopap;peHaJIIIHa B ap;peHaJIIIH. MCCJIep;OBaHlIe 3TIIX BonpOCOB B
3JIeJ\TpOUHOM MHRpOCRone IIMeeT 60JIee peaJIbHbIe B03MomHOCTII P;JUl
IIOJIyqeHHH II03IITHBHblX pe3YJIbTaTOB, oc06eHHo nplI IICCJIep;OBaHIIII
EX C nOMOIII;blO MeqeHLIX IIpep;llIeCTBeHHHROB II C yqeTOM 6HOXIIMII-
qeCKIIX p;aHUbIX.
OCHOBHOH nYTb 6IIOXIIMIIqeCROl'O CHUTe3a RaTeXOJIaMIIHOB 6LIJI
BnepBbIe HaMeqeH Blaschko (1939). hOT nYTb CBOP;HTCJI K CJIep;ylO-
III;eH cxeMe: ORHCJIeHlIe cpeHIIJIaJIaUHUa B THp03IIU H TIIp03IIHa J1.0
.3,4-P;IIOKclIcpeHIIJIaJIaUIIHa (p;ocpa); p;eRap60KCIIJIaqHJI p;ocpa B p;ocpa-
MIIU; ORIICJIeUlIe p;ocpaMIIHa B Hopap;peUaJIIIH; MeTIIJIJlqlIH Hopap;pe-
HaJIIIHa B ap;peHaJIHH. HaH60JIbmee ROJIHqeCTBO ROHeqHblX npOP;YRTOB
.CIIHTe3a (ap;peHaJIIIHa H Hopap;peuamma) onpe]:(eJIJleTCJI B rpaHY-
.JIJlpHbIX cppaKqnJlx II TeM caMblM p;ORa3bIBaeTCJI, qTO B naparaHrJIHO-
HapHblX KJIeTKax Uap;IIOqeqHHROB B3pOCJIblX mlIBOTHblX KOHeqHbIe
npop;yKTbl IIpeBpaIII;eHHJI KaTeXOJIaMIIHOB cop;epmaTCH He B BIIp;e pa-
CTBOpeHHOH B qHTOnJIa3Me cy6cTaHqnlI, a ocpopMJIeHLI B rpaHyJThI.
Tp;e me, B KaKnx CTpYRTypHblX 3JIeMeHTax IIaparaUrJIIIOHapHblX RJIe-
"TOK IIpOIICXOP;JlT npeBpaIII;eHWI, npep;llIeCTBYIOIII;IIe C03peBaHII1O ROHeq-
HblX IIPOP;YKTOB? C03p;aIOTCJI JIH OHII B Tex me rpanyJIax 11 TOJIbKO
B HlIX OP;HHX, II Torp;a, CJIep;OBaTeJIbHO, op;Ha H Ta me rpaHyJIa 110-
CTeneUHO p;oJImua MeHHTb CBOH 6HOXHMHqeCRHH aMHUHblH COCTaB,
EJIlI me qaCTb peaKqlIH, OCYIII;eCTBJIJlIOIII;HX IIOCJIep;OBaTeJIbUbIH CIIH-
Te3 aMHUOB, IIpOHCXOP;HT BHe rpaHYJI - B caMOH qHTOIIJI33Me KJIeTOR
II IIOTOM BHOBb nepeXOp;lIT B rpanyJIY?
B 1954 1'. Hillarp, Nilson, H6kfelt npep;nOJIOmHJIII, qTO ceKpe-
TopHLIe rpaHYJILI JlBJIJlIOTCJI ue TOJIbKO CTpYRTypaMlI ]:(JIH XpaHeHlIH
aMIIHOB, HO MorYT 6LITb MeCTOM pa3p;eJIbHOI'O CHHTe3a ap;peHaJIIIHa 11
Hopap;peHaJIIIHa. B IIOIICRax :mCIIepHMeHTaJIbHLIX p;ORa3aTCJlbCTB aB-
TOpbl nO]:(BepraJIH rpauYJIHpHbIe cppaRqHlI Hap;IIOqeqHlIKOB p;eHcTBHIO
:l-'a:mHqUbIX <!m31IqeCRlIX 11 XIIMHqeCRIIX areHTOB (pa3Hblx cpep;, TeM-
nepaTypHblM II OCMOTnqeCKlIM cpaRTopaM), BbI3bIBaIOIII;lIX BbJ]:(eJIeHHe
J{aTeXOJIaMIIHOB H3 rpaHYJI. OUH OmHp;aJIH nplI aTOM pa3UYIO peaR-
n;HIO co CTOpOULI P;BYX TYlnOB rpaHYJI. Pe3YJIbTaTLI 3RCnepHMeUT8
He nop;TBepp;IIJIII OmllJl;aUIiH. 06a THna rpaHYJI pearlIpOBaJIH op;lIua-
J\OBO, 0 qeM MomHO 6blJI0 CYP;IITb Ha OCUOBaHHn IIpoqeHTHoro co-
OTHOllieHIIJI ap;peHaJIHHa II Hopap;peHaJIHUa, KOTopoe OCTaBaJIOCb TaKIIM
me, KaKHM 6LIJIO P;O onblTa. Heyp;aqHOH OKa3aJIaCb II
p;pyraJl cepllJl OIIbITOB, B KOTOpOH 3TII aBTopLI IIblTaJIIICb nOJIyqIlTb
ABa TlIIIa rpanyJI nyTeM p;HcpcpepeHqllaJIbHOI'O n;eHTpHcpyrllpoBaHIIJI.
JJ;aHHbIe 3KCnepHMeHTOB n03BOJIIIJIII 11M c]:(e.rraTb 3aKJIIOqeUlie 06 OT-
CYTCTBHII cYIII;eCTBeHHOH pa3HIIqLI B CTpYRType II CBOHCTBax HOp8A-
.304
peHaJIlIH- II ap;peHaJlIIHCOp;epmaIII;lIX rpaHYJl. R TaKOMY me 3aRJlIO-
qeHlI10 IIpmneJl II Hagen (1954), nOKa3aBmlIH ocamp;eHile 0601IX
aMilHOB B op;HoIIMeHHoH cppaRQlI1I 60JIbllllIX rpaHYJl.
I1cCJlep;OBaHlIe cpepMeHTaTIIBHblX IIpOQeCCOB, yqacTuyroIII;IIX B ]:(e-
Rap60RCliJIlipOBaHlIM p;ocpa B Hap;IIOqeqHlIKaX, BbIHBIIJIO pa3HYIO CTe-
neHb aRTIIBHOCTII p;ocpa - p;eRap60RCIIJla3bl BO cppaRQIIHx rOMore-
HaTOB. Blaschko if coaBTOpbl (1955) B rOMoreHaTax M03TOBOro CJlOH
6bIKa HalllJlII 3TOT 3H3IIM n Hap;OCap;OqHOH mlIp;ROCTIl, B TO BpeMJI
RaR rpaHYJIHpHaJl cppaRQIlH OKa3aJIaCb JIlIllIeHHOH p;eRap60KCIlJIa3-
HOH aKTllBHOCTil. ABTOpbl npep;IIOJIOiI\IIJIIl, qTO eCJIIi 3TIl p;aHHbIe IIe-
peHeCTIf Ha RJIeTRY, TO He06xop;IIMO Cp;eJIaTb p;Ba p;onYIII;eHIfJl. IIep.
Boe p;onYIII;eHIIe 3aRJllOqaeTCJI B TOM, qTO 06pa30BaHIIe aMIiHOB IIpO-
IICXOP;I1T B «KJICTOqHOM CORe), a xpaHeHlIe ROHeqHblX npOP;YRTOB-
Hopa]:(peHaJIlIHa II ap;pcnmnma - B rpaHYJIax. BTopoe p;OIIYIII;eHlle, Bbl-
TeRalOIII;ee Ir3 nepBoro, 3aRJlIO'IaeTCH B nplI3HaHIIII p;BlImeHIIH aMIIHOB
B RJleTHe B ]:(BYX nanpaBJIeHIIHX: B HanpaBJIeHlI1I I13 QIITOnJIa3MLI K
I'paHYJIaM B nepllop; HOBo06pa30BaHllJl aMIiHOB II B 06paTHOM HaIIpaB-
JIeHIIII - II3 rpaHYJI B QIITOIIJIa3MY B neplIop; Bblp;eJIeHIIH ropMOHOB._
C MHeHIIeM Blaschko II coaBTOpOB COrJIaCIIJIIiCb Bertler C coaBTopa-
Mil (1960), KOTOpbIe HalliJIll, 'ITO p;onaMIiH o6pa3yeTcH II3 p;OIIa.
}];aJIbHeHIlIYIO JIOHaJIII3aQIIIO aMHHOB 110 nYTll IIX CIIHTe3a Bertler
C coaBTopaMII IIpep;CTaBJIJlJIII CJIeP;YIOIII;IIM 06pa30M. 06pa30BaBmIIHCJI
B QIITOnJIa3Me p;onaMIIH lIepeXOp;IIT HaHHM-TO nyTeM BO cppaKQlI1I
60JIbmlIX rpanyJI, I'p;e BepOJlTHee Bcero I'IIP;pOKCIIJIIIpyeTCJI B Hopap;-
peHamm. TaRoe 3aRJIIOqeHIIe aBTopOB COBnap;aeT C onblTaMIi Kirsh-
ner (1962), ROTOpblH HallleJI, qTO 3H3HMHaJl CIICTeMa, OTBeTCTBeH-
HaH 3a rlIp;pORCIIJlJlQIIIO 60ROBOH n;enll npep;meCTBeHHIIKa Iiopap;pe-
HaJIIIHa - p;onaMIIHa, CBJl3aHa C rpanyJIHpHOH cppaKQIIeH KJIeTOK.
3TOT me aBTOp B P;PyroH pa60Te COBMeCTHO C Goodall (Kirshner, Go-
odall, 1957) IIOKa3aJIII, qTO MeTIIJIJlQIIH Hopap;peHaJIIIHa B ap;peHa-
JIHH IIMeeT MeCTO B Hap;OCap;OqHOH cppaRQIIII.
ECJIII IIepeHOCIITb pe3YJIbTaTbl 6nOXllMIIqeCRIIX p;aHHLIX, nOJIyqeH-
HblX nYTeM cppaRQIIoHlIpOBaHIIJI 1'0MOreHaTOB, Ha KJIeTRY II ee CTPyK-
TYPLI, TO Hap;O COrJIaCIITbCJI C paccymp;eHIIJlMII II p;onyn:J;eHmIMIi Bla-
schko C coaBTopaMII (1955) 0 CYIII;eCTBOBaHlIlI B ap;peHaJIOBOH KJIeTKe
IIOCTOJlHHO CYIII;eCTBYIOIII;IIX «TOKOBl) RaTeXOJIaMIIHOB IIJIII, IIHaqe, p;B1I-
meHIIH IIX B rpaHYJIbl II 06paTHo II3 rpaHYJI, He06xop;IIMoro P;JIH ca-
Moro npoQecca CllHTe3a aMIIHOB. RpoMe Toro, Hap;o nplI3HaTb, qTO
OCHOBHOH CIIHTe3 RaTeXOJlaMIIHOB npOIICXOP;IIT B QIITOIIJIa3Me, a aR-
TIIBHblH MexaHII3M P;JIH ROH:qeHTpa:qIIII II oCB060m]:(eHIIJI aMIIHOB Ha-
XOP;IITCH B rpaHY JIax.
Ham 3KcnepllMeHTaJIbHLIH MaTeplIaJI He II03BOJIJleT CYP;IITb 0 IIpo-
Qeccax CIIHTe3a RaTeXOJIaMIIHOB B xpOMacpcplIHHblX KJIeTRaX, HO p;aeT
B03MomHOCTb BblCKa3aTb HeKOTOpbIe co06pameHIIH 0 p;IIHaMIIKe MeM-
6paHHbiX CTPYRTyp ROMnJIeRCa fOJIbp;mlI, HOTOpLIH pac:qeHIIBaeTCJI
HaMil B TeCHOH CBJl31I C HHTeHCllcpIIKan;lIeH HOBoo6pa30BaHIIH rpa-
HYJI. ECJIII CPYHRQIIJI ROMnJIeKCa fOJIbp;mII OrpaHII'IIIBaeTCJI KOHp;eH-
ca:qlIeH npOP;YRTOB ceKpeQlI1I RJIeTKlI, a caM CIIHTe3 npOIICXOP;IIT B
1/4 11 H. A. CMHTTeH
305
3H)J;OIIJIa3MaTllqeCIWM peTHI<YJlYMe, TO IlIlipOKOe pacnpOCTpaHeHlle
ero CTPYRTYPHMX 9J1eMeHTOB B nepllo)J; IIaHOOJlee HHTeHCHBHOH npo-
)J;yRqllll ropMOHOB B xpOMacpcplIHHOH RJleTRe (1lI0ROBOe COCTOHHlle)
OOeCIIeqHBaeT IIplIOJllImeHlle ero 9J1eMeHTOB R MeCTaM CIIHTe3a. ECJlH
me 8J1eMeHTbl ROMIIJleKCa fOJlb)J;mII yqaCTBYIOT B HeRoTopMX 3Tanax
CIIHTe3a RaTeXOJlaMIIHOB, TO yBeJllIqeHlIe nOBepXHOCTII OOeCneqHBaeT
paCllIlIpeHlIe nJlOIqa)J;1I )J;JlJI OCYIqeCTBJleHIIH cpepMeHTaTIIBHblX peaK-
IIH CIIHTe3a ropMOHOB. MM CKJlOHHbl paCCMaTplIBaTb MeJlRlle Be3II-
KYJlbl, pacnpOCTpaHJlIOIqHeCJI no BceH qlITonJla3Me B nepllO)J;M aK-
TIIBHoro CIIHTe3a 1'0pMOHOB, npHHa)J;JlemHOCTbIO KOMnJleKCa fOJlb)J;mII
(CMHTTeH, 1965a, 6).
BblcRa3aHHoe HaMil cooopameHlle 0 )J;BlImeHIIH MeJlRIIX Be3IIKYJI
OT OKOJlOJl)J;epHOH 30HM K nepHcpepli1i RJleTKII B HaCTOJlIqee BpeMH no)];-
TBepm)J;aeTCJI aBTopa)J;IIOl'pacpllqeCKIiM MeTO)J;OM (Moppert, 1966).
ITocJle BBe)];eHIIH KpblcaM )J;IIrll)];pOKclIcpeHHJlaJiaHIIHa-H 3 npocJleme-
HO 00pa30BaHHe nY3MpbKOB fOJlb)];mll B RJleTKaX M03roBoro CJlOH
Ha)J;IIOqeqHIIRa, ynJlOTHJlIOIqIlXCJI II YBeJllIqHBalOIqIlXCJI no Mepe IIX
CMeIqeHIIH Ha IIepllcpeplIlO KJleTKII. AHaJlOrlI'IHbIe )J;aHHMe IIoJlyqeHLl
Ha RJleTKaX CJlII311CTbiX meJle3, II BblCRa3aHO cooopameHHe, 'ITO ceK-
peT HeOeJlKOBOH nplIpO)J;bl CIIHTe3HpyeTCJI B KOMIIJleKCe fOJlb)];mII
(AJlOB, 1968).
TaKoBLl COBpeMeHHbIe IIpe)];CTaBJleHIIJI 00 YJlbTpacTPYKTYPHOH op-
raH1I3 IIH KJleTOK M03rOBoro CJlOH Ha,rJ;nOqeqHIIKOB BMCIDIL"'{ n03BO-
HOqHLIX II 0 MeMopaHHblx MexaHH3Max, yqaCTBYIOIqHX B ceKpeTOp-
HOM npoqecce.
r ,//,a6a X
3AKJIIOQEHHE.
3AKOHOMEPHOCTH 3BOJIIOQIIH
CHMIIA TO-A,IJ;PEHAJIOBOII CHCTEMbI
IIpHBeAeHHblll B HaCTOHID;ell paOOTe cpaKTlI'IeCKlIll MaTeplIaJI aD-
Topa no cpaBHHTeJIbHOll MopcpOJIOrHII CHMnaTH'IeCROll H xpOMacpcpHH-
Hoii CHCTeM n03BOHO'lHbIX B COt.IeTaHHH e nOJIyt.IeHHbIMII H KpllTHqe-
CRH paCCMOTpeHHblMlI JIHTepaTypHbIMH HCTOqHIIRaMII ,nalOT UCHOBa-
Hlle c]:(eJIaTb HeKOTopble 06U\IIe BbIBO,nbl H oupe,neJIIITb cpIl3HOJIOl'lfIIe-
CRill I CMbleJI r.IOP4>OJTOrllqeCROll opraHH3aqHH CIiMlIaTo-a,npeHaJIOBOll
CHCTeMbI BbICllIliX 1I03BOHO'lllbIX.
TeH,neHqHH R JIORaJIH3aqHH xpOMacpcpHHHOll TRaHH B Ha,nIIOqeqHII-
Re, HaOJIIO,naeMaJl B npoqecce SBOJIlOqIIII 1I03BOHOqHbIX, II MOpcIJo-
CPYHRqHoHaJIbHOe YCJIOmHeHlIe CHMlIaTHqeCROll HepBHon CHCTeMbI-
STO ,nBa npoqecca, ROTopble IICTOpHqeCRII B03I1I1RaIOT BMeCTe II na-
paJIJIeJIbHO pa3BHBaIOTCH, 06eCneqHBaH B3a1lMOBblro,nHbIe YCJIOBI:IJI
CPYHRqlloHIIpOBaHIIH. Cocpe,nOTOqeHHOCTb TRaHII B O,nHOM MeCTe II ee
r.lOpcpOJIOrHqeCKOe ocpopMJIeHHe B BlI,D;e CTPYRTypbl opraHHOI'O TlIlIa
03HaqalOT npem,ne Bcero naJIHqHe CJIOmHOH CIICTeMbi MecTHoro RpOBG-
ooparqeHIIH, nplICIIOC06JIeHHOrO K BbmOJIHeHlI1O CJIOmHblX CPYHKD;JlU
,naHHoro opraHa. CcpHHRTOpbl, apTepHOBeH03HbIe aHaCTOM03b1, CIIHY-
COII,nHoe CTpoeHlIe lialIlIJIJIHpHOH CIICTeMbi II ,nPyrlle COCY]:(IICTI>IC
CTPYRTypbl, ROTOpbIMH 06JIaJJ;a1OT Ha,nnOqeqHlIRII BblCllIlIX II03BOHOq.
HbIX, u6eCIIeqHBalOT nona,naHHe B RpOBb ropMOHOB B HymHbIe MOMeHTbI,
B HymHbIx ROJIHqeCTBaX 1I B HymHbIe OTpe3RlI BpeMeHH. ECTeCTBeU-
HO, qTO TaIWH MexaHH3M OOJIee cOBepmeHHbIH, qeM nOCTOHHHoe nD-
cTynJIeHHe ropMOHOB B ManlCTpaJIbHblH IipOBOTOR II3 oRpymalOlI{CU
ero xpoMacpcpllHHoH TKanH. Ho ocyrqecTBJIeHHe STOro MexaHU3Ma B03-
MomHO B YCJIOBIIHX BblCOROH MOPCPOCPYHRqHoHaJIbHOll opraHlI3aqHII
elIMnaTlIqeCKOH HepBHoH CHCTeMbl, ee CBJl3eH C HepBHblMII qeHTpa-
1\IH II C nepHcpepHqeCRIIMII aHaJIlI3aTopaMH. I1MeHHo CHCTeMa MeeT-
HOro KpOBOTORa, cHaomeHHaH COBepllleHHblMH B MopCPOJIOrlflIeCKoM
II CPII31I0JIOrllqeCKOM CMblCJIe IIHHepBaqHoHHblMII MexaHH3MaMII, ooce-
neqIIBaeT TY TOHRyrO npHCIIOCOOJIeHHOCTb cIIeqHcpHqeCRoro CPYHRqIlO-
HlIpOBaHIIH opraHa R nOTpeOHOCTJIM Bcero opraHII3Ma H OblCTpOTY
peammaqlIlI ee, ROTopaH B03MomHa TOJIbKO Ha BblCOROM ypOBHC SHo-
JIlOqllll HepBHoH CIICTeMbl. B CBOIO Oqepe,nb, HepBHaH CIICTeMa cno-
cooHa ocyrqecTBJIHTb perYJIHPYIOII{ee BJIHHHHe C HallOOJIbmeH JIer-
HOCTblO B TOM CJIyqae, Ror,na cneqHcplI'IeCRaJl CPYHRqHH SH,nORpIlHHOi
xpOMacpcpllHHOH meJIe3b1 JIORaJIII30BaHa B O,nHOM Y'IaCTKe. MopcpOJIO-
I'lI'IeCKaJl qeHTpaJIlI3aqlIJI xpOMacpcplffiHOH TRaHII B O,nHOM opraH9
1/ 2 12 H.A.CMHTTell
307
AeJIaeT Hall60JIee :JWWCKTHBHbIM «ynpaBJIeHlle) ero eJlTe:IbHOCTbIO
co CTOpOHbl HepBHoii: CIiCTeMbl, [10 cpaBHemlIO C TeM, qTO HMeJIO
6Ll MeCTO IIpli IIWWY3HOM ee pacnpe eJIeHIIII B TYJIOBIIIII;e mIiBOT-
Horo.
TaKnM 06pa30M, napaJIJIeJILHO pa3BIITliIO IICnOJIHIITeJIbHoro 3BeHa
peWJIeRTopHoii: A)'rlI, YCTaHOBJIemlIO CTPYRTYPHblX CBJl3eii: ero C MHO-
roo6pa3HLlMli peQenTopHblMII HepBHblMII OROHqaHIIHMIl II, CJIe OBa-
TeJIbHO, IIapaJIJIeJIbHO YCJIOmHeHIlIO, pacmllpeHIlIO II YTOqHeHIIIO WII-
3IIOJIOrlIqeCKoii: peryJIHQlI1I 3H ORpIIHHoii: WYHRn;1I1I IIaparaHrJIlleB,
II eT npoQecc MOpWOJIOrlIqeCRIIX npe06pa30BaHIlii: Bceii: a]:(peHaJIOBOC
(xpOMaWWIIHHoii:) CIiCTeMLl. npOIlCXO]:(HIII;lIe nepecTpoii:KII QeJIeco06-
pa3HO HaIIpaBJIeHbI Ha HalI60JIee 3wweRTIIBHoe pemeHlIe 3a;J;aq, B03-
HIIRalOIII;IIX B xo e 3BOJIIOQIIII Ha nYTII pa3BIITliJI Bblcmnx n03BO-
HO'lHbIX C IIX OqeHh CJIOmHoii: 06III;eii: opraHII3aQlleii:.
npe06pa30BaHIIJI, COBeplllalOIII;lIeCJI B IIpOn;ecce WIIJIOreHe3a 1103-
BOHOqHblX mlIBOTHblX, conpOBom aIOTCJI rJIy60RHMII 113MeHeHIIHMII B
TOIIorpawo-aHaTOMII'IeCROM pacnOJIOmeHlI1I ROMnJIeRCOB xpOMaWWIIH-
HOii: TRaHII. 113MeHeHIIJI BblpamaIOTCJI B nOTepe ROHTaRTOB ee C O -
ImMIi TRaHeBblMIi CIiCTeMaMIi II opraHaMII II B B03HIIKHOBeHIIII HO-
BLlX, HalI60JIee BbITO HbIX JIJI OCYIQeCTBJIeHIIH Ha JIemaIII;eii: 3H 0-
RpIIHHOii: a peHaJIOBoii: WYHKQIIII, CBoii:cTBeHHoii: Ram OMY II3 3TanOB
RCTOpnqeCROrO pa3BIITIDI n03BOHOqHbIX. AHaTOMlIqeCRlIe nepeMeIII;e-
IIHH MeHJlIOT CTPYRTYPY Tex opraHOB, B ROTOpblX npOTeRaIOT 3TII
npOQeCCbl, B pe3YJIbTaTe qero O;IJ;HII II3 HIL"'{ TepJlIOT B ClloeM CO-
CTllBe xapaRTepHbIe 3JIeMeHTLI xpOMaWWIIHHoii: TRaHII, PyrlIe npll-
06peTaIOT IIX, B TpeTblIX MeHHlOTCH MeCTa IIX paCIIOJIOmeHI:JH. I1Hbl-
HII CJIOBaMII, aHaTOMllqeCRlIe IIepe BllmeHIIJI BJIeKYT 3a c060ii: rn-
CTOJIOrII'IeCRYIO nepecTpoii:RY Tex CIiCTeM opraHOB, KOTopLle ORa3L1-
BaIOTCJI 3aTpOHYTblMII 3TIIM npOQeCCOM II ROTOpbIe qeTRO
06HapymllBaIOTCJI nplI CpaBHIITeJIbHG-MOPWOJIOrlI'IeCROM .D3yqeHIIII
.rOMOJIOruqHbIX opraHHblX CIICTeM Y pa3HbiX KJIaCCOB II03BOHOqHblX
mlIBOTHbIX.
