Биология. Новейший справочник - Чебышев Н.В., Гузикова Г.С., Лазарева Ю.Б., Ларина С.Н.

Author: Чебышев Н.В. Гузикова Г.С. Лазарева Ю.Б. Ларина С.Н.
Tags: биология справочник зоология естественные науки естествознание издательство махаон
ISBN: 5-18-001064-0
Year: 2007
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Jfa3apesa 10.E., J1ap1ma

Co~epmaHHe
BBE,UEIUiE
YpoBHH opraH11aanm1 >K11Boii :viwrepHH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . 10
OcHOBHhie cBoiicTBa nornhIX opraHH3MOB . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

OBIIV\H J-3110.TIOnrn:
OCHOBbl I ~I1TOJIOfl1I1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
MeTOJ\bl H3Y'ICH!1H Ko1€T!Of . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
061qce CTpOCHHe K.1€Tl{H . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
XHMW!eCKHiI COCTall T«1CTKH . . . . . . . . . • . . . . . . . . . . . . . • . . . . • . . . . . . • . . . . • .
Opranw1ecK1te BeII\CCTBa KJieTKH . . . . . . • . . . . . . . . . • . . . . . . . . • . . . . . • . . . . . . .
BeJ1KH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . • . • . . . . . . . . • . . . • . . .
<lJepMCHThl . • . . . . . . . . . . • . . • . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . • . . . .
JlHTIH,l\hl . . . . . . . . . . . . . • . . • . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . .
Yr,1eBOJ\hT . . . . . . . . . . . . . . . . . . . . . . . . . . . . • • • . . • . . . . . . . . . . . . • • . . . . . . . .
HyKJICHTIOllh!C J{HC.TIOThl . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . .
,ZJJ-IK (,l\eao1<c11p1160HyK,1emroBa.H 1rnc.1orn) . . . . . . . . . . . . . . . . . . . . . • • . . . . . . . . •
PIIK (pn6ony1G1c11HOBaR KHCJTorn) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
AT<P (a,l\eTI03HHTpmllOCQ>opna.H 1mc.10Ta) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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HeK:1cT01 111h1c cpopMbI n<H3HH . . . . . . . . . . . • • . . . . . . . . . . . . . • . . . . . . . . . • . . . . 30
B:upyChl . . . . . . . . . . . . . . . . . . . . . . . . . . . . • • . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

KJrcTO'lllhIC cpop:vib1 n<H3HH . . . . . . . . . . . . . • . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . 34
Ha,l\I~apcTBO upoKapHOThI. . . . . . . . . . . . . • • . . . . . . . . . . . . . . . . . • . • . . . . . . . . . . 34
Ha,l\napcTBoayKap110Thl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . 36
06MeH llCII\CCTB 11 IIpeBpaII\Cl!He 3Hepr11H . . . . . . . . . . . . . • . . . . • . . • . . . . . . . . . . 48
lPOTOCHTITC3 . . . . . . . . . . • • . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . • . . . . . . . . . . . 50
XcMOCHHTea . . . . . . . . . • • . . . . . . . . . . . . . . . . • . . . . . . . . . . . . • . . • . . . • . . . • . . . 55
8HepreT11qec1<11n o6:vieH . . . . • . . . . . . . . . . . . • • • . . . . . . . . . . . . . . • . • . . . . . . . . . 55
,IJ;e.1eH11e KJTeTKH . . . . . . . . . . • . . . . . . . . . . . . . • • . . . . . . . . . . . . . . . . • • . . . . . . . . 58
lIHTcpcpa3a . . . . • . . . . . . . . • . . . . • . . . . . . . . . . • • . . . . . . . . . . . . • . • • . . . . . . . . • 59
MHT03 . . . . . . . . . . . . . . . . . . . . • . • . . . . . . . . . . . . . . . • • . . . . . . . . . . . . . . . . . . . . 59
Mefio3 . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . • . . . 61
Pa3MHO>KCH11eopraHH3MOB . . . . . . . . . . . . . . . . . . . . . • . . • . . . . . . . . . . . . . • . . . . 64
Becrronocpa3:viHon<enuc • . . . . . . . . . • . • . . . . • . . . . . . . . • • . • . . . . . . . . • . . . . . . . 64
IlOJTOBOC pa3MHO>K€HHe • . . . . . . . . . . . . . . . • • . . . . . . . . . . . • . . . . . . . . . . . . . . . • 66
faMeTOI'CHC3 . . . . . . . . . . . . . . . . . . . . . . • • • . • . . . . . . • . . • . • . . • . . . • . . . . . . . • 67
I1H,ll.HBH,l\ya,1bHOC pa3BHTHC opraHH3Ma •••.......••..••...•.........•.•. 72
IlOCT3M6p11ona.1bHOe pa3BHTHC . . . . . . . . • • . . . . . . . . • . . • . ..••.••....••.... 79
fEHETJIKA
OcHOBHhlC IIOHHTHH :u onpeJJ;e,1en11H reneTHKH. 3aKOHhl r. MenJJ;e,1H .•••.....•. 81
Baa:uMoJ(eAcTB11ere110B . . . . • . . • . • • . . . . . . . . . . • . • . . . . . . . . . . • . . • • . . • . . • . 86
Cu;enJiennoc Hac.1e,l(oBaHHe. feHeT:uqec1we onpeJJ;eJTe11uc no.1a. ll;ttToreneT:uKa •.• 91
MoJieKyJIHpHaHre11cTHKa ..•..•.•..........••............•.•••..••.•.. 97
PerrJI:uKaQHH ,UHK ••.....••..•.•.........••...........••.••...•...•. 98
PeaJittaar~:uH reHeTwICcKofi 11H<)JopMau;m1 B KneTKe. Bttoc11HTC3 6eJTKa ..•..•... 99
TpancKp11rru11H ...•........•.•.••.••.......•••......••.••..••.•... 100

4

CoiJepJ1Canue

fipouecc1rnr sy1rnp1wT1P1ec1-rnx H-PHK . . . . . . . . . . . . . . . . . . . . . • . . • . . . . . • . .
TpaHc"1HUHH . . • . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . .
MenHUHHcKaareHCTHKa . . . . . . . . . . . . . . . . • . . . . . . . . . . • . . . . . • • . . . . . . . . .
OcnoBHL!e '.\1.eTO/lbI 11ayqeHH.H renen1K11 qe;10BcKa . . . . . . . . . . . . . . . . . . . . . . . . .
feHeanorH'!CCKHft MPTon . . . . . . . . . . • . . . . • . . . • . . . . . . . . • . . . • . . . . • • . . . . .
QHTOreHCTH'leCKHH MCTOn . . . . . . . . . • • . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . .
feHeTHKa COMBTH'leCKHX KJICTOK • . . . . . . . • . • . . . . . . . . . . . . . • . . . • . . . . . . . . •
MeTO.L\ .u;epMaTor1m<jlnKH . . . • . . . . . . . . . . . . • . . . . . . . • . . . . • . • . . . . . . . . . . . •
BJu13Heu.oah1tt MeTon . . . . . . • . • . . . • • . . . • . . . . . . . . . . . . . . • . . . . . . . . . . • . . •
BnoxHMH'lecKHe MeTonhr ...•.•..•.•.•..•............•••.......•.•.•.
IlonyJl.f!ll,l!OHHO-CTaTHCTl!'!CCKl!ll MeTon . • . . . . • . . • . . . . . . . . . . . . . . . • • • . • . .
MeTOALI npenaTaJihHOtt (Bo ape MR 6epeMCHHOCTH) .u;HarHOCTHKH ..•...•....•..
lfaMeH'IHBOCTh. Tmrhr H3MeH'!HBOcTH. Pe11apa1ou1 .IJ:HK •.•.....•.•...•..•..
feHeTH'leC!(HC OCHOBbI CCJ!eK.UHH )fUJBOTHLIX, pacTeHHH,
Ml!KpoopraHH3MOB. BnoTexuonorHH •••.........••..........•......•.•.•
Ce,1eKU.HH pacTenntt ..••.•..•.•....•.•....•..•...•......•.....••...•
CeJieRU.HH mnBOTHhIX ....•..•.•.•...........••......•...••..•.•....•
CeJJeRll,IIH MlmpoopraIIH3MOB (B OCHOBHOM 6aRTepttii H rpH6oa) ••••••....•...
BttoTexHonorH.H. fenHaH ilIDKeHeptt.H •..•.•.•.••...•.•.•.••........•..•
IJ;l1TOIIJI83MaTw1ec1we nacJie)J;OBaHl!e •••.•.•...•..••..•.•.•.••.•.•..•..•
3BOJUOU:ll.H
Bo3HHKnosenll.c muanu na 3eMne. Pa3Bl!Tl!e opranuqecKoro M11pa .•....•...•
Pa3Bl1Tile mtt3HH na 3eMJie ........••.....•...•.•.•......•••..•.•.•..•
OcHoBnLre :JTanbl <iBo,110u.ttu opranll.'!ecKoro Mttpa .•......••...•........•.
Kpynnei1:nrne apoMop<jlo3bI B ::iaon10r~Hll opranuqecKoro MHpa .•..•........•.
.IJ:oKa3aTeJibCTBa 3BOJII0ll,llll ...•..•.•..•.•...•.•.•.....••.•.•....•....
llaneoHTOJJOrH'ICCKHC,[\OKa3aTeJibCTBa .•......•.•.....•.•..•...•..•...•
BuoreorpacpttqecKne ;:i;oKa3aTeJihCTBa . . • . • . . . • . . • . . . . . . . . . . . • • . • . • . . . • .
Mop<jlonorll.'lecK11e;:i;oKa3aTeJILCTBa .••.•....•......••.•..••..•..•...•.
3M6puonoruqecK11e noKa3aTeJihCTBa ..••.....•...•.•.•..•....•.....•....
feHeTtt'leCKHe AOKB3aTeJibCTBa •.•...••.•...•.......•.• .- ...•..•.•...•..
BHOXHMl!'!CCKHe l! MOJICRyJIHPH0-6HOJIOI'H'!CCKlle noKa3aTeJihCTBa •.•.•..•..•
Ilpoucxomnentte qenoseKa ..•......•.•...••..•..•......•.•.•.....•...
llaneoHTOJiorll.'lecK11e noKa3aTeJILCTBa npoucxomnenu.H
'leJIOBeKa. 3BOJIIOI~HH npHMBTOB .•.•.•.•.•..•.•.......•.........•...•.•
Pa3Bl1Ttte BLrcrnnx rrpll.MaTOB •.••.•.•..••.•.••.•.•.•.•....•......•....
0CHOBHhie CTBAHH ::JBOJUOI~llll qeJIOBCKa ...•••.••.•.•.•...••.•......•....
IlttTCKBHTPOTibI (npeBHeiinme JIIO,[\ll, apxaHTpOIILI) •.•....•.•.•.•..•.•..•.•
Hean.n;epTaJihll,LI (.upeanue nro;:i;u, naneoanTponLr) .••.....••...•..•.....••
qeJIOBCK COBpeMeHHOI'O THITa, llCROnaeMblll ll cospeMeHHblH (neoaHTpOITbI) .••.
CoapeMeHHhIH 'lenoae:K u ero 3BOJIIOIJ:I1H (ueoanTponLr) ...•.••.•...•..•..•.
.IJ:o.uapBHHOBCKllll rrepHOA pa3Bl!Tll.II 6HOJIOI'llH ••..•..••.•.•..•...•.•..•••
Tpanc<jlopMtt3M ..•..•.•••.••.•••••.•.•.•...•••••.•••.•••.•.••...••
llepsa.11 u.enocTna.11 Kouu.enuu.11 :3Bon10u11u. JK. B. JlaMapK ••...••..•.•..••.•
3soJIIOll,ilounoe yqenll.e qapnaa .IJ:apsuna •...•.••••.•••.•..••.••.•...•••
Ml1KP03BOJIIOll,ll.11 ••••.••••••..•.•••••••.••••••••.•.•.••..•••••.••.•
llorryJIHll,HH - 3JieMeHT8pHa.H CllCTeMa 3BOJIIOUHH .•.•..••.••.•.•.•.••••••

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CoiJepJtCanue

cf>aKTOpbl 3BOJIIOUH11 . . . . . . . . . . . · · . • . . . . . . . . . . • . . • • · · · · · . . . . . • . . . . . .
OcHOBHOH 3Tarr 3BOJIIOUlIOHHOro npOilecca - Bll.ll . . . . . . • . . . • • . . . . . . . . • . . . .
MaKp03BOJIIOHllH . . . . . . . . . • . . . • . . . . . . . . • . . . . . . . . . . . . • . . . . . . • . . . • . . .
OcHOBHhIC nanpan.11eH11H 3BOJI]{)I~llll . . . . . • • . • . . . . . . . • • . • . • . . . . . . • . . . . . . .
OcHOBHhIC rro.11o;irnHHH Cl!HTeTnqecKott Teop1111 3BOJIIOilHH ••......•....•.•...
CaJibTailHOHnaH TeopnH ;:momou1111 .•.•........••••.•.......•....•.•..•
ConpeMeHHa.H Teopn.H 3no1110uu11 . • . . • . . . . . • . . . . . . • . . • . . • . . • . . . . . . . . . . • •

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3KOJIOfIIH
ITonHTHH «cpe.11.a 06J1TaHHH» 11 «3Ko.11or11qecKne cjJaKTOpbI» . . . . . . . • • . . . . . . • .
3K0.110f'HH rrorryJIHUllll . . . . . . . • • . . . . . • • . . . . . . . . . • . • . . • . . . . • • . . . . . . . • .
3K0.110f'J1qecKaH xapaKTepHCTHKa BH,Ua .•.......••••.••......•...•..•...
BnoreoileHoa • . . . . . • • • • . . • . . . . . . . • . . . . . . . . • • • . • • . . . . . . • . . . . • . . • . . .
CTPYKTypa6noreoueHoaa • . . • . . . . . . • . . . . . . . • • • . . . . . . . . . . . • . . . . . . . . . • .
IToKaaaTe.'ln cpyHKUHOHnponan11.H 3KOCHCTCMhI • • . . . . . . . . . . . . • . • . . . . . . . . • •
Arpouenoah1 • . . . • • • . . . . . . . . . . . . • . • . . . . . . • . . • . . . . . . • . • • . . . . . . . • . . . .
BnoccjJepa • • • . . . • • . . • . . . . . . . • . . • . . . . . • . . . . . . . . . • . . • . . • . . . . . • . • • . . .
JK1rnoe BClllCCTBO ll ero cj:>yHKUHH ..••......•.......••.•.••...•.•..••...
,Il;eHTeJihHOCTh 'IeJIOBeI<a. HooccjJepa •......•••.......•.•.•..•.•••..•...•
BmrnHHe .11.eHTCJihHOCTH •rn;wneKa Ha cTa6H"'lhHOCTh 6J1occjJeph1 ..•...•...•.•.•
OcHOBHhre 31rn110rw1ecKne rrpo6.11eMhI . . . . . . . . . . . . . . . . • . • . . • . . . . . • . . • • • .

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BOTAHIIKA
T1<aHH 11 opraHhI BhIClllHX pacTeH11il . • . • • . . . . • • . . . . • . . . . . • . • . . • . . . . . . . .
BereTaTllBHhre opraHhI (Kopen1>, cTe6e"1h, Jll!CThH) . . . • . . . . . . . • . • . . . . • • • . . .
MopcjJo.1orw1ecKoe cTpoeH11e Kopn.H. KopHeBa.H c11cTeMa . . . . . . . . . • . . . . • • . • . .
AHaTOMJ1qecKoecTpoeH11eKopHH • . . . . • . • . . . . . . • • • • . • • . . . . • . • . . . . . . . . . .
CTe6e.11b . • . . . . . . • . . . . . . . . • . . . . . . . . . . . . . . . . • • . . • • . • . . . . • . . . . . . . . . .
JlHCT .... ,,, .. ,,,, •...... ,,,, •..... ,,, . . . . . . , .•.•..... , •••.....••
feHepaTHB!Ihie opraIIhI (UBCTOK, rr.110.u, ceMH) .........••..•....•••....•••
IJ;BeTOK .. , , , ... , , , , , ...... , , , ..•.. , , , , •....... , •.•.... , ••..••.••.
fIJIO.U, , .. , , , ... , , , , , , . , .... , , , , , •.. , , , , , ..... , , , , , , , ... , , , , , ••...
CeMH . • . . . . • • . . . • • . . . • • . . . . . . . • . • • . . . • . . . • . . . . • • . . . • . . • . . . • • . . . . .
KJ1accncjJ11KaUHH 110Kpb1ToceMeHHhIX pacTeH11tt ••.......•••.•..••.....••.
OcHOBHhre ceMelicTBa K.11acca ,Il;ny.uo.11hHbre
(KpecTOilBCTHhie, PoaouneTHhie, Bo6onh1e, ITac.11eHOBb1e, C.11omHouneTHhre,
MaJihBOBhre, MapeBbie, BHHorpa,nHbre) ••.......••••.•....••.•.....••...
CeMelicTno KpecTouneTHhie (KarrycTHhre) ......••••.••...•••.........•..
CeMelicTBO Po30ilBCTHhre •..•••....••......•.•...••...•.••...•......•
CeMeil:cTBO Bo6oBbre (MoThIJThIWBhre) • . • • . . . . . . . . . . . . . . • . . • . . . . . . • . . • . •
CeMelicTBO ITac.11enoBb1e ..••......•.••••...•.••••.•..•.••.••..••••...
CeMelicTno CJ1omnouBeTHhre ..•••....•...•...••••..•....••.•.....•••..
OcHonnwe ceMeiicTBa KJ1acca 0.llHO.UOJihHhie (JI11.11eliHble, 3.11aKOBh1e) ••......•
CeMeil:cTBO Jiu.11ei1:1Ible ..•.•••.•......•••..•••...••...••.•.••...•..•.
CeMeil:cTno 3"1aKH, 11J111 MHTJIHKOBbie •.•••...•....•....•..••••...••.•.•
Ha.uuapcTno ,Il;oH.llep11b1e, HJIH ITpoKapn:oThI . . . . . • • • • . • . . . . . . . . . . . • . • . • . .
BaKTepHH •.•...•••........•.•....•••...••.•••.•••.......••.....••
IJ;apCTBO I'ptt6br ...•...•..••.•.•...•..•...••••.•••........•....•...
0TJJ;eJI Jl11lllaHHllKH .•.....••.•••...••••..•••...•..•....•.••..•..•••

1 76
183
184
185
188
192
195
195
200
201
203

204
204
205
205
206
207
208
208
209
209
210
212
216

6

Coilep:Jl(;anue

napCTBO PacTemrn. JlO[\UapCTBO HacT05111~11e flOi~OpOCJil1 . . . . . . . . • . . . . . . . . . . 218
OT l\eJI 3eJICHhIC BO,'l;opoc;rn . . . • . . . . . . . . . . • . . . . . . . . . . . . . . . . . • . . . . . • . . • 218
0T[\enBypweBOAOPOCnl1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • 220
OTJ\C.'l KpacnhIC BOl\Opocmt, 11.'IH Barpmmn . . . . . . . • . • . • • . . . . . . . . . . • . . . . . . 221
II01v~apcTBO Bhicmuc crwpoBbie pacTemrn . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . 223
OTJ\CJI MoxoBH,UHbie . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . • . . . 223
K.'!acc JI:ucTOCTe6e.'lbIIhie MXH, IIOAI<Jracc Bc.1b1e MXIL CcjiarttyM ..•.•.......• 225
0TAeJI IlarropOTHHKOBlfl\l!bie . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . • 225
0Tp;eJI TI;rayHOBHAIIbie . . . . • . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . 228
0Tp;en XBou~eBH,ll,HhIC • . . . . . . . . . . . . . . • . . . . . . . . . . . . • . . . . . . . . . . . • • . • . . . 229
CeMCHHbIC pacTcH11n ....•.....•...••.•..••.•.•..•...........•..•..• 230
0Tp;CJI foJ!OCCMeHIIbJe . . . • . . . • . • . . . . . . . . . • . . • . . . . . . . . . . . . . • . . • . . . . . • 230
0Tp;eJ! IloKpbITOCCMCIIHhIC, 11.'IH IJ)eTIWBh!C • . . • . . . . . . . . . . . • . . . . . . . . . • . • . 233
300JIOI'IUI
KJ1acc:ucjnmautt.H mttBOTHoro MMpa . . . . . . . • . . . . . . • . . . . . . • . . . . . . . . . . . . . . 237
Ilo;:i;uapcTBo O;:i;HoKJieTo'-IHhIC mHBOTHh1c, uJI:u TipocTeilurne ...•...•.....•.. 237
06rua.H xapaKTepHCTHKa npOCTCHllIHX ..•....•....•.•......•.....••..•.• 238
TttrrcapKoMaCTHrocjJophl .•...•.•.........•..•...•....•.•..•.....•..• 240
Tttn arrHKOMUJieKca •.•.•......•.•..•.•....•..•••....•....•.......•. 248
Tnrr HHcjJy3opHH (pecHH'-!Hb1e) • . . . . • . . . . • . . . . • . . . . . . . . . . . . . . . . • . . . . . • . 249
Tio;:i;u.apcTBO MnoroKJJeTO'-!HhIC . . . . . . • . . . . . . . . . . . . . . . . . • . • . . . . • . . . . . • . 251
Tttn ry6K:u . . . • . . . . • . . . . . . • . . . . • . • • . . . . . . . . . . . . . . . • . • • . . . . • . . • . . . . 251
Tttrr KHUie'IHOIIOJIOCTIIhIC . . . . . . • . . . . • . . . . . . . . . . . . • . • . . . . • . . . . . . . • . . . . 252
KJiacc fttAPOH,ll,Hhie . . . . • . . . . . • . . . . . . . . . • . . . . . . . . . . . . . . . . • . . . . . . . . . . 253
KJiacc Cu.mpOHl\Hhre, :u.'!:u C1~ttcjJOMep;y3hI . • . • . . • . . . . • . . . . . . • . • . . . . . . . . . . . 256
KJiaccRopaJ1noBwerroJittllhl . . . . . . . • . . . . • . . . . . . . . . . . . . • . . . . . . . . . . . . . . 257
Tttrr UJIOCK11e '-!CPBH • . . . • • . . . . . . . . . . . . . . . • . . . . • . • . . . . . . . . . . . . • . . • . . • 258
KJiacc PecHH'-!Hhie '-IepBtt • . . . . . • . . . . . . . . . . . . • . • . • . . . . . . . . . . . . • . . . . • . • 261
Knacc Cocanh~HKH, :u.1n Tpel\rnTOAhI. . . . . . . • . . . . • . • . . . . . . . . • . . . . . . . . . . . 263
Knacc JleHTO'IHhIC qepB11, u.rrn 1~ecTo.n;b1 . . . . . . . . . . . . • . . . . . . • . • . . . . . . . . . . . 265
Tnrr KpyrJihIC, ttJIH ncpBH'-!HonoJiocTHhJe, '-ICpBH . . . . . . . • . • . . . . • . . . . . . . . . • . 268
KJiacc HcMaTOAhI, l1JIH co6cTBCHHO J<pyrJihIC qepBH •..••..•..•.•..•.•...... 270
THrr KOJih'-laTbJC '-IepBH, HJ!lf KOJ'Ib'-ICI~bl .•.•..•..•.........•.••.•.•.....• 273
KJiacc MaJio~eTHHKOBbie qepB:u (o.'!:uroxeThI) •..•.•..•.......•.•....•.... 275
KJiacc MHoro~eTHHKOBhie '-!CpBH (rroJittXeThI) ..•.•..•..•.••••.•.....•... 278
KJiacc II11nBI<H . . . . . . • . . . . . . . . . • . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . • • . 279
T11n MOJIJIIOCKH, :u.'!H MHl'KOTe,'lh!e •.•.•.......•...•....•.•...•...•.••. 280
KJiaccBp10xoHorueMoJIJIIOCI<tt ..••.•.•.....•.........•......•.•...•.• 284
KJiacc ):J:BycTBOp'-!aThre, HJIH IlJ1aCTHH'-!BToma6epHbie .•..•...•.....•....•.. 286
KJiacc foJI0Bonor11e MOJIJIJOCKH ......•..•.•.•......••.•.......•.•.••.• 288
T11rr'-!JieH11cTonor11e •...•....••........•..•.•.•..• . •...•......•.•.•• 289
KJiacc PaKoo6pa3Hhie .••.•.••..•.•.••.•....•...•..•.....•.•.•..•..• 292
KJiacc Tiay1<006pa3HbW • . • . . . . . . . . • . • . . . . . . • . . . • . • . . . • . . . • . . . . . . . . . • 296
KJiacc HaceKOMhre ......•.....•..•.•.••.•..•.•.•..•.......•.•..•.•• 298
T11rr xopp;OBhie ..••....••.•.•..••.•.......•....••.....•...•.•..•.•. 309
JlH'-!HHO'-!HOXOp,!J;OBb!e ..•......••..•.......••...•..•........•••.•.•. 310
KJiacc RpyrJiopoTb!C •...•...•..•..•.....•....•.•.......•.••...•...•. 314

7

CoiJepJ1Canue

317
326
JCwcc Ilpec:vihIKRIOII\HCCH (pCIITH,11111) • • . . . . . . . • . • • . • • . . • . • • . • • • • . • . . • • 333
JC1acc I!THllhI • . . . • • . . . . . . . • . . . . . . . . . . . . • • • • • • • • • . • • • • • • • • . • . • . • . • • 339
KJrace M.1e1rnm1TaIOll\He (3Bep11) . . . . • • . • • • • • • . • • • . • . . • . • . . • . . . • • . • • • . . 351
CpaimttTCJibHaH attaTo:v11rn cncTeM opranoB II03BOHO'!HbIX . • . • • . . . . • • . . • • • . • 364
<PirnoreHe3 Hapym:Hh!X IIO!<pOBOB . . . . . • • . . . . . . . • • • • . • • • • • . . . . • • • • • • • • • . 364
<P11JIOl'CIIe3 Cl{C.'IeTa . • • • • . • • . . . • • . . . . . . . . . . • • • • • • • . • • • • • • • . • • • • . . . • . 369
.I(mpcpepelll\HPOBKa IIOaBOHO'lHHK:J B np01~ecce 3BOJIJO~Ifl1 . . . • . . • . . • . • . . . . • . 369
3BOJI!Olil1H '!Cpena Il03BOHO'f!Ih!X • • • . • • • • • • . • . • . . . . • . . . . • . . . . . • • . . • . . • • 369
3BOJ110lil1H pa3Bl1Tl1H IIllTHIIaJIOH IW!Ie'fl!OCTH • • • . . . • • • . . • • . . . . • • • . . • . • • • 370
<P11JioreHe3 IIHll\eBap11Te.1LIIOH CHCTC:vlbl • . • • • • • . . • • • • • • • • • • • • • • • • . • • . . • • 370
<Ptt.'!Oreuea ,ll,bIXaTe.1hl!OH CliCTC'.ll.LI . . • . . • . • • • • • • • • • • • • • • • • • • • • • . . • • . . • • 372
<PttJIOreuea KpOBeHOCllOH Cl1CTC'.ll.LI. . • . . • • • • . • • • • • • • • • • • • • • • • • • • . • • • . • • • 3 73
3BOJIIOl\IUI COCY,ll,OB . • . . . • • . . • • • . . . . • • • • • • • • • . . • • • • . • • • • . • • • . • • . . • • • 373
Ha,ll,K.'Iacc pb16b1 . • . . • • . • • . • . . • • • • • • • • • . . . . . . • . . . • . . . . . . . • . . • • • • • • . .

ICwcc 3eMHOBO/J,IIbIC (a:viqm6m1) • • . • . • • . . . . . . . . . . . . . . . . . • . . . . . . • • • • . • .

<PttJiore11e3 nepmrni1 c11cTeMbI . . • • . . • . • • • • • • • • . . • • . • • • • . • • • • • . . • • . . • • •
CTpoenne roJ10nnoro M03ra noaBOHO'!HbIX • • • • • • • • . • • • • • . • . • • • • • • . • • • . . • .
<Ptt.'IOl'Cl!C3 opral!OB '!YBCTB . . • . • . . • . • • • • • • • • . . • • • . • • . . . • • . • • . . • • • • • • •
<PttJIOl'CHC3 Bhl,ll,CJI!1TCJibHOH 11 no.1osoii CHCTeM

.••••..•.•.••.••..•••••.••

CBH3L Bhll\CJIHTCJILHOH l1 IIOJIOBOH CHCTeM • • • • • . . • • . . • • . • . • • . • . . . • • • • • . . •
AHATOM11H

11

3 74
3 74
376
377
379

<Pl13I10JIOrIUI "l!EJIOBEKA

••••••..•...•...••.••••••.•••

380
385
396
397
400
403

CTpyKTypuo-cpyHKn11onaJibHh1e qacT11 nepm10U: c11cTeMbI . • . • • . • • • • • . • . . . • • •

404

TKaHH . . . . • • . . • . • • • • • • . . . • . . • • . . . . . . • • . • • • • • • • • • • • • • • • • . • . . . • • • . •
OnopHO-,l\Bl1raTe.'lbHbIH armapaT . . • . • • . . . • • . • • • • • • • • • • • • • • • • • • • . . • . . . •
Mbunel!HaH c11cTeMa

•••••••...••.•..•.•••••.•••••••.••••••••....•••

CTpoeHHC MbillllibI • . • • • • . • . • • • • • • • • . . • • • • . • . • • • • • . • • • • • • • • • • • • . . • • •
HepmiaH c11cTeMa

...••••••••••••••••••••••.•.•••...•..•.••.•••••••

Ilepe,llalJa B036yn<,ll,eHl1H nepBHOH IWCTKOH

405
407
<PyHKlil111 OT,ll,eJJOB l'OJ!OBHOf'O :vioara • • • • • • • • • • . • • • • . • • . • . • • . • • . . • • . . • • . 408
IIepmpepn'IeCKaH IICpBHaH CHCTe:.i:a . • • . • • • • • . • . . . • • . • • . • . . • . • . . . • • • • • . . 411
CocTaB pecliJieKTopnoU: ,ll,yr11 . . • . • . . • . • • • • • • • • . • • • • . • • . . • • • • • • . . • • • • • . • 413
Bbrcll!aH HepsHaH ,ll,eHTC.'lbHOCTb • . • • . • • • . . • • • • • • • . • • . • . • • • . • . . • • • • • • • • 416
Mexatttt3M 06paa0Ban11>1 ycJIOBHoro pecjJJieKca • • • • • • . • • . • . • • • • • • • • • • • • • • . 41 7
Bo36ym,ll,eH11e 11 TOpMom:enne yc.10BHbIX pecp.1eKcoB • • • • • • • . • . • • • • • • • . • • • . • 41 7
Y'-IeH11e 0 ,ll,BYX CHrHaJILIIb!X CHCTCMax • . . . • • • • • • • • • • • • . • • • . • • • • • • . . • • • • • 419
T11rrb1 BhICllleii HepBnoU: ,ll,CHTCJibHOCT11 . . . • • • • • • • . • • • . . . • • . • • • • • • . . • • • • • 419
Ilpo~eccLI IIaMHTH • • • • • • . • • . • . • • . • • . • • • • • • • • • • . • . • • • • . • • • • • • . • • . • • • ,121
BH,ll,hI naM>1T11 . . • • • • • • • • • . • • . • • . • . • . • • • • • • • • • • • • • . • • • • • • • • • • • . • ••• 421
Oprauh1 '!YBCTB • • • . • . • • • • • • • . • • • • • • • • • • • • • • • • • • • • • • • • • • . • • • . • . • • • • 422
OpraH apeHttH . • • . . • . . • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • . • . • • • . . • • • • . • 423
BcrroMoraTeJihllbIC opraHhI rJia3a • • • • • • • • • . . • • . • • • • • • • • • . . • . • • • . • • • • • • • 425
3pttTC.'lbHLiii anaJIH3aTop • • • • • • • • • • • • • • • • . • . . • • • • • • • • • . • • • • • • . • • • • . • • 426
Oprau cJiyxa tt paBHOBeCHH • • • • • • • • • • • • • • . . • . • • • • • • • • • • • • • • • • • • • • • • . • 427
3BYKOBOCITpl1HH!\1a!Ollll1H anrrapaT BHyTpemrero yxa • • • • • . . • • • . • • • • • • • • • . • 429
Oprau paBHOBec11>1 (BecT116yJIHPHhIH anrrapaT) . . • • • • . . • . • . • • • . • • . • • • • • . • • 431
Crr11uttoii M03r • • . • . . . . • • • • • • • • . • • • • • • • . • • . . . . • . . • . • . . • . • • • • • • . • • . •
foJIOBHOH M03r

•.••....•.••.•.••..•••••••....•..•.•.••.•••.....•••

8

CoiJep:H<:anue

OpraHhI o6oHmnrn YI BKyca . • . . . . . . • . . . • . . • . . . . • . . . . . . . . . . . . . • . . . . . . . . 432
Koma . . . . . . . . . . . . . . . . . . • . . . . . . . • . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . 433
KmKHhIH ana.1Y13aTop . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . • . • . . • . . . . . . . . . 435
BHyTpeHHHH ceKpeu,1u1 . . . . . . . . . . . . . . . . • . • . . • . . . . . . . . • . . . . . . . . . . . . • . 436
06Me!I Belll,eCTB H ::rneprirn . . • . . . . . . • . . . • . • . . • . . . . . . . . . • . . . . . . . . . . . . . . 445
06MeH BOAhl H MHHepa.'IbHblX COJierr . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . 446
06'.\feH 6e,1KOB . . . . . . . . . . . • . . . . . • . . . • . • . . . . . . . • . . • . . . . . • . • • • • . . . . . . 44 7
06MeH yr,1eBOl\OB . • . . . . . . • . . . . • . • • . . • . . . . . . . • . . . . . . . . . . . . . • • . . . . . . • 448
06MeH )fU!pOB ••••.•......• · . • . . . . . . . . . • . . • . . . . . . . . . . . . • . . . . . . . . • . 449
BHT3MIU!bi • . • . • . . . . . • . . . . . . . . . . . . . . . . . . . . • . . • . . . . . • . . . . . . . . . . . . . . 449
KpoBoo6pauleHtte . . . . . . . . . . • . • . . • . . . . • . . . . . . . . . . . . • • . . . . . . . . . . . . • . 452
CTpoeHtte RpoBenocHhIX cocyn;os . . . . . . • • . . . . . • . . . . . . . • . . . . • . . . . • . . . . . • 453
KpoBHHoen;aBJietttte ..•.••......•.•..•......•......•...•..••...••••• 456
PeryJI.HU,Yl.H ABHJKCHUH r<posu IIO cocy,ZJ,aM •.•..•.•.•....••......•.•...... 456
CTpoen11e cepAila •...••.•••...•...•.....•..•...•....••..••......•.. 457
BHyTpeHHH.H cpe,ll;a opraatt3Ma .•.•.......•..•..........••.....•.••.•.. 460
CTpoea11e JIUMcPaTII'!ecRoil c11cTeMh1 .•..•.••.•....•.....•......•......• 461
KposeTBopHbleopraHhl ..•............•.••.•....•...•••.....••.•....• 462
~OpMeHHbleaJieMeHTbIKpOBH ...••.••.•...........•.•..••••.•.•..•.•. 464
MexaHH3M csepThrnaHHH Kposu . . • . . • . . . . . . • . . • . . . . . . • . • • . • • . . . . . . . . . . 467
fpyrrnhI KPOBH 'Ie;IOBeJ<a ....•..•.•.•••...•..•.•.•.....•.•.•....•.••. 468
L\b!X3HllC . . • . . . . . . . . . . . . . . . . . . • . . . . . • . . . . . . . . • . . . . • . . . . . . . • . . . . . • 469
Mexantt3M s11oxa H BhI,noxa •..•...•....•....•.•......••..•....•.....• 4 72
fa3006MeH B JICI'KHX ll Tl<aJIHX . . . . . • . . . . • . . . . . . . . . . • . . • . . • . . . . • . . • . . . . 4 7-4
IlnuleBapeHtte •••.••.•.•......••.•...•.........•.•.......•......•. 4 76
fioJIOCTh pTa .•.••.•....•....•.•........••.•.•.........••••.••...•. 477
Ilrrlll,esapIITe.'Ibllbie meJie.1bl .......•.•..•.•........•....••.••...•.••. 484
PeryJIHU,ll.JI JJHIIleBape1mH ....•.•......•....•.•......••.•.....•...•.• 486
Bhl.n;eJieHIIe .....•..•.•....•.•........•......•.....••..••...•..•.• 48 7
Pa3MHOJKeHll.e .•........•.••...•......•.•......•.•.•.....•..•.•.•. 492
MymcKa.H no,1osaH Cll.CTeMa •.•...••.•.•.•.•...••....•.•.....••••...•. 492
JKeHCKaH IlQjIOBaH Cll.CTeMa ....•.••••.••..••.....•.••.•..•.••.•.•.•.• 493
3M6pll.OHaJibHhlll nep110,ll, pa3Bl1Tll.H '!eJIOBeKa •...•.....•..•.••.•••..•..•• 496

TECTbI
06Ill,aH 6ttOJIOl'll.JI ...•.•••.....•.•.•.....•.....•..... , •.••.....•.•.• 497
il;HTOJIOI'IIH ......•.•......••....•....•.••.•.•••..•••.•••....... . • 497
feHeTHI{a ..•.......•...••.•••...•......••.•.••..•.••.......•.••.. 498
3BOJIIOU:HOHHOe y'!eHll.e .......•.••...••...•....•...•........•.•...•. 499
AHTponoreHe3 ..•..•.......•..•..•..•...•.•.....••...•...•.•.•.•.• 500
3KOJ!OI'HH .•...•..•.•..•.•..••.•••.•........••......•...•...•.•.• 501
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ycoBepweHCTBOBa1-rne B CBJI-

3H c 3THM M€TO,ll;OB ncc.1e.n;oBaHHH JI03BOJIH:IH HaMHOrO r.1y6me ll3YlfHTb
cTpyKTYPY n cpyHKIJ;HOHnpoBaHne pa3JIHlfHhIX KOMIIOHeHTOB KJieTJHL Ha ocHOBaHHH IIOJiylfeHHhlX pe3yJibTaTOB 6hIJIH ccpopMyJIHpoBaHhl OCH06Hb/e noJlOJICeHUJl co6pe.MeHHOU KJLemo•mou meopuu.
Bee JICU6bte opia1tU3Mbt cocmo.Hm u3 KJLemoK.
KJtemKa - 3Jte.MeHmapHa.H J1Cu6a.H cucme.Ma, erJuuu4a cmpoe1tu.H, .HCU.311eoe.HmeJtbHOcmu, pa.J.Mlto.HCeHUJl U UllrJU6UOya.'lbHOi?O pa;wumUJl JICU6blX opiaHU.J.M06. Bne KJtemKu 1tem JICU.JHU.
KJtemKu 6Cex opianu.JM06 cxodttbt no cmpoeHu10 u xu.MU'lCCICOMy cocmaay,
u 6 nux cxodllbt.M o6pa3o.M npome1Ca10m npo4eccbt 06.Mena Bew,ecm6, xpaHenue u peaJtu3a4uJl nacJteocm6enHou un<jJop.Ma4uu.
Ho6bte 1CJtem1Cu 60.Jttu1w10m moJtb/CO nymeM deJte1tu.H panee cyw,ecmBy10w,ux 1CJtemo1C.
KJtemo'lnoe cmpoeHue Bcex llblHe J/CuByw,ux opiaHU3.M06 - c6uoemeJtbcmBo eduncm6a ux npoucxoJ1CdeHu.H.
TaKHM o6pa3oM, B HaqaJie

XX B.,

6Jiaro.n;apJI co3.n;aHHIO KJieTOlfHOH Teopnn,

ccpopMHpOBaJIOCh npe.n;cTaBJI€HHe 0 e.n;HHCTBe opraHnqeCKOH rrpHpO,ll;bl.

I. CneTonan MHKpocKODHH:
1. <Pa3060-ICOHmpacmtta.R .MU1Cpoc1conu.H -

3TOT MCTO,ll; II03BOJIJieT pe3KO

IIOBhICHTh KOHTpacTHOCTh H3o6pameHHJI o6'beKTa.

2.

JlOJt.Hpu3a4uonHaJl .MUICpOCICOllUJl -

MeTo.n;, rrpu KOTOpOM B rro.1€ 3pe-

HH.H IIOJI.HpH3aIJ;HOHHOro MHKpOCKOIIa B pe3yJihTaTe ,ll;BOHHOro JiyT-IerrpeJIOMJI€HH.H ( «aHH30TPOIIHH») o6'h€KThl OKa3hIBaIOTCJI RPKO CB€TJIIIJ;HMHC.H Ha TeMHOM cpoHe.
3. HHmep<jJepen4uOHHaJl .MUICpOCICOllUJl -

MeTO,ll;, npH KOTOpOM CBeT

pa3.n;eJIJI€TCJI Ha .n;Ba rryqKa. 0.n;HH H3 HHX rrpoxo.n;HT qepe3 o6'heKT,
.n;pyro:H -

MHMO Hero. BcJie.n;CTBHe aToro cBeT, rrporne.n;nrnii qepe3 o6'h-

eKT, HCIIhIThIBaeT cpa30BhIM C,ll;BHr, H H306pameHH€ CTpOHTCJI TaKHM
o6pa30M, qTo yqacTKH KJieTKH, 06Jia.n;a10ru;He pa3HOH IIJIOTHOCTbIO, 6y.n;yT OTJIHlfaThC.H .n;pyr OT .n;pyra no CTerreHH KOHTpaCTHOCTH.

4.

Mu1Cpoc1Conu.H B me.MHO.M noJte -

rrptteM, rrpn KOTopoM n3o6pametttte

nona.n;aeT B o6'beKTHB B pe3yJihTaTe OTpameHHJI CBeTa OT o6'beKTa, 6Jia-

15

MemoiJbi u3y'lenun 1aiemKu

ro,napJI lJeMy MeJih'Ia:HUIHC 1.J:UCTHI~bl Bbir JIJ!,[IJ!T CBCTJ!lIJ;HMHCJ! TO'-IKRMH
Ha TeMHOM cpoHe.

5. YJ1bmpa<fJuoJ1emoea.H MUICpOCKOllUJl -

crroco6, rrpn KOTOpOM KOMITO-

HeHTbl ofrheKTa 06J1a,na10T cneu;ttcpHllecKHM rrorJio~eHHeM Y<P-J1yqei1:
c orrpe,neJieHIIOH ,ll;JIHHOM BOJ!Hbl, '-ITO II03BOJIJieT BbIJ!BHTb HX 6e3 OKpallIHBaHIIJI.

6. <PJtyopecu,eHml-taJl MUKpOCKOllUJl -

MeTO,ll; IICCJie,noBaHHJ! )KlfBbIX K.!e-

TOK B YJ1bTpacpHOJ1€TOBOM CBeTe. Il03BOJ1JleT H3YlJIITb co6CTBeHny10

cpJIIOOpecu;eHI~HIO (cBelJeHHe) Be~eCTB, a TaK)Ke HX CTpoeHHe IIOCJIC .ll;06aBJieHIIJ! B HHX crreu;HaJihHbIX KpaCHTeJiett (cpnyopoxpOMOB) • .IJ:aHHhIII
MeTO,ZJ; II03BOJIJieT orrpe,ri;eJIHTb MeCTa paCIIOJIO)KeHHJ! HYKJieHHOBbIX
KHCJIOT, BHTaMHHOB, mttpOB H .ri;p.

7. JI,ucprpepettu,uaJtb1toe yJ1bmpau,ettmpurpyzupoeattue -

MeTo.n; II3ylJe-

HH.H COCTaBa KJieTOK, OCHOBaHHbIM Ha pa3,neJieHHll KJ'IeTOlJHbIX opraHeJ'IJI H BKJIIOlJeHHH no llX IIJIOTHOCTH. IlpH 6hICTPOM Bpa~eHHH pacTBopa B crreu;ttaJihHOM rrpII6ope -

yJihTpau;eHTpttcpyre -

opraHeJIJ!hI

KJleTOK Bhma.n;aIOT B oca.n;oK, pacrroJiara.HCb CJIO.HMH B COOTBeTCTBHll
co csoeii IIJIOTHOCTbIO. 8TH CJIOH pa3.ll;eJI.HIOT H H3ylJaIOT OT,ZJ;€JlbHO.

II. 8JieKTpOHHaH MHKpocKonm1:
1. Tpanc.MuccuOHHa.H :JJleKmpoHHa.H MUKpocKonu.H -

11306pa)KeHHe Ha

cpJiyopecu;Hpy10~eM 3KpaHe 3JieKTPOHHOro MHKpOCKOIIa rrpH pacceHBaHHH rryqKa 3JleKTPOHOB o6'beKTOM. qeM 60Jihllle paccellBaIO~a.H
crroco6HOCTb Toro HJIH HHOro yqacTKa, TeM 6oJiee TeMHbIM BbirJI.H,ll;HT
OH Ha 3KpaHe.

2.

CKattupy10w,a.H (pacmpoea.H) 3JteKmpo1t1ta.H MUKpocKonu.H -

MeTo.n;

H3ylJeHH.H TpexMepHOH IIOBepxHOCTH KJieTKH .ri;ettCTBHeM rryl.J:Ka 3JleKTpOHOB no IIOBepXHOCTH 06'heKTa.

III. PettTrettocTpyRTypHLiii attaJIH3 -

H3ylJeHHe KOHcpurypau;HH MoJieKyJ1

6eJIKa, HYKJieHHOBhIX KHCJIOT ITO pacrrpe.n;eJieHllIO Ha 06'beKTe peHTreHOBCKOro
H3JiylJeHIIJ!.

IV. BHTaJILHoe (npmKH3tteunoe) H3yqettHe KJieToK -

H3yqentte cBo6o.n;Ho-

:anrny~Hx IlpOCTCMllIHX opraHII3MOB, KJieTOK KYJibTYPhl TKaHH, a TaK)Ke KJleTOK KpOBH.

lfayIJenue r/JUKCUpo6aHHblX KJlemoK ( c HCIIOJib30BaHHeM crreu;ucpHl!eCKOM
o6pa60TKH H rrpuMeHeHHeM pa3JIHlJHbIX KpaCHTeJie:H) rrpOBO,ZJ;HTC.H CJ1e,ny10~H
MH MeTO,ZJ;aMH:

1. F(umOXU.MU'ieCKUe .MemoiJbl

HCCJie,ZJ;OBaHH.H OCHOBaHbl Ha npHHIJ;IIIIe

crreu;HcpHlleCKOrO CBJ!3bIBaHHJ! KpaCHTeJie:H c orrpe.n;eJieHHbIMH KOMIIOHeHTaMH HJIH

Be~eCTBaMH

2. AemopaiJuozparjJu.H

KJieTKH.

ocHoBaHa Ha perucTpa1~HH se~eCTB, MelJeHHhIX

pa,ZJ;HOHYKJIH,ZI;aMH.

3. HMMyHozucmoxu.Mu'iecKue .MemoiJbt

ocHoBaHhI Ha orrpe.n;eJ1eHHH aH-

TureHOB B KJleTKax H Ha o6Hapy)KeHHH TKaHeBhIX aHTHTCJI.

4. [_{ umocneKmporjJOmOMempU.H

6a3HpyeTCJ! Ha onpe.n;eJieHHH KOJIIIlJeCT-

06~an

16

6uo1wiun. Oc1to6bl 1.fumo1wzuu

Ba sern;ecTB l1 l1X COCTaBHblX ::weMeHTOB B KJleTKe B 3aBIICl1MOCTl1 OT
nor Jiorn;eHJUl CBeTOBblX Jlyl.JeH onpe,neJieHHOH ,iJ;JIMHbl BOJIHbl.

5.

ll,umo<jJU3UOJl02U'leclWU .lft,emoi)

OIIHpaeTCH Ha JI3yl.JeHIIe q:nI3IIOJIOrl1·

l.JeCKHX npou,eccoB Ha OT,neJihHbIX II30Jil1pOBaHHblX KJieTKax ll,bl, HepBHbie CTBOJ1bl, raHrJIMU HT •

6. A1emoo ICYJlbmypbt m/CaHeu

'.Vlhllll-

.n;.

npe.n;noJraraeT HCC.'Ie,ll;OBaHIW JKIIBbIX

KJieTOK, IIOMern;eHHblX B COOTBCTCTBYIOID;YIO cpe.n;y, B KOTOpoii OHH
cnoco6Hbl K aBTOHOMHOMY pocTy.
IlpIIMeHHIOTCH TaIOKe

MUICpoxupypzua -

BblIIOJIHeHIIe c IIOMOID;bIO cneu,H-

aJihHblX MaHIIIlYJIHTOpOB pa3JIHl.JHblX orrepau,nii Ha KJieTKe Hee opraHOH,n;ax, l1

npUJICU3HeHIWe OICpawuBanue

JKIIBbIX KJieTOK KpaCIITeJiflMII B .n;1rnrra30He

KOHU,eHTpau;nH:, He Bbl3hIBaIOrn;nx TOKCII'IeCKOro 3cpcpeKTa.

06m;ee cTpoeuue 1meT1m
KJtem/Ca - amo 3JleMenmapnaR cmpyKmypnaa, <jJyn1Cu,uonaJ1blta.R u zeuemu'leCKa.R eiJunuu,a JICUBbtX opzanu3M06, 02panu'leH1ta.R no.llynp<muu,ae.wou
MeM6panou, cocmoaw,aa u3 aopa u u,umon.naaMbt, cnoco6naa IC caMopezyJLRu,uu, CUM006H06JleHUIO u CUM060CnpOU36eiJeHUtO.
<l>opMa KJieTKH (pnc. 1). Pa3Jin'-laIOT KJieTKn c n3MeH'IHBOtt cpopMott (aMe6hI,
JiettKOll,HTbl H .n;p.) H
Ha n

KJieTKII, cpopMa KOTOpbIX 6oJiee HJIH Mettee IIOCTOHH-

cneu,IIcpII'IHa ,ll;JIH Ka:m.n;oro nx TIIrra (ttttcpy3opmf,

crrepMaTo30II,ll;hI,

3pIITp01I,IITbl, 3IlIITeJIIIfu'IbHbie H uepBHbIC KJieTKII,
a TaKme 6oJibIIIHHCTBO pacTnTeJibHhIX KJieToK).
ct>opMa KJieTOK 3aBIICl1.T OT HX cpyHK1I,IIOHaJibHOrO rrpIIcrroco6JieHIIH, IIOBepxHOCTHoro HaTHJKeHMH M BH3KOCTM rrpOTOIIJia3MbI, MeXaHM'IeCKOI'O
B03,n:eifcTBHH rrpHJierruorn;nx KJieTOK, pnrH,n:HOCTII
(:meCTKOCTH) KJieTO'IHOH MeM6paHbI.

Pa3Mephl KJ1eToK. Pa3MephI KJieToK KOJie6JIIOTCH B rrmpOKMX rrpe.n;eJiax. Lt:ttaMeTp caMbIX
MeJIKHX JKl1BOTHbIX KJieTOK paBeH

4

MKM. 0.n;tta-

KO HeKOTOphie pacTnTeJihHhie M JKHBOTHhie KJieTKH MOJKHO Bn,n;eTh HeBoopymeHHhIM rJia30M, TaK
KaK OHH ,ll;OCTMraIOT HeCKOJihKHX caHTHMeTpOB
B ,n:waMeTpe (ttarrpnMep, Hifu;eKJieTKM).

B
a

TKattax JKMBOTHhIX o6'beM KJieTOK orrpe.n;e-

JieHHoro THila ,ll;OBOJihHO IlOCTOHHeH H He 3aBMCHT
OT o6rn;ttx paaMepos TeJla. HarrpnMep, KJieTKM

PMC. 1.
a -

C!>opMa KileTOK:

HepsHa11 l<JleTKa;

6- r-

IIO'IeK II rre'IeHM y 6h1Ka, JIOIIIa,n;M II MblillH IIMean11-

IOT rrpMMepHO 0,ll;IIH H TOT

me

pa3Mep; pa3JIH'!He

Ten11anbHb1e KlleTKL1;,l1-COeAL1HL1-

B BeJIH'IMHe opratta o6ycJIOBJIHBaeTCH 'IHCJIOM,

TenbHOTKaHHafl KneTKa; e -

a He o6'beMOM KJieTOK. 8To npaBMJIO HHOr;:i;a Ha-

KlleTKa; JK -

11iiiu,e-

Mb1we•iHa11 KneTKa

3hIBaIOT

3a/COHOM nocmORHHOZO o6oeMa.

XuMu'teCKUii cocmaB KJlemKu

17

XHMH'lleCKHii cocTan KJieTKH

B COCTUB )KllBblX opraHI:I3MOB BXO).(.HT Te me XHMHqecKne ;)JJC:\1CHThl, KOTOpbie COCTaB."IfffOT H 06'heKTbl HeiKHBOll npHpO.D;bl.
Co11epmautte n 1rn:enw xHMH'lec1rnx coe;i:uue1rnii: (6 % 1w Cblpy10 Macey)
~--

L

-----·

~~ -B~~e(-;,~a=---

--

--~--1

____He_opram1_q_ecK11e. _____ _j______ __Opra1_1w1ec:KJ:!~--- __

. Bo.n;a
I M11HepaJihHhle coJIH

70--80
1-1,5

L_ __________________

· Bemrn
JK11pb!
YrJieBOAhI

10--20
1--5
0,2---2

---~·_HYl{:l!C11H<~!3h~li!1_CJIOThl ____!_-:-_2__ ;

B 1rneTKe o6ttapymetto 6oJiee 80 x1:1M1:1qecKnx sJieMeHTOB Ilep1:10.1J;11qecKorr c11cTeMhI
IL Mett,n;eJieesa. Ilp1:16JIH3HTeJihHO 40 3J"IeMeHTOB np1:1H1:1MaIOT yqacTHe
B npou.eccax o6MeHa semecTB 1:1 o6Jia,n;aroT Bb1pameHttorr 61:10.1or1:1qecKorr aKTHBHOCThIO. 8TH 3JieMeHTbl Ha3bIBaIOTC.H 61:1oreHHbIM1l. Ilo co.n;epmaHHIO HX ,n;eJI.HT Ha
TPH rpynnh1:
1. M aKpo;ue.11-ie1-tmbt (98 '%) - 0, C, H, N.
2. Mu1Cp03JteMe1-tmbt (1,9 °/.i) - K, P, S, Mg, Cl, Ca, Na, Fe.
3. Y.llbmpaMuKpo:uteMeHmbt (0,01 'Xi) - I, Cu, Co, Zn, Mo, Br, Mn, B 1:1 .n;p.
Boi}a B cpe,n;HeM COCTaBJI.HeT 80 % MaCChl KJieTKH (.n;o 95 % - B KJieTKax MeAY3hl, 90 % - B KJieTKax 3apo.n;hrllla qeJioBeKa, 79 '% - B MhIIIIu,ax cep.n;u,a,
60 %1 - B CTapbIX KJieTKax H 10 % - B KJieTKax 3MaJIH 3y60B). Bo.n;a Haxo,n;HTC.H B KJieTKax B .n;syx cpopMax: CB06o,n;Horr - 95 'Xi H CB.H3aHHOll - 5 %i. IloTep.H 20 % BO,ll;hI cMepTeJihHa ,n;JI.H opraHH3Ma. Taxoe BhICOKoe co.n;epmaHHe BO,n;hr
B KJieTKe onpe.n;emieTCH MHoroo6paa1:1eM ee cpyHKIJ,Htt:
- cpe.n;a ,ll;Jl.H npoTeKaHHH 6HOXHMHqecKHX peaxu,1:1i1;
- YHHBepcaJihHbIH paCTBOpHTeJih;
- cpe,n;CTBO TpaHCIIOpTHPOBKH BemecTB B KJieTKax (.n;mpcpy3H.H) H B opraHH3Me
(Kposoo6pamettlie );
- o6paaosattMe KJieToqHhlX CTPYKTYP (60Jibllloe co.n;epJI{aHHe B U.HTon.1a3Me);
- onpe,n;eJI.HeT o6'beM 1:1 ynpyroCTh KJieTKI:I ( 06ecneq1rnaeT ocMOTHqecKoe
H Typropttoe ,n;aBJieHHe);
- yqaCTByeT B TepMoperyJI.HIJ,HH (6Jiaro.n;ap.H BblCOKOH TeIIJIOeMKOCTH H TellJIOIIpOBO,/l;HOCTlI);
- yqacTsyeT B peaKIJ,H.HX rM,n;poJIH3a lI cpoToCHHTe3a (HcToqHHK HOHOB H-t );
- cpe.n;a ,ll;Jl.H OIIJIO,ll;OTBOpettm1;
- y pacTeHHH o6ecrreqHsaeT TpattcnHpa11;tt10 H rrpopacTaHHe ceM.HH.
BMoJiorMqecKa.H poJih BO,n;hr orrpe,n;eJI.HeTc.H MaJiblMH paaMepaMH MOJieKyJI,
crre11;mpHKOH HX CTpOeHHH (IIOJIHpHOCTb, crroco6HOCTb o6pa30BbIBaTh B0).(0PO.n;Hhie CB.fl3H). Ilo OTHOIIIeHlIIO K BO,n;e Bee semecTBa ,n;eJIHTC.H Ha 2uapo<PU.llb1-tbte (paCTBOplIMhle B Bo,n;e) - MHHepaJihHhle COJIH, KHCJIOThl, meJioqH,

.n;.

06utaJI 6uo1wzuJI. OcnoBbt 1fUmo1wzuu

18

Buo.'loruqec1rnn po:1h HeK0Topb1x xnMH'lec11:ux JJieMeHTOB
cf>Il3IlOJIOfl1'!eCKaH pOJlb

8;ieMeHThl

Yr.wpo.n (C)

Bxo;vrT B COCTaB BO/~bl, opral!ll<ICCKilX BernecTB (6eJ1Kll, 11yK-

Bo11opo,1J; (H)

;1e11HOBbJe KllCJIOThr, yn1cno,1J;b1, JI111IIIL\hI).

R'.IlCJIOpOl\ (0)

Y<IacTsymT B c1rnTe3e opraHH'Iec1<11x uernccTB Il q1yHKl\IIHX,

AaoT (N)

ocyn1ecTB.1HCMbIX oTlfM11 opraHll'ICCKllMll BeU(eCTBaMII.
Y<IacTsyeT B npou:cccax so36yn<J(eH1u1 K;1cT101, B 11011,1J;cpma-

HaTpllll (Na)

Hllll OCMOTil'ICCKOI'O ,ll.aBJICHilH Il

pH

cpe,ll.hl, B,1IlHCT na pa6oTy

noqeK
BXO!lllT B COCTaB KOCTHOII TKaHll, HC06XO,ll;llM npn csepTbIBa-

Ra;1bll,Ilii (Ca)

Hllll KpOBll, MhIIIIe'IHOM COKpan(eHlll1
Heo6xo11HM 11.1n Bo36ym.neHilH nepBHhJX KJICTOK, rrp0Be11e1111a

RaJIIltt (R)

llMIIY-1bCOB, COKparneHllH Mbllll1I
YqacTnyeT B 11011AepJKaHilll pH JKe,1y11o'IHOI'O COKa, OCMOTil<IC-

XJiop (Cl)

CKOI'O l(aBJICHl1H IIJia3Mbl KpOBH
ct>occiJop (P)

CTpyKTYPHblll IWMITOHCHT IWCTeii 11 3y60B, BXOJ(HT B COCTaB

JI~e.'IC30 (Fe)

CTpyKTYPHh!H KOMIIOHeHT !'CMOr,10611:Ha KPOBH, MHOI'JI06HHa

ATct>, HA)l;ct>, cpoccpoJ1mrn11on

Mbllll1I, cpepMCIITOB I(errn nepenoca 3,1eKTpOHOB

Mo11 (I)
Mell.h (Cu)

BXOl\HT B COCTaB I'OpMOHOB 11IllTOBil!llIOii JKeJie3hI
-

-

--

-

-

-

--

-·

YqacTByeT B npou:eccax KpoBCTBopeHMH 11 c:11nTe3e reMOI'JI06MHa

cl>TOp (F)
Marm1if (Mg)

C1·pyKTypHblll KOMIIOHCIIT 3y6uoii TKaHll
BXOJ(HT B COCTaB XJIOpOcp11JIJia, KOcpepMCHTOB, aKTHB11pyeT
0HepreTH'IecK:11iJ: o6MCH :11 c11nTe3 AHR

Cepa

(S)

BXOJ(IlT B COCTaB aMill!OKl'!CJIOT, 6eJIKOB (FrnCyJimI) H BllTaM:rrHOB (B )
1

1l;IlHK {Zn)

K0Mnoue11T cpepMeHTOB, 11eo6xo11:rrMbIX .n;m1 uopM3.1hHOro pocTa

Ko6aJibT (Co)

Bxo.n;MT B cocTaB BttTaMMHa B 12

MapraHeu (Mn)

Heo6xo11ttM .n;Jia oKMCJICHllH m:rrpHbIX KMCJIOT, yqacTByeT
B rrpou:eccax l\hixamrn Il <PoTOCilHTc3a

MOHOcaxapII,D;hI, 6eJIKll ll .n;p. ll

zuiJpo<jJo6ttblC

(BO,[(OHepaCTBOpIIMbie) -

jKif-

pbI, IIOJIIIcaxapII,D;hl ll .n;p.

MunepaJlbflbte coJlu

cocTaBJIRIOT

1-1,5 'Yo

MaCChr KJieTKH. BoJihIIIa.H

'laCTb HeopraHII'leCKIIX Bell(eCTB HaXO,[(.HTCR B KJieTKe B BII.n;e COJieH HOH,

COJI.HHOH,

cpoccpopHoii:.

cep-

MttHepaJihHhre COJIII IIrparoT BamHyIO pOJib

B pa3BIITMII mIIBhIX opraHII3MOB. IIx He.n;ocTaToK MJIH II36hlTOK MOJKeT rrptt-

Opzanu'!ecKue 6eUfecnrna K.riemKu. £e.ri1<u

19

B€CTH K r116eJIH opraHH3Ma. COJIH :v10ryT Haxo,n:HTbCH B K.1CTKe Jll16o B BHµ:e
HOHOB, JI!160 B TBep,n:oM COCTOJIHHH. Munepa.lhHbie COJII1 B BOlWblX paCTBOpax ;:i;11cco1(1111pyKiT ua aHHOHbI 11 KaTHOHhI, y 1rncTBYIOT B no,n:.n:ep:maHnn ocMOTIIqecKoro JJ,aBJTeHHH (3a cqeT pa3HOCTH KOHI(eHTpau,Hii no o6e CTOPOHbl
MeM6paHhI KJieTKH), TIO)J,)(epJKHBaIOT c;ia6orue.10lfHYIO pH

(7,2-7,4)

B TKa-

HeBOM Jl\I:!,llKOCTH.

Oprattnqec1me Bem.ecTBa KJJeTKH
BamueiIIIIIIMII opram1qecKHMn Beru;ecTBaMn KJieTKH RBJIHIOTCR 6e"1KH,

mtt-

pbr, yrJieBO!l:bl n HYKJieHHOBb!C KHCJIOThI.
CTpyK'l'ypubre n ;::i;pyrne oco6euuocTH KJieTKH Tecuo CBJI3aHhI c ,n:JIHHHhIMH
MOJieKyJiaMH, COC'l'OJIIIJ;I:!Mll H3 TIOB'l'Opmoru;nxcJI ;)JieMeHTapHbIX C,11,HHHU., coep;HHeHHbIX KOBaJieHTHbIMH CBJl3JIMH. 3TH e,n:HHHUbl Ha3hIBaIOTCJI MOHOMPpa-

MU, a o6pa3yIOrn;uec.H MaKpOMoJieKyJihI -

no.rtuMepaMu.

OprauntJecK11e coe;::i;I:Iueu11a cocTaBJIJIIOT B cpe,n:neM
mirnoro opraHH3Ma.

B pa3Jrn:qnbre

20-30 '%

:vraccbr KJie'l'Kn

TI:!IIbI KJI€TOK BXO,ll;HT Heoµ;IIHaKOBOe KOJIHqe-

CTBO opraHuqecKnx coe;::i;nHennii. TaK, B pacTHTeJibHhIX KJieTKax rrpeo6Jla)(aJOT
CJIOiKHLie yrJieBO)J;hI -

noJincaxap11)(hI, B aornoTHhIX -

6eJIKH 11 a<Hphr. Op;HaKo

KaJK,l(aJI rpynna opraHI1'IeCKI1X Beru;eCTB B JII060M THrre KJI€TOK BhIITOJIHJieT CXO,l(Hbie QJYH1\IXl1H.

Bemrn
y pacTeHHM

6eJIKH CHHT€311PYIOTCJI B pu6ocoMaX l13 aMHHOKHCJlOT.

y

)KH-

BOTHbIX 6eJIKH IIOCTyrraJOT B opraHH3M c rrnru;eii, pacru;enJIHIOTCJI AO aMHHOKHCJIOT, KOTOphre HJI;YT na CHHT€3 co6cTB€HHbIX 6eJIKOB B COOTBeTCTBUH c retteTHqecKHM KOJ(OM.

Cocma6 6e.ri1Co6
BuonoJlUMepbl. MonoMepaMn JIBJlRJOTCH aMMHOKHCJIOThI - HH3KOMOJ1eKyJIRpH&re coe)(HHenuJI. Pa3JIHqaroT 20 aMn:HOKHCJIOT. 0614aJL <jJop.wy.ria:

R

I
NH 2 -

C -

COOR,

I

R

r.n;e NR 2 aMunorpynna ( ocHOBHhie cBonc'l'Ba), COOR Rap6oKCHJibHaH
rpynna (KHCJIOTHhre cBoncTna), R - pa,n;n:KaJI (20 BapuaHTOB). 8To aMcpoTepHhie coe,n;HHeHHR.
<l>H3HqecKue cnoi1:cTBa:

6ecl(neTnh1e,

xpucTannnqecKMe,

pacTnopnMhI

B Bop;e, HepaCTBOpHMbl B opraHH'I€CKHX paCTBOpHTeJIJIX.
AMHHOKHCJIO'l'hI, coe;::i;nH.HJICh, 06pa3y10T rrerrTH)(HYIO CBJI3h (xonaJieHTHyIO,
npoqHyIO).
-C-N-

11

I

0

R

06UfUJl 6uoJtoZuJl. Ocno6bt 1.J,Umo;wzuu

20
KJWCCU<jJUKQL{UJl 6PJlKOB."

I.

Ilo KOJIII'IeCTBY aMHHOKI1C~IOT:

1.
2.

II.

2 ,!l,O 10 a:vi11HoKucJioT);
10 .no ••. ):
co6cTBeHHO IIOJJI1IJellTH,ll,bl (OT 10 .n;o 100 aMl1HOKHC.10T) ropMOHbI;
6eJIKl1 (npoTel1Hbl)- OT 100 aMI!HOKHC.10T ,ll,O HeCKOJibKI1X Ml1.1.Jll10HOB.
O.:rnronenn1,11,b1 (OT

IloJittnenTI1,ll,bI (OT

IIo cocTaBy:

Ilpocmble 6e.il!CU ( npomPU/lbl) COCTOJIT T0.1bKO H3 OCTaTIWB aMIIHOKUCJIOT:
npoTaMHHbl H rHCTOHbl -

BXO,ll,JIT B COCTaB HyK.1enpoTeH,ll,OB. MrpaIOT

Ba:IKHYIO pOJib B pery.1Hl(HH MeTa6o.1H'feCKOM aKTHBHOCTH reHOMa;
npoJiaMHHhI H rmoTe.1n:HhI -

6e.i1KH pacTn:TeJibHoro npo11cxom,n;enn:JI, co-

cTaBJIHIOT OCHOBHYIO Macey KJie:UKOBHHhI;
aJih6YMI1Hbl n: rJio6yJIUHbl -- 6eJIKH :IKHBOTHOro npmICXQ)l{,ll,eHirn (5111'1HbIM 6eJlOK, Mbllllll;bI, MOJIOKO).

CJWJICHbte 6eJtKU ( npomPudbt) pa3.1wrn10T n: na3hIBaIOT no npocTeTn:<IeCKOM
(He 6eJIKOBoi1) rpynne:
xpoMonpoTen:,11,hI -

6eJIKH, B KOTOpb!X npocTeTH'feCKOM rpynrrofi: CJIY:IKHT

ITHrMeHT (reMor JI06HH, Il;HTOXpOMhI, ,ll,bIXaTeJihHhie cpepMeHTbI);
HYKJieorrpoTeH,ll,hI -

6eJIKH, CB513aHHhie c HyKJiel1HOBbIMI1 KHCJIOTaMn: (oc-

HOBa 51,11,epnoro Bem;ecTBa JIHITOilpOTe:H,ll,bI -

xpoMaTHHa);

coe,n;r-rneHH51 6eJIKOB n: JIHilll,ll,OB (c]JepMeHTbl ITJia3MaTI1-

qecKHX MeM6pa11);
cpoccpoupoTen:,n;hI -

coe,n;11nemrn 6enKoB 11 cpoccpaToB (B MonoKe, meJITI{e

I<ypn:noro RHu;a, ttKpe pbr6, II,HC);
rmmorrp0Te11,n;b1 -

coe,11,n:nemrn 6eJIKOB H yrJ1eso,11,oB (KoMnoneHT KJieTo<I-

noii MeM6panLr);
MeTaJIJIOIIpOTeH,ll,hI -

III.

coe.n;nnenmI 6eJIKOB II MeTam10B (cpepMeHTbI).

Ilo CTPYKType:

<Pu6pUJlJl.HpHbte 6eJtKU -

IIOJIHITeITTI1,ll,Hbie Hll'reBH,ll,Hbie u;errn, ITJIOXO pac-

TBOpllMbl B 130,ll,e (KepaTn:H BOJIOC ll poroB, MI103HH MhIITII.J;, KOJIJiareH KOCTe:H,
cpn6pn:noren KpoBn:).

I'Jl06yJtRpllbte 6eJtKU -

TI0.7II1ITeITTH,ll,Hble u;entt, CJIO:IKeHHbie B cpopMe ITiapa,

BO,n;opaCTBOpllMbie (npoTeHHhI TIJia3MbI KpOBH, cpepMeHTbI).

CTpyKTYPhI 6eJrna (ypoBHH opraHHaau;HH)

II epBu'lnaR -

(pnc.

2)

JIHHeii:na51 nocJie,n;oBaTeJihHOCTh aMHHOKHCJIOT, coe,n;uneHHhIX

,n;pyr c ,n;pyroM nerrTH,ll,HbIMll CB51351MH B ITOJIHI1eIITH,ll,HYIO u.errh (crreu;HcpH'!Ha,
3aKO,ll,HpoBaHa B .IJ;HK, OT Hee aaBMCRT CBOHCTBa ll cpyHKD;Hll 6eJIKOB).

BmoputtnaR: IIOJIHrrerrTH,ll,Ha51 u.errh, 3aKpyqeuHa51 B crrHpaJih. a-cnupaJ/,b rrpaB03aKpy<IeHHa51, B03HMKa10m;a51 B pe3yJihTaTe o6pa30BaHH51 BO,ll,OPO,ll,HhIX
CB513eii Mem,n;y C=O II NH, Mem,n;y nepBhIM u TpeThHM aMMHOKHCJIOTHhIMH 3BeHh51MH (rJI06HH);
-

CKJia,n;<IaTa51, {3-cnupaJtb, B03HHKaIOm;aa B peayJihTaTe o6pa3oBaHH51 Bo,n;opo,n;HhIX CB513ell Mem.n;y napaJIJieJihHO opn:eHTHpOBaHHhIMll yqaCTKaMH no-

21

CmpyKmypa 6enKa

JI11nenn1;i:no:H Mo.1eKyJrh1 (KepaTHtt BOJIOC);
-- Tpexq,1ettttaa cn11paJih -

crr11pa.1e-

B11.n;tta.H CTPYKTypa, B03HHKaIOIIJa.H
B pe3yJihTaTe rreperrJieTett11.H Tpex
rro.111rrerrTH.n;tthrx ri:erreii (KOJIJiareH).

Tpemu'l1taJt -

rroJI11rrerrTH.n;Hhre ri:err11

B pe3yJihTaTe rrpocTpattCTBettttOH yKJia,ll;KH o6pa3yIOT
paKTeptta

i1to6yJty (KJiy6oK). Otta xa-

,ll;Jl.H

6oJibllIHttCTBa

6eJIKOB

H rro.n;.n;epmHBaeTC.H HOHHbIMH, BO,ll;Opo.n;HbIMH CB.ff3.HMH, ,ll;HCYJih<pH,ll;HbIMll MOCTI1KaMH

(-S-S- ),

r11.n;pocpo6HhIMH B3aHMO-

6enKa:

3 -

.n;eii:CTBH.HMH. Bee rJio6yJI.HpHhre 6eJIKH -

4

2

P111c. 2.

Pa3nV14HbJe crpyKrypb1 MoneKyn
I -

nepBV14Hasi;

TpernYHasi; 4 -

2 -

BTOPV14Hasi:

YernepTVIYHasi (Ha

cpepMettTbI, attTIITeJia, ropMOttbI.

q emeepmU'llWJl -

xapaKTeptta ,ll;Jl.H 6eJIKOB co CJIO:?KttbIM CTpoettIIeM, co-

CTOHT H3 tteCKOJibKHX IlOJIHileITTH,ll;ttb!X ri:eneii:, y.n;epmHBaeMbIX B MOJieKyJie
,ll;HCYJih<pH,ll;ttbIMH MOCTHKaMH, Bo.n;opo.n;tthIMH CB.ff3HMH H r11.n;pocpo6ttbIMH B3aHMo.n;eHCTBH.HMH. HarrpHMep, MOJ1CKyJia reMOrJI06Htta COCTOHT H3 'ICTbipex rroJIHITerrTH,ll;ttblX l~erreii, Kam.n;a.H 113 KOTOpbIX CB.H3atta c o.n;tto:i1: rpynrroiI reMa,
y.n;epm11BaIOIIJero KHCJiopo.n;.
HapyllleHHe CTPYKTYPhI 6eJIKa tta3brnaeTc.H

iJe1wmypau,ueu. Otta ttacTynaeT

rro.n; .n;eHCTBHeM cpH3H'IeCKHX H XHMM'IeCKHX cpaKTOpOB (TeMrrepaTypa, B03.n;eiICTBHe KHCJIOT H .n;p.). BbrnaeT o6paTHMOH (pettaTypari:H.H) H tteo6paTHMOH (eCJIH
ttapylllaeTc.H rrepBH'Itta.H cTpyKTypa MOJieKyJihI).

<Py1ucu,uu 6eJt1Coe:
1. Cmpy1Cmyp1taJl (cTpOHTeJihtta.H) -

BXO,D;.ffT B COCTaB Bcex MeM6pattttbIX

CTPYKTyp.

2. <PepMe1tmamU6/taJl (KaTaJIHTH'IeCKa.H)
3. 3aw,um1taJl (HMMYttOJIOrH'IeCKa.H) -

-

cpepMettTbI opraHH3Ma.

aHTIITeJia,

HMMyHorJio6yJIHtt,

HttTepcpepott.

4. Co1CpamumeJtbltaJl

(.n;BHraTeJihtta.H) -

.n;BHmettHe pecttM'IeK,

mryn1-

KOB, COKpaIIJettHe MblllIIJ:.

5. I'opMOHllJlbHllJl (peryJI.HTOptta.H) - ropMOHbI (HttCYJIHH).
6. 3nepiemu'leC/CaJl - ITPH CJKHrattHH 1 r 6eJIKa BbI,D;eJI.HeTC.ff 1 7 ,6

K,Il,JK

3HeprHH.

7. TpancnopmnaJl - reMorJI061:1H.
8. Peu,enmopnaR - po.n;orrcHH rJia3a (peri:errTOphr).
9. 3anaca1ow,aR - .HH'IHhIM aJih6yMHH, Ka3eHH MOJIOKa, ::m.n;ocnepMa ceM.HH.
10. To1Ccu'lec1CaJl - R,n; 3Meii:, rrayKoB, HaceKOMhIX.
«l>epMeHTLI
<PepMeHmbt (OT JiaT. fermentum -

3aKBaCKa) -

3TO

6eJIKH, rrp11cyTCTBYIO-

Ill;He BO BCex »<HBbIX KJieTKax H HrpaIOIIIHe pOJib 61:10Jior1:1qecKHX KaTaJIH3aTOpOB.