COBepmeHHO eCTeCTBeHHO, qTO aHaTOMo-rIiCTOJIOruqeCRRC nepe-
CTpoii:RII, B OCHOBe ROTOpblX JIemlIT QeJIeC006pa3HO HaIIpaBJIeHHOe 113-
MeHeHue 3H)J;ORPUHHOii: WYHRQIIII xpOMawwuHHoii: CIICTeMLl B PH Y
n03BOHOqHLlX, OTpamaIOTCJI II Ha CTpoeH1I1I xpoMawclJuHHLlX KJIeTOR.
Hx 3a aqII MeHHIOTCJI B COOTBeTCTBUIi C 3BOJIIO IIeii: Bceii: CIiCTeMLl.
OT He06xo uMOCTIi IIOCTOHHHO CTOHTb Ha CTpame IIHTepeCOB Bcero
opraHU3Ma U BbIDOJIHJlTb 3TY 3a aqy «CaMOCTOJlTeJIbHO» (6e3 cllr-
HaJIOB CO CTOpOHbI HepBHoii: CIICTeMLl, He cnoco6Hoii: eIII;e R 3TOii:
pOJIII), xpOMaWWUHHaJl RJIeTRa IIOCTeneHHO [lpIiCIIOCa6JIIIBaeTCJI K
TOMY ypOBHIO pa3BIITIIJI, ROTOporo oCTuraeT cJIl1JIOreHeTuqecRII 60-
nee pa3BIITLlii: opraHlI3M, 06JIa aIOIII;IIii: UHOii: MopcJIOJIOruqecRoii opra-
1IR3a BeH xpoMacJIwmmoH TR8.Jm. B 3TUX HOBLlX YCJIOBBJIX cJIU3IIO-
nOrUqeCRaJl 3ap;a'la xpoMawcJllIHHoii KJIeTRB «06JIeI"laeTCH». TeIIepb
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0 BJ .---:r-.rp.r "'"'-.... L
! '1
c. ./
......,......-....rr"
- -
S"
III
Of 02 II J . 0 .J
'.
]' II ('. IIH. CXl' Ia (1IllJlOll'I\l'3a aAPl'!liUlOlIoii (xpmw4>4>lIlllloii) CIlCTeMbi y n03-
1I0!lO'IIIhlX !Iii 'H'ThIjIl'X ypOlllIflX
Ll - aIlIlTO IlI'ICCI<O I; E - rIICTO:IOrlI'lCCHOM; n - 'ITOJIOI'II'leCHOM; r - 1!J1I3HOJIOl'II'IeCHOM
( eTalio""'ICCHOM). I - Hpyr:\IIpnThIC II pbl6hl; II- 8CMHOBOgHhIC II npCCMhIHaIO HecH; III -
nrn bl II M:ICHOnI'TaIOlInlC. 3a OCHOBY HpoBoo6pa eHIIH MlIHOI' B3RTa cxeMa B. C. MaTBee-
113 (1963).
1 - apTCplIa!II.naR clIcTeMa; 2 - Rcn03HaR CHCTCMa; 3 - anpellaJIOBaR CHCTcr.ra; 4 - nOJIHO-.
enllo 4JYHl< I1()III1PYIOII\dH ;Hf':1C3I1CTaU TH31lb aAPCHaJIOBOii CIICTe I\J: 5 - PYiUlMCHTapHall.
}I{C!IC31!CT3fI THanb a:tpella.'1ouoii CHCTCMhI
( '.U. j
--
{';
o
{!
2
3
Pile. 119. CXCMa pacnpc;{e.'l"I1I1H a.1pl'I1i1:IOBOii (xpo lil4upililIlOii) TKaHII II oc-
HODHbIX cn maTII'leClmX CTDO:WII y IlIoICIIIIIX I10;mOIlO'IIIhIX. l l'HTpaJIlI.JannH
clJYHKLI,lll1 D M03rOBO 1 ,':10" I1i1;\III1'!I"I11I1IiOIl:
J - nOJIIiOIleHHO I!JYHHIUiOIll'PYIOIl\aR meJIe311CTaR THallb 3ApCnaJIOBOA CIICTCMbI' 2 - VIlII-
MCHTapHaH menC3BCTaH 'l'HaHb agpCnaJIOBoi! cllcTeMbI; 3 - CTBOJIbI II Y3JIbI CIIMllaTII'leCl<o"
OIlCTeKbI
IIX XeMOpen;eIITOpHOll «PYHKn;IDI (Driiner, 1925; de Castro, 1926,
1928), YCTaHOBJIeHHOll «p1I3HOJlOraMIi B Tex me 30Hax :..... COrJIa CHO_ (
.a'1'Oll KOHn;elIn;1I1I lIaparaHrJIHOHapHble KJIeTKH HBJIHIOTCH Mop«pOJIOrH-
"leCKIDd ey6cTpaToM, OTBeTc.TBeHHblM 3a BocnpllHTlIe C BHrOB X1U411Qe-
CKOro_ co.CTaBft npoTeKalOm;ell B cocYJUlX KpOBH. O]:(HO 113 HanpaBJIe- I
HlIll IIOCJIe]:(YIOm;ero pa3BIITllH aTIiX B3I'JIH]:(OB IIpIiBeJIO K «p1I3110JIO-
ruqeCKHM 0606m;eHHeM (Comroe, Schmidt, 1938; Schmidt, Comroe,
1940) 0 TOM, qTO BbiCOKopa3BIITaH qyBcTBHTeJIbHOCTL COCY]:(OB meiiHo- ·
rpYAHoll 06;'IaCTIi 06YC!IOBJIeHa IIX ma6epHblM npOHcxom]:(eHlIeM. B Ka-
-.:reCTBe IIoJJ;KpenJIeHIIH 8TOMY nOCJIe]:(HeMy BblBO)UT CJlymlIJIII &M6plIo-
JIOrllQeCKIIe ]:(aHHble (Boyd, 1937b) 06 06m;eM Me30]:(epMaJIbHOM IIC-
TOqHIIKe pa3BHTliH II ma6p II mellHOI'O (KapoTII)J;Horo) lIaparaHrJIHH t.
O.1HaKo «paKTlIqeCRlIe aBHble H3yqeHnJl. naparaHrJIlIOHapBoii CII-
CTeMhr B cpaDHIITeJIbHO-MoP«P0JIOruqecKoM aCIIeKTe, BCRpblBalOID;IIe B8-
KOHOMepHble H3MeHeHllH opraHOB aTOll CHCTeMbi B f!mJIoreHeae no-
. !BOHOqHb!.X, n03BOJIHIOT_roBopIITL 06 a ORpIIHHOM xapaKTepe napa-
raHr':I1IeB B npHcep]:(eqHblX cocy]:(ax ma6epHoro npOllcxo eHIIH II
'060cHoBLIBaIOT IIX peAYltllPOBaHHhle a OKpllHHble CBOllCTBa. He060c-
HOBaHHOCTL me IIpe]:(cTaBJIeHlIll 06 IIX xeMopen;enTOpHoll «PYHKn;IIH
BblTeKaeT II3 Bcex HaKOnJIeHHblX 3a 1I0CJIe]:(HlIe eCHTIIJIeTIIH CBe]:(e--
HIiH 0 qyBcTBllTeJIbHOll IIHHepBan;nll BnyTpeHHHX opraHOB, KOTOpas
HBJIHeTCH HeOT'LeMJIeMblM II a6COJIIOTHO He06xo]:(IIMbiM 3BeHOM B
4>YHKn;IIHX HepBHoll CHCTeMbi. lIB aTOro CJIe]:(yeT C eJIaTL 3aK.JIlOqe-
Hne, QTO He T C II e n; II a JI b H bI X X e :M 0 P e I{ e n TOp H hi X 0 P r a-
HOB, a e C T b X e MOp e n; e II TOp H a H «p Y H K n; II H, C BOll C '1'-
BeHHaH JII060MY opraHY II 06H3aHHaH CBOIIM C
.IIJ;eCTBOBaHHeM pe«pJIeKTOpnOll ]:(eHTeJILHOCTII HepB-
H 0 H C II C T e M bl.
Bce MaTeplIaJIbI, npllBe]:(eHHble B HaCTOHm;ell pa60Te II OCHOBaH-
Hble Ha COBpeMeHHblX MeTo]:(ax MllKpOCKOlIuqeCKoro n rIiCTOXIIMIIQe-
(;KOrO aHaJI1l30B, nOKa3b1BaIOT MOP«Po«pYHKn;llOHaJIbHOe e HHCTBO Bcex
IIaparaHrJIlIeB, npllH8J];JIemHOCTb IIX K e]:(IIHOll 8H]:(OKpnHHOll CIICTeMe
II 3aBllCllMOCTL «p1l3110JIOrIlQeCKOll aKTllBHOCTH OT ypOBHH aBOJIIOn;IIOH-
HOro pa3BllTliH nepll«peplIQeCKllX n n;eHTpaJIbHblX CBH3ell CllMnaTII-
"lecKoii HepBHOll CllCTeMbi.
t Boyd C'IHTaJI, 'lTO npeccopeIlenTopHlde SOHId P1d6, 8McJ>1I61Iii: II K8POTHJJ;HIdH
CIIHYC MJIeKOUHT8IOII\HX HBJIJlIOTCJl rOMOJIOrH'lHLIMB: o6paSOBaHIIJIMB:.
.
JIIITEP A TYP A
Aeap1we r. B. HepBHIdH annapaT Hap;nO'le'lHIdX meJIe3. M., M3P;-BO «Mep;lIl\u-
Ha», 1964.
A.ltOB 11. A. HOBlde p;aHHlde 0 cel<peTOpHoii cJ>YHKl\HH KOMnJIeKCa rOJIbp;mlI.-
"Yen. COBpeM. 6HOJI., 1968,65, 144-151. .
AHU'lKOB C. B., Be.lteHbl>UU M. JI. CDapMaKOJIOrlIJi XIIMIIOpel\enTOpOB KapoTlIp;-
HOro KJIy60'IKa. JI., «Mep;rIl3», 1962.
Belf B. A. HeCRoJIbKo 3aMeqaHHII 0 MHKpOCKOnH'leCKOM cTpoeHIIH Hap;nO'leqHII-
KOB.- "YHIIBepcnTeTcK. 1I3BeCT'IIH. RlIeB, 1864, 6, 48-60.
BoiJpoBa H. B. CpanHlITeJIbHlde II BKcnepHMeHTaJIbHO-MopcJ>OJIOrll'leCKIIe p;aH-
Hide no IIHTpaKapp;lIaJIbHOH nepBHOH CIICTeMe XOpp;OBIdX lliIIBOTHbIX.-
TpYP;1d V Bcec. C'be3p;a aHaTOMOB, rHCTOJIOrOB H BM6pIIOJIOroB. JI., 1951.
588-590.
BoiJpoBa H. B. CpaBHIITeJIbHlde p;aHHlde no HHHepBal\IIH Cepp;e'lHO-coCYP;IICTOH
CIICTeMId JIaHl\eTHHKa, P1d6, aMcJ>1I6Hii II pellTlIJIlIH. OKT. P;IICC. M., 1957.
ByiJaHlfeB A. EO. "YcTaHoBKa P;JIH 6ldCTpoii JIIIOcJ>HJIbHOH cymlUl: II 3aJIIIBKlI B
napacJ>HH flICTOJIOflI'leCKOrO MaTeplIaJIa.- IITOJIOrIlH, 1967, 9, 245-248.
ByiJaHlfeB A. EO. HeiipoceKpeTOpnoe Be eCTBO 11 KaTl'XOJIaMIIHId nepep;Hero
rlInOTaJIaMyca B 3apop;ldmeBoM pa3BIITHH. RaHp;, Allee. M., 1968.
BbZ1WB N. M. Ropa roJIOBHOrO M03ra H BHYTpeHHlIe opraHId. M.- JI., «MeArII3'>,
1947.
BUHHUI>oB H. A. Pa3BlITHe nepBHOH TPy6KH II p;HcJ>cJ>epeHl\lIpOBKa HeHpOHOB.-
"Yen. COBpeM. 6I1oJI., 1946,21,379-399.
BUHHUI>OB H. A. Pa3BIITlIe H pOCT KJIeTO'lHIdX MeM6paH.- lliYPH. BBOJIIOl\. 6110-
XHM. H cJ>H3HOJI., 1966,2, 175-182.
B.lta{)oc-llpuuMal>. 9. X. R Bonpocy 0 pa3BlITHH l<apOTIlP;HOro TeJIa. Te3HcId
P;OKJI. VI Bcec. C'be3p;a aHaTOMOB, rlICTOJIoroB, BMUplIOJIOroB. RHeB, 1958, 2,
450-451.
BOpoHlfoBa M., JIuo3Hep JI., MapKeJ1.0Ba H., llyxaJ1.bCl>all E. TpHToH H aKCO-
JIOTJIb. M., «CoBeTcKaH JIIIT-pa», 1952.
Fa.ltblfoBa 3. B. a B03paCTlWH aHaTOMIIII COCYAlICTOH CIlCTeMId Hap;nO'le'lHHKOB.
C6. «Bonpocld aHaTOMHII COCYP;HCTOH CIICTeMId pe6enKa II B3pOCJIOrQ». JI.,
JIeHIIHrp. nep;HaTp. IIH-T, 1958, 357-365.
(raTeH6u /l,:J/C.) (Gatenby J. B.). AnnapaT rOJIbp;lliIl. C6. «IIp06JIeMId l\IITOcJ>II-
31I0JIOflIH». M., MJI, 1957, 242-272.
rOBblpUH B. A. 06 OTCYTCTBHH npHMoH CIIMnaTH'leCKOH lInHepBal\HH CKeJIeT-
HIdX MIdilll\.- OKJI. All CCCP, 1965, 160, 1179-1181.
rOBblpUH B. A. TpocJ>II'leCKaH cJ>ynKl\HH CIIMnaTII'leCKlIX HepBOB cepP;l\a II CKe-
JIeTHIdX Mldml\. J1., «llaYKa», 1967.
FOBblpUH B. A., JIeOHTl,eBa r. P. K Bonpocy 0 xpoMacJ>cJ>HrrrroH TKaHH H HCTOq-
HlIRaX KaTeXlIHOBIdX aMlIHOB B cepP;l\e n03BOHO'lIIbIX lliHBOTHIdX.- BIOJIJI.
BKcnepHM. 6110JI. H Mep;., 1965a, 59, 1, 98-100.
rOBblpUH B. A., JIeoHTbeBa r. P. Pacrrpep;eJIeHIIe KaTeXOJIOBIdX aMIIHOB B MHO-
Kapp;e n03BOHO'lHbIX.- lliypn. BBOJI. 6IIOXHM. H cJ>H3HOJI., 19656, 1, 38-44.
roiJUHOB B. M. BnYTpIICTBOJIbIIOe pacnpep;eJIeIIlle II cTpoeHlle HepBHORJleTO'lHO-
ro annapaTa 6JIYllip;aIO ero HepBa Y HeKoTopldX penTHJIIIH.- Apx. aHaT.,
rnCTOJI., BM6pIlOJI., 1939, 20, 1, 109-120.
Fo.lty6 /l,. M. P83BIITIIe H8p;nO'le'lHIdX llieJIe3 H HX IIHHepB8l\IIJI Y 'leJIOBeKa H
HeKOTopldX lliIiBOTHIdX. MHHcK, M3p;-BO All BCCP, 1936.
314
rO.ll.biJUH J/. C. OHToreHe3 l\Opld OOJIbmnX nOJIymaplIu rOJIOBIIOro M03ra. ORT.
p;IICC. JI., 1940.
rO.ltbiJuH J/. C. fHcTOreHeTII'leCRne OCHOBaHIIH HeiipoHHoH TeopIIII II HX KPIITH-
Ra.- TpYP;1d MH-Ta M03ra lIM. EexTepeBa, JI., 1947, 18, 22-24.
rO.ltbiJuH J/. C. MaTepllaJIId K BMUpnOHaJIbHoMY pa3BHTHIO cmlHHoro M03ra. Cli.
«OHToreHe3 M03ra», JI., MU-T M03ra HM. B. M. EexTe p eBa 1949 16 215-
234. ' "
ropoiJuHC1>aJl. C. !:. R Bonpocy 00 apXHTeKTOHIIRe 6JIYllip;aIO ero HepBa. HepB-
nO-KJICTO'lHIdIl annapaT 6JIY)I(p;aIO ero HepB8 RpOJIlIRa.- Apx. 6HOJI. HayK,
1936, 44, 3, 5-30.
rpueopbeBa T. A. MHHe{>Bal\Hfl KpOBelIOCHIdX COCYP;OB. M., «Mep;rH3», 1954.
/(oee.ll.b A. C. [Dogiel A.j Die Ncrvenendigungen in den Nebennieren der Sau-
getiere.- Arch. Anat. Physiol., Anat. Abt., 1894, 90-104.
/(oee.Jlb II. M. CpaBHIITeJIbHaH aHaTOMHH, cJ>1I3110JIOrIlH H cJ>apMaROJlOrHH l(pOBe-
HOCIIIdX II J1HMcJ>aTH'leCRIIX COCYP;OB. Ra3anb, 1903-1904.
/(oJl.eo-Ca6YPoB B. A. aT pep;aRTopa. B RH. C M II P II 0 B A. A. «RapOTllAnaH
pecJ>JIeRCOrelIHaH 30Ha». JI., U3P;-BO «BoeHHo-Mop. Mep;. aKap;.», 1945, 3-4.
/(o.lteo-Ca6ypoe B. A. IIo IIOBOP;Y RH. T. A. fpHropbeBoH «MHHepBal\11H KpOBe-
HOCHblX COCYP;OB». M., 1954.- Apx. aHaT., rlICTOJI., SM6pIlOJI., 1956, 1, 94-
97.
/(o.lteo-Ca6YPoB B. A. MHlIepBaI\IIH BeH (3KcnepHMenTaJIbHO-MopcJ>OJIOrlI'leCROe
HCCJIep;OBaIIHe). JI., «Mep;rn3», 1958.
/(ocToeBc"uu A. [Dostojevsky A.I Ein Beitrag zur microscopischen Anatomie
der Nebennieren bei Saugethiren.- Arch. mikrosk. Anat., 1886, 27, 272-
296.
/(YPUlfblH tP. A. MaTeplIaJIId K 1I3yqeHHIO cTpoeIIHH Hap;nO'le'lHbIX llieJIe3 MJIe-
KOIIHTaIO HX. OKT. p;HCC. RYH6IdmeB, 1942.
/(YPUlfbl.H tP. A. 06 oc06eIIHocTHX RpOBocH86llieHHH Hap;nO'le'lHIIROB.- EIOu.
BRcnepllM. 6HOJI. H Mep;., 1945, 20, 10-11, 60-61.
J/(UH"UH J/. H. MIITOTII'leCKlIe (KJIeTO'lnlde) I\IIKJIId. B RH. «PYKOBOP;CTBO 110
I\IITOJIOrlIlI». M.- JI., «HaYRa», 1966, 2, 244-251.
J/(upHoea A. A. MopcJ>oJIOrlI'leCKlIe II rHCTOXIIMlI'leCRHe 1I3MeHeHIIH B MOTOp-
HLIX KJIeTKaX rpyp;Horo OTp;eJI8 cnllHHoro M03ra II II nllp8MIIp;HIdX KJIeTRaX
p;BlIraTeJIbHOH 06J1aCTII KOpld 60JIbmlIX nonymapDii RpldCId H8 p8aJIII'IHbIX
3Tanax OHToreHe3a. AnTOpecJ>. KaHp;. P;lIcc. M., 1955.
BaeapauH A. A. a MOPcJ>OJIOrlI'leCROM 3H8'leHIIII aBToHoMHOH HepBHoH CIICTe-
MLI.- Apx. 6HOJI. HayK, 1937, 45, 2, 17-36.
BaeapauH A. A. O'lepKH no BBOJIIOI\IIOHHOH rllCTOJIOrlI1I HepBHoH CIICTeMbL M.-
JI., «Mep;rn3», 1941.
BaeapauH A. A. CIIHTe3 HR II RIIHeTHRa KJIeTO'lHIdX nonyJIHI\IIH B OHToreHe-
3e MJIeKOnIITaIO IIX. JI., «H8yK8», 1967.
Ba.ltlWHO M. M. a xpoMacJ>cJ>IIHHIdX BRJIIO'leHHHX B CIIMnaTlI'leCRIIX Y3JI8X. B
KH. «TpYAId, nOCB. 40-JIerneH p;elIT. npocJ>. B. H'. TOHKOD8», 1937, 283-
292.
Bo.ltOryxuH A. C. a cocYAax H8p;nO'Ie'lHHROD.- TpYP;1d 15 C'Le3p;8 pOCCIIHCKIIX
xHpyproB. IIr., 1923,313-316.
HBaHoe r. tP. XpoMacJ>cJ>HHH8H II IIHTeppCH8JIOB8H CIICTeMa 'leJIOBeRa. M.- JI.,
foe. Mep;. H3P;-BO, 1930.
HeaHoB r. tP. HepBId II opraHId 'lYBCTD cepp;e'lHO-cocYAIICTOH CIICTeMld. M.-
JI., «Mep;rII3», 1945.
HeaHoe r. tP. R Mop(f>OJIOrlIlI II KJI8CclIcJ>IIKa H HeKOTOpLIX cJ>OPM IIHTp8pel\en-
TOpOB. C6. dJp06JIeMld KOpTHRO-BIIC paJIbUOH naTOJIOnnI». M., li3P;-DO
AH CCCP, 1949, 135-153.
H.II.bUHa B. n. 3KcnepIIMeHTaJIbHO-rnCTOJIOrlI'leCKOe IICCJIep;OBanlIe IIHHepBaI\IIH
Hap;nO'Ie'lHIIKa MJIeROnIlTaIOII{IIX. C6. «MopcJ>OJIOrIIH aBTOHOMHOH HepBHoH
CIICTeMld»' M.- JI., «Mep;rn3», 1939, 109-128.
H.ltbUHa B. n. qYBcTBIITeJIbliaH IIHHepB8I\IDl M03rOBoro Be eCTB8 H8p;nO'Ie'IHlI-
Ka. C6. «MopcJ>OJIOrIDl 'lYBCTBIITeJIbHOB HHHepB8 1I BHyrpeHHIIX OpraHOB)).
M., li3P;-BO AMH CCCP, 1947,135-148.
KapneH"o B. II. liHHepB8I\I1H XpOM8cJ>cJ>IIHHOH TR8HII CIIMnaTlI'leCKIiX YaJIOB
nOrpaHII'IHOrO CTBOJI8.- Tpyp;1d BOJIrorp8p;cK. Mep;. IIB-T8, f964, 15, 128-130.
315
l(a/fHe.ltbCOH 3. C. 1\;:\peHaJIODlde II nopap;peHaJIOBlde RJleTRII B rraAnO'le'lHHKe
CBlIHbH.- AORJI. AH CCCP, 1964, 155, 1182-1183.
l(a/fHe" bcoH 3. C. [Katznelson Z. S.]. Zur Friihentwicklung der Nebenniere des
Schwl'ines.- Z. mikrosk.-anat. Forsch., 1965, 73, 187-199.
l( aIfHe.ltbCOH 3. C. [Katznelson Z. S.]. Spathistogenese der Nebenniere des
Schwcines.- Z. mikrosk.-anat. Forsch., 1966a, 74, 193-208.
KalfHe.ltbCOH 3. C. MaTepHaJIId no panneMY rncTOrenesy nap;nO'le'lHIlRa OBl\Id.
B KH. «MaTepllaJIId 15-ii HaY'ln. KOHcJ>epeHl\HH JIenHHrp. neTepHuapnoro
IIrr-TI\), JI., H)[j()u, 181-184.
KaljHeJlbCOH 3. C. HeRoToplde IITorll IICCJIep;OBanllii: no paSBIITIIJO II rllcTocJ>JI-
SIIOJIOrllH naAnOqe'lIIHKa MJIeKOIIIITaIO HX.- C6. pa60T JIcHllIlrp. BeTepn-
napnoro IIH-Ta. JI., 1!J67a, Bldn. 29, 172-186.
KUl/HeJlbCOH 3. C. rncTorenes II opraHoreHes Hap;nO'le'lIIIIKa KpynIIoro poraTO-
1'0 CHOTa.- Cu. paooT JIeHHHrp. BeTepHnapIIoro IIU-Ta. JI., 1967, Bldn. 28,
617-626(6).
Ka/fHe.ltbCOH 3. C. Bnp;oBldc oc06enHOCTlJ pasBIITIIH lIap;nO'le'lHHKa y onI{ld. Cu.
pa60T JIenlIHrp. BeTepllIJapIIoro HH-n. JI., 1969, Bldn. 30, 229-242.
KaUfeHl>O H. [Kastschenko N.]. Das Schicksal dcI' embryonal en Schlundspal-
ten bei Siiugetil'ren. (Zur Entwicklungsgcschiche des mittlercn und
iinssel'en Ohres, der Thyreoidea und der Thymus, Carotidenanlage).-
Arch. mikrosk. Anat., 1887,30, 1-26.
l(eiJpoBCI>UU B. B. HTOJlOrlIH UeJlIWBIdX CllnTeJOB B llillBOTIIOH KJIeTKe. M.,
MSP;-BO AH CCCP, 195H.
KUCe.ll.b 3. M. R Borrpocy 0 BeHOSHIdX anaCTOMosax Hap;nO'le'lHlIKa.- TpYAId
roc. IIll-Ta Mep;. sHaulIH. JI., 1928, Bldrr. 3, 67-71.
KHoppe A. r. AHcJ>cJ>epeHl\lIpOBRa KJIeTO'lHOrO MaTepHaJIa BM6pHOHaJIbHldX sa-
'laTKOB. AOKT. p;HCC. JI., 1948.
KHoppe A. r. ihlUpHOHaJIbHOe pasBIITlIe BereTaTIIBHoii: HepBHoH CHCTeMId nos-
BOHO'lIIIdX.- Yen. COBpelll. 61I0JI., 1949a, 27, 37-62.
KHoppe A. r. R Bonpocy 06 IICTO'lHHKaX BOSHIIKHOBeHHH CIIMnaTII'leCKHX YS-
JIOB nOrpaHII'lHOrO CTBOJIa B HOpMaJIbHOM BM6pHoreHese. C6. «OHTOreHes
MOSra». JI., MH-T Mosra 11M. B. M. BeXTepeBa, 19496, 16, 235-247(6).
KHoppe A. r., CYBopoBa J/. B. OCHoBHlde BTanbl p;1IcJ>cJ>epeHqllpoBHII HeHpo-
Ha.- Apx. aHaT., rHCTOJI., BM6pHOJI., 1959, 37, 7, 3-18.
KHoppe A. r., CyeopoBa J/. B. MCTO'lHIIHH pasBHTHH HeHpoHoB CIIMnaTII'leCHHX
raHrJIlIeB nOrpaHII'lHo1'O CTBOJIa B BM6pHoreHese nOSBOHO'IHIdX.- Apx.
aHaT., rIlCTOJI., 3M6pIIOJl., 1961, 40, 5, 93-104.
l(OBPU3/CI>O H. M. B03pacTHaH rlICTOJIOrll'leCKaH xapaKTeplIcTIIR8 xpoMacJ>cJ>IIH-
Hoii CHCTeMId 'leJIOBeKa.- Apx. naTOJIOrIIH, 1962a, 5, 39-46.
l(oepu3/Cl>o H. M. IIocTHaTaJIbHaH BBOJIIOl\IIH B oc06eHHocTII cTpoeHHH BamHeii-
mllx naparanrJIHeB 'leJIOBeHa.- Apx. 8HaT., rHCTOJI., 3M6pBOJI., 19626, 43,
f2, 50-58.
l(oBPU3/Cl>O H. M. MSMeHeHIIH B xpoMacJ>cJ>IIHHOH CIICTeMe B CBHSII C BospacToM
II naTOJIorneii. ABTopecJ>. p;OHT. AHCC. RHeB, 1970.
KO.lteCHUl>OBa H. A. MHTP8MYP8JIbHlde HepBHlde II xpoMacJ>cJ>IIHHlde aJIeMeHTId
JIBMcJ>aTll'leCKOrO npoToHa 'lepenaxlI.- Apx. aHaT., rIICTOJI., BM6pIIOJI., 1963,
44,2,34-38.
KO.ltOCOB H. r. [Kolossow N.]. Zur Frage der Innervation der Nebenniere.-
Z. mikrosk.-anat. Forsch.. 1930, 20, 107-121.
KOpOTl>OB A. r. R cpaBHHTeJIbHOH MopcJ>OJIOrnlI xpoMacJ>cJ>IIHHoii TKaHII B CTeII-
Ke aopTbl. B KH. «MaTeplIaJIId 4-H IIoBOJIllicK. KoncJ>. cJ>IISIIOJIOl'OB, cJ>apMa-
KOJI01'OB II 6HOXIIMIIKOB C yqaCTlIeM MopcJ>OJIOrOB H KJIHIIIIl\IICTOB». Capa-
TOB, 1966, T. 2, 343-345. ..
KOpO'll>UH J/. 11. AHcJ>cJ>epeHl\lIpoBl\a II cTapeHlIe BereT8TIIBHoro HellpoHa. M.-
JI., «HaYRa», 1965.
KOlUTORH/f X. C. OCHOBId cpaBHHTeJIbHOH cJ>IISIIOJIOrlIlI, T. I. M.- JI., MSp;-BO
AH CCCP, 1951.
KpacYCl>aR A. A. 06 IIH xapaKTep p8SBeTBJIeHIIH COCYP;OB B meJIesax C BHYT-
peHHeH ceKpeqlleH II B 'laCTHOCTH B Hap;nO'le'lHIIRax.- TpYJ;\1d I Bcepoc-
CIIHcKoro C'besp;a sOOJIOrOB, aHaTOMOBII rIlCTOJIOrOB. JI., 1923, 171-172.
IrPYMUHR Y. H. 3M6plIOHaJIbHOe pasBIITHe Hap;nO'le'lHIIKOB 'leJIOBeKa II HeRO-
316
TOphlX MJIeKOnlITaIOIl\I1X mIIBOTHIdX. B KH. «Bonpocld l\HTOJlOrnlI, rIICTO-
. J10rHII II BMUPIIOJIOrIlII». Pura, IIS)];-BO AH JIaTB. CCP, 1960, 199-204.
R ynpusUW6 B. B. HepBllJdll annapaT COCYP;OB MaJIoro Kpyra KpoBo06pa enIlJl.
JI., «MeArHS», 1959.
.7a6peHTbe6 B. If. qYBcTBIITeJIbHaH IIHHepBal\HH BHYTpeHHIIX opraHoB. C6.
«MopcJ>oJIorIlH 'lYBcTBIITeJIbHOH HHHepBal\HII BHYTpeHHHx opraHOB». M.,
Jf3A. AMH CCCP, 1947, 5-21.
JIe6eiJe6a B. A. MexauII3MId xeMOpel\enl\lIH. M.- JI., «Hay«a», 1965.
JIe6uHcoH JI. B., JIeiiKuHa M. N. MopcJ>oJIorH'lecKoe II rHCTOXHMII'IeCKOe IIsy-
'leUHe MOTOpHIdX II 'lYBCTBHTeJIbHIdX HepBHldx KJIeTOK BM6plIoHa KYPII-
l\1d ua paSHIdX 3Tanax paSBIITHH.- OKJI. AH CCCP, 1956, t09, 201-204.
JIe6uHcon JI. B., AHaHo6a E. B. HYKJIelIHOBlde EIICJIOTId B MOTOpHIdX KJIeTKaX
CTIllHlIOro M03m II 'lYBCTBlITeJIbHIdX KJIeTKaX CnHHaJIbHOrO raHrJIllH Ha He-
EOTOpblX CTap;HHX pa3BlITHH 3M6pHoHa KypHl\Id.- OKJI. AH CCCP 1956
109, 381-384. ' ,
JIeii6coH JI. r. Caxap KpOBH. M.- JI., IISA-BO AH CCCP, 1962.
Jleii6coH JJ. r., Cra6po6cI>uii E. M. COAcpmaHHe ap;peHaJIlIHa H Hopap;peHamI-
lIa B Hap;nO'leqUIlRaX pasBIIBaIO lIxcH KypllllblX BM6plIOHOB.- <Dn3110JI.
mypH. CCCP, 1962,48, 857-863.
J7eOHTbe6a r. P. Pacnpep;eJIeHlle KaTeXOJIOBIdX aMlIlIOB B CTeHKe KpOBeHOCHIdX
COCY!\OB Y I'PyrJIOpOTIdX. pldO. aMcJ>1I6l1ii II penTll;'lIIii. iRypH. BBO.'1IOl\. 6m>-
XllM. H cJ>HSHOJI., 196Ga, 2, 31-36.
J7eOHTbe6a r. P. Pacnpep;eJIeHHe KaTeXOJIOBIdX aMlIHOB B CTeHKe KpOBeHOCHIdX
COCYP;OB TenJIOKpOBnldX mHBOTHIdX.- Apx. aHaT., rHCTOJI., BM6pHOJI., 19666,
50, 3, 36-41.
JJeoHTbe6a r. P. 06 aApeHeprll'leCKOii HHHepBal\1I1I KpOBeHOCHIdX COCy.n:OB nOB-
BOIIO'lIlIdX lliIlBOTnIdX. ABTopecJ>. KaHp;. p;IICC. JI., 196B.
.7hlceHKo6 H. K. R MopcJ>OJIOrnll cunyc-HepBa feplIHra.- <DIISHOJI. mypH. CCCP,
1939, 27, 166-169.
JlaHyxuH B. H., ByaHuK06 r. A. ROJIlI'leCTBeHuoe llsY'lenllC ap;pena:II1Ha II
Hopap;penamma B 1l:t;:lnO'le'lIlIII;:ax KYPIlHOro BM6plIoHa.- ORJI. AH CCCP,
1959, 127, 934-937.
MaccoH n. OnyXOJIII qeJIOBeKa. M., JfSP;-BO «MeAul\HHa», 19G5.
JlaT6ee6 B. C. Rypc SOOJIOrllll. M., Y'lneArIl3, 1935.
-'laT6eee B. C. DOJIbmoll npaKTHKYM no SOOJIOrUH IIOSBOHO'lHIdX. M., JfSP;-BO
MocK. YH-Ta, 1963.
JlaT.luHa a. ill., MeHbLUUI>06 B. B. COBpeMeHHlde npep;CTaBJIeHHH 0 CIIHTese D
npeBpall\eHHII KaTeXOJIaMIIHOB B opraHlIsMe.- Yen. cOBpeM. 61I0JI., 1964,
58, 321-345.
JlaT.lUHa a. ill., MeHbLUUI>06 B. B. RmlHlI'leCKaH 61I0XllMIIH KaTeXOJIaMHHOB.
1\1., «Mep;rus», 1967.
Me:JICHUH (/J. N. lIccJIeAOBaUHH no rncTorerresy H rlleTOXIIMHH Hap;nO'le'lHHKa y
p;OMaIIlHUX IITIIl\. ABTopccJ>. KaHp;. P;I1cc. TapTY, 1970.
:1IUJlOXUH A. A. 06 acJ>cJ>epeHTHOll BHHepBal\UII IIII eBapHTeJIbnOrO TpaETa. C6.
«Bonpocld MopcJ>o;lOrIIH pel\enTOpOB BHYTpeHHIIX opranOB H Cepp;e'lHO-CO-
CYP;IICTOH CHCTeMId». M.- JI., IISP;-BO AH CCCP, 1953a, 77-97.
MU.WXUH A. A. R Bonpocy 0 coucTBeHHoii pecJ>JIeKTopHoii: P;yre KHme'lHHK8.-
ORJI. AH CCCP, 19530, 93, 933-935.
JIU.lOXUH A. A. CpaBHIITeJIbHO-rIlCTOJIOrH'leCRHe Ha6JIIOp;euHH Hap; acJ>cJ>epeHT'
HOll IIHHepBal\UeH nllll\eBapllTeJIbHOrO TpaKTa B PHAY nOSBOHO'lHld:X. Tpy-
;:lid VI Bcec. C'be3p;a aHaTOMOB, rHCTOJIOrOB, BM6pIIOJIOroB. XapbRoB, 19613,
1, 837-839. v v v
},[UJlOXUH A. A. MopcJ>OJIOrIIH CHHanTB'leCKIIX CBHsen B KHme'lHOII HepBHOII CD-
CTeMe RpymopOTIdX. C6. «MopcJ>OJIOrHH MemHeHpOH8JIbHldX CBHSeH». M.-
JI., IISP;-BO AH CCCP, 19616, 76-83.
MU.ltOXUH A. A. IIHTeppe enTopld nlI eBapHTeJIbHOrO TpaKT8 HeKOTopldX Hna-
IDIIX nOSBOnO'lHIdX (CpaBHIITeJIbHO-rHCTOJIOrB'leCKOe HCCJIep;OB8HlIe). M.-
JI., «HaYKa», 1963. v A
Mopoao6 E. K. R Bonpocy 06 IIHHepBa IIB cHHo-KapoTIIp;HOII pecJ>JIeKCOreHHo
30HId y nTnl\.- Apx. anaT., rIICTOJI., BM6pIIOJI., 1969, 56, 3, 35-38.
13 H. A. C HITTeH
317
Mopoa06 E. li. OTHomenlle H3IdHO-rJIOTO'lrl'Oro rrepBa K RapoTIlp;HoMy rJIOMycy
y nTHl\.- Apx. aHaT., rIICTOJI., BM6pHOJI., 1970, 58, 1, 79-82.
BaiJee6 (/J. n., Y.ltYM6el>06 9. r. HeRoTop-lde pe3YJIbTaTId rHCTOXIIMII'leCRo1'O
1I3Y'leHIIH aRTIIBHOCTH OI{11CJIHTeJIbnldX cJ>epMeHToB B RapoTIIp;HOM TeJIe KO-
meK.- Apx. aHaT., fIICTOJI., BM6plIOJI., 1969, 57, 8, 871-874.
HacoH06 /1.. H. [Nassonov D. ]. Das golgische Binnennetz und seine Beziehun-
ge!l der Sekretion. Untersuchungen iiber einige Amphibiendriisen.- Arch.
mlkrosk. Anat., 1923, 97, 136-186.
BacoH06 /1.. H. [Nassonov D.]. Das golgische Binnennetz und seine Beziehun-
gen zu der Sekretion F.ortzetz g). .M0l1!!lOlogische und experimentele
Untersuchungen an mmgen Saugetlerdrusen.- Arch. mikrosk. Anat.
1924, 100, 433-472. '
H06UI>06 A., 9ccHep 9., rO.ltb8ifJUwep C., Xeiiec M. (Novikoff A., Essner E., et
al.). HynJIe03HP;cJ>occJ>aTa3HaH aKTHBIIOCTb KJIeTO'lHIdX MeM6paH. C6. «Y JIbT-
pacTpYRTypa II cJ>YHHl\HH RJIeTI{II». M., «Mnp», 1965, 84-111.
lIuHec JI. H. [Pines L.]. Ober die Innervation des chromaffinen Gewebes des
Sympathicus und iiber rlas Sympathico-chromaffine System im allgemei-
nen.- Arch. Psychiat. u. Nervenkrankh., 1924, 70, 636-647.
llAe'll>o6a E. li. (Pep;.). C6. «MopcJ>OJIOrHH 'lYBcTBHTeJIbHoii: IIHHepBal\IIH BHYT-
peHHHX OpraHOB». M., 113p;. AMH CCCP, 1947.
IIoiJeopHaR r. A. AHp;peHaJIoBlde II Hopap;pCnaJIOBbJe KJIeTKH B M031'OBOIII Be-
ecTBe Hap;nO'le'lHIIROB JIa60paTopHldx rpld3YHoB II XR HIdX. C6. pa60T
JIeHHHrp. BeTepllHapHo1'O IIH-Ta, 1967B, 28, 683-694.
IIoHo tape6a B. H. Pa3BlITlle IIHHepBal\1I11 HapoTIlp;Horo rJIOMyca II BRcneplI-
MeHTaJIbHIdH aHaJIH3 npHpop;1d rJIOMYCHblX KJIeTOK. RaHp;. P;I1cc. M., 1946.
IIpuuMal> 9. X. Pacnpep;eJIeHlle ap;peHaJIllna II Hopap;peHaJIlIna B KJIeTKaX Ha.
pOTHp;Horo TeJIa.- ]:(ORJI. AH CCCP, 1959, 128,618-621.
IIpuuMal> 9. X. MCTO'lHIIKII pa3BIITIIH xpoMacJ>cJ>IIHHIdX KJIeTOI{ meHHo-rpYP;lIldx
naparaHrlIHeB nTHl\.- Apx. aHaT., rHCTOJI., BM6pIIOJI., 1960, 39, 12, 81-85.
IIpuuMaK 9. X. MopcJ>ocJ>ynKl\HoHaJIbulde oc06euHocTlI RJIeTOK meiiHo-rpyp;nldx
naparaHrJIIIeB B CBH3H C IIX p;1IcJ>cJ>epeHI{lIpOBKOH.- Apx. anaT., rHCTOJI., BM-
6PIIOJI., 1962, 43, 9, 66-75.
IIpuuMal> 9. X. 8JIeHTpOHHOMIIKpOCKOmlqeCKOe I1CCJIep;OBaHIIe HepBHo1'O 3a'laT-
Ra paHHHx CTap;I1H pa3BIITHH.- ]:(OKJI. AH CCCP, 1963, 148, 477-480.
IIpuuMal> 9. X. 8JIeRTpOHHaH MHKpOCKOIIIIH TOHRIIX MexaHII3MOB Bblp;eJIeHIIH
KaTeXOJIaMHHOB npIl CTHMyJIHl\HII ceKpeTopuo1'O npOl\ecca.- ]:(OKJI.
AH CCCP, 1965a, 162, 678, 680.
IIpuuMal> 9. X. Cy6MIIKpOCKOnIl'leCKaH CTPYI{Typa xpoMacJ>cJ>I1I1HIdX HJIeTOK H<lP;-
nO'le'lHHKa l<pblC B cJ>a3e BIdp;eJIenIlH Cel{peTa.- }J;OHJI. AH CCCP, 19656, 162,
1171-1174.
IIpuuMal> 9. X. YJIbTpacTpYRTypa xpoMacJ>cJ>HHHoH RJIeTRII B cJ>a3e BOCCTaHOB-
JIeHHH KaTeXOJIaMIIHOB.- ]:(OKJI. AH CCCP, 1966, 166, 1443-1446.
IIpuuMaK 9. X. a HeKoTopldX cJ>oPMax CeKpel\llH RaTeXOJIaMIIIIOB B IIIemKJIe-
TO'lHlde II cy6BHp;OTeJIlIaJIbHlde npocTpaHcTBa.- I1TOJIorHH, 1967,9,169-174.
IIpuuMaK 9. X., CMUTUH H. A. R Bonpocy 06 IICTO'lnIlKaX pa3BIITIIH ImpOTII;:{-
HOro TeJI8.- ]:(OHJI. AH CCCP, 1959, 129,457-460.
Po6epTuc 9., H06uHcKuu B., CaiJc (/J. (Robertis E., Nowinski W., Sacz F.).
EHOJIOrHH RJIeTRII. M., «MHp», 1967.
CaKOeU'l J/. B. MHHepBal\IIH 6pIOmrro1'O aOpTaJIbHo1'O naparaHrJIHH.- Apx.
anaT., rIICTOJI., BM6pHOJI., 1964, 46, 6, 50-55.
Cal>06U'l J/. B. B03pacTHaH MopcJ>OJIOrlIH II IIHHepBaI{IIH naparaHrJIHeB 'leJIOBe.
Ra II HeROTopldX II03BOnO'lnIdX. ABTopecJ>. p;OKT. p;IICC. M., 1969.
CanuH M. P. 0 BeHax Hap;nO'le'lHIIROB II HanpaBJIeHIIHX OTTOKa RpODII OT M03-
1'OB01'O Be eCTBa BT01'O opraHa.- Apx. aHaT., rIlCTOJI., BM6pIIOJI., 1961, 1i0,
6, 82-90.
CanuH M. P. MOPcJ>OJIOrll'leCKOe HCCJIep;OBaHIIe IIYTeH TpaHcnopTa 1'OpMoHon
M03roBo1'O BeTI{eCTB8 Hap;nO'le'lHHROB B HopMe II npIl rHnepTonn'lecrwII60-
JIe3HII. C6. «Ap;peHaJIHH II Hopap;peHaJIIIHI). M., «HaYKa», 1964, 233-235.
CeMaWKe6U'l r. M. a xapaRTepe RJIeTOK, BHep;pHIO HXCH B IIHTepp£'Ha.1fOBYIO 3<1-
RJI RY Hap;nO'le'lHHKa 'leJIOBe'leCKOrO BM6pIlOHa. B RH.: «C60PHlIK HaY'lHbIX
TJ>Y;JJ;OB POCTOB H/]:( Mep;. HH-Ta». POCTOB R/]:(., 1960, tIi, 526-529.
318
C"OPU11'WR B. l'rl., Ca'l.IW lJ. H. 1\ Bonpocy 0 CTpOCHlI1I TYJIOBII HOro OTp;c.'1a 110-
rpallH'lHOrO CHMlJaTII'leclwro C'fBOJIa pbI6.- Tpyp;1d TPOBf(IWro BeTCplIHap-
1I0ro HH-Ta, 1967, 12,47-49.
C.ItUpH06 A. [Smirnow A. S.]. Dic Structur der Nervenzcllen im Sympathicus
der Amphibicn.- Arch. mikro!'k. Anat., 1890, 35, 407-424.
C.ItUpH06 A. A. HapOTIlp;lIaH pCcJ>:'IeRCOreHHaH aOHa. JI., 113,q-BO «BoeHHo-Mop.-MeA.
aKap;.», 1945.
C.ItUpH06 A. A. HOBbIC p;alllIbIe R JllOpcJ>OJIOrHII lIepBHOro annapaTa cHHoKapoTIIp;-
HOll pecJ>JIeRCOrClillOll aOllbl.'- TpYP;1d V Bcec. C'bcap;a aHaTOJIIOB, rllCTO;JOrOB,
B1I10pHOJIOroB. JI., 1951, 641-643.
C.ItUpH06 A. A. HelwTopble cpaBHHTCJIbllo-aHaTOMH'lCClmC 1{aHHIdC K IIHHcpBa-
l\JIlI KapOTlIp;I1oii II aopTaJIbuoi't pccJ>JICRCOreHHldx aOH. C6. «HcpBuan pery-
.:JHl\HH KpoBo06paI.QClIII1I II p;bIXaHIIH». 1\1., 1952, 268-274.
C.1tUpH06a H. r. l1uUCpBaI{IIH KapoTIIp;Horo rJImlyca (glomus caroticum) n Ra-
pOTIlp;Horo clIHyca (sinus caroticus) y 'leJIOBeKa. C6. «MopcJ>oJIorllH 'lYBCT-
BIITeJIblIOll 1lHlIl'pBal\1II1 BHYTpeHHllx opraHOB». M., Uap;-Bo AMH CCCP,
1948, 106-125.
C.1tUTTeH H. A. UCTO'UmI> 06paaoBalilUI XpOJlIacJ>cJ>IIHIIbTX KJICTOR 6pIOillIl0-aOP-
Ta.'1bHbIX naparallr.'1l1cB.- Am..'1. AH CCCP, 1958. 120. 886-889.
G.ItUTTeIl H. A. I'HCTOXIIJllHql'CI.oe II 6110XIIMIIQeCKOC onpc;J;e;wuIIH Hopa;:lpeuaJIu-
lIa B HaI1ITO'leqHIIRaX 'lepITblX II 6eJIldx l<pbIC.- AOKJI. AH CCCP, 1960, 135,
728-731.
C.ItUTTeH H. A. Pa3BIITile opIOlimo-aOpTaJIbHbIX rraparalIrJIHCB II xpoMacJ>cJ>IIHHalI
peaRl\H1T y paaHOITHfJlICnTlIpOBanHbIx iKHBOTHbIX.- Tpyp;1d VI Bcec. C'beap;8
auaT., rnCTOJI., BMOpIlO.1.. 19tH, 2. 248-250.
CMllTTeH H. A. [Smitten N.]. Cytological analysis of thc origin of chromaffino-
blasts in !'ome mammals.- J. Embriol. expo Morphol., 1962, to, 152-166.
C.IlUTTeIt H. A. [Smitten N.]. Cytological analy!'is of catecholamine synthcsis in
the ontogcnesis of \"Crlcbratcs and problems of mclanogenesis.- Gen.,
Compo Endocrinol., HJ63, 2. 362-377.
C.!I/UTTeH H. A. CIIIlTea 1IIep;lIaTopoB CIIJlllTaTH'leCKOH HCpBHOH CIICTCJIIId II nllrMeH-
To06paaoBalIlle B OIlToreHeae nOaBOHO'IllbIX iI<HBOTHIdX.- mYPH. 06 eii
UIIOJI., 1964,25,51-61.
G.llUTTeH H. A. CTPYKTyplIaH lIa1llCU'lIlBOCTb MeJll6paHHldx CIICTCJII II cJ>1I31I0JIOru.
'lCCKaH aKTIIBHOCTb BIIP;ORplllIllOll KJICTRII. TeaHcbI II peI}JcpaTId P;OKJI. 4-ro
HaYQHOrO COBelI aHlI1I IJO BnOJIIOI{. I}Jn3110JIOrHII naMHTII JI. A. Op6eJIII. JI.,
1965a, 240-241.
C.ItUTTeH H. A. [Smitten N.]. Cytological and ultrastructural pattcrn of the sec-
retory activity of adrcnomcdullar ccll!'.- Arch. Anat. micro Morph. exp.,
19656, M, 145-162.
C.IlUTTeH H. A. YJIbTpaCTpYI.Typa :uI6pIIOUaJIbHOH KJICTKII JllOarOBoro CJIOH HaA-
nOQC'lHHKOB.- AOKJI. All CCCP, HJ66a, 168,447-450.
C. luTTeH H. A. 3JIeRTpOnuaH JllIlI<pOCKOIJIIH II l\IITOXHJlIHH rOpMOHaJIbllOH cJ>YHR-
I{HH naparanrJIIICB B BM6pllorencae. Co. «CTaHOBJICHlle Bnp;oRpHHHblX cJ>ynK-
l\IIH B aap01{ldmeBOJII paaBIITIIII». M., «HaYRa», 19660, 118-130.