06~aJI

22

6uo.1iozuJI. OcnoBbi lfUmonozuu

CuHTe3MPYIOTC.H M3 aMMHOKMCJIOT Ha ptt6ocoMax B COOTBeTCTBMM c reneTV!'IeCKHM
KO,D;OM.
Ilo XHMH'IeCKOll npwpo.n;e ;JTO

6P.!llW (H:MeIOT CJIOJI<HOe CTpoeHH:e, qeTBepTlI'IttP6e.iK06ble KO.Mnoueflmbl (HOHbl MCT3Jlv'IOB, BHTaMHHb!).
BeJIKOBa.H qacTh cpepMeHTa Ha3hIBaeTc.H ano<jJep.Me1tm. He6eJIKOBa.H qacTh
cpepMeHTa Ko<jJaKmop - HOH MeTaJI.W HJIH HeopraHI1'IeCKOC seru;eCTBO; KOrpep.Meflm (K03H3HM) - opram1qecKoe seru;ecTBO.
CBoilcmea rpep.Menmo6:
HYIO CTPYKTypy) M

1.

BhlcoKa.H crrer~ml:mqHocTh, TO eCTb cpepMeHT H cy6cTpaT rro cTpoemuo
COOTBCTCTBYIOT )J.pyr )J.pyry.

2.
3.

,[(ettCTBHC B CTporo onpe)J.eJieHHOH IIOCJI€)J.OBaTeJihHOCTM.
YcKopettue orrpe)J.eJieHHhIX peil.KIJ;HH.

K.naccurjJuKau,u.R cpep.MeHmoe:
Cyru;ecTsyeT rnecTb KJiaccos cpepMeHTOB B 3aBHCHMOCTM OT Tuna KaTaJIH3MpyeMbIX peam:i;nfi::

1. OKcuDopeDyKmaJbl -

KaTaJIH3HpyIOT OKMCJIHTeJibHO-soccTaHOBMTeJib-

Hhie peaKIJ;HM.

2. Tpanc<jJepaJbl -

KaTaJm3MpyroT uepeHoc <PYHK1IHOHaJibHhIX rpyrrrr OT

O)J.HOro seru;ecTBa K )J.pyroMy.

3. I'uopo.na3bl -

KaTaJIH3HPYJOT rH)J.pOJIH3 -

peaKIJ;HH pacru;ernieHH.H CJIO)l{-

HhIX opraHuqecKHX seru;ecTB Ha rrpOCThie IIYT€M rrpncoe)J,HHeHH.H BO,ll;bI.

4. JI ua;Jbl -

KaTaJIH3HPYIOT

HCI'H)J.pOJIHTHqecKoe rrpncoe)J.HHeHHe HJill

OTru;errJieHHe <PYHKIJ;HOHaJihHOH rpyrrrrhI.

5. HJo.Mepaabt - H30Mepu3ar:i;u.H.
6. JI U?a3bl ( cuttmemaabl) - KaTaJIH3HPYIOT peaKIJ;HH CHHTe3a c
BaHH€M attepruu AT<l>.
<PyHKU,U.Jl rjJep.Meflm06 -

HCilOJib30-

ysemfqeHMe CKOPOCTM KJieToqHbIX peaK11Hll B MHJI-

JIHOHbI pa3.
Hu O)J.HH rrpor:i;ecc B KJieTKe He rrpottcxo)J.MT 6e3 yqacTH.H cpepMeHTOB (pe)J.yrrnnKa111rn ,[(HK, CMHTe3 PHK, 6e.JIKOB,

AT<l>,

cpoTOCMHTe3, )J.hIXaHue H )J.p.).

Jlnn~hl

y

paCT€HHH JIMIIH)J.hl CMHTe3HpyIOTC.H B KaHaJiax rJia)J.KOH

8TIC. y

JRHBOT-

HhIX IlOCTyrraIOT c rmru;ei1:, pacru;errJI.HIOTC.H )J.O COCTaBHhIX qacTeii (rJIMIJ;epMH
M mHpHbie KMCJIOTbl) H BHOBh CHHT€3MPYIOTC.H B co6CTB€HHhie JRHpbl. Co.n;epJRaHHe JRHpa B KJI€TKe COCT3BJI.H€T
CocTaB JIMIIH)J.OB.

5-15 'X,

MaCChI cyxoro seru;eCTBa.

JKupbt ( mpui.nuu,epuobt) -

CJIOJRHhie acpnphI TpexaTOMHoro

CIIHpTa l'JIH11€PHHa H BhICOKOMOJieKy.TI.HpHbIX JRHpHblX KHCJIOT.

cH 2

-

o - cp - R1

CH

-

0 -

CO-R 2

CH 2

-

0 -

CO - R 3

!

JKnpbl 6bJBaIOT HaChlIIJ;€HHbie H HeHaCblill€HHhie.

06w,a.R <jJopMy.na:

<Pep;nenmbt. JI unuilbi

H acbtuJ,eH llbU'

23

(TBCpJJ;bIC) -

cTcapttHOBaa,

naJihMHTIIHOBMI,

:vrnc;u-IHaJJ

KHC.'10Thl (rOBJ:I)KHH, CBHHOH mttp).

H e1taCbLUJ,eH Hbte

(mtt.n;Ktte) -

OjICHHOBaa, JIHHOJICBaa KIICJIOThI (no,r:i;co:r-

tteqrroe MacJ10, pb16tti1: )!{ttp).
ct>II3WlCCKHe CBOHCTB3 JKIIpOB:

JKHpbI nepaCTBOpHMbl B BO,r:i;e; paCTBOPHMbl

B opraHIFieCKHX paCTBOPIITCJI.HX (aqrnpe. CITIIpTe, 6eH3IIHe II JJ;p.).

JI unowh1 - C.'IO)KHhie scjmphl BbICOKOMOJieKyJIHPHhlX JKIIpllbIX I{IICJ'lOT
K IIHM OTHOCJITCJI IrnpOTHHOHJJ;hI (cjJOTOCIIHTCTnqeCKHC IlHI'MCHTbI),

II CIIHPTOB.

cTepOHJJ;HhIC ropMOHhl (ITOJIOBbie, I'JIIOKOKOpTllIWH,[\hI H .n;p.).

KJ1,accurjm1rnu,uJt JtunuooB:
1. IIpocmbte J1,Unu<Jb1 rrpe.n;cTaBJIJ:IIOT co6oi1: scpIIphI crrttpTa 11 :tJ{IIPHhIX KttcJIOT ( mpuz.11 uu,epUObt ):
cmepouiht - rrpOH3BO,L\Hhie crrttpTa XOJICCTepoJia (IIOJIOBhie ropMOHhI, xoJieCTepo.'I, KOpTmwcTepoII,nhl, BHTaMHH D);
mepneHbt - BCIIJ;CCTBa, B octtoBe CTPYKTYPhI KOTOphrx JiemttT rrJITIIyroJ1hHhIM yrJreBo,r:i;opo,r:i; uaorrpett (xJiopocpttJIJI, KapoTHHOH,L\hI, BHTaMHH

J1cupopacmBopu.Mbte Buma.MUHbt (A, LI;, E, K);
BOC/CG - C.'IO)KHbie scpttpbI BhICIIIHX JKHpHbIX

A);

KIICJIOT H BhICOKOMOJieKy-

mipHbIX CIIHpTOB.

2. CJLO;HC/lbte J/,UllUObl - :JTO COeJJ;HH8HHe
H JJ;pyr11x BCIIJ;eCTB. K HHM OTHOCJITCJI:
rjJOcrjJo.11unuobL -

::icpttpOB CITHpTa, JKHPHhIX KHCJIOT

coe,r:i;ttHeHIIJI I'JIHI~epmia, JKHpHh!X KHCJ'IOT H OCT3TKOB

q>0ccpopHOM KHCJ'lOThI;

crpuHzo.rtunuiJbt - coe,n;IIH8HHJI JIHIIH)J,OB H CIIHpTa CcjJHHI'03HHa;
Z.!IUKO.!IUnuiJbt - coe,n;11HeHHJI JIHilII)J.OB II yrJieBO,L\OB;
J/,UnonpomeuiJbL -- C08)J.HHCHHJI JIHilH)J,OB H 6eJIKOB;
nuz.Mellmbt (KapoTIIHOIIJJ;hI, reMOI'JI06HH, XJIOpocjJHJIJI H .n;p.).
<Py1-tKu,uu .11unuiJoB:
1. 3Hepzemu 'leCKaR - rrptt OKHCJI8HHH 1 r JIIIIIH)J,OB Bhl,L\CJ'lJICTCH 38, 9
T8IIJI3. JlttIIIIJJ;hl o6ecrreqHB3IOT

25-30 '%

K,llJK

BceH: sHeprHH, Heo6XOJJ;HMOH op-

raHII3My.

2. Cmpoume.flb/iaJl (cTpyKTypHaJI) 3. I'op.MOHaJlb/taJl ll pezyJ[,JtmOpHaR -

BXOJJ;JIT B COCT3B 6IIOMeM6paHhl.
CTepOH,L\Hhie ropMOHhl peryJ'IllPYIOT

o6MeH BeIIJ;eCTB H pa3MHOJKCHHC, BHT3MIIHhl.

4. 3anaca/OUJ,llR B )KHBOTHhIX -

B paCTHTCJihHhIX KJICTKax ceMHH IIJIOJJ;OB pacTeHHH,
JKHpoBaJI TK3Hb.

5. If cmO'lllUK ;miJoceHHOU 60iJbt 1,1

6.
7.
8.
9.

rrpH OKHCJICHHH

1r

JKIIpOB o6paayeTCJ:l

MJI BO)J.hl.

Tep.MOU.JOJ/,RU,UOHHaR - yqacTIIe B TerrJioo6Mette.
3aUJ,UmHaR - OT MexattnqecKHX IlOBpCJK)J.CHHM.
Bb1iJe.1umeJ1bliaR - y ttaceKOMhIX - «JKIIposoe TeJIO».
Cneu,uaJibllbte rjJynKu,uu - xttMnqecKtte curHaJihI (cpepoMOHhI

y HaceKo-

Mhrx), o6pa30B3HHe BO)J.OOTT3JIKHB3IOIIJ;HX ITOKphITIIH y pacTeHIIM (BOCKa), OKpacKa IIJIO)J.OB H CCMHH, yqacTHe B cpoTOCHHTC3e (rrHI'MCHThI).

06Ufa1l 6uo;wzu1l. OcnoBw ztumo;wzuu

24
YrJieBO,UbI

B

KJieTKax pacTeHHH yr;reso.n;hr CHHTe3npyIOTCH B X.i'Ioporv:racTax B npo11,ec-

ce cpoToCHHTe3a 113

C0 2

H 20

11

no.n; .n;eii:cTBHeM coJIHe•moro cseTa.

B

opraHH3M

;1rnsoTHhIX nocTyna:10T c TIIIIll,eii:. Co,n.epmamrn yrJ'Ieso.n;os B KJieTKe
(B KpOBll

0, 1--0, 12 'fr,), B paCTII'reJibHOH KJieTKe .n;o 90 'Yr,

1-5 'Yo

(1rny6HH KapToc}>eJIJI,

ceMeHa).
YrJieBO,D;hI -

:JTO yrJ1eso.n;opo.n;Hhie coe.n;IIHeHIIH. 06I11,aJI cpopMy.w: Cn(H 20)m.

KJtaccu<jJu 1cau,u.R y2JtrnoiJ06:

1.

IlpocThie yrJieBOJJ;hl -

MOHocaxaptt,n;hl.

B

3aBttCHMOCTH OT qIIcJia yrJie-

mpuOJbl (rJIHll,epaJih,IJ;erII.n;),
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B cocTaB HYKJieHHOBhIX KMCJIOT, AT<l>), zeKCOJbl (rn10xo3a, cppyKT03a).
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2.

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3TO ITOJIMMepbI, COCTOJ!Ill,He H3 MOHocaxapH,ll;OB,

coe,n;HHeHHhIX rJIMK03HJJ;HOI'i: CBH3hIO. Pa3JIIFrnroT:

• O.llUZOCaxapuiJbl -

yrJieBO.D;hI, IlOCTpOeHHhie H3 tte60JihIIIOro qucJia

(2-10)

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• iJucaxapuiJbl BOH caxap -

yrJieBO,D;hl, 06'he,D;HHHIOill,IIe ,D;Ba MOHOCaXaptt.n;a (rmw;e-

caxapo3a, COCTOMT H3 O.D;HOH MOJICKYJihl rmoK03hl H O,D;HOH

+ rJIIOK03a; JiaKT03a =
+ cppyKT03a);

MOJieKyJihI raJiaKT03hr; MaJihT03a = rmoK03a
K03a

+ raJiaKT03a;

• noJtucaxapuiJbt -

caxapo3a = rJIIOKo3a

rmo-

cJIO)KHhie BhICOKOMoJieKyJiapHhie yrJieso.n;hl, o6pa3o-

BaHHhie COTHHMH H ThIC.HqaMH MOJieKyJI MOHocaxap11,n.oB, HepaCTBOpIIMhle, He HMeIOT C.1'Ia,D;Koro BKyca. ::ho JIHHCHHhie 11 pa3BeTBJieHHhie TIOJIHMephl, MOHOMepbl KOTOphlX coe.n;ttHeHhl l'JIIIK03MJJ;HOH CBJI3hl0;

• ZOMono.riucaxapuiJbl -

COCTOHT II3 MHO)KCCTBa OJJ;HHaKOBhIX MOHOcaxa-

pH.!l;HhIX OCTaTKOB (KpaxMa.1'1, l'JIHKOl'eH, u;eJIJIIOJI03a);

• zemeponoJtucaxapuiJbl -

COCTOHT II3 MOHOCaxapH,D;OB pa3HhIX BIIJJ;OB (re-

nap1rn).

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cJio:arnhle npHpo,n;Hhie BhICOKoMoJieKyJIHpHhie coep;irneHtta,

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1-5 %

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Y i1te60dbt. II yK1teu1to6ble

KUCJWmbt. )_( / / K

25

A3or~1cToe

OCHOBaH11te
aAeHlltH, "1/1111

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4ll'IT03111H, ll'IJlH

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Ki.1CJ10Tbl

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II 0Jtutty101eomuab1 Maccoi1.

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Motto.111.epa.l1tu HYKJteu1toBbLX KucJ1om

m;tte H3 a30THCToro OCHOBaHH.H, caxapa JIOThI (pHC.

.HBJI.HIOTc.a

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cocToB-

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3).

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OCHOBaHH.H:

u; -

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B o6pa3oBaHHH HYKJieoTn:,n;oB rrpHHHMaIOT yqacTn:e ,n;Ba MOHocaxapu.n;a Jbt.

3To pn:603a -

B PHK H ,n;e3oKcn:pn:6o3a -

JieOTH,ll;OB .HB.'l.HeTc.a

ne1tmo-

B .IJ:HK. TpeThHM KoMrrotteHTOM HYK-

ocmamoK rpocrpopHou KUCJLombt -

rpocrpam. B 3aBHCIIMOCTn

OT

qn:cJia cpoccpaTHhIX rpyrrrr pa3JIH'18IOT MOHO-, ,L\H- H TpHcpoccpaTbl HYKJieOTH,11,0B.
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a c rrpn:coe,n;n:HeHHhIM cpoccpaTOM -

1tyKJteomuoa.Mu.

HyK.rteoJuaa.111.u,

Ha3BaHn:e HYKJieOTM,11,0B

o6yCJIOBJieHO BXO,ll;HIIJ;HMH B HX COCTaB a30THCThIMH OCHOBaHHHMH H o603HalJ:aeTC.H 3arJiaBHhIMH 6yKBaMH.

B

rrpn:po.n;e cym;ecTBYIOT ,n;Ba Tttrra HYKJieHHOBhrx KHCJIOT, OTJin::qaJOIIVIXCB

no COCTaBy, CTpoeHHIO H cpyHKIJ;lIHM: .IJ:HK JIOTa, PHK -

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pn:6ottyKJieHHOBa.a KHCJIOTa •

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TpexMeptta.a Mo,z:i;eJih CTpoeHHH .IJ:HK 6hIJia rrpe,nJiometta ,nm. YoTCOHOM

II

<I>.

KpHKOM B

1953

r. MoJieKy JI a ,[:t;HK cocTOHT H3 ,n;Byx rroJIMHYKJieoTH.n;-

Hhix u;errei1, 3aKpyqeHHhIX B rrpaBOCTOpOHHIOIO CIIHpaJib (pHC.
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2

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10

nap

HYKJieOTH,ll;OB. HarrpaBJieHHe u;enei1 ,IJ;HK B3aHMHO rrpoTHBorroJiomHo. Ha
KOHIJ;e BCer.n;a pacrrOJIOJKeH OCTaTOK cpoccpopHOH KHCJIOThl, Ha
60,n;Ha.H OH-rpyrrrra.

3'

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5·

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H cpoccpaTOB. Ilapbl a30TlICThIX OCHOBaHHH HYKJieOTH,ll;OB pacrroJioJKeHhl BHYT-

PII cnHpaJIH. Mem.n;y ocHoBaHH.HMH o6pa3yIOTCH Bo,n;opo,n;Hhre CBH3H. OcTOB
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Ba npHHIJ;Hrrax

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11

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26

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rrpoKapnoTnqecKoii KJJeTKe

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pacrroJJaraeTca B al];eptto:H 06;iacT11 -

(o.n;tta KOJlh1leBaH MOJieKyJia) l1 B 1llITOIIJia3Me (IIJia3MHll;bl -- Ma-

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,UHK).

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B MOJieKyJie

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HYKJieOTHAOB, TO B03HHKaIOT MyTa1IHH.

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(caMosocrrporr3Bel];ettne) -

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rrpaBJIHTh» B03HHK8IOill;IIe B ee IIeIIHX lI3MeHeHHH, TO ecTb BOCCTaHOBJiettne
rrpaBHJihHOii: nocJiel];OBaTeJihHOCTH HYKJieoTHll;OB.

PIIK (pu6ony1rneuno6aH 1mc.1wma)

27

<PylllCU,UU Jl,H K:
1. XpatteH11e HacJre,n;cTBeHHOH ttmpopMau;ttI:I
o cTpyKType cneu;I:IcpttqecKI:Ix .n;;rn opraHH3Ma 6eJIKOB.
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3. YqacTHe B pea"1113au;tttt reneTI:IqecKoil HHqJOpMaIJ;I:II:I, TO CCTb B npou;ecce Cl:IHTe3a IIOJ'IlIIIeIITll,ll,OB.

PHK

(pn6011y1~Jien110BaH KHCJIOTa)

PHK npe,n;cTaBJI.HeT co6oil: o,n;Hou;enoqequbril: .JIHHeHHhitt reTepOTIOJIHMep, COCTO.HII~llH ll3 pH60HYKJieoTll,D;OB. Hy101eomuiJ PHK cocTOHT H3 a3omucmozo ocnoeauuR (a,n;eHHH, ryaHHH, U:HT03HH, ypau;uJI), caxapa - pu6o3bt H ocmam1Ca <Poc<Popnou ICUCJtOmbt.
CB.H3h Mem,n;y HYKJieoTtt,n;aMH ocyru;ecTB,,'!.HeTC.H, KaK
H B o,n;HoiI H3 u;ene:H .[(HK, 'lepe3 yzJtepoiJ u ocmamo1C <Poc<Popuou ICUCJtOmbt. B OTJIHqHe OT .[(HK, co,n;epmmrne
KOTOpoil: B KJieTKax OTHOCHTeJibHO IIOCTO.HHHO, co,n;ep.IB:aHHe PHK KOJie6JieTc.H. 0Ho rrOBhlIIIaeTc.H B KJieTKax,
B KOTOphIX rrpottcxo,n;HT CHHTe3 6eJIKa. B 3aBHCHMOCTH
OT BhIIIOJIH.HeMhIX cpyHKIJ.llH pa3JIHqalOT HeCKOJihKO 6UPMC. 5. C08A"1H8H"18 HyKiJoe PHK:
neOT"1AOB s noni-iHyKneo1. Pu6oco.MHaJ1 PHK (p-PHK) - cocTaBJI.HeT
rnAHYIO L\8nb
85 '% OT Bce:H PHK B KJieTKe. 3To caMbre KpyrrHhre MOJieKyJihI PHK, B HX cocTaB Bxo,n;HT 3-5 Thrc. HyKJieOTH,11,os, MOJieKYJI.HpHa.H Macca ,D;OCTHraeT 1,0-1,5 MJIH. d, p-PHK CHHTe3MpyeTC.H B .H,11,pbIIIIKaX ll B KOMIIJieKce c 6eJIKaMH cpopMttpyeT cy6'he,D;HHllll;bl pll60COM.
Ha ptt6ocoMax u,n;eT CHHTe3 6eJIKa. Ptt6ocoMHa.H PHK, Bxo,n;.Hru;a.H B cocTaB
ll;llTOIIJia3MaTHqecKHX ptt60COM ayKapMOT, 60JihIIIe no pa3MCpaM, qeM
p-PHK ptt60COM npoKapMOT, MllTOXOH,D;PllH ll nJiaCTll,D;.
<PyHICU,UU p-PHK:
- cpopMHpOBaHHe aKTllBHOro u;eHTpa pH60COMhI;
- o6ecrreqeHHe B3attMo,n;eiicTBH.H ptt6ocoMhI u TpaHcnopTHoii PHK.
2. Hu<Pop.Mau,uouuaR PHK (u-PHK ), uJtu .Mampu'lHaR PHK ( .M-PHK) cocTaBJI.HeT 5 % oT Bceii PHK B KJien<e, KOJrttqecTBO ee 3aBHCHT OT cTa.n;11tt
KJieToqHoro IJ;HKJia. TaK, npu HHTeHCHBHOM CHHTe3e 6eJIKOB KoJiuqecTBO
tt-PHK ITOBhIIIIaeTC.H. Pa3Mephr tt-PHK pa3JIHqHhI u 3aBMC.HT OT o6'heMa
KorrttpyeMo:H MHcpopMaIJ;HH. CttHTe3HPYIOTC.H tt-PHK B .H.n;pe, B nporwcce
TpaHCKPHITIJ;llll Ha onpe,n;eJieHHOM yqacTKe MOJieKYJibI .[(HK - reHe.
<PyHICU,UR u-PHK:
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3. TpancnopmHaJl PHK (m-PHK) - cocTaBmreT 10 'X, OT Bce:H PHK
B KJieTKe. Cyru;ecTByeT 6o;ree 40 BH,L\OB T-PHK. MoJieKyJihI T-PHK caMhle

06Ufa1l 6uo1wzuK. Oc1to611t l{umollozuu

28

KopOTl\He, COCTOHT 113 80-100 HYKJieOTM.l(OB, MO.JieKyJIJipHaH Macca
25-30 Tb!C. d, T-PHK co.n:epmaTC.H B lV1TOIIJia3Me 10IeTKI1 l1 rrptto6peTaIOT crreurrcp11qecKyIO 1wmpopMa11,mo B BM.n:e «KJiesepttoro JIMCTKa» (cpopMa Tpl1JIHCTHmrn). Ha o,n;HoM IWHue Haxo,n;MTCJI TpMrrJieT nyK.'leoTn.n:os
(aHTHKo,n:oH), KO,n:MpyI0111,11f1 orrpe,n;e.'leHHYIO aMHHOKHCJIOTy. Ha .n:pyroM
KOHUe HMeeTC.H TpHrrJieT HyK.'leOTH,!l;OB, K KOTOpOMY rrp11coe.n:nH~IeTCH
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<PyH1W,U.H m-PHK:
TpaHcnopT aMHHOKHCJIOT K MeCTY CHHTe3a 6eJIKa - pn6ocoMaM.
JioKaJtu3au,u.H PHK B 101emKe:
PHK ttaxo.n:nTCJI B a,n;phillIKe, l!MTOIIJia3Me, pn6ocoMax, MHToxoH,n;p1rnx .
lI !lJiaCTH,11;3X.
B npnpo.n:e ecTh ellle o.n;MH BII,n; PHK - eupyc11a.H PHK. Y PHK-co.n:epmalllMX snpycon o6uapymeHa .n:syxIIerroqe•rnaa PHK, CTPYKTypa IWTopoii 6Jin3Ka
K CTPYKType ,[(HK. Y 3TMx opraHH3MOB PHK, KaK ,n;Byx-, TaK n o.n:uo11erroqetJuaa, BbIIIOJIHHeT cpyHKIIMIO xpaHeHnH: reHeTIItieCKOH nmpopMa11nn.

AT«l> (a,lJ,eH03HHTpmpoc<t>opuaa KHCJIOTa)
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B ,ll;BYX IIOCJie.D;HHX OCTaTKax cpoccpaTOB.
B COCTaB AT<l> BXO,ll;HT OCTaTOK 330THCTOro OCHOBaHnH: (aoeHUH), caxaprreHT03a (pu6o3a) H mpu ocmamKa <jJoc<jJop11ou KUCJlOmbt (pllc. 6). MoJieKyJia AT<l> oqeHh HeycTo.ifq1rna n cnoco6Ha OTllleIIJIH:Th o.n:tty nJIH ,n;Be MOJieKy-

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OCHO&aHHe

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29

+
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OTIIIeIIJieHmI MOJieKyJibl cpoccpopHOH KIICJIOTbl -

ao6om.n;eHIIeM

40

corrpoBom.n;aIOTC.H oc-

K,UJK/MOJlh aHeprMH. CB.H3II B MOJieKyJie AT<t> Ha3bIBaIOTC.H

Ma1Cpoapiu'lec1Cu.Mu II o6o3Haqa10Tc.H 3HaKOM «-». 3arracb1 AT<t> nocTO.HHHO
rrorroJIH.HIOTC.fl 3a cqeT rrpo11ecca <jJoc<jJopuJtupoeanuR, To ecTb rrpncoe.n11tteHII.fl cpoccpopHOM KIICJIOTbl K A.U<I>. <I>occpopMJIIIpoBaHIIe rrpOl:ICXO,nHT c pa3HOH l:IHTeHCl:IBHOCThIO rrpl:l .D;hIXaHHl:I (B Ml:ITOXOH.ZJ;pl:l.Hx), rJil:IKOJil:l3e (B IJ;l:ITOrrJia3Me ), cpOTOCl:IHTe3e (B XJIOpOIIJJaCTe ).
AT<t> .HBJI.fleTC.fl ocHOBHhIM CB.fl3YIOII~l:IM 3BeHOM Mem.n;y rrpo11eccaM11, corrpoBOJK.n;a10111I:1MIICH Bhl)l;eJieHIIeM Il HaKOfIJieHl:leM aHeprHH, Il rrpou;eccaMH, rrpoTeKaIOIIIHMH c 3aTpaTaMH aueprHl:I.
KpoMe AT<t>, B KJieTKax npHCYTCTBYIOT H .n;pyrIIe MOHOHYKJieoTH,D,hI, oTJIHqaIOIIIHecH CTPYKTypoii: a30THCTOro OCHOBaHH.fl: fT<t> (rya3HHTpttcpoccpopHaH KHCJIOTa); YT<t> (yptt.n;HHTpHcpoccpopHaH KHCJIOTa); UT<I> (1IHTH.ZJ;HHTpIIcpoccpopHa.H
KHCJIOTa).
3To TaKme MaKpoaprl:lqecKn:e coe.n;IIHeHIIH, :3Heprn:H rH.n;poJIM3a KOTOpbIX
HCIIOJib3yeTCH, Harrpn:Mep, rrpn: CHHTe3e 6eJIKa (fT<t>), IIOJIIICaxapII.D;OB (YT<I>),
cpoccpoJittrrII.n;oa (UT<I>). 3TII HYKJieoTM.ZJ;hI Heo6xo.n;ttMhI ,[\JI.fl cn:HTe3a PHK.
IlOMHMO MOHOHyKJieOTII)l;OB, BaJKHYIO pOJib B o6MeHe Be111ecTB ttrpaIOT iJu-

ltYKJteomuiJbt -

HA,U' (Hn:KoTn:HaMII,n,a.n;eHIIH.ZJ;HHYKJieOTtt.n;), HA,U<t>· (mmo-

THHaMI:1,n;a,neHIIH,ll;IIHYKJieOTH,n;q>0ccpaT), IIMe10111ne B CBOeM COCTaBe .n;Ba a30THCTbIX OCHOBaHII.H (a.n;eHHH II aMH,ll; HHKOTIIHOBOH Kl:ICJIOTbI), ,n;Ba ocTaTKa
PH6oab1 H ,n;Ba OCTaTKa cpoccpopHOH KHCJIOTbl (pn:c.
HA,U', HA,U<l>' -

7).

yHIIBepcaJihHhre a1Cl{enmopbt (oT JiaT. acceptor -

llHK) aaeprHII, a Hx BOCCTaHOBJieHHbie cpopMhI -

HA,UH n: HA,U<I>H -

rrpIIeMyHttBep-

CaJibllble ,ll;OHOpbl aTOMOB BO.n;opo.n;a (.n;Byx aJieKTPOHOB II o.n;Horo rrpOTOHa)
B 60JiblIIHHCTBe OKIICJIIITeJihHO-BOCCTaHOBHTeJihHbIX peaKIJ;IIH.

06u~aJl

30

6uo1wzuJI.

Ocu06bl ~umo;wzuu

Cy6CTpaT -- H 2 --r HA)], -> cy6cTpaT -r HAJI.H + H
0KHCJieHI18 pa3JIH'IHblX cy6cTpaTOB B rrpo1iecce 3HepreTWICC1Wro o6MeHa
npHBO.U.MT x Hmwrrnemno aTOMOB Bo.n;opo.n;a B B11.n;e HAJl.H (peam..1:i111 uJmJrn
Kpe6ca, OKHC01errne »01pubIX KIICJIOT HT . .n;.). HAJlH Mm!WT ncrro.'lb30BaThCH
B pa3o1I-PIHhIX peaKIJ,lUIX 6110CHHTe3a, OKHCJIHTbCH B iJ,blXaTCJibHOH uerr11 MHTOXOH,D;pHii:; ocBo6o:>K.n;eHHaH rrp11 3TOM 3Heprm:i 3arracaeTcH B Btt,n;e ATcI>.
HA,UcI>H 06pa3yeTcH nptt oKHCJIHTeJibHOM pacu~eu.1e1IHH yrJieBo;:i:on, y pacTeHHH - npH cpOTOCHHTe3e, a TaK:>Ke HCITOJib3yeTCH ;:1;.1H rro.1yc1e1nrn: 3Heprnn B )],bIXaTeJibHOH nerru MHTOXOH,n;pmI.
I\poMe HMIWTHHOBblX HYKJieOTH,ll,OB Mory'!' 6hITb rjJJta6U/-l06bl(' llljlfJlf'OmuiJbt
cI>AJl. (q>Jiamrna.n;emrn.n;rmyKJieOTH,ll;), cI>MH (cpJiaBHHMOHOIIYKJICOTH,ll;}, KOTOpble HBo'IHIOTCH npoH3BO,LJ;HhIMH BHTaMnHa B 2 (pu6oq>JiaBHHa). Onn rrpHcoe.n;nHHIOT ,n;Ba aTOMa BO,LJ;Opo,u,a ,llBa npOTOHa I1 ,llBa 3.'ICKTpona. l1x OKHCJieHHe
B .IJ;bIXaTeJibHOH u;enn MHTOXOH,llpllH TaK:>Ke conpoBo:>K.n;aeTCH BbI.U.eJieHHeM
3Heprnn, 3arracaeMoii: B Bn.n;e ATcI>.
Ilo xapaKTepy oprann3aunn H,llepnoro anrrapaTa Bee KJieTKH .n;e.1.HTCH ua
rrpoKapHOTH'leCKHe ll ::iyKapHO'l'H'leCKHe.

I lKHBM;,op<~

HeKRe~

- BllPYCbI,
- <jlartt

/~

IlpoKapllOTbl (,110$f,11epHble)

;)yxapttOThl (H.11CPHhie)

- 6aKTeptttt,

- paCTCHHH,

- u,nano6aincptttt

- lKllBOTHhie,

(C1!He3eJieHh!e BOJ\OpOC.~H)

- rptt6b1

HEK.nETO'"IHblE Cl>OPMbl >K"'13H"'1

BupycLI
BnpycbI 6hrnn OTKPhIThI B 1892 r. pyccKHM 6oTaHHKOM Jl.. l1. l1BaHOBCKHM,
KOTOPhIH BrrepBbie IIOJIY'IHJI HHtPeKUHOHHhIM 3KCTpaKT ll3 pacTeHHH Ta6aKa,
nopa:>KeHHbIX M03aH'IHOH 6oJie3HhIO. 8KCTpaKT rrponycKaJica <Iepe3 cpttJibTp,
crroco6HhIH 3a,u,ep.iKHBaTb 6aKTepHH, HO OTcpHJibTpOB3HHaa :>KH,ll;KOCTb e~e coxpamma 1rn¢eKUHOHHbre CBOHCTBa. B 1898 r. roJIJiaH.n.eu MapTHH BeiiepHHK
IIPH.llYMaJI CJIOBO « BHpyc »' qTo B rrepeBO,lle c JiaTHHCKOI'O o603Ha<IaeT «H.D; ».
To.1I1>1w c noMO~hIO &JieKTponnoro MHKpocKona y,u,aJIOCh yBn,n;eTb 3TH MeJibqa:ii:IIIue .JKHBbre cy~ecTBa.

He1oiemo'lHbLe tjJopMbL :JICU3HU. BupycbL

Bupycbt -

31

3To HCKJieTO'IHaJI cj_)OpMa :!KI13HM, cnoco6naH npomn<aTb B mirny 10

KJieTKY M pa3MHomaThCJI TOJihKO BHYTPH ee. OHM aBJIH!OTCa o6JrnraTHhIMH rrapa31naMH, HO, B OT.'IH'IHC OT .z:i;pyrHX rrapa3I:ITOB, napa3HTHPYIOT Ha retteTH'ICCKO:vr
ypoBHe. Cyw;ecTnyeT HeCKOJ'IhIW r11n0Te3 rrpoMcxom.z:i;eutta BttpycoB:
-

Bttpychl rrpott30IIlJrn OT K.1eTO'IHhrx opraHli3MOB.

-

BMpyChl npOI130IlIJil:I H3 «6er.10H» HYKJ1CHHOBOH Kl:ICJIOTbl, TO CCTb HYKJ1Cl:IHOBOH KHC.'IOThl,

KOTOpaa npuo6peJia cnoco6nocTh pen.'IHIIliPO-

BaTbCH He3aBHCHMO OT KJICTOK, B KOTOpbIX oua B03Hl:IKJia.
Bttpychl B03HMK.iIM B pe3yJ'IhTaTe .z:i;ereHepaIIMli (ni,1pom,n:enmi:) KJieTO'IHhIX opraHM3MOB.
BttpycbI pacnpocTpaHCHhl B np11po,n:e TIOBCCMCCTHO H nopa:ma!OT Bee rpynnbI
opraHH3MOB.

B HaCToaw;ee BpeMa OTIHCaHO OKOJIO 500 cj_)op::vi BHpycoB, nopama300 cj_)opM BHpycoB,

!Olll;HX TCTIJIOKpOBHhIX TI03BOHO'IHblX :!KHBOTHhIX, H 6oJiee

3apama10w;ttx BbICilIHe pacTeHHa. HayKa, H3yLia10w;aH nttpyc1>1, Ha31>rnaeTca

BupycoJwiueu.

20

Pa3Mepbl BHpycoB COCTaBJIHIOT OT
B

50 pa3

.z:i;o

3000

HM, TO eCTb B cpe.z:i;HeM OHli

MCHbllle 6aKTepHM.

Ilone,z:i;eHtte BMpycos:

o6JittraTHhre napa3HThI;

BbI3bIBaJOT 3a60JieBaHttH;

B KJICTKax-xo3Renax «BhIKJIJOLiaJOT» .IJ:HK M HCTIOJih3YJOT cnott .IJ:HK H PHK, .z:i;aJOT KJICTKe KOMaH,z:i;y CliHTC3HpoBaTb CBOM KJ'ICTKH;

nepe.z:i;a!OTCJI I13 KJICTKH

B KJICTKY B BM)J;e HHepTHhIX 'IaCTI11l.

CTpoeHHe BHpycoB

Cepau,eBuHa 6upyca

(p11c.

8)

COCTOHT H3 cj_)parMeHTa reHeTH'ICCKOro MaTepHaJia (.IJ:HK,

PHK). BttpychI co.z:i;epmaT ncer.z:i;a O)J;HH THn HYKJieHHonoii: KMCJIOTbI, npHLieM KaK
O)J;HOIJ;CIIO'IC'IHOH, TaK H .z:i;nyx1.1,eIIO'IC'IHOH, KaK JIHHCHHOH, TaK H KOJlbIICBOH.

Kancua -

3aw;HTHaR 6eJIKOBaR 060-

JIOLIKa, KOTopaR 3aw;ttw;aeT .IJ:HK, PHK
OT cpepMeHTOB-HYKJiea3 H

Y<l>-H3JiyLie-

HH:H, o6ecneLIHBaeT ocam,z:i;eHtte Bttpyca
Ha IlOBepxHOCTH KJICTKli-X03RHHa.

CynepKancua -

2

.z:i;onoJIHHTeJihHaR

JIHrronpoTeH,z:i;HaR o60JIO'IKa,

KOTOpaR

B03HHKaeT H3 IIJia3MaTH'ICCKOH MeM6paHbI KJieTKH-X035IHHa (xapaKTepHa )J;JIR
BhICOKOOpraHH30BaHHbIX

BHq,

BHpycoB

-

rpHnna, repneca).

IlOJIHOCTbIO ccj_)opMHPOBaHHaR

HH-

<l>eKIIHOHHaR 'IaCTHu;a BHe KJICTKH-X03.HHHa Ha3bIBaeTC.H

BapuOHOM

rrpoTeH,z:i;HbIH KOMrrJieKc).

(HYKJie-

EcJIH Bttpyc

P111c. 8.

CxeMarnYeCKlll~

.pa3pe3 s111pyca,

111Me10L11ero KancoMepHoe crpoeH111e: 1 -Kanc0Mepb1 (KanC'1A -

6enKOsa11 o6onoYKa);

Haxo.z:i;HTCR BHYTPH KJieTKH-X03HHHa,

2 - sHyTpeHHS111 cpeAa s111pyca; 3 - reHeT111-

TO OH cyw;ecTByeT B cj_)opMe HYKJICHHO-

YecK111~ Marep111an (,ll,HK 111n'1 PHK); 4 -

BOH KllCJIOTbI.

060-

06tqaK 6uo1wiuJl.

32

Oc1W6bt

q,umo.noiuu

060JI01ma Bllpyca ITOCTpOeHa 113 o,n;HHaKO-

1wnco-

BbIX IlOBTOP.lUOll\lfXCR cy6oe.n:1rnm.1, -

Mepo6,

KOTOpbie o6pa3yIOT CTPYKTYPhI c BhICO-

KoiI CTeneHbIO CHMMCTpllll, cnoco6Hhie KPHCTaJIJill3llpOBaTbCR.

BoJihUIHHCTBO BHpycoB

ITOCTpOeHO no O,UHOMY ll3 ,n;Byx TllilOB CHMMeTpHH -

cnupG.llbHOU

HJIH

1Cy6U'l.l'CICOU.

Ilo crmparrhHOMY THIIY CHMMeTpHII noCTpoeHO 60JibllIHHCTBO BnpycoB, nopa)KaIOll\llX pacTeHM.H, l1 HeKOTOpbie BllPYCbl 6aKTepnii: (6aKTepuocpam). Bnpyc rn6a'IHOH M03aHKH (BTM) HMeeT CilllpaJibHYIO CllMMeTplIIO,
co.n:ep)KlIT BHYTPH crrHpaJihHYIO PHK Karrcw,n;
COCTOHT

H3

2130

O)J;HHa:KOBbIX 6eJIKOBbIX

cy6'be)J;HHHI.I,, cocTaBJI.HIOll\HX BMecTe c PHK

PHc. 9. CxeMaTw-iecKoe vi306pa>t<eHvie
6aKTepvio¢ara:

1-

ronosKa c 11Koca3,a-

pwiecKOCi c11MMeTp11eii\;

2 -

nOllblei l\l11l11HAPl14€CKl1iil OTpo-

CTOK; 4 -

Yexon co cn1-1panbHOCi CillMMe-

Tpilleii\ 1'13 COKpaf1'1TellbHblX 6enKOB; 5 ·-

6 -

XBOCT0Bbl8

HYKJZeo-

Kancua.
BOJihIIIa.H

soporn11-

40K; 3 -

6a3anbHBSl nnacrnHa;

e;:i;HHYIO I.I,eJIOCTHYIO CTPYKTYPY -

IaCTh BnpycoB, BbI3bIBaIOll\lIX

1

HHQJeKI.l,HlI y qeJIOBeKa lI )KlIBOTHbIX, HMeeT
Ky6H'IeCKHH THil CHMMeTpnn. Karrcu,n; HMeeT
QJOPMY HKOcaa,n;pa THrpaHHIIKa c

12

npaBMJlbHOro ,n;Ba,UI.I,a-

BepIIIIIHaMII II

30

pe6paMII

(Bttpyc IlOJIIIOMHeJIHTa).

HJllT1'1

Baf(mepuo<Pazu -

rpynna BHpycoB, no-

pamaIOll\HX 6aKTepHH (IlO)KHpaTeJIH 6aKTepttii:). 0TKphITbl B
CKHM MHKpo6110JIOrOM <l>pe,n;epHKOM TyopTOM. ll.MeIOT

1915 r.

aHrJIHH-

UIWCa3apu'l.eCf(yHJ

?O-

Jl061CY (con;epmall\YIO reHeTH'-IeCKllH MaTepttaJI) ll X60Cm, o6Jian;aIOll\HH Cilll-

paJihHOH CHMMeTpne:H (pHC.

9).

8TH BHPYChI o6HTaIOT B KHIIIe'IHHKe 'ICJIOBeKa

ll )KllBOTHbIX, OHM IlOJie3HhI, TaK KaK rropamaIOT 6aKTepttll. B Me,[l;HI.l,HHe HX
npHMeH.HIOT ,ZJ;JI.H Jieqem-rn 6pIOIIIHOro TH<}Ja, XOJiephI.
KJiaccHcpttKaI.I,n.H BttpycoB cTponTC.H no BH,n;y H cpopMe ux HYKJiettHOBOii

KlICJIOThI,

THITY ClIMMeTplIH,

HaJIH'IHIO lIJIH

OTCYTCTBHIO BHeIIIHett

MeM6paHhI.
0TJIH'IHe BHpyca OT KJICTKH: BHPYCbl HHKOrp;a He pa3MHO.iKaIOTC.H n;eJieHHeM; B CBOCM COCTaBe HMeIOT TOJ1hKO O.D;HH THII HYKJICHHOBOii KHCJIOTbl (,Zl;HK
HJIH PHK); HHKor.n;a He pa3MHOJKaIOTC.H BHe KJieTKH-X03.HHHa.
Pa3MHO.iKCHHe (penpon;yKUHH) BKJIIO'IaeT Tpn nponecca: perrJIHKaI.I,HIO BMpycHoH: HYKJieHHOBOH KHCJIOThI; CHHTe3 BHPYCHhIX 6eJIKOB; c6opKy BHPHOHOB.

Y BHpycoB HeT rroJia.
HOH IrneTKH -

Penmrnau;H.H MOJJeKyJihI ,ll;HK (PHK) BHYTPU nopameH-

MHOr03TaI1HhIH npor,ecc, KOTOpbIH COCTOHT H3 IIIeCTH CTa)J;HH:

1. AiJcop6u,uR -

npouecc rrpHKpenJieHII.H BHpyCHhIX qacTHI.I, K rroBepxHoc-

TH KJieTKH.

2. HwoelCU,UR

(y 6aKTepuo<}JaroB) -

npoHHKHOBeHHe BHPYCHOH qacTHUhl

B KJJeTI{Y H BBe,n;eHHe HYKJieHHOBOH KllCJIOThI H3 6eJIKOBOrO Kanctt.n;a.

HeicJ1,emo'tnbte tjJopMbt :J/Cu3nu. Bupycbt

33

PHK-co11epman1He

]J:HK-co11.epmanvie

1 HHTb HYKJieH-

2 HHTH HyK-

HOBOff KHCJIOTbl

JieH:HOBOff KllC..'IOThl

·· +6aKTepHo<j:lar

1 HHTb HYK"1em1oeoii
KHCJIOTbi
3HTepoeHpycbl,
- BHpyc rpHnna,
- BHpyc 6erneHCTBa,
·- BHpyc Ta6a•moii
M03RHKH (BTM)

- a11.eHOBHPYCb1,
- BHpycHaH ocna,
- repnec

2 llH'rH HyK~'IeHHOBOJi KHCJTOThl

- OHKOreHHble
BHpycbl,

- BM'I

3. PenJtu1Cau,u.R MoJte1CyJt Bupycnou ny1CJteunoBou /CUCJtOmbt -

rrpottcxo-

,n;ttT 3a cqeT HYKJieOTH,ll;OB, HaKOIIJieHHblX B KJieTKax X03HHHa.

4. Cunme3 eupycttbtX 6eJt/COB u rpepMenmoB -

rrpottcxo,n;ttT Ha ptt6ocoMax

KJieTKH.

5. C6op1Ca eupycttbtX 'tacmuu, -

3a cqeT rropa:meHHhIX BHPYCHhIX HyKJie-

HHOBhlX KHCJIOT H BHpyCHbIX 6eJIKOB.

6. Jiu3uc -

Bh1xo,n; BttpycHhIX qacTHIJ H3 rropa:meHHOH KJieTKH.

ptt:H corrposo:m,n;aeTCH pa3pyllieHHeM (JIH3HCOM) KJieTKH.

y

Y 6aKTe-

ayKapHOT

rrpOHCXO,ll;HT rryTeM BhlIIHqHBaHHH
o60JIOqKM KJieTKH M

7

«BbITaJIKMBa-

HHeM» BttpycHbIX qacTMIJ B oKpy:maIOIIJYIO cpe,n;y.
HoBbie BttpyCHhie qacTHIJhI 3apa:maIOT
eIIJe Herropa:meHHhie KJieTKH, M IJHKJI pa3BHTHa BMpyca IIOBTOpHeTCH.

3tta'lenue eupycoe

Y JKHBOTHhix H

qeJioBeKa BHpycb1 BbI3bI-

BaIOT, cpe,n;n rrpoqnx, TaKtte 3a60JieBaHHH,
KaK 6emeHCTBO, rerraTMT,
Kpacayxy, ocrry,

OP3,

rpHIIII,

KOpb,

rroJIHOMHeJIHT, 3H-

11e<l>aJIHT, Cillf.[(, paKOBble OIIYXOJIM.

Bo36yaumeJtb CIIH!fa (pttc. 10) -

BH-

pyc HMMyHo,n;e<l>HIJMTa qeJioBeKa (Blfl:J).

o.

lfMeeT c<t>epHqecKyIO <l>opMy ,n;ttaMeTpOM

Pi.tc. 1

100-150 HM.

Ye11oseKa: 1 -

HapymHaa o6oJioqKa BHpyca

COCTOHT H3 KJieTOqHoH MeM6paHbl KJieTKH-

3 -

8111pyc VIMMYHO,D,ecj>m.(VITa
cep,D,u,esi-1Ha; 2 -

PHK;

o6parnasi TpaHcKpi-1nTa3a; 4 -

X03HHHa. B MeM6paHy scTpoeHbI pe11errTop-

nporna3a; 5, 6 - n111ni-1,1J,Hasi ,D,sycnoi:iHasi

HLie «rpH60BH,ll;HLie»

MeM6paHa; 7 - i-1Hrnrpa3a

o6pa3oBaHHH. IIo,n;

06UfaJi 6uo.1wzuJi. Ocno6b£ q,umo.11.ozuu

34

HapymHoli o6oJioqKoli pacnoJiaraeTca cep.n;:u;emrna BHpyca c reHeTuqecKMM MaTepttaJioM B BH.n;e .n;Byx Mo.wKyJI PHK (Kam.n;aa H3 9 reHoB Blll.J) H cpepMeHTa
(o6paTHaJI TpaHCKpMnTa3a). 8TOT cpepMeHT KaTaJIM3HpyeT peaKIJ;HIO o6paTHOH
TpaHCKpnn:u;nu B KJieTKax JIMMcpo:u;uToB. Bupyc nopamaeT rJiaBHhIM o6pa3oM T·
xeJtnepbt JlUM<jJo~umo6, Ha noBepxHOCTH KOTOPhIX ecTh pe:u;enTophl, cnoco6Hhre
CBH3bIBaThCH c IIOBepXHOCTHhIM 6eJIKOM Bllq. B TaKOM COCTOHHHH OH MO:>KeT
coxpaHHThCH ,ll;OJirO, He npOHBJI.fUI ce6a. lIMMYHHa.H CMCTeMa opraHH3Ma qeJioBeKa yTpaquBaeT CBOH 3aIIJ;HTHhie CBOilCTBa, H opraHH3M rrepecTaeT 6opOThCH
c JII06oii: :irncpeK:u;n:en, paKOBhIMH KJieTKaMH H rrorn:6aeT. Cpe.n;HHR npo.n;om1mTeJihHOCTh :>KH3HH HHqmrrupoBaHHoro qeJIOBeKa COCTaBJI.HeT 7-10 JieT. llcToqHHKOM 3apameHHH CJIY:>KHT TOJihlW qeJIOBeK - HOCHTeJib BHpyca HMMYHO,D;ecpn:rrHTa. CIIH,L( rrepe.n;aeTca rroJIOBhIM rryTeM, qepe3 KpOBh M TKaHn:, co.n;epmar:u;ue
BMpyc HMMYHO,D;ecpHrrHTa, OT MaTeptt: K IlJIO,ll;y.
YpacTeHHll BMpychl BhI3hIBaIOT CBOH 3a60JieBaHMH, HarrpHMep Bttpyc Ta6aqHOR M03aHKM, meJITYXH CBe.KJihl H .n;p.
Bnpychr (6a.KTepuocparn) HCIIOJih3YIOT lJ.JIH JieqeHHH ,ll;H3eHTepuu, 6p10IIIHoro TMcpa, xoJiephI.

K.nETO'"IHblE ct>OPMbl >KH3HH

Ha,11;11,apcTBO npoitapuoTbl
K rrpo.KapHOTH'IeC.KHM (,n;oa,n;epHhIM) opraHH3MaM OTHOCHT 6a1cmepuu n: cuHe3eJteHbte eooopocJtu (rruaHo6a.KTepuu). 3To caMhie npocThie, MeJih'IaRUIHe
H Han:6oJiee pacnpocTpaHeHHble opraHH3MhI. 0HH 6blJIH e,D;HHCTBeHHOll cpopMOH
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yqacTByror:u;n:e B IlOCTpoeHHH KJieToqHhIX rreperopo,n;o.K H B perrpo,n;y.KD;HH
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6aKTepHH HMeIOT :JKI'YTHKH 6oJiee rrpOCTOI'O CTpOeHHH, qeM y ayKapHOT.

35

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36

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37

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41

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XpoMocOMbL (oT rpe'L chroma rrpe,n;JiomeH B

1888 r.

ll,BeT, KpacKa, n soma -

TeJJo); TepMMH

HeMeu;KMM yqeHhIM Bn:JJhreJJhMOM BaJih,n;eilepoM) -

rrpe,n;cTaBJIJilOT co6oii KOH,n;eHCMpOB3HHhIH xpoMaTMH BO speMJI MMT033 HJIH
MeH033. OcHOBY xpOMOCOMhI COCTaBJJ.HeT ,n;syxu;erroqeq:HaH MOJieKyJJa ,D;HK, KOTOpaa no,n;sepraeTCH yirna,n;Ke,

rrpno6peTaJI orrpe,n;eJJeHHYIO TpexMeptty10

CTPYKTYPY, HJJH KOHcpopMaun:10. Bh1,n;eJ1HIOT HecKOJihKO yposHeii rrpocTpaHcTBeHHoii yKJJa,n;KM -

KoMnaKmu3au,uu /l,HK (cM. pa3,n;eJI «I'eHeTnKa» ).

XpoMocoMhI u3yqa10TcJI B MeTacpa3e MMT03a. MeTacpa3HaR xpoMocoMa coCTOI:IT n:3 ,n;syx xpoMamua (pMc.

15).

HMeeTcJI nepeu'lna.R nepemRJKKa,

HJIH u,ettmpoMepa, KoTopaa ,n;eJIMT xpoMocoMy Ha nJte'lu. HeKOTOphle xpoMocoMhl HM€lOT 6mOpU'l1-tYIO nepem.RJKICY H cnym-

HUIC (yqacTOK KOpOTKOPO IIJJetia, OT,n;eJIHeMbIH
BTOpntIHOii rrepeTHJKKoii).

KoHUbI xpoMocoM

H33hIB3lOTCH me.1wMepaMu.

<f.>yHICU,UU R8pa: xpaHeHn:e H3CJI€,Zl;CTBeHHOH
MHcpopMaUMM H nepe,n;aqa ee )l;ffqepHHM KJleTKaM
B npou;ecce ,n;eJieHHH, peryJIHIJ:HH n: KOHTpOJih npoueccos JKM3He,n;eHTeJlhHOCTH KJieTKH, yqacTHe
B CHHTe3e 6eJIK3 (TpaHCKPHIIUHJI), M€CTO o6pa30BaHMH cy6'he,ZJ;MHHU pH60COM.

PMc. 15. XpoMocOMbl:

1- MeTaL1,eHTJ)11YecKasi; 2- cy6Mernu,eHTp11yecKasi; 3, 4- aKpou,eHTp11YecK11e. CTpoeH11e

PaaJIHqHhJe THIIhI ay1mpu0Tuqec1mx .KJieTo.K
3yKapMOTHtieCKHe opraHH3MhI ,n;eJJHT Ha TpH
uapCTBa: pacTeHI:IH, :IKHBOTHhlX H rpn:6os.
PaccMoTpes o6m;Hii 1rnaH cTpoeHMJI KJieTKH

16, 17),

XPOMOCOMbl: 5 - LJ,eHTPOMepa;

(pnc.

6 - BTOpwmasi nepeTSJ)KKa; 7 CnYTH11K; 8 - xpoMaT11Abl; 9 -

Hble oco6eHHOCTH pacTHTeJihHOH H :IKHBOTHOM

TeflOMepbl

B pa3,n;eJie «BOT3HMK3» ).

KJieTKI:I

MOJKHO Bhl)l;eJIHTh OTJIMtIMTeJih-

(KJI€TKM

rpM60B

paccMaTpnBalOTC.H

47

K11,emo'lnbie </JopMw JtCUJnu. HaiJu,,apcmBo 3y1wpuomw

CpaBueuHe cTpoeHHH pacTHTeJibHOH H ;KHBOTnoii KJieTOK

PacTHTeJihHaH KJieTKa

IlpH3HaKH
IIJiaCTH/:lbI

XJiopornracTbI,

:lKHBOTHaH KJJeTKa
0TCYTCTBYK>T

xpOMOTIJiaCTbI,
JieHKOTIJiaCTbl
reTepoTpOcpHbIH

Cnoco6

ABTOTpOcpHbIH

nuTaHHH

( cpoTOTpocpHbIH'

( canpoTpOcpHbIH,

xeMOTpocpHbIH)

napa3HTH'leCKHH)

B xJioponJiacTax,

B MHTOXOH,llpHHX

CuuTe3 AT<I>

MHTOXOH,llpH.HX
Pac~enJieuHe

B xJioponJiacTax H Bcex

Bo Bcex 'laCTHX KJieTKH, r,11e

AT<I>

qacT.HX KJieTKH,

ueo6xo,11HMbI 3aTpaTbI

r,11e Heo6XO,!IHMbl

3HeprHH

3aTpaTbl 3HeprHH
KJieTO'IHhIH

y

HH3IUHX pacTeHHH

Bo Bcex KJieTKax

n;eHTp
ll;eJIJIK>JI03HMI

PacnoJiomeua cuapymH OT

KJieTO'IHa.H

KJieTO'IHOH MeM6paHbl

0TCYTCTByeT

CTeHKa
BKJIK>'leHH.H

3anaCHble TIHTaTeJibHbie
Be~eCTBa

B BH):le 3epeu

3anaCHbie TIHTaTeJibHbie
Be~eCTBa

B BH):le 3epeH H

KpaxMaJia,6eJIKa,KaneJih

KaneJih (mHpbI, yrJieBo,11

M8CJia; BaKyOJIH c KJieTO'IHbIM

rJIHKOreu); KOHe'IHbie

COKOM;KpHCTaJIJibICOJieH

npo,11yKTbI o6Meua,
KPHCTaJIJibl COJieli; nHrMeHTbl

BaKyoJIH

Kpynubre noJiocTH,

CoKpaTHTeJibHble,

3aTIOJIHeHHbie KJieTO'IHblM

TIH~eaapHTeJibHbie,

COKOM - BO,!IHbIM paCTBOpOM
pa3JIH'IHbIX

Be~eCTB,

.HBJI.HK>~HXC.H

Bbl):leJIHTeJibHble B8KYOJIH.
06bl'IHO MeJIKHe

3anaCHbIMH

HJIH KOHe'IHbIMH
npo,11yKT8MH.0CMOTH'leCKHe
pe3epayapbI KJieTRH
CB.H3b Mem,11y

I1Jia3MO):leCMbl

KJieTKaMH

Ilepeuoc qepe3 MeM6paHhI,
KJieTO'IHble KOHTaKTbl

,lJ;eJieHHe

KJieTO'IHa.H neperopo,11Ra

IlepeT.amRa 06pa3yeTC.H B

Il;HTOTIJI83Mbl npH

CTpOHTC.H B n;eHTpo6emHOM

n;eHTPOCTpeMHTeJibHOM

MHT03e H MeH03e

uanpaBJieHHH

uanpaBJieHHH

rJIHKOKaJIHKC Ha

0TCYTCTByeT

llMeeTC.H

aapymuoii:
MeM6paue

06~aJt 6uonozuJt. Ocn06bL 1fUmonozuu

48

J - -·- \

~ft=~~..-- - --- 22
-21

2

20
-J9
18

3 ..

17
16

5
6

PMc. 16. CrpoeHi.-ie pacrnTeJTbHOiii KJTeTKi.-i :

PMc. 17. CrpoeH11e :>Kl1BOTHOiii KJTerKi.-i: I -3KJou,111-

1-

roa; 2- ceKperopHbli1 nyab1peK; 3

KJTeTOYHafl CTeHKa; 2 -

flAepHafl o60JTOY-

3 - flASPHafl nopa; 4 - flAPblWKO; 5 flAPO; 6- xpoMarnH ; 7- rflaAK"1iii 3HAOnna3MaT"1Y9CK"1ill peTi.-iKynyM ; 8 - nnaJMantYeCKafl M9M6paHa ; 9 - U,"1TOnna3Ma; 10 - Ml1 KKa;

porpy604Kl1 ; 11 -

M11KPOt:fmnaMeHTbl; 12 -

annapar ronbA)f(11 ; 13 -

n113oc0Ma ; 14 -

Honnacr; 15 -

KfleTOYHbli1 COK ; 16 -

XOHAP1111 ; 17 -

rpaHa; 18 -

ponnacra;

ro-

Ml1TO-

MeM6paHa xno-

19 - xnoponnacr; 20 - p116oc0Mb1

- n11U-1esapi.-irenb4 - annapar ronbA)f(11 ; 5 - M11Kpo<f>11naMeHTb1; 6 - MmoxoH.Qp1111 ; 7 - wepoxosarb1i1 3HAonna3Marw-iecK11i1 pernKynyM; 8 - p116ocOMb1,
Ha11 aaKyonb;

CBS'l3aHHb18 c 3HAOnna3MaTll1Y9CKl1M per11KynyMOM ;

9- nna3M8Tl1Y9CKaR MeM6paHa; 10 - U.11TO-

nna3Ma ; 11 -

13 -

3yxpoMarnH ; 12 -

flAPblWKO; 14 -

o6onOYKa; 16 -

r erepoxpoMarnH ;

flA8PH8fl n o pa; 15 -

flAepHafl

AB9 U,9HTPl10!111 , pacnonmKeHHble

nOA npl'IMblM yrnOM Apyr K Apyry; 17 -

CB060AHble

c 3H.QOnna 3MarnYeCK11M pernKynyMOM; 21 -

p1160COMbl; 18 -

wepoxosarb1iii 3HAonna3MarnYecK11ill per11Ky-

20 - rnaAK11i1 3HAOnna 3M8Tl1Y9CKl1iii per11KynyM;
21 - JTl-1HOU,11T03Hbliii ny3blpeK ; 22 - n11HOU,11T03;

22 - CB060AHbl9 p1160COMbl; 23 - KJTeT04Hafl CTeHKa COCeAH11X KJTeTOK; 24 - nnaJ-

nyM;

MO.QeCMbl

J11130COMa; 19 -

M11KPOTpy6o4K11;

23 - Ml1KPOBOPCl1HKa; 24 - nornou_\eH11e 11!111 ceKpeU,11R y OCHOB8Hl1R M11KPOBOpCl1HOK; 25

-

RAPQ

06MEH BEW.ECTB M nPEBPAW.EHME 3HEPrMM
Bee JRHBhie opraHH3MhI, o6HTaroru;He Ha 3eMJie, .HBJUIIOTCH OTKphIThIMH CHCTeMaMH, ,ll;Jl.H KOTOpblX xapaKTepeH HerrpephIBHO rrpoTeKarorn;Hii o6MeH Bern;ecTB H aHepruu c 01<pyJ«a10rn;eif cpe,n;oii.
06MeH Bern;ecTB H 3HeprHH - OCHOBHOe CBOHCTBO JRHBoro. Bee rrpou;ecCbl
JRH3He,n;enTeJihHOCTH (pocT, pa3BHTHe, pa3MHOiKeHHe H ,n;p.) HeB03MQ;KHhl 6ea
IlOJiy'leHH$l H3 OKpyJ«a10rn;ell: cpe,n;bl B npou;ecce IIHTaHH.H Bem;eCTB ( CTpOHTeJihHoro MaTepHaJia) H 3HeprHH, Heo6xo,n;HMhIX ,ll;JIH. nocTpoeHMH. XMMHqecKHX coe,n;HHeHHM opraHH3Ma, a TaKJ«e ,ll;JI.H COBeprneHMJf pa3JIM'IHOH pa60Thl.
CoBoKynHoCTh pea1<u;uif o6MeHa, npoTeKa10w;ttx B opraHH3Me, Ha3blBaeTcn
Me ma60JIU3MO.M.

06Men Beiq,ecmB u npe6paiq,enue :mepzuu

49

MeTa6oJIH3M BK.moqaeT B cefor ,[(Ba rrpou;ecca:

1. A11a60JtU3M (accn::vrn:m1u;ttH,

rrJiacTn:qecm1ii o6MeH) -

coBoRynHocTh 3H.L\O-

TepMIIqecRttx peaKu;Hii 6IIOCHHTe3a, KOTOpble npHBOl\flT K o6pa3oBaHHIO
CJIO)KHhlX Beru;ecTB H3 60J1ee npOCTbIX ( CHHTC3 6eJIKa, rJIHKOf'eHa; cpOTOCHHTe3, xeMOCHHTe3).

2. Kama60JtU3M

(,n;IICCHMIIJIHD;IIH, ;mepreTIIqecKIIH o6MeH) -

COROI{yn-

HOCTh 3K30TepMIIqeCKHX peaKD;HH pacna,[(a II OKHCJieHirn BhlCOKOMo.1eKYJIHpHbIX Beru;ecTB

.n;o

6oJiee npocTbIX.

8TM rrpou;eCChl B3aMMOCBH3aHhl BO BpeMeHM M rrpocTpaHCTBe, flB.1flIOTCfl 3BeHbHMH MeTa6oJIM3Ma. Pea.KD;IIM 6MOCMHTe3a HeB03MOIBHhI 6e3 3HeprMM, KOTOpaa
BhI.n;em:i:eTCH B peaKD;llHX 3HepreTIIqecKoro o6MeHa. PeaKD;HH ,n;IIcc11M11Jinu;nn ne

MYT

6e3 <t>epMeHTOB, 06pa3y10ru;nxcH B peaKu;Max rrJiaCTMqecKoro o6MeHa.

Ilo THilY IlHT3HHH BCe )KHBhie opraHM3Mhl rro.n;pa3,n;eJIHIOT Ha:

1. Aemompo<jJ11bte -

crroco6Hhre CMHTe3MpoBaTh opraHMqecRne Beru;ecTBa

H3 HeopraHnqecKnx.
<l>OTOCHHTeTllKll (cpOTOaBTOTpO<t>hI) -

K HMM OTHOCHTC.H 3eJieHhie pacTe-

HllH ll <t>oTOCHHTe3npyroru;ne 6aKTepnII, KOTOPhie MCII0Jlh3YIOT 3HeprMIO
COJIHeqHoro CBeTa. Bee OCTaJihHble )KllBhie cyru;ecTBa MCTIOJih3YIOT 3HepI'HIO, 3aKJIIOqeHHYIO B XHMnqeCKllX CB.H3HX.
XeMOCMHTeTMKM (xeMoaBTOTPO<l>hI) -

K HMM OTHOC.HTC.H HeKOTOphie

6aKTepnn, KOTOphie llCIIOJih3YIOT 3Heprnro, BhI,ll;eJI.HIOiu;YIOC.H npM OKllCJieHllll HeopraHnqecRnx coe,n;nHeHnii ( ceposo.n;opo,n;a, aMMMa.Ka, IBeJie3a M ,n;p.).

2. I'emepompo<jJHbte IIIHHCTBO 6aKTepnii,

IBllBOTHhie, rpn6h1, He3eJieHhre pacTeHIIH, 6oJih-

He cnoco6HhIX caMOCTOHTeJihHO CMHTe3MpoBaTh opraHM-

qecKHe Beru;ecTBa ll MCIIOJih3YIOiu;MX ::meprnIO XMMlfqeCKHX CBH3eii I'OTOBhIX oprrurnqecKHX coe,n;nHeHnii.
Carrpo<t>nThI (carrpoTpo<t>Hhre opraHn3MbI) -

rrttTaIOTCH opraHnqecRnMM

Beru;eCTBaMn MepTBbIX TeJI (6aRTepnn, rpn6hI).
Ilapa3llTbl (napa3llTMqecKMe opraHH3MbI) -

IIHTaIOTCH opraHnqecKIIMll

Beru;eCTBaMn )KHBbIX opraHH3MOB (6oJie3HeTBOPHhie 6aKTepHM, rrapa3HTnqecKne pacTeHn.a, rptt6hI, IBHBOTHble).

3. Mun:compo<jJHbte opzanu3Mbl
HHH) -

(3BrJieHa 3eJieHa.a, HaceKOMO.H,D;Hhie pacTe-

MOI'YT IIMTaThCH KaK aBTOTpo<t>Ho, TaK ll reTepoTpo<J:>Ho.

Ilo cnoco6y nocTynJieHHH IlHTaTeJibHhIX Bern;ecTB opraHH3MhI
Ha

ZOJW30U1tble -

3aXBaTbIBaIOru;He rrnru;eBble qacTHI.l;hI M

pa3,n;eJIHIOT

ZOJLO<jJUmltble -

BCa-

CbIBaroru;tte pacTBopeHHble aeru;ecTBa.
Ilo OTHomeHHIO K KHCJIOpO,ll;y pa3JIMqaroT:

Aapo6nbte

opraHH3MhI (Ty6epKyJie3HaH rraJIO'IKa) -

,n;hIIIIaT Kttcnopo-

,n;oM, Heo6xo,n;HMhIM ,ll;JIH OKllCJIMTeJihHO-BOCCTaHOBHTeJihHbIX peaKD;llll
npH TKaHeBOM ,ll;bIXaHllH.

Anaapo6Hbte
OKMCJinTeJin.
H

cmpozue

opraHM3Mhl -

HCTIOJih3YIOT BMeCTO KHCJiopo,n;a ,n;pyrMe

0Hn .n;ennTca Ha

necmpozue

(,n;M<t>TepMiiHaH rranoq.Ka)

(B036y,n;nTeJIH CTOJI6HHKa, raHrpeHhI; noqBeHHhie 6aKTepMH).

06UfaH 6uo.1wzuH. Ocno6&i lfUmo.nozuu

50

TaKHM o6pa30M, rroTOK Bem;ecTB H 3Heprww o6ecneqwBaeT caMoo6ttoBJieHHe
H caMOBOCI1pOH3Be):(eHHe opraHH3MOB.
«l>oTOCllHTe3

<Pomocunme3 - 3TO rrpou,ecc npeo6pa3oBaHHH coJIHeqHorr 3HeprHH B rroTeHu,HaJihHYIO 3HeprHIO XHMHqecKHX CB.H3e:H: opraHHlfeCKHX se~eCTB. 8To TaIC>Ke:
rrpou;ecc o6pa3oBaHHH opraHMlfeCKHX sem;ecTB npw yqacTHH 3HeprHH cseTa. OH cBorrcTseHeH KJJeTKaM, co,D;epmam;HM cneu;waJihHhie cpoTOCHHTe3npyrom;ne nMrMeHThI, rJJaBHhIM H3 KOTOpbIX HBmieTC.H XJIOpocpMJIJI;
e):(MHCTBeHHhIM rrpou,ecc B 6noccpepe, Be,D;ym;nii: K 3anacaHnIO 3HeprnH 3a
cqeT ee BHernHero MCTOqHHKa;
3BeHO 6HoreoXHMHqecKHX ll;HKJIOB Ha 3eMJie M OCHOBa Bcex u;erreli IlHTaHMH. 3arraceHHaH B rrpo):(yKTax 3HeprH.H - OCHOBHOH HCTOqHnK 3Heprwn
)l;JIH qeJIOBe'IeCTBa.
EmerO)l;HO B pe3yJihTaTe cpOTOCHHTe3a Ha 3eMJie 06pa3yeTCH 150 MJIPA TOHH
opraHH'IeCKOro Bem;ecTBa (rrepsnqHaH rrpo)l;yKu;nH) M BhI):(eJIHeTCH 200 MJIPA
TOHH CB060):(HOro KHCJIOpO):(a. KpyrosopoT KHCJIOPO)l;a, yrJJepo):(a M )l;pyrHX
3JieMeHTOB, BOBJieKaeMhIX B cpOTOCHHTe3, ITO)l;):(epmHBaeT cospeMeHHhIH coCTaB aTMOCcpephI, He06XO):(HMhIH )l;Jl.H cym;eCTBOBaHH.H Ha 3eMJie COBpeMeHHhIX
cpopM mH3HH. <l>oTocnHTe3 rrpen.aTCTByeT yBeJinqeHHIO KOHu;eHTpau;un C0 2 ,
rrpe):(OTBpam;a.a rreperpeB 3eMJIH BCJie)l;CTBHe TaK Ha3hrnaeMoro « rrapHHKOBoro
3cpcpeKTa».
<l>oTOCHHTe3 B pacTeHHHX ocy~ecTBJI.HeTcH B xJioporrJiacTax. OH BKJIIOqaeT
,ZJ;Ba 3Tarra, IIOCJie,ll;OBaTeJibHO CBH3aHHhie Mem)l;y co6oii:
I. 8'rarr 110rJiom;eHnH H rrpeo6pa3oBamrn 3Heprwn ( ceemoeou npou,ecc ).
II. 8Tarr rrpeBpam;emrn Bem;ecTB ( me.Mttoeou npou,ecc).
,[(Ba rrpou;ecca cpOTOCHHTe3a BbipamaIOTC.H OT ):(eJihHhIMM o6o6~eHHhlMM
ypaBHeHHHMH.
I. 12H2 0 + 3Hepm.a cBeTa ~ 12H2 + 60 2 i + 3Heprn.a AT<l> (cBeTOBOH rrpou;ecc).
II. 6C02 + 12H2 + 3Heprn.a AT<l> ~ C6 H 12 0 6 + 60 2 i (TeMHOBoii: rrpou;ecc).
06o6m;eHHOe ypaBHeHne <PoTOCHHTe3a:
6C0 2

+ 6H 20

:JHeprHfl CBeTa

--x-JI-op_o_<jl_HJI_JI__,,.

C 6H

120 6

+ 60 2 i

Ceemoeou npou,ecc rrponcxo.zi;nT B muJta1CouiJax xJtoponJtacmoe, me.r.rnoeou - e cmpo;ne, To ecTh 3TH rrpou;ecchl rrpocTpaHCTBeHHO pa306m;eHh1.
IlJiaCTH.D;hI xapaKTepHhI TOJihKO .zi;JI.a pacTeHHM. 0Hn He Hali):(eHhI y rpu6oB
My IKHBOTHhIX, 3a HCKJIIOqeHHeM <fJoTOCMHTe3upyrom;Mx rrpocTeii:rnnx. B 3aBMCHMOCTH OT OKpaCKM, CB.H3aHHOH c HaJIHlfHeM HJIM OTCYTCTBMeM rrnrMeHTOB,
pa3JIHqaIOT TpH OCHOBHhIX THIIa IIJiaCTH)l;: XJtOponJtaCmbt (3eJieHoro u;BeTa),
xpo;nonJtacmbt (JKeJIToro, opaHJKeBoro, KpacHoro u;BeTa), JteuKonJtacmbt (6ecu;BeTHhre). 0)l;HH rrJiaCTH)l;hI MoryT rrpeBpam;aThCH B )l;pyrue. CTpoeHHe rrJiacTH,n;
MO/KHO paCCMOTpeTh Ha rrpHMepe XJIOPOIIJiaCTOB.
XJtoponJtacmbt - 3TO )l;ByMeM6paHHhie opraHOM)l;hI, BHyTpeHHHH MeM6paHa
KOTOpbIX 06pa3yeT muJtalCOUObl, qiopMnpyrom;ue epattbl, Mem,n;y KOTOpbIMM Ha-

06MeH. 6€Ufecm6 u npe6paUfenue :Jnepzuu. <Pomocunme3

51

xo,LIMTCJI cmpwna. B MeM6pattax TIVJaKOI1,LIOB Haxo,L\aTea rr11r:vieHThI (x:iopoqmJIJI, KapOTI1HOI1,L\hl 11 .11,p.). B CTpOMe co.n,epiKaTeJI cpepMeHThl H p1160COMhl. retteTWieeKHH arrrrapaT rrpe.n,eTaBJICH UHKJIHqeeKHMH MOJieKyJiaMH .II:HK. 3To
UOJiyaBTOHOMHhle opratteJIJlhl, erroeo6Hhie eIIHTe3IIpOBaTb co6eTBCHHhie 6CJ1KOBhle MoJieKyJihI. OcttoBttaa cPYHKI~Ha xJioporrJiaeToB - cPOTOeIIHTe3, a TaKJKe
e11ttTe3 AT<t> II A.II:<P (c1Joe¢op11JI11posattIIe), eHHTe3 11 rII.n,poJIH3 JIHIIIIJ(OB, aeeIIMHJIJIUHOHHoro KpaxMaJra II 6eJIKOB, OTKJia,D;bIBaIO~Hxea B CTpo:vie.
B oenoBe eTpoeHIIJI XJIOPOcPHJIJIOB JieJKHT Mg-nop<jJupu1w6btu CKeJtem. Ero
eoeTaBJrnIOT rrop¢11pnttosa.a «rOJIOBKa» e aTOMaMH Mg B uettTpe (rII,11,poclJ11J1bHa, JieJ'KHT Ha IIOBepxHoeTH MCM6paHbl) II cPIITOJihHhIH «XBOeT» (rIIJJ;poq>o6HhIH,
y,n;epJKHBaCT MOJieKyJiy XJIOpocpHJIJ1a B MeM6paHe).
Cy~ecTByeT HeeKOJibKO MO.L\HcPHRau11i1 xJiop01p11JIJIOB (a, b, e, d), OTJmqa10rn;nxca erreKTpaMH rrorJio~eHHJI. Bee paeTeHHJI B KaqeeTBe oeHOBHOI'O IIHrMeHTa eo,n;epJKaT eHHe-3eJieHhIH XJlOpO<jJUJlJl a, a B KaqeeTBe ,11,0IIOJIHIITeJihHhIX 3eJieHO-J'KeJIThlH XJlOpO<jJUJlJl b.
B xJioporrJiaeTax xJiopocpnJIJI n ,n;pyrne rrnrMeHThI, rrorpyJKeHHhie B TmrnKoH.L\hI, eo6paHhI B <l>YHKUHOHaJihHbie e,n;HHHUhl (rro 250-400 MOJieKyJI), Ha3hIBaeMhie <jJomocucmeM.a.Mu.
HMe10Tea ,n;sa Tnrra cpoToe11eTeM: <PC-I n <PC-II. Kam.n,a.a clJoToencTeMa eo,n;epmnT 250-400 MOJieKyJI rrnrMeHTOB. Bee n11rMeHThI clJoToeneTeMhI MoryT
rrorno~aTb qaeTHUhI eBeTOBOH aHeprnH (cPOTOHhl, KBaHThI eBeTa), HO TOJlhKO
0,11,Ha MOJieKyJia XJiopocpnJIJia aTOH eneTCMhI MOJ'KeT HeIIOJlh30BaTh rrorJio~eH
HYIO aHeprHIO B q>OTOXHMnqeeKHX peaKUHJIX. 8Ta MOJieKyJia Ha3hIBaeTeJI peaKU,UOliliblM. u,eHmpoM., a ,n;pyrne MOJieKyJihl IIHrMeHTOB Ha3bIBaIOTC.H aHmeHllblMU, TaK KaK yJiaBJIHBaIOT aHeprnIO eBeTa, IIO,LI06HO aJITeHHaM, ,D;JIH IIOeJ1C,n;y10~ei1 rrepe,n;aqn pea1<unoHHOMY ueHTpy.
B <PC-I pea1<unoHHhiii ueHTP o6pa3oBaH MoJieKyJioii xJiopocpnJIJia a, o6o3HaqaeMoi1 P 700 , r,L1e 700 - onTnMyM rrorJI01uettna B HM. B <PC-II peaKUHOHHhIH
ueHTP TaKme o6pa3oBaH MOJieKyJIOM XJIOPOcPHJIJia a, o603HaqaeMOH p 680' r,n;e
orrTHMyM norJio~ettna JremnT B pai1oHe 680 HM. <t>oToeneTeMhI I II II pa6oTaIOT
eHHXPOHHO ll HerrpephIBHO, HO <PC-I MOJ'KeT <l>YHKUHOHllpOBaTh OT,D;eJihHO.
CpaBHHTeJILHaJJ xapaKTepncTnKa pa60Tb1 ,11;Byx l}>oTocncTeM npu l}>oTocuHTeae
<l>OTO~OC~OpHJillpOBaHue

He11uKJmtiecKoe

ll;ttKJllltJCCKOe

°YqaCTBYIO~tte $OTOCHCTeMhl

<PoTocucTeMhI I, II

<PoTocucTeMa I

IlyTb :rneKTpOHOB

HeIIttKJitttiecKuii

littKJilltJeCKllH

IlePBhlll ,D;OHOP aJieKTpOHOB

Bo,n;a
HA,U<P.