G.'tuTTeH H. A. mCJIeallCTalI ro':THBHOCTb naparaHrJIIICB B BM6plIorcHcae. TealIcld
VII Bcec. C'beap;a aHaT., rnCTOJI., 3J116pIlOJI. T6HJIIICH, 1966B, 315.
CMuTTeH H. A. 3JICRTpOHHOJlIHKpOCKOIJIIQCCROe HCCJICp;OBaHHe ceRpcTopHbIX l\IIK.
JIOB B BM6pHorcneae ap;peUaJIOBOH KJIeTRH.- HTOJIOrI1H, 1967, 9, 162-168,
CMuTTeH H. A. [Smitten N. A.]. Fluorcscence catecholamincs in paraganglia.
(AORJI. Ha IX Memp;ynap. KOllrpecce aHaToMoB.). JI. 113;\-Bo BUHUTU, 1970,
1-4.
CMuTTeH H. A. COBpcMeHHoc COCTOHHHC Bonpoca 0 MOpcJ>OcJ>YIIKI\UoHaJILIDo1X
cBoi1:cTBaX meHno-rpyp;HIdX naparallrJIllcB. C6. «HepBHalI pcryJIlIl{H1I cJ>op-
JlI006paaOBaTeJIbnldX npol\CCCOB,). M., 1971a, 84-85.
CMuTTeH H. A. Il>IIJIOrCHCa ap;pcnaJIOBOH CIICTeJIIId B PHP;Y nOaBOHO'lHIdX mlIBooo.
IIIdX II HCROTOpblC acnCRTbi lIaTOJIOrH'leCKOH anaTOMIIII. l\IaTeplIaJIId V Be&-
COI03HOro C'be31{a naTOJIorOaHaTOMOB. M., 19716, 6-7.
CMuTTeH H. A., B.ltaiJoc-Ilpuii.llal> a. X. PaaBIITHc Hp;epHbIx II f(UTOnJIa3MaTB"I9-
CKIIX CTPYRTYP B RJICTKaX HepBHldx yaJIOB. TCSIICId p;Olm. ROOpp;BHal{JloBBo--
13* 319
to COBe anIlJI no np06JIellie «Y3JIOBlde BonpOCbI l\IITOJIOrllll,). JI., 1959,
145-147.
C.MUT7'eH H. A., Ma.ltUH06CI>aR JI. II. R Bonpocy 0 pa3BIITIIII xpoMacJ>cJ>IIHHbIX BJIe-
IIIeHTOB B 6pIOlliHOiI nOJIOCTII nTIIl\.- l];ORJI. All CCCP, 1960, 135, 445-448.
C.MUTTeH H. A., ByiJaHlfe6 A. 10. CDJIyopeCIH'IIUIIH RaTeXOJIaIIIIIHOB cHlImaTII'leCROfl:
II ap;peHaJIOBOiI CIlCTeM B 3apop;bIllieBoM pa3BIITIIII.- J];ORJI. AH CCCP, 1968,
183, 1225-1228.
COl>O.lt06 B. n. CynpapeHaJIOBaH meJIe3a y Rana temporaria H ce ce30HHbJe 113-
IIIeHeHIIH. B RH. «MaTepllaJIhI XI ROHcJ>. cTyp;eHToB II acnHpaHToB IIlOpcJ>OJI.
RacJ>ep;p II JIa60paTOpnii JIeHHHrp. ByaoB H HaY'lHD-IICCJIeA. HH-TOB, 17-
18 anpeJIH». JI., 1968, 48-49.
Cra6po6cI>UU E. M. BmIHHlIe HIICYJIHHa Ha cop;epmaliHe ap;peHaJIIlHa II Hopap;p9-
HaJIIIHa B HaAnO'le"IIIHIWX l{ypHHbIX BIIII)PHOHOB II l\bIllJIHT.- CDII3HOJI. mypH.
CCCP, 1964, 50, 365-372.
CTa6p06CI>UU E. M. CDYHHl\HH MoaroBoro Bel.QeCTBa Hap;nO"Ie'lHIIROB B OIiToreHe-
ae. ABTopecJ>. RaHA. p;IICC. JI., 1965.
CTa6p06CI>UU E. M. PacnpeACJI€Hlle aAPCHaJIIlHa II Hopam>eHaJIHHa B opraHax
6aJITHHCROH MilHorn Lampcrta fluviatilis B HopMe II nplI paaJIII'lHbIX cJ>YHR-
I{llOHaJIbHbIX COCTOHHlIHX.- mypH. BBOJIIOl\. 6110XHM. II cJ>1I3HOJI., 1967, 3,
216-221.
CY60po6a JI. B., RHoppe A. r. UCTOqHHRII paaBIITIIH IIHTpaMypaJILHIdX BereTa-
TIIBHbIX raHrJIHeB B BM6pHoreHeae n03BOnO'lHbIX.- Apx. aHaT., rliCTOJI.,
BM6pHOJI., 1960, 38, 1, 105-116.
TepeHTbe6 II. B. JIHrymRa. M., UaA-Bo «COBeTCKaH JIIIT-pa,), 1950.
TepeHTbe6 II. B. repneTOJIOrIm. 'TleHHe 0 aeMHOBop;HbIX II npecMIdRaIO lIXCH.
M., Uap;-Bo «BbIclliaH lliKOJIa», 1961.
TOl>UH n. B. COBpellleHHlde npep;CTaBJIeHlIH 0 CTpYKType II cJ>YHl{l\HflX ROMnJIeK-
ca rOJIbp;mll.- Apx. aliaT., rnCTOJI., BM6pHOJI., 1963, 45, 12, 1-22.
TpeTbRI>06 p. R. Moar neCIWpOIIRil. CI1o., TIm. CTaCIOJIeBH'lH, 1910.
TpeTbRI>06 p. R. OpraHbI "IYBCTB pe"InOH MIIIIorll. Op;ecca, THn. «TexnHR'), 1916.
TpeTbRI>06 p. R. [Tretjakoff D.}. Das periphere Nervensystem des Flu13neunau-
ges.- Z. wiss. Zool., 1927, 129,359-452.
J!XTOMCI>UU A. A. a HepBHo-rYMopaJILHLIX COOTHOITICHIIHX. B KH. A. A. Y X T 0 11I-
C K H H. Coop. CO'l., JI., 1951,2, 136-147.
X.ltOnUH H. r. 06 e6HOJIOrIl"IeClme II BI{CnepllIlIeHTaJIbHLIe OCHOBbI rHCTOJIOrnli.
JI., I13p;-no AH CCCP, 1946.
l(bl6y.ltbCI>UU H. O. a cJ>YHKI nll IIaAnoqe"IHOH meJIeabI.- BoeHHD-MeA. mypn.,
1896,186, 162-175.
'CJe60l>Cap06 M. H. a ceHpcTopnLlx HepBax HaAnO'le'lIIJnWB. 3KCnepIIMeIITaJIhHOe
nCCJIe)lOBaHHe. J];HCC. Ra3alIb, 1910.
Ye60l>Cap06 M. H. 06 OTp;eJllITe,lbIIIdX HepBax Hap;n01.le"IHHKOB.- PyccIHlii Bpa'I,
1909,8,26,873-875.
-rJe60l>CapOB M. H. a ceKpeTOpHldx HepBax Hap;nOqe'lHIIROB. M., «MeArH3'), HJ49.
qepHue06cI>uu B. H. I1cCJIeAOBaHlIe pel{enTOpOB HeKOTopblX BIIYTpeHHIIX opra-
HOB. Coo6 . 1.- CDHaHOJI. mypn. CCCP, 1940a, 29, 3-14; Co06 . 2.- Talll
me, 19406, 29, 15-25; Co06 . 3.- TaM me, 1940B, 29, 526-535.
'lepHue06CI>UU B. H. AcJ>cJ>epeHTHbIe CHCTeMbI BHYTpeHHHx opranoB. RHpOD.
Map;-Bo «BoeHHo-lIlOp. MeA. aRaA.'), 1\)43.
QepHue06C1>UU B. H. Pel\enTOpbI CepAC"IHo-COCYP;HCTOH CIlCTeMbI.- Yen. cOBpelll.
6110JI., 1947,23, 215-240.
QepHue06C1>UU B. H. UHTep0l{enTOpbI. M., «MeArn3», 1000.
Q3/Coy IJu-iJa, qaHb-MuHb-xyau. rIICTOXII!IIII'leCRHe HCCJIep;OBaHHH HaAnO'le'lHU-
ROB Rana nigromaculata HalloweIl.- Acta anat. sinica, 1965, 8, 2, 213-219.
qu",uHaiJae H. A. UHTpaoprallHoc pacnpCAeJIeHlle apTCpHaJIbHLIX COCYP;OB HaJ"{-
nO'le'lHIIKa. TpYP;bI I1n-Ta ,mcnepliM. MOpcJ>OJI. AH rpya. CCP, 1962a, 137-
138.
qu",uHaiJae H. A. 3RCTpaopraHlIbIe apTepHII Hap;nO'le'lHHKa.- TaM me, 196215.
148-151.
qu",rmaiJae H. A. MopcfJoJIorHH BeHoaHoH CIICTeMbI Hap;nOqe'lHOiI mCJIeald.-' Ta {
'-Ke,I962B,310--312.
320
UleHifJaUH A. A. COCYP;bI Hap;nOqeqmma II HX CBH3H.- TPyp;bl RHITIHHeBcK. roe.
Mep;. HH-Ta, 1949, 1,44-54.
Wu.)[I>e6u", B. Rypc cpaBHIITeJIbnoii aHaTOMl1II n03BOHO'UILIX iKIIBOTHIdX M.- IIr.,
HJ22.
W""a.ltbeayaeH 11. n. OpranH3M Iml, D;eJIOe B IIHp;uBIlp;yaJIbHOM II IICTOplI1JeCKOM
pa3BIITHII. M.- JI., 113p;-Bo AH CCCP, 1938.
mJota.ltM!ayaen n. n. IlYTII II 3aKoHoMepHocTII BBOJIIOl\1I0HHOro npo:o;ecca. M.- JI.,
I13P;-BO AH CCCP, HJ40.
mJ.ta.ltM!ayaen n. n. AYTOCTHJIIIH II npeo6pa30BaHHH BepxHHx OTp;eJIOB nepBblX
nneD;epaJibliLIX p;yr HII3lIIHX naaeMHblX n03BOHO'lHbIX.- 300JI. lliYPH., 1953,
32, 30-42.
my6un A. C. I1HHepBal\llH apTeplI0-BeH03Hldx aHaCTOM030B. M., «Mep;11I3»,
1952.
IOuu,e6u", H. JI. ROlllTIJIeKC rOJIbp;llili II MHTOXOHP;plIlI B ceRpeTOpHoM D;IIKJIe xpo-
lacJ>cJ>IIHHOH KJIeTRII B cJ>a3e BLlp;eJIeIUlH.- EIOJIJI. BKcneplIM. 61I0JI. MeA.,
1966a, 61, 3, 108-112.
IOw"e6u", H. JI. ROMnJIeRC rOJIbp;llilI II MIITOXOHP;pHII B ceupeTopHoM l\HRJIC xI'I-
lIIaq)(IJlIIIHOH KJIeTKH B cJ>a3e BOCCTanOBJICHIIH.- EIOJIJI. BKcnepHM. 6110JI. IIWj.:\.,
19666, 62, 12, 105-108.
H2.1t06 B. B. IITOJIOflI'IeCKOe II rHCTOXHMUqeCKOe HCCJIep;OBaHlIe ceKpeTopHoii
- p;eHTeJIbHOCTH xpoMacpcJ>HIIHblX KJIeTOK M03rOBoro Be CCTBa nap;n01Je1JHHKa.
ABTopecJ>. Kallp;. p;lICC. M., 1966a.
H2.1t06 B. B. rnCTOXlIMllqeCROe IICCJIep;OBaHHe ap;peHaJIHHa, nopap;peHaJIIIHa n
.. aCKop611HOBOH RIICJIOTId B Hap;nOqe'lHUKaX IwrneR npll HHCYJIliHOBOH KOMe.-
Apx. anaT., rnCTOJI., BM6pIIOJI., 19666,51, 10, 122-125.
HC60un r. B. a 3Ha'leHHII TaK na3blBaeMbIX TeMHblX RJIeTOR.- Yen. COBpeY.
6IIfJ:I.. 1939, 11, 430-441.
HC60un r. B. TelllHble n CBeTJIbIe RJIeTKII. M., 113p;-Bo AMH CCCP, 1948.
HXOHT06 R. M. R yqeHHIO 0 xpoMacJ>cpHHHoii CHCTeMe (p;06aBo'iRbIe opranbl co-
'lYBCTBeHHoro HepBa).- PYCCKHH Bpa'l, 1913a, 12, 527-529.
HXOHT06 R. M. R Y1JeIIlIIO 0 xpoMacJ>cJ>HHIwH CIlCTeMe. CTpoeHHe p;06aBo1JRbIX
opraHoB CHllmaTII1JeCIWrO IIepBa Y 1JeJIOBeKa.- Ra3aH. Mep;. iRYPH., 19136,
13.394-444; 463-507.
HX01tT06 R. M. R Y1JeRUIO 0 xpoMacpcJ>HHHoH CHCTeMe. R Bonpocy 0 cTpOeHIIJI
COHHOH llieJIe3bI Y 'leJIOBeKa.- Ra3aH. Mep;. lliYPH., 1914, 14, 137-149.
Abraham A. Die Innervation der Blutgefiisse.- Acta bioI. Acad. sci. Hung.,
1953, 4, 69-160.
Abraham A. Electron microscopic investigations on the human carotid body
(preliminary communication).- Z. mikrosk.-anat. Forsch., 1968, 79,
309-315.
Abraham A. Elektronenmikroskopische Untersuchungen an menschlichen Ka-
rotiskorpern.- Z. mikrosk.-anat. Forsch., 1970, 81, 413-453.
Adams W. E. The cervical region of the Lacertilia. A critical review of certain
aspects of its anatomy.- J. Anat. (Lond.), 1939, 74, 57-71.
A d1.ms W. E. Observations on the lacertilian sympathetic system.- J. Anat.
(Lond.), 1942, 77, 6-11.
Adams W. E. The carotid sinus complex and epithelial body of Varanus va-
rius.- Anat. Rec., 1952, U3, 1-27.
Adams W. E. The carotid arch in lizards with particular reference to the origin
uf the internal carotid artery.- J. Morphol., 1953, 92, 115-155.
Adams W. E. On the possible homologies of the occipital artery in mammals,
with some remarks on the philogeny and certain anomalies of the subcla-
vian and carotid arteries.- Acta Anat., 1957a, 29, 90-113.
Adams W. E. The carotid bifurcation in Chamaeleo.- Anat. Rec., 1957B, 12g,
651-663.
Adams W. E. The comparative morphology of the carotid body and carotid
sinus. Springfield USA, C. C. Thomas, 1958.
Adams W. E. The development of the adrenals and sympathetic paraganglia
in certain Madagaskar insectivores.- J. Embryol. expo Morphol., 1958a, 6,
575-585.
321
Adams W. E., Pickerscill K. W., Unu"rwood G. The drvelopment of the carotid'
arch in Spherodactylus argus Gosse, with some rrmarks on «Inselbildun
gen,> in the aortic arches.- J. Morphol., 1957, 101. 399-424.
Agduhr E. A., Hammar J. A. Demonstration showing «Wachsplatten- und glas-
rekonstruktionnen, zur Beleuchtung der friiheren embryonalen Entwick-
lung des Glomus carotidt'um».- Anat. Anz. Ergiinzungsh., 1932, 75.
233-234.
Al-Lami F. Light and electron microscopy of the adrenal medulla of l\Iacaca
mulata Monkey.- Anat. Rec., 1969, 164,317-332.
Al-Lami f'.. Murray R. G. Fine structure of the carotid body of normal and ano-
xis cats.- Anat. Rec., 1968a, 160,697-718.
Al-Lami F., Murray R. G. Fine structure of the carotid body of Macaca mulata
Monkey.- J. Ultrastruch. Res., 1968b, 24, 465-478.
Allen W. F. Distribution of the spinal nerves in polistotrema and some special
studies un the development of spinal nerves.- J. Compo Neurol., 1917, 28,
137-213.
Angulo y Gonz.:Jlez A. W. Hystogenesis of the monopolar neuroblast and the
ventral longitudinal path in the albino rat.- J. Compo Neurol., 1939, 71 (2),
325-360.
Arnold J. Ueber die Structur des Ganglion inlercaroticum.- Virchow's Arch,
1865, 33, 190-209.
Arnold M., Hager G. Funktionrntwicklung dl'r Nl'lwnniere beim Goldham-
ster.- Z. Zellforsch. mikrosk. Anat., 1967,83,117-132.
Arvy L. Histo-enzymologie des Glandes endocrines. Ganthier - Villars, Paris,
1963.
Arvy L., Bonichon M. Contribution a l'histo-enzymologie de Ia glandr surrenalc
chez Crocodilus niloticus Laur.- Z. Zellforsch. mikrosk. Anal., 1958, 48,
4 HI-435.
Ask-Upmark E. The carotid sinus and the cerebral circulation. An anatomical.
experimental and clinical investigation including some observations on
rete mirabile caroticum. Lund: Berlingska Boktryckeriet.- Acta psychiat.,
Copenhagen, suppl. 6,1935, 1-374.
A ugustinsson K. B., Pange R., Johnels A., Ostlund E. Histological, phisiologica I
and biochemical studies on the heart of two cyclostomI's, Hagfish (Mixinl')
and Lamprey (Lampetra).- J. PhysioI., 1956, 131, 257-276.
Axelrod J. The metabolism, storage and release of catecholamines. Recent Pro-
gress Hormone Res. N.- Y.-L., 1965,21, 597-622.
.Axetrod 1., Hertting G., Patrick R. Inhibition of H3-norepinephrine release by
monoamineoxidase inhibiLors.- J. Pharmacol. expo Therap., 1961, 13
325-328.
Ayers H. Amphioxine nerve in myxinoids.- J. Compo Neurol., 193U, 50, 469-478.
Bacnmann R. Die Nebenniere.- Handb. mikrosk. An.lt. Mensch. Berlin, Sprin-
ger. 1954,6. Teil 5, 1-952.
Baecker R. Uber die Nebennieren der Teleostier.- Z. mikrosk-anat. Forsch.,
1928. 15, 204-273.
Baker R. C., Graves G. O. The behaviour of the neural crest in the forebrain
region of Amblystoma.- J. Compo Neurol., 1939, 71 (2), 389-402.
Balfour F. M. The development of elasmobranch fishes.- J. Anat. PhysioL. 1877.
11. 406-490.
Balfour f'. M. Uber die Entwicklung und die morphologie der Suprarenalkor.
per (Nebennieren).- BioI. Zbl., 1881-1882, 1, 136-138.
Banerji T. K. Alteration in the adrenomedullary hormones in a teleostean fish
following rescrpine treatment.- Proc. zool. Soc. (Calcutta), 1970, 23, 73-
81.
Bander A. Uber zwei chromoffine ZeIItypen im Nebennierenmark und ihre Be-
ziehung zum Vorkommen von AdrenaIin und ArterenoI.- Arch. expo
Pathol., Pharm., 1954, 223, 2, 140-147.
Battaglia G. Ricerche ultrastrutturali sui glomo carotideo del ratto.- Arch. Ita!.
Anat. EmbrioI., 1968,73, 155-187.
Battaglia G. Osservazioni ultrastrutturali suI glomo carotid eo denervato di ral-
10.- Boll. Soc. !tal. BioI. Sperim (omaggio), 1970, 46, 20, 1-3.
322
Battaglia G., Mariani G. Osservazioni ultrastrutturali suI glomo aortico - pot-
monare di coniglio.- Bool. Soc. !tal. BioI. Sperim. (omaggio), 1970, 46,
:!O, 1-3.
BenecLeczky J. Electron-microscopic differentiation of cell types in the frog's
adrenal medulla.- Acta morphol. Acad. sci. Hung., 1967a, 15, 271-285.
BenecLeczky 1. Electron-histochemical localization of acid phosphatase acti-
vity in the adrenal medulla of rats.- Nature, 1967b, 214, 1243-1244.
Benedeczky I., Puppi A., Tigyi A. Histochemical and electron microscopical stu-
dy of the adrenal medulla of the Grass snake (Natrix natrix).- Acta bioI.
Acad. sci. Hung., 1965, 15,271-284.
Benedeczky J., Puppi A., Tigyi K., Lissak K. Electron microscopic study of adre-
naline and noradrenaline secretion of the adrenal medulla.- Acta bioI.
Acad. sci. Hung., 1965,15,285-298.
Bennett H. S. Cytological manifestations of secretion in the adrenal medulla
of the cat.- Amer. J. Anat., 1941,69,333-381.
Bennett H. S., Kilham L. The blood vessels of the adrenal gland of the adult
cat.- Anat. Rec., 1940, 77, 447-472.
Benoit A. Recherches sur l'origine et la signification du ganglion carotidien
(souris).- Arch. bioI. (Paris), 1928,38, 219-247.
Berchtold J. P. Mise en evidence histochimique d'adrenaline et de noradrenaline
dans Ie tissu chromaffine de Triturus marmoratus Latr. et de Salamandra
salamandra L. (Urodcles Salamandridae).- C. r. Acad. sci., 1966, ser. D263,
t8, 1337-1340.
Berkelbach v. H. Nebenniere und Paraganglion. Handb. vergl. Anat. Wirbeltie
reo Berlin - Viellne, eds. L. Bolk, E. Goppart, 1934, 2, Part, 777-816.
Bernard C. Lecones de pathologie experimentale. Paris, 1871. (PyCCK. nepeBop;,
f937) .
l'ertltr A., Hillarp N. .4.., Rosengren E. Some observation on the synthesis anrl
storage of catecholamines in the adrenaline-containing cells of the sup-
rarenal medulla. Ciba found. sympos. «Adrenergic mechanisms», 1960,
100-102.
Bianchi S. Sur la signification des neurones fluorescents chez la sangsue Hi-
rudo medicinalis.- Ann. Endocrinol., (Paris), 1969, 30, 545-548.
Biedl A. Innere Sekretion. 1913, teil I (313-534); tell II (1-82).
Biedl A., Wiesel J. Ober die funktionelle Bedeutung der Nebenorgane des Sym-
pathicus (Zuckerkandl) und der chromaffinen Zellgruppen.- Pflug. Arch.,
1902, 91, 434-461.
Biscardi A. M., Wassermann G. F. Release of catecholamines from adrenalino-
gcnic and noradrenolinogenic tissue by the action of nicotine <<in vitro».-
Arch. Intern. pharm. therap., 1966, 159,424-433.
Biscoe T. J., Lall A., Sampson S. R. On the nerve endings associated with the
carotid body glomus cells of the cat.- J. Physiol., 1969, 200, 619-620.
Biscoe T. J., Steh bens W. E. Electron microscopic observations on the carotid
body.- Nature, 1965,208,708-709.
Biscoe T. J., Stehbens W. E. Ultrastructure of the carotid body.- J. Cell BioI.,
1966, 30, 563-578.
Biscoe T. 1., StehbenB W. E. Ultrastructure uf the denervated carotid body.-
Quart. J. expo Physiol., 1967,52,31-36.
Blaschko H. The specific adion of I-dopa decarboxylase.- J. Physio!., 1939, 96,
50-51.
Blaschko ll., llagen P., Welch A. D. Observations on the intracellular granules
of the adrenal medulla.- J. Physiol., 1955, 129, 27-49.
Blaschko H., Hagen J. M., Hagen P. Mitochondria enzymes and chromaffin gra-
nules.- J. Physio!., 1957, 139,316-322.
Blessing M. Glomera im Suprakardialraum eines 4 Wochen alten Kindes.--
Klin. Wschr., 1963, 41, 1025-1026.
Bloom G., Ostlund E., Euler U. S. V., Lishajko F., Ritzen M., Adams-Ray 1.
Studies on catecholamine-containing granules of S r. ecific cells in cyclo-
stome hearts.- Acta physiol. Scand., 1961 53, Supp. 185.
Bliimcke S., Rode J., Niedorf H. R. The carotid body after oxygen deficiency.-
Z. Zellforsch. mikrosk. Anat., 1967, 80, 52-77.
323
BWmcke S., Niedorf H. R., Rode 1. Das Glomus caroticum nach CO 2 - Erho-
hung in der Atemluft.-- Z. ZeHforsch. mikrosk. Anat., 1967, 81. 557-570.
Boissezon de P. La trifurcation carotidienne et Ie corpuscule intercarotidien du
cheval.- Ann. d'anat. pathol. med. chir.,_ 1936, 13, 733-747.
Boi sezon de P. Le corpuscule carotidien contient - il des cellules chromaffi-
nes?- J. Med. (Bordeaux), 1939a, 116,341-342.
Boissezon de P. Le labirinthe carotidien de la grenouille rousse adulte.- Bull.
Soc. histoire. natur., Toulouse, 1939b, 73, 145-152.