<PoTocucTeMa I

IlocJie,n;ttuii aK11enTop
Ilpo,n;yKThl peaKuuu

AT<P, HA,U<PH, KllCJIOPO,L\

AT<P

--

--

<PoTocncTeMa I

I. CeemoBaR <jJaaa rrpoTeKaeT Ha eBeTy Ha MeM6paHax THJiaKOH,L\OB npH yqaCTHH XJiopocpmrna, 6eJIKOB - rrepeHoeqHKOB aJreKTPOHOB n clJepMeHTa - AT<PCllHTeTa3hI (pne. 18).

06Ufan 6uo1wzu11,.

52

Oc1W6bt

1fUmo1wzuu

¢OTOHbl

csera

Cl>oTO~~~:.e,Ma II

.........

:.:~((.::=r:·: Q)c~~%b1
.:-~.· -~: ......
PMc. 18. Caernaal'I cj:>a3a cj:>orno1Hrn3a
BKmoqaeT cJie.n;yrow.ne npou;ecchr: ueu,u1ow'lec1we <jJomo<jJoc<jJopuJLupoea·
Hue ( Z-cxe.ina ); u,uKJLU'l.ec1we <jJomo<jJoc<jJopuJLupoBaHue; <jJomo_,w3 BOObl.
Bee MOJieKyJihl IIHrMeHTOB yJiaBJIHBaIOT ::meprHIO COJIHelJHOro CBeTa H nepe.n;aIOT ee Ha MOJieKyJihl peaKIJ;HOHHOro u;eHTpa p680' IIPH 3TOM 2e 11epeXO,ll,.HT Ha
6oJiee BblCOKHH 3HepreTH'leCKHH ypoBeHh H yJiaBJIHBaIOTC.H aKu;eIITOPOM

Q.

OT aKu;enTopa Ha'IHHaeTC.H aJieKTpOHHhIH IIOTOK, B KOTopoM aJieKTpOHhI «C11ycKaIOTC.H » 110 aJieKTpo1rno-TpaHc11opTttoii u;e11n K cpoTocncTeMe
3TOH u;e11H HOJibHhie

I.

KoMIIOHeHThI

IJ;HTOXpOMbl (6eJIKli, co.n;epmaw.He me.'le30 H cepy), XHHOHbl (cpe-

coe.n;irneHH.H)

IlpH 3TOM IIPOHCXO,ll,HT

n

rrnacTou;naHHH

(6e.'IoK,

<jJomo<jJoc<jJopuJlUp06aHue,

TaTKa cpoccpoptto:H KHCJIOThr MOJieKy JI hr

co.n;epmaw.nH:

Me.n;h).

TO eCTb CHHTe3 H3 A.IJ:Cl> Hoc-

AT<I>.

MoJieKyJia p680• IIOTep.HBIIIa.H CBOH 3JieKTPOHbI, BOCCTaHaBJIHBaeTC.H 3a
c'leT aJieKTpottoB .n;oHopa, KOTOphrM .HBJ'I.HeTC.H Bo.n;a. Ilponcxo.n;HT

JLU3 -

<Pomo-

CBeT03aBHCHMOe pacw.e11JieHne BO,ll,bI. Cl>epMeHTbI, OTBeTCTBeHHhre aa

cpoTOJIH3 BO,ll,hl, pacnoJiaraIOTC.H Ha BHyTpeHHeii CTOpOHe MeM6paHhI THJiaKOH,ll,OB. Cl>OTOJIH3 BO,ll,hI yqacTByeT B C03,D,aHHH rpa.n;neHTa IIpOTOHOB qepe3
MeM6paHy' r.n;e BhICOKa.H KOHIJ;eHTpau;n.H IIPOTOHOB OKa3hIBaeTC.H BO BHYTpeHHeM IIpOCTpaHCTBe THJiaKOH,ll,OB. IlpOTOHHbIM rpa.n;HeHT crroco6cTByeT
CHHTeay

AT<I>.

B <PoTOCHCTeMe
IJ;HOHHhrii u;eHTP

I

atteprH.H cBeTa OT aHTeHHhIX 11nrMeHTOB 11ocTyrraeT B peaK-

P 700 .

Boa6ym.n;eHHhre aJieKTPOHhl nepe.n;aroTC.H Ha :rneKTpoH-

06;n,en Beiq,ecmB u npe6paiq,enue ;mepzuu. <l>omocunme3

53

HbIH aK11,enTOp P 430 (6eJIOK, co.n;epmaI11,11ii: JKeJie3o 11 cepy). P 4a0 nepe11awr cB011
:3JieKTpOHbI Ha .n;pyroil: JKeJie30Co.n;epJKaI11,MH 6eJioK - cpeppe.n;OKCHH, a IIOCJieJJ;HMM Ha KocpepMeHT HA,ll.<t> , KOTOpbIM BOCcTaHaBmrnaeTc.H /IO HA.D:<t>H 2 . MoJieKyJia P700 B xo.n;e rrpo11,ecca OKHCJI.HeTC.H, HO 3aTeM BOCCTaHaBJIHBaeT IIOTep.HHHbie :3JieKTpOHbl 3a cqeT :3JieKTpOHOB, IIOCTynaIOill,MX TIO :3JieKTPOHHO-TpaHcnopTHOM 11,enu OT <PC-II.
0,n;HOHarrpaBJieHHhIM IIOTOK Ha3hIBaeTC.H fteU,UICJlU'leC/CUM nomo/CO.llt 3Jle/Cmpo1t06, a o6pa3oBaHtte AT<t> - 1teu,u1CJtU'leC1CUM <Pomo</_Joc<PopuJtupoea1tueM.
<t>C-I MOJKeT pa6oTaTb He3aBMCMMO OT <t>C-II. 3TOT rrpo11,ecc Ha3hIBaIOT 11,l:IKJIRqecKRM IIOTOKOM 3JieKTpOHOB. B xo.n;e Hero He rrpoMCXO,ll;MT cp0TOJIM3 BO,ll;hl,
BbI,n;eJieHRe 0 2 M o6pa3oBamrn HA,ll.<t>H 2, o.n;HaKo 06pa3yeTC.H AT<t>. TaK KaK
cnHTe3 AT<t> CB.H3aH c U,UICJlU'leCICU.M nomo/Co.M 3Jte1Cmponoe, ero Ha3hIBaIOT
u,u1CJtU'leC1CU.M <Pomo<Poc<PopuJtupoea1tue.M. Y ayKapttoTuqecKttx KJieToK oH
ocyill,eCTBJI.HeTC.H pe,n;Ko, o,n;HaKO y cpOTOCMHTe3I:IpyIOI11,l:IX 6aKTepttii: JieJKMT
B ocHOBe rrpo11,eccoB aHaapo6Horo cpoTOCMHTe3a.
TaKttM o6pa3oM, 1Co1te'l1tbt.Mu coeaune1tuJl.Mu ceemoeou <Pa3bt .HBJI.HIOTc.H:
AT<t>, BoccTaHOBJieHHhIM HA,ll.<t>H2' 0 2•
CBo6o,n;HhIM KMCJiopo.n; rrocTyrraeT B aTMoccpepy, AT<t> M HA,ll.<t>H 2 TpaHcnop™PYIOTC.H B CTPOMY XJIOpOIIJiaCTa M yqacTBYIOT B npo11,eccax TeMHOBOM cpa3hl.
IL Te.M1toea.R <Pa3a rrpoTeKaeT B cTpoMe xJioponJiacTa n MOJKeT nponcxo,n;HTh
He TOJibKO Ha cBeTy, HO u B TeMHOTe. XnMnqecKa.H aHeprn.H (B Btt.n;e AT<t>), 3anaceHHa.H B xo.n;e CBeTOBOM peaKll,HR, HCIIOJih3yeTC.H JJ;JIP, BOCCTaHOBJieHH.H yrJiepo.n;a. PeaK11,nn TeMHOBo:H cpa3hI rrpe,n;cTaBJI.HIOT co6oii: 11,erroqKy rrocJie.n;oBaTeJibHhIX npeo6pa30BaHMH yrJieKMCJioro ra3a, IIPHB0,11;.Hill,YIO K o6pa30BaHHIO rJIIOK03bl H ,11;pyrHX opraHttqeCKMX BeI11,eCTB.
B cocTaBe xJioporrnacToB HMeeTc.H cpepMeHT (pH6yJio3o-l,5-6ucpoccpaTKap6oKcttJia3a), KOTOphIH KaTaJIH3HpyeT coe.n;I:IHeHtte MOJieKyJihI yrJieKHCJIOro ra3a c rrpott3BO,ll;HbIM II.HTHyrJiepo.n;Horo caxapa - pu6o3hI c pu6yJto.3o6u<Poc<Pamo.M (PB<P). 0.n;Ha MOJieKyJia PB<t> coe,11;HH.HeTc.H c MOJieKyJio:H C0 2
c o6pa3oBamrnM wecmuyiJtepoa1tou .MoJte/CyJtbt [C 6 ], KOTopaa HerrpoqHa
H 6hlcTpo pacrra,11;aeTc.H Ha ,11;Be TpexyrJiepo,11;Hhie MoJieKyJihI <Poc<PoiJtu4epu1weou ICUCJlOmbt, <PI'K (C3)· 3aTeM rrpOHCXO.D;MT U.HKJI peaKll,MM, B KOTOpbIX
ttepe3 pa,11; npoMeJKyToqHhIX rrpo.n;yKTOB cpoccporJIH11,epI:IHOBa.H KHCJIOTa npeo6pa3yeTC.H B ,11;Be MOJieKyJihI <Poc<PoiJtuu,epoaJtbaeiuaa (<PI'A), 3aTeM o,11;Ha
MOJieKyJia <t>fA tt,11;eT Ha pereHepa11,n10 PB<t>, a BTopaa Ha o6pa3oBaHtte
l'JIIOK03hl. B :3THX peaK11,H.Hx MCIIOJih3YIOTCH 3HeprmI AT<t> H HA.D:<t>H, o6pa30BaHHhIX B CBeTOBOH cpa3e. lJ;MKJI 3TRX peaKll,MH IIOJiyqHJI Ha3BaHne «11,MKJI
KaJibBHHa» (pttc. 19).
CyMMapttoe ypaBHeHtte CHHTe3a l'JIIOK03hI:
6C0 2 + 12HA,D;H + 18AT<t> -i- C 6 H 12 0 6

+ 12HA,ll.+ + 18A,ll.<t> + 18<t> + 6H20

KpoMe rJIIOK03hI, B rrpo11,ecce cpoToCHHTe3a H3 <t>fK o6pa3yIOTC.H .n;pyrne MOllOMephI CJIOJKHhIX opraHnqeCKHX COe.D;HHeHHH - aMHHOKHCJIOTbI, rJIH1J,epHH,
'KHPHhle KBCJIOTbl, HyKJieOTM,ll;bl.

06UfaH 6uo;wzun. OcnoBbt lfUmoaozuu

54

HecTa6ttJihHhI:
rrpoMemyToqHhiii rrpo,n;yKT

co

P:u6yJio3o-l,56mpocct>aT (PB<I>)

<Pocct>orJim:i;ep:uHoBaR
KllCJIOTa (<l>I'R) 2C3

pereHepau;11JI
2AT<I>
42AT<I>

_ ___, <I>ocq>0rJI11u;epoaJib,n;ertt,n; (<I>I'A) 2C3

I'JIIOK03a

+ 2<I>

r

2HA,ll;<I>H2
'-+2HA,l:J;<fl+

~-___,<flpyKT03a

Pitc. 19.

l.J,111KJ1 KanbBlllHa

<l>aKTOpbl, BJIIUIIOITI;He Ha cpoTOCHHTe3

Ceem -

cKopocTh cpoTOCHHTe3a npHMO nponopu;noHaJihHa HHTeHCHBHOCTH

CBeTa; npH ITOBbiilleHHH HHTeHCHBHOCTH CKOpOCTb cpOTOCHHTe3a 3aMe,n;JIHeTCH,
TaK KaK pa3pyrnaroTCH MOJieKyJihI XJiopocpHJIJia.

BpeMR:

qeM 6oJibrne nepno,n;

OCBern;eHHOCTH, TeM ,ZJ;OJibIITe npo,n;oJimaeTCH cpOTOCHHTe3.

Ko11u,e11mpau,uR yzJLelCUCJlOZO za3a

0,03-0,04 % );

npn KOHD;eHTpau;HH

(B HOpMe B aTMoccpepe co,n;epmHTCH

C02 ,n;o 0,1 °/c,

ITOBbIIITaeTCH CKOpOCTb TeM-

HOBhIX peaKD;HH cpoTOCHHTe3a.

BofJa -

cy6cTpaT cpoTOCHHTe3a; npn yMeHhIIIeHHH co,n;epmairnH BO.D;hI 3a-

C0 2 He MomeT ,n;ncpcpyH,n;npoBaTb.
onTttMaJibHaH TeMnepaTypa ,n;JIH cpoToCHHTe3a

KphIBaroTCH ycThHD;a, H

Te.Mnepamypa -

+25 °C

(,n;JIH

pacTeHHH cpe,n;Heii ITOJIOCbI); CHHmeHne npHBO,ll;HT K 3aMe,n;JieHHIO pa60Tbl cpepMeHTOB.
,UocTaToqHaH

ICOHU,enmpau,uR XJlOpO<jJUJlJlU.

Heo6xo,n;HMo ,n;ocTaTOqHoe MHHepaJihHOe nnTaHne

Koc.Mu1leC1CaR pOJlb pacme11uu
CH B TOM, qTo pacTeHHH -

(onncaHa

K. A.

(Mn,

Fe,

Mg,

NH ,n;p.).

THMttpH3eBhIM) 3aKJIJOqaeT-

e,n;HHCTBeHHbie opraHH3Mhl, ycBaHBarorn;ne COJIHeq-

HYIO ::rneprHIO H aKKYMYJIHpyrorn;ne ee B Btt,n;e ITOTeHu;HaJihHOH XHMfiqecKOH

0 2 no,n;,n;epmnBaeT mtt3lfa KHCJiopo,n;a o6paayeTCH 030H,

::rneprHH opraHHqecKHX coe,n;nHeHHH. Bhl,n;eJIHrorn;niicH
He,n;eHTeJihHOCTh Bcex aapo6HhlX opraHH3MOB.

KOTOphlii aarn;Hrn;aeT Bee mHBoe OT YJihTpacpHoJieTOBhIX Jiyqe:H. PacTeHHH nc-

C0 2, tt36b1ToK KOToporo co3,ZJ;aBaJI «napHHKOBhlH acpcpeKT»' H TeMnepaTypa nJiaHeThl ITOHH3HJiaCh ,n;o Hhl-

ITOJih30BaJIH H3 aTMoccpepbI rpoMa,n;Hoe KOJIHqecTBO
HeIIIHHX 3HaqeHHH.

06Men 6e14ecm6 u npe6pa14enue ;mepzuu. XeMocunme3

55

XeMOCHHTe3
KpoMe cpoTOCHHTe3a, cyw,ecTByeT ew,e o.n;Ha cj:>opMa aBTOTpoctmoil accn::vmJI.HUfllI - xeMOCMHTe3, tta6;no,uaeMhIH y HeKOTophIX 6aKTepvril. TaK11e 6aKTepttH Ha3bIBaIOTCJI XP.MOCUltmemuKa.Mu.
XeJnocuHme.:1 - 3TO cnoco6ttocTh xeMocnHTeTHKOB np11 OKl1CJIPH11n neopraH11qecKHX Beu~eCTB 11CilOJih30BaTb BbI,UeJI11BlIIYIOCJI 3Hepr11IO ,UJIH o6pa30BaHHfl
opram1qeCKl1X Bew,eCTB, 11.UYID;l1X Ha IIOCTpoeHne IGieTOK CBOero TCJia. XeMoCHHTeTlIKH, B OT.111'-Ilie OT cpOTOaBTOTpocpoB, 11CilOJih3YIOT B npouecce ClIHTe3a
He COJIHeqHbIH CBeT, a 3Hepr11IO 3K30TepMnqecKIIX peaKU11M.
PyccKIIH MHKpo6IIoJior C. II. Bnttorpa.n;cK11ii B 1889 r. BnepBhie .n;oKa3a.1
cyru;ecTBOBaH11e B rrpIIpo.n;e aBTOTpOcpHbIX He3eJieHhlX opraH113MOB (Hl1Tp11cp11UHPYIOW,IIX 6aKTep11i1) n on11caJI cyru;HOCTh rrpouecca xeMOCIIHTe3a.
1. Humpu</Juu,upy10w,ue 6a1tmepuu. PacnpocTpaJieHhI B no<rne n n pa3JI11qHhIX
BO,UOeMax . .D:o6hIBaJOT 3HeprIIIO nyTeM OKIICJieHII.H aMMIIaKa 11 a30T11CTOH Kl1CJI0Thl, 11rparoT namuyro poJih B KpyrosopoTe a3oTa B npIIpo.n;e. Bh1.n;eJI.HIOT Bo
BHeIIIHIOJO cpe.n;y cpepMeHThl, KOTOpbre KaTaJIII3IIPYIOT OKIICJieH11e cy6cTpaTOB:
- aMM11aK (cy6cTpaT) OKIICJlaeTCJI .n;o a30TIICTOH KHCJ10ThI 11 3Hepnrn:
2NH 3 + 30 2 ~ 2HN0 2 + 2H 20
-

+ E;

a30THCTaJI Kl1CJIOTa OKIICJiaeTCJI .n;o a30THOH KHCJIOTbl II 3Hepr1111:
2HN0 2 + 0 2 H

2HN0 3 +

E.

2. Becu,Bemttble cepo6aKmepuu. PacnpocTpatteHhI B Bo.n;oeMax, Bo,ua KOTOpb1x co.n;epmnT cepoBo.n;opo.n; . .D:o6hrnaroT atteprn10 .UJI.H CIIHTeaa oprattnqecKHX coe.n;IIHeHIIH 113 yrJieKIICJIOro raaa, OKIICJIJl.H cepOBO.UOPO.U.
2H 28

+ 0 2~

2H 20

+

28

+E

IlpII ue.n;ocTaTKe cepoBo,uopo.n;a cepo6aKTepII11 rrpOII3BO.U.HT .n;aJihtteiiIIIee OKIICJieHHe CB060.n;HOI1 cepbl .n;o cepHOH KIICJIOThl.
28

+

30 2 + 2H 20 ~ 2H 2801 +

E

3. JKeJte306a1emepuu. 06paay10T 6oJIOTHYIO meJieattyro PY.UY.
4Fe2·

+ 0 2 + 4H'

~ 4Fe3•

+ 2H 20 + E

4. BoiJopoiJHbte 6a1emepuu. PacrrpocTpatteHhr B rroqBe. OK11CJI.HIOT BO,uopo.n;,
06paay10ru;HMC.H rrpII attaapo6HOM pa3JIO)KeHHII opraHWieCKIIX OCTaTKOB
M11KpoopraHII3MaMH IlO'lBhl.
2H 2 + 0 2 ~ 2H 20

+E

3nepreTH'leCKHii o6MeH
3ttepiemu'leCICUU 06.MeH ( icama60JlU3.M, UJlU i}uccU.MUJlJlU,UJ/.) 3TO COBOKYilHOCTb peaKuIIil cpepMeHTaTHBHOro pacru;errJieHu.H opraHnqecKux Bew,ecTB,

06~all

56

6uo1wiull.

Oc1W6bl

1tumo1wiuu

conpoBom.n:a10:w;11xcH Bhrp;eJieHHeM ::rneprttu. Kone'-IHhIMH npo.n:yKTaMH pacna.n:a HBJUIIOTCH co~, H20, ATcD. 3Hepl'HH, Bhr.n;eJIHIOIIJ;aflCH IIpH paapyrueHHH XllMHqecKHX cBH3eH:, 3anacaeTcH B MaKpoapruqeCKHX CBH3HX MOJieKyJ1 ATcD
" .n;pyrwx BhICOK03HepreTttqecKHx coe.n;HHeHHtt. ATcD - yuwBepcaJihHhrtt ucToqHHK aHeproo6ecneqeHHH KJieTKH. CttHTe3 A Tell nponcxo.n;HT B KJieTKax Bcex
opraHH3MOB B npouecce ¢ocipopuJzupo6a1tuR - npncoe.n;ttHeHr-ur HeopraHttqecKoro cpoccparn K AJJ:cD.
IlpeBpa:w;eHHe Be:ru;eCTB l1 3HeprHH B npouecce .ZJ:HCCHMHJIHUHH BKJIIOqaeT
B cefoc cJie,n;y10:w;11e 3Tarrhr:
I aTan - no11.roTOBHTeJ1bHhIH. CJIOJRHhie opram1qecKtte Be:w;ecTBa no.n:
,n;ettCTBHeM cpepMeHTOB pacna,n;aIOTCH Ha npoCTbie (MOHOMepb1): 6eJl}(U iJo
aMUIWKUCJlom; JICUpbt 00 ?Jluu,epul-la u JICUpllblX }(UCJlOm; yiJLeBOiJbt (KpaxMQJl) oo .M011ocaxapuoo6 ( ?JllOK03bt ); nyKJteunoBbte KUCJtontbt iJo 1-lYKJLeomuiJoB Um. 0.
Ilp11 3TOM BhI,!l;em1eTCH 3Heprl1H B BH,n;e TeIIJia.
II aTarr - rJ1HKOJ1H3 (6eCKHCJiopo,n;HhIH, aHaapo6Hbrtt) - MHorocTyrreHqaThitt <lJepMeHTaTHBHhIH npouecc pac:w;enJieHmI I'JIIOK03hI ,n;o .ZJ:BYX MOJieKyJI IIHpOBHHOrpa,n;HOH KHCJIOThI (IIBK, IIHpyBaT), ,n;Byx MOJieKyJI ATcD 11 BOCCTaHOBJieHHOH <lJopMhI nepettocq11Ka Bo.n;opo,n;a HA.UH, c o6pa3oBantteM npoMeJRyToqHhIX rrpo,n;yKTOB. Ilpouecc ocy:w;eCTBJIHeTCH B UHTOIIJia3Me 6e3 yqacTIUI
KllCJIOpo,n;a, c MeM6paHaM11 He CBJI3aH.

ITBK

I' ..'1101{0:13

Ilp11 3TOM 40 'Xi ::rneprtttt aanacaeTcH B Bn,n;e ATcD, a 60 'Y,, 3Heprnn paccenBaeTcH B BH,n;e TenJia.
III. llpu omcymcmBuu KUCJLOpooa:
y ,n;pommeH: 11 pacTeHHH rrpo11cxo,n;HT CIIl1pTOBOe 6pomeHHe, nptt KOTOPOM CHaqaJia o6pa3yeTCH yKCyCHhIH aJib,!l;erH,n;, a 3aTeM 3Tl1JIOBhIH CIIHpT:

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B peayJihTaTe rJI11KoJIH3a o,n;HOH MOJieKyJihI rJIIOK03hI BhICBo6om,n;aeTcR

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06Men 6PUfeCm6 u npe11paUfenue anepzuu. :J11epzemu'leCK1tii o6Men

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c o6pa30Ban11eM

36

MOJICKYJI AT<f>: .n;Be Mo.'IeKyJlhI o6pa3yIOTCR B pemni;ttRx

u;irn.11a Kpe6ca (B MaTpttKce MttTOXOH.ll,pttil) H

34

:vro.'IeKyJihI -

B rrpouecce

OKHC.'lHTCo'IhHOrO cpoccpopH.'"!llpOBaHttR (Ha BHYTPCHHett MeM6paue MHTOXOH.ll,·
pn:ii,

B KOTopyIO BcTpoeui,1 cpepMeHThI .ll,bIXaTeJihHOll ILerrtt). Ha6.'lIOJJ;aeTcH

U,UKJl mpuKap601-w6blX Kuc.1wm ( u,uKJL Kpe6ca ); 3aKJIIOq11TeJibHYIO cTa.n;n:IO OKHCJieHna - ;J.'teKmpo11Ho-mpancnopm1ty10 u,enb.
TOJlbKO y aspo6Hb!X opraHH3MOB. 8TOT npouecc BR.JIJOqaeT:

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BhICOKosuepreTttqecKoil CBH3hIO. 8To coe.n;n:Hentte BOBJieKaeTca B umrn Kpe6ca uenh nocJie,1i;oBaTeJihHhIX peaKUHll. AueTHJI-KoA coe,n;ttHHeTca c MOJieKyJioil w;aBeJieBoyKcycnoil KMCJIOThl (mYK), rrpu 3TOM 06pa3yeTCH TPMKap6oHOBaH JIHMOHHaH KHCJIOTa. B xo,n;e IIOCJie.n;yJOw;MX cpepMeHTaTMBHhIX peaKUHll JIHMOHHaH
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3

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ttcnoJih3yeTca ,n;JIH cpoccpopmmpoBaHn:H A,U<f> H o6pa30BfilIHH AT<f>. JlttMOHHaH
KHCJIOTa TCpJJ.eT .n;Ba yrJiepO.ll,HhIX aTOMa, 3a cqeT KOTOpb!X o6pa3YJOTCH .n;Be MOJICKYJlhl C0 2 • B cyMMe, B pe3yJihTaTe ceMM nocJie.n;oBaTeJihHhIX cpepMeHTaTHBHhIX peaKUHll, JIMMOHHaH KHCJIOTa rrpeBpaw;aeTCH B mYK. 06pa30BaBlllaRCH MOJICKyJia
mYK COC.ll,HHHCTCH C HOBOM MO.'lCKy.1oll aI~eTHJI-KoA, H UHKJI IIOBTOpHCTCH.
B COCTaBe JIHMOHHOM KHCJIOThl croplleT rrpHCOC.ll,MHMBlllHMCH OCTaTOK aueTHJIKoA. IlpM 3TOM o6pa3yIOTCH yrJieKHCJIL!H ra3, aTOMhI Bop;opo.n;a; sJieKTPOHhI rrepeHOCHTCH Ha aKUCIITOphI

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mpa1tcMe.w6paHHbtu nome1tu,uaJL. B MeM6paHy BCTpoeH cpepAT<P-cu1tmema3a. Bero MOJieKyJie OTKphrnaeTcH KaHaJI, qepe3 KOTOphIM

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200 MB.

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Ha BOCCTaHOBJICHHC Q2-), H COC.ll,HHHJOTCH c rrpOTOHaMH.

0614aR 6uo.rwzuR.

58

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06lll;ee IWJII1'-!8CTBO :meprttM, BhI,n;eJrnIOlll;8HCH Ha 8TOM 8Tane, ---

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AHa60JI1P1eCKMH rrpou:ecc, B

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pe3yJihTaTe KOTOporo H3 rrpoCTbIX

pe3yJibTaTe KOToporo l\IOJICKYJlbl

HeopraHM'IeCKHX coe,n;HHeHHM

yrJieBO,ll;OB pacru:errJIHIOTCH

CHHTe3HpyIOTCJl MOJieKy Jibl

npOCTb!X HeopraHH'leCKHX

yrJieBO,Zl;OB

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110

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IlpOHCXO,ll;HT yMeHb111e1rne

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opraHH'leCKOH MaCCbI

Y syKap110T rrpou:ecc rrpoTeKaeT B

Y syKapttoT rrpou:ecc rrpoTeKaeT B

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IlpoHCXO;IHT TOJibKO B KJieTKax,

IlpOHCXO,ll;HT BO BCex KJieTKax B

co,n;epmaru;ttx XJiopocpHJIJI, Ha cseTy

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J

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KJLemo'lllbtit u,uKJL -

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Ta ee IIOHBJI8Hl1H .n;o TIOCJie,n;yIOlll;ero ,n;eJieHtt.H HJil1 CMepni:. ll;:ttKJI ttMeeT pa3HYIO rrpo,n;OJI:JKl1T8JlhHOCTb B 3aBHCHMOCTl1 OT TKaHeBOM rrp:ttna,n;Jie:>KHOCTl1
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( uHmep<jJa3y) H co6cTBeHHO ,n;eJieHHe - Mumoa ( cpe,n;HHH rrpo.n;omK11TeJih-

penenue x:nemx:u. Hnmep</Ja3a. Mumo3

59

IIHTepcI>aaa
H ttmep<jJa3a - rrep1rn,n; Jini::me,n;eJITeJibHOCTH KJienrn Mem.ny .D;C.'IemrnM 11 .
3aHHMaeT 90 <;(, BCero BpeMeHH KJieTO'-IHOro IJ;HKJia.
IIepttO.D;hl HHTepcpa3bI:

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OH BKmoqaeT: CHHTe3 6eJIKOB, cpepMeHTOB, tteo6xo,n:ttMhIX .D;JIH o6pa 30 _
BaHHH PHK, H Beru;ecTB, Bxo,n;Hru;ttx B cocTaB .D:HK; CHHTe3 PHK; o6pa30BaHtte ptt6ocoM; CHHTe3 AT<P; o6pa3oBatttte opraHeJIJI - MHTOXOH,n;pttii, XJIOponJiaCTOB (y pacTeHHM), 8IIC, JIH30COM, annapaTa fOJib,[\JKl1,
BaKyoJieii, IlY3blpbKOB; pocT KJieTKH ,n;o «KPHTH:qecKOM MaCCbI ll;HTOIlJia3MbI»; o6pa30BaHtte Beru;eCTB, no,n;aBJIHIOIIl;HX HJIH CTHMYJIH:py10ru;ttx HaqaJio cJie,n;y10ru;eii: cpa3hI.
XpoMOCOMa rrpecttHTeTHqecKoro rreptto,n;a - oottoxpoMamuiJtta.R, c retteTttqecKHM Ha6opoM 2n2c, r,n;e n - qHCJIO xpOMOCOM, c - KOJIHqecTBO .D:HK.
2. Cuttmemu'leCKUil, uJtu S-nepuoiJ, B KOTopoM npottcxo,n;ttT perrJIHKau;ttH
,n:HK IIOJIYKOHCepBaTHBHbIM cnoco6oM (CM. pa3,n;eJI «feHeTHKa» ).
B HaqaJie rreptto,n;a xpoMoCOMhl - oiJttoxpoMamuiJttbte (o6pa3oBaRbI o,n;ROM xpoMaTtt,n;oii, 2n2c). IIocJie perrJittKau;tttt .D:HK - iJByxpoMamuiJHbte (cOCTOJIT H3 .D;BYX O,[\HHaKOBhIX xpoMaTH,[\, 2n4c).
CttHTe3 6eJIKOB (rttCTOHOB), yqacTBy10ru;ttx B cpopMttpoBaRHH CTPYKTYpbI xpoMOCOM.
3. Ilocmcuttmemu'leCKuit (npeiJ.Mumomu'leCKuit), HJIH G2 -nepuoiJ, B KOTOpoM rrpOHCXO,[\HT rro,n;rOTOBKa KJieTKH K ,n;eJieHHlO.
BKJIIOqaeT B ce6JI: CHHTe3 6eJIKa; o6pa3oBatttte MHTOTttqecKoro anrrapaTa; CHRTe3 AT<P; CHHTe3 PHK; CHHTe3 6eJIKOB, orrpe,n;eJIJIIOru;ttx ,n;eJieHtte KJieTKH; yBeJittqeHHe MaCCbI IJ;HTOIIJia3MhI; pe3KOe B03pacTaHtte
o6'heMa H,n;pa.
XpoMocoMa - ,n;ByxpoMaTtt,n;HaJI, c reHeTttqecKHM tta6opoM 2n4c.

MHT03
M umo3 - aTo tterrpHMOe ,n;eJieHtte KJieTKH, rrptt KOTopoM npottcxo,n;ttT paBRoMepaoe pacrrpe,n;eJieatte ttacJie,n;CTBeHHoro MaTepttana Me:>K,n;y ,n;oqepHHMH KJieTKaMH
(pRc. 201.
Oco6ettttocmu .Mumo3a: ocHOBHOM crroco6 ,n;eJiemrn ayKapttoTHqecKoii: KJieT1m; BepeTeRo ,n;eJieRttH 06pa3yeTcH; y,n;Boett11e tt paBHOMepttoe pacrrpe,n;eJiemrn
reaeTHqecKoro MaTep11ana; paBHOMepttoe ,n;eJieHHe u;RTOIIJia3MhI.
BuoJtoZu'lecKoe 3Ha'lettue .Mumoaa - o6ecrreqeHHe Toqttoii rrepe,n;aqH ttacne.n;cTBeHHOii HHQJOpMaIJ;HH Ka:>K,[\OMY H3 ,n;oqepRHX JI,n;ep. qHCJIO xpOMOCOM
B 06e11x ,n;oqepHHX KJieTKax paBHO qHcJiy xpoMOCOM MaTepRHCKOM KJieTKH.
MRT03 BKJIIOqaeT qeThipe c]la3hI: npo<jJa3a, .Mema<jJa3a, attarjJa3a, meJto<jJa3a. fio BpeMeHH 3aHHMaeT B cpe,n;HeM 10 °lo OT BpeMeHH MHTOTHqecKOrO I.J;HKJia.
1. llpo<jJa3a ( 2n4c) - CaMaJI rrpo,n;OJIJKHTeJibHaJI c]la3a MHT03a, B KOTOpOH rrpoHCXO,[\HT KOR,n;eHcau;m1 xpOMOCOM H c]lopMRpoBaHHe MHTOTHqecKoro arrrrapaTa.

06~aR

60

6uo.1wzuR. OcnOBbt 1.j.Umo.1wzuu

Te.w(\1aail
)H'.110on1<rn K.:ienrn

A

Pfll.:THTe:1bHaa K.1Pih'.ll

6

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a-13 AByx xpoMarnA nocne G2-nepa-10Aa 2n4c; Ill - A04epHa-1e xpoMoCOMbl 2n2c.
5 - cxeMa M"1T03a B )1("1BOTHoi;i "' pacrnrnnbHoi;i KneTKax: 1 - nepeT5l)l(Ka; 2 - KneT04Ha5l neperopOAKa

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o60JIOqKM M .H,ll;pbIIIIeK; cpopMHpOBaHMe MMTOTMqecKoro arrrrapaTa: pacxom,n;eHMe
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BepeTeHa ,n;eJieHH.H KJieTKH.
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MMTOTMqecKoro anrrapaTa; B KJieTKe HaXO,ll;HTC.H ,D;Ba ,ll;HIIJIOH,ll;HblX Ha6opa.
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K rrepncpepnn) B 3KBaTopHaJibHOH IIJIOCKOCTH.
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,aenenue Knem1eu. ftfeii.o.J

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61

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3.

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A.Mumo3 -

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pacrrpe,neJieHMf! reHeTWieCKoro MaTepMaJia He rrpOl1CXO,L\MT (ll3 0.D;HOH KJieTKll
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B «II8TOJIOr11qecKH M3MeHeHHbIX» M o6peqeH-

3

HhlX Ha rtt6eJib KJieTKax, He crroco6Hhrx B ,na.'lhHeiiUieM ,n;aTh KJieTKM IIOJIHOU:eHHbie (BOCIIaJieHMfl, 3JIOKa'IeCTBeHHbIH pOCT rrpM orryxOJif!X).

4

Me:Ho3
M euo3 -

3TO pe;:i:yI<U:MOHHOe ,neJieHMe,

B pe3yJJhTaTe KOToporo rrpoucxo,nnT pe,n;yKu;tta
'IllCJia XpOMOCOM c .D;llIIJIOll,[(HOrO

(2n)

.no rarr-

JIOll,[(HOro (n).

KOToporo CBR3aHO c IIOf!BJieHneM IIOJIOBOrO pa3-

PMc. 21. CxeMa Mel1o3a: 1 - MaTePHHCKHe xpOMOCOMbt; 2 - xpOMOCO-

MHOateHua (C03peBaHMe raMeT), II03TOMY ero

Mbl, COCT05Hl.IH8 H3 ABVX xpoMaTHA no-

Ha3hlBaIOT ,D;eJieHneM co3peBattua. OH rrpoucxo-

cne G2-nep1-10Aa: 3 - KOHb10ra1..11-1S'I;
4 -· Kpocc1-1Hrosep; 5 - nepsoe Mel10rn4ecKoe AeneH1-1e; 6 - sTopoe Mel1o-

Meiio3 -

cnoco6 ,neJieHna, B03HllKHOBeHwe

.n;HT IlpH o6pa30BaHMM raMeT

y

)Kl1BOTHbIX

H cnop y pacTeHHH, rptt60B 11 BKJIJO'IaeT ,ll;Ba IIOCJie,D;oBaTeJihHhIX .ne.11eHMH a,npa.

B

Kam,n;oM

TH48CKoe AeneH1-1e

62

06UfaH 6uo.1wzuH. OcnoBbl l{Umo1wzuu

,n;e.rremu1

BhI.n;e;:r.HIOT <IeThipe cTa,z:urn:: rrpo<l_:>a3a, MeTacpa3a, attacpa3a, Te.11oc}ia3a

c 1rn.n;eKcaM11I11.1111

II (pnc. 21).

Hep Boe .n;eJieHHe MeH:oaa (pe;:i;y1m;Ho1111oe)

H Hmeprpaaa I ( 2n2c----+ 2n4c)

-

rrpo11cxo,n;11T yBeJ1ttt1ett11e 'lHc.w opratte.ru1,

CHHTe3 BeliICCTB H HaKOIIJieHHe 3HeprHH, Heo6xo,n;HMhIX ,ll;Jl.ff ,[(CJICHIUI, yBeJIH'leHHe pa3MepoB KJieTKH, y,n;BoeHHe I!eHTpHoJiett, penJIHKanmi .D:HK. KaJK,[(a.H
xpoMocoMa npe,n;cTaBJieHa napoii: xpoMaTH,n;, coe.n;11tteHHhIX u;ettTpoMepoi1.

IlporjJa.Ja I ( 2n4c) -

caMa.H rrpol(OJIJKHTeJihHa.a <l_:>a3a, BKJIIO'laronwa IIHTh

CTa,[(HH:

1. Jlenmome1-w -

Ha'lHHaeTCH CIIHpa.'lH3aI!Hff xpOMOCOM. XpoMOCOMhl CTa-

HOBHTCH BH,[(HMhIMH B MHKPOCKOII, KaK ,[(JIHHHhie TOHKHe HHTH. Kam,n;aa xpOMOCOMa COCTOHT H3 ,n;Byx xpoMaTH,[(.

2.

3uiomena -

B

a,n;pe BH,D;eH ,[(HIIJIOH,[(HhIH Ha6op xpoMOCOM.

rrpo,n;oJimaeTca crrupaJIH3aIJ,MH

1(0/lo/O?aU,U.Fl roMOJIOrH'lHhIX
I'O.MOJWZU'lHblMU H33hIBaIOTCH xpoMOCO-

xpOMOCOM H rrpoHCXO,[(HT
xpOMOCOM.

Mhl, HMeIOliIHe O,[(HH3KOBYIO cpopMy H pa3Mep: O,n;Ha H3
HHX IIOJiy'leHa OT MaTepH, ,n;pyraa -

OT OTLI,a. foMOJIO-

rH'lHhie xpOMOCOMhl rrpHTHrHBaIOTCH H rrpHKJia,D;bIBaIOTCH ,n;pyr K ,n;pyry ITO BCCH ,[(JIHHe. U:eHTpOMepa 0,[(HOH
H3 rrapHbIX xpOMOCOM TO'IHO rrpMJieraeT K IJ,eHTpOMepe

61AeaneHT

,n;pyroii:, H KaJK,n;aa xpoMOMepa rrpm:reraeT K roMOJIO-

np011CXOA11T KOHbtora41-1si

rH'lHOH xpoMOMepe ,n;pyroil. llapa KOH'hIOrlipOB3BIIIHX
roMOJIOrH'lHhIX xpOMOCOM Ha3hrnaeTCH

6ueaJleHmO.M.

Kam,n;hIH 6HBaJieHT cocTOHT H3 'leTBepKH xpoMaTH,n; -

TeTpa,n;hI. "llttcJio 6ttBaJieH-

TOB paBHO raIIJIOH,[(HOMY Ha6opy xpOMOCOM.

3.

Ilaxumena -

rrpottcxo,n;RT .D;aJihtteii:IIIaa crr11paJIH3aI!MH. TectthIH KOHTaKT

Mem,n;y xpoMaTH,[(3MH ,n;aeT B03MOJKHOCTh o6MeHMBaThCH H,[(CHTH'lHbIMH Y'JaCTKaMH roMOJIOrH'lHhIX xpOMOCOM. 8To HBJICHMe Ha3bIBaeTCH

4.

,l(unJlomena -

KpOCCUH206epo.M.

xapaKTeptt3yeTCH B03HI1KHOBCHHeM CMJI OTTaJIKHBam·m.

XpOMOCOMhl, COCT3BJIHIOliIHC 6MBaJieHT, Ha'lHHalOT OTXO,[(HTh ,n;pyr OT ,n;pyra B 06JiaCTH I!CHTpOMep. llpH pacxom,n;eHMH IIJIC'l xpoMOCOM B HCKOTOPhIX MCCTax 06HapyJKMBaeTCH HBJICHHe rrepeKpecTa I1 cu;errJieHHH -

XHa3MhI, KOTOpbie CJiyJKaT

CTPYKTYPHhIM BhipameHtteM KpoccttHroBepa.

napa cecn>HHCKllX
xpoMaTl1,!I

X11a3Mbl

~a

ceen>HHCKllX

><pOMaTl1,!I

npo11CXOAl1T KpOCC11HrDBep

nax11neHa

,lJ,11nnornHa

,ae;ieuue K1£em1m. M euo3
5. l(ua1wHe3 -

63

xapaKTeptt3yeTCJI MaKCHMaJihHOM crm:paJitt3au;ttei1, yKopo-

qeHneM H yTOJilll;eHHeM xpOMOCOM. 0TTaJIKHBaHtte xpOMOCOM rrpo,n;OJIJKaeTCJI,
HO OHH OCTalOTCJI coe,n;HHeHHhlMH B 6ttBaJieHThl CBOHMH KOHIJ;aMH. 51.n;phlIIIKO
H a.z:i;epHaJI o6oJioqKa paCTBOPJIIOTCJI. IJ;eHTPHOJIH pacxo,n;JITCJI K IIOJIIOCaM, 06paayIOTCJl HHTH BepeTeHa ,n;eJieHHJI.

Mema<jJa3a I (2n4c). BttBaJieHThr xpoMocoM pacnoJiararoTcn no 3KBaTopy
KJieTKH. K HHM rrpttKpenJI5IIOTCJI HHTH BepeTeHa .n;eneHH51.

AHa<jJa3a I ( 2n4c). HttTH BepeTeHa ,n;eJieHHJI coKpaw;aroTcn, HK rro.JirocaM KJieTKH
xpoMocoMbt, a He xpoMaTH,D;hI, 1mK nptt MHT03e. B .z:i;oqep-

pacxo.n;nTcn roMo.JiorttqHhre

mrn KJieTKH nona.n;aroT TOJihKO no O,D;HOH H3 rraphl rOMO.JIOrHqHbIX xpoMOCOM.

TeJto<jJa3a I ( 2n2c -

n2c

B

1caJ1CiJou KJtemKe ). qHCJIO xpoMOCOM B Kam,z:i;oil: K.JieT-

Ke cTaHOBHTCJI rarrJIOH,IJ;HhIM, HO KaJK,n;an xpOMOCOMa COCTOHT H3 ,D;BYX xpoMaTH,IJ;.
Ha KOpOTKOe BpeMJI o6paayeTC.H H,n;epHan o6onoqKa, rrpottcxo.n;ttT ,n;eJieHtte u;ttTOIIJiaaMbI.

Y MHOrHx pacTeHHH KJieTKa H3 atta<Pa3hI I cpaay me nepexo.n;ttT B MeTallJaay II.

Merncpa3a I (2n4c)

AHa(j)a3a I (2n4c)

Te11ocpa3a I
(2n2c - n2c a Ka)l(,ll,oC1 KilSTKe)

BTopoe .n;eneuHe Me:Hoaa (a1rnau;Houuoe)
HHmepfjJa.Ja JI, uJtu uHmepKuHe3 ( n2c) -

xapaKTepHa ,n;n.H JKHBOTHhIX KJie-

TOK. IIpe,n;cTaBJI.HeT co6oii KOPOTKHH rrepephIB Mem.n;y nepBhIM H BTOphIM MeiioTH'IecKHMH .n;eneHH.HMH, BO BpeM.H KOToporo He rrpOHCXO,D;HT perrmrnau;tt.H ,n:HK.

IIpofjJa3a II ( n2c ). XpoMocoMhI yToJiw;aroTc.H H yKopaqttBaroTC.H. 51,n;pbIIllKO
H SI.n;epHa.H o6oJio•ma paapyillaIOTC.H. 06paayeTc.H BepeTeHo .n;eneHH.H.

M emafjJa3a I I ( n2c).

,n:ByxpoMaTH.D;Hhre xpoMOCOMhI BhICTpaHBaIOTCJJ B aK-

BaTopttaJihHOH IIJIOCKOCTH KJieTKH (MeTa<PaaHaJI IIJiaCTHHKa), K HMM rrpttKperrJISllOTCSI HHTH BepeTeHa ,n;eJieHH.H.

AHafjJa3a II (2n2c ). ,n:ByxpoMaTH,D;Hhre xpoMoCOMhI .n;enSITCJI Ha ,z:i;oqepHtte
XPOMOCOMbI (6bIBillHe xpoMaTH,D;bI), KOTOpbre pacxo,n;.HTCJJ K rrpOTHBOIIOJIOJKHbIM IIOJIIOCaM KJieTKH.

TeJtofjJa3a II (2n2c -

nc B KaJ1CiJou K.nemKe). XpoMOCOMhI .n;ecrrttpamrny-

IOTc.H, CTaHoB.HTC.H HeBH.D;HMhIMH. 06paayeTC.H .H,n;epHa.H o6oJIO'IKa. Kam.n;oe JJ,n;po co.n;epmttT rarrJiott,n;Hoe qttcJio xpoMocoM. IIpottcxo,n;ttT ,n;eJieHtte u;ttTorrnaaMhl, Ma ttcxo.n;Hoii KJieTKH o6paayroTC.H

'lembtpe ian.nouOHbte KJieTKH.

06~aJt

64

npoc)la3a II (n2c)

AHac)la3a II (2n2c)

6uo1wzuJt. Ocno6bL ifumo,11,ozuu

Merac)la3a II (n2c)

Te11oc)la3a II (2n2c - nc s Ka>KAO~ KlleTKe)

3Ha'!eHHe Mei1:03a
1. IIpe,1:i;oTBpa:u.~aeT y,n;BoeHtte qttcna xpoMOCOM B KaJK,n;OM HOBOM noKoneHHH.
2. 06ecneqHBaeT retteTttqecKoe pa3Hoo6pa3He raMeT, cmrnaHHoe c KpocCHHroBepoM H pacxom,n;eHHeM B attacpa3e I pa3HhIX Ha6opoB HeroMoJiorttqHhIX xpoMOCOM.
3. 5!BmieTCH OCHOBOH KOM6HHaTHBHOH H3MeHqlfBOCTH.

PA3MHO>KEHHE OPrAHH3MOB
Pa3.M1t0J1Cenue - CBOMCTBO JKHBhIX opraHH3MOB npoH3Bo,n;HTh ce6e no,n;o6HhIX. Cyru;ecTByeT ,n;Ba ocHOBHhIX cnoco6a pa3MHOJKeHHH - 6ecnonoe H nonoBoe.

BecnoJioe paaMuoarnuHe
BecnoJtoe pa3.M1-toJ1Ce1-tue ocyru;ecTBJIHeTcH npH yqacnm o,n;Hoii po,n;ttTeJibCKOH
oco6u H npottcxo,n;HT 6e3 o6pa3oBaHHH raMeT. Oco6H o6pa3yIOTCH H3 o,n;tto:i1 HJIH
HeCKOJlbKHX COMaTnqecKHX KJieTOK po,n;HTeJI.H ITYTeM MHTOTHqecKoro ,n;eneHHH.
IloTOMCTBO HBJIHeTCH reHeTHqecKHMH KOITHHMH po,n;HTeJibCRoro opraHH3Ma. Hatt6onee urnpoKo pacupocTpatteHo cpe,n;H npo:KapttoT, rpu6oB H pacTeHHH. Bttonortt'!ec:Koe 3HaqeHtte 6ecnonoro pa3MHOJKeHHH: ycttJIHBaeT pOJih cTa6ttJIH3ttpyroru;ero eCTeCTBeHHOro OT6opa, cnoco6cTBYeT coxpaHeHHIO npHCIIOC06JIHeMOCTH
K nocToHHHhIM ycnoBHHM cpe,n;hl o6HTaHHH.
Crroco6h1 6ecnoJioro pa3MHOJKeHHH:
1. ,l{eJtenue - cnoco6 6ecnonoro pa3MHOJKeHHH, xapaKTepHhIH ,D;JIH o,n;HoKJieToqHhIX opraHH3MOB, rrptt RoTopoM MaTepttHCKaH oco6h ,n;eJIHTCH Ha
,D;Be HJIH 6oJibIIIee KOJIHqecTBO ,n;oqepHHX KJieTOK.
Bhl,n;eJIHIOT: rrpocToe 6nttapttoe ,n;eneHHe (rrp0Kaptt0Tb1); MHTOTHqecKoe
6HHapttoe ,n;eJieHHe (npocTeHIIIHe, 0,D;HOKJieToqHbie BO,ZJ;OpOCJIH); MHO-

pa 3 MnoJtCenue opzanu.1M06. Becno1we pa3M1toJ1Ce1tue

65

PaaMHO»WHHC

- .u.eJ1eHHe (6H11apnoe, MHTOTlfliecKoe, m113oron11H),
-

II0l!KOB8HJIC,

- cjlparMe11rn111rn,

c OOJ10)10'1'BOpemieM:

-

KOH'bJOraLU1fl'

-

-

-

KOIIY~1.HIUU1

IIOJIH3M6pHOHH.H,

H30raMH.H,

- cnopoo6paaona1me,
- aereTaTHBHOe,

- reTeporaMIUI,

-

Bea onJio.a,oTRopenHH:

- ooraMH.H.

KJ"I0HHpOB8HHC

118pTCHOfCHC3

.meCTBeHHOe .n;eJieHJfe, MJIM IUH30J'OHM.H (MaJI.HpMMHhIM IIJia3MO,Zl;MM).
fl.n;po .n;eJIMTC.H c o6pa30BaHJfeM ,ZJ;BYX HJIM 60Jihlllero 'IMCJia .a.n;ep, KOTOphle OKpy.maIOTC.H CJIOeM II;HTOIIJia3Mhl, If B pe3yJihTaTe <jlopMnpyeTC.H
caMOCTO.HTeJibHhlH opraHM3M.

2.

IlO'llWBattue -

cnoco6 6ecrrOJIOI'O pa3MHOJKeHM.H, rrpM KOTOPOM HOBhie

oco6M o6paayIOTC.ff B BM,ZJ;e BhlpOCTOB Ha TeJie pO,ZJ;MTeJihCKOM oco6M • .n:oqepHMe oco6tt, OT,ZJ;eJI.rr.rrch, rrepexo.n;.rrT K

caMOCTO.HTeJihHOMy o6paay

.IKM3HM (rtt.n;pa, .n;po.mmtt) MJIM MOI'YT OCTaBaTbC.H rrpnKpernieHHhIMH,
o6paay.rr KOJIOHMM (KopaJIJIOBbie IlOJIMilhI).

3. <PpazMettmau,u.H -

crroco6 6ecrrOJIOI'O pa3MHOJKeHM.ff, rrptt KOTOpOM HOBbie

oco6M o6paayIOTC.H M3 <jlparMeHTOB, Ha KOTOpbie pacrra.n;aeTC.ff MaTepHHCKa.H
oco6b (peCHM'IHhJe, KOJih'IaThJe qepBM, MOpCKMe 3Be3,ZJ;hl, CIIMpOrMpa).

4. IIOJtU3M6puottu.H -

crroco6 6ecrroJioro pa3MHomeHtt.rr, rrptt KOTopoM HO-

Bh1e oco6tt o6paayIOTC.H M3 qacTeii:, Ha KOTOpb1e pacrra.n;aeTc.a 3M6pttoH
(6poHeHOCII;hl, MOH03MI'OTHhie 6JIM3HeII;hl qeJIOBeKa).

5.

BezemamuBttoe pa3MHOJ1Ce1-tue -

crroco6 6ecnoJioro paaMHomeHn.H,

nptt KOTOpOM HOBhie oco6tt o6paayIOTC.H M3 qacTeii: BereTaTMBHOI'O TeJia
MJIH H3 oco6bIX CTpyKTYP (KOpHeBmIJ;e, KJiy6eHb, JIYKOBHII;hI, OTBO,ZJ;KH,
ychl, JIMCTh.H M .n;p.). MomeT 6bITh ecTeCTBeHHhlM H ncKyccTBeHHhlM.
XapaKTepHo ,ZJ;JI.rr pacTeHHM H HCIIOJih3yeTC.H B ceJieKII;HH (HcKyccTBeHHoe BereTaTHBHOe pa3MHOJKeHHe).
6.

Cnopoo6pa3oBattue -

pa3MHomeHHe aa C'IeT o6paaoBaHn.rr crrop. Xa-

paKTepHo ,ZJ;JI.H pacTeHMM (BO,Zl;OpOCJIH, MXH, rrarropOTHHKH), rpH60B (neHHII;HJIJiyM, IUJI.HilO'IHhJe rpH6hI), rrpocTeMIUHX (crropOBHKH).

7. KJto1-tupoBa1-tue -

ttcKyccTBeHHhIM MeTo.n; pa3MHomeHH.H, KOTOphIH 110-

.HBMJic.a B Ha'IaJie 60-x rr. XX B. OcHOBaH Ha noJiyqeHMH HOBoro opraHH3Ma H3 O,ZJ;HOM KJieTKH HCXO,ZJ;HOI'O.

B OCHOBe o6pa30BaHH.ff KJIOHa Jie.IKHT MH-

T03 • .Il:JI.rr KJIOHHPOBaHH.H pacTeHHH KJieTKH o6pa30BaTeJihHOM TKaHM BhlP~HBaIOT Ha cnerr;HaJihHhlX IIHTaTeJihHhlX cpe.n;ax (MeTO,Zl; HCIIOJih3YeTC.ff

,Zl;JI.H noJiyqeHn.rr HOBhIX copTOB pacTeHnii:).

B 1997 r.

KJIOHnpoBaHneM

6hlJia IlOJiyqeHa OBe'!Ka .Il:oJIJIH, KOTOpa.H He OTJIH'IaJiaCh OT ,ZJ;OHOpa .rr.n;pa,
BO CHJihBO OTJIH'IaJiaCb OT perr;nrrneHTa • .D:oHopoM 6bIJIH KJieTKH MOJIOqHOM

)f<CJie3hl 0.L(HOH rropO.L(hI, a peI.1,I:IITI:IeHTOM -

.HMIJ,eKJieTKI:I c npe.nBapI:ITeJib-

HO pa3pymeHHhIMH a.npaMM OBIJ.hI .npyroii: rropo.nhr.

B

HacTomu;ee BpeM.H

Be.L(yTC.H pa60Thl no KJIOHI:IpOBaHI:IIO IIeJIOBeKa, 'ITO Bhl3hIBaeT 6ypHbie crropbl He TOJihKO cpe.L(I:I YlJeHhIX, HO II y pa3JII:IlIHhIX rpyrrrr HaceJieHI:Ifl.

lloJIOBOe pa3MHO~eHHe

IT oJwBoe paaM1wJ1CeHue

ocyru;ecTBJI.HeTc.H rrpM yIIaCTI:IH .ll:BYX po.nMTeJil>CKHX

oco6eii, conpOBOJK.L(aeTC.H IIOJIOBhIM rrpou;eccoM, rrpH KOTOpOM rrpOHCXO.L(HT CJIH.HHHe .L\BYX IlOJIOBhIX KJieTOK (raMeT).

y

HOBhIX oco6eii COIIeTaIOTCfl rrpH3HaKH

.L\BYX pO.L(HTeJieii. IlO.HBHBIIrnec.H IIOTOMKH OTJII:IIIaIOTC.H reHeTH'IeCKH .npyr OT
.npyra. 3TO rrpHBO.L(HT K IlO.HBJieHHIO HOBhIX KOM6HHaIJ,Hll reHeTHIIeCKOro MaTepHaJia. IIoJIOBoe pa3MHomeHHe ,ZJ;eJiaeT opraHH3M 6oJiee a.narrTHpoBaHHhIM K H3MeH.HIOIIJ,HMC.H ycJIOBI:IHM 0Kpyma10ru;eii cpe.L(hI.

B

npou;ecce 3BOJIIOIJ,HH rroJioBoe

pa3MHOJKeHHe 6oJiee rrporpeccHBHO. He.nocTaTKH noJioBoro pa3MHOJKeHH.H: Heo6xo.nnMo yqacTne .L(Byx oco6eii (KpoMe napTeHoreHe3a) H Tpe6yeTC.H 6oJihIIIHM
npoMeJKyTOK BpeMeHH, IIeM npH BereTaTHBHOM pa3MHOJKeHHH, He06XO.L(HMhlll

B

.ZJ;JI.H noJIHoro cpopMnpoBaHH.H opraHH3Ma.

3aBHCHMOCTH OT oco6eHHOCTH

CTpoeHH.H raMeT BhI,ZJ;eJI.HIOT CJie.L(yIOIIJ,He cpopMhl IlOJIOBOro pa3MHOJKeHH.H:

1. 1130ZaMUH (OT rpeII. isos -

paBHhIM H

ga'mos -

6paK) -

rrpHMHTHBHa.H

cpopMa IIOJIOBOro npou.ecca, rrpH KOTOpOM CJIHBaIOTC.H .[(Be O.ZJ;HHaKOBhie
ITO BeJIH'IHHe MaJihie IlO.L(BHJKHhie raMeThl (paBHOJKryTHKOBhie, B0.[(0pOCJIH, XHTpH.L(HeBhie rpH6hI).

2. I'emepozaMuH

(oT rpeII.

he'teros -

HHOii, .npyroii, H

gamos -

6paK) -

npHMHTHBHa.H cpopMa IlOJIOBoro rrpou.ecca, npH KOTOpOM CJIHBaIOTC.H ,ZJ;Be
IIO.ZJ;BHJKHhie co mryTHKaMH KJieTKH pa3Horo pa3Mepa (BO.L(OpOCJIH, XHTpH,ZJ;HeBhie rpH6hl).

3. OozaMUH

(oT rpeq.

oo'n -

.Hiiu;o) -

Han6oJiee pacnpocTpaHeHHa.H cpopMa

IIOJIOBOro npou;ecca, npH KOTOpOM JKeHCKHe raMeThl -

.HMIJ,eKJieTKH -

He

no,ZJ;BHJKHhI H 6oJiee KpynHhie no cpaBHeHHIO c MY.IKCKHMH raMeTaMH crrepMaT030H,ZJ;aMH (MHOrOKJieTO'IHhie JKHBOTHhie, rpH6hI, BO.D;OpOCJIH, BCe
BhlCIIIHe pacTeHH.H).

4. KoH010za4ua

(OT JiaT.

conjugatie -

coe,ZJ;HHeHHe) -

npou;ecc coe,ZJ;HHe-

HHH .ZJ;BYX oco6eii, rrpH KOTOpOM npOHCXO.ZJ;HT o6MeH HaCJie,ZJ;CTBeHHhlM
MaTepHaJIOM, B pe3yJihTaTe qero IlO.HBJI.HIOTC.H oco6H, reHeTH'IeCKH OTJIH'IHhie OT pO,ZJ;HTeJihCKHX opraHH3MOB (HHcpy3opH.H-TycpeJihKa, KHIIIelIHafl naJIOIIKa).

B

pe3yJihTaTe KOH'hIOrau;HH He nponcxo.nttT yBeJIH'IeHH.H

'IHCJia oco6eii.

5. KonyJta4ua (oT JiaT. copulatio -

CJIH.HHHe) -

noJioBoii npou;ecc y O.ZJ;HO-

KJieTO'IHhIX opraHH3MOB, npH KOTOpOM IlOJIHOCThIO CJIHBaIOTC.H KOilYJIHpyIOIIJ,He oco6H, BhIIlOJIH.HIOIIJ,He cpyHKIJ,HH IlOJIOBhIX KJieTOK (raMeT).
IIpn aToM rrpoHCXO.ZJ;HT peKoM6HHaIJ,H.H (o6MeH) HacJie,ZJ;cTBeHHoro MaTepnaJia (MaJI.HpHMHhIM IIJia3MO.ZJ;Hll).

6. IlapmeHozeHe3
.neHne) -

(oT rpeq.

parthe'nos -

.neBCTBeHHHIJ,a H

genesis -

pom-

cnoco6 ynporu;eHHoro noJioBoro paaMHOJKeHH.H, npn KOTopoM

no1wBoe pa.1MHOJ1CeHue. I'aMemoiene3

67

aapO.ll;billl pa3BlIBaeTCH 113 HeOil.'IO)J;OTBOpemwii HllrJ,eKJieTKII. I1CTO'IHlfKOM HaCJie;:J;CTBeHHOro MaTepHaJrn }l,JIH pa3BHTllH HBJ'IReTCR ):t:HK H:ihi;eKJ'IeTKll

(iw-wielie3). IIapTettorettea npe;:i;cTaBJIReT co6oii Mo;:i;mp1maumo

IIOJIOBOrO pa3MHOiKeHHR B npouecce 3BOmOIJ,HH onpe;:i;eJ'IeHHbIX BH)J;OB
iKllBOTHbIX. Illupmw pacnpocTpaHeH y 6ecn03BOHO'IHbIX (TJ'IH, OCbI, Il'IeJihI, .n.aqmttH) Hy II03BOHO'IHbIX iKllBOTHblX (npeCMhlKaIOIL\HecR, IlTHU.bI).
PaaJrn'IaIOT

ucKyccmBeflliblU tt ecmecmee1tlibtU (TJIM, oc1~r, nqeJibI) nap-

Tettorettea. IIapTeHorettea BKJIIOqeH B iKll3HeHHhre UMKJihr :'.\l:HorHx napa3IITOB. Pa3MHOiKeHne, npn KoTopoM paaBHTHe Riiua npottcxo,n;HT aa c'IeT
reHeTu:qecKoro MaTep11UJi:a cnepMaToao11;:i;a ( TaK KaK H;:J;po HH1J.eK"'IeTKI1
rror116aeT), HaahrnaeTcR
pe,n;Ko (y iKI1BOTHb!X -

I'epMa</JpoiJumbt -

aniJpo2e1tea. B np11po;:i;e BCTpeqaeTCH Kpalitte

Hae3)J;HllKll, y pacTeHllH -

BII.D;hI, y K0Topb1x o.n;tta M Ta

KyKypyaa).

me oco6b cnoco6tta npoHa-

Bo,n;nTh n iKeHCKHe, M MYiKCKtte raMeThl. fepMa<lJpo,n;tt:TH3M CtJMTaeTcR caMo:i1:
rrpttMllTllBHOH <lJopMOH IlOJIOBoro pa3MHOiKeHMR H CBOHCTBeHeH rrpHMHTHBHhlM
opraHII3MaM (npocTeiillltt:e, nJiocKne, JieHTO'IHhre tJepBM, yJIHTKH H .n;p.). IIpe.n;cTaBJIHeT co6oH: rrpncnoco6JieHHe K Cll,UR'IeMy, MaJIOilO,ll;BHJKHOMy, rrapa3HTHqeCKOMY 06pa3y iKll3Hll.

I'aMeToreHe3
I'aMemoie1tea -

3TO npouecc <lJopM11poBaHHR raMeT, OCHOBHbIM 3TaIIOM KO-

TOporo RBJIReTCR MeH03. Pa3Bl1Tlle raMeT y MHOrOKJieTO'lHbIX iKHBOTHhIX rrpo-

cnepMamo3ouiJbt) o6pa3yIOTCR B CeMeHHMKax, meHCKHe IlOJIOBhie KJieTKll ( JlUl{eKJtemKU) - B HH'lHHKax. I'aMembl - 3TO BbICOKO,n;mp<lJepeHU.HPOBaHHbie KJieTKH,

HCXO,ll;HT B noJIOBhlX meJieaax (rotta,n;ax). MymcKHe noJIOBhie KJieTKH (

HMeIOru;He ranJIOH,ll;HhlM tta6op xpOMOCOM. Ilpou.ecc o6pa30BaHHR MyiKCKHX IIO-

cnepMamoieHe3, a meHCKHX - 060ZeHe3.
CnepMamoienea no,n;paa,n;eJIReTcR Ha qeThrpe <lJaahr:
1. B </JaJe pa3MHoJ/CeHuH tt:3 KJieToK cnepMaToreHHOH TKaHH (roHOU.HTOB)
B pe3yJibTaTe MHT03a o6pa3yIOTCR ,ll;HilJIOH,UHbie cnepMamoioHUU
(2n2c).
2. <Pa3a pocma COOTBeTCTByeT HHTep<lJa3e I Merroaa (no.n;roTOBKa KJieTOK

JIOBhlX KJleTOK Ha3bJBaeTCR

K .n;eJieHHIO), B KOTOpoii npOHCXO,D;HT penJIHKaUHH ):t:HK. CnepMaToroHHll yBeJIHl{HBaIOTCH B paaMepax, H 113 Kam,n;orr KJieTKH pa3BHBaeTCH

cnepMamo4um I nopRiJ1ca (2n4c).
<jJaabt co3peeaHuJl KJieTKH .n;eJinTcn Mett030M. B pe3yJihTaTe
I ,n;eJ1eHtt:H Meiloaa o6pa3yIOTCH .n;Be KJieTKH cnep.1namo4umbt II nopRiJKa (n2c), a IIOCJie BTOporo ,n;eJieHHH pa3BHBaIOTCR qeTbJpe O,ll;HHaKOBble no BeJIH'lHHe KJieTKH cnepMamuiJbt c ranJIOH,UHhlM aa6opoM
XpOMOCOM (nc).
4. B </JaJe </Jop..MupoeaHUJl npoHcxo,n;tt:T KJieTOtJHa.ff ,n;n<lJ<lJepeauHpOBKa
H 06pa3yroTCH c nep.1namo3ouiJbt c rarrJIOH.D;HhlM aa6opoM xpoMOCOM ( nc) ·
ropMoH, 06ecne'lHBa10ru;Hii crrepMaToreHe3, - mecmocmepoH. Kam,n;hre 2-3 Meca~ cnepMaT030H,Uhl o6HOBJIHIOTCH, H cnepMaToreHe3 npoMCXO,UHT HenpepbIBHO.

3.

Bo BpeMn

68

06UfaR 6uo1tozu1i. Ocn06bL 4umo1tozuu

reueT11- '

3011bl

'IeCKHii

CnepMaTore11e3

; ( cjia:-ia )!{!13HeHHOro

0Bore11e3

ua6op

IJ,HK"'la)
cnepMaToro111111
Pa3MHO:iKfHmH
(MHT03)

i

II

•

I

!

I-

III

l

2n2c

- --· - \. ····-· i-cnepMaTori.uT I nopH,n;Ka

I

PocTa
(HHTep<lJaaa)

i

•

." e

'e
f> .:

c{
·t

OBOrOHl111

·-- ·-- ,--- -

·-r·

--·+
OBOlJ,HT lnopH,D;Ka

2n4c

crrepMaT,OlJ,HT II'nopR,n;Ra I

Co3peBaHHH
.
(Meii:oa)

I

OBOlJ,HT,

pe):(.¥Kll,HOHHOe

II rropRJ(Ka

teJJbu,e

/"

I

......
......

""0

·=::s •
Q)

[ __

f----~

IV

--

•

)

\,j

n2c

. ..

HHIJ,eKJJe'i\Ka
:
'peJJ,yKU,HOlll!b~e

•• •

nc

crrep1:'4aT03oH,n;hl

I
'

(

,,--'-

,

\___)

'_

TeJJbll,3

I --- ---· --- -- -

<t>opMHpOBaHHH

L__

--

i

i

nc

'-·

Cmpoenue cnep.Mamo.10uiJa (pttc. 22, 23)
3peJiblH crrepMaT030H,ll; HMeeT rOJIOBKY' rneii:Ky H XBOCT. B ZOJl06Ke HaXO,ll;HTCH
RiJpo (c rarrJIOH,ll;HblM Ha6opoM xpOMOCOM) H aKpOCO.Ma (BH,ll;OH3MeHeHHhlH KOMIIJieKC roJih,ll;:>KH, co.n;epmam;nii: <PepMeHThl, pacm;ern1mom;ne o60JIO'!KY HMIJ;eKJieTKll ). B rneii:Ke Haxo.n;HTCH u;eHTPHOJIH (coe.n;HH.mom;tte oceByro qacTh c H.n;poM roJioBKtt), B rrpoMe:>KyTO'!HOH (cpe.n;Heii:) 'laCTH - MHTOXOH,ll;pHll (CHHTeaupyrom;ue
ATcf>). Xeocm rrpe.n;cTaBJieH o.n;HHM, y HeKoTophIX su.n;os .n;syMH H 6oJiee J1aymuKa.Mu (o6paaoBaHHbIMH MHKpOTpy60'!KaMH) • .Il:JIH ,ll;BH:>KeHllH :>KrYTHKOB llCIIOJib3yeTCH :meprHH MaKposprtt'!eCKHX CBHaeii: ATcf>. UuTOIIJia3Ma crrepMaToaott.n;a
rrpe.n;cTaBJieHa TOHKOH o60JIO'!KOH.
Oeoiene3 rro.n;paa.n;emreTc.H Ha Tpu ipaabl:
1. B </Ja3e pa3.MHOJtCenuR H3 KJieToK osoreHHoii: TKaHH .HH'!HHKOB (roaou;ttTOB) B pe3yJibTaTe MllT03a o6pa3yJOTC.H ,ll;HIIJIOll,ll;Hbie 060ZOHUU (2n2c ).
2. <Pa3a pocma cooTBeTCTByeT HHTep<Paae I Meii:oaa (rro.n;roTOBKa KJieTOK
K .n;eJieHuro), B KOTopoii: rrpoucxo.n;HT perrJiuKan;HH .IJ:HK. 0Boroattu yBeJill'IHBaJOTC.H B pa3Mepax, BCJie.n;CTBHe HaKOrrJieHll.H IIllTaTeJibHbIX Be-

pa 3 .Mnom:enue opzanu3M06. I'aMemozeneJ

69

B
P111c. 22.

CnepMaTOreHe3

y MOPCKOiii

CBHH-

P111c. 23.

CrpoeH11e cnepMarn3011Aa: A -06-

KH. Y4aCTOK nonepe4HOro pa3pe3a 113B11roro

t11Hiil BHA; 5 -

KaHaJlbU,a ceMeHHHKa: 1 -

uieiilKa; 3 -

2-

cnepMaTOu,HT I nopsiAKa;

1..!HT II nopsiAKa; 4 -

cnepMaTOIOHHS1;

3-

cnepMaro-

cnepMarnAa; 5 -

coMa); 5 -

cxeMa crpoeHHS1. 1 xBocr; 4 -

fOllOBKa; 2 -

annapar ronbAJKH (aKpo-

S1APO; 6 - MHTOXOHAPHH; 7 -

HHTb (>KryrnK);

ocesasi

8 - L!eHTPHOll11; 9- npoMe>KyT04-

HaS1 4aCTb

cnepMaro30HA

w;ecTB B Btt.n;e ace.TJTKa, H H3 Kam.n;oiI KJieTKH pa3BHBaeTcsr oBou,um

3.

p.Rihca (2n4c).
Bo BpeMsr </Ja3bt co3pe6anu.R nepe.n; oByJimi;neiI OBOD;HT
JIHTCH nyTeM Metto3a. B pe3yJihTaTe

I

I

I no·

nopsr.n;Ka .n;e-

.n;eJieHHH Metto3a 06pa3y10Tcsr: 60-

Jiee KpyrrHaH K.JieTKa c IIHTaTeJihHblMH Bern;ecTBaMH H opraHeJIJiaMH -

oBou,um II nop.Ra1ca (n2c) H o.n;Ha MeJIKasr KJieTKa - nep6oe noJt.Rpttoe
B KOTopoiI ttMeeTCH
(peay1Cu,uottttoe, ttanpa6umeJtb1toe} meJtbU,e, TOJihKO sr.n;po. BTopoe .n;eJieHtte Meilo3a ,n;oxo,n;HT .n;o cTa.n;tttt MeTacpa3hI,
HO He rrpo.n;oJimaeTCH ,n;aJihllle .n;o Tex rrop, rroKa He rrpott3oiI.n;eT orrJio.n;oTBopeHtte, IIOCJie KOTOporo OBOD;HT
pa3yeTCH KpyrrHasr K.JieTKa -

II rropSI.n;Ka 3aBeplllaeT ,n;eJieHHe H

06-

.Ritu,e1CJtem1Ca (nc), a TaKme Bmopoe no-

Jl.Rpnoe meJtbU,e. Ifa rrepBoro rroJIHpHoro TeJihn;a 06pa3y10TCH .n;Ba MeJIKHX BTOpHqHbIX. Bee OHH B .n;aJibHettllJeM pa3pylllaJOTCH.
0BoreHe3 rrpoHCXO,ll;HT IIO,IJ; KOHTpOJieM ropMOHa npoiecmepotta. B OTJIHqtte
OT o6pa3oBaHH.H crrepMaT030H,IJ;OB rrpon;ecc o6pa3oBaHH.H .Hiin;eKJieTOK y qeJioBeKa liaq1rnaeTCH B aM6pHOHaJibHOM rreptto,n;e I:l TeqeT rrpepbIBHCTO.

y

3apO,ll;billJa

llOJilIOCTbJO ocyw;ecTB.JI.HJOTC.H cpa3bl pa3MHO:lKeHHH, pocTa H HaqHHaeTC.H cpa3a
coapeBaHHH.