Boissezon de P. La zone reIlexogi'>ne sinusale et Ie corpusculc carotidien.-
BioI. med. (Paris), 1943a, 33, 36-48.
Boissezon de P. Les apparcils de reglage de la circulation du corpu!'cule caroti-
dien.- Bull. d'histol. et de technol. microscopique, 1943b, 20, 136-140.
Boissezon de P. Les vaisseaux, les nerfs et les ganglions nerveux dan!' la struc-
ture du corpuscule carotidien.- Bull. d'histol. et de technol. microscopique_
1944, 21, 54-64.
Bok S. T. Das Ruckenmark I. Embryologic gleichzeitig Einbeitung zur Synapto-
logie.- Handb. mikrosk. Anat. Menschen, Berlin, Springer, 1928, 4, Teil 1.
478-482.
Bottir 1., Pribek L. Corpuscule paraganglionnaire dans l'orbite.- Ann. d'anat.
pathol. med. chir., 1935, 12, 227-228.
Bottazzi F. Untersuchungen iiber das viscerale Nervensystem del' Selachier.-
Ztsch. BioI., 1902, 25, 372-442.
Bouckaert J. J., Dautrebande L., Heymans C. Dussociation anatolllo-physiologi-
que des deux sensibilites du sinus carotidien: sensibilite a la prcssion et
scnsibiliLe chimique.- Ann. Physiol. Physicochim. BioI., 1 31, 7, 207-210.
Boyd J. D. The development and significance of the amphibian carotid body.--
J. Anat. (Lond.), 1933, 68, 154-155.
Boyd J. D. Development of the human carotid body.- J. Anat. (Lond.), 1934-
HJ35,69, 144-145.
Boyd J. D. Observations on the human carotid sinus and its nerve sllpply.-
Anat. Anz., 1937a, 84, 386-399.
Boyd J. D. The developmcnt of thc human carotid body.- Contr. Emhryol. Car-
neg. Inst., 1937b, 26, 1-31.
Boyd J. D. The nerve supply of the branchial arch arteries in Vipera herus.-
J. Anat. (Lond.), 1942, 76,248-257.
Boyd J. D. Origin, development and distribution of chromaffin cl'lls. Ciba found.
sympos. «Adrenergic Mechanismu!',). London, 1960, 63-82.
Brachet J. Biochemical cytology. 1957. N.-Y. Acad. Press Inc. (PYCCR. nepeBop;p
M., 196U).
Brandt W. Das Darmnervensystem yon Myxine glutinose.- Z. Anal. EnLw.
Gesch., 1922, 65, 284 ( HT. 110 l'\icol J., 1952).
Brauer.4. Entwicklung der Exkretionsorgane der Gymnophionen.- Zool. Jahrb..
Abt. Anal. Ontogen., 1902, 16, 133-139.
Brauer .4. A topographical and cytological study of the sympathetic nervous
components of the suprarenal of the chick embryo.- J. lnrphol.. 1932.
53, 277-325.
Brauer A. The differentiation of the sympathetic nervous components of the
chick suprarenal in chorio-allantoic grafts.- Anat. Rec., 1934, 50, 60-61.
Brundin J. Studies on the preaortal paraganglia of newborn rabbits.- Acta
physiol. Scand., 1966,70, SuppI. 290.
Burman S. O. The chemoreceptor system and its tumour-the chemodectoma.-
Surg. Gyn. Obst., 1956, 102,330-341.
Callingham B. .4., Mann M. Adrenalinc and noradrenaline content of the adre-
nal gland of the rat following depletion with reserpine.- Nature, 1958.
181, 423-424.
Callingham B. .4., Mann M. Depletion and replacement of the adrenaline amI
noradrenaline contents of the rat adrenal gland following treatment with.
reseprine.- Brit. J. Pharmacol., 1962, 18, 138-149.
Campenhout E., van. Contribution to the problem of the development of the
sympathetic nervous system.- J. expo Zool., 1930, 56, 295-320.
324
Caravita S., COSCkL L. Les cellules chromaffines du coeur de la lamproie (Lam-
petra zanandreai). Etude au microscope electronique avant et apres un
traitement a la reserpine.- Arch. bioI. (Paris), 1966, 77, 723-753.
. Carman J. B. The carotid labyrinth in Hyla aurea, with a note on that in
Leiopelma hochstetteri.- J. Anat. (Lond.), 1955, 89, 503-525.
Ca.spersson T. Studien iiber del' Eiweillumsaz der Zelle.- Naturwissenschaften,
HJ41, 29, 33-44.
Caspers son T. Cell growth and cell function. N.-Y., 1950.
Castro de F. Sur la structure et l'innervation de la grande intercarotidienne
(glomus caroticum) de l'homme et des mammiferes, et sur un nouveau sy-
steme d'innervation autonome du nerf glossoph'1ringien etudes anatomi-
quI's et I'xperimenlales.- Trab. Lab. Invest. bioI. Univ. Madr., 1926, 24,
365-432.
Castro de F. Sur la sLrul'lure et l'innervation du sinus carotidien de l'homme
et. des mammifi>res. Nouveaux faits sur l'innervation et la fonction du glo-
llIUS caroticum. Etudes anatomiques et physiologiques..- Trab. Lab. Invest.
bioI. Univ. ladr., 1928, 25, 331-380.
Castro de F. Uber die Slruktur und Innervation des Glomus caroticum bein
l\Ienschen und bei den Siiugetieren. Anatomischexperimentelle Untersu-
chungen.- Z. Anal. En!w. Gesch., 1929,89,250-265.
CoJ$tro de F. Sur la strucLure tip la synapse dans les cheIllocepteurs: leur meca-
nisme d'pxciLation l'l r61e dans la circulation sanguine locale.- Acta phy-
siol. Scand., 1951, 22, 14-43.
Celestino da Costa A. Origine et developpement de l'appareil surrenal et du sy-
slt'me nl'rveux sympathique chez les cheiroptcres.- l\1em. Soc. Portug.
Sci. Natur., 1917,4,1-103.
Celestino da Costa A. Noll' sur Ie developpement de la surrenale du herisson.--
C. r. Soc. bioI., 1920, 83, 878-879.
Celestino da Costa A. Le tissu paraganglionnaire.- Bull. d'histol. et de technol.
microscopique, 1926, 3. 10-25.
Celestino da Costa A. Sur les rapports entre les ebauches du corpuscule caro-
tidien l't sympathique cervical chez les cheiropteres.- C. r. Ass. Anat.,
30th reunion, 1935a, 90-94.
Celestino da Costa A. Paraganglia and carotid body.- J. Anat. (Lond.), 1935b,
69, 479-483.
Celestino da Costa A. bur ll's elements paraganglionnaires des embryons des
mammiferes.- C. r. Ass. Anat., 31st reunion, 1936a, 96-102.
Celestino da Costa A. Les paraganglions cervicaus des embryons de cheiropte-
res.- C. r. Soc. bioI.. 1936b, 122, 242-244.
Celestino da Costa A. Signification histologique et embryologique du corpusculc
carotidien.- C. r. Ass. Anal. reunion, Budapest, 1939, 30-47.
Celestino da CostoJ. A. Paraganglions et sympathique.- Ann. Endocrinol. (Pa-
ris), 1!JiG, 1,337-357, 4..j9-464.
Celestino da Costa .4. La notion de metaneurogonie.- C. r. Ass. Anat., 41st
reunion, 1955, 644-653.
Chakravorti B. P. Are Lhere chromaffin cells inside the ventricle of frog's
hearl?- Indian J. med. Sci., 1959, 13,311-314.
Chen I-Li. Yates R. D. Electron microscopic radioautographic studies of the
carotid body following injections of laheled biogenic amine precursors.-
J. Cell BioI., 1969,42, 794-803.
Chen I-Li, Yates R. D., Duncan D. The effects of reserpine and hypoxia on the
amines-storing granules of the hamster carotid body.- J. Cell BioI.,
1969,42,804--816.
Chester lones I. The adrenal cortex. Camhridge, University Press, 1957.
Chester Jones I. Adrenocorticosteroid. The hiology of myxine. Ed. A. Brodal a.
R. Fange, 1963, 488-502.
-Chevrel R. Sur l'anatomie du Systeme nerveux grand sympathique des elasmo-
hransches et des poissons osseux.- Arch. Zoo!. expo et gen., 1889, 5,
Suppl. 1.
325
Chevrel R. Recherches anatomiques sur Ie systeme neryeux grand sympathi-
que de l'esturgeon.- Arch. Zool. expo et gen., 1894, 2, Ser. 3, 401-444.
Paris.
Chiarini C. Observations sur les glomus carotidien et la glomus aortico-pulmo-
naire des mammiferes avec Ie microscope a la fluorescence. IX MelliI{YHa-
pOP;HbIH ROHrpecc aHaTOMOB. Te311CbI AORJlap;OB. 1970, 34. JIo. lJ:3p;. BlIHIITII.
Chiocchio S. R., Biscardi A. M., Tramezznni J. 1I. Catecholamines in the carotid
body of the cat.-- Nature, 1966,212.834-835.
Chowdhary D. S. The carotid labyrinth of Rana tigrina.- ;.\;ature, 1951, 167,
1074.
Chowdhary D. S. A comparative study of the carotid body and carotid sinus
of vertebrates, II. The carotid body and «carotid sinus» of the fowl (Gal-
lus domesticus). Doctorat.e thesis. Edinburgh, HJ53 58 pp. (Typescript).
( IIT. no Adams W. 1958).
Christie R. V. The function of the carotid gland (glomus caroticum). 1. The
action of extracts of a carotid gland tumor in man.- Endocrinology, 1933a,
17, 421-432.
Christie R. V. The function of the carotid gland (glomus caroticum). II. The
action of extracts of Lhe carotid gland of the elasmobranch.- Endocrinolo-
gy, 1933, 17, 433-444.
Ciaccio C. Rapporti ist.ogenetici tra il simpatico e Ie cellule cromaffini. Ricerche
istologichl'. Arch. Hal. Anal. Embriol., 1906, 5. 256-267. ( IIT. no Bach..
mann R.. 1954).
Clark A. J. Comparative physiology of the heart. Cambridge, University Press,
1927.
Clementi P., Zocche G. P. Morphological and pharmacological effects of reser-
pine, given alone or after iproniazid, on the catecholamines of the adrenal
glands of the rat.- J. Cell. BioL. 1963, 17,3.587-596.
Comroe J. H., jr. The peripheral chemoreceptors.- Handb. Physiol., 1964. 1.
Sec. 3, Respiration I, 557-583.
Comroe J. H., jr., Schmidt C. F. The part played by refle es from the carotid
body in the chemical regulation of respiration in the dog.- Amer. J. Phy-
sioL. 1938, 121, 75-97.
Coupland R. E. The prenatal development of the abdominal paraaortic bodies
in man.- J. Anal. (Lond.), 1952,86,357-372.
Coupland R. E. On the morphology and adrenaline-noradrenaline content of
chromaffin tissue.- J. EndocrinoI.. 1953,9, 194-203.
Coupland R. E. Post-natal fate of the abdominal paraaortic bodies in man.-
J. Anal. (Lond.), 1954.88,455-464.
Coupland R. E. The developmcnt and f8te of the abdominal chromaffin tissue
in the rabbit.- J. Anat. (Lond.), 1956, 9<J, 527-537.
Coupland R. E. The catecholamine content of the adrenal medulla of the rat
following reserpine - induced depletion.- J. Endocrinol., 1959, 18, 15 -
161.
Coupland R. E. The post-natal distribution of the abdominal chromaffin tis-
sue in the guinea-pig, mouse and white rat.- J. Anat. (Lond.), 1960, 94,
244-256.
Coupland R. E. The natural history of the chromaffin cell. Longmans, 1965a.
Coupland R. E. Electron microscopic observations on the structure of the rat
adrenal medulla. II. Xormal innervation.- J. Anal. (Lond.), 196;)b, 99.
255-272.
Coupland R. E., Pyper A. S., Hopwood D. A method for differentiating between
noradrenaline- and adrenalinestoring cells in the light and electron micro-
scope.- Nature, 1964, 201, 1240-1242.
Coupland R. E, Hopwood D. The mechanism of the differential staining reac
tion for adrenaline- and noradrenalinestoring granules in tissue fixed in
glularaldehyde.- J. Anat. (Lond.), 19G6, toO, 227-243.
Coupland R. E., Weakley B. S. Developing chromaffin tissue in the rabbit:
an electron microscopic study.- J. Anal (Lond.), 1968, t02, 3, 425-
455.
326
Coupland R. E., Weakley B. S. Electron microscopic observation on the adrenal
medulla and extra - adrenal chromaffin tissue of the postnatal rabbit.-'
J. Anat. (Lond.), 1970, 106,213-231.
Couvreur E., Duculty J. L'innervation cardiaque chez les crocodiliens.- Arch.
into Phy iol.. 1924,24, 104-111.
Cowdry E. The development of cytopla mic constituens of the nerve cell in thl1
chick.- Amer. J. Anat., 1914, 15, 389-419.
Crowder R. E. The development of the adrenal gland in man with special re-
ference to origin and ultimate location of cell types ad evidence in favor
of the «cell migration,) theory.- Contr. Emhryol. Carneg. Inst., 1957, 36,
H13-21O.
Dalton A. 1. A study of the Golgi material of hepatic and intestinal epithelial
cells with the electron microscope.- Z. ZeIlforsch. mikrosk. anat., 1952,
36. 522-5 0.
Dalton A. J. Golgi apparatus and secretion granules. In: «The Celli>. N.-Y.- L.,
1961,2, 603-619.
D:mielopolu D.. Manescu E. Reflexogene Zoncn der Carotis. I. Ahhandlung. Ein-
leitung.- Z. Ges. expo med., 1928, 63, 139-142.
Datta B.. Ghosh A. .\ dll'lI1o<'vtologicnl investigation of the avian adrenal me-
dulla.- Cytologia, 1965, 29, 226-232.
Dawson A. B. Histochemical evidence of early differentiation of the suprare-
nal gland of the chick.- - J. Morphol., 1953, 92, 579-595.
Detwiller S. R. Observations on the migration of neural crest cells, and upon
the development of tlw spinal ganglia ami vertebral arches in amblysto-
ma.- Amer. J. Anat.. 1937,61,63-94.
Dittus P. Interrenalsystelll und chromaffine Zellen in Lebensablauf von Ichthyo-
phis glutinosus L.- Z. wiss. Zool., 1936, 147, 459-512.
Dolezel S., KOl'al{;ik r., Kriska M. Monoamines in the glomus pulmonale.-
Ex}'erientia, 1968, 24, 442-443.
Douglas W. JJ'. The mechanism of release of catecholamines from the adrenal
medulla.- Pharmacol. Revs, HJ66, 18,471-480.
Driiner L. Deber die anatomischen Unterlagen del' Sinusreflexe Herings.-
Dtsch. med. Wschr., H125, 51, 559-560.
Duncan D. An electron microscop study of the embrionic neural tube and
notochord.- Techas. Repts. BioI. Med., 1957, 15(3), 367-377.
Duncan D., Yates R. {;hrastructure of the carotid body of the cat as revealed
by various fixatives and the use of reserpine.- Anat. Rec., 1967, 157,
667 -682.
Eade N. R. The distribution of catechol amines in gomogenates of thc bovine
medulla.- J. Physiol., 1958, 141, 183-192.
Ecker A. DeI' feinerL' Bau der Nebennieren beim Menschen und der vier 'Yir-
helthierklassen.- Braunschweig, 1846.
Ecker A., Wiedersheim R. Anatomic des Frosches auf Grund eigener Unter-
suchungen durchaus neue Bearbeitung von E. Gaupp. Abt. 2. Lehre vom
Nerven und Gefiissystem, 1899.
Ehinger B., Falck B., Myhrberg H. E. Biogenic monoamines in Hirudo medi-
cinalis.- Histochemie, 1968, 15, 140-149.
Elfvin L. G. The fine structure of the cell surface of chromaffine cells in the
rat adrenal meduIla.- J. Ultrastruct. Res., 1965, 12, 263-286.
Elfvin L. G. A new granule containing nerve cell in the inferior mesenteric
ganglion of the rahbit.- J. Ultrastruct. Rl's., 1968, 22, 37-44.
Elliott T. R. The innervation of the adrenal glands.- J. Physiol., 1913 46
285--290. ' ,
Enemar A., Falck B., Nakanson R. Observations on the appearance of norepi-
nephrine in the symvathetic nervous ystem of the chick embryo.- De-
velopm. BioI., 1965, 11,268-283.
Engstrom H. On the double innervation on the sensory el}ithelia of the in-
ner ear.- Acta Otolar., 1958, 48, 109-118.
Eriinko O. Histochemical evidence of the presence of acid-phosphatase-
positive and negative cell islets in the adrenal medulla of the rat.- Na-
ture, 1951, 168, 250-251.
327
E:riinko O. Distribution of adrenalin and noradrenalin in the adrenal medul-
la.- Nature, 1955a, 115, 88-89.
Eriinko O. Effect of insulin on chromaffin reaction fluorescing islets and cat-
hochol amines in thc adrenal medulla of the rat.- Acta pathoi. mil' 1'0-
bioI. Scand., 1955B, 36, 219-223.
Eriinko O. Histochemistry of noradrenaline in the adrenal medulla of rats
and mice.- Endocrinology, 1955c, 57, 363-368.
Eriinko O. Radioautographic demonstration of noradrenalin and adrenalin in
the adrenal medulla.- J. Histochem., Cytochem.. 1957, 5, 408-4.13.
Eriinko 0., Hauninen L. Electron microscopic observations on the adrenal me-
dulla of the rat.- Acta pathol. microbiol. Scand., 1960, 50, 125-1::J2.
Eriinko 0., Hiirkonen M. Noradrcnaline and acetylcholinesterase in sympathe-
tic ganglion cells of the rat.- Acta physiol. Scand., 1964, 61, 299-300.
Eriinko 0., Hopsu V. Effect of reserpine on the histochemistry and content of
adrcDalin and noradrenalin in the adrenal medulla of the rat and mou-
se.- Endocrinology, 1958, 62, 15-23.
Eriinko 0., Hopsu V. Distribution conccntration of adrenalin and noradrena-
lin in the adrenal medulla of the rat following reserpine indused deple-
tion.- Acta physiol. Scand., 1961, 51, 239-246.
Eriinko 0., Hopsu V., Riiisiinen L. Effect of prolonged administration of nico-
tine on 1he medullary volume and the distribution of noradrenaline in
the adrcnals of thc rat.- Endocrinology, 1959a, 65, 293-297.
Eriinko 0., Hopsu V., Riiisiinen L. Changes induced by prolonged administra-
tion of nicotinc in the distributions of cholincsterases and acid phospha-
tase in the adrenal medulla of the rat.- J. NeUfochem., 1959b, 4., 332-
337.
Eriinko 0., Palkama A. Differential staining of the adrenaline and noradrena-
line - containil!J adrenomedullary cells.- Acta pathol. microhiol. Scand.,
1959, 47, 357-363.
Eriinko 0., Riiisiinen L. Cytophotometric study on the chromaffin rcaction the
iodate reaction and formalin induced fluorescence as indicators of adre-
naline concentracion in the adrcnal medulla.- J. Histochem., Cytochem.,_
1961, 1, 54-58.
Essner E., Novikoff A. Cytological studies on two functional hepatomas (In-
terrelation of endoplasmic reticulum Golgi apparatus and lysosomes).-
J. Cell. BioI., 1962, 15, 289-312.
Euler U. S. v. Noradrenaline. Springfield, Thomas, USA, 1956.
Euler U. S. v. Adrcnergic ncurogormones. In: Comparative Endocrinology,
_ N.-Y.- London, Acad. Press, 1963, 2, 209-238.
Eyzaguirre C., Koyano H. Effects of hypoxia hypercapnia and pH on the che-
moreceptor activity of the carotid body in vitro.- J. Physiol., 1965a, 178,
385-409. .
Eyzaguirre C., Koyano II. Thc effects of some pharmacological agents on che-
moreceptor discharges.- J. Physiol., 1965b, 178,410-437.
Eyzaguirre C., Koyano H. Effects of electrical stimulation on the frequency
of chemoreceptor discharges.- J. Physiol., 1oo5c, 178,438-462.
Eyzaguirre C., Koyano H., Taylor J. R. Presence of acetylcholine and trans-
mitter release from carotid body chemoreceptors.- J. Physiol., 1965d,_
178, 463-476.
Falck B. Observations on the possibilitics of the cellular localization of mo-
noamines by a fluorescence method.- Acta Physiol. Scand., 1962, 56,
Suppl. 197.
Falck B., Ow man Ch. A detailed methodological description of the fluoresce'}-
ce method for the cellular dcmonstration of biogenic monoaminl's.- Acla
Univ. Lundensis, 1965, Sec. 2, 7, 1-23.
Falck B., Hiiggendal J., Owman Gh. The localozation of adrena\inl' in adrener-
gic nervcs.- Quart. J. EXI)tl. PhysioI., 1963, 4.8, 253-257.
Fancello O. Interrl'ne, surreni c ciclo sessuale nei selaci ovil'ari.- Pllhbl.
Staz. Zool. Napoli, 1937, 16, 80-88.
Fedele M. Sulla innervazione del labirinlo carotiedo nei halraci.- R. C. Acad.
Ital., 1941, Ser. 7, 3, 159-173.
it'8
Fedile JI. La innervazione del tronco arterioso nei batraci anuri.- Arch.
ital. Anat. Embriol., 1943, 48, 84-121.
Fillenz M., Woods R. Some observations on the rabbit carotid body.- J. PhysioL
1966, 186, 39P--40P.
Foxon G. E. N. The mode of action of the heart in the frog.- New. Bioi..
1952, 12, 113-127.
Francis /!'. T. B. The anatomy of the salamander. Oxford, 1934.
Froriep A. Die Entwirklung und Bau des autonomen Nervensystems.- Metl.-
naturw. Arch., Hl07, 1, 301-321.
Gabe M., Martoja M. Contribution a l'histologie de la glande surrcnale des
squamata.- Arch. Anat. micro:;c., Morphol. exptl., 1961, 50, 1-34.
Gabe .1/., Martoja J1. Contribution a l'histologie de la glande surrenale des
cheloniens (reptilens).-Arch. Anat. microsc., Morphol. exptl., 1962, 51.
107-128.
Gabe M., Martoja M., Saint-Girons H. Etat actuel des connaissances sur la
glande surrenale des reptilcs.- Annce bioI., 1964, 3, 303-376.
Gabe M., Rancurel P. Contribution a l'histologie de la glande surrenale de
crocodylus niloticus Laur.- Arch. Anat. microsc., Morpho!. exptl., 1964,
53, 225-240.
Gabe M., Saint-Girons H. Particularites histologiques de la glande surrenale
chez sphenodon punctatus gray (reptule rhynehocephale).- C. r. Acad.
sci., 1964,258, 3559-3562.
Gannou B. J., Bumstock G. Excitatory adrener:gic inncrvation of the fish he
art.- Compo Biochem. Physiol., 1969,29 765-773.
Garner C. M., Dunca' D. Observations of the fine structure of the carotid bo-
dy.- Anat. Rec., 1958, 130,691-700.
Gaskell 1. F. The distribution and physiological action of the suprarenal m
dullary tissue in Petl'omyzon fluviatilis.- J. Physiol., 1912, 44, 59-67.
Gaskell J. F. The chromaffine system of annelidis and the relation of the sy-
stem to the contractile vascular system in the leech, Hirudo medicina-
lis.- Phil. trans. Roy. Soc. (Lond), 1914, Ser. B, 205, 153-211.
Gastun J., Douglas W. Appearance of catechol amines during development of
Rana pipiens.- Developm. BioI., 1962, 5, 468-481.
Gastpar H. Die Tumoren des Glomus carolicum, Glomus jugulare, Glomus
thympanicum und Glomus vagale. Stokholm, 1961.
Gaupp E. Anatomie des Frosches. 1899.
Gerard P. Sur l'existence d'un dispositif glomique dans l'aorte de certains te-
Ieosteens.- Arch. bioi. (Paris), 1943,54, 75-92.
Ghosh A. A comparative study of the histochemistry of the avian adrenals.-
Gen. Compo Endocrinol. Suppl., 1962, 1, 75-80.
Ghosh A., Datta B. Effect of reserpine on the release of adrenomedullary ca
techolaminl's and their role in glycemic and pressor responses in two avi-
an specics.- Gen. compo Enuocrinol., Suppl. 2, 1969, 354-357.
Ghosh I., Ghosh A. A comparative cytochemical study on the avian suprare-
nal medulla.- Proc. Zool. Soc. (India), HJ62, 15, 135-149.
Ghosh I., Ghosh A. InHuencp of age on cytochemically demonstrable avian
cotecho] hormonps.- Ztsch. Bioi., 1964, 114, 400-404.
Giacomini E. Sopra la fine struttura delle capsule sUITenali degli Anfibii.-
Monitore Zool. ital., 1898a, 9, 86-91.
Giacomini E. Sur les terminaisons nerve uses dans les capsules sUITenales "Ies
oiseaux.- Arch. ital. bioI., 1898b, 29, 482-483.