R

MOMeHTY poac.n;eHH.H .n;eBoqKI:l B ee .HHqHHKaX HaXO,ll;.HTC.H COTHH

TLicaq oson;ttTOB

I

rropsr.n;Ka, ocTaHOBHBIIIHXCH Ha cTa.zi;HH au1Cmuome1tbt (Bo

Bpel\rn KOTOpoiI xpOMOCOMbI HMeJOT BH,IJ; «Jla.Mn06blX w,emolC») npo</Ja3bl

3a. B

I .111.eito-

nepwo.n; no.Jiosoro co3peBaHHH Metto3 B0306HoB.JI.HeTc.H, H Ka:lK,ll;hIH Mee.Hu;

no,n; ,n;eiicTBHeM IIO.JIOBbIX ropMOHOB O,ll;HH H3 OBOD;HTOB ,IJ;OXO,ll;HT ,IJ;O MeTacpa3bI

II

06UfaH 6uo.rwzun. Ocno6&t lfUmo.rwzuu

70

Meiio3a. B

.n;aJihHeiiweM Meiio3 MomeT

H.D;TYI .n:o KOHu;a npn ycJIOBllH OilJIO,ll;OTBOpeHHH, ec.1n onJIO.D;OTBope:f!ne He nponcxo.n;ttT, OBOUHT

II

nopH.n;Ka norn6aeT

H BbIBO,ll;HTCH H3 opraHH3Ma.

Cmpomue .HUU,eJC.llemJCu (pttc.

24).

ct>opMa OKpyrJiaH, pa3Mepb1 OT HeCKOJibKHX MHKpOMeTpOB .n;o HeCKOJibKMX
caHTMMeTpoB. KJieTKa Heno.n;BumHa.a, co.n;ep.m11T BCe K.JieTOtfHble opraHe.JI.JibI, MOpcpo.Jior11treCKH ocpopM.JieHHoe .a.n;po, H.n;epHO-UHTOnJia3MaTUtfeCKOe COOTHOllleHne
C.ll;BHHYTO B CTOPOHy I.J;HTOn.Jia3Mbl 38 CtfeT
PMC.

I -

4-

24. 'rlillu,eKJleTKa MJl8KOm1Tat0Ll.\ero:
2 - 5lApo; 3 - 060110YKa;

uvirnnna3Ma;

3anaca nuTaTe.JihHbIX Bem;ecTB -

PHK,

¢01lllVIKYll5lpHbl8 Kll8TKVI

me.JIT-

Ka. JRe.JITOK co.n;epmnT mnpbI, yrJieBO,ll;bl,
MUHepaJibHbie Bem;ecTBa, 6eJIKH

(JinnonpoTen.n;hI, rJIHKonpoTen.n;hI). KoJIHtfeCTBO nnTaTeJibHbIX Bem;ecTB 38BHCHT OT ycJIOBHM, B KOTOpbIX npoHCXO.ll;HT
pa3BHTHe 3apO.lJ;blllla. IIpn pa3BHTHH .aiiu;eKJieTKH BHe oprattn3Ma MaTepn (nTn-

95 '% OT ee o6'heMa, a BHYTPH opram-135 'X,' TaK KaK nnTaTeJibHhie Bem;eCTBa, Heo6xo.n;n-

IJ;hI, perrTHJIHH) meJITOK COCTaBJI.HeT 6oJiee
Ma (MJieKonnTaIOm;He)- MeHee

Mbie .D;JIH pa3BIITMH, 3M6ptt0HbI no.JiyqaIOT OT MaTepn.
Ha.n; n.Jia3MaTntrecKoii MeM6paHoii nMeIOTCH .n;onoJIHHTeJihHhie o6oJIOtfKH,
pa3JIHtraIOm;necH no nponcxom.n;eHnIO:

nep8U'lllbte ( JJCe.TLmO'lltbte) -

o6pa3yIOTCH H3 noBepXHOCTHOro CJIOH He-

3peJIOM noJIOBOM KJieTKH (OBOI.J;HTa); npoHH38Hbl MHKpOBOpCHHK8MH
H 0Tp0CTK8MH cpOJIJIHKYJIHPHhIX KJieTOK, no KOTOpbIM nocTynaIOT nMTaTeJibHhle Bem;ecTBa. BhlnOJIHHIOT 3am;HTHYIO cpyHKI.J;HIO, o6ecnetrHBa.a
BH.ll;OBYIO cneu;mpwrnoCTb npOHHKHOBeHHH cnepMaT030IJ,ll;a.

y

MJieKO-

nnTaIOIIJ;HX 3Ta 060.JIOtfIW Ha3hIBaeTCJI 6.JieCTJim;eii;

BmopU'lllbte ( xopuon) -

COCTOJIT H3 cpoJIJIHKYJI.HpHbIX KJieTOK HJIH Bbl-

.n;eJIHeMbIX HMH ceKpeToB. BTop:ntrHa.a o6oJIOtfKa H:nu; HaceKOMbIX co.n;ep.m:nT KaHa.JI -

MHKponn.Jie, trepe3 KOTOphIM cnepMaT030H.D; npomrna-

eT B Hiiu;eKJieTKy;

mpemu'l.nbte -

cpopMnpyIDTCH BO BpeM.a npoxoa<,n;eH:nH Hiiu;eKJieTKH no

.aiiu;eBo.n;aM H3 Bem;ecTB, CeRpeTnpyeMhlX meJie3aMH CTeHOK .aiiu;eBO,ll;OB.
Xopolllo pa3BHTLI y penTHJIHM n nT:nu;, Hanpn.Mep, TpeTWIHhlMH o6oJiotrKaM:n IlTHI.l; HB.JIHIOTCH 6eJIKOBaJI, no.n;cKOpJiyno:aaJI lI CKOp.JiyrroBaJI 060JIOtfKH. Riiu;eKJieTKH MJieKon:nTa10m;:nx TpeT:ntrHoii o6oJIOtfKH He nMeIOT.

Oce.Menenue -

3TO npou;ecc, 06ycJiaBJIHBa10m;:nii BCTpetry meHcK:nx n Mym-

CK:nx raMeT y muBOTHblx. PaaJiutraIOT ecTecTBeHHOe u 11cKyccTBeHHoe oceMeHeH:ne (lll:npoKo 11cnoJIL3yeTca B pL160Bo,n;cTBe n m:nBOTHOBo.n;cTBe). HapyJJCnoe oce.Menenue BCTpetraeTCH y MHor:nx muBOTHLIX, 06uTa10m;ux B Bo,n;e. B 3TOM cJiyqae
.aiiu;a ll cnepMaT030H,ll;hl Bbl,ll;eJI.HIOTCH B OKpyma10m;y10 cpe.n;y. r.n;e npOHCXO,ll;HT

pa3MnoJ1Cenue opzanu.1M06. I'aMemoiene3

HX cJIMJIHMe.

BHympe111-tee oce.11te11e/-lue

71

xapaKTepHO /1.•1R o6I1TaTeJ1eH cywn, r.ne

OTCYTCTBYIO'f yc.'IOBIUI ,ll;JIR. coxpaHeHMJI H BCTpeqlI raMeT BO BHenrnei1: cpe,11;e.
ITplI 3TOH cpopMe oceMeHeHHR cnepMaT030H,ll;hI BBO.D;RTCR B rro.10Bbie nyT11 caMKH.
OceMeHeHMe npe/1.rneCTByeT c.1e)l,y10~eMy npou,eccy -

On1wrJomeope1tue -

OIIJI0/1.0TBopeHHIO.

3TO npou,ecc c.111JIHHR MymcKoll: 11 meucKoii no.10Bhix

J{JieTOK (raMeT), B pe3yJibTaTe KOTOporo 06pa3yeTCR. OIIJIO,ll;OTBOpeHHOe HMU,O,
HJIH

:mioma,

.na10~aJI

HaqaJio HOBOMY opraHI13MY.

ITpou;ecc OIIJIO,D;OTBOpeHHR MO/KHO pa3,D;CJII1Th Ha HCCKO.'lhKO cpa3 (p11c.

1. <Paaa c6.iuJ1CeHUR.

BcTpeqe

raMeT

cnoco6cTBYIOT raMOHhI -

25):

x11M11qe-

CKMe Be~ecTBa, BhI,ll;eJIJieMhie IIOJ'IOBhIMH K.'leTKaMM. CnepMaT030M,ll;hI
BhI,D;eJIRIOT aH,n;poraMOHhl, a J:lMU,eKJieTKH -

I'MHOraMOHbl. 8TH Be~eCT

Ba RKTHBH3HpyIOT /J.BM/KeHHe H peryJIHPYIOT npOHHKHOBeHHe cnepMaT030H,ll;OB B .aiiu,eKJieTKY.

2. <Pa3a aKmueau,uu.

IIpottcxo,n;HT KOHTaKT cnepMaT030tt,n;a H JIHll,eKJieTKH.

B o6oJioqKe Jittll,eKJieTKH p.a.n;a mttBOTHhIX cy~ecTByeT OTBepcT11e - MHKponnJie, qepe3 KOTopoe npOHHKaeT cnepMaT030H,ll;. y 60JihIIIHHCTBa BH,ll;OB
MHKPOilHJie OTCYTCTByeT H npOHMKHOBeHne cnepMaT030H,D;a
eTCJ:l 3a cqeT

aKpoco.M1toit peaKu,uu.

ocy~eCTBJIJ:l

IIo.n; .n;eiicTBHeM cpepMeHTa (rnaJiypo-

HH,n;a3LI) H .n;pyrHX 6HOJIOI'HqecKH aKTMBHhIX Be1u;ecTB, OCB06om.n;aIO~HX
CJ:l H3 aKpOCOMhl crrepMaT030M,ll;a, pa3pyrnaIOTCJ:l o60JIOqKJ1 J:llill,eKJieTKM.
AKpocoMHaJI HHTh rrpom,rnaeT qepe3 pacTsopeHHYIO 30HY JIHll,eBLix

CrrnJLrnat111~ di.
cnep~IJTOJOH~!a

c nepm1•11w11 /
060.10'1KOl1

\

flpmnn.:.uone1mc

'1CpC3

ncpnn'lHYIO

IlrraJMJHl 1fCCKaR --

MeM6paHa
si:HucKJJeTJG.f'

Ilepe11•-mafl - - - - o6oJ101..fKa

•

Jlapo H'1ltCK.rJCTKl1

"""T'

Pr.tc. 25. CxeMa npm.iecca on110,nornopeH111H: 1-4- nocne.11oearenbHb1e crn.11111111 npol\ecca

06UfaR 6uo11,ozua . Ocno6bl, 1tumo11,ozuu

72

o6o.TioqeK u cJiuBaeTCH c MeM6pattoM: .Hiiu.eKJieTKH.

B noM MecTe H3 U.HTO-

rrna3Mhl Hiiu.eKJieTKH 06pa3yeTc.H BocnpttHHMmorn;ttif 6yropoK.

3. <l>a3a npoHUfCH06f'llUJl.

IlpOMCXO,ZJ;HT npOHHKHOBeHMe rOJIOBKH H KJieToq-

HOro u.ettTpa B .HifU.eKJieTKY.

B

aaBMCMMOCTH OT KOJI11qecTBa npom·m aro-

.Mo11ocnep.Mu1<J (nponoJLucnep.MUIO (B U.HTOnJia3MY

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CKa~ Me3o,aepMa

(6)

me.rw6.Ttacmbt -

HaqttHaroT ,n;eJIHThCH H ,n;aroT Haqa-

JIO Me3o,n;epMe.

2. 3Hmepou,e.!lbl-ibtu

cnoco6 xapaKTepeH

,n;JIH

BTO-

pttqHopoThIX. KJieTKJI, cpopMHPYIOII.I;Jie Me3o,n;epMY, o6oco6JIHIOTCH B BH,n;e KapMaHOB nepBJiquoii:
KlllllKJI. IloJIOCTJI KapMaHOB npeBpaII.I;aIOTCH B

.!lO.M.
KH -

u,e-

Me30,n;epMa .n;eJIJITCH Ha OT,n;eJihHhre yqacT-

COMUmbt,

H3 KOTOphIX o6pa3yIOTCH onpe,n;e-

JieHHhie TKaHJI JI opraHhI. IlocJie o6pa3oBaHHH Me3o,n;epMhI HaqHnaeTcH npor.r;ecc rJicTo-

H

opraHoreHe3a.

CHaqaJia cpopMttpyroTcH oceBhie opraHhI

-

HepBHaH Tpy6Ka, xop,n;a. 3aTeM Bee OCTaJihHhie.

H eupy.!lRl(UR -

aTo o6pa3oBaHue 3apo.n;hmrn c oceBhIMH opraHaMH: HepBHan Tpy6-

Ka, xop.n;a, KllllleqHaH Tpy6Ka. 3apo,n;hilll Ha 3TOH CTa,ll;Hll Ha3hIBaeTCH

y

Heupy.wu.

JiaHI.I;€THIIKa ll3 3KTO,n;epMbI Ha CITJIHHOH CTOpOHe 3apO,D;bIIlla 06pa3yeTCH

Hep6Ha.fl mpy61w.

OcTaJihHan 3KTo,n;epMa cpopMJipyeT KOiKHhIM anuTeJIJIM JI ero

xopiJa.
Me.JoiJep.Ma

npOJI3BO,ll;Hhie. l13 3HTO.n;epMbl rro.n; HepBHOM Tpy6KOM cpopMJipyeTCH
Ilo,n; xop,n;oii: Haxo,n;MTCH

COMUm06,

ICUlUe'lHaR mpy61Ca.

IIo 6oKaM OT xop,n;hr -

KOTopaH ,ll,McpcpepeHr.r;JipOBaHa CJie,n;yIOII.I;JIM o6pa30M: HapymHaH qaCTb

iJep.Momo.MO.M. I13 Hee cpopMMCIC.!lepomo.M - 06CKJiepoTOMOM Haxo,11;HTCH .Muomo.M, ,11;aro-

COMMTa, npHJieraIOII.I;aH K 3KTo.n;epMe, H83hIBaeTCH

pyeTCH CO€,ll;HHJITeJibHaH TK8Hb KOiKM. BttyTpeHHHH qacTh paayeT CKeJieT. Mem,11;y ,n;epMOTOMOM n

II.I;MH HaqaJio nonepeqHO-IIOJIOCaTOH MYCKyJiaType. Ilo,11; COMJITaMJI HaXO,ll;HTCH
HOiKKJI ( He<Ppomo.M), JI3 KOTOpblX cpopMHpyeTCH MOqeIJOJIOBaH CHCTeMa. l_{e.!lOMU'leCICUe MPlUICU cpopMMPYIOTCH CJIMMeTpwrno TIO 6oKaM. CTeHKM II.eJIOMHqeCKHX M€IllKOB, o6paII.I;eHHble B CTOpoHy KHIIJeqHHKa, Ha3bIBaIOTCH cn.!lal-iXl-iOll.!le6pou, B CTOpoHy 3KT0,11;epMbl - COMamon.!le6pou. 8TJI JIMCTKH yqaCTBYIOT B o6pa30BaHHH cep,11;eqHo-cocy,11;HCTOH CHCT€MbI, IIJieBpbI, 6pIOIIIMHhI, rrepJIKap,11;a.
IIocTpoettHe OT,ll;eJihHhIX oprattoB npoHCXO,ll;HT 3a cqeT npeo6paaoBaHMH aaPO,ll;hIIlleBhIX JIHCTKOB 3KTo,11;epMhI, Meao,11;epMhI H 3HTo,11;epMhI.
8Ta RJiaCcHcpmcar.r;HH HOCJIT ycJIOBHbIH xapaKTep, TaK I-l:aR B IlOCTpoeHJIJI
RaiK,11;oro opraHa yqacTByeT HeCKOJibKO 3apO,ZJ;bllll€BbIX JIHCTKOB.

flocm3M6pU01tll/lbltOe pa:wumue

79

IlpoH3BO;'lHbie 3ap0,1bIIIIeBbIX JIHCTIWB

··---·

""'""'"" ~

Meao11epMa

aKTO,ll;epMa
HepBHa.a Tpy6Ka

,Il;epMa;

3mtTemtif cpe;:i;uei1: KlfllIKif,

(roJIOBHOll If CIUIHHOll

coeti;IfHifTeJihHall TKaHh;

me,'Iyti;Ka;

M03r);

CKCJ'ICT;

1rnw;eBaplfTC,'1bHbie iKCJIC3bl

noKpOBbl TCJ'Ia

Mb!Illllhl;

(rreqeuL,rro;:i;meJiy;:i;oqHaJI

(ami:;:i;epMMC If ero

KpOBCHOCHall, JllfM<lJaTif-

iKCJ1C3a);

rrpOM3BOtJ;Hble: HOrTM,

qecKall, BbltJ;CJIMTeJibHall,

3IllfTCJ1Ifll JierKifX lf

KOrTif, rrepbll, BOJIOCbl,

IlOJIOBall ClfCTCMbl;

B03tJ;YXOHOCHh!X rryTett;

3MaJib 3y6oB, KOiKHbie

u;eJIOM, IIJICBpa, 6pIOlIIMHa,

rrepe;:i;mrn:, cpe.l(HRR ,l(OJIM

lf MOJIOqHblC iKCJIC3bl

neplfKap;:i; If ;:i;p.

rlfrrO<lJMaa; lll;lfTOBiftJ;Hhie,

HT. ti;.);

rrapaw;IfTOBiftJ;HhIC iKCJIC3hl,

opraHhl apeHnH, cJiyxa,

TlfMYC lf ti;p.

o6oHllHMll;
3ap;Hllll ti; OJI JI rMrro<PMaa,
3Illf<lJM3;
rrepeti;Hnli M 3MHMli
OTtJ;CJibl KMIIIeqHMKa

IIocTaMopnoHaJibHOe pa3BHTHe
Pa3JUPiaIOT ,n;Ba nrna rrocTaM6pttoHaJibHOro pa3BMTmr:

1. HenpH.Moe pa3eumue -

HenpH.Moe

II

npH.Moe.

JiMqirnoqHoe, xapaKTepn3yeTc.H HaJinqtteM o.z:1-

HOH mm HecKOJihKMX JIH'IMHO'IHhIX cTa,n;nil:. Ilponcxo,n;MT c MeTaMop<Po30M (xapaKTepHO .D;JI.H HaceKOMbIX) (p1IC.
IlpM

28).

llO.!lHO.M .Mema.Mop<jJo3e II3 aH:u;a BbIXO.D;HT JIHqHHKa,

OTJIHqaroiu;a-

HC.H OT B3pOCJIOH oco6H HaJIIIqMeM crreu;MaJibHblX opraHOB. 0Ha IlMTaeTC.H,

a -

B.ipocJtaJl rpop.Ma
6 -

R3poc.wR rpop.Ma

PMc. 28. MeraMopq,03 y HaceKOMb1x: a - ttenOJTHbll'i MeraMopcPo3 y K06b1JTK11; 6 - nonHbJYi MeTaMOP<1>03

Y 6a6o~K11

06ui,aR 6uo;wzuR. I'enemu1w

80
pacTeT, JIHHJJeT II rrpeBpa~aeTCH B KYKOJIKY

(y

6oJiblIIHHCTBa HeIIO,ll,BIIJK-

HaH cTa,'l;HJI). OpraHhI JIWIIIHKH pacTBOpmoTCH, coxpam1eTc.H TOJlhKO HepBHaH CHCTeMa, 3aqaTKH IIOJ10BbIX JKeJie3 II n:viarHHa.;IbHhie ,ll,IICKII, 3a C'leT
KO'l'Opb!X cpopMIIPYIOTCH opraHhl B3pOCJIOro HaceKOMOro. lfa KyKOJ'IKH 06pa3yeTCH B3pOCJiaH oco6h II

IIMaro. Ilp11Mep: IKYKII, MYXH, 6a60'IKII, KOMapbI

llP·
51ihrO-')> Jlll'lIIHKa

Ilpn

---+

KyKOJIKa

---+

n:Maro

He no.rt HOM MemaMoprpoae Il3 Hliu;a BbIXO,ll,HT Jin:qnHKa, IIOXOJKaH Ha

B3pOCJIYIO OC06h, HO MeHhlllllX pa3MepoB. JlH'lllHKa .'IHHHeT HeCKOJlbKO
rrpenpa~aeTCH

pa3, ,ll,OCTHraeT 3peJIOCTil Il

BO B3p0CJ10e HaceKOMOe.

Ilpn:Mep: KY3He'IIIKII, TapaKaHhI n: .n.p.
51li:u;o

2. Jlp.HMOe pa.wumue

---+ JIM'lHHKa ---+

nMaro

rrpon:cxo,ll,HT B ,n;nyx cpopMax.

He.rtU'lUIW'lHa.H rfJopMa (pb16b1, perrTn:Jin:n, IITHIJ;hI). 51H:u;a aTnx JKMBOTHhIX 6oraThI JKeJITKOM • .[(JIH ,ll;hIXaHMH, BhI,ll,eJieHHH, IIMTaHMH eCTb BpeMeHHhie (rrpOBII30pHb!e) opraHhl.
51H:u;o

---+

n:Maro

BHympuympo6Ha.H rpopMa (MJieKorrnTaro~n:e, qeJioBeK). Bee cpynKu;n:n:
3apO,ll,b!IIIa

ocy~eCTBJIHIOTCH

cneu;11aJihHoro oprana -

3a C'leT opraHII3Ma MaTepn: c

IIOMO~bIO

nJiau;eHThI. XapaKTepuayeTca pa3BIITHeM npoBM30pHhIX (npeMeHHbIX) oprattoB: JKeJITO'lHbIM MellIOK, aMHMOH, xopHOH, aJIJiaHTOMC.
Xopn:oH II aJIJiaHTOIIC cpopM11pyroT nJiau;enTY (pIIc.

11

29).

lfayqenne 3M6pHOHaJibHOro Il IIOCT3M6pIIOHaJihHOro pa3BHTHH JKMBO'l'HbIX .n.a.10 B03MOJKHOCTb HaHTH 06~11e qepTbI B 3THX npo-

3aKoH
cxoocmea 3apOObtllleU (I\. Bap): «3M6pHOHhl

u;eccax 11 ccpopMyJinpoBaTh 3aKOHhr:

IIMeIOT 06~11e CXO,ll,CTBa B npe.n.eJiax THIIa Ha'lHHaH c CaMbIX paHHllX CTa,n;nii pa3BHTIIH»;

PMc. 29.

CxeMa aapogb1wa npV1Mara,

pa3BVIB3IOUJ,erocs:i BHYTPVI OKp~alOUJ,VIX
era

2noM;

4 -

6 -

HOlKKa

XOPVIOHa;
1..\eHra;

3 -

aapogb1w;

BHe3apOAblW8Bbl~ 1..1e-

aMHVIOH;

nonoCTb;

7-

1

o6onoYeK:
XOPVIOH;

5 -

aMHVIOTVIY8CK3Sl

nynoYHbl~

annaHTOVICa; 8 -

g -

10 -

KaHan1K;
BOpCVIHKlil

MaTepVIHCKa!l TKaHb nna)1(8nTOYHbl~

11 - lKenTOYHbl~ MeWOK

CTe6eneK;

6uoieHemu'lecKuu

.JaKon

(3. feKKeJih,

<I>.

MIOJIJiep):

«0HToreHea ecTh KpaTKoe

H

6hrcTpoe

noBTopeHn:e

cpn:Jiorene3a,

TO eCTb llH,ll,IlBII,ll,yaJihHOe pa3BllTIIe opraHII3Ma IIOBTOpHeT IlCTOpllIO pa3BHTIIH CBOero BII,ll,a». Pa6oTbI

A.H.

Cenepu;ona noano-

JIMJIII ,ll;OIIOJIHIITb II yTO'IHIITb 3TOT 3altoH:
«3apO,D;blllI B CBOeM OHTOreHe3e npoxo,n;n:T
cpopMbl He B3pOCJibIX npe,n;KOB, a IIX aapo,ll;hillleH ».

OcnoBnwe nonRmuR u onpeiJe;ienuR ienemuKu

81

rEHETlllKA
OcHOBHLie noH.HTHH H onpe,IJ;eJieHHH reHeTHKH.
3aKOHLif.M:eH,IJ;eJI.H

I'ettemuKa

(OT rpe'-I.

genesus -

ttayKa, rr3yqa10~aa 3a-

rrporrcxom.n;emrn) -

KOHOMepHOCTH HaCJie,n;CTBeHHOCTH lI H3MeH'-IlIBOCTH opraHlI3MOB.
KaK B .i'II06oii TO'-IHOH nayKe, B reHeTIIKe cy~eCTBYIOT CBOH 'IeTKlie IIOHaTHH, orrpe,n;eJieHHH H 3aKOHhl, KOTOpbIMH OHa orrepn:pyeT.

HacJLeacmBeHttocmb -

cBoiicTBO opraHH3MOB rroBTopaTh B pa,n;y rr0Ko.i1e-

HHH cxo,n;Hhie rrpH3HaKH rr o6ecrre'-IrrBaTh crreu;mpH'-IecKHH xapaKTep HH,!l;HBHµ;yaJihHOro pa3BHTHH B orrpe,n;eJieHHhIX ycJIOBHHX cpe,!l;hl.

H 3MeH'lUBOCmb -

crroco6ttocTh opraHlI3MOB H3MeHHTh CBOH rrplI3HaKH H

CBOMCTBa.

I'eH -

CTPYKTyptto-$yHKIIHOHaJibHaa eµ;HHHI1a ttac.'le,n;cTBeHHOCTH.

MH'-IecKoil: TO'-IKH 3peHHH

zeH -

C

xrr-

3TO yqacTOK MOJieKyJihl ,[(HK, KO,!l;lipyIO~HM

HH$OpMaI1HIO 0 KaKOM-JIH6o 6eJIKe (rromrneIITH)J;e).
COBOKYIIHOCTh reHOB, co,n;epma~HXCH B raIIJIOH,!l;HOM Ha6ope xpo-

I'eHOM -

MOCOM KJieTKH.

/(HK -

µ;e30KCHpH60Hy1rneHHOBaa KHCJIOTa -

MOHOMepoM KOToporo HBJIB:eTCH HYKJieOTH,D; (pHC.

HyKJLeomua

MaKpOMOJieKyJia, IIOJIHMep,

30).

= caxap + a30THCTOe OCHOBattrre + OCTaTOK <PoccpopHOH KHCJIOThI.

(IIHTHyrJie-

(A,n;eHHH,fyaHHH,

5 °C;

THMHH, l!HT031IH)

pO,D;HhlH -

p

pH6oaa HJIH ,n;eaoKcttptt6oaa)

.D;HK Koaupyem UH<jJOpMGU,UIO -

TpHIIJieThl HyKJieOTH,ll;OB orrpe.n;eJIHIOT IIOCJie-

,ll;OBaTeJibHOCTh aMHHOKHCJIOT B IIOJIHIIeIITH.n;e.

AJiJLeJlb -

Ta IIJIH HHaa pa3HOBH,!l;HOCTh (aJibTepHaTHBHaa <PopMa) reHa.

OH

CH,
0

OH

H

OH

H

-cH,

~·.
~

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0

I

I

OH

H

P1tc. 31. Hy1<J1eo311A- 6e3 ocTaTKa
c)>occ)>opHOiil Kl!ICnOTbl

06UfaR 6uo;wzuJi. I'enemu1w

82

AnJieJihHhie reHhI pacnonomeHhI B o,D;:mrnKOBhIX JIOKycax roMoJiori1qHhIX
xpoMOCOM (OTBeqaJOT 3a pa3HOBH,ll;HOCTb 0,!J;HOrO rrpH3HaKa, o603HaqaJOTCJI O,ll;HOM 6yKBOM).

I'oMO.!l02U'lHble xpOMOCOMbl -

rrapHbie xpoMOCOMhI, O,ll;HHaKOBhie no pa3Me-

paM, <Pop:-.ie, cop;epmaHHIO reHOB (pttc.
MHoJ1Cecm6eH1tbtu a.fl.fle.flu3.M -

32).

cym;ecTBOBaHtte reHa B BH,n:e HeCJW.JihKHX

anneneM: y s11,11;a H rronynmi;1111. B rrpOIIecce 3BOJIIOIIHH B03HHKaeT B pe3yJihTaTe
H3MeHeHHJI nocne,D;osaTeJibHOCTH HYKJieOTH.D;OB s reHe. IloJIBJIJieTCJI s nony;rnIIHH opraHH3MOB, TOr,D;a KaK y O,ll;HOro ,ll;llrnIOll,ll;HOrO opraHH3Ma cym;eCTByeT
TOJibKO ,D;Ba aJIJieJIJI O,ll;HOro reHa.
IlpuMepbI MHOJKeCTBeHHOrO aJIJieJIH3Ma:

1. Tptt
IA; IB; I 0
y

anneJIJI

rpynrr

y nonyJIJIIIHH,

-

KposH

I 0 IA

s

H ,D;p. -

cttcTeMe

A

B

0

y Bllp;a.

O,D;Horo opraHll3Ma 3a ITPll3HaK OTBeqaJOT ):\Ba aJIJieJIJI (MaTepu H OTIIa).

Hanp11Mep, rpynrrhI RposH

I (0) 0
II (A)

III

reHoTHilhI

I0 I 0

MeM6paHhI 3pHTPOIIHTOB «rOJibie»

o-< A

Ha MeM6paae aHTHreH A

IA IA IA IO

B

Ha MeM6paae aHTHreH B

IB pi IB IO

(B)CK

c:\~

IV (AB)

Ha MeM6paae aHTHreHhI A H B

2.

IAIB

AJIJieJIH ORpaCRll rnepCTH y RpOJIHKOB.

I'OM03U20mHOCmb -

B rOMOJiornqHbIX xpOMOCO-

MaX IIpHCYTCTBYIOT ,D;Ba O.D;HHaROBhIX aJIJieJIJI O,ll;Horo
reHa:

AA HJIH

aa.

I'emepo3u2omnocmb -

B rOMOJIOriftIHhIX xpoMoco-

Max rrpncyTCTBYIOT ,D;Ba pa3HhIX aJIJieJIJI O,ll;HOrO reHa:
Aa.

I'aMembt -

IlOJIOBhie KJieTRll opraHH3Ma, HMeIO-

m;tte rarrJIOll,ll;HhIM Ha6op xpoMOCOM.

«qHCThie

JIHHHH » -

JIHHHH pacTeHHH H JKHBOT-

HbIX, noJiyqeHHhie B pe3yJihTaTe ,ll;JIHTeJihHOrO caMOOIIhIJieHHJI H.JIH 6.JIH3KOpO,ll;CTBeHHOro CKpem;nsaHHJI.

I'u6puOO.!l02U'leCKUU .Memo() 5
PMc. 32. roMonorwlHble xpoMOCOMbl c annenbHb1M111 reHaMll1:

1-

napa roMonorw1HblX

XPOMOCOM;

2 - nOK)'C; 3 -

annen111; 4 COCTO~H111e;

MeTO,ll; CRpem;nsa-

HllJI.

rGM03111rDTHOe

5 -

rDTHOe COCTO~Hllle

reTepo3111-

3HaJI reHOTHII opraHH3MOB H

raMeTbI, KOTOpbie

yqaCTBYIOT B OilJIO,ll;OTBOpeHHH, MO)KHO rrpep;cKa3aTh
H npoaHaJIH3HPOBaTh, qTQ IlOJiyqHTCJI B IIOTOMCTBe.

I'u6puo -

oco6h, noJiyqeHHaa B peayJihTaTe CKpe-

m;ttsaHHR.

I'enomun -

CllCTeMa B33HMO)J;eHCTBYIOm;Hx reHOB

opraHH3Ma (RJieTKH).

3aKOH bl I'. 111 ettiJe.nR

83

!Ipa6ll/IO 6blflUCbl6QllUR ?QMem: '-IMCJIO THilOB raMeT, 06pa3yeMhIX oprmma-

MOM, onpe;1,eJIHeTcH no cpopMy.'le

AA

1)
raMeTb1:

4

Aa

2)

n =

O; 2°

n =

1; 2 1 = 2

=

2",

r,n:e n -

1 TMn

'-IHCJ10 reTepo3MroT.

@

raMeT

@

THna raMeT

©~

cw
AQBb

3)

AaBbCc

4)

CITD

@@

n. = 2; 22 = 4 TMna raMeT
(fil!reTepo3uroTa)

n =

3; 23 = 8

0
@ @

THilOB raMeT

(TpHreTep03HrOTa)

3aKOHLII'.M:eHJ.J;e~H
3aKOHhI 6hIJIH OTKphITbI B

1865

r. aBCTpaJIMHCKMM ecTeCTBOMCilhITaTeJieM

fperopoM Mett,n:eJieM. «BHOBh OTKphIThI» H no,n:TBepm,n:eHhI B

1901

r. roJIJian,n:-

IJ,eM fyro ,n:e <I>pn30M, 3aTeM HeMeIJ,KHM yqeHhIM KapJioM KoppeHcoM n aBcTpnii:IJ,eM 3pHXOM cpoH qepMaKOM. 3aKOHhI 6bIJIH BhIBe,n:eHhI:
c HCIIOJih30BaHH€M rn6pn,n:oJiorM'-I€CKoro MeTo,n:a (CKpell.l,HBaHirn);
B pe3yJihTaTe HCIIOJih30BaHH.R «qHCThIX JIHHHH» pacTeHHH ropoxa, He ,n:aroll.J,MX pacll.],eIIJICHHR B IIOTOMCTBe rrpu caMOOilhIJ1€HHH M MMeIOll.l,MX ,n:Ba roMOJIOrMtIHhIX retta;
c ucnoJih30BaHneM cTpororo MaTeMaTnqecKoro no,n:cqeTa n attaJin3a.

I

u

II

3QICOHbl ocHo6aHbl HQ M01wiu6puiht0M c1Cpew,u6QHuu (aHaJIH3 i-rncJie-

.n;oBaHM.R no O,ll;HOMY rrpH3HaKy).

III

3QICOH oc1wBaH HQ iJuiu6puiJHOM u noJtuiu6puiJHoM c1Cpew,uBQHUU (oco-

6u OTJIM'IaJIHCh no ,n:ByM HJIH HeCKOJihKHM rrapaM rrpH3HaKOB).

I aaRoH -

aaKOH e;::i;ttttoo6paam1 rtt6ptt;::i;oB I rroKOJieHHH

n!'rPH3H3H
A
a

iK€JIThIH (,n:oMnHaHTHbIH)
3eJieHhIH (peu;ecCHBHhlH)

P:
G:
Fi:

~AA x

®
Aa

aa

cf'

0
100 'Yr)

<PopMyJtup061(Q 3Ql(OHa: rrpH CKpell.J,HBaHHH ,n:Byx oco6eii:, pa3JIM'Ial0ll.J,HXCR

no O,lt;Ho:H nape aJibTepHaTHBHhIX rrpH3HaKOB, B rrepBOM IlOKOJieHHH y rn6pn,n:OB

06Uj-aK 6uo1wzuK. I'enemu1w

84

Ha6mo,n;aeTC.H e,n;HH006pa31le rrptt3HalWB H npo.aBJI.HeTCfl ,Zl;OMHHaHTHhlH npn3HaK.

II 3aKOH - 3aKoH pacm;eIIJieHH»
P: (_jl Aa x er Aa

@0 @0

G:

F 2:

AA Aa Aa

3 m.

aa

: 1 3.

<Pop.MyJLup061ca 3alW1ta:

rrpH CKpern;irnaHHH rH6pH,D;OB rrepBoro IlOKOJieHHfl

Mem,n;y co6oft BO BTOPOM IlOKOJieHHH IIOHBJifllOTCfl oco6H KaK c ,ll;OMHHaHTHhIMH,
TaK H c pe:a;eCCHBHbIMH npH3HaKaMH; npOHCXO,Zl;llT pacrn;ern1emrn no Q.JeHOTHny
B COOTHOllleHHH

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Cq,en.nennoe nac.nedoBanue. l'enemu'tec1we onpedenenue no.na.

~umozenemu1w

91

IletteTpaHTHOCTb H :3KCnpeccHBHOCTb HBJIJUOTCH pe3y.'IhTaTOM BJIHHHMH BCero reHOTttna H cpe,n;h1 Ha onpe,n;e.'JeHHhllI reH.

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1. OpiaHU3.MeHHblU - OCHOBhl 3aJIOiKeHbl r. Mett,n;eJieM B 1865 r.
2. KJtemo'lHbtu, mrn u,umoienemu'lecKuu - ocHOBhI 3aJiomeHhI B ;rn6opaTOpHH T. Mopratta B 1910 r.
3. MoJteKyJt.H,PHbtu - Ha'!aJI pa3BHBaThCH c OTKphITHeM CTPYKTYPhI MOJieKYJihI .JJ:HK (1953 r.).
K Ha'IaJIY xx B. c pa3BHTHeM MHKpOCKOilH'IeCKOH TeXHHKH HaKOilHJIOCb MHOmeCTBO ,n;aHHhIX o noBe,n;eHHH xpoMOCOM. B 1902 r. Y. CeTTOH conocTaBHJI noBe.n:eHHe xpOMOCOM H nplrnHaKOB npH MHT03e H Meiio3e H o6HapyiKHJI, 'ITO XpOMOCOMbl H npH3HaKH «BeJIH ce6H» aHaJIOrH'IHO. BnocJie,n;cTBHH yqeHhiii npe,n;noJIOiKHJI,
"llTO reHhI Haxo,n;HTCH B xpoMocoMax. K 1910 r. B Jia6opaTopnn aMepHKaHcKoro reHeTHKa T. MopraHa B pe3yJihTaTe onhITOB Ha nJio,n;oBoii MYIIIKe ,n;po3ocpHJie 6h1Jia
ccpopMyJittpoBaHa xpo.11toco.11tna.H meopu.H nacJte8cm6eHHocmu.
OcHOBHLie noJio:meHHH xpoMOCOMHOH TeopHH uacJie,n;cTBeHHOCTH

1. HacJie,n;cTBeHHhie e,n;HHHllhI, HJIH reHhI, JIOKaJIH3YIOTCH B xpo¥OCOMax.
2. Kam,n;hlii reH B xpoMocoMe 3aHHMaeT onpe,n;eJieHHOe noJiomeHHe - JtoKyc.
feHbl B xpOMOCOMe pacnOJIOiKeHhl JIHHeHHO.
3. feHbl, Haxo,n;mu;neCH B O,D;HOH xpOMOCOMe, HacJie,n;yIOTCH c11enJieHHO H o6pa3yIOT rpynny c11erIJieHHH. qHcJio
rpynn c11enJieHHf! paBHO ranJIOH,n;HOMY Ha6opy xpoMocoM.
4. Mem,n;y roMOJiorH'IHhIMH xpoMocot
MaMH B03MOiKeH o6MeH yqacTKaMH - 1CpOCCUHZ06ep (pHC. 36).
5. PaccTOf!HHe Mem,n;y reHaMH H3MepHeTcH B MOpraHH.n;ax HJIH npo11eHTax KpoccnHroBepa: 1M=1 % .
6. PaccTOf!HHe Mem,n;y reHaMH nponop11ttoHaJihHO npo11eHTaM KpocI \
I \
CHHroBepa, a CHJia c11enJieHHf!
Mem,n;y reHaMH o6paTHO nponop11nOHaJII:.Ha paCCTOf!HHIO Mem,n;y HHMH.
B Jia6opaTopnH MopraHa 61:.1JIH npoBePi.tc. 36. Kpocci-1Hrosep: 1 - ABe roMono.Zl;eHhl pa3JIH"l1Hble :3KcnepHMeHThl.
rw1Hb1e xpOMOCOMbl; 2 - l'IX nepeKp8CT BO
OnhIThl MopraHa, ,n;oKa31:.rna10I11He
BP8M5l KOHb10rau.i-1i-1; 3 - AB8 H0Bbl8 KOM61'1cn;enJieaaoe aacJie,n;oBaHHe (pttc. 37):
HaLJ,l'!H

'®

2JW~

QD 00
<DCD CDCD

06UfaR 6uo1wzu11.. I'enemuica

92

!'•.
I I

h• • b

t>

I I

., • • v

' •• v

"i ib
~·

•

v

6 r,
h •• h

BA ••

v •• "

vl:j

I I

:tfr

.
'
-1-~
..,,

!v

~

50'7"

HeHpoccoeepbl 83

'-----v--Hpoccoaepbl 17 %

%

a
PMC.

37.

CL1en11eHHOe Hac11eAOBaH11e reHOB 0KpaCKl1 rnna 11 COCTOS1Hl1Sl KpblllbeB y nllOAOBOiil Myx11

(Drosophila melanogaster): a coM y caMuoe AP030¢lL111bl: pp KOKpblllblM caMuOM
KpblllOiil caMKOiil
TOMKOB:

6

(BBW) CKP01J.10Ha c YepHblM KOpOT-

(bbw); Fi - cepbliil caMeu c AJ111HHblMl1 Kpb11lb5lMl1 (BbVv) CKP01J.10H c YepHOiil KOPOTKO-

(bbw); F2 -

50 % -

no11Hoe cuen11eH11e reHOB scneACTe11e orcyrcre115l nepeKpecra xpoMocaMKa cepa5l c AJ111HHblM11 Kpb11lb5lMl1

nOCKOJlbKY y caMua He npo11CXOAl1T KpOCC11HrDBepa, n05lB5lTC5l ABa B11Aa no-

YepHblX KOpOTKOKPblJlblX 11

50 % -

c AJll1HHblMl1 KpblJlb5lMl1
AJ111HHblM11 KPb11lb5lM11

6 - HenOJlHOe
y caMOK AP030¢>11Jlbl: pp - CaMKa

cepblX c HOPMallbHblMl1 KPblJlb5lMl1;

(YaCrn'IHOe) cuen11eH11e npl13HaKOB BCJ10ACTBL1e nepeKpecra xpOMOCOM

(BBW) CKpelJ.leHa c yepHblM KOPOTKOKPblJlblM caMuOM (bbw); F1 - cepa5l caMKa c

(BbVv) CKpelJ.leHa c YepHblM KOpOTKOKPblllblM caMUOM (bbw); F2 - noCKOJlbKY y caM-

K11 npo11CXOAl1T KPOCC11HrDBep rDMOJ10rl1'1HblX xpOMOCOM, o6pa3ylOTC5l YeTblpe rnna raMeT 11 n05lBSlTC5l YeTblpe B11Aa nOTOMKOB: HeKpoccosepbl -

cepble c AJ111HHblMl1 KPblJlb5lMl1

(BbVv) 11 YepHble KOpOTKOKPblJlble

(bbw), Kpoccosepbl - YepHble c AJll1HHblMl1 KPblJlb5lMl1 (bbVv), cepble KOPOTKOKpblJlble (Bbw)

rea
B

b (black)

v

v (vestigial)

II waaaK
cepoe TeJio
qepaoe TeJio
HOpMaJibHbie
KpbIJib.H
aaqaToqab1e
(KopoTKHe)

B H V aaxo,n;HTCH

Bn 7
V

'%

(M)

B

I aTan

P:

G:

o.n;aoii xpoMocoMe

:.,B+tB x b t

v-~ -t-v

·tb/

v T i-v

c:v ~
B++b

F1:

v_l
I

-t_

IV

cepoe TeJio,
HOpMaJihHble
KPhlJiblI

93

Clfen11,ennoe nac11,ea0Banue. I'enemu'lec1we onpeae11,euue no11,a. l{umozeuemu1w

II aTaII
Peu;u11pmrnoe c11:pem;uBa11ue - CKpern;:rrnamrn caMu;OB (a) n caMoK (6)
c peu;ecCHBHOH rOM03HrOTHOH oco6bIO.

a)

____;__ b

B_ I

P:

G:

I

vr-

I

1

b ) ~b

x
v

.

I

--v ·-,.I

6)

.

B +---!

P:

V

~ (5) ~

G:

B+ j-b

b-t

+b

vl -t v

v~-

lv

v+-1 v

-r-·

v1

'lepnoe TeJio,
KOpOTKHe
KpbIJib.H

50 %

50 'X,

II

v
y cf ,n;po30!l>HJibl IIOJIHOe cu;e11Jie11ne (HeT KpOCCHHrOBepa) =>
reHbl HacJie,n;yIOTCH cu;e11JieHHO.
B pe3yJibTaTe aToro rroJiyqHJIHCh TOJibKO po,n;nTeJihCKHe
coqeTaHHH reHOB.

iV

Q) ~ CD
83 '%

neKpoccoaepttwe -

(!D

w
17 %

KpoccoeepHbie --cepoe TeJio,
HOPMBJibtthle
Kpb!Jlb.H

rb

-

-

F2:

b bl
x

F2:

BT

r

ht

v+ . v

cepoe TeJio,
HOpMBJlbHbJe Kpb!Jlb.H

j-b

vi- +v

'lepttoe TeJJo,
KOPOTKHe Kpb!Jib.H

HeKpoccoeepw -

83 %

bl
VT
cepoe TeJio,
KOpOTKHe Kpb!Jib.H

qepttoc TeJJo,
HOPMBJibHble Kpb!Jib.H

Kpoccoaepw --

17 %

Y <.J? ,n;po3o!l>HJibI cu;errJieHne 11e110JI11oe.
IIpou;eHT 11:poccn11r0Bepa 3aBHCHT OT
paCCTOHHHH Mem,n;y reHaMH.

06~a11,

94

6uo1wzu11,. I'enemu1w

f eHeTHqec1we orrpe.IJ;eJieHHe rroJia
c;IO)KHhIH rrpH3Ha:K, HacJie,D,OBaHHC KOToporo orrpe,D,CJUieTCH pa3BI1-

[fOJl -

THeM 20/WO.
Bhr,D,eJrnJOT HeCKOJihKO THITOB orrpe,D,eJieHHH rroJia B 3aBHCHMOCTH OT tIHCJia
11 cocTaBa 110.'IOBbIX xpoMOCOM (pHc.

1.
2.
3.

38):

1IeJJOBeK, MJieKOilHTarow;tte H ,D,p.

XX

Bo,D,ano:i1: KJIOII Protenor
HeKOTOphre 6a6otIKH,

IITHIIhl, pbI6bI, 3eMHOBO)J,Hhie

4.

XY

XX

IlepenoH'IaTOKphIJihie

XO

zo
zw

zw
zz

2n

n

(32 xp.) (16 xp.)
Il'Ie,'Ihl

foMoraMeTHhIH IlOJI -

TpyTHH

O,ll,HHaKOBhie xpoMOCOMbI.

feTeporaMeTHhlH IIOJI -

pa3Hhle XpOMOCOMhI.

Y qeJioBeKa orrpe,D,eJiarow;ttM B pa3BHTHH rroJia HBJIHeTcH Ha.'IHtIHe Y-xpoMocoMhl. B Hett, rro-BH,D,HMOMy, Haxo,D,HTCH KJIIO'lesoii: reH, ttrparou~n:i1: pOJih peryJIHTopa. Y MyTaHTOB rrptt JIJ06oM KOJIHtieCTBe X-xpoMocoM ,D,06aBJie1rne Y-xpoMOCOMhI IIPHBO,D,HT K cpopMHPOBaHHIO MymcKoro cpeHOTHrra.
XXY XO -

CHH,D,pOM KJiaiiHcpeJihTepa.
CHH.D.POM IIJepuiescKoro-TepHepa (

XXXoY~
XYY

J

2 ).

71

0
Cu;em1eHHoe c IIOJIOM HacJie;i:i;oBaHHe

IlpU31W1C cu,enJLell c nOJLO.M -

reH, orrpe,D,CJIHIOUJ;HH pa3BHTHC oTOro rrpH3Ha-

Ka, HaXO)J,HTCH B IIOJIOBOH xpoMOCOMe.

x -

cu;errJieHHhie npll3HaKH.

y-

cu;err-

JICHHhie rrpH3HaKH (roJiaH,D,pnqec-

x x

x y

11

I1

,~~"CD

~

r!f

ii~.~
x x

x y

Koe HaCJie,D,OBaHMe OT OTIJ;a K ChlHy).

BoJihIIIMHCTBO

6oJie3HeH -

rrpH3HaKOB

peIJCCCHBHbie.

l.J;HTOreHeTHKa

Kapuomun -

aTo Ha6op xpo-

MOCOM B COMaTH'leCKOH KJieTKe
opraHH3Ma ,D,aHHoro BH,D,a. XapaKTepn3yeTC.H

rrpaBMJiaMH

XpOMO-

COM:

P111c. 38.

XpoMoCOMHbli'i MexaH1<13M o6pa-

P: Cf' 44 ayrocOMbl +XX x 44 ayrocOMbl + XY d'

G: 22 ayTOCOMbl

rrpaBHJIO IIOCTO.HHCTBa 'lllCJia
xpoMOCOM;

30BaH1-tfl nona:

'/>< ;; :::~~~~~: +

X

v~+Y

F 1: 44 ayrocOMbl + XX 44 ayrocoMbl + XY

rrpaBHJIO rrapHOCTH roMOJIOrMqHhIX xpoMOCOM (y )J,HilJIOH,ll,HblX
BH,ll,OB; 0,D,Ha xpOMOCOMa IIOJiyqeHa
OT OTIJ;a, ,D,pyraH -

OT MaTeptt);

Cq,en;iennoe nac;ieiJoBanue. I'enemu"'ec1we onpeiJe;ienue no;ia. L{umozenemu1w

95

TipaBirnO IIH,D;IIBII,D;yaJibHOeTII - OTipe,n;eJieHHOe eTpOeHIIe XPOMOeOM (pa3Mep, cpopMa, co,n;epmaHIIe reHoB);
TipaBIIJIO HeTipepbIBHOeTII - Tiepe,n; ,n;eJieHIIeM KJieTKII .IJ:HK y,n;BaIIBaeTeJI TIO
npIIHD;IITIY KOMnJieMeHTapHOeTII.
Bee xpoMoeoMhI Tio,n;pa3,n;eJIJIIOT Ha ayToeoMhI II TIOJIOBhie xpoMoeOMhI.
II OJL06bte - aTo xpoMoeoMhI, OTipe,n;eJIHIOID;IIe cpopMIIpoBamrn MyJKeKoro
Ii meHeKoro TIOJIOB.
AymocOMbt - Bee xpoMoeoMhI B KJieTKax, 3a HeKJIIOqeHIIeM TIOJlOBhIX.
IlpuMep:
1) KapIIOTIITihl qeJIOBeKa - 46 xpoMoeoM, 23 napbl xpOMOeOM
22 TiaphI ayToeoM +XX IIJIII XY; 22 A+ XX; 22 A+ XY;
2) KapIIOTHTI ,n;p030cpIIJihl - 8 xpOMOeoM;
3) KapIIOTIITI aeKapH,D;hl - 2 xpoMoeOMhI;
4) KapIIOTIITI eo6aKII - 78 xpoMoeoM.
B 3aBHeIIMOeTII OT paeTioJiomeHIIH u;eHTpoMephl xpoMoeoMhI 6hrnaroT:
1. Memau,eHmpu'lecKue - paBHOTIJieqHe.
2. Cy6Memau,eHmpu'lecKue - HepaBHOTIJieqIIe.
3. AKpou,eHmpu'lecKue - TianoqKoBII,D;Hhie.
4. TeJtou,eHmpu'lecKue - u;eHTpoMepa paeTioJiomeHa Ha KOHu;e
xpoMOeOMhI.
PaaBIITIIe MeTo,n;oB oKpaeKli II MIIKpoeKoII
nIIpOBaHIIH TipliBeJIO K TIOJIBJieHIIIO MeTo,n;a
au<P<PepeHu,UaJlbliOZO OKpautu6aHUH. Kam,n;aJI
xpoMoeoMa IIMeeT eBoii: IIH,D;HBli,n;yaJihHhIH pIIeyHOK paeTionomeHIIH ayxpoMaTIIHOBhIX II reTepoxpoMaTIIHOBhIX yqaeTKOB.
B 1968 r. mBe,n;eKIIii: D;IITOJior T. KacTiepeeoH
IV
e KOJIJieraMII, a B 1971 r. IIapIImCKaJI KOHcpepeH~H KJiaCCIIcpIIu;IIpOBaJIII MeTO,ZJ;hl CTaH,n;apTII3a~H xpOMOeOM (paa6HBKa TIO rpyTITiaM): BCero
Ce.Mb zpynn - no BeJiffqHHe II TIOJIO;KeHHIO u;eHTs
poMepbI (pHe. 39).
Ilpwnep.
1. fpyTina A - N2 1, 2, 3 - KpynHhle MeTaD;eHTpHqeeKIIe.
2. X-xpoMoeoMa - rpyTina C, epe,n;HHH
cy6MeTau;eHTpIIqeeKaJI.
PMc. 39. <t>opMbl xpoMocoM:
3. Y-xpoMoeoMa - rpyTITia G, MeJIKaJI aKI - TenoL1eHTpw-1ecKafl; II - aKpOpou;eHTpHqeeKa.H.

Yponuu KOMIIaKTHaau;uu (y1tm1,a:KH) xpoMocoM

B PMHhie cpaahl KJieToqHoro D;HKJia (HHTepct>aaa HJIH MHT03 (Meii:oa) xpOMOeOMhl HMeIOT
Pa3Hoe CTpoeHHe.

L1eHTp1114ecKafl; Ill -

cy6MernL1eHT-

p1114ecKafl; IV - MernL1eHTp1114ecKafl.

1 - L\eHTPOMepa; 2 - cnyrHlllK; 3 KOPOTKOe nne40; 4 - AJJlllHHOe nne40; 5-xpoMaTlllAbl

06UfaH 6uo1wiuI£. I'enemu1<a

96

B npou;ecce HHTepcpa3hI B KJieTKe 6o"'lhIIJaH qacTh HacJie,n;cTseirnoro MaTepnaJia aKTHBHa, 1103TOMY ,n;ecnHpaJIH30BaHa H Ha3hIBaeTCH ayxpoMamUftOM;
qacTh HeaKTHBHoro AHR KOMIIaKTHO co6paHa H 06pa3yeT zemepoxpoMamu ft
(HarrpHMep, TeJihu;e Bappa - KOMIIaKTH30BaHHaH X-xpoMocoMa y S? ).
IlpH ,n;eJieHHH H3 xpoMaTHHOBhIX HHTeH o6pa3yIOTCH xpOMOCOMhl.
y qeJIOBeKa ,l(JJHHa xpoMOCOM B 10 ThlC. pa3 MeHbllle HCXO,l(HOH ,l(JIHHhl MO"'leKYJihl AHR.
Bh1,n;eJIHIOT HeCKOJihKO ypoBHe:H KOMrraKTH3aUHH AHR:

1. AHR (d

=

2 HM).

~
2. HyKJieocoMHhIH -

~
3. CoJieHoH,n; -

I2HM

«6ycw no HHTKe» (d"" 10 HM).

IllHM

«CIIHpaJih» (d"" 20 - 30 HM).

I

30HM

4. «IleTJIH»- HHTepcpa3HaH xpoMOHeMa (d"" 100 - 300 HM).

I
I

I

300 HM - neTJIH na qm6pttJIJibI
300 HM -

c6mnKeHne neTeJib

5. MeTacpa3HaH xpoMaTH,n;a (d"" 500- 600 HM)- BH,l(Ha B CBeTOBOH MHKpocKon.

~

I

700 •• - OAHaxJ>O•''""

1400 HM -

xpoMoCoMa na ,11syx xpoMaTH,ll

RoMnaKTH3aUHH xpoMocoM Mo:m:eT HCIIOJih30BaThCH KaK peryJIHTOP aKTHBHOCTH reHOB: CIIHpaJIH30BaHa - He aKTHBHa, ,n;ecnHpaJIH30BaHa - aKTHBHa.
IlpuMep. 0,n;Ha H3 ,n;Byx X-xpoMocoM y meHin;HH B HOpMe He aKTHBHa => crrnPa.T.IH30BaHa, 06pa3yeT reTepoxpoMaTMHOBoe TeJihu;e Bappa (rroJIOBOH X-reTepoxpoMaTHH).
Y meHiu;MHhl c KapMOTMIIOM 46, XX - 1 TeJihu;e Bappa.
Y My:m:qHHhl c KapnoTHIIOM 46, XY - 0 TeJieu; Bappa.
Y My:m:qnHhI c cnH,n;poMOM RJia:HBcpeJibTepa 4 7, XXY - 1 TeJihu;e Bappa.
II oJtumenna.R. xpoMocoMa - xpoMocoMa, o6pa3YIOJUaHCH B pe3yJihTaTe
MHOroKpaTHOH peIIJIHKal.l;HH HHTH ,ll;HR 6e3 ee pacxom,n;eHHH. B pe3yJihTaTe o,n;-

97
Ha xpoMOCOMa MO:>KeT COCTOHTh H3 6oJiee

,UHIC

1

ThIC . .11,Byxr~eIIO"Y:e'!HbIX M0.1eKy.1

TaKtte xpoMOCOMhr BCTpe'IaIOTCH B KJICT·Kax CJIIOHHhIX me.1e3 ttaceirn-

MbIX, B KJI€TKax HX MaJibIII1I'I1€BhlX cocy,n:on.
IloJittTeHHbie xpOMOCOMhI xopowo Bll.11,Hhl B CB€TOBOH MttKpOCKOil, Korn;a
KJieTKa HaXO,ll;ttTCH B HHTepcpa3e. feHeTHKH I1CIIOJih3YlOT 3TI1 XpOMOCOMbl ,ll;.'1.H
cocTaBJieHHH reHeTH'l€CKHX KapT H, CJie.11,0RaTe.1hHO, .11,JIH onpe,n:eJieHHH TIOJIO;KeHHH reHOB B xpoMOCOMax.

MOJl EKYJlSI PHASI rEH ETll1 KA
MaTeptta.'lhHhIM cy6cTpaToM HacJie.11,cTBeHHOCTH H tt3MeH-Y:HBOCTH HBJIHIOTca
HYKJiettHOBhre KttCJIOThI. PaaJittY:aIOT ,n;sa Btt,n,a HyK.1ettHOBhrx KttcJioT: llHK
(,n,eaoKcttptt6oHyKJieHHOBaH KHCJioTa) H

PHK (ptt6oHyKJieHHOBaH KHCJioTa).

XnMtt-Y:eCKH Hatt6oJiee ycTOM-Y:HBa .IJ:HK. MoJieKyJia .UHK HBJIHeTCH cy6cTpaTOM HaCJie,D,CTBeHHOCTH H
PHK

-

H3M€H-Y:HBOCTH B

60JihillHHCTBe :>KHBhIX CHCTeM.

ocHOBa HacJie,n,cTBeHHOCTH y MHornx Bttpycos (snpyc Ta6a-Y:HOM Moaa-

HKH, peTpOBttpychI II03BOHO-Y:HhIX).

,r:(HK

H

TH,D,hI (pHC.

PHK 40).

3TO noJIHMephI, MOHOMepaMH KOTOphIX HBJIHIOTCH HYKJieo-

Coe,n;ttHeHHe HYKJieoTtt,n,oB B u;errtt rrponcxo,n;ttT aa cqeT o6paaoBamrn cpoccpo3cpHpHoil: CBH3H Mem,n,y OH-rpyrrnoil:

(*)

tt cpoccpaTHOM rpyrrrroil: cne,n,y:ioru;ero

HYKJieOTH,n,a. CxeMy cTpoeHHH .IJ:HK CM. B paa,n;ene

« 11;ttTOJIOrHH » •

Ha rrpHHIJ;HIIe KOMIIJieMeHTapHOCTH OCHOBaHhI MeXaHU3Mbl MampU'lliOZO

cu11.me3a:
1) PenJLu1cau,uR .UHK (y,n;soeHHe).
2) TpattCl{:punu,uJl - CHHTea PHK Ha MaTpttu;e .UHK.
3) TpaH.CMlU,UJl - CHHT€3 6eJIKa Ha ptt6ocoMe.

-

aaOTHCTbie OCHOBaHHH

(A,n;eHHH,THMHH,fyaHHH
IJ;HT03HH)

*OH
IieHT03a - ,11;e30KCHptt603a (HJIH p11603a)

06UfUR 6uoJWZUR. I'enemux:a

98

Pe1rnHI>:al\HH

5

,IJ;HK

.U.mr Bcex ay1rnpwon1qecKttx opraHH3MOB
penJIHKaU:MJI .II.HK rrpottc2
XO,[\MT IlOJiyKOHCepBaTMBHhIM Mes·
TO,l\OM.
II oJLy1wncepBamuBllbtit Me moo
pen.1lu1cau,uu 3aKJIIOqaeTCH B TOM, '!TO
IlOJiyL!eHHhie ,z:i;oqepHHe MOJieKy.'Ihl
5
.II.HK
co,z:i;epmaT O,D;HY u;eIIOLIKY OT Ma6
TepttHCKOM MOJieKyJihl .II.HK (KOHCep5'
BaTlrnHyro), a BTopyro - BHOBh CMHPr.tc. 41. Penm1Kau,w1 ,[\HK: 1 - ,[\HK-renviTe3ttpoBaHHyIO.
Ka3a; 2 - ,[\HK-rnno1113oMepa3a; 3 - PHKPenJLu1cau,ua ,[(HK cJioJKHhrii:
npaViMa3a; 4 - t:1ecw6111n1113111py10ll.(111e 6enrn;
rrpou;ecc, rrpottcxo,a;aru;ttil: B a,a;pe
5 - ,[\HK-nOJlV1Mepa3a o6pa3yeT ,[\HK, YAJ1Vlc y'laCTMeM ¢epMeHTOB.
H5l8T ¢parMeHTbl 0Ka3aKVI, y):lanHeT p11160HyKOcuoBHhre ¢epMeHThr, yqacTByroneoTV1Abl; 6 - ,[\HK-nma3a cwvisaer ¢parru;tte B penJIMKau;tttt .Il;HK, M MX pOJih
M8HTbl 0Ka3aKVI
(pttc. 41):
.IJ;HK-reJittKa3a - pacnJieTaeT crrttpaJih;
.Il;HK-TOIIOM30Mepa3a - pacKpyqttBaeT crrttpaJih;
.IJ;HK-rroJittMepa3a - ocyru;ecTBJIHeT cttnTe3 ,z:i;oL!epueil: u;errtt;
.Il;HK-Jittra3a - 4CUIMBaeT» cpparMeHThI 0Ka3aKM •
.IJ;oqepmrn u;errh .Il;HK CMHTe3ttpyeTCH TOJihKO B HarrpaBJieHHM 5' ~ 3'.
JluiJupy10w,aa u,enb - MaTepttHCKaa u;enh .Il;HK, na KOTopoil: tt,z:i;eT uerrpephIBHhIH CMHTe3 C 3' ~ 5'.
3ana3iJbl6a10w,aa u,enb (OTCTaroru;aH) - MaTepHHCKaH u:enh .Il;HK, Ha KOTOpo:H M,D;eT npepbIBMCThIH CMHTe3 c o6pa30BaHHeM cpparMeHTOB 0Ka3aKM M KaJK,l\hIM <lJparMeHT HMeeT HanpaBJieHHe c 3' ~ 5'.

I

ori

ori

ori

-

:60
on

Pr.tc. 42. Penn111KaU,V15l y 3yKapvior. Penn111KaL.\V15l 111H111u,111111pyercH s cneu,111¢111YeCKOM yYacrKe ,[\HK,
Ha3b1eaeMOM T04KOVi «Ha4ana" pennviKau,111111 (ori). Pacryll.(vie u,envi o6pa3ylOT penm1KaTV1BHb1e s11nK11.
CVIHT83 HOBblX u,eneVi ,[\HK npOVICXOAVIT B penn111KaH18HblX BVIJlKax, nepeMell.(alOll.(VIXC5l B ABYX
HanpaBJl6HVl5lX

peanuJaUrufl zenemu<tec1wu u1upopMaituu

PeIIJII1KaI~IUl

1aiem1w. liuocunmeJ 6e1rn:a

99

y :JyrrnpHO'l'OB rrpo11cxo,n;H'l' B HeCKOJlhH:HX MCCTax Ha Kam.n;oi1:

3'1'0 noJtupen.llUIWH/lblU MeXallU.:iJlt.

xpoMOCOMe -

y

6

rrpoKapHOT, HMelOI.QHX KOJibUeByIO '.\1.0JieKYJIY ,UHK, pacrrpocTpaneH JIW-

HOpenJlU/COHHblU .11exaliU.:iM, T. e. Cl1HTe3 Ha'UII!aeTC.H B O)J;HOH TOqKe (p11c.
PeaJIH3aD;HH reHeTH11ecKoH:

42).

Hm:l>opMan;HH B KJieTKe.

6HOCHHTe3 6eJIKa
Bee Mttoroo6pa3tte 6e.>IKOBhIX MO.TieKy.1 B JKHBhIX opraHI13Max 3aKo.n;wposaHO B MO.WKYJiaX ,UHK (PHK).

I'eHemu'leC/CUU /COO -

3TO IIOCJ1e.n;oBaTeJihHOCTh TplIIIJieTOB HyKJ'IeOTlI)J;OB B HYltJie-

HHOBbIX KlIC.'IOTax, 3a.n;a10ru;a.H COOTBeTCTBYIOI.QMH rrop.H.n;OK IL'VIMHOKlICJIOT B 6e:max.
Ilpe.n;rrOJIOJKeHMe 0 cyrr~eCTBOBaHMlI reHeT11qecKoro KO,n;a 6bIJIO BhICKa3aHo
B

1954

6hIJI

r. aMepHKaHCKMM cpH3HKOM r. raMOBhIM. B

pacwwcpposau

C. Qqoa H

113
43 =

X.

aMepHKaHCKMMH

1959

r. reueTifqecKHH KO,[(

M.

6MoxMMMKaMH

Httpeu6eproM,

Kopauo:i1:.

qeThrpex HYKJieOTH)J;OB, KOM6MHIIpy.H MX ITO TpM, MOJKHO COCTaBHTb

64

TpHIIJieTa. }fa

64 TpHrrneTOB TpH .HBJI.HlOTC.H 6eCCMhICJieHHhIMH, HJIH
- OHH He IIIMcppylOT aMHHOKHCJIOTy; rrpH cqHThIBa-

«HOHCeHC-TplIIIJieTaMH »

HHlf MHcpOpMaUHH Ha ypoBHe M-PHK BbIIlOJIH.HlOT cpyHKUHIO CTOII-KO,[\OHOB TpttnJieThI ATT, AD;T, ATIJ; (s M-PHK HM coOTBeTCTBYIOT Ko,n;ouh1 Y
TaJihHhie TpHn.1eThI

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(61)

Yr A, Y Ar). Oc-

KO.UIIPYIOT onpe.n;e-

JieHHbie aMHHOKMCJIOThI.

lbepHaJ!

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CBoiicTBa reueTn11ec1wro Ko11;a

1. y HHBepcaJibHOCTh -

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+

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raHH3MOB (3a HeKOTOpblMM MCKJIIOqeHH.HMlf -

2.

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.UHK
PHK

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0

0()

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3.

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1

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JIOTa, KPOMe MeTHOHMHa, KO.UHpyeTC.H He-

OpouecCHHr PHK

c:::::::=::J

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cqHThIBaIOTC.H KaK OT,[\eJibHhie caMOCTO.HTeJibHbie e,n;HHHilhI.

4.

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l

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pery JIHPYIOIIl;HX TpaHCJIRUHIO.

B CJIOJKHOM

npouecce peaJIH3aUHM reueTwqe-

CKoli HHcpOpMaIIHH BhI,D;eJIRIOT qeTbipe OCHOBllLIX aTana (pMc. 43): mpaiicKpunu,uJO, npou,eccunz, mpaHcJtau,uJO, nocmmpancJtau,uonnbte
U3MeHeHua 6eJtKa.

P1o1c. 43. CxeMa npouecca sKcnpecc111111

reHeTl!14eCKO~

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06UfaR 6uo;wzuR. I'enemuwa

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,

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All,Y, All,ll,, All,A, AU,r

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AcrraparHHOBafl KIICJIOTa

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I' JiyTaMIIHOBafl KIICJIOTa

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--

·-

TpaHcKpmrn;m1

TpanC1cpunu,uH - aTo CHHTe3 PHK Ha MaTpmi;e ,n:HK.
B rrpou;ecce TpaHCKPHIIIJ;HH yqaCTByroT MHOrHe ipepMeHTbI, KOTOphie ocy~eCTB
JUUOT pernrnKau;mo (.Il:HK-reJIHKa3a, .D:HK-TorroH30Mepa3a, PHK-rroJIHMepa3a).
B CJIOJKHhIX 6HOCHHTeTHqecKux rrpou;eccax TpaHcKpHIIU:HH H TpaHCJI.Hll;HH
BhI,n;eJI.HIOT HHHD;Hall;HIO, :3JIOHrau;uIO lI TepMHHall;HIO (puc. 44).
Hnuu,uau,u.H. - CB.H3hIBaHHe PHK-rroJIHMepa3hI (ipepMeHTa, ocy~ecTBJI.HIO
~ero CHHTe3 PHK) c rrpoMoTopoM H o6paaosaHHe rrepso:H CB.H3H Mem.n;y .n;syMa
HYKJieoTu.n;aMH. IIpoMomop - yqacToK ,n:HK, co.n;epma~HH BhlCOKOKOHcepsaTHBHYIO rrocJie,n;osaTeJihHOCTh HYKJieOTH.n;os TATA, KOTOphIH CJIYJKHT .n;Jia yaHaBaHH.H H CB.H3hIBaHH.H c PHK-rroJIHMepa3oif.

101

,llsoAHaR crmpam. JlHK
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He-rpmcKpH6H PYeMIUI uem.

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u

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uerm JlHK

BHOBL

coeAHHRIOTCR

:3.Jtonza4uJ1 -

yp;mrneHHe rroJIHHYKJieOTHp;Hoii n;errH PHK. 8JI0Hran;m1 PHK

IIPOHCXO)J;HT )J;O TepMHHHpy101n;e:H TpaHCKPHIID;HIO IIOCJie)J;OBaTeJihHOCTH.

Tep.Mutta4uJ1 -

oKoHqaHHe CHHTe3a PHR.

B pe3yJihTaTe TpaHcKpHrrn;HH
1. p-PHK (pu60CO.MHaJ1):

o6pa3yIOTCH HeCKOJihKO BH)J;OB PHK:

a) CHHTe3ttpyeTCH c rroMOlll;hIO PHK-rroJIHMepa3hl

I;

6) o6pa3yeTCH Ha yqaCTKax .IJ:HK B 30He H)J;pbllllKOBOro opraHH3aTopa HJIH
B o6JiaCTH BTOpHqHoii rrepeTH:>KKH (xpOMOCOMhI:

13, 14, 15, 21, 22);

B) HBJIHeTCH CTPYKTYPHhIM KOMIIOHeHTOM ptt6ocoM;
r) o6ecrreqHBaeT CBH3hIBaHHe ptt60COM c HYKJieOTH)J;HOll IIOCJie)J;OBaTeJihHOCTblO M-PHK (IIOCJie)J;OBaTeJihHOCTb HYKJieOTH)J;OB lllaiiHa-.IJ:aJibrapHa B JIH)J;epHOM yqaCTKe y rrpoKapHOT H aHaJIOrHqHaH y 3yKapHOT).

2. m-PHK (mpancnopmttaR):
a) CHHTe3HpyeTCH c rroMOlll;hIO PHK-rroJIMMepa3w

III;

6) TpaHCIIOpTMpyeT COOTBeTCTBYIOlll;Me aMMHOKHCJIOTbl K MeCTY CHHTe3a
6eJIKa, T. e. K pH60COMaM;
B) B

ll;HTOIIJia3Me KJieTKH o6Hapy:>KHBaIOTCH pa3JIHqHbJe T-PHK )J;JIH

TpaHCIIOpTa

20 aMHHOKHCJIOT;

06UfUH 6UO/WZUH. I'enemu1w

102

r) I<aiK.[(OM T-PHK COOTBeTCTBYCT CBOM crreu;mpw-1ec1rn::il: cpepMeHT KO,[\a3a
(aM1rnoar~IIJI-T-PHK-cIIHTeTa3a). Ko.na3a Y3HaeT aHTIIKOAOH T-PHR:
II rrpucoe,n,IIHHeT K ne:il: ttymtty10 aMIIHOKIICJ!OTY.
CTpoettIIe T-PHK: pa3Mep 75-100 HYKJICOTtt,n;oB; HMeeT KOMITJieMeHTapHhre y'laCTKII, 3a cqeT KOTOpbIX 06pa3yeTCH CTPYKTypa « KJieBepHoro Jill CT a»'
IIJIH «TpIIJIIICTHIIKa»; K 3' -OH KOHIJ;Y rrpttcoe;:i;IIHHeTCH aMHHOKIIC.'IOTa.

3. u-PHK (wupopMaU,UOHHaR):
a) CIIHTe3IIpyeTc.H c rroMOI.IJ;hIO PHK-rroJIIIMepa3hI II;
6) 06pa3yeTCH Ha MaTpIIu;e Ko.noreHHOH u;errII. KoiJoieHHaR u,enb - o.n;tta H3
ABYX u;erre:il: B MOJieKyJie .IJ:HK, KO.[(IIpy10m;a.H orrpe.neJieHHbIM IlOJIIIrrerrTII.[(.
CTpoeHIIe II-PHK:
Y syKapIIoT: Ko.n;IIpy10m;IIe yqacTKH (3K30HhI), HeKO,n;IIpy10m;IIe yqacTKII (uHmpmtbt}, peryJI.HTOpHbre yqacTKII (JiuiJepnaR II mpeuJiepnaR no-

c;ieiJoeame,ib1wcmb ).

y rrpoKapIIOT B 60JiblIIIIHCTBe CJIY'laeB II-PHK COCTOIIT npaKTIIqeCKII u;eJIIIKOM II3 K0,1J;IIpy10m;eil IlOCJie.[\OBaTeJibHOCTII. feHhI rrpoKapIIOT He IIMeIOT IIHTPOHHOH CTPYKTyphr, HO co.nepmaT CBOII peryJI.HTOpHhie y'laCTKII.

4. MR-PHK (.MaJlaaaiJepnaR PHK):
a) co.n;epmIIT - 50-200 HYKJieoTIIAOB;
6) BXO,ll;HT B COCTaB M.H-PHK, ocym;eCTBJl.HIOI.IJ;IIX CITJiaHCIIHr;
B) co.nepiKIIT KOHIJ;eBbie yqacTKII, KOMITJieMeHTapHbie cailTy CITJiaHCIIHra
(Bbipe3aHII.H) IIHTPOHOB.
Bee cosoKyrrttocTII MOJieKyJI PHK B .H.[(pe 06pa3y10T rryJI - reTeporeHHYIO
.H,IJ;epHyIO CIICTeMy PHK (r.H-PHK). B cocTaB STOro nyJia r.H-PHK sxo.n;.HT TaKme
MOJieKyJihI PHK, no.nsepra10m;IIec.H «C03pesaHIIIO», IIJIII npou;eccIIHry nepe.n;
BbIXO,ll;OM II3 .H.ll;pa B IJ;IITOITJia3MY.

II po:u;eccHHr ayKa pHoT111rncKHX H- PHK
Ilpou,eccuni - 3TO «C03peBaHIIe» nepBII'IHOrO TpaHCKpIInTa II-PHK II npeBpam;eHIIe II-PHK B M-PHK, KOTOpoe BKJIIO'IaeT CJie.ny10m;IIe 3Tanhr:
1. CnJiaucuni - Bhipe3attIIe IIHTpOHOB II CIIIIIBaHIIe SK30HOB (HTO - HeTpaHCJIIIpyeMhie o6JiaCTII, JIII,ll;epHa.H II TpettJieptta.H).
CnJJailcHHr
.
3
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A.rrbTepHaTIIBHhIH CITJlaMCIIHr - cym;eCTBOBaHIIe pa3Jlll'IHhlX cnoco6os Bblpe3aHII.H IIHTpoHoB II3 o.nttoil II TOM me II-PHK. B pe3yJihTaTe o.n;Ha II-PHK MomeT o6pa3oBaTh HeCKOJihKO pa3HhIX M-PHK. llcnOJih3yeTC.H ,ll;Jl.H YBeJIII'IeHII.H pa3H006pa3II.H CIIHTe3IIpyeMblX 6eJIKOB, HanpIIMep ,ll;Jl.H o6pa30BaHII.H M-PHK pa3Jill'-IHblX aHTIITeJI.
2. K:mupoeanue - 3aI.IJ;IITa 5' KOHu;a M-PHK (3a c'leT npIIcoe.n;IIHeHIIa K HeMY ryaH03IIHTpmpoccpaTa B npoTIIsonoJIOiKHOM HanpaBJieHIIII).
3. IloJiuaiJenuJiupoBanue - 3am;IITa 3' KOHu;a M-PHK nyTeM npIIcoe.n;IIHemrn u;enO'IKII II3 100-200 HYKJieOTII.[(OB A.

103

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1

B 3peJ10il .11-PHK Bhr.n;eJunoT K8II, TpattcJrnpyeMyro o6JracTh (CIIIHThre B o.nHO u;eJioe ;:)1{30Hbl), HeTpaHCJIMpyeMLie ofornCTM (HTO) M ITOJrna.n;eHMJ10BhIH
«XBOCT».
4. MemuJtupoBaHue - M3MeHeHtte rrepBbIX nyKJieoT.11,noB Tpanc:KpttrrTa.
Ilpou;ecc1rnr 3aBepIIIaeTc.H coe,nttHeHIIeM M-PHK co crreu;mpwi:ecKMMU 6eJIKaMM M BhIX0,11,0M B UMTOIIJia3MY :KJ!eTKlf qepe3 .11,11,epHbie rroph!.

TpaacJI.mi;m1
TpaHc.Mtl{U.H - aTo rrpou;ecc 6IIocttHTe3a rronIIrrerrTII,11,HOH u;errII Ha MaTpIIu;e
M-PHK, rrpoIIcxo,11,.11m;IIii B pII6ocoMax (rrepeBo,11, «.H3hIKa» HYKJieOTII,11,0B Ha
«H3hIK» aMIIHOKIICJIOT).
Ilpou;ecc TpaHCJIHIJ;IIII MOJKHO pa3,11,eJIIITb Ha TPII :3Tarra (pttc. 45):
1. HHu4ua4ua - c6op:Ka pII6ocOMhI (Tpe6yeT rrpwcyTCTBIIH Mg' II cpaKTopoB TpaHCJIHIJ;IIM):
rrp.11coe,11,IIHeHIIe MaJIOH cy6'be,ll,MHIIIIhI pII60COMhI I{ M-PHK;
B3aIIMo,11,eiicTBIIe nepBoro (cTapToBoro) Ko,notta M-PHK AYr c T-PHK,
Hecym;eii: aMIIHOKIICJIOTY MeTMOHIIH;
rrpIIcoe,11,IIHeHIIe 6oJihIIIOii: cy6'he,ll,IIHHIJ;hI.
2. aJt0Hia4u.H - y,11,Jinnenne noJI11nerrT11,nttoH: uerrn:
HatIHHaeTCH c o6pa30BaHHH rrepBoii rrerrTH.l(HOH CBH3H MeJK,ny aMIIHOKHCJIOTaMn;
3a:KaHtIHBaeTCH rrpn «npOtITeHHM» IIOCJie,11,0BaTeJihHOCTII M-PHK .no
CTOII-Ko.n;ona na PHK;
OCHOBHOH cpepMeHT B pII6ocoMe, yqaCTBYIOIQIIH B o6pa30BaHIIII rrerrTII,Zl;HOH CBH3II, - rrerrTII,11,IIJITpaHccpepa3a.
3. TepMuHau,ua - 3aBepIIIeHIIe CIIHTe3a:
rrpoIIcxo,n;i1T y3HaBaHIIe cTorr-Ko,noHa (YAA, YAr, YrA);
I{ IIOCJie,nHeii: aMHHOKIICJIOTe B IIOJIIIIIeIITII.l(e rrpncoe,nIIH.HeTC.11 BO.n;a,
II OHa OTIIJ;eIIJIHeTCH OT T-PHK;
rrenTII,11,Ha.H uerrh OT,ll;eJIHeTCH OT pII60COMhl;
PII6ocoMa pacrra,11,aeTc.11 Ha .n;Be cy6'he.l(IIHIIIJhI.
IlOCTTpaHCJUID;HOHHbie H3MeHeHH.H 6emwB

HeK0Toph1e CIIHTe3IIpoBaHHhie 6eJIKH He aKTIIBHhI 6ea rrocJie,nyrom;ero «coapenaaII.11» - Mo.n;IIcpIIKaIJIIII. HarrpIIMep, IIHCYJIIIH IIcxo,11,no cIInTeaIIpyeTc.11
B KJieTKax no.n;JKeJiy,11,0LIHOH JKeJie3bl B BII.n;e ,ll;JIIIHHOrO ITOJIIIIIeIITII.n;a rrpe-npoIDICyJIHHa, nocJie oTm;errJieHIIH N-KoHu;eBoii, 3aTeM cpe,nneii qacTII nerrTII,na on
npeapam;aeTC.H B IIHCyJIIIH.

06utaR 6uo.1wzu1i. I'enemu1w

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DpllCQeJOIHHT1, HOByto

MOlleJcyny MentOHHHa

PHc. 45. CxeMa fa10c11HTe3a 6enKa Ha p116oc0Me
Perymu~un

pa6oTLI renoB (renHaH aKcnpeccu11)

Ilp01capuombt.

l.JTO «BKJIIOqaeT» HJUI «BblKJIIoqaeT» reHhI? IJoqeMy Haqlf-

HaeTCH TpaHCKpHIIUHH, a 3aTeM If TpaHCJUfUHH? 3ttaHJfe OTBeTOB Ha 3TH BorrpoCbl II03BOJIHJIO 6w pelIIHTb MHOrHe npo6JieMbI, CBH38HHhie c pa3BHTHeM If 6oJie3H.HMH opraHH3Ma. OJJ;Hoii H3 rrepBhix nonhITOK perneHMH aToro oqeHb CJiomHoro Bonpoca cTaJio co3JJ;att1:1e ~wae.1tu

B 1961

r. cppaHUY3hI

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E.coli

(KmrreqHaH rraJioqKa) B oTcyTCTBHe JiaKT03hI He CHHTe3H-

pyeT cpepMeHTbl JJ;JI.H ee IIOTpe6JieHHH. Ilptt JJ;06aBJieHHH JiaKT03bl Haq1:1HaeTC.H
TpaHcKpHITUH.H H TpaHCJI.HUH.H Hya<HhlX cpepMeHTOB (pHC.

46):

105

Pezy.1uHfUR pa6ombt zeno6

- JiaKT03a CBH3hrnaeTCH c 6eJIKOM-penpeccopo!'v1;
- penpeccop yxo.n.wr co csoero MecTa Ha MOJieKyJie ,[{HK (orrepaTope):
- oTKphrnaeTcH «ITYTh» .n;JI.H pa6oThr PHK-rroJinMepa3hr;
- CHHTe3HPYIOTC.H M-PHK H COOTBeTCTBYIOrnHe cpepMCHThl ,D;JLH paCIIJ,CII.TICHHH JiaKT03hI;
- JiaKT03a pacrnerrJI.HeTc.a;
- c pacrnerrJieHneM rrocJie.n;He:il: MOJieKyJihI JiaKT03hI 6eJIOK-penpeccop
BHOBh CTaHOBHTC.H CB06o.n;HhIM, «Ca,Z:VITC.H» Ha )lHK-onepaTop, 6JIOI\'.HpyeT TpaHCKPHTIIl,HIO.
IIpn .n.o6asneHHM "i:aKT03hI xo.n. co6hrTuM CHOBa rroBTOpHeTc.H. Be"10K-penpeccop Ko.n;wpyeT oT.n;eJihHhIM reH peryJI.HTop. 8Ta cxeMa pa6oThI reHa
y 6aKTepttn rro"1yqnJia Ha3saune ::mcnpeccttn.
Onepon - 3TO rpynna CTPYKTYPHhIX reHoB (Il,HCTpOHOB), uaxo.n.HrnnxcH rro.n.
KOHTPOJieM o6rnnx peryJI.HTOpHblX 3JieMeHTOB (rrocJie,n,oBaTeJihHOCTe:il: ,[{HK
H 6eJIKOB-peryJI.HTOpOB).
CuuTe3npyeMa.H M-PHK y rrpoKapnoT rroJIHil,MCTpOHHa, T. e. HMeeT tteCKOJihKO CTpyKTYPHhIX reHOB.
9y1Capuomb1. CnHTe3upyeMhre M-PHK .HBJI.HIOTC.H MaTpwu;aMn ,Zl,Jl:.H oT,n,e.TJ:hHhIX neIITH,D;OB, T. e. OHH MOHOil,HCTPOHHbie. KaJR,D;hlM reH B orrepoHe MMeCT
csoIO cucTeMy peryJI.Hil,HH. K peryJinpyIOIIVIM 3"i:eMeHTaM rrpou;ecca 3I<crrpeccun OTHOC.HT )lHK-rrocne.n.osaTeJibHOCTH H 6eJIKH-peryJI.HTOpbI.
)lHK-rrocne.n.osaTCJibHOCTH:
npo;;wmopbt - yCHJIHBaIOT TpaHCKpHIIIl,HIO B nec.HTKH H COTHH pa3;
3HXancepbt - HaXO,D;HTCH Ha 60JiblliOM paCCTO.HHHH OT rena;
CllUJl€HC€pbt ( «rJiyIIIMTeJIH») - HaXO,D;.ffTC.H Ha 3HaqnTeJibHOM paCCTO.HHHM, CHHJRaIOT TpancKpHTIIl,HIO.
BenKn-peryJI.HTOphr - 3TO 6eJIKOBhre q>aKTOphr TpaHcKpunu;ttn, KOTOphre
MoryT 6bITh o6rnMMH ,D;JI.H pa3HhIX rpyrrrr reHOB.
AKTMBHOCTh reHOB MOJReT TaKJRe perynnpoBaTbC.H rryTeM H3MeHeHM.H cTerreHH KOMrraKTu3au;nu xpoMaTuHa (crrnpaJIH3au;uu).
Penpeccop

CrpyKTVPHble reHbl
npaMOlOp
.. oneparop ,--------__,.,_----~

f-1oc1+P-0-+- /1eZ-+4cY+/acA-£
~~.
M-PHK

lac- M·PHK

/ac-penpeccopa

lac- penpeccop

fJ· ranlKTO:tHA33a

f3· ralUIKTOlMAnepMea3a

(J. ranaKT03M.ATP8HC81,leTMna~a

PMc. 46.

naKT03Hb1~

onepoH

06utaR 6uo.rwiuR. l'euemu1w

106

ME,D,"1Ll,"1HCKASI rEHET"1KA
Oco6eHHOCTH reHCTHKII 'IeJioBeKa cocTOHT B CJICJlYIOU~eM:
II03,llHee IIOJIOBOe coapeBaHIIe II ,llJIIITeJibHaH CMeHa IIOKO.'leHIIH (-

30 ;:reT );

Ma.'loe KOJIII'IeCTBO IIOTOMKOB;
C.'IOJKHhIH KapIIOTIIII (MHOro xpOMOCOM H rpynn CIIern:reHIIH);
Ha JIIO,llax aanpe~eHhI :mcnepmv1eHTaJihHhre 6paK11;
COIIIIaJ'lbHOe HepaBeHCTBO.

Cy~eCTBeHHhIM cpaKTopoM, 06JrerlJaI01I111M aa,lla'III II3y'IeHIIH reHeTIIKII 'IeJIOBeKa, CJiymaT MHOrO'IIICJieHHbie II MHOrOJ'leTHIIe ,llaHHbIC no aHa.1M3Y pa3JIII'IHhIX llplI3HaKOB.

OcHOBHLie MeTO.IJ;LI H3yqeHHH reHeTHKH 11eJI0BeKa

reHeanon11'i0CKMH M0TOA
MeTO,ll BBe,lleH B KOHIIe

XIX

B. aHrJIHHCKlIM y'IeHhIM

<I>.

raJ'lhTOHOM. OcHO-

BaH Ha COCTaBJieHHM lI aHa.'III3e pO,llOCJIOBHhIX. YcJIOBHhie o603HalJeHHJI BBe,lleHhl B

1931

r. r . IOCTOM (lICIIOJih3YIOTCH B HaCTOR~ee BpeMH) (pIIC.

D-0-0paK

rerepo311rorHbM!
IJ f)--HOCMTem•
pe4ec-

['.]

D=O-

KJlOBHOPOACTBeHHblii

0paK

4 7).

0
0 !-

CllBHOl"O reHa

n11'4HO o0clleAOBaHHblB

00-0-noeTopHblM OpaK

P111c. 47. YcnOBHb1e o6o3Ha'leH111si np111 coCTasneH111111 poAocnoBHblX nor. IOcTy

Meau1fu1tc1wR. zenemmw. I'enea1wzu'LeCKuii MemoiJ

107

Cyw;HoCTb MeTo.n:a:
COCTaBJ'IeHIIe pO,ll,OCJIOBHOH (rryTeM orrpoca) c lICIIOJib30BaHIIeM CHMBO-

J'IOB.

Ilpo6aHO -

qeJ'IOBeK, pO,ll,OCJ'JOBHYIO KOTOporo H3yqamT;

reHeaJIOrlfqecKHH aHaJIH3.

Il03BOJI.HeT BbIHCHHTb THII HaCJie,ll,OBaillf.H

MHOrHX rrpH3HaKOB.
OcHOBHhre THIIhI HacJie,D,OBaHH.H rrpH3HaKoB:

1. Aymoco.M1w-oo.MuHaHmHbLU mun Hac.rieooea1wJ1 (reH

Haxo,nHTC.H B ayTo-

coMe, .HBJI.HeTC.H JJ;OMHHaHTHbIM).
XapaKTepHhre rrpH3HaKH ayTocoMHo-,n;oMHHaHTHoro Tttrra HacJie,n;oBaHH.H:
npH3HaK rrpO.HBJIReTC.H B KaJKJJ;OM IIOKOJ'JeHHII (rro BepTHKam:1);
rrepe,ll,aqa rrpttanaKa (aa6oJieBaHH.H) OT po,ll,HTeJieil: ,n;eT.HM;
o6a IIOJia rropaJKaIOTC.H c O,ll,HHaKOBOH qacTOTOH.

Ilpu.Mepbl

HacJie,ll,OBaHH.H no ayTOCOMHO-,ll,OMHHaHTHOMY THrry:

CHH,ll,pOM Mapq:iaHa ( «rrayqhlf» rraJ'IhIIhI, aHeBpttaMa aopThI, ITO,ll,BbIBHX
xpycTaJIHKa);
TIOJIII,ll,aKTllJIH.H (rneCTb rraJihl!eB);
6paXH,D,aKTHJIHa (IWpOTKHe naJibIIbI).
lfHor,ll,a rrptt auaJittae po,D,OCJIOBHOH o6ttapyJKHBaeTc.H oco6a.H CHTYaIIH.H
,ll,OMHHaHTHbIH reH He rrpo.HBJI.HeTC.H, T. e. He rreHeTpHpyeT. IlpH 3TOM BHJJ; po,n;oCJIOBHOH MeH.HeTC.H.

IleHempaHmuocmb -

aTo qacToTa rrpo.HBJieHH.H reHa cpe,D,H HOCHTeJ'Jeil:

,n;aHHOro reHa, eCJIH reH rrpO.HBJI.HeTC.H, TO OH rreHeTpaHTeH. IleHeTpaHTHOCTb
npe,ll,CTaBJI.HeT co6oH:

OTHOlIIeHIIe oco6eH:,

HMeIOru;Hx ,ll,aHHbIH npH3HaK,

K oco6.HM, HMeIOru;HM ,n;aHHbIH reH. BhrpaJKaeTc.H B npoIIeHTax. HarrpHMep,
neHeTpaHTHOCTb ctttt,n;poMa Mapcpana CeTqaTKH) -

30 'X,;

peTI·rn06JiaCTOMhI (orryxoJih

80 '%.

ihccnpeccueHocmb -

3TO cTerreHb BbrpaJKeHHOCTH retta. CTeneHh BbrpaJKeH-

HOCTH IIPH3HaKa TaKJKe yqHThIBaIOT rrpH COCTaBJieHHH po,n;OCJIOBHOH.

2. Aymoco.MHo-peu,eccueHbtu mun Hac.rieooeauuJl

(ren ttaxo,n;HTC.H B ayToco-

Me, .HBJI.HeTC.H peIIeCCHBHbIM).
XapaKTepnbre rrpH3HaKH ayTocoMHo-peIIeCcMBHoro TMrra ttacJie,n;oBaHM.H:
npM3HaK rrpO.HBJI.HeTC.H He B KaJK,ll,OM ITOKOJieHMll (qaw;e npll 6JIM3KOpo,n;CTBeHHhIX 6paKax);
60JibHhie ,ll,eTH poJK,n;aIOTC.H OT cpeHOTllIIMqecKH 3,ll,OpOBbIX pO,ll,MTeJieH (HOCMTeJieH reHa);
o6a IIOJia HaCJie,ll,yIOT rrpH3HaK c O,ll,MHaKOBOH qacTOTOH.

Ilpu.Mepbt

HaCJie,n;oBaHM.H ITO ayTOCOMHO-peIIeCCMBHOMY THrry:

aJib6MHll3M (HapyrneHMe o6MeHa aMMHOKMCJIOT ll CllHTeaa rrMrMeHTa MeJiaHMHa);
cpeHMJIKeTOHypH.H (HapyrneH o6MeH cpeHMJiaJiaHMHa

:::> rrpOHCXO,ll,llT Ha-

KOITJieHHe B Moqe rrpo,ll,yKTOB ero OKHCJieHH.H, ,ll,eHCTByrom;Hx Ha IIeHTpaJibHYIO HepBHYIO CllCTeMy).

3. X-cu,enJleHHbtu oo.MunanmHblU mun uac.Tleooeanua
B X-xpoMocoMe, aBJI.HeTca ,ll,OMMHaHTHhIM).

(reH Haxo,n;HTC.H

06Ufa1l 6uo;wzun . I'e1temu1w

108

I3 X-xpoMocoMe ttaxo11,wren 60,1 ee 200 1·ettoll. Y meHII.urn aHOMa: cbHbIH re11
MOJKeT H8XO.ZU1ThC.H B 011,HOii (reTepo:rnroTa) IL111 B 06e11x (rOM03IffOTa) X-xpOMOcoMax; y Mymq11H TOJihKO B 011,noii: X-xp0Moco.:-.1e. Y MY:>K'll1H ucer.ua uptteyTeTBYeT TOJibKO o.n;MH a;:uieJib reHa (reMH3J.ffOTHOeTb), cuenJieHHoro c X-xpoMocoMoii:.
I'e.wu.wzomHOCmb - aTo npueyTCTB11e B opnrn11aMc TOJihH:o o,r.uroro a.1.rteJIH
1wKoro-.::iu60 rena.
Ilpu.¥Lep:
rCHhI B X-xpoMoeoMc y MYJK'Il1H - XY;
reHhI B Y-xpoMoeoMe y My:>Kq11H - XY;
reHhI B X-xpoMoeoMe y memu:mr - XO (euHp;poM IllepemeBeKoro---TepHepa).
XapaKTep11hre npM3HaKH X-euern1eHHoro 11,o:vi:1111a11Tttoro nma 11acJie,[\0Bam.u1:
3a60.1eBaHMe npoeJieJKHBaeTe.H B Kam.n;oM IlOKOJieHMH;
ee,1u 60JieH OTeu, TO see .n;oqcpu 6y11,yT 60.'JhHhIMH, a ebIHOBb.fl - 3.llOPOBhIMM;
eeJiu 6oJihHa ~vrnTh, TO sepo.HTHOeTh pomp;etnrn 6oJihHoro pe6eHKa paBHa
50 •y., He3aBHeHMO OT no.'Ia;
60,1e10T KaK MyiKqHHhI, T8K l1 .mem:.u;uHbI (6oJihHhlX mem.u;11H B eeMLe B ,n:sa
pa3a 60JibUie, qeM 6oJihHhIX My.mq11H).
IlpuMPpbt X-c.u:enJieHHoro ,n:oMHHaHTHoro Hae.1e,n:osaHu.H:
KOpuqHeBa.fl OKpaeKa :JM8JIH 3y60B;
BHTaMMH D-pe3MCTCHTHbIH paxH'r.
4. X-cu,enJLPltH.btu peu,eccuettbtii mun 11ac. iedo6anuR (reH Haxo,n:MTea s X-xpoMoeoMe, HB.'IRCTC.fl peueceHBHh!M).
XapaKTepHhie nptt3HaKw X-c1~enJieHH01·0 peueccttBnoro Tnna HacJie,n:oBaHHR:
6oJieIOT npeMMYlllCCTBCHHO My)KqHHhr;
60JihHLie ,n:eTH pom,naIOTCJl OT QleHOTHilHqeci-m 3,l(OpOBLIX po,n:HTeJiei1,
HO M8Tb 6oJihHOro .flBJIHeTCH reTep03HI'OTHOH HOCHTeJihHl1UCH COOTBCTeTBYIOU~ero reHa;
6oJihHhre Mymq11Hb1 He nepe,n:aI-OT 3a6oJiesaHne ChIHOBhJ:IM, HO nee ,n:oqepn - HOCI1TeJihHHU:hI COOTBeTeTBYIOlllero reHa.
IlpuMPpbt X-euenJieHHoro pe~eeeMBHoro HaeJie.n;oBann11:
reMOQlMJIHJJ A (HeCBepThrnaeMoeTh KpoB11) (p11e. 48);
MHOnanrn )lrorneirna (aTpOQlH.H Mhlllle<JHhlX BOJIOKOH);
~8JihTOHH3M.

5. Y·cu,enJteunoe 1-tacJLeao6aHue (reH Haxo,n:MTCJJ s Y-xpoMocoMe; Bcer,n:a
B I'CMH3HPOTHOM COCTOJ:IHHH, Il03TOMY seer.n:a npO.HBJIJ:leTeH).
XapaKTeptthle nptt3HaKn Y-cuerrJieHHoro HaeJie,n:osaHMH: reHhI, JIOKaJIH3oBaHHbie B Y-xpoMoeoMe, nepe,n:aIOTCH TOJihKO ebIHOBhHM (roJiaH~pnqeeKoe
HacJie.n;oBarrne); ,n:oqepu Bcer,n:a a.n:opOBhI, T. K. mrnor~a He noJiyqa10T Y-xpoMOCOMy.
IlpuMep Y-euenJieHHoro HaeJie,n:oBaHm1: rnnepTpMX03 MoqKM yxa ( «MOXHaThie yrnn »).
6. I'oJLa1tapu'lec1we HacJLeaoeaHue - 3TO HacJief\OBaHMe rrpM3HaKa TOjihKO
no MymcKOH JIHHl1H (nepe.n;aqa OT OT~a K ChIHy).

L1,umozenemu•1ec1wu 1tzem0<J

109

II

6

III

6
P111c. 48. PoAocnosHasi ceMblll c reMocjm11111ew A

MeTo,n; BKJIIoqaeT ;mcrrpecc-.n;mi.rttocT1rny rroJia (onpe,11,eJrentte X- 11 Y-xpoMaTHHa) H

1Capuomunupoea1tue -

orrpe,n;eJieHHe KOJiuqecTBa If KatieCTBa xpo-

MOCOM c IJ,eJihIO ,11,HarHOCTHKH xpOMOCOMHhlX 6oJie3Heii (reHOMllhlX MyTaIJ,HH,
xpOMOCOMHhlX a6eppaI.1,llH).
MaTepua.'loM ,11,JIH I.l,llToreneTHtfecKoro HCC"'le,n;oBaHmI MoryT 6hITh KJieTKH
rrepmpepttqecKoii KpOBll (JIHMcpOI.1,HThI); KJieTKH, noJiyqeHI!hie npn aMHHOII;eHTe3e HJIH 6HOHCHH xopHoHa.

:J1Ccnpecc·iJua21-wcmu1Ca -

::>TO ttccJie,11,oBaHtte noJioBoro X-reTepoxpoMaTH-

Ha (TeJihI.1,3 Bappa) B H,11,pax KJICTOK CJIH3HCTOM o60JI0'!I{H pOTOBOH IIOJIOCTH HJIH
B .n;pyrHX KJieTKax.

B

ttopMe y mettru;HH, MMeIOru;ttx ,n;Be X-xpoMOCOMhI, o.n;na MHaKTHBI1pyeTc.H,

o6paay.H reTepoxpoMaTMHOBYIO rJihI6Ky B .H.n;pe -

TeJihIJ,e Bappa.

Y

MYJKLJHH

(XY) ee HeT.
MymcKoii Y-rro.'!oBoii: xpoMaTIIH (F-TeJihUe) orrpe.n;eJI.HIOT rryTeM cneIJ,HaJihHoro OKpallIMBaHIUI H rrpoCMOTpa B JIIOMHHecueHTHOM MHKPOCKO!le . Y-xpoMaTHH BhI,!l;eJI.HeTC.H KaK CHJihHO CBeT.Hlll;8.HCJI TOqKa. qHCJIO Y-TeJreu COOTBCTCTBYeT q11cJiy Y-xpoMocoM B KapHOTHne.

Kapuomunupoeanue -

3TO anaJIH3 cpoToKapttorpaMMhI. ,n:JI.H anaJittaa Ka-

PHOTHrra xpOMOCOMhl pacrroJiaraIOT IIO rpymraM B BH,n;e H,!l;HOrpaMMhl (pHC.

HiJuozpa.M.Ma -

49).

aTo cttcTeMaTH3ttpoBaHHhiii KapHOTHrr.

CorJiacno ,n:ettBepcKoii KJiaccmpttKaUHH (,n:ettBep, CIIIA,

1960 r.)

xpoMoco-

Mhl pacIIoJiaraIOT IIOIIapHO IIO Mepe y6hIBaHHH HX BeJIHqHHhl, c yqeTOM pacrroJlO:i«eHHH ueHTpOMephl, HaJIHqllfl BTOPH'IHhIX rrepeTH:i«eK l1 CIIYTHHKOB. IloJIOBhle XPOMOCOMhl BhI,D;e.'!JIIOTCJI OT,D;eJihHO.
IlapII:i«CKa.H KJiaCCHcpl1K8I.l,H.H
aJihHOH OKpacKH xpoMocoM.

B

(1971 I'.)

OCHOBaHa Ha MeTo,n;e .n;mpc}lepeHI.I;H-

Kam,uoii: nape xpoMOCOM IIpOHB.'!JieTcH xapaK-

TeptthIH TOJlhKO ,!l;JIJI Hee YHHKaJihHhIH rrop.H,!l;OK qepe,n;oBaHH.H TeMHhlX H CBeTJlhIX IIOJIOC reTepo- H ayxpOM8THHa.

06utaJL 6uo:wzun. I'euernuKa

110

lJ

A

6-12

c

f)"Al>Ac')I\
13-15

XX IA IA

1J

F:

!6

17

:i::

'----v------'

AA

AA

.t

~'r'

F

G

P11c. 49.

Kap~ornn

4enoseKa

XpoMoCOMhI qeJIOBeiw pa3,n;eJIHIOT Ha ceMh rpyrrrr:

1. A - KpyrrHhie MeTaueHTpIIqecKIIe (1, 2, 3).
2. B - Kpyrrtthie cy6MeTauettTpIIqecKtte (4, 5).
3. C - cpe,n;ttIIe cy6MeTaueHTpttqecKtte (6-12).
4. D - cpe,n;HIIe aKpoue11TpIIqecKIIe c BTopwrnoii rrepeTJDI<Koii
(13, 14, 15).
5. E - 6oJiee MeJIKHe cy6MeTaueHTpw.recKIIe (16, 17, 18).
6. F - Memrne MeTauettTpw1ecKIIe (19, 20).
7. G - MeJIKIIe aKpouettTpttqecKtte c BTOpttqttoil rrepenrnrnoil (21, 22).
IlOJIOBaH X-xpoMOCOMa OTHOCHTCJI K rpynrre
CKaJI; Y-xpoMocoMa -

c,

cpe,n;HJIJI MeTaueHTpll'!e-

MeJIKIIH aKpouettTpIIK.

reHen1Ka COM3TM"l0CKMX KneTOK
8Ta ,ll;HCD;IIIlJIHHa II3yqaeT HaCJie,n;CTBeHHOCTb H H3MeHqHBOCTb COMaTHqeCKHX KJieTOK, HCilOJib3Y.H KyJihTypy KJieTOK pa3JIIIqHbIX TKaHeii H opraHOB.
Cyr.u;ecTBYIOT cJie,n;y10ru;IIe MeTo,n;hI reHeTHKH coMaTttqecKIIX KJJeTOK qeJioBeKa:

1. Ilpocmoe KYJtbmuBupoeattue.
2. I'u6pu{}u3au,uH - CJIIIJIHIIe KJieTOK

pa3HbIX TIIITOB (HanpIIMep, KJleTKH

«qeJIOBeK-Mbllllb» IlOCTeneHHO TepHIOT xpOMOCOMbI; MOiKHO ycTaHaBJIIIBaTb
rpyIIIIbI cuerrJieHH.H no O,ll;HOBpeMeHHO ucqe3aIOru;HM rrpH3HaKaM).

3. KJtOHUpoeattue PH,ll;OM.

I'u6puaoMa -

IIOJiyl.J:eHIIe IIOTOMKOB O,ll;HOH KJleTKH, narrpHMep rtt6:3TO KJieToqHbIH rtt6ptt.a;, IIOJiyqeHHbIH rryTeM CJIH.HHIIJI

HOpMaJibHOro JIHMc}>OUHTa H orryxoJJeBOH KJieTKII; rrpOH3BO,ll;HT MOHOKJIOHaJibHbie aHTIITeJia O,n;Horo TJIIIa.

4. CeJteKu,uH -

oT6op KJieTOK c 3apaHee 3a,n;aHHbIMH cBoiicTBaMH.

MeiJuijunc1wK zenemuica. E1iu3neqo6btU ;nemoiJ

111

MeTOA AepMaTormu:f>1r1K1r1
8ToT MeTo.z:i; rrpe,n;cTaB;uieT co6o:i1 i-13yqeHtte narrttJI.·urpHhrx y3opoB rraJihr~eB,
Jia,L(OHett, CTOil •
.IJ:epMaTOrJittqmqecKtte Y30pbl llH,!1;11Bl1,!J;YaJibHhl H ue MemnOTCH B Tel!eHtte Bce:i1
)Kll3HH • .IJ:epMaTOrJittqml!eCKHH amL1H3 ll03B0.1.HCT orrpe,n;eJIHTb 3MI'OTHOCTb 6J1H3HeUOB (MOHO- HJIH 1(11-), HeKOTOphIX rCHOMHhIX l1 xpOMOCOMHhlX MYTaUHH (60Jre3HH .IJ:ayHa, IlaTay ll .z:i;p.), npoBeCTH tt,z:i;eIITHq:mKar~HIO Jllll!HOCTH B KPHMHHaJIHCTHKe.

6nlr13He~OBblH M0TOA
MeTo.z:i; rro3Bo.1JieT oueHHTh po.Jih Hac.'le,z:i;cTBeHHOCTM H cpe.z:i;hr B pa3BMTHH rrptt3HaKa.
BJJH3HeUhI 6hrnaroT MOH03HrOTHhIMH H .n;tt3ttrOTHhIMH.
Motto3ttrOTHhie 6JJtt3HeI~hI pa3BttBalOTCJI H3 o.z:i;Hott orrJio,n;oTBopeHtto:i1 RiiueKJieTKH (3ttrOTbl) B pe3yJibTaTe ee pa3.z:i;eJleHHJI Ha,!l;BOe (Ha CTa,!l;Hl1 .n;po6JJeHttB:)
c o6pa30Bam1eM ,n;Byx sM6pnottoB. MoH03ttroTHhie 6.'IH3Heuhr HMeroT o.z:i;ttHaKoBhie reHOTHilhI, Bcer.z:i;a 0,ll;HOro IlOJJa; HMelOT BhICOKYIO CTeneHh CXOJJ:CTBa (KOHKOp,z:i;aHTHOCTb) no MHOrHM rrptt3HaKaM.

Pa3JJttqne ttx rrpH3HaKoB 3aBttCHT

TOJJhKO OT <})aKTOPOB BH€IIJHeil cpe,n:hI •
.IJ:n3ttroTHhre 6JJH3Heuhr pom.n;aroTCJI H3 ,z:i;Byx HJ'IH 60.JJee o,n:HoBpeMeHHO oByJittpoBaBlllttX H OIIJJO,!J;OTBOpeHHhIX pa3HhIMH crrepMaT030HJJ:aMH JIMU€KJJ€TOK.
0HH HM€IOT pa3Hhie reHOTHTihI, MoryT 6bITh 0,!l;HOro H.'IH pa3HOro IIOJJa. 0HH
cxo,n:HhI Mem.z:i;y co6oH: He 60.JihWe, qeM 6paThJI H cecTphr, pom,z:i;eHHhie nopo3Hh.
XapaKTeptt3yIDTC.H .z:i;ncKop.z:i;attTHOCThlO -

Hecxo,n:cTBOM rro :viHorttM rrpH3Ha-

KaM. BJJaro.z:i;apB: O,!l;HOBpeMeHHOMy pom,n;eHHIO H BOCTIHTaHHIO HM€IOT o6n~He
cpe,z:i;OBhie <})aKTOPhI. Pa3JIHqne npH3HaKOB B OCHOBHOM CBJI3aHO c reHOTHIIaMH •
.I1:JJJI Jl:OKa3aTeJJbCTBa po.JIM Hac.1e,z:i;CTBCHHOCTH cpaBHHBalOT ,!J;O.'IIO

('%)

KOH-

KOp,n:aHTHhIX nap (OJJ:HHaKOBhIX no KOHKpeTHOMY npH3HaKy) cpe,n:H MOHO- H ,!l;H3HrOTHhIX 6JIH3HeuoB •
.r:i:aHHhre 0 KOHKOp,n;aHTHOCTH 6JIH3H€UOB no H€KOTOpbIM rrpH3HaKaM np1rne,!1;€Hbl B Ta6Jittue.

Ilptt3HaKH

I

I

KoHKOp,11aHTHOCTh (cxo,11cTBO),
MoH03IffOTHhie

.IJ:tt3IffOTHhie

100

46

99,5
97

28
23

KocoJiarrocTh

32

3

3rr11Jierrcm1

67

3

llil130l}lpeHIUI

70

3

Ty6epKyJie3

fpyrrrra KpOBll
ll;BeT rJia3
IJ;BeT BOJIOC

I

'Xi

37

15

ICopb

98

94

CaxapHhlii: ,1111a6eT

65

18

06UfaR 6uo1wzuR. I'enemu1w

112

BJJaro,n;apH 6JJtt3HenoBoMy MCTo,n;y 6b1.rrn BhIHCHeua nac.'le,n;cTsemrnH 11pe1~
pacrroJ10:t1<enHocTh qeJJOBeKa K p.H,n;y 3a6oJJeBaHm'i: lllH30(l)peHIIH, 3IlllJICIICHII,
caxapHOMY ,LJ;Ha6eTy H ,n;p.

6MOXMMM'teCKllle MeTOAbl
3TH MeTO,ll,hI OCHOBaHhI Ha u3yqeHHII o6M~Ha Bern,eCTB (qlepMeHTOB, aMHHOKHCJIOT II T. ,n;.) MCTO,n;aMH 6HOXHMHH. 0HH l103BO.'l5HOT o6HapyiRHTb HapyllleHHH B o6Mene BCll~CCTB, BhI3BaHHhre H3MCHeHlUIMH renoB, II, KaK CJ!C.D;CTBHC,
aKTHBHOCTH pa3JIIIqHhIX qlepMeHTOB. KpoMe Toro, no3BOJJJIIOT ycTaHOBHTh reTepo311roTHoe HOCIITe.'lhCTBO HCKOTOpbIX 3a6oJJeBaHHH.
HacJJe.n;cTBCHHhre 6oJJe3HII o6Mena Bern,ecTB rro.n;pa3,n;eJJJIIOTCJI Ha 6o.'le3HH
yrJJeBO,ll;HOro o6MeHa (caxapHhlll .n,11a6eT), o6Mena aMHHOKHC.'IOT, JIHTIH,ll;OB
II T. ,n;.

IlpuMep.

ct>ennJJKeTOHypnJI -- 60J'Ie3Hh aMnHOKHCJIOTHoro o6MeHa. BJJOKH-

pyeTcJI npeBparn,eHIIe HC3aMCHHMOH aMIIHOKHC.'IOThI tjleHIIJJa.'laHnHa B Tnpo3IIH, npII 3TOM tjleHHJJaJJaHHH OKHCJIJICTCJI B tjlCHIIJITIHpOBHHOrpa,n;HyIO KHCJJOTy, KOTOpaJI BhIBO,ZJ;HTCJI c Moqoii:. 3a6oJJeBaHJie npHBO,ll;HT K pa3BHTHIO CJJa6oYMH51 y ,n;eTeii. PaHHee BhrnBJieHIIe n ncKJIIO'-IeHne tjleHIIJiaJiaHnna II3 nnTaHnH
II03BOJIJIIOT npMOCTaHOBHTh pa3BHTHC 3a60.1eBaHH5l.

XHMH'lec1me npo11ecch1 B opraHH3Me npu cI>eHHJIKeTOHypuu
tjlepMCHT tjleHtt.1aJJaHIIHrn.n;poKCHJia3a (,n;etjleKTHhrii)

I )( •I

tjleHnJiaJiaHIIH

THp03IIH

I X •I

MeJiaHnH

O=C-CH-CH-

1

I

OH NH 2

2

-OH

tjleHnJiyKcycnaJI KHCJIOTa
¢ennJIMOJIO'-IHaH K-Ta
¢ettnmmpoB1rnorpa,n;Haa K-Ta

nonymll.,\lllOHHO·CT3TlllCTlll'teCKlllM MeTOA
3To MCTO,ll, lI3Y'-ICHHH pacnpocTpaHeHHJI nacJie,D;CTBCHHblX npn3HaKOB (naCJIC,D;CTBCHHblX

aa6oJieBaHnii) B

nonyJJHl1lIHX.

Cyrn,eCTBeHHbIM

MOMCHTOM

npn HCTIOJib30BaHHH 3Toro MeTo,n;a HBJIJICTCH CTaTHCTlI'-ICCKaa o6pa60TKa
,Zl;8HHbIX.

M eiJu~uncKax zenemuKa. 1Wemodbl npenama1tbnoii iJuaznocmu1m

113

IlollyJUIUUOHHO-CTaTIICTtttieCKlIH MeTO,ll; IT03BOJIJ!eT pacctrnTaTh B nonyJrn:UlIIf ll3CTOTY HOpMaJihHhIX lI ITaTOJIOrIItieCKlIX reHOB lI reHOTlIITOB: reTepo3HroT, roM031IrOT ,ll;OMIIHaHTHhIX lI peuecCHBHhIX.
"lJacToTa rettOTIInos II cJlettoTttnoB pacctIHThrnaeTcJI ITO <jJopMyJte XapauBauH6epia:

p2 + 2pq + q2

=

(p

+ q)2

=

1,

r,n;e p -- qacTOTa .n;oMHHaHTHoro retta A,
q - qacTOTa peueccttBHoro reHa a,
q2 -- t.IaCTOTa rOM0311l'OT ITO pe1~eCClIBHOMY reHy aa.
Bamtto, tITo no 3TOtt cpopMyJie Momno pacctIIITaTh qacTOTY reTepo311roT no naTOJiornqecKIIM peuecCIIBHhlM reHaM, KOTOphlC HaX0,11;.HTC.H B CKphlTOM COCTO.HHIIH.
JipuMep. "llaCTOTa cpeHIIJIKeTOHypIIII (aa) B norryJI.Hl[llll 1: 10000,
3HatIIIT, q 2Ha = 0,0001; q = 0,01

p+q

=

1,

Tor,n;a p = 1 - q = 1 - 0,01 = 0,99
2pqAo = 2 x 0,99 x 0,01 = 0,0198 - 0,02 m:rn 2 'Xi.
3HatIIIT, qacTOTa reTep031IrOT ITO reny cpeHIIJIKeTOHypIIH B II3ytiaeMOM IIOITyJI.HUlilI COCTaBJI.HeT 2 '% .
MeTOAblnpeHaTanbHOM
(BO epeMSI 6epeMeHHOCTlll)
AlllarHOCTlllKlll

AMHuou,eHme3 - OT6op (rrpo6a) aMHHOTMtiecKott mu,n;KOCTH c ueJihIO UHTOreHeTHtiecKoro HJilI 6IIOXHMHlleCKOro HCCJie,n;oBaHH.H ,ll;JI.H ,n;11arHOCTMKH HaCJie,n;cTBeHHhIX 3a6oJieBaHHM.
Il03BOJI.HeT BhUICHHTh reHHhie 6oJie3HH o6MeHa Bell[eCTB lI xpOMOCOMHhle 6oJie3HH.
0T6op aMHIIOTMtiecKo:H mtt,n;KocTu rrpoBO,n;HT rro.n; KOHTpoJieM yJihTpa3ByKoBoro HCCJie,n;oBaHH.H Ha 14-16-:H He,n;eJie 6epeMeHHOCTH.
OnpeaeJteHue 6eJ11w aJtb<jJa<jJemonpomeuHa ( A<PII) B ChrnopoTi~e KPOBH 6epeMeHtto:H HJIH B aMHHOTttt.IecKott mu,n;KOCTH. A<l>II - rJinKorrpoTen.n;, KOTOPhrtt npo,n;yuupyeTc.H rretieHhIO nJio,n;a.
HccJie,n;oBaHtte npoBo,n;HT Ha 14-16-:H tte,n;eJie 6epeMeHHOCTH.
YpoBeHh 6eJIKa A<l>II nOBhIIIJaeTC.H npn p.H,n;e attoMaJIHtt pa3BHTHH ITJio,n;a,
TaKwx KaK aH3ttuecpaJIHH (oTCYTCTBMe M03ra), BHyTpHyTpo6Ha.H rn6eJih rrJio,n;a.
CHIDKettwe A<l>II rrpowcxo,n;HT rrptt xpoMOCOMHOtt naToJiornw (cIIH,n;poM .[(ayHa,
3.n;Bap,n;ca II ,n;p.).
YJtbmpa3ByKoBoe uccJteaoBaHue nJtoaa ttan6oJiee pacrrpocTpatteHHhltt
MeTo.n;. Ott OCHOBaH Ha crroco6HOCTH YJihTpa3BYKOBOM BOJIHhl OTpamaThC.H OT
UOBepxHOCTH pa3,n;eJia .D;BYX cpe.n;, OTJIHllaIOil[HXC.H JllllIHOM IIJIOTHOCThlO, tITO
noaBoJI.HeT noJiytIMTh H3o6pameHwe Ha 3Kpatte. HccJie,n;oBaHHe rrpoBo,n;HT Ha
PaaJIMqHhrx cpoKax 6epeMeHHOCTH.
MeTo,n; II03BOJI.HeT .n;warHOCTnposaTh rpy6hre rropoKH M03ra, rropoKH KotteqllOCTeii, rroqeK H .n;p.

06UfaR 6uo;wiuR. I'enemu1w

114

1113MEH'-llllBOCTb. Tlllnbl 1113MEH'-llllBOCTlll. PEnAPAU.lllSI AHK
H3Meli'IU6ocmb -

8TO CBOHCTBO oprairn.3MOB ,n;aBaTh OTKJIOHeH11H B npH3Ha-

Kax rro.n; .n;eil:CTBl1eM BHelliHl1X 11 BHyTpeHHl1X cpaKTOpOB cpe,n;hI.

He1Iae.1eµ;cTsettua.H
(MOAlll<j:>lllKal\lllOHHasi
111n111 <j:>eHomnw-iecKasi)

Hac.'le;:i;cTBeHHa.H
(reHomnw-iecKasi)

J{o.1116unamu6HaJ1

I'e111taJ1
nepecmpouJCu

I) oeJ1e4u.11;

I) ianllOUOU.ll;

2) oyn.w1Ca4u11;

2) no;wn.wuou.11

3) u 1rnepc11J1;
4) mpa11c.w1Ca4u11

ll3MeH•IHBOCTb opraHl13MOB MO:>KHO pa3,n;eJil1Tb Ha .n;Ba B11.n;a: HeHaCJie,n;cTBeHHaH (Mo,n;11cp11Kau;110HHaH 11Jil1 cpeHOTl1IIH'IeCKaH) 11 HaCJie,n;CTBeHHaH (reHOTl1II11qecKaH).

Moau<PU1cau,uoHHaJl U3Melil/U6ocmb -

:JTO H3MeHeH11H oco611, B03H11Kru11e

no.n; B03,n;eil:CTBl1eM cpe,n;hI, He npl1BO.D;HIIJ;l1e K 113MeHeHl1HM B reHOTl1IIe 11 He nepe.n;a10ru;11ecH no HacJ1e,n;cTBY.
XapaKTep11cT11K11 Mo,n;11cp11Kau;110HHOil: H3MeHq11BocT11:

1)
2)
3)

HeHaCJie,n;yeMOCTb 113MeHeHHOro npl13HaKa;
MaCCOBOCTb (npOHBJIHeTCH y MHOr11x oco6eil: B IIOIIYJIHIJ;1111);
a,n;eKBaTHOCTb (Bcer.n;a COOTBeTCTByeT 113MeHeHl1HM BHeruHeil: cpe,n;bl: qeM

60Jihllie B03,D;eHCTBl1e, TeM 60Jibllie OTBeTHaH peaKIJ;11H);

4)

a,n;aIITl1BHOCTb (nOBhIIllaeT np11cnoco6JieHHOCTb opraH113Ma);

5)

.n;11arra30H 113MeHeHl1H .D;JIH oco6eii: ,n;aHHOrO B11.n;a o6yCJIOBJieH reHeT11qe-

CKl1 11 xapaKTep113yeTCH HOpMOM peaKIJ;1111.

HopMa pea1Cu,uu -

8TO reHeT11qecK11 o6ycJIOBJieHHhIH rrpe,n;eJI pa3BHTHH

IIpH3HaKa.
HopMa peaKIJ;HH MomeT 6hITh
pocT, 11HTemreKT) HJIH
rpyrrrra KpOBH).

y31Cou

wupo1Cou

(KOJIHqecTBeHHhie rrpH3HaKH -

(KaqecTBeHHhie npH3HaKH -

Bee,

u;BeT BOJioc, rJia3;

y 3 Men'lu6ocmb.
II pwnep1>1

Tun bi u.1.11ien'lu6ocmu. Penapau,,uR

If II K

115

Mo.umlrn1rn11;i10H11oii n3MewnrnocnI:

113MeHeH11e III1I'MeHTaU,J1l:I KO./Kl:I IIOCJie 3arapa;
IlOBbIWeHI:IC ypOBIUI reMOI'JI06mrn B KpOBl:I B 1\Ba pa3a IIpl1 l(.111'reJJbHOM
HaXO./K,'(CHHM B ropax Ha BhICOTe

4000

M.

Mow1qn11rn11,Ho1111a.H 113MCH'II:IBOCTh MHorI:Ix npH3HaKOB y pa3Hhrx opra1111 3 MOB IIO,[('IHII.HeTC.H o6II~HM 33KOIIOMepHOCT.HM. ,[(aHHb!M Bl:Ij( II3MeH'll1BOCTI1 l13yqaeTCH B 6oJihIII11X rpyrrrrax opraHI13MOB. CTerreHh rrpOHBJieHHH IIp113HaKa pa3JII1LIHa y 'lJlCHOB 3TOH rpynrrhr. BoJihWHHCTBO IIMeeT cpel(nee 3Ha'Ien11e (poC'r,
Bee 11 T.

lJ:.).

M11HnMaJihHOe II MaKc11MaJibHOe rrpoHBJieHHH npH3HaKa na6J110,z:i;a-

IOTCH TO.lbKO

y

OT1\eJ1bHhIX oco6eiL

311a11eH11e 3HKOHOMepHOCTeii: MO,ll;HQJHKall,HOHHOH II3MeH'Il:IBOCTl1 II03B0Jl.HCT
npe,D;cKa3hIB3Tb CTerrenb BbipameHHOCTl:I IIPII3HaKa B 3aBI1ClIMOCTH OT yCJIOBHH
cpe,D;hl II HMeeT 60JibWOe 3Ha'ICHIIe B ceJICKll,IIII (coJie-, 3acyxo-, MOp030YCTOH'Il1BOCTb).

II

HaCJ1eiJcm6eHHaJ1 UJMe1l'tueocm1> rro,D;pa3l(eJ1.HeTCH Ha 1wM6ullamu6Hy10
Mymau,uo11Hy10.
IlpH'II1H3Ml1 KOM6HH8TIIBHOH II3MeH'IIIBOCTl1 .HBJI.HIOTCH: KpOCCIIHl'OBep; He-

3aBIICIIMOe pacxom;~eHIIe xpoMOCOM B MeH03e; He3aBIICIIMOe CO'IeTanne raMeT
rrpII OIIJIO.[(OTBOpeHIIII.

IIpuMepo.'lt KoM6nHaTHBno:ii: H3MeH'IHBOCTn HBJIHeTCH TpeTIIii 3aKoH Mett;i:eJIH (,D;eTII O,ll;HIIX pO,ll;IITeJJe:iI OTJIH'l3IOTC.H ,lJ;pyr OT ,ll;pyra HOT pO,ll;HTCJie:ii:).

Mymau,uo1tHaJ1 U3MPH'lU6ocm1> B03H111rneT B pe3yJibTaTe pa3JIII'lHhrx H3MeMymau,uoH1ta.R meopu.R 6bIJia cqiop:viy;rn-

HeHIIH HacJJel(cTseHHoro MaTepIIaJia.
poBaHa B

1901

r. I'OJIJiaHl(CKHM 6oTaHIIKOM ryro l(e Cl>pII30M.

Mymau,uu -

aTo pe3Kne cKa11Koo6pa3Hhie H3MeHeHn.H reHOT11na. MyTall,IIH

H3MeHHIOT HaCJie,D;CTBeHHhIH MaTepnaJI H MoryT rrpHBO,ll;HTb K pa3.1H'lHbIM rrpo.HBJieHH.HM Ha ypOBHe cpeHOTIUia.
CBottcTBa MyTaii:11rr:
- B03HHKaIOT BHC38IIHO;
- He HarrpaBJieHbI -

MOJKeT MYTHPOE3Th JII06aH qacTb reHOTHrra;

- 60JihllIHHCTBO MyTau,n:H -

peu,eccttBHhre, pe,ll;KO -

,ll;OMHHaHTHhIC;

- ,ll;JIH opraHH3Ma 60JihllIHHCTBO MyTall,HH Bpel(Hhie, pe,ll;KO -

HeHTpaJihllbie

IIJIH IIOJJe3Hbie;
- MYTa11,11n nacJre,ll;yIOTc.H;
- BhI3bIBHIOT CTOHKne H3MeHeHII.ff HacJie,ll;CTBeHHOrO MaTepIIaJia;
- BhI3bIB8IOT Ka'!eCTBeHHbie H3MeHeHH.H rrpn3HaKa, He o6pa3yIOill,He HerrpePhIEHoro p.H,ll;a BOKpyr cpel(HeiI BeJIII'IIIHhr;
- MoryT B03HHK8Tb B reHOTHIIe IIOBTOpHO.
MYTa11,II11 MOryT B03HHK3Th ITO,ll; ,ll;e:iicTBHeM BHellIHHX n BHyTpeHHHX cpaKTopoB.

MymaieH1>1 -

3To cpaKTOpbr BHellltte:H cpe,D;hI, BhI3bIBaIOill,He MyTaU,HII.

MyTareHhr rro,D;paa,ll;eJIHIOTCH B 33BHCHMOCTH OT rrpttpO,ll;hI B03l(e:ttcTBI1H tta:
XBMnqeCKne (a30THCTa.ff KHCJIOTa, IIIIPIIT, HHTp030MeTIIJIMO'IeBIIH8 II ,lJ;p.), cpnali'lecKne (pa,ll;Hall,11.H, BbICOKa.H TeMrrepaTypa), 6nOJIOI'II'IeCKI1e (a.ueHOBHPYCbI,
PeTpoBnpychI).

0614aH 6uo1wzuH. I'enemu1w

116

CoBpeMetmoe onpe.ueJICHHe

.A'Lymau,uil:

cj:lopMall,HH .uaHHOro opraHH3Ma,

3TO H3MetteHHa retteTHlJ:eCKOtt HH-

KOTOphie npH perrJIHKaIJHI1 3aKpenJIJl!OTCa

H nepe,1.1a10TC.H no HaCJIC,L(CTBy.

K.rnccwjJuKUU,UJl .wyma1~uil

--------Ilo

CIIOHTaHHhie
(B np11po.ue; 'IaCTOTa 10

-----

HH11YI~npyc:\.10CTli

:, - 10 ,;

Ha 0,1\Hll ren B IIOKOJICHHH)

lfH,l.lyI~HpOBaHHhIC

(.ucticTBHC MyTal'CHOM; '!a~e
B 10-100 pa3 11 60,'ICC)

Ilo BJllU!IlllIO Ha )K113HCCIIOC06HOCTh

tteH:TpaJihHhie

YCJIOBHO-JICTaJlbHblC (rrpII
orrpe,UCJICJ!Hh!X
yc,1omrnx)

no,1y,'leTaJlhHhIC

JICTaJihHbie
(Bhl3bIBaIO~He rII6CJib)

------------- ------Ilo MCCTY B031IIIl<HOBemu1 B opraHH3:vIC

renepaTHBHh!C
(B IIOJIOBhIX K.'ICTKax)

COMaTII'ICCKIIe
(B K,'leTKax Te,1a oco6rr)

Ilo xapaKTepy H3MeneHHH reHCTlI'ICCKOro MaTepIIaJia

reHHhie

xpo:VIOCOMHhIC

I'ettnbte Mymau,uu,

mo'le'lnbte

HJIH

rcHOMHhie

(TOlJ:KOBhie) npe,11,cTaBJIJIIOT co6oli: H3Me-

HeHHH B OT,L(CJihHbIX reHax.
THrrhI reHHhIX MyTail,Hli:

Bea caeuia paNKU C'lUmbWGHU.R

1.(20 %

/

3aMeHa

TpaH3IIIl,HlI (nypIIn ---)> rrypmr
ITHplIMll,!\lIH---)> IIIIpl!MlI,UIIH)

OCHOBaHFUI~

BCCX MyTail,HH)

(KOJIHlJ:CCTBO HYKJICOTM,11,0B He MCHJlCTCH):

~

[AT: u;r --

TpaHCBepCIIH (rrypl!H ---)> IIllpIIMII,!\lIH
Illlpl!Mll,l\HH ---)>)

2.

llHBepcHH -

H3MCHCHHe rropH,11,Ka HYKJICOTH,l.IOB.

IIVpHHhl'

l

rr~pHMH~IIHbl J

#JM€ttl/,U6ocmb. Tunbt U3M€ttl/,U6ocmu. Penapa~uR )~llK

117

Co crJBuzoM pa.MKU c'lumbtBaHUJl (KO.JIIFieCTBO HYKJ'IeOTH.ZWB B reHe MemieTc.fl): ~e.'leUH.fl - Bhma~eHHe; ~yrwmrnun.H - Y~BOCJIHe; Hl!Ccpu;HJI - BCTaBKa.
Ilpu.Mepbt:
1. 3aMeHa no THrry TpaHcBepcnH
CepnoBH~HOKJieTo<rna.H aHeMH.H
6-ll aMHHOKHCJIOTa
HbA - r;1yTaMHHOBaH
HbS - HaJIHH --- l.J;AT

}
KHCJIOTa -

IU'T

f3-u;euL ,UHK

2. 3aMeHbI mo6oro Tnna MoryT npHBO~HTb K ITO.HBJieHHIO CTOil-KO~OHOB.

B pe3YJihTaTe TpaHCJI.Hu;n.H 6eJrna o6pbrnaeTcH. 06pa3yeTCH H3MeHeHHhIH (yKopoqeHHhitt) IlOJ'IHITeITTH~. r~-TaJiacceMHH - HbA HMeeT HOpMaJihHYIO a-cy6'he~i-rnnu;y' HO yKopoqeHHYIO 13-cy6'he~HHHI.I;Y.

Xpo.Moco.MHbte Mymau,uu - 3TO H3MeHeHHH CTPYKTYPhI xpoMocoM.
1. ,[(eJieu;nn - Bhma~aeT qacTb xpoMOCOMhI.
Ilpwnep. CnH~poM «KOIIIaqhero KpnKa» - ~eJieun.H KopoTKoro rrJieqa
Il.HTOH XpOMOCOMbI.
2. ,[(yrrJinKauIIn - y~soeune yqacTKa xpoMocoMhI.
IIpuMep. ,[(yrrJinKauna reHa Bar B reHoTnne Myxn ~po3ocpnJihI rrpnBo~HT K yMeHhIIIeHHIO qncJia cpaceTOK r.,1a3.

3. llHBepcnn

rrapau;eHTpnqecKIIe
----------- (He 3aTparnBaJOT u;eHTPOMepHbIH paii:oH)
-----rrapaueHTpnqecKne (3aTparirnaJOT pattoH r~eHTpoMepbI)

c::::JIM::l._ . /

c::
____,.
____
:::>

llpu.111ep. Ilpe~ITOJiaraeTC.H, qTo BTOpaH xpoMOCOMa y qeJIOBeKa - 3TO pe3YJibTaT o6'he~nHeHHH, u;eHTpnqecKoro CJIHHHH.H 12-ii: H 13-ii: y rrpe~Ka.
5. TpaHcrro3nu;11n -- rrepeMeru;eHHH Mo6nJihHhIX reHeTnqecKnx 3JJeMeHTOB.
llpuMep. TpaHcrro30HhI 6aKTepnn, As n Ds 3JieMeHThI y KYKYPY3hI,
M,[(r (Mo6nJibHhie ~ncnepcnpoBaHHhie retthr) ~po3oqrnJihI.
I'eno.Mnbte Mymau,uu - 3TO n3MeHeHne qncJia xpoMocoM (B rarrJIOH~HOM Ha6ope - reHoMe).

06UfaH 6uonozun. J'enemu1rn

118

I'eltO.M - ITOJIOBl:IHHhlH Ha6op ,UHK l:I nOJIOBl:IHHhIH Ha6op reHeTI:I'IeCKoro
MaTepI:IaJ:ra.
1. 3yrr:IOHl\HH -- yBeJIH'IeHne Ha6opoB xpOMOCOM, Kpanwe n.
IIpu.Mep. 3n -- TpI:Irr;10n.n:h1
4n - TeTparnrnM,n:hI II .up.

2. Attayrr.110M,[(MH - He Kpanroe n yBeJIM'ICHI:Ie MJIM yMeHhllleHI:Ie 'IMCJia
xpoMOCOM.
-2n + 1 - Tpl:ICOMI:IH.
IIpu.Mep. TpMCOMHH no 21-n xpoMocoMe - cnH.n:poM ,UayHa (47; + 21-a).
XXY - Tpl:ICOMHH y MYiK'IMH no X-xpoMocoMe - CMH,n;poM KJiaiimpeJihTepa.
-2n - 1 - MOHOCOMUH.
Ilpu.Mep. C1rn.n:poM lllepellleBcKoro-Tepttepa: 45, XO. JKettru;MHa c rroJIOBhrMM xpoMocoMaMM XO.
IlpH'IMHa amiynJIOl:I,ll;l:Il:I - Hepacxom,n:eHI:Ie IlOJIOBhIX xpoMOCOM BO BpeMH
Menoaa.
1. Hepacxom.n,em1e noJIOBhIX xpoMocoM BO BpeMH Melioaa y MaTepM:
P:
~ 46, XX
x
46, XY cf'
~- -"
r.-~
,;;;~
G: 24,(_~_:X,,/
24,1,Q;
23,Q\.)
23,\¥/
F: 47, XXX
47, XXY
45, XO
23, YO
TpMCOMMH no
Cntt,n:poM
CMH,n;poM
fM6eJih anrOThI
X-xpoMocoMe KJiaiimpeJihTepa llleperneBcKoro-TepHepa
2. Hepacxom.n:em1e noJIOBhIX xpoMocoM so BpeMH Melioaa y 0T1.1,a:
P:
Y46, XX
x
46, XY cf'
24 ,,:::- -"'
2 r.o-;'\
G
2 r;~

:

.<~-~

3,\_)0

F: 47, XXY
CI:IH,n:poM
KJiai'rn¢leJihTepa

2,\___)

45, XO
CnH.n:poM
lllepellleBcKoro-TepHepa

3aJWH roMOJIOrH11ec1rnx p.H)J;OB HaCJie)J;CTBeHHOH
H3MeH11HBOCTH

H. II.

BaBHJIOBa

I'e1temu'lecKu 6Jtu3Kue euobt u poabt xapaKmepu3y10mc.R «cxoo1tbt.Mu p.Roa.Mu ltllCJleocmee1t1tOU U3.MClt'lU60cmu». 31ta.R U3MCH'lU60Cmb OOltO?O euoa, MOJICHO npeocKa3amb noReJie1-we napaJIJICJlbltbtX <jJopM y (}pyioio pooa uJiu euaa.
H. M. BaBMJIOB npocJie,n;MJI M3MeH'IMBOCTh MHomecTBa npI:IaHaKoB y aJiaKOB: pmn, rrrneHHI!hI, oBca, H'IMeHH, KYKYPY3hl.
3aKoH H. M. BaBMJIOBa MMeeT 6oJihrnoe aHa'IeHI:Ie ,n;JIH cem~Kl.l,MH. OH noaBoJIHeT aapaHee npe,n;BI:I,UeTh, KaKI:Ie npH3HaKI:I H3MeHHTCH y Toro HJIH l:IHOro BH.n:a B peayJihTaTe B03,ll;eHCTBHH Ha Hero MyTareHHhIX ¢>aKTOPOB.
B HacToaru;ee BpeMH aaKoH roMoJiorM'IecKMX pa,n;oB no,n;TBepm,n:eH He TOJihKO
Ha 3Jl8Kax, HO TaKme Ha npnMepe rpM60B, Ml:IKpoopraHH3MOB, iKl:IBOTHhIX.
IlpH'IHHhI CXO,ll;CTBa MyTal.l,l:IH 3aKJIIO'IalOTCH B CXO,UCTBe xapI:IOTl:ITIOB y 6JII:I3Kl:IX BH,ll;OB l:I B CXO,ll;CTBe paCTIOJIO;KeHI:IH aJIJieJihHhIX reHOB B xpOMOCOMax.

penapau,un ;n,011,eKyllbi!{HK

119

Penapau;HH MOJieI~YJihI

,r:t;HK

B ICICTKe cyw;ecTByeT CHCTC'Ma 3RIIl;llThl HRCJie.n;cTBeHHOro MRTep11a.'Ia ll,HK
OT B03HlllO!OBeJHIH MyTau;tt:IT ll IIOBpem.n;eHHM - ClICTeMa penapaILMll.
Penapa41ut - 3TO BoccTaHoB.'IeHHe noBpem.n;eHHOM CTPYKTyphr ,UHK (c;romHhIM qiep:vieHTaTHBHbIII npouecc ).
BhIJ~eJIHIOT ttecKO.'IhKO pa3HOBM/::(HOCTe:ii: penapaIIMH:
I. cf>OTopenapaumi.
II. TeMHOBaH penapmi;HH, tt;rn 3KCIIH3HOHHaa (Bh1pe3arow;aH) peuapaIIMH,
BIUilO'IalOIIl;aa: .n:opCIIJ1HK3THBHyro penapauttro; IIOCTpeIIJIMKaTHBHYIO
penapaumo; penJittKaTHBHYJO penapauttro.
III. SOS-penapari;HH.
I. «l>0Topenapau;m1 (pHc. 50).
IIo.n .n:efrcTBMeM yJLbmparjJuoJZemoeblX JLy'leil ceema B IIen11 ,II.HK Mem.n:y
,nByMH ocHoBaHHHMH T-T o6pa3yroTCH xnMttqecKMe CBH3M (B03HMKaeT T-T
,nttMep), Hapyma10ni:11ii: cqHThIBaHMe MH<l:>opMaIIMll M penJIMK3I::(l1lO .UHK.
Ilo.n: ,nettCTBHCM euiJU.MO?O ceema aKTMBMpyeTCH: <I:>epMeHT (<i:>OTOJ1lla3a), KOTOpblll pacw;enJ'IH:eT 3TM ,n:OIIOJIHMTeJibHbie CBH:3H. 8TOT IIPOIIecc H33hIBaeTCH:
rpomopenapa4uR.
II. TeMHOBaH, HJIH 3Kcu;uauottHaH (Bb1peaarorn;an), penapau;un.
8Ta penapari;tt.fl xapaKTeptt3yeTC.fl ocHOBHhIMH npoueccaMtt:
cpepMeHT (:::m.n;ottyKJiea3a) «Y3HaeT» IIOBpem.n;eHHhIM yqacTOK HMTll
,UHK;
cpepMeHT (3K30HYH:J'lea3a) «Bh!pe3aCT» IIOBpem.n;eHHhIM yqacTOK;
cpepMeHT (,UHK-noJIHMepa3a) CHHTC3HpyeT .UHK no npHHI!HIIY KOMIIJieMeHTapHOCTH;
<I:>epMeHT (.'1Hra3a) «ClllIIBaeT» KOHI!hl BHOBb CMHTe3MpOBRHHOrO yqacTKa C OCHOBHOH HHThlO ,[{HK.
Ilo BpeMeHM ocyw;eCTBJieHHH TeMHOBa.fl penapau;H.fl MOJKeT npoHCXO.ll:MTh .n:o
HJIH IIOCJie peIIJIHK3I!MM ,UHK M BO BpeM.fl penJIMKaIIMM.
1. nopenJtUICllmU61-taJl penapa4UJl - 3TO BOCCTaHOBJieHMe IIOBpem,n:eHHOH
HHTM ,UHK .n:o ee y,n:BoeHM.fl B G 1 nep110.n:e KJieToqttoro IIMKJ'la.
2. llocmpenJLu1CamueHa.H penapa4u.H - BoccTaHOBJieHMe ,UHK nocJie penJIMKa!IMM B G 2 nep110.n;e.

I I I I I I

AfAAl..ll
II Ill

Ill II

T Ll T=T f A
I I I \
PMc. 50.

BHAHMbU1 C88T

t t t +

<l> o 10pea n111 ea It l/ljl
<l>eJ>MeHT

I I

I I I I I I

AfAAl..ll
II Ill II II Ill II

ll..lllfA
I I I

<t>ornpenapa1..wm. TlilM"1HOBbl8 A"1M8Pbl pacw,enmHOTC51, "1 BOCCTaHaBn"1B3IOTC51 BOAOPOAHble CB513"1

A-T

06UfaR 6uo.rwzuR. I'enemuica

120

IlpoHCXO)lHT

«Y3HaBaHHe»

n curnBaH:ue KOHI~OB

.JJ:HR.

IrnpymeHtt.H,

«Bhipe3aHHe»

,11,ecpeKTa B

.JJ:HR

l1nor,11,a 6e3 ,11,oCTpa:uBaH:u.H tte.n,ocTa10ruero yqacTKa

CTPYKTypa reHa MOJKeT H3MemIThCR.

3. PenJtwcamuenaa penapau,u.R -

BoccTanoBJ:re1me .JJ:HR B xo.n;e penJJHKaI.J;Htt.

IlpOHCXO,ll,HT y,ll,aJJeHtte H3MeHeHHhlX HYKJJeOTH,ll,OB B XO,ll,e pocTa uerru,
HO B HeKOTOph!X CJJyqaax perrJJHKautt.H MO:>KeT H,11,TH no H3MeHeHHOll MaTp:uue.
CTpyKTypa re Ha MomeT MeH.HThC.H.
CyruecTBYIOT MyTau11tt (y qeJJoBeKa -

reHHhte 6oJJe3Htt), BhI3hIBa10ru:ue tta-

pyruemrn rJJaBHhIX cpepMeHTOB perrapau:utt. BocCTaHOBJJeH:ue

.JJ:HR

B Tairnx

CJJy'la.HX CTaHOBHTC.H HeB03MO'.IKHhIM.

Ilpu.Mepbt
1. Il:urMeHTHa.H

Kcepo,ll,epMa. Ilp:u rr:urMeHTHOH KCepo.n,epMe B KJJeTKaX OT-

cyTCTByeT cpepMeHT (,ll,e3oKcmrnpHMH,ll,HHcpOTOJJHa3a), Heo6XO,ll,HMhlH ,ll,JJ.H perrapau:u:u

,IJ;HR, noBpem.n;eHHOH yJJhTpacp:uoJJeTOBhlMtt JJyqaM:u. Ilo)l )lettcTBlieM

COJJHeqHoro CBeTa IlO.HBJJ.HIOTC.H pacruttpeH:ue Karr:uJJJJ.HpOB, oporoBeHUe 3IlH)lepMHCa, rropameH:ue rJJa3, pa3BHTHe paKOBhIX orryxoJJett KOmH, KOTOphre rrpnBO,ll,.HT K rrpem)leBpeMeHHOH cMepTH.

2.

IlpOTHBOilOJJOJKHhIM np:uMepoM qpe3Bhl'l8HHO BhlCOKOH crroco6HOCTH

Micrococcus radiodurans.

8TH 6aK-

Tep:u:u MoryT BbI)KHBaTh B yCJJOBH.HX YJJhTpacpuoJJeTOBOH pa,ll,:uauHH, B

1000 pa3

KJieToK K perrapautttt .HBJJ.HeTc.H opraHU3M

rrpeBbllliaIOruei1: CMepTeJJbHYIO ,ll,03y ,ll,JJ.H ,ll,pyr11x MHKpoopraHH3MOB.

m.

SOS-perrapaum1 -

Bpem)leHHH CTpyKTYPhl

3TO 6bICTpa.H penapau:irn np:u 3Haq:UTeJJhHOM 110-

.JJ:HK.

Jl:JJ.H Hee xapaKTepHO HeToqHoe co6JJIO)leHHe

KOMITJJeMeHTapHOCTH.

rEHETM"4ECKME OCHOBbl CEnEKLl,MH >KMBOTHblX,
PACTEHMM, MMKPOOPrAH"13MOB.
6"10TEXHononu'

CeJte1w u.R 1

HayKa o coa,ll,aHHH HOBhIX :u yJJyqrueHUH cyruecTBYIOJUHX no-

po,lJ, ;KHBOTHhIX, copTOB pacTeHHH, IlITaMMOB MHKpoopraHH3MOB.
TeopeT:u'leCKOH OCHOBOH ceJJeKu:u:u RBJJ.HeTcf! reHeT:uKa.

3ada'lU ceJle/CU,UU -

C03)laH:ue HOBhlX H yJJyqrueH:ue cyrueCTBYIOJUHX no-

PO,ll,, copToB, IIITaMMOB pa3JJHqHhIX opraHH3MOB.
Jl:JJfl peIIIeHH.H aa.n;aq ceJJeKUHH JJI06hIX opraH:U3MOB Heo6xo.n,HMO:
3HaHHe 3aKOHOMepHOCTeli HaCJJe)lOBaHH.H np:U3HaKOB;
H3y'leH:ue HacJJe,ll,CTBeHHOH H3MeHqHBOCTH;
H3y'leHHe BJJHf!HH.H cpe.n,bl Ha pa3BHTHe npH3H8KOB (MO,lJ,HcpHKan;:uomrn.fl
H3MeH'lHBOCTh);
nay'leHne copToBoro, BH.D;OBOro, po.n.oBoro paattoo6paa:ua KYJihTyp;
paapa60TKa CTpaTerHH H MeTO,ll,OB HCKYCCTBeHHOro OT6opa.

Ilopodbt, copma, mma.M.Mbt -

3TO ncKyccTBeHHO co3.n,aHHh1e qeJJoBeKoM no-

nyJif!D;HH opraHH3MOB. Jl:JJR HHX xapaKTepeH Ha6op HHTepecyIOJUHX qeJJoBeKa
npHaHaKoB, aaKpenJJeHHhIX ttacJie,n;cTBeHHO.

I'enemu<tecKue ocnoBht ce;ieKlfUU

CeJie1n~m1

.[(.'I.fl

121

pacTeHHii

perneHH.H 3a_naq CeJ1emJ,Hl1 paCTCHMM o'reHh BaJKHO COpTOBOe l1 BH,l(OBOe

pa3H006pa3HC.

H. H.

BaBHJIOBhIM 6b1Jia co6paHa KOJIJieKIJ,M.H copToB KYJihTYPHhIX pacTe-

HHM lI HX ,nHKHX rrpe,nKOB co Bcero 3eMHOI'O rnapa; 6bIJII1 OTKpbl'fbI CCMb IJ,eHTpOB ,npeBHCI'O 3eMJie,neJIH.H, HJIH IJ,eHTPOB MHOroo6pa3I1.H I1 IIPOI1CXOJK,l(eHttH
KYJihTypHbIX pacTeHI1M (HMCHHO B 3THX IJ,eHTpax Heo6XO,l(HMO HCKaTb reHOTHIlbl ,nJI.H ceJieKIJ,HH HYJKHhIX rrptt3HaKoB).