. Giacomini E. Contributo alIa conoscenza delle capsule surrenali nei Ciclosto-
mi, Sulla capsule sUl'renali dei Petromizonti.- Monitore Zool. ital., 1902,
13, 143-162.
Giacomini E. Sulla esistenza dell a sostanza midollare nelle capsule surrenali
dei Teleostei.- Monitore Zool. Hal., 1902b, 13, 183-189.
Giacomini E. Contributo alIa conoscenza delle capsule sUITenali dei Ganoidi e
particularmente Sull'esistenza delle coro sostanza midollare.- Monitorc
Zool. ital., 1904, 15, 19-32.
Giacomini E. Contributo alIa conoscenza del sistema delle capsule surrl'nali
dei Teleostei. Sulla sostanza midollare (organi soprarenali 0 tessuto cro-
329
maffine) di Amiurus catus L.- Rend. Accad. Sci., CI. Sri, Fis., natur. Ser.,
(1st. Bologna). 1904-1905,9, 183-189.
Giacomini E. II sistema interrenale e il sistema cromaffine (sistema feocro-
mo) nelle anguiIle adulte, nelle cienche e nei leptocefali.- Mem. Rend.
Accad. Sci. (1st. Bologna), Cl. Sri, Fis., 19U8, Ser. 6, 5, 407-441.
Giacomini E. II sistema interrenale e il sistema cromaffine (sistema feocro-
mo) in altre specie di Murenoidi.- Rend. Accad. Sci., Cl. Sci. Fis., n.l-
tur. Ser., (1st. Bologna), 1908-1909, 13, 87-98.
Giacomini E. II sistema interrenale e il sistema cromaffine (sistema feocro-
mo) in altre specie di Murenoidi. Ml'm. II. Parte I. Mem. Rend. Accad.
Sci. (1st. Bologna), CI. Sci. Fis., 1909, Ser. 6, 6, 415-456.
Giacomini E. II sistema interrenale e il sistema cromaffine (sistema feoc-
romo) in alcune specie di Teleostei con rene cefalico (pronephros) per-
sistente. Caratteri differenziali fra interrenale anteriore e corpus coli di
Stannius. Cenno sullo sviluppo diquesti ol1gani nei Salmonidi.- Rent!.
Accad. Sci., Cl. Sci. Fis., natur. ser., (1st. Bologna). 1909-1910, 14, 86-
103.
Giacomini E. II sistema interrenale e iI sistema cromaffin(' (sistema feocro-
mo) in altre specie di Murenoidi Mem. II. Parte 2. Mem. Rend. Accad.
Sci. (1st. Bologna), CI. Sci. Fis., 1910, Ser. 6, 7, 373-410.
Giacomini E. Anatomia microscopic a e sviluppo del sistema interrenale e del
sistema cromaffine (sistema fl'ocromo) dei Salmonidi.- Rend. Accad..
Sci., CI. Sci. Fis., natur. Ser., (1st. Bologna), 1910-1911, 15, 107-108.
Giacomini E. Anatomia microscopia e sviluppo r(1'1 sistema interrenale e del
sistema cromaffine (sist{'ma feocromo) dei Salmonidi. PurteI: Anatomia
microscopica.- Mem. Rend. Accad. Sci. (1st. Bologna), CI. Sci. Fis., 1911,
Ser. 6,8, 367-387.
Giacomini E. SuI sistema interrenale e suI sistema chromaffine di alcuni Te-
leostei abissali (Argyropelecus e Scopelus).- Rend. Accad. Sci., Cl. Sci.
Fis., natur. Ser. (1st. Bologna), 1920-1921,23, 130-t35.
Giordano-Lanza G. Ricerche istologiche ed istochimiche sulla midollare dell
surrene nel corso dello sviluppo emhrionale. «Quaderni di Analomia Pra-
tica», 1965, Ser. 21, N 1-4, 1-41.
Glees P. DeI' periphere und zentrall' Anteil des sympathischen N ervensystems
der Selachier. Utrecht, N. V. A. Oosthoek, 1940.
Goodrich E. S. Studies on the stmcture and development of vertehrates. N.-
Y.- London, 1958, 1, Chapt. 1-8; 2, Chapt. 9-14.
Goormaghtigh N. Organogenese et histogenese de la capsule surren:\le el de
plexus coeliaque chez les oiseauxet chez les mammifercs.- Ann. So!:.
med. gand., 1914, 5, 24-46.
Goormaghtigh N. Organogencse et histogenes(' de la capsule surrenale el du
plexus coeliaque.- Arch. bioI., (Paris), 1921, 31, 83-172.
Goormaghtigh N. Le probleme de l'hypertension arterielle considere du point
de vue anatome phthologique.- Ann. Soc. Sci. med. nat. Drux., 1930,72-90.
Goormaghtigh N. Sur l'existence de paraganglions vagaux.- C. r. Soc. hiol.,
1935, 120, 1348-1350.
Goormaghtigh N. On the existence of abdominal vagal paraganglia in thl?
adult mouse.- J. Anat. (Lond.), 1936, 71, 7;-90.
Goormaghtigh N., Pannier R. Contribution a la localisation anatomique de la
zone vasosensible de la region cardioaortique.- C. r. Soc. bioI., 1936,121,
882-884.
Goormaghtigh N., Pannier R. Les paraganglions du coeur et des zones vaso-
sensibles carotidienne et cardio - aortique chez Ie chat adulte.- Arch.
bioI. (Paris), 1939, 50, 455-533.
Gorbman A., Bern H. A textbook of comparative endocrinology. N.-Y.- Lon-
don, 1962.
Gottschau M. Structur und embryonale Entwicklung dcI' Nebennieren bei Sa-
ugetieren.- Arch. Anat. Physiol., L i z., 1883, 535-536.
Gouder B. Y. M., Desai R. N. Studies 0 the carotid body in the frog Rana tig-
rina Daud.- Naturwissenschaften, 1966, 53, 535-536.
330
Grignon G., Hatier R., Guedenet J. C., Dollander A. Aspects ultrastructuraux
dp la glande surrenale au cours du developpement chpz l'embryon de pou-
let.- C. r. Soc. bioI., 1966(1967), 160, 1654-1657.
Gray E. G., Guille,.y R, W. Synaptic. morphology in the normal and degenera-
tiIll5 nervous system. In: "Intern. Rev. Cytology". N.-Y., Acad. Press Inc.
HJ66, 19, 111-182. '
Grimley P. M.. Glenner G. G. Ultrastructure of the human carotid body.-
Circulation. 1968,37,64.8-665.
Groschuff K. Dl'merkungen zu der vorlaufigen Mitteilung von Jacoby: ueber
die Entwicklung der Nehendriisen der Schildrtise und der Carotidendrti-
se.- Anat. Anz., 1896, 12, 497-512.
Grynfeltt E. Structure dps corps suprarenaux des plagiostomes,- C. r. Acad.
f;ci., 1902a, 135, 373-374.
Grynteltt E. Sur Ie corps interrenal des plagiostomes.- C. r. Acad. Sci., 1902b,
135, 439-441.
Grynfeltt E. Les organes chromaffines. MontpelIier rued., 1903a.
Grynfeltt E. Sur la capsule sUlTl'nale des amphibiens.- C. r. Acad. Sci., 1903b,
137, 77-79.
Grynfeltt E. Sur la capsule surrenale des amphibiens. Paris. 1904a.
Grynfeltt E. Notes histologie sur la capsule sUITenale des amphibiens.- J.
Anat. PhysioI., 1904b, -10, 180-220, 425-465.
Hagen P. Thp distribution of adrl'naline and noradrenaline in ox adrenal me-
dulla.- J. Physiol., 1954, 123, 53-54.
Hagen P., Barrnett R. J. The storage of amines in the chromaffin cells. Ciba
found. sympos. «Adrenergic Mechanisms», 1960, 83-99.
Hall B. K., llnghes H. P. Response of host embryonic chicks to grafts of ad-
ditional adrenal glands. II. Ultrastructure of normal and host adrenal
Iglands.- Z. Zellforsch. mikrosk. Anat. 1970, 108, 1-16.
Hamburger V. Life history of a nerve cell.- Amer. Sci., 1957, 45, 263-277.
/lammar J. A. A quelle epoque de la vie foetale de l'homme apparaissent les
premiers signes d'une activite endocrine?- Upsala Liikareforen forhandl.,
1925,30, 375-480.
llammar J. A. Konstitutionsanatomische Studien tiber die Neurotisierung des
Menschenembryos. I. Zur Bildungsgeschichte des Glomus caroticum.- Z.
mikrosk.-anat. Forsch., 1934, 35, 602-630.
Hammond W. S., Yntema C. L. De r letions in the thoraco-lumbar sympathe-
tic system following removal 0 ncural crest in the chick.- J. Compo Neu-
rol., 1947, 6, 37-265.
Hanke W., Weber K. Histophysiological investigation on the zonation activi-
ty, and mode of secrction of the adrenal gland of the frog, Rana tempo-
I'aria Linnacus.- Gen. Compo Endocrinol., 1965,5,444-455.
Harde$ty I. On the development and nature of the neuroglia.- Amer. J. Anat.,
1904,3, 230-268.
Harman M. T., Derbyshire N. C. The development of the suprarenal glands in
the guinea-pig (Cavia Cobaya).- Amer. J. Anat., 1932, -19. 335-347.
Harrison R. G. Kl'ue Versuche und BeobachtuIllgen iiber die Entwicklung des
peripheren Nerven del' Wirbeltiere. Bonn, Sitzgsber. d. Niederrhein. Ges.
Nat. Heilkunde, 1904.
Harrison R. G. Furthes experiments on the developmcnt of peripheral ner-
ves.- Amer. J. Anat., 1907,5, 121-131.
Harrison R. G. Thl' adI'l'nal circulation in the rabbit.- J. EndocrinoL 1957, 15,
64-71.
Harrison R. G., lIoey M. J. The adrenal circulation. Oxford, Blackwell,1960.
Hartman F. A., Brownell K. A. The Adrenal Gland. London, Henry Kimpton,
1949.
lIartman F. A., Knoff R. A., ]',fcNntt A. W., Carver 1. E, Chromaffin patterns
in bird adrenals.- Anat. Rec., 1947, 97, 211-221.
Hattori H. Morphologische und entwicklungsgeschichtliche Untersuchungen
der Schildrtisp, dcI' EpitplkOrperchen, der Thymus, des postbranchialen
Kiirpel's und dcI' Carotisdruse von Hynobius nebulosus (japanisch). Japan.
J. med. sci., 1934, 5, 77-78.
231
Hays V. J. The development of the adrenals in birds.- Anat. Rec., 1914, 8,
451-473.
Hebard W. B., Charipper H. A. A comparative study of the morphology and
histochemistry of the reptilian adrenal gland.- Zoologica (USA), 1955,
40, 101-123.
Heinleth von. Beitrag zur Histologie des Perithelioma glandulae caroticae.-
Zbl. allg. Path., path. Anat., 1900, 11, 599-603.
Held H. Die Entwicklung des Nervengewebes bei den Wirbeltieren. Leipzi6'.
1909.
Hempel K., Kraft M. Zum Brenzkatechinamin-Stoffwechsell der Nebenniere.
Licht- und elektronenmikroskopisch-autoradiographische Untersuchungen
mit H3-Dopa Miiusen.- Z. ZeIlforsch. mikrosk. Anat. 1968, 85, 322-
342.
Henle 1. Obcr das Gewebe del' Nebennieren und der Hypophyse.- Ztschr. rat.
Med., 1865,24, 143-152.
Hering H. E. Der Sinus caroticus an dcI' Ursprungsstelle lieI' Carotis interna
als Ausgangsort eines hemmenden Herzreflexes und eines depressorischen
Gefiissreflexes.- Miinch. Med. Wschr., 1924, 71, 701-704.
Hering H. E. Die Karotissinusreflexe auf Herz und Gefiisse. Dresden - Leip-
zig, Th. Steinkopff, 1927.
Helant M. Le rOle de l'appareil de Golgi dans la secretion glandulaire.-
BioI. merl., 1964, 53, 593-619.
Hertl Ill. Zum Nicleolus-Problem.- Z. Zellforsch. mikrosk. Anat., 1957, 46,
18-51.
Hess A. The experimental embryology of the foetal nl'l'VOUS system.- BioI.
rev., 1957,32, 231-260.
Hett V. Ein Beitrag zur Histogenese der menschlichen Nebenniere. -Z. mik-
rosk.-anat. Forsch., 1925, 3, 179-282.
Heynzans C. (XeihraHc C.). qYBcTBIITe!lblllde II pecJ>JleJ{corellll1J cocyglIcTlde
30HId aopT'b1 II sinus caroticus, IiX 3Ua'If'IfIfl' B !f>1I3I10JIOI"IIII, naTOJlOflIlI n
cJ>apIllaROJIOrHlI.- Yen. cOBpeIII. 6IIOJI., 1934, 3, 3(j2-372.
Heymans C. ,Bouckaert J. J. Sinus caroticus and respiratory reflexes.- J.
Physiol., 1930, 69, 254-266.
Heymuns C., 130uckaert J., Dautrebande L. Wcitere Untersuchungen libel' die
Blutdruckziiglel' und die reflektorische Atmungsrelation durch innere
Druck- und chemische Reize.- Pflug. Arch., 1932, 230, 283-298.
Heymans C., Bouckaert J., Regniers P. Le sinus carotidien ct la zone homolo-
gie cardio-aortique... Paris, 1933.
Heymuns C., Neil E. Reflexogenic areas of the cardiovascular system. London,
Churchill LTD, 1958.
Heymuns J. F., Heymans C. Action stimulante reflexe de l'anemie sur la res-
piration de la tCte «isoli(») du chien.- C. r. Soc. bioi, 1926, 94, 1255-1257.
Hillarp N. A. Isolation and somc biochemical properties of the catechol ami-
ne granules in the cow adrcnal medulla.- Acta physiol. Scand., 1958,43,
82-96.
l/illarp N. A., Hokfelt B. Evidence of allrenalin and noradrenallin in separate
adrcnal medullary cells.- Acta PhysioI. Scand., 1953, 30, 55-6B.
Hillarp N. A., Hokfelt B. Citological demonstration of noradrenalin in the sup-
rarenal medulla under condition of varied secretory activity.- Endocri-
nology, 1954, 55, 255-260.
Hillarp N. A., /Iokfelt B. Histochemical dl'monstration of noradrenaline an-(
adrenaline in the adrcnal medulla.- J. Histochem. Cytochem., 1955, 3,
1-5.
Hillarp N. A., Nilson B., [[okfelt B. The cytology f adrenal medullary ?ells
with special reference to the storage and secretion of the sympathomlmL
tic ilmines.- Acta Allat., 1954, 21, 155-167.
Hirsch G. C. The external secretion of the pancreas as a whole and the com-
munication between the endoplasmic reticulum and the Goigi bodies. In:
«Biological Structure and Function». Lood.- _-Y., Acad. Press. Inc.
1960, 1, 195.
332
Hirt A. Die vergleichende Anatomie des Sympathischen Nervensystems.-
Handb. vergl. Anat. WirbeItiere, Berlin-Vienna, eds. L. Bolk, E, Gop-
past, 1934, 2, Part. 1, 685-777.
His W. Die Neuroblasten und deren Entsetzung im embryonal en Mark.-
Arch. Anat. Physiol., Anat. Abt. Lpz., 1889, 249-300.
His W. Histogenese und Zusammenhang der Nervenelemente.- Arch. Anat.
Physiol., Anat. Abt, Lpz., Sup pl. B, 1890, 95-117.
His W., jun. Die Entwickelung des Bauchsympathicus beim Huhnchen und
Menschen.- Arch. Anat. Physiol. Anat. Abt. Lpz., Suppl. B, 1897, 137-170
Hoffmann C. K. Znr Entwicklungsgeschichte des Sympathicus bei den Urode-
len.- Z. wiss. Zool., 1902,74, 71-73.
Hoffman H., Birrell J. H. W. The carotid body in normal and anoxic states:
an electron microscopic study.- Acta Anat., 1958, 32, Suppl. 30, 297-311.
Hoffmeister H., Lickfeld K., Ruska H., Rybak B. Secretions granulaires dans
Ie coenr branchial de Myxine glutinosa L.- Z. Zellforsch. mikrosk. Anat.,
1961, 55, 810-817.
Hoglund R. An ultrastructural study of the carotid body of horse and dog.-
Z. Zellforsch. mikrosk. Anat., 1967, 76, 568-576.
Hokfelt B. Noradrenaline and adrenaline in mammalian tissues.- Acta Phy-
siol. Scand., 1951, 25, Sup pl. 92, 1-34.
Hollinshead W. H. The innervation of the adrenal glands.- J. Compo Neurol..
1936, 64, 449-467.
Hollinshead W. H. The innervation of abdominal chromaffin tissue.- J. COml).
Nenrol., 1937, 67, 133-143.
Hollinshead W. H. The origin of the nerve fibres to the glomus aorticum of
the cat.- J. Compo Neurol., 1939, 71, 417-426.
Hollinshead W. H. The innervation of the supracal'dial bodies in the cat.'-
J. Compo Neurol., 194.0a, 73, 37-48.
Hollinshead W. H. A note on the blood supply of the supracardial bodies in
the kitten.- Anat. Rec., 1940b, 76, 283-289.
Hollinshead W. fl. Chromaffin tissue and paraganglia.- Quart. Rev. bioI.,
1940c, 15, 156-171.
Hollinshead W. H. A comparative study of the glomus coccygeum and the
carotid body.- Anat. Rec., 1942,84, 1-16.
Hollinshead W. H. A cytological study of the carotid body of the cat.- Am(,r.
J. Anat., 143, 73, 185-213.
Hollinshead W. H. Localization of chemoreceptor reflexes in the abdominal
body of the rat.- Anat. Rec., 1946, 94, 470-471.
Holtz P. Kurze Originalmitteilungen.- Naturwissenschaften, 1939, 27, 724-
725.
Holzbauer M., Vogt M. Depression by reserpine of the noradrenaline concen-
tration in the hypotalamus of the cat.- J. Neurochem., 1956, 1, 8-11.
Howard .4., Pelc S. R. Synthesis of DNA in normal and irradiated cells and
its relation to chromosome breakage.- Heredity, 1953, Suppl. 6, 261-269.
Hunstman M. E. The effect of certain hormono-like substances of the iso-
lated heart of the skate.- Contr. Canad. Bioi., 1931, New ser. 7, 31-38.
Hyden H. Protein metabolism in the nerve cell during growth and functi-
on.- Acta Physiol. Scand., 1943,6, Suppl. 17.
Inoue .II. On the spinal root pathway of the adrenalin-secretory fibers to
the adrenal gland in toads.- Tohoku J. expo Med., 1958, 68, 379-383.
Ishii Kosei, Ishii Kazuke. Adrenergic transmission of the carotid chemorecep-
tor impulses in the toad.- Tohoku J. expo Med., 1967, 91, 119-128.
Ishii Kosei, Ishii Kazuko. Efferent innervation to the chemoreceptor of the
carotid labyrinth of the toad.- Tohoku J. expo Med., 1970, 102, 113-119.
Ishii K., Ishii K., Honda K. Evidence for adrenergic transmission of the caro-
tid chemoreceptor impulses in the toad.- Nature, 1966a, 210, 1057-1058.
.Ishii K., Honda K., Ishii K. The function of the carotid labyrinth in the to-
ad.- Tohoku J. expo Med., 1966b, 88, 103-116.
.Ishii K., Osaki T. Electron microscopy of the chemoreceptor cells of the ca-
rotid labyrinth of the toad.- Nature 1966, 212, 1499-150Q,
333
Ishii K., Oosaki T. Fine structure of the chemoreceptor cell in the amphibian
carotid labyrinth.- J. Anat. (Lond.), 1969, 104, 263-280.
Ishida S. So-called carotid body of the amphibia.- Igaku Kenkyuu, Fukuoka,
1954, 24, 1024-1050 (Thesis in Japanese language). ( IIT. no Rogers,
1966).
Ito I. Postnatal histogenesis of the adrenal mcdulla of the hamster with spe-
cial reference to the functional structure.- Okajomas Folia anat. jap.,
1957, 30, 239-256. ( IIT. no Adams, 1958).
Ito I. Embryological studies on human adrenal rgland and Zuckerkandl's pa-
raganglia.- Arch. Histol. jap., 1959, 17, 279-2114.
Izquierdo J. J. On the influence of the extracardial nerves upon sino auricular
conduction in the heart of Scyllium.- J. Physiol., 1930, 69, 29-47.
Jacoby M. Ober die Entwicklung der Nebendriisen der Schildriise und dcr
Carotidendriise.- Anat. Anz., 1896, 12, 152-157.
Jensen D. Cardioregulation in an aneural heart.- Compo Biochem. Physiol.,
1961, 2, 181-201.
Johnels A. G., Palmgren A. «Chromaffin» cells in the heart of Mixine glutino-
sa.- Acta Zool., 1960, 41, 313-314.
Johnston J. B. Additional notes on the cranial nerves of petromyzonts.- J.
Compo Neuroi., 1908, 18,569-618.
Jones D. S. Origin of the sympathetic trunks in the chick embryo.- Anat.
Rec., 1937, 70, 45-65.
Jones D. S. Studies on the origin of the sheath cclls and sympathetic ganglia
in the chick.- Anat. Rec., 1939, 73, 343-357.
Tones D. S. Further studies on the origin of the sympathetic ganglia in the
chick embryo.- Aoat. Rec., 1941, 79, 7-15.
Juba A. Untersuchungen iiber die Entwicklung des sympathischeo Grenztran-
ges. Z. Zellforsch. mikrosk. Anat., 1937, 26, 3115-425.
Julin Ch. Recherches sur l'appareil yasculaire et Ie systeme ncrveux periphe-
rique de I'Ammocoetesl) (Petromyzon planeri). Arch. bioI., (Paris), 1887,
7, 759-902.
Jung F. Physiologische Versuche iibcr Pressorezeptorcn an der Karotistei-
lungsstelle bei Vogeln.- Z. Kreislaufforsch., 1934, 26, 328-334.
Jungueira L. C. U. Nota sobre i morfologia dai adrenais dos ofidios.- Rev.
brasil. bioi., 1944, 4, 63-67.
Kaushiva B. S. Thc arterial system of the pond-tI\l'tlc. Lissemys punctata
(Bonnaterre).- Proc. Ind. Acad. Sci., 1940, 12, 84-94.
Keene M. F. K., Hewer E. E. Observations on the development of the human
. suprarenal gland.- J. Anat(Lond.), 1927, 6t, 302-324.
Kingsburg B. F. The term «chromaffin system» and the nature of the chro-
maffin reaction.- Anat. Rec., 1911, 5, 11-16.
Kirshner N. .Uptake of KA by a particulate fraction of the adrenal medul-
la.- J. BIOI. Chem., 1962,237,2311-2317.
Kirshner N., Goodall M. The formation of adrenaline from noradrenaline.-
Biochem. biophys. acta, 1957, 24, 658-659.
Klein U., Kracht J. Ober Hormonbildungsstatten im Nebennierenmark.-
Endockrinologie, 1958, 35, 259-280.
Klump W., Eggerd B. Die Schilddriise und die branchigenen Organe yon let-
hyophis glutinosus L.,- Z. wiss. Zool., 1934, 146, 329-381.
Knab J. Untersuchungen iiber den histochemischen Nachweis von Vitamin C
in der Niere und Nebenniere yon SaI\l'opsiden.- Z. mikrosk. anat. .Forsch.,
1942,52,418-439.
Knoche H., Schmitt G. Ober chemo- und pressoreceptorenfelder am Coronar-
kreislauf.- Z. Zellforsch. mikrosk. Anat., 1963,61, 524-560.
Koch E. Die reflektorische Selbststenerung des Kreislaufs, Dresden. Lpz., Th.
Steinkopff, 1931.
Kock de L. L. Histology of the carotid body.- Nature, 1951, 167,611-612.
Kock de L. L. The intra-glomerular tissues of the carotid body.- Acta
Anat., 1954, 21, 101-116.
Kock de L. L. The carotid body system of the higher vertehrates.- Acta
Anat., 1959, 37, 265-280.
334
Kock de L. L., Dunn A. E. Ultra-structure of carotid body tissue as seen in
serial sections.- Nature, 1964, 202, 821-822.
Koelle G. B. The histochemical differentiation of types of cholinesterases and
their localizations in tissues of the cat.- J. Pharmacol. expo Therap.,
1950, 100, 158-179.
Kohn A. Die Nebennieren der Selachier ncbst Beitragen zur Kenntnis de\'
Morphologie dcI' Wirbeltiernebenniere im allgemeinem.- Arch. mikrosk.
Anat., 1898,53,281-312.
Kohn A. Ober den Bau und die Entwicklung der sogenannten Carotisdriise.-
Arch. mikrosk. Anat., 1900, 56, 81-148.
Kohn A. Das chromaffine Gewebe.- Ergebn. Anal. Entwicklunggesch., 1902,
. 12, 253-348.
Kohn A. Die Paraganglien.- Arch. mikrosk. Anat., 1903, 62, 263-365.
Kohn A. Ober die Entwicklung des sympatischen Nervensystems der Saiige.
tiere.- Arch. mikrosk. Anat., 1907,70, 266-317.