U:euTpbI npoucxoJK;:i:eu011 RyJILTYPHhIX pacTenuu

,------

---

U:eHTpbl
pm-1cxoJK,aeH1u1
-

1. IOarnoA3ttaTCKtttt

MecTonoJioJKeHtte

--·-·1·--·----------·1
KyJ1hTYPHh1e pacTeHttH

-+-----

Tporrtt'!ecKaH HH,attH,
Hu,aoKttTaii, o-sa IOro-

~~~:~~~· -1 ::~~:~·~
A3ttaTCKtttt

BocTO'!Hhlii Kn:Tatt,
.HrroHttH, KopeH,
TaftsaHh

3. IOro-3arra,11HoA3ttaTCKHii

MaJiaH A3HH, Cpe,aH.11.11
A3H.H, HpaH,
Acprann:cT~ H, l0ro3arra,11Han f H,!lttH

Puc, caxapHhltt
TPOCTHHK,
1\ttTpycosh!e,
6aKJia:>KaHbl 11 AP·
( OKOJIO 33 °/.1
KYJihTYPHhIX pacTe11n:H:)
CoH, 11poco, rpe'ln:xa,
lIJIO,!IOBhle H OBO~Hble
KYJihTypw - c.111sa,
BHlllH5l 11 AP·
(20 'X> KYJihTYPHhIX
pacTeHttii)
IlIIICHHl\a, pO:>Kb,
6o6osbie KYJihTYPhI,
JieH,KOHOlIJIH,perra,
'!eCHOK, BHHOrpa,11 H
,lip. (4 % KYJibTYPHhlX
pacTeHtttt)

!----------+--------------+~-------·--~

4. CpC,!IH3CMHOMOpCKHH

CTpaHhl no 6eperaM
Cpe,an:JeMHoro Mopa

KarrycTa,caxapHaH
CBCKJia, MaCJIHHbl,
KJICBep (11 %
KYJihTYPHhlX
paCTeHHH:)

!-----·----+---------------- ----------

5. 3c}lHOlICKHH

6.

U•H,PM>H~·

aMepnKaHCKHH

7. AH,11HttCKHii

A6HCCHHCKOe narophe
AcppHKH

IIO*•"' Mm~.,
I

3arra/(Hoe rro6epemLe
IOarnoii AMep1rnn:

Tsep,11aH rrllleHttua,
H'!MeHh, 6ananb1,
RocpeH:Hoe ,n;epeso,
co pro

KYKYPY"

KO<Oo

'w"

ea, Ta6aR,'xJiorr'l;THH' H
KapToc}leJIL, ananac
XHHnoe ,11epeso

~-------~-----------~-------·-

I

06~aJt

122

Ben Hoe
cKpell.\HBaII11e)

6uo:wzuIL. J'euemu1rn

rrepe1\pecTHOonhr.1aeMhIX)

AyT6pur11rnr

I'H6pHJJ;H3a:u;m1
H H6puiJuni (6JIH3Kopo,n;cTBeHHoe

CKperu;ttBam1e) HCIIOJih3YIOT .ll;JJ.H no:ry1.1c-

HH.H TaKHX JIUHHH, KOTOphie IIPH IIOCJie,n;yJOru;eM CKpen11rnaHIIH ,n;aIOT a<:\Jc}_)eKT
reTepo3Hca.

I'emep03UC ( «rtt6ptt.zi;HaH CMJia») -

HBJieHMe, nptt KOTOpOM rtt6pH.ll;bl no

PH-

,n;y npM3HaKOB M CBOHCTB rrpeBOCXO,ll;JIT po,n;ttTeJihCKMe C\JopMhI. lfaq1rna.H co BTOporo noKOJie1ma reTepo3HC 3aTyxaeT.

Aym6puiJuHi -

cKperu;ttnaHtte Hepo,n;cTBeHHhIX c}_)opM o,n;noro BIIJJ;a (HeT 06-

ru;ttx npe,n;KOB B rrpe,n;eJiax qeThipex -

OmiJaJteHHaJl w6puiJu3a4uH -

IIIeCTM IIOKOJieHHH).

CKperu;1rnamre oco6eiI (paC'reHHH), OTHOCH-

IQMXCH K pa3HhIM mr,n;aM.
OT,n;aJieHHhie rtt6pn,n;hr o6hPIHO cTepmihHhr, TaK KaK y HMX HapyrnaeTcH
MeH03 (,n;Ba raIIJIOll,Zl;HhIX Ha6opa XpOMOCOM pa3HbIX Bll.ll;OB He MOryT IWH'hlOfIIpoBaTb), II raMeTbI He o6pa3yIOTC.fI.

B 1924 r.

r.

,I(.

KaprreqeHKO C03,n;aJI KarrycTHO-pe,n;eqHbIM rII6pn,n;, IIOHBH-

JiaCh rrepBaH :\WTO.ll;HKa rrpeo.zi;oneHHH 6ecrrJIO.ll;llH y OT.ll;aJ"IeHHbIX rH6pu,n;oB.
YqeHhIH cKpecTHJI pe.n;hKY
raMeTbI -

n

=

9).

(2n

=

18;

n

raMeTbI -

=

9)

n KarrycTy

(2n

=

18;

Ho B rrpou.ecce Meii:oaa xpoMocoMbI He KOH'bIOrnpoBaJIH ~ rH-

6pn.zi;hr CTepMJibHbI.
3aTeM c IIOMOllJ;blO KOJIXIII1HHa (6JIOKHpyeT o6pa30BaHtte MMKPOTpy6oqeK uepeTeua ,n;eJieHHH) y,n;BOHJI xpOMOCOMHblH Ha6op rtt6ptt,n;a
BOHJIHCb KaK «pe,n;eqHhie»

(9 + 9), TaK n

«KarrycTHhie»

(4n

=

36),

IIpll oTOM

(9 + 9) xpoMOCOMhI.

y;:i;-

KoH'hKJ-

ral11'IH B03MOJKHa ~ raMeTbl o6pa3yIOTCH, ll ~ IlJIO.ll;OBIITOCTb BOCCTaHOBJ1eHa.

CeJieKIJ;HH mHBOTHLIX

B

ceJieKI111H JKMBOTHbIX MCIIOJib3YlOTC.H Te

nw

OCHOBHbie MeTO.ll;bI -

r116pH-

.ll;II3al1II.fI ll OT6op.

Oco6ennocmu ceJlelCU,UU JICUBOmllbtX: B OCHOBHOM IIOJIOBOe pa3MHOJKeHne;
HeMHOfOqMCJieHHOe IIOTOMCTBO; pe,n;KaH CMeHa IlOKOJieHHH.

Ce.neKijUR. MuKpoopianu3M06. 6uomexno11,ozuJ1.

123

CoepeMeHHbte (iJonoJtuumeJ1bllbte} Jvtemoabt ceJteKu,uu:

11cKyccTBemwe oce-

J.14eHeH11e, ropMOHa.'IhHaH cyneposy.'IHIJ,IIH If TpaHcrrnaHTaIJ,IIH 3N16pMOHOB, OT'AaneHHaH rII6pHJJ;II3all,HH.
,Il.oCTHmeHIIH ceneKIJ,HH:
rII6pHJJ; il:opKIUMpCKOH 6e.'IOH CBHHbH
CBHHhH

(?) -

(cf')

H 6ecnopOJJ;HOH yKpaMHCKOH

6e.irnH CTenHaH CBHHhH;

r116pMJJ; K06hinl1IJ;bI c OCJIOM -

Myn: BbIHOCJIMB, JJ;OJII'O :>KIIBeT, HO 6ecn.'IOJJ;CH.

HHorn;a raMeTorette3 rrpoTeKaeT HOpMaJihHO II nonyqaJOTCH IIJIO,lJ,OBIIThre r 11 6pHJJ;hI.

llpuMep.

ApxapoMepHHOChr (apxap 11 Mepattoc), 6ecTep (6enyra co cTep.w:-

'AhIO), XOHOP11K (xopeK 11 HOpKa).

CeJie1m;m1 MHKpoopraHH3MOB (B OCHOBHOM oaKTepHH H rpHOOB)
OcHOBHhrM11 MeTon;aMII ceneKIJ,11H MIIKpoopraHM3MOB HBJIHIOTCH 3Kcnep11l\'leHTaJihHhr:ii: MyTareHe3 II OT6op BhICOKOrrpon;yKT11BHhIX IIJTaMMOB.
Oco6eHHOCTH ceneKIJ,1111 M11KpoopraHH3MOB:
reHOM 6aKTep11:ii: rarrJI011,1J,HhIH (on;Ha KOJihll,eBaH MOJieKyJia ,UHK), II03TOMY JII06hre MyTaIJ,HH rrpOHBJIHIOTC.H yme B nepBOM IIOKOJieH11II;
oqeHh BhICOKa.H HHTeHCHBHOCTh pa3MHO:>KeHHH o6JierqaeT IIOHCK MyTaHTOB.

llpuMep. B

peayJihTaTe HcKyccTBeHHoro MyTareHeaa 11 oT6opa 6hrJI noJiyqett

mTaMM rp116a neHIIIJ,IIJIJia no aHT116IIOTHKY neHHIJ,MJIJIHHY. IIpon;yKTMBHOCTh
6hIJia yBeJI11qeHa B

1000 paa.

BnoTexuoJiorHH. reuuan HHmeuepH»
BuomexttoJtoZu.fl -

ucnOJih30BaH11e :>KMBhIX oprattH3:vIOB u

11x 611onoruqe-

CK11x npoIJ,eCCOB B npOM3BOJJ;CTBe Heo6XOJJ;HMhIX qeJIOBeKy Belll,eCTB If npon;yKTOB.
06'beKThI 6HOTeXHOJIOI'HH -

6aKTepuH,

rpII6hr,

KJieTKH paCTMTeJihHbIX

H '1CHBOTHbIX TKaHeil. Hx Bhrpalll,HBaIOT Ha IIHTaTeJihHhIX cperi;ax B cneIJ,MaJihHhIX ycJIOBHHX B 6IIopeaKTopax.
0CHOBHhie MCTOJJ;hl 6HOTCXHOJIOI'MM:
KJICTOqHa.H HH:>Kettep11.H (COMaTHqecKaH r116p11,1J,H3all,HH -

CJIHHHHe co-

MaT11qecKHX KJICTOK, ceJICKll,IIH KJICTOK);
XpOMOCOMHaH HH:>KeHepHH;
reHHaH HH:>KeHepMH.

I'eHHa.R UHJICeHepu.fl -

COBOKYIIHOCTh MeTOJJ;HK,

II03BOJIHIOII~HX

BhIL(eJIHTh HyJK-

HLiii reu II3 reHoMa on;ttoro opraHH3Ma 11 BBeCTH ero B reuoM n;pyroro opraHH3Ma.

TpattcieHHbte opiaHU3Mbt -

:>KMBOTHhie II pacTeHHH co «BCTpoeHHhIM» re-

HOM.

Tpattc<jJopMupoeaHHbte opiaHU3Mbt -

6aKTep1111 11 rp116hr c BBen;eHHhIM qy-

)kepo.n;HhIM reaoM.

lipuMep.

K11IIJeqttaH rraJioqKa

E. coli,

m1rnylll,a.H B KttIIJeqtt11Ke qeJioBeKa,

ROCJie Tpaacq>0pMaIJ,1111 rettaMH HHCYJIHHa HJIH ropMoHa pocTa rrpIIo6peJia cno006aocTh c1111Te3IIpoBaTh ropMOHhI MHCYJIMH IIJIH coMaTOTporrIIH. HcrroJih3yeT-

ea KaK npo.n;yIJ,eHT ropMOHOB.

06UfaJl 6uo;wzuJl. I'enemu1rn

124

UHTOnnA3MATH~ECKOEHACnEAOBAHME
HeKoTopaH -.:racT1> aac.1e,11,cTBe1rnoro MaTepnaJrn KJieTKII aaxo,11,IITCH B UIITon.rra3Me. 8To: KOJibUeBaH .IJ:HK MlITOXOH,11,pII:ii:; KOJihIJ;eBaH .IJ:HK IIJiaCTH,11,; BHeH,11,epHbie reHeTnqecmrn sJieMeHThI (nJia3MH,ll,hr).
HacJie,11,oBaHHe UIITon.rra3MaTH-.:recKIIX reHOB ae no,11,qHHHeTcH Mett,11,eJieBCKIIM
3aKOHOMepHOCTHM. HacJie,11,oBaHHe UHTOJUia3MaTnqecKHX renoB npoMcxo,11,MT no
MaTepHHCKOM JilIHlilI, OHM nepe,11,aIOTCH OT MaTeplI KO BCeM ee IlOTOMKaM.
IlplI OilJIO,ll,OTBOpeHHH reHeT11qecKHM COCTaB il;HTOnJia3Mbl B 3IIrOTe onpe,n;eJI.HeT-

CH UlITOilJia3MOM .aii:ueKJieTKlI, il;lITOilJia3Ma cnepMaT030H,ll,a B onpe,n;eJieHIIH reHeTMqecKoro COCTaBa npaKTII'leCKlI He yqac·rByeT.

llpuMep.

HacJie,n;oBaHHe necTpOJIHCTHOCTII y

MHorttx pacTeHHtt CB.H3aao

c MyTaUH.HMH B .IJ:HK XJ'IOponJJaCTOB II aapyrneHHeM CHHTe3a XJIOpoqmmrn.

y

qe.JIOBeKa MHTOXOH,ll,pHaJibHbIM reHOM npe,n;cTaBJieH KO.JihUeBOH .IJ:HK, co-

CTO.HI.UeM H3

16 569

HyKJieOTlI,ll,OB. qaCTOTa MyTaUlIM B MHTOXOH,11,pliaJibHOM

.IJ:HK B ,11,eC.HTb pa3 BbIIIIe, qeM B .a,n;epHOH. MyTaUHlI B MHTOXOH,ll,pHaJihHOH
.IJ:HK HBJI.HIOTCH np11q1rnoii: 6oJiee

100

6o.'Ie3ae:H • .IJ:tta6eT, snHJienc1rn, oTcpo-

qeHaaa Kap,11,HOMHOilaTIIH, 6oJie3Hb IlapKlIHCOHa rrepe,n;aIOTCH OT MaTepII KO
BCeM ee ,n;eTHM.
CpaBHlrnaa MlITOXOH,ll,pHaJibHYIO .IJ:HK rrpe,n;KOB lI IIOTOMKOB, MOiKHO ,ll,OKa3aTh po,11,CTBO KaK Mem,11,y OT,ZJ;e.JibHbIMH oco6HMH, TaK lI Mem,11,y IIOIIYJI.H!IlIHMlI.
TaKHM o6paaoM no HCKorraeMhIM ocTaHRaM 61>rJia n,11,eHT11qmu11poBaHa ceMh.H
pyccKoro uapa HHKOJiaH

II.

Bo3HUKIW6enue :JICU3nu na .1eM.rie. Pa.16umue opianu'leCKOU> Mupa

125

3BOJ1IOLl,l1SI
Bo3HHKHOBeHHe JICH3HH Ha 3eMJie.
Pa3BHTHe opraRH'lec1wro MHpa
Bee rn:rroTe3bI o B03HMKHosemn1 iKM3IBI Ha 3el\e1e :vromHo pa3ri;eJIMTh Ha TPM
rpynnhl:

1.
2.
3.

TeopMR 6ttoreHe3a -

.iKMBOe npottcxo,11,MT TOJihIW OT .iKMBOro.

TeopMR a6ttoreHeaa Teopmi: naHcnepMMII -

.iKMBOe noRBJIHeTc.H OT ttemttsoro.
.iKM3Hh Ha 3eMJIIO aatteceHa M3 KOCMoca.

B HaCTO.HIUee speM.H Hatt6oJiee npII3HaHHOH .HBJI.HeTCJI rttnoTeaa, BhICKaaattaa.H poccttiicKMM 6noxIIMIIKOM A.
CKHM 6ttoJioroM

II.

OrrapIIHhIM B

1924-1928

rr. H attrJIIIrr-

.n;m. XoJI,n;e:HttoM. CyTh rttnoTe3hI B TOM, qTo IIO.HBJieHIIIO .iKII3-

HH aa 3eMJie o6.H3aTeJibHO npe,n;mecTBOBaJIO a6Horemrne o6paaosaHtte opraHtt'leCKHX coe,n;IIHeHHH H3 HeopraHHqecKHX. licToqHMKOM SHeprIIH ,D;JI.ll 3Toro
cJiyIBHJIO yJihTpaqmoJieTosoe II3JiyqeHHe CoJIHn;a. AMIIHOKHCJIOThI,
H ,n;pyrHe coe,n;MHeHM.ll HaKailJIHBaJIIICb
CTeHI:J;HIO

«rrepBMqHoro

,n;o

caxapa

Tex nop, IIOKa He rrptto6peJIII KOHCII-

6y JlbOHa ». lIMeHHO B HeM B03HIIK.'Ia .iKH3Hb. 3Ta rHIIO-

Te3a .llBHJiaCb OCHOBaHIICM ,D;JI.ll C03,ll;aHHJI TeOpHII 6IIOII033a.

TeopuR 6uonoa3a c<t>opMym1posaua B 1947 r. aHrJIIIttCKHM yqeHhIM
,ZJ;m. BepHaJIOM. OH Bh1,n;eJIIIJI TPH CTa,ll;HH 6Horrosaa:

1.
2.
3.

A6ttoreHHoe B03HHKHOBetttte 6ttoJiorttqecKIIx MOHoMepos.
06pa30BaHIIe 6ttOJ'IOrHqec1rnx IIOJIHMepoB.
ct>opMttposamrn MeM6paHHhIX CTPYKTYP H rrepsttqHhrx opraHH3MOB (rrpo6HOHTOB).

IlepBbie ,n;Ba 3Tarra OTHOC.HT K rrepIIo,n;y XHMHqec1w:H 3BOJIIOUHM.

c TpeTbero

aTana Ha'IHHaeTC.ll 6IIOJIOrII'leCKaJI 3BOJIIOUH.ll.

1. A6uoiei-1Hoe 603HU/CfWBeHue 6uoJtoiu'lecKux .MOHO.Mepo6.
4500 MJIH JICT Ha3a,n; aTMOC<l>epa 3eMJIH

Ilo ,l];aHHbIM KOCMOJIOrHH,

COCTO.HJia

H3 raaoB: OKCII,ll;OB yrJiepo.n;a (CO, C0 2), rrapoB BO.D;hI, MeTaHa (CH 1), aMMIIaKa
(NH3) II ,n;p. CBo6o,n;HhIH KIICJiopo,n; (02) OTCYTCTBOBaJI. Ilo,11; ,n;ell:CTBIIeM KOPOTKOBOJIHOBOro YJibTpa<l>IIOJieTOBoro II3JiyqeHII.ll II rp030BblX pa3p.H,D;OB MOrJIII
npOHCXO,D;HTh XIIMII'IeCKIIe peaKUIIII, npIIBO,D;.llJUIIe K o6pa30BaHIIIO aMIIHOKHCJIOT, caxapoB H ,n;pyrHX 6IIOJIOrtt'leCKHX MOHOMepoB.

B 1953

r. aMepIIKaHcKne IICCJie,n;oBaTeJIII MIIJIJiep II IOpII CKOHCTpyttpo-

BaJiu crreu;naJihHhiii arrrrapaT, npoBeJIII sKcrrepIIMeHT II ,n;oKaaaJIH STY B03M:omHocTb.

2. 06paaoBaHue u 360JlfOU,UR 6uonoJLu.Mepo6.
,[t;aJihHell:rua.H XIIMIIqecKa.H 3BOJIIOUII.ll rrpOIICXO,D;IIJia rrpII pe,n;KOM coqeTaHHII
6Jiaronp1rn:THbIX YCJIOBIIH CIIHTeaa 6IIOIIOJIHMepoB.
·

AMep1maHCKHll: yqeHhIH C. ct>oKc B

1957

r. orrhITHhIM rryTeM ,n;oKaaaJI B03-

llomHocTh o6paaoBaHII.ll rrenTII,D;HblX CB.H3eH B pacTBope aMIIHOKIICJIOT. 3KcrrePllllleHTaJibHO 6b1JI0 rroKaaaHo, qTo B BO,D;HOM pacTBope aJiaHIIHa B npHCYTCTBHH
OC<>6oro BII,n;a r JIIIH03eMa n A T<I> 06paay10TC.H uenoqKII rroJIIIaJiaHIIHa.

06~aJl 6uoJWZUJl. ;)6(1/1,l()l{UJl

126

Oco6oe 3Ha'!e1rne JJ;.1H 611orroJUIMepoB HMeeT 06paa0Batt11e
Mo.1eKy.1,

T.

qeK B

cnoco6tthIX K

1982

caMoBOCIIpo113BOJJ;CTBy.

r. oTKph1.1 ptt603HMhI -

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PHK

PHK,

)Ke BpeM.R MOJieKyJia

JJ;HK

PHK

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o6Jial(aIOil.\He 1rnrn.111-

TM'!ecKoii aKTHBHOCTLIO. Ott OIIHcaJI .RBJiettHe caMocnJiaiicnttra -yJJ;a.1eHHH HHTpOHOB H3 MOJ1eKYJ1hI

11

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B

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MCHHOI'O xpaHeHH.R HmpopMaD;MH, TaK KaK HBJU:ICTCH 60J1ee YCTOH'!HBOH.

3. <Pop.Mupoea1-tue .ne.M6paHllbtX cmpy1Cmyp u 3eo.1t10u,wi npo6urmmoe.
Cw,n:yIOil.\HM rnaroM cTaJio ofrLe,n;MHeHMe no.'IttMepos B .n;ncKpeTHh1e rpyrrIIbI
c o6pa3oBaHHeM qacTttu; 6oJiee KpynHoro pa3Mepa -

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npomo6uo1-tmbt (:u:JIH npo6uo1-tmb1).
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Bhie opraHH3MhI -

K caMOBOCIIpo-

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Pa3BHTHe :tKH3HH Ha

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B pa3BHTHH :>RH3HH Bbr,n:eJIHIOT TPH 3Tarra:

1. 8BOJ1lOJJ;HH npoKapHOT.
2. 8BOJ'll0ll;HH O)J;HOKJieTOtIHhlX 3yKapHOT.
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I. 3eoJlJOU,UR npoicapuom.
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4

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rro.n;u;apcTBa: apxe6aKTep1rn, 3y6aKTepHH (BKJIIO'!a.H 11.Hairn6aKTep11n), 3yKapnOTbI (rpn6bI, pacTeHHH, rrpocTeiiurne, :>RHBOTHbie).

Apxe6aicmepuu -

3TO IIpoKapHOTbI, KOTopbre HMeIOT oco6b1e csoiicTBa:

a) 3K30H-ttHTpOHHYIO CTPYKTYPY rettos;
6) HX KJieTOtIHhie MeM6paHbl COCTOHT H3 H30IIpeHOH,ll;OB (3cjJHpbl yrJieB0,11,0po,n;oB), He BCTpeqaJOID;HXCH y .n;pyrHX opraHH3MOB;
B) o6HTaJOT OHH B 3KCTpeMaJibHbIX YCJIOBHHX -

B KOHI.J;eHTpnpoBaHHbIX pac-

TBopax, ropHtIHX ByJIKaHH'IeCKHX HCTOtIHHKaX, 6eCKHCJIOpO,n;HhIX CJIOHX·
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Y HHX:

3TO npoKapHOTbl, lllHPOKO rrpe,n:cTaBJieHHhie B COBpeMeHHOli

pa 3 Bumue JICUJnu na 3eM/l,e

127

a) reHOMhI He IIMeIOT :JK30H-HHTpOHHOH CTPYKTyphr;
6) OHII qpe3Bhl'l3HHO pa3H006pa3Hhl no MeTa60JIII3MY.
BamHhIH apoMopcpo3 - B03HIIKHOBeHIIe cpoTOCIIHTC3a.
3a C'l8T .n:e.HTeJibHOCTII <PoTOCIIHT83IIpy10ru;nx rrpoKapIIOT' BhI,1.\8.'IRIOil~lIX
1<IICJIOpo.n:, OKOJIO 2 MJIP,ZJ; JI8T Ha3a.n; c<PopMIIpOBaJiaCb aTMoc<Pepa, rro.n:o6Ha.H coBP8M8HHOH.
rrpHHIJ;IIrrIIaJihHO HOBhIH 3Tan B 3BOJIIOI~IIII

OihWKJtemo'lHbte ayKapuombl -

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CorJiacHo meopuu cuM6uoie1te.Ja, :JyKapIIOThI B03HIIKJIII 6Jiaro.n:ap.H CIIM6II03y Kpyrrttoii aHaapo6ttoii KJieTKII (apxe6aKTepIIII c <PepMeHTaMII rJilIKOJIII3a)
c aapo6HhIMII 6aKTepIIRMII. Attaapo6Hhre 6aKTepIIII, crroco6Hbie K OKIICJilITeJrhaoMy qiopMIIpOB3HIIIO, .n:aJIII Ha'laJio MIITOXOH,ZJ;PIIRM. CHM6II03 c IJ;IIaHo6aKTepn.HMH rrpnBeJI K IIORBJieHIIIO xJioporn:racTOB. CnM6Ho3 c 6aKTepmIMH Tuna
crrnpoxeT - K o6pa3oBaHuIO JKryTHKOB. B rroJih3Y rHnoTe3hI CIIM6uorette3a roBOPHT CTpOeHIIe opraHeJIJI (.n;ByMeM6paHHOCTb), CTpOeHue IIX ,UHK II 6eJIOKCHHTe3npy10ru;ero arrrrapaTa, a TaK:>Ke crroco6 pa3MHOJKeHIIR II qyBCTBHTeJihHOCTh
IC TeM me aHTn6noTnKaM, 'ITO n 6aKTepnu.

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8ToT npou;ecc CBR3aH c p.H.n;oM Ba:arneH:urnx apoMop<Po3oB: noRBJieHneM MnToaa, IlOJIOBoro rrpou;ecca, B03HllKHOB8HHeM MHOrOKJI8TO'lHOCTil (puc. 51).
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B KOJIOHnn KJieToK B03HnKaJio pa3.n;eJiettue <PYHKIJ;MH (nnru;eBapettne n: nepe,ZJ;BnmeHne) (pnc. 52).

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128

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Ocno6nMe 3manbi 36011-10u,uu opzanu'lec1wzo Mupa

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aa.H :IKH3Hb» ). 80Hbl .n;eJIHTCH Ha 3pbl.
KpunmOJou: apxeif (.n;peBHeifruaH )l{M3Hh); rrpoTepo3oii (paHHHH )l{H3Hb).
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8pbI ,n;eJIHTCH Ha rrepuo,n;b1. HaaBaHHH rrep110,n;oB cooTBeTCTBYIOT 11a3BaHHIO MeCT, r.n;e OHH 6bIJIH OIIHCaHbl (rrepMCKMM, KeM6pMHCKMH), JIH6o Bamaeii:llIMM rrpOHBJieHMHM iKM3HH B 3TOT rrepHO,ll; (KaMeHHoyroJibHhIH, MeJIOBOH).
HcTopHH 3eMJIM 11ayqaeTcH rro ocTaTKaM .n;peBHHX opraHH3MOB B paaJI11qHbIX reoJior11qecK11x cJIOHX. Coa.n;aeTCH rraJieOHTOJior11qec1rnJI J!eTorr11ch ocHoBHbIX 3BOJIIOil;HOHHhIX co6h1T11ii:. OrpoMHbIH BKJia.n; B pa3BHTHe rrnJieOHTOJIOrH11
c,n;eJiaJI aHrJI11ii:cx11ii: yqeHhIH q. Jia:H:TeJib (1797-1875). OH ccpopMyJmpoBaJI
nputtu,un aKmyaJtU3Ma, corJiaCHO KOTOpOMy Ha 3eMJie IIOCTOHHHO .n;ettCTBYIOT
cxo,n;Hbie cpaKTOpbI cpe.n;h1. OcHOBHbie co6hlTMH rrMeOHTOJior11qecKoii J!eTorruc11
rrpe,n;cTaBJieHbl B Ta6JI. aa c. 130.

Kpy1IHeiinme apoMopcp03bI B aBOJIIOD;HH opraHH'lec1wro MHpa
ApxeucKaR 3pa (3500-2500 MJIH JieT Ha3a,n;)
1) IloHBJieHHe m11aH11. iKH3Hh rrpe,n;cTaBJieHa rrpoKap110TaM11. Crroco6 rrHTaHHH reTepoTpOcpHbIH, aHa3po6HbIH. 06MeH Beru;eCTB - rJIMKOJIH3 (Ha1160Jiee .n;peBHHH cpopMa).
2) IloHBJieHHe aBTOTpOcpHoro crroco6a ITHTaHHH - cpOTOCHHTe3a I1 xeMOCHHTe3a, cp11xcan;1111 aTMoccpepHoro aaoTa (oKOJIO 3 MJip.n; JieT Ha3a,n;).
3) IlOHBJieH11e OKCHreHHOro cpOTOCHHTe3a l1 B pe3yJihTaTe HalWrr.1eH11e KHCJIOpo.n;a B aTMoccpepe.
4) BoaHHKHOBeH11e ,D;bIXaHHH - OKMCJIMTeJibHOro cpoccpop11JI11poBaHHH.
5) CcpopMHpoBaJIHCb .u;Ba n;apcTBa 6axTepHif: apxe6aKTep1rn H ::iy6aKTepHH
(HCTHHHhie 6aKTepHH).
Ilpomepo3oiJcKaR apa (2500-570 MJIH JieT Haaa.u;)
l) IlOHBJieHHe ayKapHOT B peayJibTaTe CJIOiKHOro CHM6H03a pa3HblX rpyrrn
IIPOKapHOT.
2) IloHBJieHHe MHT03a, Me:H:oaa, rroJioBoro rrpon;ecca.
3) BoaHHKHOBemrn MHOrOKJieToqHoCTH (l,3-1,4 MJip.u; JieT Haaa.u;). cI>opMHpoBaH11e OCHOBHbIX n;apCTB: rrpOTHCTOB, iKHBOTHbIX, pacTeHHH (B0).(0POCJieii), rpH6oB.
4) Bb1xo.u; rp1160B 11 Bo.u;opocJieii: Ha cyruy B BH.u;e JIHlllaHHHKOB. Pa3BHTHe
IIPHMHTHBHbIX coo6ru;ecTB Ha cyrue (6aKTepHH H n;11aH06aKTep1111).
5) y iKHBOTHblX IIOJ:!BJIHeTCH BHyTpeHHJ:!J:! IIOJIOCTb l1 CKeJieT (HapyiKHbIH,
BHyTpeHHHH, r11.u;pocTaT11qecKHH).

130

06w,aJt 6uO/WZUJt. 360/l/OlfUJt

OcuoBHhIC JTanhI naJieOHTOJiorn•rnc1wii JieTonucn

-----1
Spa
I

0
I
Kall:uo3oll:cKaH

I

50

Me.no Boil
Me303ofrcKaJI

IOpcKHil
TpHacoBhill:

I

Co6h1T11e

3BOJUOll,HH qeJIOBeKa
Heoreu
( qeTBepTHqHhIH) I <PopMHPOBaHHe COBpeMeHHhlX
ceMeHCTB IlTHI.\ H
MJieKOIIHTaIOill,HX
na.'!eoreu
(TpeTHqHhlH)

100

200

IlepHO,[\

>:S:
0
I?
0
A

300

@

IlocJieAHHe AHH03aBphI
TiepBhie npHMaThl
IlepBhie 11,BeTKOBh!e pacTeHHH
)J;HH03aBphI
TiepBhie IITHI!hl
TiepBbie MJieKOIIHTaIOill,He
Be.nHKoe BhIMHpaHHe MopCKHx
opraHH3MOB

KaMeuuoyroJihHhlll
(Kap6ou)

TiepBhie perrTHJIHH
CeMeHHhle rrarropoTHHKH

)J;eBOHCKHH

TiepBhle aMqm6HH
YBeJin:qeuHe pa3uoo6pa3HH phr6

400

CH.nypHllCKHll

TiepBhie Ha3eMHb1e cocy,1v1cThle
paCTeHHH

450

OpAOBHKCKHll

B3phrn pa3Hoo6pa3HH B
ceMellCTBax MHOrOKJieToqHhIX
JKHBOTHhlX

'9
350

IlaJie030HCKaH

500
550

KeM6pn:iicKn:il

650

>:S:
0
0

..."'
~

:s:A

700

~

TiepBhie JJieMeHThI cKeJieTa
JJ;oKeM6pHiiCKaH

3AHKapCKHH

I

I

I

TiepBhle ph!6hr
TiepBhie XOPAOBbre

600

I
I
II

flepMCKHH

Cl)

'

,[(HBepreHI_J:HH M.'!eKOIIHTa!Olll,HX 1

fOCIIOACTBO peIITHJIHll

250

I

I

TiepBhle MHrKOTeJih!e
MHOrOKJieToquhre JKHBOTHhie
TiepBhle CJieAhI JKHBOTHhIX

l

I

I

I

1Ia.1teo3ouc1caR apa (570-230 MJIH JieT Haaa,n;).
1)
2)
3)
4)

IlO.HBJieHIIe xop,n;OBhIX (KeM6ptt:H).
Bhrxo;:i; pacTeHHM (cocy;:i;ttcThIX) Ha cyruy (cmiyp).
06paaoBaHtte aMqm6tt:H (cTerou;e<t>aJioB) (,n;eBOH).
<l>opMHPOBaHHe penTHJIHM (Kap6oH) H IlO.HBJieHHe ceMeHHhIX nanopoTHHKOB.

131

Me303ouc1cafl Jpa (230-67 MJIH JieT Ha3a;:i;)

1)

KpynHhIM ap0Mopcj_Jo3 -

B03HttKHOBemre TenJioKpOBHOCTH. IlonBJUIJOT-

c.H nepBbie MJieKOTIHTalOII~He H HaCTO.Hru;tte KOCTHCTbie pbI6bI.

2)

PacnBeT roJioceMeHHhIX, noaBJieHtte nBeTKOBb1x pacTeHttll: • .IJ:aJlhHe:Hmee pa3BHTHe iKH3HeHHbIX cjJOpM H yBeJill'IeHHe TeMTIOB 3BOJIJOntttt.

Kai1H030Uc1caJ1 Jpa (65 MJIH JieT Ha3a;:i; -

ttacTom1~ee BpeM.H)

1)

I'JiaBHhIH apoMopcjJ03 -

2)

IlpoHCXO)J;HT cjJopMn:poBaHHe npeCMhIKaroru;n:xc.H, l'OJIOBOHOrn:x MO.JIJIIO-

aHTponoreHe3, KOTOpbIM npHBe.1 K IIO.HBJieHHJO

COBpeMeHHOI'O qeJIOBeKa.
CKOB n: ;:i;pyrux rpyrm ;:i;peBHHX mHBOTHhIX. PacnpocTpaaa10Tc.H MJieKonn:Taroru;ue.

3)
4)

IllttpOIW pacceJI.HIOTC.H O,ll;HO,ll;OJibHble TpaB.HHHCTbie pacTeHH.H.
0TCTYTIJieHHe JieCOB, CB.H3aHHOe c IlOXOJIO;:J;aHHeM H cyxoCThlO KJIHMaTa,
cTn:Mynn:poBaJio 3Bomonn:ro qeJioBeKoo6pa3HhIX o6e3h.HH (roMn:Hn:;:i;),
'ITO npttBeJio K IlO.HBJieHHIO aBcTpaJIOTIHTeKOB H apxaHTpOilOB.

,IJ;oKa3a TeJibCTBa aBOJIIOD;HH
3BOJIIOnHOHHhIM nponecc n:3yqaeTCSI pa3JIH"<IHhIMH MeTo;:i;aMn:. Kam;:i;hIH
H3 MeTO,ll;OB npe,ll;CTaBJI.HeT CBOH ,ll;OKa3aTe.JlhCTBa.
0CHOBHLie /J;Olm3aTeJILCTBa
(MeTO/];hl H3yqeHHJI) 3BOJIIOD;HH

1.

IlaJieOHTOJIOI'H"<IeCKHe: HCKOnaeMhle nepexo,D;Hhie cjJopMhl; naJieOHTOJiorn:qecKtte p.H,ll;hl.

2.

Blrnreorpacpn:qec1me: cpaBHeHn:e
qJJiop H cpayH KOHTHHeHTOB; peJUiKTbI.

3.

I CTA)Ull

5JJ)~8}5)

~t~~~
Ill CTA.llH.11

MopcpoJiorttlfeCKHe:

~~~

roMoJio-

ruSI oprattoB; py,n;ttMeHTh1; aTaBH3Mhl.

4.

3M:6pttOJIOI'H"<IeCKHe (pttc.
-

53):

«3aKOH 3apo,n;bIIIIeBOI'O CXO,!J;-

CTBa~ K. B3pa;
-

npttttnurr

peKamrTy JI.Hnuw

3. I'eKKeJISI

IIOBTOpeHue

pM6a

ca,·u1MaHJ(pa 4epeuaxa

KpbICa

l.IC.'lOBeK

npe,n;KOBbIX cpopM B OHTOreaeae

5.
6.
7.

53.

reHeTH'leCKI1e.

P111c.

BHOXHMHlJeCKHe H MOJieKyJISip-

3M6p1110Hbl scex n03BOHOYHblX Ha paHHlllX 3Ta·

5lsneH111e 3apOAblWesoro CXOACTBa.

H0-6nonorH"<IeCKHe.

nax pa3Bt.1Tt.1Sl 6onee CXOAHbl APYr c APYrOM,

1Iapa3IiTOJIOI'H"<IeCKHH MeTo,n;.

YeM Ha 6011ee no3AH111X (no 3. reKKeJllO, 1866)

132

06UfaJt 6uoJWZUJt. 360./1,IOU,.UJt

Ila.JieOHTOJIOrH'leCKHe ,ll;OKa3a TeJihCTBa

H cKnnae.MbW nepexoihtbte rpop.'llbt -

q>opMbI opraHII3MOB, cotieTaJOn:J,tte

rrptt3HaKH 6oJiee µ;peBHIIX tt MOJIO.ll,bIX rpyrrrr. Haxo.ll,KII H orrttcaHtte TaKttx
cpopM II03BOJIHIOT BOCCTaHaB.'IHBaTh qmJioreHe3 OT.ll,eJibHbIX rpyrrrr.

JipuNepbl. 1.

HxTHOCTera -

IICKOrraeMaJI cpop:vm, IWTOpaJI II03BOJIJieT CBJI3aTb

p1>16 c Ha3eMHhIMII II03BOHO'IHI>IMII.

2.

ApxeorrTepttKc (rrepBorrTttI.J.a) -

rrepexo.n-

HaJI cpopMa OT perrTMJmii: K rrTttu;aM JOpcKoro rrepttoµ;a. IIptt3HaKM perrTuJmi1:
,ll;JllIHHhIH XBOCT c Hecpocrrrn.MIICJI II03BOHKaMII, 6pJOII!Hbie pe6pa, pa3BIITbie 3y6bI.
IIptt3HaKH IITHu;: TeJIO IIOKpbITO rrepbRMH, nepe,ll;HIIe KOHe'IHOCTII rrpeBparn;eHhl B THIIH'IHbIC KpbIJlbJI.

II a.neonmo.noiu'iecKue pRObL -

aTo pR.ll,hI ttcKorraeMhIX cpopM, CBJI3aHHhie

µ;pyr c µ;pyroM B rrpou;ecce 3BOJIIOI.J.IIH tt oTpamaJOrn;tte xoµ; qm.noreHe3a.

Ilpu.Mep.

8BOJIIOI.J.IIOHHhIH pR.ll, JIOII!a,ll;eH Ha'laT B HCCJie.ll,OBaHHJIX pOCCIIH-

CKOrO naJieOHTOJiora B.

0.

KoBaJieBcKoro

(1842-1883).

HaJIII'IHe MHOrttx rro-

cJie;:i;oBaTe.'IhHO CMeHRIOll.J.HX µ;pyr µ;pyra cpopM II03BOJIIIJIO IIOCTPOHTh cjJHJioreHeTII'leCKHH PR;:\ OT aorttrrnyca .ll,O COBpeMeHHOH JIOII!a.ll,H.
IIaJieoHTOJiorntieCKHe .ll,OKa3aTe.TihCTBa MoryT 6hITh rrpttMeHeHhr K 113yqeHHIO MaKp03BOJIIOI.J.IIH (npoTeKaIOrn;e:H B MaCII!Ta6ax COTeH ThICJI'l JieT) H He HCIIOJ!h3YIOTCJI rrpH H3y'leHHII MIIKp03B0JlI0lJ.HII.

Bnoreorpac}>nqecKne .uoKa3a TeJihCTBa
BttoreorpacpttR no3BOJIJieT aHaJIH3ttpoBaTh 3BOJIIOI.J.IIOHHhie JIBJieHHJI pa3HhIX MaCII!Ta6oB OT MHKPO- ,Zl;O MaKp03BOJIIOIJ.MM.

Cpaaneuue rp.nopbt u rpayHbt.

Pa3JHI'IIIJI HJIII cxoµ;cTBa cocTaBa cjJJiophI 11 cpay-

HhI MoryT 6hITh CBJI3aHbl co BpeMeHeM reoJIOrII'leCKOro pa3,ll;eJieHHJI MaTepttKOB.

IIpuMepbt. 1.

ABcTpaJIHJI na rrpOTJIJI<eHHII 6oJiee

Jiach c µ;pyrttMH MaTepuKaMtt. B

120

MJIH JieT He coeµ;ttHR-

3TOT rrepttoµ; rrpoucxoµ;HJIO cpopMttpoBaHue

oco6oi1: cpayHhI, pa3BHBa.'IttCh cyM'laThie 11 KJIOatIHhie MJieKorr11Ta10rn;tte.

2.

CJie-

,ll;hI reoJiorutiecKoro e,n;ttHCTBa IOmHorr AMepttKM, AcppHKII, ocTpoBa Ma,n;aracKap coxpaHJIIOTCJI B coBpeMeHHOH ¢ayHe. HarrpttMep, Rrn;epttIJ.hI-ttryaHhI Ma,n;aracKapa u IOmHott AMepttKII.

Pe.nuKmbt.
coM

PeJIIIKTOBhie cpopMhI -

IlpI13HaKOB,

aTo HhIHe m11Byrn;11e Btt,n;hr c KOMITJieK-

xapaKTepHhIX .ll,.;'IJI ,n;aBHO Bh!Mepll!HX

rpyrrrr

rrpOIIIJihIX

arrox. PeJIHKTOBhie cpopMhI cB11,n;eTeJihCTBy10T o cjJJiope 11 <Payne ,n;aJieKoro
rrpOIIIJIOro 3eMJIH.

IIpuMepbt. 1.

faTTepttR -

perrTIIJIIIJI, 0611Ta10rn;aJI B HoBoi1 3eJiaH,n;HH. 8TOT

BH,11; JIBJIJieTCJI e,n;HHCTBeHHhIM HhIHe JI<HBYll.J.HM rrpe,n;cTaBIITeJieM IIO,ll;KJiacca
IIepBOJirn;epoB B KJiacce PerrTttJIHH.

2.

JlaTHMepHR (u;eJioKaHT) -

KIICTerrepaJI

p1>16a, 06ttTa10rn;aJI B rJiy6oKOBO,n;HhIX ytiaCTKax y 6eperoB BocTOtIHOH A¢pttKH.
E,n;ttHCTBeHHhIH rrpeµ;cTaBttTeJih oTpJI.n;a KttcTerrephIX pb16, Hatt6oJiee 6JIH3KHH
K Ha3eMHhIM II03BOHO'IHhIM.

3.

fttHKro ,ll;BYJIOIIaCTHblll -

peJIHKTOBOe pacTe-

HHe. B HaCTOJirn;ee BpeMJI pacrrpocTpaHeHo B KttTae 11 .HrroHHH TOJihKO KaK

.n;e-

133

yopt/JOlWZU'tec1>ue iloKa3amellbCm6a

icopaTMBHOe pacTeHMe. 06JIMK rMHKrO II03BOJUieT rrpe,ncTaBl1Th JT,peBeCHhie <PopJ\{bl, BhIMepume B IOPCKOM rrepMO,D,e.

.

BMoreorpaqmqecKMe MeTO,ll;hI no3BOJUIIOT rrnyqaTh npol!eCchI KaK MaKpo;rnoJ:IJOI!MH, TaK II MMKpmmOJIIOl!MM.

Mop<l>oJiorH11ecKHe ;:i;oKa3a TeJILCTBa
Ba3MPYIOTCH Ha OCHOBHOM rrpIIHIJ;IIne: rJiy6oKoe BHyTpeHHee CXO,ll;CTBO opraHH3MOB noKa3hIBaeT po,z:i;CTBO cpaBHMBaeMhIX <j)opM.

I'oMOJWZUJl opzaH06.

31'0 opraHhI, IIMe!Oll~IIe CX0,11;-

fOMOJIOrIIqHhie opraHhI -

Hblff nJiaH CTpOeHIIH, BhITIOJIHHIOIQIIe KaK CXO,ll;Hhie, TaK H pa3JIIIqHhie <PYHKl!HH
H pa3BHB3IOIQHeCH II3 CXO,ll;HhIX 3aqaTKOB.

IIpuMepbt. 1.

Pa3JIJiqHhre no BHeIII-

ueMY BH,ll;Y II cpyHKIJ;IIHM KOHeqHOCTII
MJieKOIIHT3!0IQIIX IIMeIOT CXO,ll;HhIH IIJ13H
CTpOeHIIH H cpopMIIPOBaHIIH: KOCTII IIJieqa, npe,z:i;rrJieqh.H, 33IIHCThH, IIHCTH, cpa-

2.

JiaHI' naJihIJ;eB.
qepena

B

pH,z:i;y

lfayqeHIIe aHaTOMHII
HII3IIIHX

II

A

BhICIIIIIX

Il03BOHOqHhIX II03BOJIIIJIO ycTaHOBIITh
rOMOJIOrIIIO KOCTeif qepena y pb!6 H CJiyXOBhIX KOCToqeK
(pHC.

y

MJICKOIIHT3!0IQIIX

54).

fOMOJIOI'flqHhie opraHhl CJie,z:i;yeT OT-

B

JIHqaTb OT aHaJIOJ'HqHhIX.
AliaJiorHqHhle opraHhl c BHeIIIHHM CXO,ll;CTBOM,

aTo opraHhI

BhIIIOJIHHIOIQHe

CXO,!J;Hhle <l>YHKIJ;HH, HO HMeIOIQIIe pa3HOe
npoucxom,z:i;eHIIe. ,[{JIH ycTaHOBJieHHH po,n;-

B

CTBa H nyTeH 3BOJIIOIJ;HH OHM He HMe!OT
3HatieHHSI.

IlpuMepbt. 1.

KomoqKH

aKau;m1,

6ap6apHca, 60.HphIIIIHHKa H emeBMKH -

P1tc. 54.

3TO 3HaJIOrHqHhie opraHhl, rrpOHCXO)J;HT

cnyxosbrx KOCTOYeK cpeAHero yxa. CpasH1-1-

•a

TenbHO-aHaToM1-1YeCK"1H

paaJIHqHhIX

3aqaTIWB

H

TKaHeH,

II() BbIIlOJIHSl!OT CXO,ll;HhIC cpyHKIJ;HH 33-

JIUITbI pacTeHHH OT noe,z:i;aHHH ;KHBOTlllil:r.t:u.

2.

fJia3a qeJioBeKa H KaJihMapa

A-

AaHCKOro,

qTo 6bIJIO IIOK333HO cneu;H-

. . . .H HCCJie,n;oBaHHHMH.

~ttPyiJuMenmbt.

l'a&LI

-

3TO

opraHhl,

yTpaTHBIIIHe

5 -

ll.111X;

MOnOTOYeK;

CTpeMeYKO;

6-

H. H. lllop-

KBaApaTHaSl KOCTb -

HaKOBanbHSl MneKOn11TalOL1.!HX;

Ball KOCTb -

npeCMblK8IO-

MneKon1-1Tat0ll.lero (113

1979). I -

Ha!l KOCTb pe -

Py,z:i;HMeHTapHhle op-

KOCTHOH pb16b1;

8-

llteMOTpSI Ha nopa3HTCJlhHOe CXO)J;CTBO

cpaBHHTCJihHO-aHaTOMHqe-

nosson11eT

HOBn8H"1Sl cnyxOBblX KOCT048K Mn8KOn"1TaIOL1.!"1X:

ll.18rGC!l;

·~emm,

p11A

BOCCTaHOB"1Tb 3BOnlOL\"10HHblH nYTb CTa-

bJISIIOTCSI 3H3JIOrHqHhIMH opraHaMH,

lllh.abIMH

roMononl!Sl opraHOB. r0Monon111

4-

3-

2-

C04118HOB-

r110MaHAH6y1111-

3y6Ha!l KOCTb;

5-

ymo-

6apa6aHHaSl KOCTb MJ18KOnHTalO-

rHOHA

134

06Ufa1l 6UOJWZUJl. 360/UO!fUJt

B qrnJioreHe3e CBOe 3Hal.Jemrn H cpymrn;mo H OCTaIOrn;HeCH y opraHH3MOB B BH.n;e He.n;opa3BHThIX o6paaoBannH:.

Ilpu.Mepbl. 1.

Py,n;nMeHTapHhie KOCTOl.JKH y KI1T006pa3HblX Ha MeCTe Ta3oBoro

noaca yKa3hIBaIOT Ha npmi:cxo:tK,n;e1rne KHTOB H ,n;eJibqlllHOB OT THIIH'IHbIX 'IeTBepOHOI'HX.

2.

Py.n;nMeHTapHbie aa,n;Hne KoHel.JHOCTH nHTOHa CBH,D;eTeJihCTBYIOT o ero

nponcxo:tK,n;eHHH OT opraHH3MOB c paaBHTbIMH KOHe'IHOCTHMH.

3.

Py.n;nMeHTap-

Hbie opraHbl y l.JeJIOBeKa: yrnHbie MblllIIJ;bl, MbIIllIJ;bl «BOJIOCHHOro IlOKpOBa».
Py,D;HMeHTapHbie opraHbI MOryT ,D;OCTHraTb y OT,D;eJibHbIX oco6eii: BH,n;a TaKHX pa3MepoB, 'ITO HanOMHHaIOT 0 npe.n;KOBblX cpopMax, -

AmaBu.J.Mbl.

ATaBHCTH'leCKHii: opraH -

Tor.n;a I'OBOPHT 06 aTaBH3Max.

aTo opraH (HJIH CTpyKTypa}, noKa-

3brna10rn;nii: « B03BpaT K npe.n;KaM »' B HOpMe He BCTpe'!aIOIIJ;HMCH y COBpeMeHHblX cpopM.

!Ipu.Mep. y

qeJioBeKa aTaBH3Mbl -

3TO XBOCTOBOM npn.n;aTOK, Uieii:HaH qmc-

TyJia (Heaapocrnaa BTopaa :tKa6epHaa rn;eJib).
0TJIHl.JHH py.n;nMeHTOB OT aTaBH3MOB:
py.n;HMeHTbl BCTpet.JaIOTCH

y

BCeX oco6eii: nonyJIHIJ;HH, aTaBH3Mbl -

y OT-

.n;eJibHbIX HH,D;HBH,D;OB;
py,D;HMeHT Bcer.n;a HMeeT onpe.n;eJieHHYIO cpyHKIJ;HIO (anneH,D;HKC y qeJIOBeKa -

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Horo pa3BHTHH HCC.'le,D;yIOTCH, TeM 6oJibille CXO,ll;CTBa o6Hapy:tKHBaeTCH Me:tK,n;y
pa3JIH'IHbIMH opraHH3MaMH.

Ilpu.Mep.

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l.JHHO'IHaH CTa,D;HH (rycennu;a -

JIH'IHHKa) HaIIOMHHaeT t.JepBeii:.

135

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B HeKoTopbIX

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no 3eMJIC. 3HalJHTeJihHhlll BKJia,11; B nOHHMaHHe l1CTOpI1I1 qeJIOBeKa KaK BH.n;a
c.n;eJiaJI q, ,IJ;apsnu. B pa6oTe «Ilpo11cxom.n;em1e tJeJioBeKa 11 noJiosoil oT6op»
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nponcxom.n;euna qeJioBeKa.
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qeJIOBeKa H BhICIIIHX (qeJIOBeKoo6pa3HhIX) o6e3hHH. IIp.H.Mbte iJ01ca3ameJLbCm6ll - KOCTHhie OCTaHKH HCKonaeMoro qeJIOBeKa n: ero npe.n;rneCTBeHHHKOB. KoC6eHHbte i)o1ca3ameJLbcmBa - cpaBHHTeJihHo-auaToMn:qecKHe, <Pnan:oJiorn:tJecKne, 6noxn:Mn:qecKne, reueTn:qecK11e .n;aHHhie, a TaKme HCCJie.n;osaun:e py.n;nMeHTapHhIX oprauoB n aTaBn:3MOB. Ha ocuoBaun:n: 3THX .n;aHHhIX Bud l/e.1106eK
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Tun XopiJo6bte
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qeJIOBeKy npn:cyrn;n: Bee qepThI CTpOeHHH H <Pn:3HOJIOl'HH, xapaKTepHbie ,ll;Jlfl
.n;aHHhIX TaKCOHOMn:qeCKHX rpynn.
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B CTpoeun:n: TeJia.
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paaBn:TaH KOpa 60JihillHX noJiyrnapnil l'OJIOBHOl'O Moara, TPH CJIYXOBhie KOCTOqKH cpe.n;uero yxa, ceMh rneHHhIX noaBOHKOB, uaJIHqHe uapymuoro yxa,
iKHBopom.n:euHe H .n;p.

137

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90 'Xi

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138

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H A3HIO. ,l1;pI1orruTeKaM 6hrJI rrpucyrn; rroJiytta3eMHhr:ii:-rroJiy,n;peBecHhIH o6pa3

mu3-

HM, rrepe,n;BHllieHMe Ha qeThlpex I{OHeqHOCT.HX II IlllTaHHe paCTIITeJihHOH IIHI11eil:.
IIpe.n;IIIecTBeHHIIK JIIO,n;eil: -

paManumeK.

OH OTJinqaJic.H yKopoqeHHoii: rre-

pe.n;He:ii: qacTbIO qeperra, I1MeB11rero, KaK rrpe.n;noJiaraeTc.H, rrptt3HaKH cxo,n;cT:lla
c qeJIOBeqecKHM, B03MOiKHO, nepe,n;BI1raJIC.H na ,ll;Byx HlliKHIIX KOHeqHOCT.HX.
llCITOJib30BaJI opy,n;H.H Tpy.n;a.

0

BpeMeHH pa3,n;eJieHll.H 3BOJIIOllMOHHblX IIOTOKOB COBpeMeHHbIX aHTporroll-

,ll;OB ll rOMllHll,ll; ceftqac npIIH.HTO cy,n;HTb ITO ,n;aHHbIM aHaJIH3a MllTOXOH,ll;pllaJibHOH

.l1;HK.

CoBpeMeHHhre rraJieOHTOJIOrH OTHOC.HT rrpMMaToB K Kpyry rrpe,n;KOB

aBCTpaJIOITHTeKOB (caMbIX paHHllX roMMHll,ll;).

0CHOBHble CT3A111111 3BOnlOl-'111111 "lenoaeKa
Bhl,n;eJI.HIOT qeThipe CT8,ll;HM aHTporroreHe3a, xapaKTepM3YIOillMec.H onpe.n;eJieHHhIM TllIIOM MCKOIIaeMoro qeJIOBeKa, ypOBHeM pa3BMTM.H MaTepttaJibHOH KYJibTypbI:

[lalleonmon,oiu..,,ecKue iJoKa3amen,bcm6a npoucxo:HCiJeuun 'len,o6eKa

139

AecmpaJtonumeKu (npe,n;IIIeCTBeHHHKH qe.'IOBeKa);
[JumeKaumponbt (,n;peBHeiiIIIne mo,n;H, apxaHTporrbr);
HeattiJepma.nbU,bl (,n;peBHHe JIIO,ll;H, rraJieoattTporrhr);
qeJtoBeK c0Bpe.Me111wzo muna, ucKonae.Mbtu u coBpe.MeHHbtU (HeoattTpOrrbr).
AecmpaJtonumeKu ( «IOmHaH o6e3bHHa») rrpe,n;cTaBJIHIOT co6oii o,n;Hy H3 rrepBhlX CTa,IJ;HM 3BOJII011,HH 'leJIOBeKa. Ilpe,n;IIIeCTBeHHHKH scex HCKOrraeMbIX H coBpeMeHHbIX JIIO,n;e:M:. BBe.'l TepMnH attrJ'IHMCKn:H aHaTOM

P.

)lapT, Hame,n;mnii

B 1924 r. Ha rore AcppnKaHcKoro MaTepnKa OCTaTKH ,n;eTcKoro qeperra. B HacTo.amee BpeMH n3BeCTHO HeCKOJibKO coTeH acppnKaHcKnx aBcTpaJiorrnTeKoB. oTo
npoMemyTO'lHbIM THI! Mem,n;y qeJIOBeKoo6pa3HbIMH o6e3bHHaMH H 'leJIOBeKoM.
Oco6eHHOCTH aBcTpaJionnTeKoB: o6HTaJIH Ha OTKpbITbIX TeppnTopmi:x, xoponio 6eraJIH, BblCB06o,n;nB nepe,n;HHe KOHe'lHOCTH, BeJin CTa,ll;HblM o6pa3 lliH3HH.
lfa-3a )R'.H3HH B caBaHHe H3MeHnJiaCb TepMoperyJIHll,HH: pe,n;y11,npoBaJiaCb
niepCTb, yBeJIH'lHJIOCb IIOTOOT,IJ;eJieHne, a TIOTepH c IIOTOM HOHOB HaTpHH KOMneHcnpoBaJiaCb nepexo,n;oM K XHill,HH'leCTBy HJIH IIOTpe6JieHneM TIOBapeHHOH
coJIH. He6oJihIIIne KJihIKH n pe311,hI He BhICTynaJm Ha,[\ 06I11,nM ypoBHeM 3y60B.
tJeJIIOCTH aBCTpaJIOTIHTeKOB 6bIJIH MaCCHBHee, 'leM y COBpeMeHHOro qeJioBeKa.
HB..z:16poBhH BhICTynaJin Bnepe.z:1, o6'beM M03roso:H noJIOCTn He6oJihIIIOH nee 600 CM 3; 3aTblJIO'lHbIH OT,[leJI 3aKpyrJieH.

He 60-

B CBH3H c rrpHMoxom,n;eHneM aBcTpaJionnTeKn rrpno6peJin « qeJioBe'leCKne » 3a6oJieBaHHH, a 3aTpy,n;HeHHe po,n;oB rrpnBeJ'lO K TOMY' 'ITO rrpenMylll,eCTBO IIOJiyqaJIH '
caMKH, pomaBinHe He,n;oHoIIIeHHhIX ,n;eTe:H. oTo rrpHBeJio K y,n;mrneHnIO rrepHo,n;a
,ll;eTCTBa H crroco6CTBOBaJIO yBeJin'leHHIO cpoKa o6yqeHHH MOJIO,n;oro TIOKOJieHHH.
BpeMH CTaHOBJieHHH aBCTpaJIOIIHTeKOB OTHOCHT K nepno,n;y OT

9 ,n;o 5

MJIH.

JieT Ha3a,n;. Ilo npO,ll;OJilliHTeJibHOCTH 3TOT nepno,[I 3Ha'lHTeJibHO ,ll;JIHHHee, qeM
BCH IIOCJie,n;yroI11,aH ll,HBHJIH3all,nH.
Ce:Hqac xoporuo H3yqeHbI ocTaTKn o,n;Horo n3 paHHHX aBcTpaJionnTeKoB act>apcKoro, Ha:H,n;eHHOro B oqmorrHn. 3TO OTHOCHTeJibHO He6oJihIIIaH o6e3bHHa:
,ll;JIHHa TeJia 110-120 CM, Bee 30 Kr, o6'beM M03ra 410 CM 3, npHMOXO,ll;Hlll,aH,
ay6bl noxomH Ha qeJioBe'leCKne, o6HTaJia

3,5-4

MJIH JieT Ha3a,n;. BnocJie,n;cT-

BHH aBCTpaJIOTIHTeKH pa3,n;eJIHJIHCb Ha ,ll;Be JIHHHH:

aecmpaJtonumeK MOW,HblU KopeHacToro TeJiocJ10meHnH, ,n;JinHa TeJia

150-155

CM, Bee

50-70

Kr, qeperr MaCCHBHhIH, o6'beM M03ra

510

CM 3; 3y6hl

KPYITHhle, BereTapnaHe11,;

'.

aecmpaJtonumeK a</JpuKallcKuu nMeJI MeHhIIIne pa3Mephr, ,n;m·rna TeJia

~:20-130

CM, Bee ,ll;O

40

Kr, o6'beM M03ra

450

CM3; BCeH,[IHhIH; TIOJIOmeHne TeJia

'4>11 X0,1J;h6e 6hIJIO 6oJiee BbIIlpJIMJieHHbIM.

·lfu ABcTpaJiorrnTeKn 3aHHMaJinch co6npaTeJibCTBOM n oxoTo:H, HCIIOJih30BaJIH
,ll;PY'!Hhie opy,n;HH: naJIKH, ,n;y6nHhl, KaMHH H KOCTn. CTpOHJIH rrpHMHTlIBHhre
LITHH -

maJiaIIIn H HaBeCbI. OrHH He 3HaJin. Bee OHH IIInpoKo pacnpocTpa-

. . JIHch no Acl>pHKaHCKOMY MaTepHKY (B03MOlliHO, rrpOHHKJIH B A3nIO, HO Bbl,.. JIH 1 MJIH JieT Ha3a,n;).
:. - · 2-3 MJIH JieT Ha3a,n; mHJin cyI11,eCTBa,

eI11,e 6oJiee 6Jin3KHe K qeJioBeKy. Homo

06utan 6uo1wzun. 3601uol{un

140

OcttOBHhie cTa.n:uu :JBOJIIOQUU qeJioBe•rn

Dryopithecus

Australop1t/1ecus

Homo erectus

06e3bRHOnO.O.o0HOe

18-9 MllH ner

Yenoee•ooOpaaHao
o6e3bRH8; eoapacr
OKOJJO 5 MJlH ner

2 MllH-500 TblC. ner

J:lpMonITTeK

Aecrpa.nomueK

PaaMePbl He6011bw11e;
11aaa10U1Me MXOIU'UlMe
e nonysblnpRMneHHOM

Poer 120-150CM,
Macca 20-50 Kr;
npsu.AOKOJK,£1ett"'e.
pyKa - xearaTenb-

>KHBOTMOe; B03pacr

'ienoae• np•MoXOARWMH; eoapacr

J:lpeaHei1w11ii 4enoee• (n11reKaHrpon,
CHHaHrpon,
rellAMb6eprc<1<il
'(enoeeK HAP·)

Homo sapiens
'ienoee• paayMHblH;
aoapacr
500-30 TblC. ner

Yenoee• paaYMHb1'1;
aoapacr

40 TblC. ner

J:lpeeH11o::i 4enoee•
HeaH.Qeprane1.1

YenoeeK
pa3yMHbl•;
lOTblC. neT

CoepeMeHHblii
ltenoeeK

.,;l
>S

s

:i:

3

cc"'

:t

nOflQ)K6Hl1rt

HeeblCOKHl1 pocr.
MaCCl'IBHbtl1
KOCTRK

HblVI opraH

HeeblCOKMH pocr
(155-165CM),
MaCCHBHbl~ KOCTRK,
KOpeHaCTbl'1,
nOXOAK8 COrHyraR

'ienoee•oo6pa3Hble
liepen H~3KH~. KOCH'! 'iepen 6onee BblCO•
3y6bl noQ>opMe
4epTbl B CTPQeHMM cxo,o,Hbl c 'te11oee4e- Ol{eHb TOflCTble, BblKlo1'1, Jlo6 HH3KMl>1,
~enlOCTll!
cicMMH
pa><eHHble HaA6POB- CKOWeHHbl", Clll.flbHO
Hble AyrM, Macc111e- pa3BMTbl0 HaJl6poaHb1e '"lefllOCT\11.
Hble JlyrH.
06beM M03ra
06beMM03ra
06beM MOJra
700-1250 CM 3
430-550CM3
OKOJIO 1400 CM3

Poer AO 180 CM,
¢>\11JH'(ecnu::i nm

coepeMeHHOIO
yenoeeica

'-iepen BblCOKMH, no6
np•MOH, 6ea cnnowHOro HaA6pOBHOro
sarwura, pa3svir noA6opoJl04Hbli< Bblcryn.
06beM M03ra
1000-1600 CM 3

06heM MC3f'a

1000-lBOC CM'

OpyAM• TPYJl•
OTCyrCTBylOT

C111creMam-.ecK111

"'

~

"2l.
8

CTaJlHblH o6pa3
>ICM3HM

lt13roroemi.io1 np1t1MM- r13rorosm:nor o6pa-

lltCOO/lb3Yf()T ecTeCT-

TlltBHbre 11:aMetfHble

eeHHble npeJlMeTbl
<a• opyJ:ll<e TPYJl•
(KOCTb. nanKa "T.Jl.)

OPYAH• rpyJla
(KOnbe, CKOnOTblH
KaMeHb, cKpe6oK,
py6HJ10)

Cr2JlHbli1 o6paa lKH3· 06u;er.reeHHbli< 06HM, COBMeCTH8s:i
OXOTa

pa3 >KM31iH, noAQep>KaHHe OfHQ

lt13roroemnor cno:«V13roroa11nio1
opy- C/10)1(He"1lUHe JPYJl~s:i
11•• rpyw. M3 KOCTM " rpyAa" MexaHMJMbl
KaMH•

6ora11Hb'e KaMeHHble Hble COCT86Hbfe

OPYA"" TPYA"
(OCTpM•. CKpe6KM,
HO>KH MAP·)

06w.ec1aeHHbrl>1 o6PoAOBa• o6u;HHa. Ha1.1eblcw1o1e 1locn1pa3 )I013Hlo1; HCnOllbCrpo1o1re11bcreo no- *-0HMR B H8yKe, rex3088HMe OrH•. Crpo- ce11eH1<1'1. no>ieneHHe H\.'!Ke, MCK)'C crne
HT8JlbCTBO O"i8rD8 M o6pRJl.OB. B03H'1KHOoeHHe '1CKyCC1Ba,
lKHJlb•. nepeble
aaxopoHeHMR
fOH"i8PHOro Aena,
3eMJ10AellMR

flumeKaHmponbt. JleaniJepma;iblfbl

141

habilis - qe.10BeK yMeJ1hiii HM('Jl l\J1HHY Te.1a llO 120 CM, nee 40---50 Kr, OfrhCM
660 CM3, BCeH,ll;HbIM. 8TH cyru,cCTBa npno6peJrn cuoco6HOCTb 113l'OTaBJII1-

JVI03ra

BaTb O,ll;HO opy,nue c IIOMOlll,bIO ,L\pyroro, a He c IlOMOU~bIO py1-: 11 3y60B, Kai{ 3TO
):\eJiaIOT o6e3hHHhI. 8To 6bUIH caMhie npocTeiiume opy;111a JCa. Ha3BaHHe opy,n;w:Htto:H KYJihTYPhI -

061w.10Tan ra.'Ib-

oJr.D;yBai1cKaH HJIH raJie'maH (TRK Ka1'

B peqHhIX ,L\OJIHHax CblpheM p;;rn H3I'OTOB:IeHHH opy.rr,ttii c.1ymtt.ria raJihKa). Opy,!1.HH qeJIOBeKa yMeJIOI'O OKOH'JaTeJihHO BbIBeJIH ero ll3 Mttpa :tKHBOTHhIX.

nMTeKaHTpOnbl (Ap0BH0MWM0 fllOAM, apxaHTpOnbl)
Homo erectus -

qeJioBeK np.RMOXO.L\Hlll,HM. BrrepBbre KOCTHhie cjJparMeHThr

nnTeKaHTpona o6Hapy:>KHJI ro.1Jiatt.zi;cKHM Bpaq

8.

,[(J06ya Hao. 5:1.Ba B

1891

r.

nume1canmpon
(o • .HBa), cu11a11mpon (KHrnli), zeitiJPJ1b6ep2c1Cuu 'LP.wBelC (I'epMamm), amJWHmpon (Am1mp), meJ1a11mpon (IO:arnaH AcjJpHKa), OJ1dyeaitc1wil nume1-caHmpon

IfaBeCTHO HeCKOJihKO cjJopM p;peBHeH:umx qeJioBeKoo6pa3Hhix:

(1..I;eHTpaJibHaH AcjJpHKa). BpeMH cyru,ecTBoBaHHH

y
a

1,5-0,5

MJIH JieT Ha3a.D;.

paHHHX apxaHTpOIIOB coxpaHHJIHCb IIPHMHTHBHbie o6e3bHHbH rrpH3HaKH,

y

II03,ll;HHX ITOHBJIHJIOCb 60Jibllie qeJIOBeqecKHX qepT.
Oco6eHHOCTH cTpoeHHH: pocT

150-175

70-80 Kr; yse.11wqHJIHCh
750-1400 CM 3 , JieBoe ITOJIY·

CM, Bee

OT):\eJihI I'OJIOBHOI'O M03I'a, o6'heM M03I'8 ,llOCTHI'aJI

mapHe 60JihIIIe rrpaBoro; Y3KHH IIOKaTbIM JI06, MOlll,HO pa3B11TbIH Ha.u,6pOBHblll
BaJIHK, MaCCHBHhie qemocTH 6e3 no,n;6opo.z:1oqfforo BbICTyna. IlOHBHJiaCb rrpaBOpyKOCTb, PYKH 6oJiee pa3BHThI; o6pa30B8J'ICJI He60.'IbUIOH CBO)l; CTOIIhI; II03BOHQqHHK rrpuo6peJI Re60JibUIOM H3I'H6; peqh Ha HaqaJibHOH CTa,n;un: cjJopMn:poBaHHH B BH,n;e JieneTa (IIOHBJIHIOTCH OT,D;eJibHhie CJIOBa: «,n;aii:», «Ha» HT. ,n;.); iKHJIH CTa,D;aMn:; 3a60THJIHCb 0 6JIH:>KHHX.
IluTeKaHTponbI He 6bIJIH TOJihKO co6l:rpaTeJIHMH. Ha oxoTHHqbnx cTo:H6wru,ax Ha:li:.n;eHhI KOCTH KpynHhIX iKHBOTHhIX. OxoTa Tpe6oBaJia TecHoro cnJioqeHHH ITJieMeHH. T:11rr OXOTbI -

o6JiaBhI H 3aca.n;bI. CTOHHKH pa3HhIX TIIIIOB CBn:.n;e-

TeJibCTBYlOT o cyru,ecTBOBaHHH pa3.u,eJieHHH Tpy.n;a. Opy,llHH Jia, KOJIYHhI,

KOIIbH (3aOCTpeHHbie rraJIKH).

KaMeHHbJe py6II-

JIIO,n;H 3aHHMaJIHCb pa3HbIMH

BH,D;aMH Tpy.n;a. OncpbITbie THIThI CTO.HHO.fC: OXOTHHqbn: Jiarep.H, MacTepCKHe ITO
,n;o6biqe KaMHH, ,n;oJiroBpeMeHHhie CTOHHKH. JKHJIHlll,a CTPOIIJIH ua OTKPhITbIX
MecTax H B rreru,epax. OcTaTKH oqaroB roBopHT 06 wcrroJih30BaHHH rrnTeKaHT·
POIIaMH OI'H.H.

HeaHAepTanb1'bl (APeBHMe nJOAH, naneoaHTponb1)
CBoe BH,ZJ;OBOe Ha3BaHirn OHM IIOJiyqn:JIH 6Jiaro.z:1apH Haxo.n;Ke B
Jiuae HeaR,L\epTaJib (fepMaHHH). 0611TaJIH

200-35

1856

r. B fJ,O·

ThIC. JieT Ra3a,n; B EBpone,

.A3uu u AcjJpuKe. BpeM.!I cyru,ecTBOBaHHH II03.L\HHX HeaH,n;epTaJihI1eB cosna,n;aeT
c nocJie.n;HHM oJie.n;eHeHHeM. HeaH,n;epTaJihll;hI 6hIJIH Heo,n;Hopo.n;Ho:ii rpyrrno:H.
Oco6eHHOCTH CTpoeHHH oTHX .n;peBHHX JIIO,n;e:H: o6'heM M03ra

1200-1700

CMa,

1leperr 6oJiee BbICOKHM H OKpyrJibIH, JI06 HH3KHH, CKOIIIeHHbIH, CIIJihHO pa3BH·
Thi Ha,ZJ;6poBHbie .z:IYI'H, HaJIHqHe rro.n;6opo.n;Ka; ,Zl;JIHHa TeJia
CllBlloe TeJIOCJIOJKeHue.

155-165

cM; MaC-

142

TexHoJionrn: o6pa6oTKH KaMHH HeaH,n;epTaJihCKHM qeJIOBeKoM OT JIHqaJiaCh
OTHOCMTeJibHO BbICOKHM ypoBHeM. 0HH C03,ll;aJIH KYJihTYPY CKpe6eJI H HaKOHeqHHKOB. TexHHKa IIOJiyqeHHH 3aroTOBOK opy,n;HM Tpe6oBaJia OIIhITa, TeXHHqeCKHX 3HaHHH, TOqHOll KOOp,ll;HHau;HH ,ll;BH)KeHHM, 6oJihIIIOrO BHHMaHHH. OrrhIT
II03BOJIHJ, ,n;peBHeMy qeJioBeKy COKparu;aTb KOJIHqecTBO BpeMeHH, Heo6xo,n;HMOro

,n;.nH

H3rOTOBJWHHH opy,n;Hii.

lfa

,n;epeBa H3rOTOB.ITHJIHCh cocy,n;hI

,n;.nH

BO,ll;hl

H <JJieMeHThl )KH.,llru;.
HeaH,n;epTa.IThllhI 6hIJIH HCKYCHhIMH oxoTHHKaMH. Ha HX CTOHHKax o6HapymeHhl CKOIIJ,eHHH KOcTeii: KpyrrHhIX mHBOTHhIX. C.nomHhie oxoTHHqhH ,n;eii:cT-

BHH

6br.nH rro,n; CHJIY cor.nacoBaHHoll: rpyrrrre HeaH,n;epTa.nhu;eB. llcrro.nh30BaJIHCh

rrpneMhI 06.naB H.ITH roHa mHBOTHhIX, TYIIIH y6HThIX )KHBOTHhIX pa3pe3aJIH
n rrpomapHBa.nH Ha orHe. ,Il;e.na.nH MexoByro rrpocTyro o,n;em,n;y. 06HapymeHHhre
3epHOTepKll ll3 KaMHH II03BO.ITHIOT rrpe,n;rroJraraTh, qTO cyru;ecTBOBaJia rrpHMHTHBHaH o6pa60TKa 3epHa.
H3MeHHJICH xapaKTep rroceJieHHii. 3aceJIH.'IHCh HaBechr, rpOThI, rreru;ephl.

,Il;JIH 3aiu;MThI KOCTPOB OT BeTpa ycTpaHBaJIHCh BeTpOBhie 3aCJIOHbI. IloHBHJIHCh
rrorpe6eHmI -

rrepBhre ,u:ocToBepHhre cJie,U:hl 3axopoHeHHH yMepIIIHX.

HaqHHaH c py6ema

40

ThIC. JieT HeaH,u:epTaJihIIhI cocyru;ecTBOBaJIH c BIIOJIHe

CJIO)KllBIIIHMHCH JIIO,ll;hMH cospeMeHHOro THIIa.

'ienoeeK coepeMeHHoro nma, MCKonaeMblM
M coepeMeHHblM (HeoaHTponb1)

Homo sapiens

-

qeJioBeK pa3yMHhIM. MaKCHMaJihHhIM reoJiorHqecKHH B03-

pacT IIOJIHOCThIO CQJOPMHpOBaBIIIHXCH B 3BOJIIOllHH HCKOrraeMbIX HeoaHTpOIIOB
paHee ou;eHirnaJicH B
Ha3BaHHe

40

ThIC. JieT.

«Kpo.MaflbOHU,bt» (TaK o6o3HaqaroT B JIHTepaType HCKonaeMhIX

HeOaHTpOIIOB) o6H3aHO H3BeCTHOMY Q:>paHIIY3CKOMY rraMHTHHKY BepxHero rraJieOJIHTa -

rpoTy Kpo-MaHhOH. ApeaJI pacrrpocTpaHeHHH KpoMaHhOHCKoro

qeJIOBeKa orpoMeH.

c ero IIOHBJieHHeM BH,ll;OBaH 3BOJIIOIIHH qeJIOBeKa 3aBep-

IIIaeTCH.
IlpH3HaKH OTJIHqHH qeperroB KpOMaHhOHII,eB OT HeaH,n;epTaJibII,eB: MeHee BhICTyrraroru;HM Brrepe,n; JIHlleBOM OT,n;eJI, BhICOKOe BhIIIYKJIOe TeMH, BbICOKHM rrpHMOH JI06, OKpyrJihIH 3aTbIJIOK, MeHhIIIHe no pa3MepaM qeThipexyroJihHhre rJra3HHIIhI, MeHhIIIe o6ru;.He pa3MephI qeperra. CQ:>opMHpoBaH rro,u:6opo,n;oqHhIH BhICTyrr qeperra: Ha,u:6poBHhIH BaJI.HK OTCYTCTByeT, qeJIIOCTH pa3BHThI CJia6ee,
3y6b1 HMeIOT MaJiyIO IIOJIOCTh. 06'beM M03ra 1000-1600 CM 3 ; pOCT ,n;o 180 CM.
CTpoeHHe cKeJieTa Ty.nos11ru;a HCKorraeMhlX HeoaHTporros He OTJIHqaeTcH OT coBpeMeHHoro THIIa, XOTH KOCTH ero 6oJiee MaCC.HBHhI.
mHJIH KPOMaHhOHllhI B cpe,n;HeM

30-50

JieT. TaKaH

me

rrpO,U:OJI)K.HTeJih-

,n;o

Cpe,u:H.HX BeKOB.

HOCTh m.H3H.H coxpaHHJiach y qeJIOBeKa coBpeMeHHoro THrra

KpoMaHhOHII,bI H3roTaBJI.HBaJIH 6oJiee cJiomHhIH Ha6op opy,u:11ll:: HOmH, HaKOHeqH.HK.H Kon11ll:, .HrJihI, raprryHhI HT. ,u:. KocTHHhie opy,u:HH 6hlJIH rrpoqHhIM.H
.H 60Jiee ,ll;OJirOBeqHbIMH, qeM KaMeHHhie. I'aprryHhI fl03BOJIHJIH 3QJQJeKTHBHO )l:06hIBaTb phl6y. IlpHeMhI OXOThI c HCIIOJih30BaHHeM 3HaHHH MeCTHOCT.H II03BOJIH-

\

'

143

CoBpeMennbiii •ie1wBeK u ezo i16011104uR (neoanmponbt)

Jiff y6HBaTh 3BepeH TbICffY:aMJL 1Ie.1oneK rro.1ylJMJI B03MO)KHOCTb npo:anrna1rna
B TPY.z:IHO.z:IOCTYIIHhIX paiiottax.
Jl\HJIH B rreUiepax, KOTOpbie HCIIOJih30BaJIHCb KPYI'JihIH ro.n; I1JII1 ,L\Jla Ce30Haoro npomttBaHM5L TaKme cTpOMJIM mMJIHUia B pe'IHhIX .z:IOJrn1-rnx, r.z:1e CKJiaJ~hr
BaJIH M3 I\aMHH CTeHbI, a KpbilllY I13I'OTOBJIHJIM M3 IIJKyp. Koqyror.u;ne OXOTHJIKM
cTpOMJIM nerKtte lllanaurn:. CypoBhre KJIMMaTH'leCKMe ycJIOBHH KpoMaHhOHI~aivr
noMoraJia nepeHOCMTh Ten,1aa o.z:1em,11,a.
Bo3JIHKJIO HCKyccTBO -

cKy.TJhIITYPhI, ttacTeHHaa JKMBOIIHCh B nelllepax. fpa-

BIOPhl JKHBOTHbIX H JII0,11,eH 6hIJU1 c,11,eJiaHhI Ha KaMHHX, KOCTHX, OJieHhMX porax.
Ilo.HBHJIHCh o6p.H,11,hI, Bepa B 3arpo6Hy10 iKH3Hh, pttTyaJihHhre rrorpe6em,rn.

CoepeMeHHblH

~enoeeK

1r1 ero aeomou,1r1s:i (HeoaHTponb1)

CyllleCTBYIOT ,[\Be OCHOBHhre TO'IKH 3peHna nponcxom.z:1etttta qeJioBeKa coBpeMettHoro THrrn:
mrroTe3a noJlu~e11mpU3Ma: qeJIOBeK pa3YMHbill B03HHK B HeCKOJibKMX :vrecTax
IIJiaHeThI H3 pa3HbIX rrpe,n;KOBbIX cpopM, rrpMHa,n;JielliaUIMX K rraneoattTporraM;
rttrroTeaa

.11w11o~e11mpu3Ma:

cyIIIeCTBOBaJio e.n;ttttoe MecTo B03HHKHOBeHna

qenoBe'leCTBa OT KaKoro-To o6r.u;ero rrpe.z:1Ka, nocne qero HalJaJIOCh aKTHBHoe
pacceneHHe no rrJiaHeTe.
Bee coBpeMeHHoe qenoBelJeCTBO rrpttHa,[IJieJKHT K O,L\HOMY Btt,11,y. 06llltt:H ypoBeHh cptt3HtiecKoro H yMCTBeHttoro paaBMTHH O.z:IHHaKOB y Bcex n10.z:1eii. BttyTpM

Homo sapiens BhI,11,en.HIOT
11y10, .Mo1120Jtouo11y10. Kam,n;a.H

BH,[la

TPH 6oJihlllMe pachi:

11ezpouo11y10, e6poneouo-

113 HHX .z:1enMTCH Ha MaJihie pacbI. PaanH'IHH

Me:m.z:1y pacaMM CBO,[IHTC.H K oco6eHHOCTHM ~BeTa KOJKM, BOJIOC, rJia3, cpopMbI
Hoca, ry6 M T • .z:I. Bo3HMKnH ::JTM pa3JIH'IMH B rrpo~ecce rrpttcrroco6Jiemrn qenoseqecKHX IIOIIyJIH~HH K MeCTHhIM rrpHpOJJ;HhIM ycJIOBHHM. IlpoMCXOJK,[leHHe
pac OCHOBhIBaJIOCb Ha npoaBJieHMH rJiaBHbIX MexaHlI3MOB J13MeHeHlIH reH0$0H.z:1a H IIOIIYJIH~lIH -

ecTeCTBeHHOI'O OT6opa lI JJ;pefapa reHOB.

3BOJIIO~lIOHHaa aaBepllleHHOCTb qeJIOBeKa OTHOClITeJihHa JI He 03HaqaeT
IIOJIHoro npeKpallleHMH II3MeHeHIIH ero 6nonornqecKnx CBOHCTB. AHTporronorH Ha OCHOBaHIIlI aHaTOMlI'leCKHX Ha6JIIO,[leHlIH rrporH03lIPYIOT IIOHBJieHMe

Homo futurus -

qeJIOBeKa 6y.z:1yUiero. BttoJiornqecKa.H

opraHnaa~n.H

coBpe-

MeRHoro qeJIOBeKa .z:1aeT B03MOJKHOCTb 6earpaHlilJHOH CO~MaJihHOH ::JBOJIIO~HM.
ct>aKTOphl aHTporroretteaa

Bo3HUKHOBe11ue npR.MOXOJ1COe11uR

rro3BOJIHJIO ocB060.z:1MTb rrepe,ri;Hne KOHe'l-

aocTn.

TpyiJoBaR iJeRmeJlbHOCmb
CmaiJ11btU o6pa3 JICU3HU -

cnoco6cTBOBaJia pa3BMTMIO Moara.
coBMecTHa.H ,L\e.HTeJihHOCTb rroaBoJiaJia coBeplllaTh

CJIOJKHylO T.HJKeJiyIO pa6oTy' KOTOPYIO HeB03MOJKHO 6bIJIO BbIIJOJIHMTb O,[IHOMY.

Bo3nUKllOBenue pe'lu u pa36umue R3btKa

crroco6cTBOBaJIM rrepe,ri;aqe MHcpop-

Man;1u1 n Koop.z:1nHa~nn ,ri;e.HTeJihHOCTH.

' OmKpbtmue u ucnoJLb306anue OZHR

rrpnBeJin K aKTMBHOMY ynoTpe6JieHMIO

Blactto:H: IllillllI, KOTopaa 6oraTa He3aMeHlIMbIMM aMHHOKMCJIOTaMH, crroco6cT-

BYio~nMn

pa3BMTMIO M03ra.

144

06UfaH 6uoJWZUH . .360/UO!fUH

Pa.Jeumue ebtcweu HepeHou aeameJLb1wcmu cnoco6cTBOBaJio o6yqemuo.
Pa.16umue a.'tbmpyucmu'lecicux HaK.WHHocmeu -- 6ec1wphICTHaJI 3a6oTa
0 6Jiare .LIPYl'HX H I'OTOBHOCTb mepTBOB8Tb ,D,JIJI ,D;pyrHX CBOHMH JIH'lHblMH HHTepecaMH.

,[l;olJ,a pBHHOBCKHH rrepHOlJ, pa3BHTHH 6HOJIOrHH
Yme B ,D;peBHOCTH aHTH'lHhie qnrnococphI o6ocHOBhIBaJIH tt.n,ett o B03MOJKHOCTH pa3BIITHJI rrp11po,l.lbI. B Tpy,D;ax ApHCTOTeJIJI

(IV B • .LIO H.

3.) H fepaKJIHTa

(VI B.

,l.10 H. 3.) I'OBOPHTCJI 0 e,D;HHOM IIJiaHe CTpoeHHJI BCero JKllBOro. B CO'lllHeHHJIX fepaKJIHTa Bbl,ll,BHraeTCJI MbICJlb 0 IIPOMCXOJK,D,eHHH BCero JKHBOI'O ll3 HeJKHBOI'O.
,l];JIJI cpe,D;HeBeKosoro neptto,D;a xapaKTepHo

HUe 0 npupoiJe:

Mema<jJuau'lecicoe npeiJcmaeJLe-

rrpttpo,D;a Hell3MeHHa, IIOCTOJIHHa, ll3H8'18JlbHO C03,l.laHa BoroM.

MayqeHtte npttpOJ.lhI, HaKorrJieHtte cpaKTH'leCKHX .n,aHHhIX, orr1IcaHtte HOBhIX
o6'heKTOB JKllBOTHOrO ll pacTHTeJihHOI'O Mttpa rrpttBeJill K IIOJIBJieHHIO CllCTeMaTllKll.
KapJI JlttHHell:

(1707-1778) -

BeJIHKHH rrrne.n,cKHH HaTypaJIHCT, spaq, co-

3.n,aTeJih CHCTeMaTHKH. B pa6oTe «CHcTeMa npttpo,n;hI»

(1735

r.) OH o6o6:w;aeT

CBOH B3r JIJI,D,bl ll C03,n;aeT rrepByIO Hay'IHYIO KJiaccmpmtau;IIIO JKIIBblX opraHH3MOB. JlIIHHell: coa,n;aJI

cucmeMamuicy - HayKy o rrpIIHIJ;ttnax KJiaccmpttKau;tttt
HOMeHKJlamypy - npaBHJia H8HMeHOB8HIIJI.

JKllBbIX opraHM3MOB, ll

KapJI JlHHHell: C'lIITaJI, 'ITO «BII.LIOB CTOJihKO, CI<OJihKO HX coa,l.laJio BecKoHe'lHOe cy:w;ecTBO».
BrrepBbie rrpe,n;JiomeHHaJI B

1623

r. KacrrapoM BayrIIHOM ,n;JIJI o6oaHatieHHJI

pacTeHIIH 6HHapHaJI HOMeHKJiaTypa 6bIJia yaaKOHeHa JlHHHeeM KaK o6.H3aTeJihHOe YCJIOBIIe .LIJI.H OIIHC8HH.H BH,n;a. JlHHHell: ycTaHOBllJI npHHII;llII tt:epapXll'lHOCTll CHCTeMaTH'leCKHX KaTeropH:H (no.n,BH.LlhI ~ BH.LlhI ~ po,n;hI ~ OTPHJ.lhI ~
KJl8CCbI). JlHHHeH BBeJI B IIOH.HTlle BH,n;a MOpcpOJIOI'll'leCKHH ll Qlll3llOJIOI'H'leCKllH KpHTepHll (B03MOJKHOCTb IIOJiy'leHH.H IlJIO,D;OBllTOI'O IIOTOMCTBa).

Tpanc<I>opMH3M
M,n;eH TpaHccpopMHCTOB orrRpaJIHCh Ha 11.n;eaJIHCTH~rncKHe sar JI.H.LlhI o soaHHKHOBeHHH MaTepllll: MaTepH.H coa,n;aHa Tsopu;OM, OH me rrpH,LJ;aJI e:H .LIBHJKeHHe; rrptt:po.n;a crroco6Ha H3MeHJIThCJI, TpaHccpopMHposaThC.H. HpKHM npe,n;cTaBIITeJieM TpaHCcpOpMll3Ma 6hIJI cppaHII;Y3CKllH 6HOJIOI'-3HII;MKJIOile,D;HCT jKopm
Jlytt: BIOcpcpoH

(1707-1788),

BhicTyrraBIIIHH npoTHB ,n;HcKpeTHOCTH BH,n;oB

B noJih3Y HenpepbIBHhIX PJIJ.IOB H3MeHJIIOIIl;HXC.H cpopM.

IlepBaH n;eJIOCTHaH KOHn;enn;HH 3BOJIIOD;HH.
matt BaTHCT JlaMapK

(17 44-1829)

m. B. JlaMapK

ccpOpMyJIHPOBaJI nepByIO 3BOJIIOII;IIOH-

HYIO KOHu;erru;HIO. CorJiaCHO rrpe,n;cTaBJieHHJIM

m. B. JlaMapKa, Tsopeu; coa,n;aJI

MaTepHIO II npttpo,n;y; Bee OCHOBHhle JKHBbie H HeJKHBbie o6'heKTbl B03HHKJIH II3

360JH01fUOIHWe y'l,e1tue 'lapaJa 4ap6una

145

MaTepHtt IIOJJ: soaJJ;e:licTBHCM rrpn:pOJJ:bI. YcJJo:arnettn:e opraHM33UMH mn:BhIX cyrn;ecTB -

pe3yJlhT3T aBOJUOUlIII.

CTyneHll 3BOJllOUllll, ee «rpaJJ;aUIIII»' .11emaT He Ha rrp.HMOM o'IIIHIIll, a llMe!OT
MttomecTBO seTseii II OTKJIOHeHwii. Bce:vi: mIIBhIM opraHII3MaM IIPIICYliJa IIaMeH'lllBOCTh Jia 3BOJIIOUHH -

3TO He06XOJJ;llMhIM c}:>aKTOP 3BOJIIOUllll. 0JJ;H3KO JJ:BH)KYliJaa CHee rJiaBHhIM cpaKTOP no

m.

B. JlaMapKy -

3TO, BO-rrepBh!X,

sHyTpeHHee CTpeMJieHne K pa3BllTHIO (T. e. meJiaHne opraHII3Ma), BO-BTOpbIX,
ynpamttemie HJm HeyrrpamHeHtte opraHOB. Ilo JlaMapKy, JJ;JIHHHa.H llleH

y

NIHo-

rnx 60.JIOTHhIX IlTIIJJ; eCTh pe3yJihTaT JJ:BYX rrpOIJeccos:

1.

TpeHHpOBKH Ha ynpailrneHna IIIeII

y

ITTHU, He meJiaBlllnx Hhip.HTb JJ:JI.H

Toro, 'lT06bI JJ:OCTaBaTh KOpM co )];Ha.

2.

:/KeJiamrn YJJ:JIIIHIITh llleIO, BbI3hIBaIOliJee npHJIHB KPOBll :H «Apyrttx cpo'IIOHJJ:OB» II YCHJieHHhIM pOCT IIIell.

OcHOBHaH TaKCOHOMII'lecKaH KaTeropII.H -

BuiJ. B KJiaccIIc}:>IIKaunn Jlm1-

He.H 6hl.JIO IIRTh TIIIIOB T3KCOHOB: pa3HOBIIJJ;HOCTh, BIIJJ:, po)];, OTPHA, KJiacc.

Bu1tap1ta.R noMeHKJtamypa -

)J,BOMHoe Ha3BaHIIe c yKa3aHIIeM po)J,a II BIIJJ:a,

HarrpIIMep AcKapn,ll,a 'leJioBe'lecKaJI

(Ascaris lumbricoides), -

TaKme 6hwa

BBe)J,eHa JlIIHHeeM.
B CIIcTeMaTIIKe JlnHHeH qeJioseK 6b1JI rrpaBHJihHO OTHeceH K oTp.HJJ:Y rrpIIMaTOB.
0)1,HaKO 3Ta CllCTeMa 6h!Jia IICKYCCTBeHHOM, OCHOBhlBa.JiaCb Ha CXOJJ:CTBe TO.JlbKO BHelllHlIX rrpll3HaKOB H He OTpama.Jia peaJihHbIX CB.H3eii B opraHII'leCKOM Mllpe. Ilo CBOHM lI)];eOJIOrlI'leCKHM B033peHlIJlM JlIIHHeii IIPIIJJ:epmlIBa.JIC.H

HU3Ma -

Kpeau,uo-

lIJJ;elI 6omeCTBeHHOro COTBOpeHlI.H )Kll3HH II ee HeII3MeHHOCTlI.

c HaKOIIJICHIIeM )J,aHHhIX lI pa3BlITlICM HaYKlI rrapa.JIJieJibHO KpeaIJ;lIOHlI3MY ocpopMlIJIOCh MlipOB033peH1!eCKOe HarrpaBJieH.11e

mpancrpopMU3M -

)RlI3Hh II3Iiaqa.JibHO co-

3JJ;atta BoroM, HOB Heii 3aJIOili.CHO CTpCMJieHlIC K pa3BlITlIIO H COBepIIIeHCTBOBaHlIIO •

m. B. JlaMapK B pa6oTe «<l>HJIOC0(1809 r.) H3Jl0ili.HJI 0Cli06bl nep60ZO u,eJlOCmHOZO 360JllOU,UOltliO-

.flpKlIM rrpe)J,CTaB.11TeJib TpaHccpopM.113Ma
$.11.H 300.JIOrH.II»

io y'lenu.R. Ha 3Ha'l.11TeJibHOM MaTepnaJie JlaMapK JJ:OKaaaJI, qTo auiJbt U3MeHR10mcR. Ilp.11q.11Ha lI3MeH'llIBOCTII - CTpeMJieH.11e K pa3BHTMIO, aaJiom.eHHOe BoroM. MexaHH3M H3MeH'lHBOCTH -

ynpam.HeH.11e n.JI.11 Heyrrpa)lrnemie opraHOB.

CorJiacHo JlaMapKy, JKM3Hh B03HHKJia B cpopMe rrpocThIX aJieMeHTOB, aaTeM
Haqa.JIC.H )J,JIIITeJihHhIM rrpouecc rrocTerreHHOro yCJIOili.HeHH.H 1I pa3B1IT.IIR OT Hll3lIIHX ct>opM K BhICllllIM. 3TOT rrpouecc 6hIJI Ha3BaH

«ZpaiJau,ueu». (JlaMapK TaK-

lKe BBC.JI TepMHHhI «6HOJIOrlI.H» 1I «6Hoccpepa».) 0JJ;HaKO JlaMapK cqHTaJI BlIJJ:hl
II ,ll,pyrHe TaKCOHOMHqecKHe KaTeropHH ycJIOBHhIMH.

3BOJIIOD;HOHHOe yqeHHe qapJI3a ,ll;apBHHa
qapna ,z::i:apBlIH

(1809-1882) -

Be.JIIIKHH aHrJIHMCKIIH yqeHbIH, 3HUHKJIO-

lle,zi;ncT, C03)J,a.JI OCHOBhl COBpeMeHHOH TeOpllll 3BOJIIOUIIll. B CBOeM Tpy,ll,e «llpollcxom,zi;eHHe BlI)J,OB rryTeM ecTeCTBeHHOro OT6opa»

(1859

r.) OH IIHCaJI, qTO

t. t'JI8.BHoii )J,Bllili.YliJeM CHJIOH 3BOJIIOUlIH JlBJI.HeTCJl ecTeCTBeHHhIM OT6op Ha OCHO-

l'ae HaCJJe)];CTBeHHOH lI3MCHqlIBOCT1I.

Ii

06uta11, 6uo1wzu11,.

146

360ll101fUR

Teopm1 :lBOJIIOIJ;HH OCHOBaHa Ha ,n;Byx 110JIO:>KeHliHX:

1.

CBoii:cTBOM :>KHBbIX opraHH3MOB HBJIHeTCH H3MeH'IIIBOCTh. fJiaBHYIO poJib

ttrpaeT

1tacJteocmeett1taR u3Mett'lueocmb.

IlpM H3MeHeHnn ycJIOBHii: cpe,n;b1 Ha-

cJie,n:cTBeHHhie II3MeHeHHH MoryT 011pe,n;eJIHTb cy,n:h6y OT,D;eJibHbIX oco6eii: H 11011yJim1Hii:. HacJie,n;cTBeHHaH H3MeH'IHBOCTb -

2. Ecmecmeettllbtit om6op -

MaTepHaJI ,n:JIH 3BOJIIOIIHH.

aTo pe3yJihTaT «6opb6b1 3a cy~ecTBOBaHHe»

H BhI:>KHBaHHe HaH6oJiee 11pnc11oco6JieHHbIX oco6eii:, Hx pa3MHO:>KeHHe.

C ,n;py-

roii: CTOpOHbl, 3TO rH6eJib HaHMeHee 11pnc11oco6JieHHbIX, HX HCKJIIO'IeHHe H3
aBOJIIOIIHOHHoro 11pon;ecca. ,r:i:apBMH BbI,ZJ;eJIHJI

eoeattue:

mpu <jJopMbt 6opb6bt 3a cyw,ecm-

BHYTPHBH,n:0By10, Me:>KBH,n:oByIO, 6oph6y c He6Jiaro11pHHTHhIMH ycJio-

BHHMH cpe,D;bl.
3Ha'!eHHe ytieHHH

q,

,r:i:apBHHa:

nepeoe ecmecmeettttottay'lttoe o6'0RC1tettue 360Jl/OU,Uu;
ycTaHOBJieHhI oeuJKyw,ue cuJtbt: ttac.!leocmeeH1taR ua.MeH'lueocmb u ecmecmeettttbtil om6op;
11pe,n;Jio:>KeHo

11pe,n;Jio:>KeHO o6'bHCHeHHe BH,l(006pa30BaHHH H

MHOroo6pa3HH BH,l(OB

B 11pHpo,n;e;
06'bHCHeH 11po11ecc C03.D;aHHH qeJIOBeKOM HOBbIX copTOB pacTeHHH H 110po,n; :>KHBOTHblX.

MmtpoaBOJIJOD;H.H

MwcpoaBOJllOU,UR -

3TO 3BOJIIOIIHOHHhie H3MeHeHHH, KOTophie 11p0Mcxo,n:HT BHY-

TpH BH,n;a, 11PHB0,ZJ;HT Kero ,n:mp<PepeHIIHPOBKe H 3aBeplliaIOTCH BH,n;006pa30BaHHeM.
TepMMH «MMKpoaBOJIIOIIHH» 6hIJI BBe,n;eH B
OT,ZJ;eJieHMH 11pon;eccoB MaKpOMacrnTa6a H

HOM CMbICJie 3TOT TepMMH 6bIJI 11pe,n;Jio:>KeH
H

H. B.

THMo<PeeBhIM-PecoBCKHM
Ilonym1n;HH -

IIonyJtRU,UR -

(1938

1927 r. IO. A.

<l>HJIHl1'IeHJw ,n;JIH

BH,n;oo6pa30BaHHH.

<I>. I'.

B

,r:i:o6p:maHCKHM

COBpeMeH-

(1937

r.)

r.).

aJieMeHTapHaJI e,IJ;HHHn;a :lBOJIIOIJ;HH

3TO 3JieMeHTapHaa, ,n;aJiee Hepa3.n;eJIHMaa e,n;MHHn;a, Ha KOTO-

poii: MOJKHO HCCJie,n;oBaTb 11pon;ecc 3BOJIIOIIHH.

IIonyJt.HU,U.H -

MHHIIMaJihHaa caMOBocnpoM3Bo,n;a~aacH rpynna oco6eii: o,n;-

Horo BH,n;a, Ha npOTH:>KeHHH 3BOJIIOIIH0HHO )l;JIHTeJibHOro BpeMeHH HaCeJIJIIO-

~aJI onpe,n;eJieHHOe npocTpaHCTBO, 06pa3y10~aH caMOCTOJITeJihHYIO reHeTH'IeCKYIO CHCTeMy H <PopMHpy10~aJI co6cTBeHHYIO :lKOJIOr:nqecKyIO mi:rny.
0CHOBHLie :lKOJIOro-reHeTH'leCKHe xapaKTepHCTHKH nonyJIHIJ;HH
3HaJI xapaKTepHCTHKH nonyJIHIIHH H ,l(HHaMHKY HX H3MeHeHHH, MO:>KHO
011eHHTb 3BOJIIOIIHOHHble npo11ecCbl B 11011yJIHIIHH.
0CHOBHbie 3KOJIOrH'IeCKHe xapaKTepHCTHKH 11011YJIHIIHH: nonyJtRU,U01l1lblU
apeaJl (CM. pa3)l;eJI «8KOJIOrHH» ); 'lUCJlettttOCmb OC06eu 6 nonyJlRU,UU; OUttaMUKa nonyJlRU,UU (KOJie6aHHJI pa3MepoB H 'IHCJieHHOCTH 11011YJIHIIHH); 603pacm1lOU u noJtoeou cocmae.

M UKpO:J60/t101fUJl
K

OCHOBHbIM

147

zeHemu'ieCICUM xapa1Cmepucmu1CaM MO.iKHO OTHeCTll l}JeHOTl1-

mi:qec1ure xapaKTepMCTMKH (Mopcp0cpH3HOJIOrHqecKHe ):

1. lJ acmoma ecmpe'lae.Mocmu mozo UJ1U u11020 <Pe11omuna ( OTpamaeT

qac-

TOTY Toro HJIH HHOro aJIJieJIH IlOIIYJIHI.J;HH).

IlpuMep. IlpH HCilOJib30BaHHH ,ll;JIH 6oph6hI c KpbICaMH aHTHKOaryMeHTa Bapcpapmrn B AHrJIHH (60-e rr.

xx B.) npOH30IIIJIO H3MeHeHHe COCTaBa IlOIIYJIH-

1.J;HH. IIocJie 3HaqHTeJihHOro CHHJKeHHH q1:1cJieHHOCTH cTaJIH B03HHKaTb nonyJIHI.J;HH c ycToHqHBOCTblO K 3TOMy H,n;y. IlpOH30IIIeJI 6eCC03HaTeJibHblll OT6op Ha ycTonqHBOCTh. Bapl}JapHH CTaJI Heacpl}JeKTHBeH.

2.

I'e11emu'lec1CaR 2emepo2en11ocmb nonyJLRU,UU, KOTopaH B03HHKaeT B pe-

ayJihTaTe MyTaI.J;HOHHOH H3MeHqHBOCTH HJIH KOM6HHaTOpHKH reHOB B xo,n;e IIOJIOBOro pa3MHOJKeHHH.
3HaqeHHe MyTaI.J;HOHHOH H3MeHqHBOCTH no,n;qepKHBaJI

(1926

r.) -

c.c. qeTBepHKOB

O,ll;HH H3 OCHOBaTeJierr nonyJIHI.J;HOHHOH reHeTHKH. IIonyJIHI.J;HH Ha-

CbIID;eHa MyTaI.J;HHMH, KaK «ry6Ka» (THIIbl MyTaI.J;HH, HX KJiaCcHl}JHKaI.J;HIO CM. B
paa,n;eJie «I'eHeTHKa» ). CoBOKynHoCTh reHoT1:1noB Bcex oco6ell: nonyJIHI.J;HH Ha3hIBaIOT

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A. C. Cepe6poBCKHii:. OT ypoBHH reHeTttqecKoro paaHoo6paaHH aaBHCHT HHTeHCHBHOCTh npoI.J;eCCOB, npoTeKarom;n:x B nonyJIHI.J;HHX.

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150

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1) Cma6uJtu3upy10w,uu om6op -

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151

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BHJI:

aTo Ka'-1.eCTBeHHhril: aTarr npo11,ecca 3Bon1011,rr11.
3TO coBoKyrrttocTh oco6eil:, 06I:1TaIOIIJ;I:1x B orrpe,n:eneHHOM apeane,

ttMeIOIIJ;I:IX 06r.11,1:1e MOpclJoqH13HOJ10rl:l'IeCKI:le rrptt3HaKtt, CB060,n:Ho CKpCIIJ;l1Ba10llJ;I:IXCJI Mem.n:y co6oil: 11 rrpe,n:cTaBJIJIIOIIJ;HX ycToil:q1:1Bh1e retteT1:1qecK11e c1:1cTeMh1.

Buo -

B

aTo coBoKynttocTh nonymu.i;1:1il:.

rrpe,n:enax O,ll;HOI'O BH,n:a Mem,n:y nonyJIHIJ,HHMI:I B03MOlliHO He60JlhllIOe OT-

JII:lqrrn no rpyrrrre rrpI:13HaKOB. lJaCTO 3TO MOmeT 6bITh CBH3aHO c reorpaclJttqe.
CKI:IM pacrrpe,n:enett1:1eM nonyn.a11,tttt.

KJtuHaJtbHaR U3MNi'lU6ocmb -

3TO reorpaclJHqecKa.H H3MeHq1:1BOCTh npH-

3HaKOB B npe,n:enax BH,n:a. BHJJ;hI HMeIOT qeTKO orpaHHqHBaIOIIJ;ttil: tta6op rrptt3HaKoB • .II:ame 6JIH3KOpo,n:cTBeHHbie BH,ll;bl OTJIHqaIOTC.H ,n:pyr OT ,n:pyra.

Xuamyc -

3TO «pa3phIB» B HenpepbIBHOM p.H,D;Y npH3HaKoB, yKa3bIBaIOIIJ;Hil:

Ha pa3JIHqHe MeJl<,n:y BHJJ;aMtt.

Kpumepuu euoa .

.II:n.H pa3,11;eJreHH.H (Bh1,n:eneHH.H) BH,ll;OB cyIIJ;eCTByeT Ha-

6op KPHTepMeB: Mop<PonorHqecKMii:; cPM3Monoro-6MOXMMMqecKMil:; aKonororeorpaclJMqecKMil: H reHeTMqecKMH -

M op¢0Jtoiu'lec1Cuu 1Cpumepuu
peHHero

CTpOeHH.H

rJiaBHbIH Kpl1Tep11ii: BM,n:a.

onpe,n:eJI.HeT cxo,11;cTBO BHellIHero H BHYT-

oco6eil: ,11;aHHOI'O BH,D;a • .II:oJiroe BpeM.H

6bIJI rJiaBHbIM

H e,D;HHCTBeHHbIM, HO OKa3aJIC.H He,n:ocTaToqHbIM, TaK KaK cyIIJ;eCTBYIOT BH,ll;hl-,ll;BOHHHKM.

IlpuMep.

CyIIJ;eCTByeT KOMITJieKC

BH,n:OB-,n:BOMHHKOB,

BllIMXC.H no.n: Ha3BaHMeM «MaJI.HpHHHbIH KOMap»

paHee o6'be,D;HH.H-

(Anopheles maculipennis).

B O,ll;HHX perttoHax OH KOpMHTC.H Ha qeJIOBeKe, B ,n:pyrMX -

Ha ,ll;OMallIHHX .iKH-

BOTHhIX; pa3MHO.iKaeTC.H B COJIOHOBaTOH BO,D;e, B HCKOTOpblX perMOHax pa3MHOmaeTC.H I:ICKJIIQqHTeJihHO B npecHOH BO,n:e. lfayqemre IlOKa3aJIO, qTo cyIIJ;eCTBYeT IlICCTb Bl1,ll;OB-,ll;BOHHI1KOB, npaKT11qecK11 Hepa3JII1ql1MbIX Ha CT3,ll;I111 11Maro,
HO OTJII1qaIOIIJ;I:IXCH no CTpOeHI1IO .HI111,, Jll1ql1HOK.

<Pu3UOJl0?0-6UOXUMU 1teCICUU 1Cpumepuu.

<f>H3110JIOr0-6HOXI1MI1qecKoe CX0,11;-

CTBO Ha6JIIo,n:aeTCR 60Jihille y 6JIH3KI1X BH,ll;OB, qeM y ,D;aJieKHX.

IlpuMep.

Pa3BHTHe MOJieKyn.HpHoil: 6110JiorHH no3BOJII1JIO ycTaHOBHTh cTpo-

eHHe nHrMeHTOB KpoB11 y p.H,n:a .iKMBOTHhIX. CTpoeHHe reMorJio6HHa oKa3aJIOCh
oqeHh CXO,ll;HhlM y CI1CTeMaT11qecKH 6JII13Kl1X BH,ll;OB.

IlpuMep 6Hympu6U0060U 6uOXUMU'leCICOU U3MeH'lUBOCmu. y

co6aK KOHeq-

HbIM npo,11;yKTOM a30THCTOI'O o6MeHa .HBJIHCTC.H aJIJiaHTOI1H, KOTOpbIH Bbl,ll;CJI.HeTC.H B Moqy. Ho y O,ll;HOH H3 nopo,11; B Moqe HaKaIIJII1BaeTC.H Moqesa.H KHCJIOTa,
:'ITO xapaKTepHO ,ll;JI.H qeJIOBeKa 11 BbICillHX npHMaTOB.

81C0Jtoio-ieoipa¢u'lec1Cuu 1Cpumepuu.

Ka.m,n:hr:H BI:l,11; xapaKTep113yeTc.H cso-

HM reorpaclJ11qecKHM apeaJIOM 11 co6CTBCHHOH cpe,n:oil: o6HTaHH.H -

3KOJIOrHqe-

CKOH HHIIIe:H. 0,n:HaKO cyIIJ;eCTBYIOT BH,ll;bl c COBrra,11;aIOIIJ;I1MH apeaJiaMI1 II BH,ll;blKOCMOilOJII1Tbl (,n;OMOBa.H Myxa, BOp06bH).

I'enemu'lec1Cuu 1Cpumepuu -

ocHOBHOH Kp11Tep11ii: s11,n;a. Ka.m,n;hrii: BH,n; xa-

paKTep113yeTCH onpe,n;eJieHHbIM Ha6opOM xpOMOCOM:

q11cJIOM,

napHOCTblO,

06pa3oBanue Buao6 -

pe3y11,bmam Mu«po:J6011,104uu

153

cTpoeHIIeM xpOMOCOM, BbIHBJieHHhIM c IlOMOlll;hlO ,n;m:})cpepeHIJ;Ha.JbHOH OKpacKfl, -

KapHOTHilOM.

llpwnep.

AttaJIH3 xpoMOCOMHoro Ha6opa no3BOJill.'I pa3,n;e.1HTL e,n;HHhitt Btt,n;

noJieBKH o6bIKHOBeHHOH (

46 ( Microtus arvalis)
Bn,n;

H

M icrotus arvalis)

Ha ,n;sa no,n;sH,n;a, co,n;epma1QHe

54 ( Microtus Sub-arvalis)

xpoMocoMhr.

reHeTttt-IeCKH ycTOH'IHBa.H cHcTeMa, coxpatt.H10ru;a.H cB010 IJ;e"1ocT-

HOCTh.
06pa30BaHHe BH)l:OB -

Buaoo6pa3oeattue -

pe3yJihT3T MHKp03BOJIIOIJ;HH

aTo pa3,n;eJieHHe (so speMeHH JI npocTpaHcTse) npem,n;e

e,n;1rnoro BH,n;a Ha ,n;sa JIJIH HeCKOJihKO. BH,n;oo6pa3oBaHHe npoHcxo.n;HT B pe3y.'IhTaTe nOCTO.HHHO COBeprna10ru;HXC.H BHyTpH Bll,ll;OB

npor~eCCOB

MHKP03BOJIIOIJ;HH.

BbI,n;eJI.HlOT TPJI OCHOBHhIX nyTH JI ,n;sa cnoco6a BH,n;oo6pa3oBaHH.a.
BoJihIIIOH BKJia,n; B JI3yqettHe BJI,n;oo6pa3oBaHJIH Bttec 3. Mailp. Ott BhI,n;eJIHJI

56):
1. <PuJtemu'leCICOe euaoo6pa306GHUe. BJI,n; A 11peBpaw;aeTC.H B xo,n;e 3BO.'Il0IJ;JIJI B BJI,n; B. qHCJIO BJI,n;os He H3MeH.aeTc.a.
2. I'u6puaozeHHOe euaoo6pa306GHUe. 3TOT llYTh CB.H3aH CO CJIJI.HHHeM ,ll;BYX
cyw;ecTByrow;Hx BJI,n;OB AH B H o6pa3oBaHJieM BH,n;a C.

TPJI OCHOBHbIX nyTJI, Be,n;yw;Jie K 110.HBJieHHIO HOBbIX BH,ll;OB (pile.

llpuMep.

KyJihTypHa.H CJIHBa -

ecTecTBeHHhIH TeTpa11JioH,n;, B03HJIKIIIHH

JI3 TepHa H aJihl'IJI B pe3yJihTaTe llOCJie,n;y10w;eil llOJIJif!JIOH,n;H3aIJ;JIJI llPH
CJIJI.HHJIJI raMeT CJIJIBbl H Teptta.

3. /(uBepzeHmHOC euaoofipa.J06GHUe (paCXOJ/CaeHUe).

lfa O,ll;HOrO BJI,n;a B pe-

3YJihTaTe o6pa3yIOTC.H ,n;Ba. 3TOT llYTh, 110-BJI,ll;HMOMy, HaJI60Jiee pac11poCTpaHeHHhIH.
3TOT llYTb 11epBbIM yKa3aJI q, ,n:apBJIH, Bhl,ll;BJIHYB

quu:

BJI,ll;hl

CTpeM.HTCH

K

npUHL{Un aueepieH-

MaKCJIMaJibHOMY 3allOJIHeHJilO MeCT o6HTaHHH

H c11oco60B cyw;ecTBOBaHH.H. IlocTO.HHHOe o6pa30BaHJie OTJIJI'IHH H 11pHCllOco6JieHJIH K

pa3HhIM MeCTaM o6HTa-

HH.H COCTaBJIHeT OCHOBY 11pou;ecca ,D;HBepreHu;HH (pHc.

c

A

B

A

57).

OcHOBHbte cnoco6bt

BH,n;oo6pa3oBa-

HHH 6a3HpylOTC.H Ha B03HHKHOBeHHH
Pa3JIH'IHo:ll H30JIHIJ;HH BHYTPH BH,n;a.

1. CuMnampu'leCICOe euaoo6pa3o6llHUe (OT JiaT. sym - O.D;HHaKOBhlH, patria - po,n;HHa). BM,n;oo6pa30BaH.He llPOJICXO,ll;JIT B

O,n;Horo apeaJia. lfaOJI.HIJ;HH JIOr.HqecKa.a,

3KO-

qaw;e reHeT.H'lecKa.a

(B03HHKHOBeHHe xpOMOCOMHbIX 11e-

B

A

A

P111c.

56. TpH OCHOBHblX nyrn, BeAYlllHe K

nm1sneH1<110 HOBblX BHAOB. Cnesa Hanpaso <1mnernyecKOe BHAOo6pa3osaHHe, rn6PHAHOe npOHCXO~AeHHe BHA8, npoHC-

pecTpoeK).

llpuMep.

A

11pe,n;eJiax

Ilp.H B03HHKHOBeH.HH

TeTpa11JioH,n;Horo pacTeH.HH ( 4n)

XO~AeHHe

BHAa

( pacxo~eHHfl)

nyTeM

AHBepreHL\HH

154

06U{UK 6UOJWZUK. J60JUOlfU.ll

7

P1tc. 57.

A111sepreH1.1111si nepeAHVIX KOH84HOCTe~ no3BOH04Hb1x .>KVIBOTHblX: 1 -

raHTCKoro 6poHeHOCL1a;

3-

Pbl.>K8~ B8Y8PHVll.lbl;

4-

roplMlllbl;

5-

KPOTa;

6-

<t>111Hsana; 2 -

CVIByYa;

7-

rn-

llOW(!AVI

np.>KeBaJlbCKOrD

rH6pH,IJ;bI

c

( Nicotiana)

(2n) 6y,n;yT cTepm1bHhl. B po,n;e Ta6aKa
Solanum) ttcxo,n;Hoe 'IHCJIO xpoMocoM paBHO 12,

,n;HrrJioH,n;HhlM
H KapTo<PeJia (

24, 48, 72 xpOMOCOMaMH.
2. A.nJlonampu'lec1eoe 6uaoo6pa3o6anue (oT JiaT. altos -

HO eCTb BH,IJ;bl c

pa3Hbiii:,

patria -

po,n;HHa). 3TOT crroco6 BH,n;oo6pa30BaHHH OCHOBaH Ha rrpocTpaHCTBeHHOH
H30JIHD;Hli. HOBbie BH,IJ;hl MOryT B03HHKHYTh B peay.TihTaTe pacrra,n;eHHH IIIHpoKoro HCXO,IJ;Horo apeaJia Ha HeCKOJihKO ,n;pyrHX, MeHbIIIHX.

Ma1tpoaBoJ11ou;ua
Ma1epoaBOJl/OU,UR -

3TO aBOJIIOD;HOHHhle rrpou;eccb1 Ha Ha,n;BH,IJ;OBOM ypoBHe,

IIPHBO,ll;HID;He K o6pa30BaHHIO TaKCOHOB Ha,n;BH,IJ;OBOro paHra.
Ilpou;ecc MHKpoaBOJIIOD;HH rrpo,n;oJimaeTca 6ea KaKoro-JI1160 rrepephIBa H Ha
MaKp03BOJIIOIJ;HOHHOM ypOBHe. lfaMeHaeTca TOJibKO B03MOiKHOCTh CKpeiu;1rnaHHH Mem,n;y BH,n;aMH. HoBhie BH,ll;hl MoryT BCTyIIaTh B MeiKBH,IJ;OBbie OTHOIIIeHHH,
'ITO BJIHHeT Ha <PaKTOphl 3BOJIIOD;HH H, B CBOIO O'!epe,n;h, ,n;eiiCTByeT Ha MHKp03BOJIIOIJ;HOHHhle rrpou;eccw. Co6wTHH MaKpo3BOJIIOIJ;HH oxBaThlBaIOT MHJIJIHOHhl JieT, H HX H3y'!eHHe OCHOBaHO Ha ,n;pyrHX MeTo,n;ax (rraJieOHTOJIOrHH, Mop<lJo<PH3HOJIOrHH, CTpoeHHH

.D:HK H

6eJIKOB pa3JIH'!HbIX rpyrrrr).

155

OcnoBnbte nanpaBRenuJl 360RIOIJUU

<PuJIO?ellCJ -

3TO IICTOpIIqecKoe pa3BIITIIe opraHII3MOB, KaK B u;eJIOM TaK-

coHe, TaK II B OT,ll;eJibHhIX TaKCOHax. BhI,ll;eJI.lUOT qeTbipe OCHOBHhie cpopMbl <PIIJIOreHe3a:

1. <PuJiemu'leCICa.H 360JI,IOU,U.H -

3TO II3MeHeHII.H, ITpOIICXO,ll;.HIIJ;IIe B O,ll;HOM <PIIJIO-

reHeTIIqecKOM CTBOJie (6e3 yqeTa B03MO./KHhlX ,ll;IIBepreHTHbIX OTBeTBJieHIIM).

IlpuMep.

Pa3BIITIIe ITpe.n;KoB JIOIIrn.n;eii ITO rrp.aMoi1: JIHHIIH: aorIIrrITyc -

MHOrHrrITyc -

rraparHrrITyc -

2. J1ueepienu,u.H -

coBpeMeHHa.H JIOIIIa,ll;h.

pacxom.n;eHIIe BeTBeii .n;peBa .JKH3HH OT e.n;irnoro CTBOJia

ITpe.n;KOB.

IIpuMep.

Bo3HHKHOBeHIIe paaHoo6pa3HhIX ITO Mop<PoJiorIIqecKHM oco6eHHO-

cT.HM BblOpKOB, HarrpHMep Ha faJiarraroCCKHX OCTpoBax, OT e.n;IIHOro rrpe.n;Ka.

3. KoHeepieHU,U.H -

aTo rrpou;ecc qiopMnpoBaHH.H cxo.n;Horo <PeHOTHIIIIqe-

cKoro o6JIHKa oco6ei1: .ll:BYX HJIII HeCKOJihKHX rpyrrrr.

IIpuMepbl.

<l>opMHpOBaHHe CXO,ll;HhIX <PopM TeJia y aKyJIOBbIX, HXTH03aB-

poB H KIIT006pa3HhIX. Ilpe.n;cTaBHTeJIH pa3HbIX <PIIJIOreHeTIIqecKHX rpyrrrr
HMelOT 60JihIIIOe CXO,ll;CTBO B CTpOeHHH B CB.H3H c IIPIICIIOC06JieHHeM K ./KII3HH B BO,ll;HOM cpe.n;e.
Cxo.n;cTBO cTpoeHH.H rJia3 MJieKOIIIITaroiu;nx II roJioBoHornx MOJIJIIOCKOB.

4. IIapaJIJI,eJIUJM -

qiopMHpoBaHIIe cxo.n;ttoro o6JIHKa H3HaqaJihHO .n;HBep-

reHTHhIMH, HO reHeTIIqecKII 6JIH3KHMH rpyrrrraMH. IlapaJIJieJIH3M -

3TO

KOHBepreHIJ;H.H Ha OCHOBe roMOJIOrHqHhIX opraHOB.

IlpuMep.

Pa3BHTHe ca6Jie3y6oCTH B CTBOJie KOIIIaqhIIX B03HHKaJia Heo.n:-

HOKpaTHO.

OcHOBHLie HanpaBJieHHH aBOJIIOIJ;HH
KpyrrHeMIIIIIe poccniicKIIe SBOJIIOIJ;HOHIICThl A.

H.

1.
2.
3.

IL IL
58):

CeBepu;oB H

rayaeH Bbl,ll;eJIHJIH TPH rJiaBHbIX HarrpaBJieHH.H 3BOJIIOIJ;IIH (pHC.

IllMaJih-

AporeHe3 (apoMop<Po3).
AJIJioreHe3 (n.n;noa.n;arrTau;nn).
KaTareHe3 (o6iu;a.a .n:ereHepau;n.a).

ApoieneJ -

KpyrrHhle aBOJIIDU:HOHHhle H3MeHeHH.H, Be.n;yiu;ne K o6iu;eMy yc-

JIO.JKHeHHIO CTpoeHH.H H <PYHKIJ;IIH opraHH3MOB, B pe3yJibTaTe qero 3aTpy.n;H.HeTC.H OCBOeHIIe HOBbIX ITpHpO,ll;HhIX 30H, IIO.!IBJieHHe KPYIIHbIX rrpHCIIOC06JieHHH.

Apo.;nop<jJ03bl -

KpyrrHwe a.n;aITTaU:IIH, rrpHBO,ll;HIIJ;He Ha rryTh aporeHe3a (Ha

CXeMe BhIXO,ll; H3 O,ll;HOH IIJIOCKOCTH B .n;pyryro).

Ilpu.Mepbl.

IIOHBJieHHe JierKHX y Ha3eMHhIX II03BOHoqHbIX II03BOJIHJIO OCBO-

RTb HOBYIO cpe.n;y o6IITaHHH, cyIIIy. B SBOJIIDU:HII MJieKOITHTaroiu;nx K apoMopll>o3aM MO./KHO OTHeCTH IIO.HBJieHHe IIIepcTHOro IIOKpoBa, ./KHBOpom.n;eHIIe,
BCKapMJIHBaHHe .n;eTeHbIIIIeii MOJIOKOM.

AJIJI,o?ene3 -

pa3BHTHe rpyrrrrhr BHYTPH o.n;tto:H a.n;arrTHBHOM 30Hhl, a.n;arrTa-

n;a:H o.n;Horo MaCIIITa6a.

H iJuoaiJanmau,uu -

a.n;arrTaU:HH o.n;Horo MaCIIITa6a rrpH

ll>opMnpoBaHHH pa3Hhlx rpyrrrr B rrpe.n;eJiax o.n;Ho:H a.n:arrTHBHOH 30HhI (pacxo.n:aIIJ;aeca BeTBH Ha O,ll;HOH IIJIOCKOCTH).

06~aJl 6UO.!WZUJl. ;J60.!llOlfUJl

156

8peMR
P111c. 58. CxeMa pa3B111rn5i rpynnb1 no nyrnM annoreHe3a BHyrp111 C1Aam111BHOH 30Hbl (Ha nnocKOcrn) 111 aporeHe3a -

IlpuMepbt.

c BbiXOAOM B HOBYIO CJAanTlllBHYIO 30HY (B HOBYIO nnOCKOCTb)
IloaBJiemrn KphIJia -

apoMopcpoa. llaMeHemfe ¢opMhI KphIJia

B CBH3H c oco6eHHOCTHMH IIOJieTa IITHll Ka -

H.!l;HOa,!l;aIITa1IHH. IlOHBJieHHe llBeT-

KPYIIHhIM apoMopcpo3. ll3MeHeHHe B cpopMe u;BeTKa B CBH3H co c11oco60M

OilbIJieHHH -

H.!l;HOa,!l;aIITau;HH.

KamaieHe.3 -

aBOJIIOIJ;HOHHhrii: rrpou;ecc, IIPHBO.!l;HillHH K ynpo111eHHIO opra-

Hlfaau;HH.

0611.{aJl iJeieHepaU,UJl -

11cqe3HOBeHHe CHCTeM opraHOB H CTpyKTyp, TO eCTb

MOpcpocpH3HOJIOrHqecKaH ,!l;erpa,!J;au;HH.

IlpuMepbt.

Ilapa3HTJfqecKHe cpopMbI COCTaBJIHIOT

4-5 °Ai

OT 06111ero q11cJia

COBpeMeHHhIX BH.!l;OB :>KHBOTHhIX. Ha Kpa6ax rrapaaHTHpyeT paKoo6paattoe caKKYJIHHa. Bo B3pOCJIOM COCTOHHHH, HaXO.!l;HCb Ha TeJie Kpa6a, OHa HMeeT BH.!I;
MeIIIKa, Ha6HTOro IIOJIOBhIMH npO.!J;YKTaMH, c OTXO.!l;HillHMH OT Hero OTpOCTKaMH.

ll

TOJlbKO H3yqeHHe JIHqHHO'lHblX CTa,!l;HH 1103BOJIHJIO BhlHCHHTb npHHa,!l;-

Jie:>KHOCTb aToro BH,!l;a K paKoo6pa3HhIM. "'YTpaTa IIHilleBapHTeJibHOH CHCTeMhl
y KJiacca JleHTOqHbie qepBH IlpOH30IIIJia B peayJibTaTe 11apa3HTHpOBaHHH B TOHKOM KHIIIe'lHHKe X03HHHa.

BHoJiorH11ecKHH nporpecc H perpecc B aBomou;HH

BuoJloiu'tecKuu npoipecc -

aTo TaKoii: X0.!1; aBOJIIOI.J;I:IOHHoro npou;ecca,

np11 KOTOpOM: IIOCTOHHHO HapacTaeT 'lHCJieHHOCTb oco6eli B IIOIIYJIHIJ;HHX; pacIIIHpHeTCH apeaJI; yBeJIH'lHBaeTCH pa3H006paa11e a,!l;anTaIJ;HH.
B110Jior11qecKHH nporpecc opraHH3MOB.

aTo ycnex 11 npou;BeTaHHe KaKoli-Jin6o rpynnhr

Ocno6Hbte no.1toJ1Ce1tuR cunmemu'iec1wii meopuu 360.!tlO!!UU

157

Han6o;:iee qacTo 6ttoJior11qecKHH uporpecc o6ecneqHBaeTcH 3a cqeT nOHBJieHMH apoMopcpo3oB. 0.n;HaKo cyr.u;ecTBYIOT rpyrrilhI, npeTepuemrme .n;ereHepaIJ;MIO CMCTeM H opraHOB, HO 3a cqeT 3TOro npttcrroco6HBIIIHCCH K HOBhIM YCJIOBHSiM M urnpOKO pacnpocTpaHMBIIIHeCH.
BuoJtoiuttecKuu peipecc - aTo rrpo1~ecc upoTttBono.1omHoro HarrpaBJieHMSi. ,l(JIH Hero xapaKTepHO yMeHhIIIeHHe '-IHCJieHHOCTH oco6eil: B llOIIYJIHI~H
S{X, COKpar.u;eHHe apeaJia, COKpar.u;eHHe qHCJia BH,ll;OB (BIIJIOTb ;:i:o 3JIHMHHaIJ;MM).
BnoJiorttqecKHiI perpecc qacTo CB.H3aH c .n:ereHepautte:iI M Bcer.n;a rrpttBOJJ:HT
K yua.n;Ky. B 3TOM 3aKJIIOqaeTcH oTJIHqtte 6ttoJIOI'H'-IecKoro perpecca OT o6u~eil:
,ZJ;ereHepauHH. BttoJiortt'-IeCKHH perpecc MOJKeT 6hITh He CBH3aH c .nereHepautteii:. HaupHMep, BhIMMpaHtte JJ:HH03aBpoB CBH3aHo c Herrpttcrroco6JieHHOCThIO
K H3MeHHBIIIeHCH cpe.n;e, 6e3 .nereHepauHH KaKHX-JIH60 CHCTeM.
OcnoBHbie rroJio~eHHH CHHTeTHqecKoii TeopHH 3BOJIIOD;HH

1. MaTepttaJI aBOJIIOIJ;HH -

2.
3.
4.

5.
6.

7.

8.

HacJie,ncTBeHHhie H3MeHeHMH (MyTauHH). MyTauHOHHaH H3MeHqHBOCTh HOCHT c.'Jyqaii:HhIH H HeHarrpaBJieHHhIH xapaKTep.
E,n;HHCTBeHHhIH ,nBHJKyr.u;ttii: cpaKTop - oT6op cJiyqaiIHh1x MyTaunii:, B03HHKaror.u;ttx B reHocpoH.n;e llOIIYJI.HuHH.
HattMeHhIIIaH e.n;ttHHua 3BOJII011HH - uorryJIHliH.H.
3BOJll011HH HOCHT JJ:HBepreHTHhlll xapaKTep, TO eCTh OJJ:HH TaKCOH MOJKeT
CTaTh upe,nKOM HeCKOJlhKHX ,n;oqepHHX TaKCOHOB.
06pa3oBaHHe HOBoro BH.n;a - aTarr aBOJIIOliHOHHoro upouecca. Btt,noo6pa3oBaHHe upottcxo.n;HT uocTerreHHO H ,nJIHTeJihHO. Btt,noo6pa3oBaHtte IlOCJie,ZJ;OBaTCJihHOCTh CMeHhl uorryJIHliMH.
Btt.n; - peaJihHO cyr.u;ecTByIOr.u;aH e.n;HHMua. Btt.n; o6Jia,naeT .n:irnaMHqHocThIO (TOJihKO BHYTPH Bcer.n;a B03MOJKeH o6MeH aJIJieJIHMH). Btt.n; o6Jia,naeT
ueJIOCTHOCThIO H 3aMKHYTOCThIO: cyr.u;eCTBYIOT MexaHH3Mhl rro.n;.n;epJKaHHH ueJIOCTHOCTH Ha YPOBHe 11011YJIH11HH (reorpacpH'-IeCKaH, 6HOJIOrWieCKa.H, reHeTttqecKaH H30JIH11HH) H Ha ypoBHe reHocpoH.n;a (rpyrrrrhI cuerrJieHHH, CHCTeMbl peuapauHH).
Peupo,n;yKTHBHaH H30JIH11HSi KaK KPHTepttii: BH,n;a HerrpttMeHHMa K rpyurraM opraHH3MOB 6e3 IIOJIOBOI'O upo11ecca (HeKOTOphle rpH6hI), uapTeHOreHeTHqHhIM, arroMHKTHqHhIM cpopMaM. 3ToT KPHTeptti1: HerrpttMeHHM
.ll:JIH BhIMepIIIHX rpyuu opraHH3MOB. ,l(JI.H HCKOrraeMhIX cpopM rrpHMeH.HIOT TepMHH «cppaTpHH».
MaKpoaBOJIIOIJ;H.H, TO ecTh o6pa3oBaHJrn Ha.n;BH.ll:OBhIX TaKCOHOB, upoHcxo.n;HT Ha OCHOBe Tex JKe 3aKOHOMepHOCTeii:, qTO H MHKp03BOJIIOUl'IH.

9. Jlro6oii: peaJihHhIH TaKCOH HMeeT MOHOcpHJieTifqecKoe npoHCXOJK.n;eHHe.

10.

3BOJIIOuHH Henpe.n;cKa3yeMa, HMeeT HeHarrpaBJieJIHhIH xapaKTep K HeKOH
KOHe'-IHOH ueJIH.

158

CaJILTau;HoHHaH TeopHH 3BOJIIOD;HH

Ca.llbmau,uoH1taR meopuR
B

1899 r.

(OT JiaT.

pyccKHM 6oTaHHKOM C.

H.

saltatio -

paaoBaHHe u,11,eT aa cqeT pe,11,KHX cKaqKoB ,n;aM rrpHUieJI B HaqaJie

cKaqoK) 6hrJia BhI,11,BHHYTa

Kopm1rncKHM. Ee cyTh B TOM, qTo BH,11,006MyTa:u;ttii:. K aHaJioruqHhIM Barmi-

xx B. roJIJiaH,11,CKHM 6oTaHHK fyro ,11,e <l>pH3, OCHOBOIIO-

JIO)RHHK MyTaD;HOHHOM TeOpHH. ,ZJ;eii:cTBHTeJihHO, BH,ll,006pa30BaHHe y pacTeHHM MomeT c 6oJihUIOii: BepOHTHOCThIO rrpottcxo,11,HTh rryTeM IIOJIHIIJIOH,11,HH, 'ITO,
0,11,HaKo, He rrpoTHBopeqHT CHHTeTHqecKoii: TeopHH 3BOJIIOD;HH CT8, paccMaTpHBaroru;eii: BH,ll,006pa30BaHHe KaK CMeHy reHHbIX qacTOT II0,11, ,11,eii:CTBHeM OT6opa.

B

pH,11,e rpyrrrr opraHH3MOB ,11,0

90 %

cJiyqaeB BH,11,006paaoBaHHH HaqttHaeTCH

c xpOMOCOMHblX rrepecTpoeK.
KoH:u;err:u;ttH CT8 H caJihTaD;HOHHaH TeopHH ,11,orroJIHHIOT ,11,pyr ,11,pyra.

CoBpeMeHHaH TeopHH 3BOJIIOD;HH
TeopHH 3BOJIIOD;HH Haxo,11,HTCH cero,11,HH B cTa,11,HH ,n:aJihHeii:Uiero pa3BHTHH
H CTaHOBJieHHH.

1.

CoBpeMeHHaH TeopHH 3BOJIIOD;HH OCHOBhIBaeTcH Ha rrpHHD;Hrrax, aaJio-

meHHhIX reHirnJihHbIM

2. B Hell

l.J.

,ZJ;apBHHOM.

coxpaHHIOTCH OCHOBHhle IIOJIO)ReHHH CHHTeTHqecKOM TeopHH 3BO-

JIIOD;HH.

B

TO me BpeMH COBpeMeHHaH TeOpHH 3BOJIIOD;HH IIpHHHMaeT BO BHHMaHHe

Bee HOBbie ,n:aHHhle, OCHOBaHHble Ha COBpeMeHHbIX HCCJie,n:oBaHHHX, paccMaTpHBaeT pOJib rrepeHoca reHOB B rrpe,n:eJiaX O,ll,HOro IIOKOJieHHH (rryTeM TpaHccpopMa:u;1m, TpaHc,n;yKD;HH c yqacTtteM BttpycoB H peTpOBHpycoB), CHM6ttoreHeaa
H ,n:pyrttx HBJieHHM.

UonJ/,mUJ/, «cpeiJa o6umaHUJ/,» u «a1w1wzu<tecKue <jJaKmopbt»

159

3KOnOn1SI
TepMHH «3KOJiormI» 6bIJI BBe.n;eH B
«HayKa 0 MeCT006HTaHHH».

1866

31WJlOZU.Jl -

r. 8. feKKeJieM H orrpe.n;eJieH KaK

6HOJIOrHqecKaa HayKa, H3yqawrn;aa

opraHH3aU:HIO H cpyHKU:HOHHpOBaHHe Ha.n;opraHH3MeHHbIX CHCTeM: rrorryJiau;HH, 6HOU:eH030B (coo6rn;ecTB), 6Horeou:eH030B (3KOCHCTeM), 611occpepbr.

JlOZU.Jl

31w-

orrpe.n;eJiaeTCH TamI<e KaK HayKa 0 B3aHMOOTHOIIIeHHHX opraHH3MOB

Mem.n;y co6o:H H c OKpymamrn;eii: cpe.n;oii:.

IIpeaMem u3y'lenu.Jt a1wJtozuu -

.

npHHU:HIIhI cTpoeHHa 11 cpyHKU:HOHHpoBa-

u11a pa3JIHqHbIX Ha.n;opraHH3MeHHbIX CHCTeM. 8KOJIOrHa H3yqaeT a<HBbie CHCTeMbI ua norryJIHU:HOHHO-BH,ll;OBOM ypOBHe, 6Horeou:eHOTHqecKOM H 6HoccpepHOM.

3aaa'lU 3/COJlOZUU

(COOTBeTCTBYIOT ypOBHHM 113yqeHHH a<H3HH)

Ha rronyJIHU:HOHHO-BH.n;oBoM ypoBue:
H3yqeHHe BJIHHHHH Ha opraHH3M pa3JIHqHbIX cpaKTOpOB cpe.n;hr;
.n;HHaMHKH q11cJieHHOCTH rrorryJiau:11ii:;
H3MeHeHHH CTpyKTYPbI (COCTaBa) rronyJIHU:HH.
Ha ypoBHe 611oreou:eH030B:
H3yqeHHe B3aHMo.n;eHCTBHH Mea<.n;y IIOIIYJIHU:HHMH pa3JIHqHbIX BH,ll;OB;
IIOTOKa 3HeprHH qepe3 jKHBhie CHCTeMbI;
IIOTOKa Bern;ecTBa qepe3 jKHBbie CHCTeMbI;
3aKOHOB pa3BHTHH H CMeHbI coo6rn;ecTB.

B

3KOJIOrHH Bhr.n;eJIHIOT pa.n; pa3.n;eJioB: rrorryJIHU:HOHHaa aKoJior11a, aKoJio-

rHa coo6rn;ecTB, 3KOJIOrHa BO,ll;HbIX CHCTeM (r11.n;po6110Jior11a) H .n;p.
8KOJIOrHa .n;eJIHTCH TaKme Ha ayT3KOJIOrHIO, .n;eM3KOJIOrHIO H CHH3KOJIOrHIO.

Ayma/COJlOZU.Jt -

H3yqaeT oT.n;eJihHhre opraHH3MhI H 11x npHcrroco6JieHHa

K oKpyarnrorn;e:H cpe.n;e.

l(eM3/COJlOZU.Jl Cuna1CoJLozu.Jt -

3KOJIOrHH nonyJIHU:HH.
aKoJior11a coo6rn;ecTB; oHa HCCJie.n;yeT rpyrrrrhr opraHH3MOB

B HX B3aHMo.n;eii:CTBHH co cpe.n;oii: H Mea<.n;y co6oii:.

IlOHHTHH «cpe;::i;a o6HTaHHH» H «3KOJIOrH'leCKHe cl>aKTOpbI»

Cpeaa o6umanu.Jt -

aTo KOMIIJieKc 0Kpymarorn;11x ycJioBHH, B03.n;eii:cTByro-

UJ;Hx Ha opraHH3M (oco6H o.n;Horo BH.n;a, nonyJIHU:HH .n;pyr11x BH.n;os, JII06hie HemHBhie 06'beKTbI, cp11311qecKHe H XHMHqecKHe rrpon;ecCbI H HBJieHHH).

31COJLOZU'leC1CUe <jJa/CmOpbl -

3TO KOMilOHeHThI cpe.n;hI 0611TaHHH, BJIHHIO-

IIJ;He Ha opraHH3M.
3KOJIOrHqecKHe cpaKTOpbI rro.n;pa3.n;eJIHIOTCH Ha:

I. A6HoT11qecK11e.
II. BHoTHqecK11e.
III. AHTponoreHHbie.
Bee c}>aKTOphr MOa<HO pa3.n;eJIHTh Ha

PecypCbl -

ycJLOBU.Jl

11 pecypcbt.

3TO KOMilOHeHTbI cpe.n;hI o6HTaHHH, KOTOpbre MoryT IIOTpe6JIHTb-

Cst OpraHH3MOM, rrpH 3TOM CHHmaeTca HX ,ll;OCTYIIHOCTb ,ll;JIH .n;pyrHX (nmn;a).

06uta Jl

160
YcJl06UJl -

6uoJWZUJl. :-iiwJWZUJl

3TO cpaKTOpbI, B03i'J;CHCTBHe KOTOpb!X He 38BHCVIT OT VIX IIOTpe6-

JieHHJI .n;pyrHMU opraHU3M8MVI (TeMnepaTypa).

I.

A6HOTHqec1me Q>aKTOPLI -

3TO Bee cpaKTOphI neJKHBOH npnpo.n;hr. Ba%-

HeMIUHMH JIBJIJIIOTCJI CBeT, TeMnepaTypa, BJiaJKHOCTh, aTMOCqlepa, COCTaB TIO'lBhI H .n;p.

Co.JUte'lflbtii cBem -

HCTOTfHUK 3ueprn:n .n;JIJI JKIIBLIX opraHH3MOB. BnoJiorn-

qecKoe .n;eMCTBUe 3aBHCHT OT i'J;JIHHhI BOJIHbI, HHTeHCHBHOCTH, neptt:Oi'J;Hl'.JHOCTH,
(puc.

59). Y.1bmpa<jJuo.llemo6bte .lly'lu

xapaKTeptt3YIOTCJI caMoli BLICOKOH 3Hep-

ruei1: KBaHTOB. CTnMyJiupyIOT y JKHBOTHLIX 6uocmITe3 BnTaMHHa

D.

Bocnp11-

nuMaIOTCJI 3pentteM HeKOTopLIX HaceKOMLIX. MoryT cTaTL up11qrrnoii MyTau;n:OHHLIX U3MeHeHnii.

BuiJuMbtu ceem o6ecneTJHBaeT cpOTOCHHTe3. Heo6xo.n;ttM

,D;JIJf 3peHHJI H optteHTaIJ;UH B npocTpaHCTBe. 06ecnel'.JUBaeT QJOTOTiepHO,ll;H3M.

<PomonepuoiJuaM -

peaKIJ;UJf JKHBbIX opraHH3MOB Ha H3MeHeHne ,ll;JIHHhI CBeTo-

BOro ,LJ;HJI. PeryJinpyeT cyTol'.JHhie u ce30HHhie pttTMLI JKHBLIX opraHH3MOB.

JipuMepbl. y

JKHBOTHhIX:

MHrpau;uJI TITHIJ;,

CMena IUepCTHOro TIOKPOBa

y MJieKOTIHTaIOI.QHX, Bna.n;eHHe B CTIH'!Ky, 6pa'IHOe TIOBe.n;eHne.

y

paCTeHn:i1:

IJ;BeTeHne KOpOTI:W,LJ;HeBHhIX H ,LJ;JIHHHO,LJ;HeBHhJX pacTeHHH JieTOM H oceHbIO, OTKPLIBaHne H 3aKpbIBaHHe IJ;BeTKOB H .n;p.

HH<jJpa1CpaCHbte .llY'lU -

HCTO'IHHK Ten.JIOBOM 3Heprtttt. TepMoperyJIJIIJ;HH

XOJIO,LJ;HOKPOBHhIX JKHBOTHbJX ( 6ecn03BOHO"l!HhIX, HH3IUHX TI03BOHO"l!HbIX).

TeMnepamypa BJIH.HeT Ha CKopocTL 6noxnMn"l!ecKnx npou;eccoB B opraHH3Me. ,Zl;ttana30H TeMnepaTyp ,LJ;JIH lliH3He.n;eHTeJILHOCTH 60.JibllIHHCTBa opraHH3MOB OT

0-4

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.n;o 50 "C.

OT.n;eJibHbie BHi'J;hI 6aKTeptti1: H CHHe3eJieHhIX Bo.n;opoc-

JieM MoryT o6uTaTb B rop.H'IHX HCTOl'.JHHKaX npH TeMrrepaType
JIJIPHLie BO,LJ;bI c TeMnepaTypoi1: OT

0 ·. c .n;o -2

80-90 'C.

Ilo-

'C Hace.'IeHbI 6ecno3BOHO"l!HbIMH,

pbI6aMH, BO,LJ;OpOCJIJIMH.
Ilo cnoco6HOCTH K no.n;.n;epmaHHIO TeMnepaTyphI Te.Jia JKHBOTHhie .n;eJIJITCJI
Ha

XO.llOiJflOICp06HblX (noMKH.JIOTepMHbIX) H men.llOICp06HblX (rOMOMOTepMHhIX).

PeHTreHOBCKltle
PaJJ,111oeo11Hbl
An1t1Ha
BOJlHbl

< 0, 1 HM

fblCSl'H1 MeTpOB

8HJJ,HMbliii CBeT
Q>HoneTOBbliii CHHHiii rony6oiii 3eneHbliii )l(eJlTbJH OpaH)f(eBbliii KP8CHblM

750

380 HM
P1tc. 59. ,ll.nL-1Hbl

Yet> -

BOJlH COJlHe'-tHOro L-13Jly'leHL-1S1

y11bTpa¢L-1011eT;

lllK -

()KY<!> -

)1(8CTKL-1H YllbTpacjmo11eT;

L-1Hcj>paKpacHoe L-1311y'-teHL-1e)

HM

flo1t1tmUJl «CpeiJa 06uma1tUJl» U «3KOJWZU'l-eCKUC <jJaKmOpbt»

161

TenJL01cpoB11bte (roMottoTepMttbre) -- 3TO IITMILLI n MJieKorrnTa10w;11e. TeMrrepaTypa TeJia y HHX rroeTOJIHHa, He 3aB:ne:nT 0-T OKpymaIOw;eH: epe,ZJ;bl. l1MeIOT
rrp:nerroeo6JieH:na B eTpOeH:n:n (nepbCBOH IIOKpOB, mepeTL, IIO.ll;KOJKHLIH :arnpoBOH eJIOH H ,n;p.) H npnenoeo6.JieHH.H B o6MeHe Bew;eeTB (IIHTeHeHBHOeTb OKHeJieHH.H, «.ll;BOHHOe» .ll;hIXaHne y IITHI!, 6ypbIM m:np y MJICKOIIHTaIOII~IIX H ,n;p.).
XoJLoihtoKpOBHbte (noi1KmroTepMHhre) - 3TO 60JihIIIHHeTBO oprattn3MOB:
Bee THIIhI 6eerr03BOHQqHhIX H qaeTb II03BOHQqHhIX (pbI6hI, aMcp:n6nn, npeeMLIKaIOlll;Heea). TeMnepaTypa :nx TCJia 3aBnenT OT oKpyJirnrow;eil epe,n;hr, otta MomeT IIOBhIIIIaThC.H BO BpeM.H BhICOKOM MhIIIIeqHQM aKTHBHOeT:n.
IIpucrroeo6Jiett:na K nepemirnaHnIO ne6Jiaronp:naTHhIX TeMnepaTyp:
a) 3HMttaa enaqKa (3:nMtt:nli coH) - nepettecett:ne 3HMHero rrepno,n;a;
6) atta6:no3 - BpeMe1rnoe eoeTO.HHHe, rrp:n KOTOpOM Bee )KH3HeHHhie rrporrecehI 3aMe,n;JieHhI ,n;o MHHHMyMa II OTeyTeTBYIOT Bee BH.ll;HMbie np:n3HaK:n
)f{H3HH (Ha6JIIO,ZJ;aeTe.H y XOJIO.ll;HOKpOBHbIX 3HMOH :n B mapKoe BpeM.H ro,n;a);
B) MOp030eTOHKOeTb - erroeo6HOCTh rrepeHOeHTb HH3K:ne OTpnrraTeJibHbie
TeMrrepaTyphr;
r) 3HMHHM IIOKOM - eBoii:eTBO MHOI'OJieTHHX paeTeHntt: OTM:npattne Ha,n;3eMHhIX qaeTett paeTett:na, ona,n;attne JineTbeB;
,n;) JieTHHM IlOKOH - xapaKTepeH .ll;Jl.H paHHerrBeTyw;:nx pacTeH:ntt nyeTbIHh
n rroJiynyeTbIHh (TIOJihrraH, KpoKyc :n ,n;p.).
BJtaJ1C1wcmb - oettOBHOH KOMIIOHeHT B eoeTaBe :>KHBhIX opraHH3MOB. He,n;oeTaTOK BJiarH rrp:nBO.ZJ;HT K B03HHKHOBeHHIO pa3JIHqHhIX rrpHenoeo6JieHHH
y pacTeH:nll :n Jf<HBOTHbIX.
Ilo np:nerroeo6JieHHOeTH K MeeToo6:nTatt:nIO B pa3JIHqHbIX yeJioBnax yBJiamHett:na paeTett:na ,n;eJiaTca Ha:
r:nrpocp:nThI - paeTeHna BJiamHhIX MeeT o6:nTali:na (KaJiymtt:nrra 6oJIOTHaa, rrarr:npye, pnc, rranopoTHHKH, opx11,n;Hhre). l136hIToqttaa BJiara BbI,n;eJiaeTca e IIOMOlll;bIO BO.ll;5IHbIX yeTblIII - iuiJamoiJ;
Me30<}>HThl - pacTeH:n.H, np11enoeo6JieHHbJe K Jf<H3HH B yeJIOBH.HX yMepeHHOro yBJiamttett11a. B oeHOBHOM 3TO paeTeHH.H Jiyra n Jieea. K Me30<}>11TaM OTHOe.HTC.H 3<}>eMepbl H 3<}>eMepOH.ll;bl, T. K., 6y,n;yq11 o6nTaTemIMH
nyeTbIHHhJX o6JiaeTeM, OHH 3aKaHqHBaIOT BereTarrHOHHhIM rrepno,n; BeeHOH. 3aeyIIIJIHBOe BpeM.H OHH rrepem:nBaIOT B Bn,n;e eeM.HH (3cpeMepbI),
KOpHeBiflll; H JIYKOBHI! (3<}>eMepOH.ZJ;bI);
Keepo<}>HThI - paeTeHH.H eyxnx MeeT 0611Tatt:na, erroeo6HhJe rrepettoeHTh
He,n;oeTaToK BJiaru. Ott11 HMeIOT rrpuerroeo6JieHH.H, yMeHhIIIarow;ne rroTePIO BO.ZJ;bl HJIH II03BOJI.HIOlll;He ,n;o6bIBaTb BO.ZJ;Y. y Bep6JIIOJI<Leli KOJIJOqKH
Kopetth ,n;oeTuraeT rJiy6HHhI 15 M. KopoBaK HMeeT enJihHOe orryIIIettue
JIHCTheB. ToJieTa.H KYTHKyJia, 3arraeattue BO.ZJ;hl - y KaKTyea.
Ilpncnoeo6JieHH.H y )KHBOTHhIX:
a) IIJIOTHhJe IIOKpOBhJ (XHTHH, qeIIIy.H): Bbl,ZJ;eJI.HIOTea HepaeTBOpHMhJe npo.ll;YKTbJ (MO'IeBa.H KHCJIOTa, ryaHHH), rrpouexo,n;HT 3KOHOMHH BO,D;hl;
6) rop6 Bep6JIIO,D;a, IIO.ZJ;KO)f{HhIM mup rpb13YHOB o6eerrequBaIOT o6pa30BaHHe
BO.ZJ;bI 3a eqeT ru,n;poJIH3a mupa.

162

06Uj-llll 6uo/WZUJl. :J1W/WZUJl

KucJ1opod. ,Ll;Jrn:

6oJihIIIMHCTBa JKMBhlX opraHM3MOB KttcJiopo.n; JKM3Hemrn He-

o6xo,n;ttM. KttcJiopo.n; HBJIHeTcH KOHe'-IHhlM ai-rn;enTopoM ;rneKTPOHOB B ,D;h1xaTeJihHOH u;entt. B 6eCKMCJIOpo,n;HOH cpe.n;e MOryT pa3BMBaTbCH TOJibKO aHaapo6Hble 6aKTeptttt M aH,n;onapa3ttThI. KttcJiopo.n; B aTMoccpepe o6pa3oBaJICH nyTeM
OKCttreHHOrO cpOTOCMHTe3a. IlorJIOII.J;eHMe KllCJIOpo.n;a ll3 BHeurnei1: cpe,D;bl npoMCXO,[l;llT qepe3 BCID TIOBepXHOCTb TeJia (npOCTetturne, qepBM), c TIOMOII.J;bID opraHOB ,D;hIXaHMH (Tpaxett, ma6pb1, JierKtte). KttcJiopo.n; XMMtt'-leCKM CBH3hrnaeTCH
tt nepeHOCMTCH no opraHM3MY cneu;ttaJihHhIMM nttrMeHTaMn KPOBM -

reMorJio-

6ttHOM (no3BOHO'-IHhie), reMOD;MaHMHOM (MOJIJIIDCKM, paKoo6pa3Hhie).
Ilptt He,D;OCTaTKe KllCJiopo.n;a B cpe.n;e y opraHM3MOB Bhlpa6oTaJIMCb COOTBeTCTBYIDII.J;Me npttcnoco6JieHMH:

TIOBbIIIIeHHaH KllCJIOpO,n;HaH eMKOCTb KPOBM,

60JihllIOH o6'beM JierKMX, 6oJiee '-laCTble ll I'Jiy60KMe ,[l;blXaTeJibHhle ,[l;BMJKeHMH.

II. Bn0Tn11ec1me <l>aKTOPLI -

3TO <t:>aKTOphI, B03,n;ei1:cTBYIDII.J;tte Ha opraHtt3M

co cTopoHhI .n;pyrnx JKMBhlX cyw;ecTB. K 6ttOTHtieCKMM <t:>aKTopaM cpe,n;hI OTHOCHTCH KOHKypeHD;llH, XllII.J;HM'-leCTBO, CMM6M03 (pa3JIM'-1Hhle Bll,[l;bl).

Ko1ucypenu,u.H -

aTo B3attMOOTHoIIIeHHH Mem.n;y oco6HMM MJIH nonyJIHil;MH-

MH O,ll;HOrO BM,D;a (BHYTPMBll,[l;OBaH) llJlll pa3HbIX Bll,[l;OB (MeJKBll,ll;OBaH), 6opIDII.J;llXCH 3a o,n;Htt n Te me pecypchI (nnTaHtte, cBeT tt .n;p.). KoHKypeHu;ttH oKa3hIBaeT OTpuu;aTeJihHOe BJillHHMe Ha ee yqacTHMKOB.

3mwn ( npunu,un) Kowcypenmnozo ucKJllO'lettu.H ( npunu,un I'ay3e)
MYJIHpOBaII POCCMHCKMM yqeHbIM
ttH<l>y3opnHx-Ty<l>eJihKax

r. <t>.

fay3e B

(Paramecium).

1934

6hIJI c<t:>op-

r. ll ,[l;OKa3aH B OilbITax Ha

Y'-leHhIH Ha6JIID,ll;aJI H3MeHeHtte '-IMC-

JieHHOCTM B o,n;HOBH,n;OBhIX M ,n;ByXBM,ll;OBhIX KYJihTypax. Ilttrn;eii CJIYJKMJIM 6aKTepttn M .n;pommtt, .n;o6aBJIHeMhie nopu;ttHMM. EcJitt ,n;Ba Btt,n;a c o,n;ttHaKOBhIMM aKoJiortt'-lecKttMH TIOTpe6HOCTHMll OKa3hIBaIDTCH B O,[l;HOM coo6w;ecTBe, TO 6oJiee CllJihHhIH KOHKypeHT BhlTeCHllT 6oJiee CJia6oro. BoJiee CllJibHhIH KOHKypeHT o6Jia,n;aeT

CoBMecmnoe o6umanue 6JlU3KopodcmBettttblX 6Ud06 603MOJICHO, eCJIU OllU 3UllUMalOm pa31lble 3KOJIOZU'leCKUe llUUlU.
3KOJWZU'leCKll.H llUUW - 3TO KOMTIJieKC <t:>aKTOPOB, KOTOpbie Tpe6yIDTCH ,[l;JIH
6oJihIIIei1: cKopoCThID pocTa n pa3MHomeHMH.

cyw;ecTBOBaHMH Btt,n;a, B TOM '-lllCJie CBH3ll c .n;pyrttMll Btt,n;aMM B coo6w;ecTBe.