Kohn A. Morphologie dpr inneren Sekretion und der inkretorischen Organe.-
Handb. normal. pathol. Physiol. mit Beriicks. expo Pharm. Berlin, Sprin-
ger, 1930, 16, (Paris), 3-66.
Kohno Sh. Zur vergleirhenden Histologie und Embryologie der Nebenniere
der Siiuger und des Menschen.- Z. Anat. Entw. Gesch., 1925, 77,419-480.
Kose W. Obe\' das Vorkommen «chromaffineI' ZeUen» in Sympathicus des
Mensche und der Siiugetiere. Sitzungsberichte des dtsch. naturwiss.- me-
diz. Vereins fUr Bi.ihmem. «Lotos» in Prag, N. F., 1898, 8, 224. ( IIT. no
. Kose, 1907).
J(ose W. Uber das Vorkommen einer «Carotisdriise,) und der «chromaffinen
ZeIlen» bei Vogeln.- Anat. Anz., 1902, 22, 162-170.
Kose W. Ober die «CarotisdriisCl) und dcI' «chromaffin en ZeUen» bei Vo-
geln.- Nebst Bemerkungen iiber die Kiemenspaltenderivate. Anat. Anz.,
1904,25, 609-617.
Kose W. Die Paragan,glien bei den Vogeln.- Arch. mikrosk. Anat., 1907, 69,
563-790.
Kraus R. Cytology and cytochemistry of the chemoreccptor cell in man.-
Folia morpho!., 1966, 14, 404-412.
Kraus R., Martinek 1. A contribution to the cytology of the carotid body.-
Fol. morphol, 1967, 15,392-395.
Krause R. Mikroskopische Anatomie der Wirbeltiere. IV. Teleostier, Plagiosto.
men, Zyklostomen lInd Lcptokardier. Berli. - Leipzig, Walter, Gruyter
a. Co., 1923, 674-675.
Kuntz A. The development of the sympathetic nervous system in certain fis-
hes.- J. Compo Neurol., 1911, 21, 177-214.
Kuntz A. The development of the adrcnals in the turtle.- Amer. J. Anat.,
1912, 13, 71-89.
Kuntz A. The development of the sympathetic nervous system in man.- J.
Compo Ncurol., 1920,32, 173-229.
Kuntz R. The role of cells of medullary origin in the development of the sym-
pathetic trunks.- J. Compo Neurol., 1926, 40, 389-408.
Kuntz A. The autonomic nervous system. Philadelphia, Lea a. Febiger, 1954.
Laignel - Lavastine M. Application de l'impregnation argentique de Cajal a
l'etllde histo-chimique de la cellule medullosurrenale.- C. r. Soc. bioI.,
1905, 58, 14, 661-663.
Lakshmun A. B. A descriptive study of the cytology of the adrenal glands of
south Indian anurans.-.Endocrinol. japon., 1964, 11, 169-176.
Landau E. Zur Morphologie de Nebenniere. IV Mitt.- Intern. Monatschr. Anat.
Physiol., 1907, 24, 431-446. . . .
Langer E. Beitriige zur Orthologle und Pathologle des Glomus carotlcum. (Pa-
raganglion caroticum.).- Beitr. pathol Anat., 1952 112 251-288.
Langley J. N. The simpathetic innervation of the skin of the frog.- J. PhysioI.,
1910,40, 63 P.
Langley J. N., Orbeli L. A. Observati?n on the sympathetic and') sacral autono-
mic system of the frog.- J. PhyslOl., 1910-1911, 41. 450-48 .
335
Lampinen M. Extra-adrenal chromaffin tissue of the rat and the effect of
cortical hormones on it.- Acta physiol. Scand., 1964, 62, Suppl. 231.
Laurent P., Krieger M. L'innervation des cellules epithelioides du labyrinthe
carotidien des batraciens.- C. r. Acad. sci., 1965, 261, 1565-1568.
Lawton F. E. The adrenal-autonomic complex in Alligator mississipiensis.-
J. Morphol, 1937, 60, 361-370.
Leriche M., Fontaine R., Froehlich F. L'enervation sinucarotidienne, est - elle
permis all point de vue physiologique?- Presse Med., 1935, 61, 1217-
1220.
Lever J. D. Electron microscopic observations on the normal and denervated
adrenal medulla of the rat.- Endocrinology, 1955, 57, 621-635.
Lever J. D. Fine structural appearances in relation to function in certain sec-
retory organs.- Electron Microscopy in Anatomy, 1961, 8, 207-225.
Lever J. D., Boyd J. D. Osmiophilie granules in the glomus cells of the rab-
bit carotid body.- Nature, 1957, 179, 1082-1083.
Lever J. D., Lewis P. R., Boyd J. D. Observations of the fine structure and hi-
stochemistry of the carotid body in the cat and rabbit.- J. Anat. (Lond.),
1959, 93, 478-4.90.
Leydig F. Beitrage zur mikroskopischen Anatomic und Entwicklungsgeschich-
te der Rochen und Haie. Leipzig, 1852, 127 pp. ( I1T. no Bachmann, 1954).
Leydig F. Anatomisch-histologische Untersuchungen iiber Fische und Reptili-
en. Berlin, Reimer, 1853.
Linden P., van der. RecherdlCs sur Ies reflexes sino-carotidiens chez les
oiseaux.- Arch. int. PhysioI, 1934, 40, 59-82.
Liu H. Ch., Maneely R. B. Some aspects of adrenal histology and histochemi-
stry in the soft-shelled turtle of South China (Chelonia amyda).- J. En-
docrinol., 1959, 19, 1-£1.
Lofts B., Phillips J. G. Some aspects of the structure of the adrcnal gland in
snakes.- J. Endorrinol., 1965, 33,327-328.
Luschka H. Ober die drusenartige Natur des sogenanaten Ganglion intercaro-
ticum.- Arch. Anat. Physiol., Lpz., 1862,405-414.
Lutz B. R. The innervation of thc heart of the elasmobranch, Scyllium cani-
cula.- Bioi. Bull., 1930a, 59, 211-216.
Lutz B. R. The visceral afferent pathway in the elasmobranch, Scyllium cani-
cula.- BioI. Bull., 1930b, 59, 217-221.
Lutz B. R., Case M. A. The Beginning of adrcnal function in the chic.k emb-
ryo.- Amer. J. Physiol., 1925, 73, 670-678.
Lutz B. R., Wyman L. C. The chromaphin tissuc and interrenal bodies of elas-
mobranches and the occurence of adrenalin.- J. expo Zool., 1927, 4.7,295-
307.
Lutz B. R., Wyman L. C. Reflex cardiac inhibition of branchiovascular origin
in the elasmobranch, Squalus aeanthias.- Bioi. Bull., 1932, 62, 10-16.
Mackay M. E. The action of some hormones and hormone - like substances
on the circulation in the skate.- Contr. Canad. BioI., 1931, New ser., 7,
17-29.
Marchad F. Beitragc ZIIr Kcnntniss dcI' nOI'malen und pathologischen Anato-
mie der Glandula carotica und der Nebennieren.- Int. Beitr. wiss. Mcd.,
1891, 1, 535-581.
Marcus N. Ober den Sympathikus.- S. B. Ges. Morph. Physiol., Miinchen,
1910, 25, 119-131.
Marques M., Serrano I. Adrenalina () noradrenalina na supnwnal de tarlarllgas
normais e hipofisoprivas.- Rev. brasil. bioI., 1960, 20, 331-356.
Martoja M., Duguy R., Saint-Girons H. Donnees histologiques sur les variati-
ons de la glande surrenale au COllI'S du cycle annue] chez Vipcra aspis.
L.- Arch. Anat. microsk. Morphol. exptl., 1961, 50, 233-249.
Meiyling H. A. The glomus caroticum and sinus caroticus of the horse.- Proc.
Acad. Sci., Amst., 1936,39, 706-713.
Meiyling H. A. Bau und Innervation von Glomus caroticum und sinus caro-
tic us. Eine Untel'Suchung unter Benutzung spezifischer N ervenfarbungs-
methoden.- Acta neerl. mornh., 1938, 1, 193-288.
336
Mihalik P. V. Unhw-suchlmgen tber die Entwicklung des sympathischen Ner-
vensystem.- Anat. Anz., 1940,39,241-296.
Milcou St. M. Peut-on classer Ie ganglion carotidien parmi les paragangli-
ons?- C. r. Soc. bioI., 1930, 105, 55-58.
Miller E. H. Development of the epinephrin content of the suprarenal medul-
la in early stages of the mouse.- Amer. J. Physiol., 1926, 75, 267-277.
Miller M. R. The normal histology and experimental alteration of the adrenat
of the viviparous lizard, Xantusia vigilis.- Anat. Rec., 1952, 113, 309-323.
ftlitsukuri R. B. On the development of the suprarenal bodies in Mammalia.-
Quart. J. microsk. Sci., 1882,22, 17-29.
Moore B., Vincent S. The comparative chemistry of the suprarenal capsules.-
Proc. roy Soc. (Lond.), 1898, Ser. B. 62, 280-283.
Moppert J. DeI' intrazellulare Bidungsort der phaeochromen Granula im Ne-
bennierenmark der Ratte. Eille licht unci elektronenmikroskopisch
auwradiographishsche Untersuchung.- Z. Zellforsch. mikrosk. Anat., 1966,
74, 45-52.
Morita E., Chiocchio S. R., Tramezzani J. H. The carotid body of the WedeU
seal (Leptonychotes weddelli).- Anat. Rl'c., 1970, 167, 309-328.
Morita E., Ghioocchio S. R., Tramezzani J. H. Four types of main cells in the:
carotid body of the cat.- J. Ultrastruct. Res., 1969, 28, 399-410.
Mi.iller E. Beitrage zur Kenntnis des autonollien Nervensystems. 1. Oeber die
Entwicklung des Sympathicus und des Vagus bei den Selachiern.- Arch.
mikrosk. Anat., 1920,94, 208-247.
Mi.iller E., Ingvar S. Oeber dpn Urspnmlr des Sympathicus bei den Amphibi-
en.- Upsala LiikareWren forhandl1921, 26, 1-15.
Miiller E., lngvar S. Oeber den Ursprung des Sympathicus beim Hiihnchen.-
Arch. mikrosk. Anat., 1923,99, 650-671.
Miiller J. Die Nebennieren von Gallus domesticus und Columba livia domesti-
ca.- Z. mikrosk. anat. Forsch., 1929, 17, 303-352.
Mulon P. Reaction de Vulpian au niveau des corps surrenaux des plagiosto.-
mes.- C. 1'. Soc. bioI., 1903, 55, 1156.
Mulon P. Specifite de la reaction chromaffine: glandes adrenalogenes.- C. 1'..
Soc. bioI., 1904a, 56, 113-115.
Mulon P. Lps ,glandes hypertensives on organes chomaffines.- Arch. gen. me d.,
1904b, 81, 3265-3277.
Muratori G. Recherches histologiques et experimentales sur !'innervation du
tissu paraganglionnaire (pheochrome) annexe au systeme du Derf vague-
des amniotes (glomus carotidien; paraganglions extravagaux et intrava-
gaux).- C. r. Ass. Anat., 1932a, 27th. reunion, 40H-415.
Muratori G. Ricerche istologiche e sperimentali sulI'innerazione del tessuto.
paragangliare annesso al sistema del vago. (Paraganglio carotico; para-
gangli estravagali od intravagali).- Boll. Soc. ital. BioI. Sper., 1932b, 7,.
137-142.
Muratori G. Contributo all'innervazione del tessuto paragangliare annesso at
sistema del vago (glomo carotico, paragalllgii estravagali ed intravagaIi)
e all'innervazione del zeDO carotideo.- Anat. Anz., 1932c, 75, 115-123.
Muratori G. Recerche istologichp sull'innervazione del glomo carotico.- Arch.
ital. Anat. Embriol., 1932d, 30, 573-602.
Muratori G. Conseguenze di lesioni sperimentali del vago sull'innervazione del
glomo carotico.- Monit. zool. ital., 1933a, 43 (suppl), 87-89.
Muratori G. Osservazioni sulla base istologica dei riflessi aortico e caroti-
deo.- Atti Soc. med.- chir., Padova, 1933b, 11,213-217.
Muratori G. Ricerche sperimentali sulla zona glomo-carotidea degli ucceli.-
Atti. Accad. Sci. Pad ova, 1933c, 49, 257-271.
Muratori G. SulI'esistenza di un rifles so carotideo negli ucceli.- Boll. Soc.
ital. BioI. Sper., 1933d, 8, 1273-1276.
Muratori G. Connessioni tratessuto paragangliare e zone recettrici aortiche in
vari mammiferi.- Monit. Zool. Hal., 1935, 45, 300-310.
Muratori G. Osservazioni istologiche e considerazioni embriologiche sui re-
cettori aortici degli amnioti.- Anat. Anz., 1937, 83, 367-379.
331
Muratori G. Istofisiologia comparata delle zone aortico-arteriose pressorecetti-
ve e degli annessi dispositivi chemorecettori (paraganglio carotico, para-
ganglio succlavio, paraganglio aortico-polmonare).- Atti. Soc. ital. anat,
1959, Suppl. 68, 17-84.
Muratori G. Histological observations on the cervicothoracic paraganglia of
amniotes.- Arch. into Pharmacodyn., 1962, 160, 217-226.
Jluratori G. Caratteristiche istoanatomiche doll paragangli trac.heo-bronchiali
e della vena cava 5uperiore nel Igatto.- Bioi. Sper., 1964, 40, 892-894.
Muratori G. Nidi chromaffinie anastomisio artero-venose nella parete bronchi-
ale.- Boll. Soc. ital. BioI. sper., 1965, 41, 1186-1188.
Muratori G. Microparagangli, anastomosi artero-venose peribronchiali e ana-
stomosi polmo-bronchiali nel gatto.- Arch. ital. anat. Emhriol., 1968,73,
133-154.
Muratori G., Battaglia G. Ossorvazione sulla reazione iodaffine nel paragang-
Iio carotico.- Boll. Soc. ital. BioI. Sper., 1960, 36, 400-405.
Muratori G., Battaglia G., Modunesi G. Dimostrazione istochimica della Nomd-
renalina nel cromaffine cervicotoracico e aortico-addominale degli amni-
oti. I. sservazioni al microscopio ottico.- Boll. Soc. ital. BioI. Sper., 1965a,
41, 1182-1185.
Muratori G., Battaglia G., Modonesi G. Dimostrazione istochimica della Norad-
renalina nel cromaffine cervicotoracico e aortico addominale degli amnio-
ti: II. Osservazioni al microscopio elcctronico.- Boll. Soc. ital. BioI. Sper.,
1965b, 41, 1185-1186.
Muratori G., Chiarini G., Battaglia G. Dimostrazione delle catecholamine nel
chromaffine cervico-toracico dei mammfieri mediante la microscopia a flu-
orescenza.- Rivista istochim. norm. palol., 1966, 12, 1-10.
Muscholl E., Rahn K. H., Watzka M. Nachweis von Noradrenalin im Glomus
caroticum.- Naturwissenschaften, 1960,47,325.
Muscholl E., Vogt M. Perfusion of extramedullary chromaffine tissue.-
J. Physiol., 1963, 169, 93P.
Muscholl E., Vogt M. Secretory responses of extramedullary chromaffin tis-
sue.- Brit. J. Pharmacol., 1964,22, 193-203.
Nagel M. Ober die Struktur den Nebennieren.- Muller's Arch. Anat. Physi-
01. wiss. Med., 1836, 365-383.
Naik D. V., George 1. C. Histochemical demonstration of changes in the ac-
tivity of alkaline and acid phosphatases in the adrenal of a migratory
starling.- J. Histochem. Cytochem., 1964, 12, 772-776.
Nandi J. The structure of the interrenal gland in teleost fishes.- Univ. Calif.
Puhl. Zool., 1962, 65, 129-211.
Neil E., Strom L., Zotterman Y. Action potential studies of afferent fibres in
the IX-th and X-th cranial nerves of the frog.- Acta physiol. Scand., 1950,
20, 338-350.
. Nicol J. A. C. Autonomic nervous systl'm of the ratfish.- Nature, 1950a, 165,
854-855.
Nicol J. A. C. The autonomic nervous system of the chimaeroid fish Hydrola-
gus colliei.- Quart. J. microsk. Sci., 1950b, 91, 379-399.
Nicol J. A. C. Autonomic nervous system in lower chordates.- BioI. rev.,
1952,27, 1-49.
Nikiforowsky P. M. On depressor nervp fibres in the vagus of the frog.-
J. Physiol., 1913,.45, 459-461.
Noble G. K. The biology of the amphibia. N.-Y.- London, 1931.
Nonidez J. F. The aortic (depmssor) nerve and its associated epithelioid bo-
dy the glomus aortieum.- Amer. J. Anat., 1935a, 57, 259-301.
Nonidez J. F. The presence of depressor nerves in the aorta and carotid of
birds.- Anat. Rl'c., 1935b, 62, 47-73.
Nonidez J. F. Observations on the blood supply and the innervation of the
aortic paraganglion of cat.- J. Anat. (Lond.) 1936, 70, 215-224.
Nonidez J. F. Identification of the receptor areas in the venae cavae and
pulmonary veins which initiato reflex cardiac acceleration (Bainbringe's
reflex).- Am. J. Anat., 1937a, 61, 203-232a.
.338
Nonidez J. F. Distribution of the aortic nerve fibers and the epithelioid bodieS'
(supracardial «paraganglia») in the dog.- Anat. Roo., 1937b, 69, 299-317..
Nonidez 1. G. Arterio-veneous anastomoses in the sympathetic chain ganglia
of th dog.- Anat. Rec., 1942,82, 593-608.
Norberg C. A., Hamberger B. The sympathetic adrenergic neuron some chao.
ractl,ristic revealed bay histochemical studic in the intra neuronal distri-
bution of the transmittet.- Acta physiol. Scand., 1964, 63, Suppl. 238.
Ocheterena I. Estudios neurologicos. XXIX. Acerca del seno y del glomus ca-
roticum.- Ann. Inst. BioI., Univ. Mexico, 1936,7, 397-.H4.
O'Donoghue Ch. H. The blood vascular system of the Tuatara, SphenodeD
punctatus.- Phil. Trans. Roy. Soc. (London), 1921, Ser. B. 210, 175-252.
Ogata T., Ogata A. Henle's reaction of the chromaffin cells in the adrenals
and the microscopic test for arlrenalin.- J. expo Med., 1917, 25, 807-818.
Oguri M. Studies on the adrenal glands of teleosts. Ill. On the distribution of
chromaffin cells and interrenal cells in the head kidneys of fishes.-
Japan. Soc. Sci. Fisheries Bull., 1960, 26, 443-447.
Okuda 111. The adrenalin content of the suprarenal capsule in the chick emb-
ryo.- Endocrinology, 1928, 12, 342-348.
Oliver G., Schafer E. A. The physiological effects of extracts of the suprare-
nal capsules.- J. physiol., 18 5, 18, 230-276.
Olivereau 11/. Mise en evidence d'adl'enaline et de noradrenaline dans Ie tis-
sue chromaffine d'un poisson selacien la roussette.- Ann. Endocrinol.
(Paris) 1959, 20, 645-652.
OnQdi A. Ueber die Entwickelung des sympathischen Nel'vensystem.- Arch.
mikrosk. Anat., 1886, 26, 61-80.
Onozato H., Yamamoto K. On the fine structure of adrenal chromaffin cell,>-
of the goldfish with special reference to the distribution of two catecho-
lamines, adrenaline and noradrenaline.- Ann. Zool. japon., 1966, 39, 30-38.
Ostlund E. The distribution of catecholamines in lower animals and their
effect on the heart.- Acta phiosol. Scand., 1954, 31, Suppl. 112, 1-67.
Ostlund E., Bloom G., Adams-Ray J., Ritzen M., Siegman M., Nordenstam H.,
Lishajko F., Euler U. S., von. Storage and release of catecholamines an.J.
the occurence of a specific submicroscopic granulation in cyclostoIlle he-
art.- Nature, 1960, 188, 324.-325.
Overbeeke A. P., van. Histological studies on the interrenal and the phaeoch-
romic tissue in Teleostei. Amsterdam: van Munster's Drukkerijen. N-Y.,
1960.
Painter T. Do nuclei of living cells contain more DNA than is revealed by
the Feulgen stain?- Texas Repts. BioI. Med., 1955, 3, 659-666.
Palay S. L. Synapses in the central nervous system.- J. Biophys., Biochem.
Cytology, 1956,2,4, 2, 193-202.
Palay S. L. Tho morphology of secretion Frontiers in Cytology. New Haven,
Yale Univ. Press, 1958, 305-342.
Palkama A. Histochemistry and electron microscopy of the carotid body.-
Ann. med. exptl. et bioi. fenniae, 1965,43,260-266.
Palme F. Die paraganglien iiber dem Herzen und im End gungsgebiet des.
Nervus depressol'.- Z. mikrosk.-anat. Forsch., 1934,36,391-420.
Paltauf R. Deber Geschwiilste der Glandula carotica nebst einem Be,itrage
zur Histologie und Entwickelungsgeschichte derselben.- Beitr. pathol.
Anat., 1892, 11, 289-300.
Palumbi G. La fine vascolarizzazione ed innervazione del glomo carotideo del
mammiferi.- Comment. pontif. Acad. Sci., 1940, 4, 289-320.
Pankratz D. S. The development of the suprarenal gland in the albino rat
with a consideration of its possible relation to the origin of foetal move-
ments.- Anat. Rec., 1931, 49, 31-49.
Pannese E. Observations on the morphology submicroscopic structure and bio-
logical properties of satellite cells (s. c.) in sensory ganglia of mamals.-
Z. ZeIlforsch. microsk.-Anat., 1960, 52, 567-597.
Pannier R. Donnees generales sur Ie systeme jganglionnaire' et paraganglion-
naire du coeur.- C. r. Soc. bioi., 1935, 120, 1350-1353.
339'
Penitschka W. Dber den Bau des Ganglion cervicale uteri des Menschen mit
Beriicksichtigung der mehrkernigen Ganglienzellen und des chromaffinen
Gewebes.- Anat. Anz., 1929, 66, 417.
Penitschka W. Paraganglion aorticum supracardiale.- Z. mikrosk.-anat.
Forsch., 1931, 24, 24-37.
Pessacq T. P. La nevroglie du glomus carotidien.- Arch. Anat. microsc., Mor-
phol. exptI., 1961, 50, 289-298.
Pettinari CxeMa reHeaa xpoMacJ>cJ>HHHIdX II CHIIInaTII'leCKHX KJIeTOK. 1953 ( IIT.
no Giordano-Lanza, 1(65).
Pettit A. Sur les capsules surrenales et la circulation porte surrenale des rep-
tiIes.- Bull. Soc. Zool. France, 1895, 20, 233-237.
Pettit A. Recherches sur les capsules surnJnales.- J. Anat. Physiol, 1896, 32,
301-362, 369-419.
Peyrin L., Cier J. F.. Peres G. La methylation de la noradrenaline et les in-
terrelations corticomedullaires chez la roussette.- Ann. Endocrinol. (Pa-
ris), 1969, 30, 1-38. .
Philip pot E., Goffart M., Dresse A. Le systeme surrenale sympathique chez
Ie paresseux (Choloepus hoffmanni Peters).- Arch. intern. Physiol. Bio-
chim., 1965, 73, 476-504.
Picard D., Vitry G. Histophysiologie de la medullo-surrenale. 5 e Reunion En-
docrinolog de Langue Francaise, Paris, cd Masson 1959, 5, 234-293.
Picard D., Vitry G. Cytophysiologie de la cellule medullo-surrenalienne.-
Arch. bioi. (Liege), 1964,75, 453-496.
Piezzi R. S. Two types of chromaffin cells in the adrenal gland of the «Bufo
arenaruml) Hensel.- Acta physio1. latinoamer., 1965, 15, 96-100.
Piezzi R. S. Two types of synapses in the chromaffin tissue of the toad's ad-
renal.- Acta physioI. latinoamer., 1966, 16, 282-285.
Pischinger A. Dber die Entwicklung und das Wesen des Carotislabyrinthus
bei Anuren.- Z. Anat. Entw. Gesch., 1934, t03, 45-52.
Pischinger A. Kiemenanlagen und ihre Schicksale bei Amnioten.- Hand:).
vergl. Anat. Wirbeltiere, Berlin-Vienna, eds. L. Bolk. E. Goppart. 1937, 3,
279-334.
Pitzorno M. Sulla struttura dei gangli simpatici nei selaci.- Monit. Zool.
Ita!., 1910, 21, 53-61.
Pool H. Die vergleichende Entwicklungsgeschichte der N ebennierensysteme
der Wirbeltiere.- Handb. vergl. expo Entwicklungslehre Wirbelt. Jena,
ed. Fischer, 1906, 3, 443-616.
Poll H., Sommer A. Dber phaeochrome Zellen im Centralnervensystem des
Blutegels.- Arch. Anat. u. Physiol. Physiol. Abth., Lpz. 1903, 549-560.
Rabl C. Dbe,r die Entwicklung des Urogenitalssystem der Selachier.- Morph.
Jahrb., 1896, 24, 623-767.
Rabl H. Die Entwicklung und Structur dcI' Nebennieren bei dem Vogeln.-
Arch. microsk. Anat., 1891, 38, 492-523.
Rabl H. Die Entwicklung der Carotisdriise beim Meerschweinchen. Arch. mik-
rosk. Anat., 1922, 96, 315-339.
Radu V. Les glandes surrenales des selaciens.- C. r. Soc. bioI., 1932, 11 J,
119-121.
Rahn K. H. Morphologische Untersuchungen am Paraganglion caroticum mit
histochemischem und pharIDacologischcm Nachweis Yon Noradrenalin.-
Anat. Anz., 1961, 110, 140-159.
Ramon y Cajal S. Geneses de las fibras nerviosas del embrion y observatio-
nes contrarias a la teoria cateraria.- Trab. lab. bioI. Univ. Madrid, 1906,
4, 227-294.
Ramon y Cajal S. Nou\"cHes observations sur l'evolution des neuroblastes,
avec quelques remarques sur l'hypothese neurogenetique de Hensen-
Held.- Anat. Anz., HJ08, 32, 1-25; 65-87.
Randall D. J. Functional morphology of the hl'art in fishes.- Amer. Zoo!.,
1968, 8, 17()-189.
Rau A. S., Johnson P. H. Observations on the development of the sympathe-
tic nervous system and suprarenal bodies in the sparrow.- Proc. Zoot
Soc. (Lond.), 1923, 741-768.
340
Rau A. S. Observations on the anatomy of the heart, lungs and related parts of
Ceratophrys.- J. Anat. (Lond.), 1924,58, 299-327.
Ray H. C. On the venus of the common Indian Rat - Shake, ptyas nmco-
sus.- J. Morphol., 1936, 59, 571-547.
Ray I., Ghosh A. The chemical cytology of the adrenal medulla in the domes-
tic pigeon.- Acta histochem., 1961, 11, 68-77.
Reese A. M. The vascular system of the, Florida alligator.- Proc. Acad. nat.
Sci. Philad., 1914, 66, 413-425.
Reese A. M. The ductless glands of Alligator mississipiensis.- Smithson mis-
cell. Collect, 1931, 82, 1-14.
Reis D. 1., Fuxe K. Adrenergic innervation of the carotid sinus.- Amer. J.
PhysioI., 1968, 215, 1054-1057.
Remak R. Dber ein Selbstiindiges Darmnervensystem, Berlin, ed. V. G. Rei-
mer, 1847.
Retzius G. Die sensiblen Nervenendigungen in der Haut des Petromyzon.-
BioI. Untersuch., 1892, 3, 37-40.
Retzlaff E. W. The histology of the adrenal gland of the aIIrgator lizard, Ger-
bonotns multicarinatns.- Anat .Rec., 1949, 105, 19-34.
Ribaupierre de F., Siegri$t G., Dunant J., Dolivo M. Etude de la morphologie
et de la function probable des cellules chromaffines du ganglion Sympa-
thique cervical superieur.- J. physiol., 1966, 58, 601.
Riegele L. Die Nerven des Glomus caroticum beim Menschen mit kurzer Uber-
sicht iiber den histologischen Aufban des Organs.- Z. Anat. Entw. Gesch.,
1928, 86, 142-167.
Robertis de E. Submicroscopic morphology and function of the synapse.-
Exp. Cell. Res., Suppl. 5, 1958,347-369.
Robertis de E., vaz Ferreira A. Electron microscope study of the excretion of
cathechol- containing droplets in the adrenal medulla.- Exp. Cell. Res.,
1957a, 12, 568-574.
Robertis de S., vaz Ferreira A. A multivesicular eathechol-containing body
of the adrenal medulla of the rabbit.- Exp. Cell Res., 1957b, 12, 575-
581.
Robertis de E., Sabatini D. Submicroscopic analysis of the secretory process
in the adrenal medulla.- Fed. Proc., 1960, 19, Suppl. 5, 70-78.
Rogers D. C. The development of the rat carotid body.- J. Anat. (Lond.),
1965, 99, 89-101.
Rogers D. C. A histological and histochemical study of the carotid labyrinth
in the anUl'an amphibians. Bufo marinus, Hyla aurea and Neobatrachus
pictus.- Acta Anat., 1966,63, 249-280.
Rogers D. C. The structure of the carotid bifurcation in the Lizards tiliqua
occipitalis and Trachysaurus rugosus.- J. Morphol., 1967, 122, 115-
130.
Romanoff A. L. The Avian Embryo (Structural and functional development),
N.-Y., ed. Macmillan, 1960.
Ross L. A cytological and histochemical study of the, carotid body.- Anat.
Rec., 1957, 129, 433-455.
Ross L. Electron mikroscopic observations of" the carotid body of the cat.-
J. Biophys. Biochem. CytoI., 1959, 6, 253-262.
Rugh R. Experimental Embryology, Techniques a Procedures. Burgess, Min-
neapolis, Minnesota, 1962.
Rude S. Monoamine-containing neurons in the central nervous system and
peripheral nerves of the leech, Hirudo medicinalis.- J. compo Neurol.,
1969, 136,349-372.
Rybak B. Existence d'un automatisme contractile dans les verticules du coc-
ur branchial de Myxine glutinosa.- C. S. Acad. sci., 1960, 251, 1185-1187.
Rybak B., Hoffmeister H., Ruska H. Etudes sur la physiologie ultrastructura-
Ie de l'oreillette du coeur branchial de myxine glutinosa.- Life Sciences,
1962, 4, 109-114.
Rybak B., Ruska H., Ruska C. Ultrastructures nerveuses dans Ie ventricule du
coeur de grenouille.- Experientia, 1966, 22, 735-737.
341
Saint - Girons H. Donnees histophysiologiques sur Ie cycle annuel des glan-
des endocrines et de leurs effecteurs chez 1'0rvet, Anguis fragilis L.-
Arch. Anat. micro Morphol. expo 1963,52, 1-51.
Bauer F. C. Mitoses in the neural tube.- J. Compo Neurol., 1935a, 62, 377-
405. ..p-e.
Sauer F. C. The cellular structure of the neural tube.- J. Compo Neurol.,
193563(1), 13-21b.
Bchaper A. Die huhesten Differenzierungsvorgange im Zentralnervensystem.-
Roux' Arch. Entwickl. Wiss. Mech., 1897,5,81-132.
Schneider F. H., Smith A. D., Winkler H. Secretion from the adrenal medulla:
biochemical evidence for exocytosis.- Brit. J. Pharmacol. Chemotherapy,
1967, 31, 94-104.
Schonberger W. Dber die paraganglien in der plica ventricularis bei neugebo-
renen.- Anal. Anz., 1966, 119, 296-304.
Schonberger W. Das Paraganglion laryngeum beim Gorilla.- Anat. Anz., 1967,
120, 323-327.
SchOnberger W. Die Entwicklung des Paraganglion la1'yngeum beim Mensc-
hen.- Anat. Anz., 1969, 125, 512-524.
Bchumacher S. Ober die Bedeutung der arteriovenosen Anastomosen und der
epithelioden Muskelzellen (Quellzellen).- Z. mikrosk.-anat. Forsch., 1938,
93, 107-130.
Schumann H. 1. The distribution of adrenaline and noradrenaline in chromaf-
fin granules from the chicken.- J. Physiol., 1957, 137, 318-325.
Schwarz - Karsten H. Die Entwichklung des Paraganglion caroticum und der
Paraganglion am Hel'Zen des Schweines. Z. Anat. Entw. gesch., 1944, 113,
39-65.
.Bedgwick A. On the inadequacy of the cellular theory of development and on
the early development of nerves, particularly of the third nerve and of
the sympathetic ganglia in elasmobranchii.- Quart. j. microsk. Sci., 1893,
37, 87-101.
Seto H. Dber zwischen Aorta und Arteria pulmonalis gelegene Herzparagang-
lien.- Z. ZelIforsch. mikrosk.-anat., 1935, 22, 213-231.
.Shepherd D. M., We$t G. B. Noradrenaline and accessory chromaffin tissue.-
Nature, 1952, 170,42-43.
Shepherd D. M., West G. B. Differentiation of chromaffin cells.- J. Pharma-
col. (Lond.), 1953, 5, 216-216.
Shepherd D. M., West G. B., Erspamer V. Chromaffin bodies of various spe-
cies of dogfish.- Nature, 1953, 172, 509.
Siegrist G., Dolivo M., Dunant Y., Foroglou-Kerameus C., Ribaupierre de F.,
Rouiller Ch. Ultrastructure and function of the chromaffin cells in the su-
perior cervical ganglion of the rat.- J. U1trastruct. Res., 1968, 25, 381-
407.
Sidky Y. A. The carotid sinus in lizards with an anatomical survey of the
ventral neck region.- J. Morphol., 1967, 121, 311-321.
Singer M. The sympathetics of the branchial region of the urodele, Tritu-
rus.- J. Comp. Neurol., 1942, 76, 119-143.
Smith Ch. The origin and development of the carotid body.- Amer. J. Anat.,
1924, 34, 87-131.
.Soulie A. H. Recherches sur Ie development des capsules surrenales chez les
vertebres superieurs.- J. Anat. (Paris), 1903, 39, 197-293.
.Spanner R. Oeber die WUl'Zelgebiete dcI' Nieren - Nebennieren - und Leber-
pfortader bei Reptilien.- Morphol. Jahrb., 1929, 63, 314-358.
.Stannius H. F. Das peripherische Nervensystem der Fische, anatomisch and
physiologisch Untersucht. Rostock, 1849.
Stilling H. A propos de quelques experienses nouvelles sur la mala die d'Ad-
dison.- Rev. merl., 1890, 10, 808-831.
.Stilling H. Zur Anatomie der Nebennieren. II. Mitt.- Arch. microsk. Anat.,
1898,52, 176-195.
Stilling H. Die chromafillen Zellen und Korperchen des Sympathicus.- Anat.
Anz., 189, 15, 229-233.
..342
Slznder-Plassman;; P. Untersuchungen iiber den Bulbus carotJidis Lei Mensch
und Tier im Hinblick auf die (,Sinusreflexe» nach H. E. Hering: ein
Vl'rgleich mit anderen Gdassstrecken: die Histopathologie des Bulbus ca-
rotidis; das Glomus caroticum.- Z. Anat. Entw. Gesch., 1930, 93, 567-
622.
Swammerdam 1. Bibel der Nature. Leyden: I Severin us, 1738, 2, 832 ( nT. 110
. Bachmann, 1954).
Swinyard C. A. The innervation of the suprarenal glands.- Anat. Rec., 1937,
68, 417-427.
Tallbe H. W. L. De Vera Nervi Intel'Costalis Origine Unaug.- Diss. Gottingae,:
A Vandenhoeck, 1743, 20 pp. ( HT. 110 Adams W., 1958).
Taxi J. Contribution a l'etude des connexions des neurones moteurs du syste-
me nerveux autonome.-Ann. Sci. natur. zool. (Paris), 1965, Ser. 12, 7.
413-674.
Temi T. II corpo ultimobranchiale degli uccelli Ricerche embriologiche anato-
miche e istologiche su Gallus dom.- Arch. ital. Anat. Embriol., 1927, 24.
4U7-531.
Terni T. II simpatico cervical{' degli amnioti (Ricerche di morfologia compa-
rata).- Z. Anal. Entw.-Gesch., 1931, 96, 28!J-426.
Terracoll. Le Iglomus jugulare, Paris, 1956.
Tigyi A., Benedeczky J., Puppi A., Lissak K. Identification of the secretory
granules in the adrenal medulla.- In: "Electron microscopy", Prague,
Czeskosl. Acad. Sci., 1964, Vol. 8., 491-492.
Tramezzani 1. H., Chiocchio S., Was$ermann G. F. A technique for light and
electron microscopic identification of adrenalin and noradrenalin-storing-
cells.- J. Histochem. Cytochem., 1964, 12, 890-899.
Trifar6 J. M. The secretory process of the adrenal medulla (Review).- Endo-
crinol. exper., 1970, 4, 225-271.
Trinci G. Sulla esistenza di un paraganglio cardiaco e di un paraganglio ca-
rotico (Glandula carotica) nei l'ettili.- Monit. Zool. ltal., 1909, 20, 286-
289.
Trinci G. II sistema cromaffine cardiaco-cervicale nei sauri.- Arch. ital.
Anat. Embriol., 1912, to, 197-260.
Truex R. C. Chromaffin of the sympathetic ganglia and heart.- Anat. Rec.,
1950, t08, 687-697.
Uchida S. Oeber die Entwicklung des sympatischen Nerwensystems. I Mit-
teilung. Deber die Entwicklung des Sympathicus bei dem Vogeln.- Acta
scholae medicinalis in Kioto, 1927a, to, 63-94.
Uchida S. Deber die Entwicklung des sympathischen Nerwensystems. II Mit-
teilung. Deber die Entwicklung des sympathicus beim Siiugetieren.-
Acta scholae medicinalis in Kioto, 1927b, to, 95-126.
Uchida S. Oeber die Entwicklung des sympathischen Nerwensystems. III. Mit-
teilung. Oeber die Entwicklung des Sympathicus beim Reptilien.- Acta
scholae medicinalis in Kioto, 1927c, to, 127-136.
Vaida A., Duca S., Lazar A. Zur innervation des Glomus caroticum des Mens-
chen.- Anat. Anz., 1966, 118, 171-177.
Valentin G. Oeber das Canglion intercaroticum.- Heckers Ann. Gas. Reilk,
1833, 26, 398-407.
Verdun P. Surles derives branchiaux du poulet.- C. r. Soc. bioi., 1898, 5,
243-244.
Vincent S. The suprarenal capsules in the lower vertebrates.- Proc. Birm.
Nat. Rystol. Physiol. Soc., 1896, to, 1-26.
Vincent S. On the morphology and physiology of the suprarenal capsules in
fishes.- Anat. Anz., 1897a, 13, 39-48.
Vincent S. Contributions to the comparative anatomy and histology of the
supprarenal capsules. The suprarenal bodies in fishes, and their relati-
on to the so-called head-kidney.- Trans. Zool. Soc. (Lond.), 1897b, 14,
41-84.
Vincent S. The comparative histology of the suprarenal capsules.- Intern.
Mschr. Anat. Physiol., 1897c, 15,282-303,305-326.
345
Vincent S. The comparative physiology of the suprarenal capsules. Proc. roy.
Soc. (Lond.), 1897d, 61, 64-73.
Vincent S. The carotid gland of mammalia and its relation to the suprarenal
capsule, with some remarks upon internal secretion and the phylogeny
of the latter organ.- Anat. Anz., 1900, 18,68-76.
Vincent S. The ehromaphil tissues and the adrenal medulla.- Proc. roy. Soc.
(Lond.), 1910, Ser. B, 82, 502-515.
Vincent S. Internal secretion and the ductless glands. (2nd ed.). London, Ed.
Arnold and Co, 1922,250-253.
Viveros O. H., Arqueros L., Connett R. 1.. Kirshner N. Mechanism of secretion
from the adrenal medulla. III. Studies of dopamine f\-hydroxylase as a
marker for catecholamine storage vesicle membranes in rabbit adrenal
glands.- Molec. Pharmacol., 1969a, 5, 60-68.
Viveros O. H., Arqueros L., Connett R.I., Kirshner N. Mechanism of secretion
from the adrenal medulla. IV. The fate of the storage vesicles following
insulin and reserpine administration.- Molec. Pharmacol., 1969b, 5, 69-
82.
Vivien 1. Les glandes endocrines.- Traite zool. anat.-systematique bioI., 1958,
13 (fasc. 2), 1470-1544.
.Vulpian M. Note sur quelques reactions propres a la substance des capsules
surrenales.- C. r. Acad sci., 1856, 43, 663-665. .
Wakisaka N. Histological studies in the adrenal gland of teleosts, Part I. Op-
tical and electron microscopie observations on the adrenal tissue of rain-
bow trout.- Bull. Yamaguchi Med. School., 1963a, to,99-112.
Wakisaka N. Histological studies in the adrenal gland of teleosts. Part. II.
Optical and electron microscopic observations on the adrenal tissue oi
carp.- Bull. Yamaguchi Med. School., 1963b, 10, 113-122.
Waldeyer W. Dber einige neure Forschungen im Gebiete der anatomie des
Zentralnervensystem.- Dtsch. med. WochSischr., 1891, 17, 1213-1218;
1244-1246; 1267-1269; 1287-1289.
Waring H. The development of the adrenal gland of the mouse.- Quart. J.
microsk. Sci., 1936, 78, 329-366.
iWassermann G. R. Tramezzani 1. H. Complete separation of adrenaline and
noradrenaline-secreting cells in a snake's adrenal gland. Acta physiol. la-
tinoamer., 1961, 11, 148-149.
Wassermann G. F., Tramezzani J. H. Separate distribution of adrenaline amI
noradrenaline-secreting cells in the adrenal of snakes.- Gen. Comp.
- Endocrinol., 1963, 3, 480-489.
-Watzka M. Dber die Verbindungen inkretorischer und neurogener Organe.-
C. r. Ass. Anat., 1930, 25th reunion 385-386.
Watzka M. Ober die Verbindungen inkretorischer und neUl'ogener Organe.
Anat. Anz., Erganzungsh, 1931, 71, 185-190.
Watzka M. Vergleichende Untersuchungen tiber den uItimobranchialen Kor-
per.- Z. mikrosk.-anat. Forsch., 1933, 34, 485-533.
Watzka M. Yom Paraganglion caroticum.- Anat. Anz., Erganzungsh., 1934,
78, 108-120.
Watzka M. Dber die Elltwicklung des Paraganglion caroticum der Saugetiere.-
Z. Anat. Entw. Gesch., 1938, t08, 61-73.
Watzka M. Die Paraganglien. Handb. mikrosk. Anat. Menschen. Berlin, Sprin-
ger, 1943,6, Teil 4, 262-308.
Watzka M. Dber die Paraganglien in der Placa ventricularis des menschlichen
Kehlkopfes.-Acta Anat., 1966, 63, 300-308.
Werner F. Amphibia.- Handb. Zoologie (Kukenthal's), Berlin, Lpz, 1930, 6,
HaUte 1, 2-38.
West C. B. The comparative pharmacology of the suprarenal medulla.- Quart.
Rev. Bioi., 1955, 30, 116-137.
West G. B. Insulin and the suprarenal gland of the rabbit.- Brit. j. pharma-
col.Chemotherapy, 1951, 6, 289-293.
West G. R., Shepherd D. M., Hunter R. B., Macgregor A. R. The function .of
the organs of Zuckerkandl.- Clin. Sci., 1953, 12,317-325.
:344
Wetzstein R. Elektronenmikroscopische Untersuchungen Nebennierenmark
yon Maus, Meerschweinchen und Katze.- Z. Zellforsch. mikrosk-anat.,
1957, 46, 517-576.
lWeymann M. F. The beginning and de1velopment of function in the supprare-
nal medulla of pig embryos.- Anat. Rec., 1922, 24, 299-308.
!White E. Die Structur des Glomus caroticum, seine Pathologie und Physiolo-
gie, und seine Beziehung zum Nervensystem.- Beitr. pathol. Anat., 1935,
.96, 177-227.
lVhiting H. P. Nervous structure of the spinal cord of the young larval Brook-
Lamprey.- Quart. J. microsk. Sci., 1948, 89, 359-383.
Wiesel 1. Dber die Entwicklung der NebennierCi des Schweines, besond€l's del'
l\larksubstanz.- Anat. Hefte, 1901, 16, 115-150.
Wiesel 1. Chromaffine Zellen in Gefiisswiinden.- Zbl. Physiol., 1902a, 16, 2-
3.
Wiesel 1. Beitrage zur Anatomie und Entwickillug der menschlichen Neben-
niere.- Anat. Hefte, 1902b, 19,481-522.
.Wiesel 1. Deber die Erkrankungen der Koronararterien im Verlaufe akuter
Infektionskankheiten.- Weiner. Klin. Wschr., 1906, 723-725.
Willier B. H. A study of the origin and differentiation of the suprarenal gland
in the chick embryo by chorio-allantoic grafting.- Physiol. Zool., 1930, 3,
201-225.
Willis A. G., Tange J. D. The argentophil cells of the human carotid body.-
Amer. j. anat., 1959, 105, 141-163.
Wilson G., Billingsley P. The innervation of the carotid body.- Anat. Rec.,
1923, 25, 391-4.00.
Winckler J. Zur Lage und Function der extramedlllliiren chromaffinCtn Zel-
len.- Z. Zellforsch. mikrosk. Anat., 1969, 96, 4 0-494.
Winilcarter H. de. Contribution a l'etude de 1'0vaire humain.- Arch. bioi.,
(Paris), 1910,25,683-756.
Winiwarter H., de. Origine et signification du ganglion carotidien. C. r. As;!,.
Allat. 1934, 29 th reunion, 519-528.
Winiwarter H., de. Or gine et devcloppement du ganglion carotidienne. Ap-.
pendice: participation de l'hypoblaste a la constitution des ganglions cra-
niens.- Arch. bioI. (Paris), 1939,50,67-94.
. Wirtz B. Dber das Vorkommen yon Glomera im Herzeu des Menschen der
Taube und des Huhns.- Arch. Kreislaufforschung, 1968, 56, 89-105.
Wood J. C. Identification and observation on epinephrine and norepinephri-
ne containing cells in the adrenal medulla.- Amer. J. Anat., 1963, 112,
285-303.
Wood J. G., Barrnett R.I. Histochemical demonstration of norepinephrine at
a fine structural level.- J. Histochem., Cytochem., 1964, 12, 197-209.
Wrete JlI. Be,itrage zur Kenlltnis der Entwicklung des chromaffinen Geweues
der Bauchregion :"eim Menschen.- Z. mikrosk.-anat. Forsch., 1927, 9,
79-98.
Wright A., Chesler lones I. Chromaffin tissue in the lizard adrenal gland.-
Nature, 1955, 175, 1001-1002.
Wright A., Chester Jones T. The adrenal gland in lizards and snakes.- J. En-
docrinology, 1957, 15, 83-99.
Wurtmann R., Axelrod 1., Tramezzani 1. Distribution of the adrenalineforming
enzyme in the adrenal gland of a snakes Xenodon nwrrimii.- Nature,
1967, 215, 879-880.
Wyhe 1. W., van. Dber die Mesoderm-segmente des Rumpfers und die Ent-
wicklung des Exkretionssystem bei dem Selachiern.- Arch. mikrosk. AnaL.,
1889,33,461-515.
Yaron Z. The chromaffin and interrenal cells of Acanthobrama terrae-sanctae
(Cyprinidael, Teleostei).- Gen. compo Endocrinol., 1970, 14, 542-550.
Yates R. D. An electron microscopic study of the effects of reserpine on ad-
renomedullary cells of the syrian hamster.- Anat. Rec., 1963, 146, 29-45.
Yates R. D., Shen 1. L., Duncan D. Effects of sinus nerve stimilation on caro-
tid body glomus cells.- J. Cell. mol., 1970,46, 544-552.
345
Yates R. D., Wood J. G., Duncan D. Phase and electron microscopic observati-
ons on two cell types in the adrenal medulla of the syrian hamster.- Te-
xas, Repts. BioI. Med., 1962, 20, 494-502.
Yntema Ch. /., Hammond W. Depletions and abnormalitis in the cervical sym-
pathetic system ow the chick following extirpation of neural crest.- J.
exptl. Zoo!., 1945, tOO, 237-259,
Young J. Z. On the autonomic nervous system of the teleostean fish Urano-
scopus scaber.- Quart. J. microsk. Sci., 1931, 74, 492-535.
Young J. Z. The autonomic nervous system of selachians.- Quart. J. microsk.
Sci., 1933, 75, 571-624.
Young J. Z. Partial degeneration of the nerve supply of the adrena1. A study
in autonomic innervation.- J. Anat. (Lond.), 1939, 73, 540-550.
Young J. Z. The life of vertebrates. Oxford, Clarendon press, 1950.
Zapata P., Hess A., Bliss E., Eyzaguirre C. Chemical electron microscopic an.-I
physiological observations on the role of catocholamines in the carotid
body.- Brain Res., 1969, 14, 2, 473-496.
Zettergren L., Lindstrom 1. Glomus tympanicum. Its occurence in man and its
relation to middle ear tumour of carotid body type.- Acta pathol. micro-
bioI. Scand., 1951, 28, 157-166.
Ziller-Perez H. V. On the chromaffin cells of the nerve ganglia of Hirudo me-
dicinalis Linn.- J. Compo Neurol., 1942, 76, 367-401.
Zimmermann W. Ober die Karotisdruse von Rana esculenta. Diss. Berlin, 1887.
( IIT. no Adams, 1958).
Zuckerkandl E. Dber Nebenorgane des Sympatiens in RetroperitonE...alraum des
Menschen.- Anat. Anz., 1901, 19, 95-107. 1901, 19, 95-107.
Zuckerkandl E. The developml'nt of the chromaffin OI'gans and of the supra-
renal glands. Manual of Human Embryology. USA, Lippincot, ed. Keibel
and Mall's, 1912,2, 157-179.
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