IIpuMepbl.

1. B

OIThITax

r. <t>.

fay3e B Te'-leHMe ,[l;JIMTeJibHOro BpeMeHM MOr-

Jlll cyw;ecTBOBaTb TOJibKO Te Bll,[l;hl, KOTOpbie MMeJIH pa3Hhie «TIMII.J;eBble» Hlllllll
ll ITIITaJIMCh JIH6o 6aKTepMHMll, JIM6o .n;pommaMM.

2.

Tpo<l>tt'-leCKtte JIMHMM y ITYCThIHHhIX BM,D;OB MypaBheB. B o,n;HOM MecT00611-

TaHIIII BbI,D;eJIHIDT TPM rpyITITbI MypaBheB ITO TMITY TIHTaHHH: KapITo<t:>arM ITHTaIDTCH ceMeHaMM, 3oo<t:>arn -

JKllBbIMll HaceKOMhlMll, HeKpoq:iartt -

OCTaHKaMH

JKllBOTHblX.
BL1,n;eJIHIDT TpII OCHOBHhIX CITOC06a pa3,n;eJieHHH 3KOJIOrtt'-leCKMX HMIII: B ITpOCTpaHCTBe, BO BpeMeHM, ITO pecypcaM. 3TII CITOC06bl MoryT CO'-leTaTbCH. B pe3YJihTaTe KOHKypeHil;llll MOJKeT 6h1Tb BhITecHeHne 0,ll;HOI'O Btt,n;a .n;pyrttM, pa3HaH 3KOJIOI'll'-leCKaH CITeu;naJin3all;MH Bll,[l;OB.
3Ha'-leHMe KOHKypeHil;Mll: KOHKypeHTHhie B3aMMOOTHOIIIeHMH <t:>opMttpyIDT
Bll,[l;OBOH COCTaB coo6w;ecTBa, pacITpe,n;eJieHtte Bll,[l;OB B rrpocTpaHCTBe, '-lllCJieHHOCTh Bll,[l;OB.

163

flOILJtfflUJl «CpeiJa o6umaHUJ£» U «31WftOZU'l€CKUe t}JaKmOpbL»

Xuw,ttu'lecmBo -

3TO cpopMa B3aMMOOTHOIIIeHMM, KOr)J;a opraHM3M O,ll;Horo

BM)J;a MCTIOJih3yeT opraHM3M ,ll;pyroro BM,ll;a B KatJ.eCTBe MCTOl'.J.HMKa TIMIIJ,M (JIOBMT, y61rnaeT, TIOe,ll;aeT). B3aMMOOTHOIIIeHMH XHI.QHMK-;1rnpTBa IIInpoKo pacTipocTpaHeHhI B TipMpo)J;e. XMI.QHMKH BCTpetJ.alOTCH B mnBOTHOM u,apcTBe (TITMn;hI, pbI6hI, peTITMJIMM, MJieKOTIMTalOI.QMe), cpe,ll;M n;apCTBa rpM60B H pacTeHHll
(pocHHKa,

:ampHHKa,

xmUHMtJ.eCTBa) -

BeHepnHa MyxoJIOBKa n ,ll;p.).

KaHHM6aJIM3M (cpopMa

TIOe)J;aHMe oco6eii: CBoero BM)J;a. KaHHM6aJIM3M BCTpetJ.aeTcH

y xMI.QHbIX pbI6, HaceKOMbIX, TiayKoo6pa3HhIX.
XMI.QHMtJ.eCTBO cJiyiKMT MexaHM3MOM peryJIHIJ,MM tJ.MCJieHHOCTM TIOTIYJIHU.Mii::
B TIPHPO)J;e XMI.QHMKM YHMl'.J.TOiKaIOT HaM6oJiee OCJia6JieHHbIX, 60JibHhIX oco6eii:.

Cu.11t6uo3 (oT rpeq, simbiosis -

coBMeCTHaH iKM3Hh) -

3TO pa3JIMl'.J.Hhie cpop-

MhI COBMeCTHOro cyI.QeCTBOBaHMH ( COiKMTeJibCTBa) opraHM3MOB.
CMM6M03 MOiKeT cyI.QeCTBOBaTh Ha OCHOBe pa3JIMl'.J.HhIX TMTIOB B3aMMOOTHOIIIeHMH:
TpOc}:>Ml'.J.eCKMX (TIMTaHMe O,ll;HOro 3a Cl'.J.eT cpOTOCMHTe3a ,ll;pyroro, MCTIOJib30BaHMe OCTaTKOB TIMI.QM ,ll;pyrMM TiapTHepoM);
TipOCTpaHCTBeHHbIX (COBMeCTHOe MCTIOJib30BaHMe Hop, MypaBeHHMKOB,
TioceJieHMe Ha TIOBepxHOCTM TeJia).

<jJaKyJtbmamuBHbt.M ( tteo6.H3ameJtbHbt.M) M o6JtuzamHbt.M
( o6.H3ameJtbHbt.M ). HaTipMMep, o6JIMraTHhIH CMM6Mo3: JIMIIIaHHMK (rpM6 M BoCMM6M03 6blBaeT

,ll;OpocJih), MMKOPM3a (rpM6 M pacTeHMe); cpaKyJihTaTMBHblH CMM6M03 -

COTpa-

Tie3HMl'.J.eCTBO (rMeHbl M JibBbl).
Bhl,ll;eJIHIOT HeCKOJibKO TMTIOB CMM6M03a: a) KOMMencaJIM3M: HaxJie6HMl'.J.eCTBO, KBapTMpaHCTBO; 6) Tiapa3MTM3M M TIOJiyTiapa3MTM3M; B) MyTyaJIM3M.

KoM.MeHCaJlU3.M -

3TO TaKOH TMTI CMM6MOTMl'.J.eCKMX OTHOIIIeHMH, KOr,ll;a )J;e-

HTeJibHOCTb O,ll;HOrO BM,ll;a opraHM3MOB ,ll;OCTaBJIHeT TIMI.QY MJIM y6emMI.Qe ,ll;pyroMy BM,ll;y (KOMMeHCaJiy); TIPM 3TOM KOMMeHCaJibl O,ll;HOCTOpOHHe MCTIOJih3YIOT
,ZJ;pyroii: BM,ZJ;, He TIPMHOCH eMy BhirO,ll;hl MJIM 3aMeTHoro Bpe,ZJ;a. KoMMeHcaJIM3M

11axJ1e6HU'lecmeo. HaTipMMep, B3aMMOOTHOIIIeHMe rMeH M JihBOB, pb16-TipMJIMTiaJI M aKyJI.

MOiKeT 6bITb OCHOBaH Ha TIOTpe6JieHMM OCTaTKOB TIMI.QM X03HeB -

KOMMeHCaJIM3M, TipM KOTOpOM O,ZJ;MH BM,ll; MCTIOJlh3yeT ,ll;pyroil: KaK cy6cTpaT
,Zl;JIH 0611TaHMH MJIM TpaHCTIOpTa, Ha3hIBaeTCH

IC6apmupaHCm60.M (CMHOHKMeH).

Bhlro,ZJ;y TIOJiyqaeT O,ll;MH BM,ZJ;, ,D;pyroMy 3TO 6e3pa3JIM1'.J.HO. HaTipMMep, MaJihKM
HeKOTOPhIX pb16 TIPHl'.J.YTCH B TIOJIOCTM opraHOB ,Zl;hlXaHHH rOJIOTypMH:; B MypaBeHHHKax, TepMHTHMKax, nopax rpb13YHOB TIOCeJIHIOTCH MHOrMe BM,ll;bl qJieHMCTOHOrMx, rp1160B, MCTI0Jlh3YH KaK y6emMI.Qe MJIM MeCT006MTaHMe; B pacTMTeJihHOM

~apCTBe pacTeHMH-3TIHc}:>MTbl TIOCeJIHIOTCH Ha ,ll;epeBhHX, KOTOpbie CJiymaT MecTOM IlPHKperIJieHmI, TIMTaHMH, cpoTOCHHTeaa (HaTipMMep, JIMIIIaHHMKM).

llapa3umu3M -

cpopMa B3aMMOOTHOIIIeHMH MeiK,ll;Y opraHM3MaMM pa3HblX

BR,ZJ;OB, np11 KOTOpOM O,ll;MH opraHH3M (rrapa3HT) MCIIOJlh3yeT ,ZJ;pyroil: (X03HMHa)
B Ka'IeCTBe IIOCTOHHHOH MJIH BpeMeHHOH cpe,ll;hl o6MTaHMH, MCTOl'.J.HMKa IIMI.QM.
-

IlapaaHThl 6hrnaroT

nocmo.HHHbte M epe.MeHHbte. IlocTOHHHhie rrapaaMThI

B:e MoryT CyI.QeCTBOBaTh BO BHeIIIHeH cpe)J;e BHe X03HMHa (acKapM,ll;a, 6hll'.J.MH

~eneab,

BIIIH). BpeMeHHhie Tiapa3MThI Haxo)J;HTCH Ha xoaHMHe BpeMeHHO,

164

06UfUR 6UO/WZUR. :JKO/WZUR

,n;Jrn rrIITa111u1 (KOMaphI, KJieu~tt, KJIOIIhI tt .n;p.). ITapa3ttTOB no.n;pa3.n,eJ1.HIDT Ha

aKmo-

II

a1-uJonapa3umoa. 3Kmonapa;wmbt
3HiJonapa3Umbt o6IITaIDT B

mIIBYT Ha TeJie X03.HIIHa (BIIIII,

KJieruII, 6JIOXII).

ITOJIOCT.HX IIJIH BHYTPII TeJ1a X0-

3.HIIHa (MaJI.HpIIMHblM ITJia3MO.D;IIH, 3XIIHOKOKK, acKapII.n;a, OCTpIIua, TPIIXIIHeJIJia II .n;p.).

B TeJie

rrapa3IITOB MoryT o6IITaTb co6cTBeHHhre rrapa3IIThI -

3TO

ceepxnapa-

.mmu3.JIL
,IlJI.H napa3HTOB xapaKTepHa pe.n;yKUII.H HeKOTOpblX CIICTeM opraHOB, HarrpIIMep IIIIIQeBapIITeJihHOH y JJeHTOl.J:HbIX qepsett:, MHOI'IIX opraHOB qyscTB y acKaPII.ZJ: II .n;p. B To me speM.H pa3BIIThI npIIcnoco6JieHIIH .D;JIH y,11,epmaHII.H B TeJie xo3HIIHa: KOJIIDIQe-cocyIQIIM poToso:ii armapaT, npIIcocKII, KpIDl.J:hH II ,11,p. ITapa3IITII3M IllIIPOKO pacnpocTpaHeH cpe,11,II )KIIBbIX cyIQeCTB.

FlpuMepbt. 1.

BaKTepIIII -

napa3IIThI, rpII6h1.

2.

:IB:IIBOTHhre -

secb TIIn

CnopoBIIKII (no,11,r:rapcTBO IlpocTettlllIIe); KJiacc CocaJibIQIIKII, JleHTOl.J:Hhie II3
TIIna IlJIOCKIIX qepsett:; TIIn qJieHIICTOHOI'IIe (Bnrn, 6JIOXII II ,11,p. ).
Bbie pacTeHIIH napa3IITbl II noJiynapa3IITbl -

3.

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xapaKTepII3YIDTCH pe,11,yr:rIIpOBaH-

HbIMII KOPHHMII, OTCYTCTBIIeM cnoco6HOCTII K cpOTOCIIHTeay (neTpOB KpecT, 3apa3IIXa).
Te3y'

PacTeHIIH-no.nynapa3umbt

HO

paCTBOpbl

cnoco6nhr K caMOCTOHTeJibHOMY cpoTOCIIH-

MIIHepaJibHbIX

C0"'1eH

6epyT

II3

KCIIJI€MHOI'O

CO Ka

pacTeHIIH-X03.HIIHa (norpeMOK, OMeJia 6eJia.H, Mapb.HHHIIK .n;y6paBHbIM).

Mymya.nu3.M

(oT JiaT.

mutuus -

saaIIMHhitt) -

cpopMhI o6JIIIraTHoro, B3aII-

MOBh1ro,11,Horo CO;KIITeJibCTBa ,11,BYX BII.D;OB.

IIpu.Mepbt. 1.

KJiy6eHbKII Ha KOPHHX BhICillIIX pacTeHIIH, o6pa3oBaHHhie

B pe3yJihTaTe B3aIIMo.n;ettcTBIIH c 6aKTepIIHMII po.n;a PIIao6IIyM.
(rpII6oKopeHb).

3.

JlIIIllattHIIKII.

4.

2.

MIIKOpII3a

CIIM6IIOTII'-leCKIIe 6aKTepIIII, IIH<Py3op1m,

0,11,HOKJieTOl.J:Hbie mryTIIKOBble, KOTOpbie nOMOI'aIDT )KIIBOTHOMY-X03.HIIHY nepepa6aTbIBaTb paCTHT€JibHYID nIIIQy, Bbipa6aTbIBa.H cpepMeHT ,11,JIH pacIQenJieHHH
KJieTl.J:aTKH.

5.

B

TeJiax HeKOTOphIX «npo3pal.J:HhIX» mttBOTHhIX noceJIHIDTCH

MHKpOB0,11,0pOCJIII; HanpIIMep, nOJIIInhI o6pa3yIDT CHM6II03 c B0,11,0pOCJIHMH 300KCaHT€JIJI8MH.

III. AHTponoreHHhie <1>aKTOph1

-

3To pa3JIII'IHhie ipopMhI ,11,eaTeJihHOCTH qe-

JIOBeKa, OKa3bIBaIDIQHe B03,D,eMCTBHe Ha ;KHBbJe opraHII3Mbl H KOMnOHeHTbl OKpymaIOIQeii: cpe,11,h1.
,IleHTeJibHOCTb qeJIOBeKa npHB0,11,HT B OCHOBHOM K OTpIIH;aTeJihHbIM nocJie.n;CTBHHM ,11,JIH )KHBOM np:ttp0,11,bl. 3To pa3pyllleH:tte ecTeCTBeHHblX 6~0TOnOB (Bblpy6Ka JI€COB, ocyllleH:tte 6oJIOT, CTpO:ttTeJihCTBO nJIOTHH H ,11,p.), COKpaIQeH:tte
apeaJIOB BH,11,0B H l.J:HCJieHHOCTH BH,11,0B pa,11,a )KHBOTHbIX, 3arp.H3H€HHe OKpymaIDIQe:H cpe,11,hr (aTMoc<Pephr, r:tt.n;poc<Pephl, JI:ttToc<Pephl).

3azpR3HJllOW,Ue eew,ecmea -

3TO BeIQeCTBa, CKanJIHBaIDIQHeCH B OKpymaID-

IQeH cpe,11,e B KOHUeHTpar:r:ttHX, HexapaKTepHbIX ,11,JIH ,ll,8HHOM MeCTHOCTH, H np:ttB0,11,HIQHe K HapyllleHH.HM B )KH3He,D,e.HTeJibHOCTH )KHBbIX opraHH3MOB.
Bhl,11,eJIHIDT pa3JIHl.J:Hhie rpynnhI 3arpH3HHIDIQHX se1r:recTs: no x:ttM:tt'-leCKOM
rrp:ttpo,11,e (opraH:ttl.J:ecKtte seIQeCTBa, THmeJibie MeTaJIJihI :tt .n;p.); no cTeneH:tt TOKCB'-IHOCT:tt; no MyTareHHOMY 3cpcpeKTy.

,'Ja«o1wMepnocmb iJeiicm6uJL .1Ko1wzu'leC«ux </Ja«mopo6 na opzanu.J;n

165

8TII semecTB8 MOI'YT Bbl3bIB8Tb rM6eJib opraHH3MOB H IIplIBO,ll,HTb K B03HHKHOBeHHI0 MYT8IJ;MM. Pe3yJihTaTOM MOJKeT CT8Tb aeo6p8THMoe HapymeHMe ;monorwiecKoro paBHOBeCH.H, rM6eJib 3KOCHCTeM HJIH 3IW.'10l'J1qecK8.H K8T8CTpocpa.
3aKOHOMepHOCTh ;::i:eii:CTBH.H :JKOJIOrttqecKHX <f>aKTOpOB Ha opraHH3M
P83JII'l'JHhie cp8KTOphr OK83bIB8IOT p83HOe BJIII.HHIIe Ha opr8HII3M: o.n:ttn
TOPhI -

CHJibHOe BJIIIRHIIe, .n;pyrMe -

ql

8 K6oJiee CJJ86oe; HO MOJKHO Bbl.HBI:ITh 06-

60).

rn;yIO cxeMy .n:eiicTBl:IR 3KOJIOI'HqecKI:IX cp8KTOPOB (pttc.

!Juana.mu ycmoil'lUBOcmu, HJJH mo;iepaHm1-wcmu -

3TO rrpe.n:e"'lhHbie (MM-

HMM8JJhHhie 11 MaKCHMaJibHbie) 3HaqeHH.H cpaKTOp8, Mem.n:y KOTOPbIMH opraHH3M MOJI<eT BhIJKHTh.

OnmuMy.M (Haxo.n;I:ITCR B cpe.n;Heii: qacTH .n;ttarra3on3) -

3TO TaKHe yCJIOBI:IR, B KOTOpbIX oco6H ,n;aHHOrO BH,[\8 HMeIOT H811JIYLIII111e IIOK838TeJI11 JKH3He.n;eRTe.'IhHOCTl:I.
qame roBOPRT 0 30He OIITIIMyMa H npn ne,n,ocTaTKe HJII:I 1136hITKe 3KOJIOI'HtJeCKOI'O cpaKTOpa Bhl.HBJJ.HIOT 30HY neCCI:IMyMa HJIH CTpecca. 3a npe.n;eJiaMII 30Hhl BblJKHBaHHR H8XO,ll,.HTCR

JlemaJlbHbte (cMepTe.TlhHhie) 3HaqeHII.H cpaKTOpa.

T8Ka.H 3aKoHoMepHoCTh peaKu;HM opraHH3MOB Ha B03.n;eii:cTBHe 3KOJIOrI:ItJeCKHX <l>aKTOpOB -

BaJKHblM 3KOJIOrtttJeCKIIM rrpIIHIJ;HII: ,ll,.'IR Kam.n:oro BH,ll,8 mn-

BOTHblX HJIII pacTeHHM cymecTByeT OIITIIMyM, 30H8 yrHeTeHMR H npe,n,eJibl BblHOCJIHBOCTH no OTHOilleHHIO K KaJK,ZJ;OMY cpaKTOPY cpe,D,hl.
B

aaBHCMMOCTM OT ruttpIIHhI .n;ttarraaotta ycToii:tJnBocTH K onpe,n;eJieHHhIM

<f>8KTopaM cpe,n;b1 opraHM3MhI ,n;e.TI.HTCR Ha

31COJlOZU'leCKll.fl nJLacmU'lHOCmb -

3Bpu6uoHmHbte H cmeHo6uoHmHbte.

3TO cnoco6HOCTh opraHH3MOB a.n.anTnpo-

BaThC.H K H3MeHeHH.HM cpaKTOpOB cpe,D,hl.
8BpII6HOHTHhie BH,ll,hl 3KOJJOI'HtJeCKH 60"1ee IIJI8CTlltJHhI, cnoco6Hhl BblfKHBaTb H OCTaB"'l.HTh IIOTOMCTBO nptt H3MeHeHIIH YCJ'lOBHM, HMeIOT IIIHPOKHe apeaJlhl pacrrpocTpaHem,rn.

J!u.11tumupy1-0w,uu rpaicmop -

3To

$aRTop, yponeHh KOToporo nptt6.TittJKa-

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rrpe,n,eJI8M TO.rrepaHTHOCTH H.TIH

ae
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npa6UJlO .MU HU.My.Ma JI u6u-

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paHTHOCTH, r,ll,e B03M0)KH0 pa3MHO)K8HH8)

166

06utaH 6UOJWZUH. :JKOIWZUH

BbillJe OTJIOMaHHOH ,ZJ;OCKH Bo,n;y He HaJibellih, ,ZJ;JIHHa
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me

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TpaHCIIHpau;MH (ttcrrapeHH.ff BO,ZJ;hl) BbIIIIe rrpn BbICOKOH TeMrrepaType M rrptt BeTpeHo:ft rroro,n;e. KoMrrJieKc
<PaKTOpOB, KOTOpbie Tpe6yIOTC.ff ,ZJ;JI.ff cyru;ecTBOBaHHJI
BH,n;a, BKJIIOqaJI ero CB.ff3M c ,n;pyrHMll BM,n;aMH B coo6-

P111c. 61.

«504Ka n1116111xa».

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ru;ecTBe, Ha3hIBaeTCJI

KOni1148CTBa BO-

Abl, KOTopoe MQ)KHQ Ha-

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.n;eifcTBHe

CFC1 3, CF 2Cl 2 (ttcrroJih3yeMbIX B xo;rn.n;ttJihnoii npo-

MhIIIIJiennocTtt u rrpott3Bo.n;cTBe a3po3o;ieil:). 3Ttt coe.n;ttneHtt.H pa3JiaraIOTC.H c o6pa30BaHtteM aTOMOB XJIOpa, KOTOpbie KaTaJlM3MPYIOT rrpeBpa~eHHe

030Ha B KllCJiopo.n;.

3a2p.H31te1tue npupo01tbtX eoi) u ucmow,e1tue no'lebt
flcToqHllKH 3arpH3HeHM.H BO,[\ -

c6pOCbl rrpOMbIIIIJieHHblX rrpe,n;rrptt.H-

THH, CMblBbl c ceJibX03yro.n;ttil:, B MOp.HX -

aBapllM necpT.HHbIX TaHKepoB.

3arp.H3Henue rroqB .HBJI.HeTc.H pe3yJihTaTOM rrpoMhirn"-:cenHhix c6pocoB
M HeCOBeprneHCTBa ceJibCKOro X03.HHCTBa.

C0Kpaw,e1tue 6uopa31too6paau.H rrpo6JieMa.

B

3TO e~e o.n;na Bamna.H 3KoJior11qecKa.H

pe3yJibTaTe pacqucTKH rrJio~a.n;eil: .n;JI.H ceJihX03yro,n;uil:, 3a-

rp.H3HeHH.H cpe.n;hI, Bblpy6KM Tporr11qecKHX JieCOB rrpottcxo.n;HT BhlMttpaHHe MHOl'HX BH;'l;OB JKMBOTHbIX MJIM MX IIOJIHOe ynnqTomeHMe.
BonpocbI o pernennn 3KOJioruqecKMX rrpo6JieM cTaB.HTC.H Ha rocy.n;apcTBenHOM M Mem.n;ynapo.n;HOM ypOBHe.

60TAH"1KA

BomaHUIW

(OT rpet.I.

botane -

pacTemrn, Tpasa) -

3TO ttay1rn, n3yqa10rn;aH cTpo-

ettne, iKH3He.n,eHTeJihHOCTh, pacrrpocTpattettHe pacTettnii:. BoTaHHKa o6'be.n,HHHeT

PM ttayK o

pacTeHHHx: attaTOMHIO, MopcpoJiorHIO, QlH3HOJ10rn10, 6noxHMHIO H .n,p.

3apo.m.n;eHHe HayKH 0 pacTeHHHX rrpoM30IIIJIO B JJ:pestteii: fpeUHH. «0TI~OM
6oTaHHKH» C'-IHTaIOT .n;pestterpeqecKoro yt.Iettoro TeocppacTa. «0TI~OM cospeMettttoii 6oTaHHKH» c•mTaeTCH IIIBe,ZJ;CKHii HaTypaJJHCT

XVIII

B. Kapoi JlHHtteH:,

KOTOpbIH pa3pa60TaJI CHCTeMaTHKY H BBeJI 6HHapny10 HOMeHKJiaTypy (,ZJ;BOHHOe
Ha3BaHIIe, T. e. po.n; H BH,11,).

PacmeHuJl -

1.
2.
3.
4.
5.

3To iKHBhie opraHH3MhI, HMerorn;ne cJie.n;yrorn;ne oco6ettttocrn:

ABTOTpOcpHbIH crroco6 IIHTaHH.H (B OCHOBHOM).
IlpMKpeIIJIHIOTCH K cy6cTpaTy.
HeorpaHHt.IeHHhlii pocT (T. e. pocT B Te'-IeHHe sceii iKH3HH).
CttJihHOe pacqJJeHeHHe TeJia .n;JIH yseJIM'-IeHHH KOHTaKTa c BHeIIIHeii cpe.n;oil:.
PacnpocTpaHJIIOTC.H c rrOMOID;hIO cnop, rrJio.n;os, ceMaH.
CTpoeHHe paCTMTeJihHOii: KJieTKH:

IIJIOTHaJI KJieTO'-!HaJI o60JIO'-!Ka, COCTOHID;aJI H3 ueJIJIIOo103bI (B OCHOBHOM);
HaJIH'-IHe IIJiaCTli,ZJ;: XJIOponJiaCTOB, xpOMOTIJiaCTOB, JieHKOTIJiaCTOB;
KPYTIHa.H BaKyOJih c KJieTO'-IHhIM COKOM;
OTCYTCTBHe UeHTpMOJieM H KJieTO'-IHOrO n;eHTpa B KJieTKaX BhICIIIHX paCTeHHM H HeKOTOphIX so.n;opocJieii;
3anacaHHe yrJieso.n;os B sn.n;e KpaxMaJia.

T1mHH H opraHhI Bhicmux pacTeHHii:
TKaHb -

3TO rpyrrna KJJeTOK, HMeIOID;HX CXO)J,HOe CTpOeHHe, BhlIIOJIH.HlO-

rn;nx O)J,HHaKOBhie cpyHKIJ;HH H CXO,ZJ;HhlX no npo11cxom.n;eHHIO. TKaHh MOiKeT
6bITh rrpOCTOH HJIH CJIOiKHOM.

177

Ticanu U opzanbl. 6bl.CIUUX pacmenuii

PMc. 63. BepxyweYHasi Mep111creMa no6era an0Ae111 : a npOAOnbHbl~

npoAonbH1:>1~ pa3pe3;

6-

sHewH111~ BlllA 111

pa3pe3 KOHyca HapacTaHlll5l; B - KneTKlll nepBlllYHO~ Mep111CTeMbl; ,- - napeHXlllMH35l Kner-

Ka n111cra, 33KOHYl'1BW35l Alll<l><l>epeHL\1'1POBKy. 1 - KOHYC HapaCT3H1115l ; 2 - 3aYaTOK n111cra

IIpocmaR mKaHb COCTOHT H3 KJleTOK O,ll,HHaKOBhIX no cpopMe H cpymrn;mIM
(napeHXHMa, CKJiepeHXHMa, KOJIJieHXHMa).

CJlOJIC1WJI. mKUHb COCTOHT H3 KJ16TOK pa3JIH'lHhIX no cpopMe I1 cpyHKI.l,H.flM,
HO HMeIOJ.I.\HX 061.l\ee npOHCXO>K,ZJ,eHJie (RCHJieMa).
Teno BhIClllHX pacTeHHii o6paaoBaHo TKaH.flMH.
JIHlIHhie opraHhI pacTeHHH.

B

Ifa

HHX cpopMHPYIOTC.H pa3-

aaBHCHMOCTH OT oc-

HOBHOH lPYHKI.l,IU'I BhI,ll,eJI.HIOT OCHOBHhie BH,ll,hl TKaHe:ii: o6pa30BaTeJihHhie TKaHH (MepHcTeMhl); OCHOBHhle TKamr; IlOKPOBHhie TKaHH; BhI,ll,eJIHTeJibHbie
TKaHH; MeXaHI.f'leC:KHe T:KaHH; npOBO,ll,.Hl.l\He TKaHIL

06pa30BameJtbllble mKanu ( MepucmeMbt) (pHc.

63-64).
<l>yH:KI.l,HH: ,n;eJieHHe H ,ll,HlP!PepeHI.T,HpOBKa.
XapaKTepHCTHKa: KJieTKH MeJIKirn, He,n;nipipepeHI!HpoBaHHhre, mHBhre, c TOHKOH KJieTOlIHOH
060JIO'IKOH; BaKyoJieH o6bllIHO HeT, '!aCTO ,n;eJI.HTC.11 MHTOTH'leCKH,

cpopMHPYIOT

Bee OCTaJibHhie

3

TKaHH pacTeHHH.

llo BpeM.enu

8031tU1CflOBeHU.R BhI,ll,6JI.flIOT nepBHlI-

Bhle n BTOPH'lHhie MepHCTeMhI. IIepBHlIHhre MepncKOTOpbie 3aKJia,ll,blBaIOTC.H

PMc. 64. CxeMa pacnonolKe-

B TeJie aapo,n;hmra. 8To 3apoJJ.hIIU ceMeHH, 30Hhl poc-

Hlll5l pa3nlllYHblX Mep111CT6M B

TeMh! -

MepHCTeMbl,

Ta CTe6JI.H B BblCOTY H 30Ha pocTa KOpH.H. BTopHlIHhle

pacTeH111111: 1 -

'5'.epHcTeMbl -

(am1KanbHa5l) ;

3aKJia,ll,hIBaIOTC.H no3,n;Hee, o6ecneqH-

sepxyweYHasi

2 -

111HrepKa-

8aloT POCT CTe6m1 B TOJIJ.I.\HHY (KaM6Hii cocy,n;HCTO-

nsipHasi (BCT3BOYH3Sl), 3 -

BOJIOKHHCThIX IlY'lKOB H npo6KOBblH KaM6:nii).

KOsasi(naTepanbHasi)

60-

BommtuKa

178

II o pacnoJLOJlceHuJO

BhI,D;e"i:moT:

BepxymeqHyIO (amrnaJibHYIO) MepHCTeMy -

B 30He pocTa KOpm'l H KOHyc

HapacTamu1 rro6era;
60KOBhie (KaM6Hii:, rrepm.i;HKJI) -

o6ecrreqnBaIOT pOCT CTe6Jieii: M KOpHeii:

B TOJIIIJ;MHy;
BCTaBOqHhie -

OCTaTOK BepxyrueqHoii: MepMCTeMhI; pacrrOJIOiKeHbI B OC-

HOBaHMH :vrem.n;oy3JIHll CTe6JIH l1 MOJIO,D;hIX JIHCTheB H 06ecrreq11BaIOT pOCT
Mem.noy3J1Mll, o6pa30BaHHe rro6eroB, JIMCTbeB, rrpn.n;aTOqHhIX KOpHeii:;
paHeBhie -

o6pa3yIOT 3aIIJ;HTHblll CJIOii: (rrpo6Ky) Ha.n; MeCTOM IIOBpem.n;e-

HMH TKaHeii.

0cH06HaR m1calib ( napeHxu.111a)
XapaKTepMCTMKa: KJieTKM KPYIIHhie, HerrpaBMJihHOii: cpopMhr, Mem.n;y HHMM
6oJihlllMe MemKJieTHMKM.
Ilo BhlIIOJl:HHeMhIM <}:>yHKIJ;MHM BhI,ZJ;eJIHIOT HeCKOJihKO TMIIOB rrapeHXMMhI:
a) aCCMMMJIHIJ;HOHHaH rrapeHXHMa: OCHOBHaH cpyHKIJ;HH -

cpOTOCHHTe3; co-

.n;epmHT XJIOpOIIJiaCThI; pacrroJiomeHa B JIMCThHX H MOJIO,ll;hIX CTe6JIHX;
6) 3arracaIOrri;aH: HaKorrJieHHe 3anacHhIX Berri;ecTB (6eJIKH, mnphr, KpaxMaJI). IlpHCYTCTByeT BO Bcex opraHax pacTeHMH, B OCHOBHOM B CTe6JIHX, KOpHHX, KOpHeIIJIO,ZJ;ax, KJ1y6HHX, JIYKOBMri;ax, IIJIO.n;ax -

B JIMCThHX MeHhllle.

KJieTKH KPYIIHhie; crreri;HaJIH3MpoBaHHaH 3anacaIOrri;aa rrapeHXMMa ceMeHM Ha3hrnaeTCH ::rn.n;ocrrepMoM;
B) B03,ll;YXOHOCHaH: OCHOBHaH cpyHKIJ;HH -

BeHTHJIHIJ;MH; Mem.n;y KJieTKaMM

oqeHh 60JihlllMe Me)KKJieTHMKM, CBH3aHHhie c BHelllHeii: cpe.n;oii: qepe3 OTBepCTMH yCThHIJ; H qeqeBMqeK;
r)

BO,ll;OHOCHaH: crroco6cTBYeT y.n;epmaHMIO BJiarH pacTeHMHMH; KJieTKH co-

.n;epmaT B BaKyOJIHX C.HM3HCTbie Berri;ecTBa, y.n;epmHBaIOrri;Me Bo.n;y; xoporuo pa3BMTa y BO,ll;HhIX pacTeHHll My CYKKyJieHTOB (KaKTYChI, aJI03, araBhI);
.n;) BCaCbIBaIOrri;aH (3IIH6JieMa, Ha3hIBaeMaH ·raKme PH30,ZJ;epMOll, MomeT
6bITh OTHeCeHa K rrepBHqHoii: IIOKPOBHOll TKaHH KOPHH): KJieTKH 3IIM6JieMhI
HecyT KOpHeBbie BOJIOCKM, c IIOMOIIJ;hIO KOTOpbIX npoHCXO,ll;HT rrorJiorri;eHHe
MHHepaJibHhIX Berri;ecTB l1 BO,ZJ;hl H3 rroqBhI; o6pa3yeTCH B 30He BCaCbIBaHHH
KOpHeii:.

II OKPOB!ibte mKallu
BwrroJIHHIOT 3a11i;ttTHyIO cpyHKIJ;HIO. 3arri;Hrri;aroT OT BhIChIXaHMH, rreperpeBa,
rroe.n;aHMH mHBOTHhIMM, 3apameHMH 6aKTepMHMH H rpH6aMH. Bo3HHKJIH c BhIxo.n;oM pacTeHHii Ha cyllly (HaqMHaH c rrcMJIOcpMTOB B rraJieo3oe). Pa3JIHqaroT:
rrepBHqHyro rroKpoBHYIO TKaHh BTOpHqHyIO IIOKPOBHYIO TKaHb TpeTHqHyIO IIOKPOBHYIO TKaHb -

:JnuiJepMa ( KOJICUU,a ):

:muiJep.My;
nepuiJep.My;
KOpKy, MJIM pumuiJO.M.

KJieTKH mMBhie, IIJIOTHO npMJieraIOT .n;pyr K .n;pyry,

JIHUieHbI XJIOpOIIJiaCTOB, IIOKpbIThI KYTHKyJioii:; CJIOmHaH TKaHb COCTOHT M3:
3IlH,ZJ;epMaJibHhIX KJieTOK
(y ,ll;BY,ZJ;OJihHhIX Hhie, pa.n;aMH);

-

eCTb Jie:HKOIIJiaCThI, HeT XJioponJiaCTOB

M3BHJIHCTOH cpopMhI, y O,ll;HO,ll;OJihHhIX -

rrpHMoyroJih-

TKanu u opzanbL 6bLCuiux pacmenuii

179

3aMb1Ka10IIJ;l1X KJieTOK yCTbHD; -

HMelOT 6o60BH,z:t;HYIO cpopMy, Mem.n;y HH-

MH yCTbH'IHa.H w;eJih;
OKOJIOYCTbH'IHbIX ( rro60'IHbIX) KJieTOK;
Tpl1XOM -

pa3JIH'IHbIX ITO cpopMe, CTpOeHl1IO l1 cpyHKD;l1.HM BblpOCTOB

KJieTOK 3ITl1,z:t;epMbl (BOJIOCKl1,
Hble y ceM.HH XJIOII'laTHl1Ka -

Ycmbuu,e -

u~eTl1HKH, 'leIIIyHKH; Ha1160Jiee ,z:t;JIHH-

5-6 CM).

3TO MemKJieTHHK Mem,n;y ,z:t;BYMH 3aMb1Ka10w;11M11 KJieTKaM11 6o-

6oBl1,z:t;HOM cpopMbl. lIMelOTC.H B OCHOBHOM Ha Jil1CTb.HX, eCTb Ha CTe6Jie. 06bI'IHO
paCIIOJI0'1<eHbl Ha HHJKHeH CTopoHe Jll1CTa, y BO,z:t;HbIX pacTeHl1H -

IIepuiJepMa -

Ha BepxHeii:.

cJiomnaH noKpOBHa.H TKaHh CTe6Jieft, KOpHeB11w; l1 KOpHeft

NIHoroJieTHl1X pacTeH11ii (peme -

o,n;HoJieTH11x). CMeHHeT an11,n;epMy. Ilep11,n;ep-

Ma 06pa3yeTcH 113 Mep11cTeMbI -

cpeJIJioreHa. <l>eJIJioreH 3aKJia,z:t;hrnaeTc.H B oc-

HOBHOH rrapeHx11Me no,n; an11,n;epMoii:. KJieTKl1 cpeJIJioreHa ,n;eJI.HTc.a: Hapymy OTKJia,ZJ;bIBaIOT KJieTKl1 npo6KH (MepTBbie, nponnTaHHhie cy6epl1HOM (T11na BOCKa),
a BHYTPh -

:>KHBbie napenx11MHbie KJieTKl1 cpeJIJio,n;epMbI c xJioponJiacTaMH.

TaKHM o6pa30M, nepH,n;epMa cocTOl1T 113 cpeJIJiorena, npo6KH (cpeJIJieMbI)
H <t>eJIJIO,z:t;epMbl.

llpo6Ka -

COCTOHT H3 MepTBbIX KJieTOK, KJieTO'IHa.H CTeHKa npon11TaHa JKH-

pono,n;o6HblM cy6ep11HOM. KJieTKl1 HMeIOT npHMoyroJihHYIO <t>opMy, TIJIOTHO
npHJieraIOT .n;pyr K ,n;pyry, pacnoJiomeHhI p.a,n;aM11. Ilpo6Ka coxpan.HeT TKann OT
IlOTepH BJiarH,

npOHl1KHOBeHl1.H Ml1KpoopraHH3MOB,

o6pa3y.H

«cpyTJI.Hp».

,lJ;JIH raaoo6Mena no,n; yCTbHD;eM Ha MeCTe B03,z:t;YXOHOCHOH IlOJIOCTH <t>eJIJroreH
OTKJia,ZJ;bIBaeT iRl1Bble, pbIXJIO pacnoJiomeHHbie napeHXHMHbie KJieTKH (BbIIlOJIH.HIOIIJ;a.H TKaHb), KOTOpbie pa3pb1BalOT 3IlH,z:t;epMy H C03,z:t;aIOT B03MO:>KHOCTb ra3006MeHa H TpaHCilHpau;n11 c BHeIIIHeM cpe,n;oii:. 8To 06paa0Ban11e Ha3bIBaeTC.H

'te'le6U'lKOU.

0'1eHb KpynHble 'le'leBl1'1Kl1 (.n;o

15 CM) o6paayIOTC.H y

6epe3bl (no-

nepe'IHbie IlOJIOCKH l1 qepTO'IKH).

KopKa (pumuiJoM) -

TpeTH'IHa.H noKpOBHa.H TKaHh. 06paayeTCH y Mnoro-

JieTHHX pacTeHHM B KOpHe, CTe6Jie, KOpHeBnw;e. Kam,z:t;hIH ro,n; cpopMHPYIOTC.H
6onee rJiy6oKne CJIOH nepn.n;epM, napymHhie cJIOH H pacnoJio:JKeHHhie Mem,n;y
HHMH TKaHH OTMHpaIOT. KopKa o6hI'IHO Tpew;HHOBaTa.H H HepOBHa.H, B OTJIH'IHe
OT rJia,n;Koii: nep11,n;epMbI. KopKa aaw;Hw;aeT OT MexaHH'leCKHX noBpem,n;eHHH,
pe3KOH CMeHbl TeMnepaTyp, npOHHKHOBeHH.H MHKpoopraHH3MOB.

BbtdeJtumeJtbHbte mKanu
CJiymaT ,ZJ;JIH naKonJieHHH H BbI,ZJ;eJieHHH Bew;ecTB H3 opraHH3Ma pacTennii:.
Ilpe,n;cTaBJieHbI TKaH.HMH napymnoii: H BHyTpenneii: ceKpeu;nn. KJieTKH BhI,z:t;eJIHTeJibHhIX TKaneii: no cpopMe o6bI'IHO napeHXHMHhie, TOHKOCTeHHbie. 0HH
.ll;OJiro OCTalOTC.H :>KHBblMH, BbI,z:t;eJI.H.H ceKpeT.
TKann
llHHMH:

Bnympenneu ceKpeu,uu

KJieTKH-H,z:t;H06JiaCTbl -

npe,n;cTaBJieHhI paaJIH'IHhIMH o6paaoBa-

OT,z:t;eJihHbie KJieTKH, co,n;epmaw;ne KpHCTaJIJihl

OKCaJiaTa KaJib~H.H, TepneHOH,z:t;bl, CJIH3H;
BMecTHJIHw;a BbI,z:t;eJieHHM -

302eHHbte 6.MecmuJtuw,a,

IlOJIOCTH B TOJiw;e ,n;pyrnx TKaneii:. lIMeIOTC.H

cxu-

KOTOPbie B03HHKaIOT B BH,z:t;e MemKJieTHHKOB, 3a cqeT

Eomanu1w

180

pacxom,n;ettiui JKHBbIX 1rneTOK. CeKpeT KJreTOK BhI,D;eJrneTca n IIOJIOCTh MemKJJeTmma (3qmpttbre :-.rncJia, CMOJ!bI, CJIH3b) H JIU3Ucel-tltbte 6MeCmUJIUU{G, B03mmaIOilIHe B pe3yJihTaTe pacrra,lla (JIH3nca) KJieTm< nocJie HaKoIIJiemrn ceKpeTa B MeJKKJ":leTHHKe (HaIIpHMep, :Jq:mpHhie MacJia y !IHTPYCOBblX);
CMOJIHHbie XO,llhI -

o6pa3yIOTCfl CXH3ore1rno, H3HYTPH BhICTJiaHhI ceKpe-

THPYIOI!IHMH KJieTKaMH. MoryT BeTBHThca, co,n;epmaT CMoJiy;
MJieqHHKH -

c11cTeMa noJiocTeii, npoHH3hIBaIOilIHX see pacTeHne. Co-

,n;epmaT M.rreqtthrii coK -

3MYJihCHIO 6eJioro, peme -

opattmesoro !IBeTa

(qncTOTeJI), co,n;epmailIYIO CMeCh pa3JIHqHbIX BeilieCTB (HaIIpHMep, KayqyKOHOChI, '!eBeJI, KOK-carbr3).
Bhr,n;eJIHTeJihHhie TKaHH e11eurneu ceKpeu,uu (csaaaHhI c noKposaMn pacTemrn):
meJie3HCTbie BOJIOCKH (ttanpnMep, repatth :m:eJie3KH -

o,n;Ha HJIH HeCKOJihKO ceKpeT11pyIOilIHX KJieTOK

sqmpHhre MacJia, coJieBhr,n;eJreHtte);

MHoroKJieToqttaa ceKpenrpyrou,a.H CTPYKTypa, Bhr,n;emuo-

IIIaa sc}mptthre MacJia;
HeKTapHHKH -

pa3Hoo6pa3Hhie CTPYKTYPhI, BhI,n;eJIHIOilIHe HeKTap: ca-

xapncThIM COK (paCTBOp caxapos, 6eJIKOB, CIIHpTOB, apoMaTHqeCKHX BeilieCTB). PacIIOJIOJKeHbI B OCHOBHOM B !IBeTKax (Ha qarneJIHCTHKax, JieneCTKaX, CTeHKax 3aBH3H, !IBeTOJIOJKe).

Mexa11u•tec1Cue mKaHu
06ecne'lHBaIOT onopttyro cpyHK!IHIO, npM,n;a10T npoqttoCTh opraHaM pacTeHHii (Hatt6oJiee pa3BHThI B cTe6Jie). KJieTKH HMeIOT attaqMTeJibHO yToJIilieHHhie KJieToqHbie o6oJioqKH.
Pa3JIHqa10T ,n;sa THna Mexatt1111eCKHX TKaHeii:

1.

KoJt.1ie11xuMa -

TKaHb, KJieTKH KOTopoii HMeIOT ttepasHOMepHo yTo.,1-

ilieHHhie KJieToqHhre cTeHKH. KJieTKH BhITHHYThI s,n;oJih OCH opraHa; KJieTKH
JKHBbre, co,n;ep:m:aT XJioporrnacThr. B no,n;aeMHhIX oprattax He BCTpeqaroTcH; KJieT011HaH CTCHKa COCTOHT H3 Ile.,1JIIOJI03hl, reMH!IeJIJIIOJI03hl, neKTHHOBbIX seilieCTB.
B 3aBHCHMOCTH OT THila YTOJiilieHHH KJieTOqHblX CTeHOK BbI,D;eJIHIOT ,n;sa THna KOJIJieHXHMbl:
a) yroJIKOBa.H KOJIJieHXHMa -

KJieTKH HMeIOT cpopMy rneCTHyrOJibHOro MHO-

rorpaHHHKa. YTOJiilieHMe o6oJIO'lKH rrpottcxo,n;MT c «yroJIKOB» KJieTKH.
BcTpeqaeTca B cTe6Jiax ,n;sy,zJ;OJihHhIX pacTeHMii, B qepernKax JIHCTbeB no
o6eHM cTopoHaM KPYITHhIX JKHJIOK JIHCTa;
6) IIJiaCTHH'laTaH KOJIJieHXHMa -

KJieTKH HMeIOT BH,D; rrapaJIJieJieIIHITe,n;a,

YTOJiilieHHyro KJieToqHyro o6oJioqKy ,n;syx rpaHeii. Haxo,n;MTCH B cTe6JIHX
,n;peseCHblX pacTeHHM.

2.

CKJtepeHXUMa -

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10 - KOpHesoi'i BOllOCOK
KJieTKH MHoroyrOJihHhle, IIJIOTHO COMKHYThI; KJieToqHbJe

CTeHKH MOryT npOIIHThIBaThC.H cy6epHHOM (onpo6KOBeHHe) HJIH JIHrHHHOM (o,n,peBeCHeHHe) H BhIIIOJIH.HTh 3ar:QHTHYIO <l>YHKD;HIO;

2)

Meao,n;epMa -

o6paayeT OCHOBHYIO qacTb nepsHqHOM KOphI, COCTOHT H3 na-

peHXHMHhIX KJieTOK; MOmeT BbIIIOJIH.HTh aanacaIOI:QYIO <l>YHKD;HIO If <l>YHKD;HIO
npoBe,n;eHH.H;

3)

aH,n;o,n;epMa -

o,n;Hop.H,ll,HhIM cnoil: KJieTOK. KJieTKH HMeIOT oco6hie yTOJI-

I:QeHH.H Ha pa,n;HaJibHbIX KJieToqHhIX CTeHKaX -

IIO.HCKH KacnapH.

c IIOMOI:QhlO

3THX yTOJII:QeHHM nepeKpbIBaeTC.H nepe,n;BH:>KeHHe pacTBopoB B,ll,OJih KJieToqHhIX
CTeHoK. ToncTOCTeHHhie KJieTKH .HBJI.HIOTC.H OTMHpaIOI:QHMH, :>KHBhie CKHhle KJieTKH -

npony-

e,n;HHHqHhie KJieTKH c Heo,n;peBecHeBIIIHMH cTeHKaMH. Ocy-

I:QeCTBJI.HIOT CB.H3h Mem,n;y nepBH'IHOM KOpoil: If oceBhIM D;HJIHH,ll,pOM.

Oceeou U,UJlunop COCTOHT l'.13 HeCKOJihKHX CJIOeB KJieTOK:
nepHD;HKJI .HBJI.HeTC.H

(<

KOpHepO,ll,HhIM CJIOeM » ,ll;JIJl o6pa30BaHH.H 6oKOBhIX

KOpHeil: (KJieTKH :>KHBble, TOHKOCTeHHhie, pacnOJIO:>KeHbl B O,ll,HH pa,n;);
npoKaM6HM -

60KOBa.H MepHCTeMa;

cocy,n;HCTO-BOJIOKHHCTbIM IIY'JOK, COCTO.HI:QHM H3 nepBifqHOM <t>noaMhl
H nepBHqHOM KCHJieMhI.

AnamoMU'tec1we cmpoenue 1wpna

187

l\:CIIJieMa H cpJI03Ma pacnOJIO)IWHbl qepe.n;yIOIU;HMHCH JiylJaMH H3 u;eHTpa. 06pa3yeTCH pa.n;HaJihHhIH npoBo.n;Hru;tt:H nytJoK. B -oT.'rnqHe OT CTe6.11H Kop1rn He
nMeIOT cep.n:I1eBHHhI.

y

O,D;HO,ll;OJlhHhIX pacTeHHH CTpoeHHe KOpmI OCTaeTCH 6e3 3Ha'-IHTeJibHh!X I13-

MeHeHHH B TelJeHHe Bce:H )KH3HH pacTeHHH, y .n;By,ll;OJlhHhlX npOHCXO,ll;HT nepexo.n: K BTOPHlJHOMY CTpOeHHIO.

Bmopuiuwe cmpoeuue IWpliH y a6yaoJlbliblX pacmeuuu.

Pa.n;Ha.'lbHOe pacuo-

JIO)KeHHe npOBO,ll;HIU;HX TKaHe:H CMeHHeTCH KOJIJiaTepaJibHbIM. 3To CBH3aHo
c ,n;e.aTeJihHOCThlO KaM6H.H,

KOTOpbIH B03HHKaeT Mem.n;y JiylJaMH KCHJieMhl

n cpJI03MhI. 06pa3yeTCH e.n;HHhr:H CJIOH KaM6HH. R u;ettTpy KaM6HH: OTKJia.n;hrnaeT KJieTKH BTOpHtIHoii KCHJieMhI, a K rrepttcpepHH -

BTOpHtIHo:ii cpJI03MhI. Kpo-

Me Toro, KJieTKH KaM6HH o6pa3yIOT IIIHpOKHe pa.n;HaJibHhie CBeTJihie JiyqH napeHXHMhI Mem.n;y T.HfKaMH BTOPHlJHOH rrpOBO,D;.HIU;e:H TKaHH -

cepou,eBUlillble JlY'lU.

3TO rrepBHlJHbie

0HH o6ecrrelJHBaIOT cpH3HOJIOrHlJeCKyIO CBH3h ueHTpaJih-

HOH qacTH KOpH.a c rrepBwrno:H Kopo:H. IIepBHlJHaH Kopa OTMHpaeT n c6pachrnaeTC.H. IloKpOBHOH TKaHhIO cTaHOBHTC.H rreptt.n;epMa. RoMrrJieKc TKatte:ii cHapy-

BmopU'lliOU KOpoii.
MunepaJLbHoe numauue. Bo.n;y H MHHepaJihHhre

jlCH OT KaM6lrn Ha3hIBaIOT

Beru;ecTBa KopeHh Bcachrna-

eT H3 IIOlJBhI c IIOMOID;hlO KOpHeBhIX BOJIOCKOB. Bo.n;a IIOCTyrraeT B KOpHeBOH BOJIOCOK 3a ClJeT OCMOCa. 3aTeM Bo.n;a npoxO,D;HT no )KHBhlM KJieTKaM rrepBHlJHOH
IWphl KOpH.H H rrorra.n;aeT B KCHJieMy u;eHTpaJihHOro oceBoro IIHJIHH,ll;pa KOPHH.
MHHepaJihHhre Beru;ecTBa BcachrnaroTc.a B xo.n:e rraccHBHoro HJIH aKTHBHoro
TpaHcrropTa qepe3 KJieTOlJHYIO MeM6paHy. B cocy.n;ax pa3BHBaeTc.a rroBhIIIIeHHoe OCMOTHlJeCKOe .n;aBJieHHe. KopHeBoe .n;aBJieHHe H TpaHCilHpau;HH (HcrrapeHHe) o6ecrrelJHBaIOT ,D;BH)KeHHe BO,ll;hl B TeJie pacTeHHH.
BamHyIO pOJih HrparoT KaIIHJIJI.HpHhie CHJlhI:
MeJK.n;y MOJieKyJiaMH BO,ll;hl H

ai)ze3UJl

K02e3UH -

CHJia CI1errJieHH.H

BO,ll;hl K CTeHKaM COCy,ll;OB •

.D:JI.a IIHTamrn pacTeHHIO Heo6xo.n;ttMhI:
MaKpo;:iJieMeHThI: a30T, cepa, cpoccpop, KaJitt:H, MarHHii, KaJihll;HH, meJie3o;
MHKp03JieMeHThI: 6op, Me,ll;h, MapraHeII, K06aJihT, IIHHK H .n;p •
.D:JI.a YJIYlJIIIemrn ypoma.a BHOCHT y.n;o6peHH.H. Y,n;o6peHHH pa3.n;cJI.HIOT Ha opraHHlJeCKHe (Topcp, HaB03 H .n;p.) H MHHepaJihHhie (ceJIHTpa, MOlJeBHHa H .n;p.).
BH.n;OH3MeHeHH.H

KopnenJLOObl -

KOpHe:H:

BhIIIOJIHHIOT 3arracaroru;yIO cpyHKIIHIO. Bo3HHKaIOT KaK

YTOJIIU;eHHe rJiaBHOro KOpH.H (MOPKOBh, CBeKJia, perm H .n;p.).

KopneKJLy6nu - 3arracaroru;tte rrptt.n;aTOlJHhie KOpHH (6aTaT, reoprHH).
BmHzuewow,ue Kop1w - ocyru;ecTBJI.HIOT norpyJKeHHe B rrotJBY (JiyKoBHqHhle pacTeHH.H).

Bo30YUllible KOpllU -

BCaChIBaIOT Bo,n;y H MHHepaJihHbie Beru;ecTBa H3

BJiaJKHOro B03,n;yxa (y pacTeHHH-3IIHcpHTOB -

opxH.n;en).

j(blXameJlbllbte 1wpnu IIO,D;HHMaIOTCH Ha.n; IIOBepxHOCThlO IIOlJBhI
(y pacTeHHH 60JIOT).
XooyJLbHbte Kopnu - y MaHrpoBhIX pacTeHnii Ha JIHTopaJIH Mop.a. YKpenJI.HIOT pacTeHH.H B rpyHTe.

188

BomanuKa

a
Pt.tc. 70.

reHeparnsHasi nOYKa 6y3111Hbl; B nlllCTbeB;

6

CTpoeH111e pa3nw-1HblX rnnos no4eK:

3-

a -

6

serernrnsHasi nOYKa AY6a:

reHepaTYIBHaR nOYKa BlllWHlll. 1 -

6 -

serernrnsHo-

nOY8YHbl8 Yewy111; 2 -

3aYaTKlll

3aYaTOK u,BeTKa

CTeoeJIL
Cme6eJlb -

aTo oceBa.a qacTh rro6era pacTeHH.H, cocTo.arn;a.a H3 Y3JIOB

H MeJK,n;oy3JIHH. CTe6eJih qarn;e Bcero HMeeT u;mmtt,n;pttqecKyIO cpopMy H B rrorrepeqHOM ceqeHHH MOJKeT 6bITh OKpyrJihIM, yrJIOBaTbIM, MHOrorpaHHhIM
H ,n;p.
<I>yHKI.J;HH CTe6JI.a: TpaHCIIOPT BO,Zl;hl H MHHepaJibHhIX Beu~eCTB H3 KOplrn.
B JIHCTh.H, opraHHqecKHX Bern;eCTB H3 JIHCTheB B KopeHh; yBeJIHqeHHe IIOBepxHOCTH pacTeHH.H rryTeM BeTBJieHn.a; o6pa30BaHne JIHCTheB ll JIHCTOpacrrOJIO:>KeHHe; o6pa30BaHHe I.J;BeTKOB; 3arracaHHe BO,ll;hl H IIHTaTeJihHbIX Bern;ecTB; BereTaTHBHOe pa3MHO:>KeHHe.

Ilo6ez -

aTo cTe6eJih c JIHCTh.HMH H rroqKaMH. Ilo6er pa3,n;eJieH Ha y3JihI,

B KOTOphIX pacrroJioJKeHhI JIHCTh.H H MeJK,n;oy3JIH.H. MeJK,ZWY3JIH.H 6brnaIOT yKopoqeHHhle H y,n;JIHHeHHbie.

IlO'llCa -

3TO 3aqaToqHbIH, He pa3BepHyBilIHHC.H rro6er. I'JiaBHhIH rro6er 06-

pa3yeTC.H H3 BepxyllleqHo:H rroqKH, 6oKOBhie rro6ern -

H3 rra3yllIHhIX noqeK.

IJoqKH 6hIBaIOT eezemamuBHbt.Mu, eezemamueHo-zeHepamumtbt.MU H zeHe-

pamuBHbt.MU (pnc. 70).
lfa eezemamUBHOU llO'llCU o6paayeTC.H CTe6eJib c JIHCTb.HMH H rroqKaMH.
BezemamuBHo-zeHepamueHaR no'l/Ca o6pa3yeT rro6er c u;BeTKOM HJIH cou;BeTHeM.

I'eHepamuetta.fl llO'l/Ca o6pa3yeT O,ll;HHQqHbIH I.J;BeTOK HJIH COI.J;BeTHe.
BeTBJieHHe no6era
BeTBJieHne rro6era Heo6xo,n;HMO ,n;JI.a yaem1qeHH.H rrJiorn;a.n;H TeJia pacTenmr.
Bhl,n;eJI.HIOT ,n;Ba THrra BeTBJieHH.H rro6era:

1. Bepxyme'lHoe eemeJteHue.

KoHyc HapacTaHH.H ,n;eJIHTc.a Ha ,n;Be qacTH,

KaJK,n;a.a H3 KOTOphlx 06pa3yeT rro6er. TaKoe BeTBJieHHe Ha3hIBaeTc.a TaKme

xomo.Mu'leCKoe (rrJiayHhI).

iJu-

Cmpoeuue cme6nR

o(JiwaO/l,bllblX

pacmeuuii

189

PMc. 71. Ta.mbl BSTBJ18Hlll51: a -AlllXOTOMlllYeCKOe BeTB11ett111e (BOAOPOCJlb AlllKTlllOTa); 6 - MOHOnOAlllaJlbHOe BeTBJlett111e (no6er COCHbl);
Myx111); r -

B -

ClllMnOA1118JlbHOe BeTBJlett111e no rnny MOHOX8311151 (no6er Yepe-

c111MnoA111a11bttoe seTsne>me no T111ny A11txa311151 (no6er c111pett111). L.J,111cppaM111 o6o3ttaYett

nop51AOK no6era

2. Bo1C0Boe BemBJteHue.

II06er11 06pa3y10TcH H3 rra3yIIIHhIX rroqeK.

BoKoBoe BeTBJiem1e 6hrnaeT Tpex THITOB (pHc.

71):

1) MOHOITO,D;HaJibHOe -

KOHyc HapacTaHHH coxpaHHeTCH BCIO JKH3Hh, 06ec-

lfa

rra3yIIIHhlX rroqeK o6pa3ylOTCH 60KOBbie rro6erH (BeT-

rreq11BaeT pOCT.

BH). 3TOT THIT xapaKTepeH .D;JIH rOJIOCeMeHHhlX (eJih, COCHa H .n;p.),
.D;JIH HeKOTOphlX IlOKphlTOCeMeHHhIX (.n;y6, 6yK H .n;p. );
emero.n;Ho BepxyIIIeqHaH rroqKa OTMHpaeT, a poCT

2) CHMITO,D;HaJibHOe -

H.D;eT 3a cqeT 60KOBOH rroqKH (TOITOJih, 6epe3a H .n;p.);

3) JIO:>KHO.D;HXOTOMHqecKoe (BapHaHT CHMITO,D;HaJibHOro) -

BepxyIIIeqHaH

rroqKa OTMHpaeT, a pOCT H,D;eT 3a cqeT .D;BYX 6oKOBhIX rroqeK (cHpeHh).
Ilo Timy pocTa rro6ern 6brna10T: rrpaMocToaqne (Bee BpeMH pacTeT BBepx) OOJihIDHHCTBO pacTeHHif; rrp11rro.n;H11Ma10m;Heca (rrJiayH 6yJiaBOBH.D;HhIH); cTeJI10lqHeca (ecJin OH yKopeHaeTCH, TO Ha3hIBaeTCH IlOJI3yqHM, HarrpHMep KJieBep IlOJlQyqH:H, 6y.n;pa IlJIIOW:eBH.D;HaH); BhlOW:HeCH (BblOHOK); u;errmnom;HeCH (xMeJih).
CTpoettHe cTe6Jrn: o.n;tto.n;oJILHLIX pacTettu:H

B

attaTOMHqecKOM cTpoeHHH cTe6JIH TpaBHHHCThIX pacTeHHH Bh1.n;eJIHIOT:

no1CpoBHy10 m1CaHb ( anuiJepMa ); nepBu'lny10 1Copy; u,eHmpa.rtbHbtil oceBoil u,u. 4'UniJp (pHc. 72).

Bomanu1w

190

1. J1o1Cp06Ha.H mlCUHb

TOHKI:IH

c.1oii: TIJIOTHO COMKHYThIX BhlTJIHY-

IlepBH"1-IHa.JI Kopa

ThIX KJieTOK, l:IMeeT

2. llepBU'lHa.H !Copa -

yCThIU~a.

o6pa3oBana no.n

3IlI:I,ZJ;epMOH TpeMJI Bl:I,ZJ;a:MI:I TKaneii::
MexaHI:IqecKOH KJieTKI:I;

UempanLHh!H:

TOJICTOCTeHHhie

OCHOBHOH

napeHXl:I-

MOH (cpoTOCI:IHTe3Mpyro~aJI); 3ana-

oceeoH. uw..1HHJlP

caro~eii: napeHXl:IMOM ( 0,Zl;l:IH CJIOH
KJieTOK).

3. IjeHmpaJlbHblU oceBOU U,UJlUHOp

CO-

.nepJKI:IT cocy ,Zl;l:ICTO-BOJIOKHl:ICThie
nyqKII (scer.na 3aKphIThie). TipocTpattCTBO Mem.ny nyqKa:Ml:I 3am:ITO napeH-

B

XHMOH.
PMC.

72.

cocTaB Kam.noro nyqKa

BXO,Zl;JIT BOJIOKHa Mexami:qecKOH TKaAHaTOMi1148CK08 CTp08Hi118 CTe6-

HH H

ml OAHOAOnbHOro paCT8Hi115l Ha nonepe4HOM cpe3e: 1 -

3n111AepMa; 2 -

xnopoqmnnoHOCHaH;
pi11U,111Kni1148CKafl;
Tbl~ ny40K

4 -

3 -

napeHx111Ma

B

CKnepeHX111Ma ne-

TKaHh ,ZJ;BYX THTIOB:

<jJJl03Ma.

cepiJu,eeuna -

CKOTIJieHI:Ie

KPYTIHhIX KJieTOK c TOHKHMI:I CTeHKa-

(4a - cj>no3Ma, 46 - KC111neMa) e

5 -

l:I

u;eHTpaJihHOH qacTII cTe6JIJI pac-

noJiomeHa

COCYA111CTO-BOnOKHll1C-

CKnepeHXll1MHO~ 06KnaAKe;

npOBO,ZJ;JI~aJI

ICCUJleMa

MH.

napeHXll1Ma

y

3.JiaKOB Ha MeCTe OTMeprneii:

cep.nu;eBI:IHhl

ocHOBHaH, 3anaca10w,a51

06pa3yeTCJI B03,ZJ;YIIIHaJI

IlOJIOCTh (CTe6eJih -

COJIOMHHa).

CTpoeHHe cTe6m1 ;:i;peuecH&IX pacTeHHii
,n:epeBhJI H KYCTapHHKl:I HMeIOT MHOroJieTHHe o.npeBecHeBIIIHe CTe6JIH, cnoco6Hhle yTomu;aThCJI. YTomu;eHne, HJIH BTOpnqHhIH pocT CTe6JIJI, nponcxo.n11T
3a cqeT ,ZJ;eJITeJihHOCTH o6pa30BaTeJihHOH TKaHH -

B

KaM6Irn.

CTe6Jie MHOrOJieTHero .npeBeCHOro pacTeHHJI BhI,ZJ;eJIJIIOT OCHOBHhie qacTH,

nepuiJepMa, !Copa (nepBnqHaJI n BTopnqHaJI), 1CaM6uu, iJpeeecuHa, cepiJu,eeuna (pnc. 73).
llepuiJepMa COCTOHT H3 HeCKOJlhKHX CJIOeB: OTMeprnaJI ann.nepMa (O,ZJ;HH

aTo:

CJIOH KJieTOK); npo6Ka (nepn.nepMa) -

MepTBhie KJieTKH, rrporrnTaHHhie BOCKO-

IIO,ZJ;06HhlM Be~eCTBOM cy6epHHOM.

I1ep6U'iHa.H !Copa

BKJllOqaeT: IIJiaCTHHqaTyIO KOJIJieHXHMY (MexaHnqecKaJI

TKaHh); XJIOpocpHJIJIOHOCHYIO rrapeHXHMy; O.Zl;HOPJI,ZJ;HhlH CJIOM KpaxMaJIOHOCHhlX KJieTOK -

(<

KpaxMaJIOHOCHOe BJiaraJIH~e ». 8TOT KpaxMaJI pacTeHHe He

pacxo.nyeT, IIO-BH.Zl;HMOMy, OH nrpaeT pOJih B no.n.nepmaHHH paBHOBeCHJI pacTeHHJI.

BmopU'lHa.H !Copa

BKJIIOqaeT CJIOH Jiy6a, COCTOJI~HH H3 Jiy6JIHhIX BOJIOKOH

(MexaHll'IeCKaJI TKaHh) H

CHTOBH.Zl;HhlX Tpy6oK c

KJieTKaMH-CilYTHI:ID;aMH,

a TaKjKe Jiy6aHyIO rrapeHXHMy.

KaM6uu -

JICY

CJIOH MepncTeMaT11qecKHX KJieTOK, KOTOphIH OTKJia,ZJ;hIBaeT

CTe6JIJI 3JieMeHThl

emopU'l1WZO

Jiy6a,

BHympb -

Hapy-

3.JieMeHThl BTOpllqHoii .npeBe-

BuiJou3MeJteHUH no6eza

191

cuHbI. KJieToK .n;peBeCHHhI aHa'lttTeJihHO
6oJihllle.

fpatttt:u;a KOPhI rrpoxo.n;nT no

KaM6HIO.

Jl,peeecuna

COCTaBJIJieT OCHOBHYIO

qaCTb CTe6mI H B OCHOBHOM o6paaoBaHa 3JieMeHTaMH
Mbl -

rrpOBO,ll;HIIJ;e:i1

cocy,n;aMH.

fO,ZJ;H'IHhie

CHCTeKOJih:u;a

emerO,ZJ;HhIH rrp:ttpOCT .n;peBeCHHhI 3a

o,n;uH BereTaIJ;HOHHhIH rrep:tto,n;. Bectto:H::
KaM6HH OTKJia,ZJ;hIBaeT .n;peBeCHHY' KO-

6

Topaa COCTOHT H3 IIIHPOKHX, TOHKOcTeHHbIX KJieTOK (pbIXJiaH, CBeTJiaH).
K cepe,n;:tttte JieTa OTKJia,ZJ;hIBaIOTCH MeJIKHe TOJICTOCTeHHhie KJieTKH

c

60JihIll

IllHM KOJIH'leCTBOM ,n;peBeCHHHhlX BOJIOKOH (MexaHH"'l:eCKaH TKaHh). Ilo ro,!l;HqHbIM KOJihIJ;aM MOiKHO orrpe.n;eJIHTh
B03paCT pacTeHH.H.

Cepau,eeuna

cocTOHT Ha napeHXHM-

HbIX KJieTOK. B :u;ettTpe 3TH KJieTKH 60-

BH,!J;OH3MeHeHHH rro6era (pHc.

y

CrpoeHvie APeseCHoro cre6m1

(Ha npviMepe cre6m1 nvinb1): I -

74)

KaM6vii;;;

MHOrHX pacTeHHH B noqBe o6paay-

IOTC.H no.n;aeMHhie rro6erH,

B KOTOphIX

1 - orMepwvii;; 3nvi.QepMvic; 2 - nepvi.QepMa;

3- KOnneHXVIMa; 4 -

5-

peHXVIMa;

Hhle Beru;ecTBa. KpoMe Toro, tta,n;aeMHhie

BVIHHbli;; nyy;

no6erH MoryT BH,ZJ;OH3MeH.HThCH B opra-

.n;.

Kapa; II -

Ill - .QpesecviHa; IV - cep.Qu.em·1Ha.

HaKaITJIHBaIOTC.H aanaCHhie IIHTaTeJih-

Hhl IlPHKpenJieHH.H, aaru;HThI HT •

9

IV

P111c. 73.

Jiee KPYIIHhie H qacTo OTMepIIIHe.

aCCVIMVlmlU.VIOHH35l na-

yYaCTKVI ¢n03Mbl;

7-

HOrD KOnbu.a;

6- C8PAU.8-

OC8HH5l5l AP8B8CVIH3 rD,QVIY-

8 - B8C8HH5l5l AP8B8CV1Ha ro9 - nepBVIYHaA KcvineMa

,[\VIYHOrD KOnbu.a;

1. IIoa3e.MHbte no6eiu:
KOpHeBHIIJ;e (nanopOTHHK, JiaH,ZJ;bIIII) -

MHOrOJieTHHH rro6er, HMeIOIIJ;HM pe-

.n;yn;HpoBaHHbie JIHCTh.H B BH,ZJ;e qeIIIyeK, B naayxax KOTOphIX HaXO,ZJ;.HTC.H naayrnHhle noqKH. Ha KopaeBttru;ax o6paayIOTCH np:tt,n;aToqHhle KOPHH. PaattoBH,ZJ;HOCTLIO KOpHeBHIIJ; .HBJI.HIOTC.H CTOJIOHhl KapTocpeJI.H (He,ZJ;OJirOBe'IHhI);
KJiy6HH (KapTocpeJih) -

qiopMHpyIOTC.H KaK KOHIJ;eBhie HJIH rraayIIIHhie yTOJI-

ID;eHH.H no.n;aeMHhIX no6eroB (cTOJIOHOB). Ha KJiy6tt.HX pacno;rometthI rro<JKH
(rJiaaKH). K;ry6HH c;rymaT ,ZJ;JI.H BereTaTHBttoro paaMHomeHH.H;
KJiy6HeJiyKOBH:a;a (rJia,ZJ;HOJiyc) -

3TO JIHCTBeHHhIH KJiy6eHh, T. e. KJiy6eHh,

IIOKphlThlH qeIIIyeBH,ZJ;HhlMH cyxHMH ITJieHqaThIMH JIHCTh.HMH;
JIYKOBH:a;a -

ca ,!J;OHD;eM.

yKopoqeHHhlH rro6er, CTe6JieBa.H qacTb KOTOporo Ha3hIBaeT-

K ,n;oH:a;y np:ttKpenJI.HIOTC.H JIHCTh.H H qeIIIyH. Ha ,n;omi;e pacrroJia-

raeTc.H BepxyIIIeqHa.H noqKa, H3 KOTOpoii pa3BHBaIOTC.H Ha,n;aeMHhie JIHCTb.H

It ll;BeTOHOCHa.H CTpeJIKa. JlyKOBHIJ;hI CJiyiKaT ,ZJ;JI.H BereTaTHBHOrO pa3MH03teHH.H.

Bomanu1w

192

•

.

.

3

2

1

5

9

7
PMc. 74.

COU,B8TMfl) BMHOrpaAa;
nb1pes:i;

1-

BMAOM3MeHeHMfl (MeTaMopcj:>03bl) no6era:

TblH no6er KaKryca c peAyu,11posaHHblML'1 JlMCTbflMM;

7-

5-

3 -

o6bl4Hb1Ci YAflMH8HHb1Ci no6er;

IlYKOBMU,a nyKa;

4 -

38/leHblH 6e3IlMCTHblH cj:>OTOCMHTe3MPYIOW.MH no6er APOKa:

8-

cj:>MJl/lOKilaAMM MrllMU,bl;

2. HaiJ3e.M1tbte no6eiu:

KO/ll04Ka meAM4MM;

9-

2-

Ms:ic11c-

YCbl (BMAOL'13MeHeHHbl8

6-

KOpHeBMW,a

Kily6eHb Kaprocj:>ens:i

KomoqKJf (60.HphIIIIHHK H ,11;p.), ycHKH (BHHOrpa,11;

H ,11;p.), JiyKOBHD;hI (qecHOK H ,11;p.), YChI (3eMJI.HHHKa H ,11;p.), CTe6JieBhie
KJiy6HH (D;HKJiaMeH H ,11;p.), KoqaH (KarrycTa H ,11;p.) -

rHraHTCKa.H noqKa.

JIHCT

Jlucm -

::!TO H8,11;3eMHhIM, BereTaTHBHhIM opraH pacTeHH.H (pHC. 75).

06Jia,D;aeT ,D;BycTopoHHeii CHMMeTpHeii. PacTeT orpaHHqeHHoe BpeM.H 3a ctieT
OCHOB8HH.H JIHCTa.

<Pylllw,uu JlUCma:

cpoTOCHHTe3, TpaHCIIHpau;H.f! (HcrrapeHHe BO,ll;hI), ra3oo6-

MeH, 3anaca10ru;a.H <PYHKD;H.H; BereTaTHBHOe pa3MHO:>KeHHe.
OcHOBHhie qacTH JIHcTa:

ma, npuJtucmHUICU

JtucmoeaR nJtacmu111Ca, 'lepewoK, oc11oea11ue Jtuc-

(BhipOCThI OCHOBRHH.H JIHCTa).

"IJepeIIIKOB H npHJIHCTHHKOB MO:>KeT H He 6hITh. 0CHOB8HHe JIHCTa MOJKeT pacIIIHp.HThC.H H OXB8ThIB8Th CTe6eJih, IIPH ::!TOM o6paayeTC.f! BJiaraJIHru;e. JlHCThH
c qepemKaMH H83hIB8IOTC.f!

'lepeULIC06bl.MU,

6e3 qepeIIIKOB -

CUdJl'lU.MU.

"IJepem-

KH opHeHTHPYIOT JIHCTOBYIO nJiaCTHHKY no OTHOIDeHHIO K HCTOqHHKY CBeTa, co3,D;aBa.H JIHCTOByIO M038HKY -

pa3Meru;eHHe JIHCTheB Ha rro6ere, rrpH KOTOpOM

OHH He 3aTeHHIOT ,11;pyr ,11;pyra. JlHCTb.H pa3JIHqaroT no p.H,D;Y npH3H8KOB: no pa3MepaM (OT HeCKOJihKHX MHJIJIHMeTpOB ,11;0

20 M

-

y naJibM), no JKHJIKOBaHHIO,

no

cpopMe JIHCTOBOM nJiaCTHHKH, no KpaIO JIHCTOBOM nJiaCTHHKH, no pacnoJioJ:KeHHIO Ha CTe6Jie.

193

)ICuJ1,1'06anue Jl,UCma. <PopMa J1-ucma

JKHJIKOBaHHe JIHCTa
3TO rrpOBO;:J;HIIVie rryqKH JIHCTa. JKHJIKOBaHHe 6hrnaeT:

)/(UJl/CU -

,nyroBoe (JiaH;:J;bIIIJ H ;:i;pyrHe O;:J;HO;:J;OJihHhie);
rrapaJIJieJihHOe (JIHCT 3JiaI<a);
rraJihqaToe -

BCe )KHJIKH CXO,ll;HTCH B O;:J;HOM TOqKe (KJieH TaTapCKHM

H .n;pyrHe ;:J;BY;:J;O.lJbHhie);
rrepHCTOe -

Bhipameua u;euTpaJihHaH )KHJIKa (qepeMyxa H ;:i;pyrHe ,ll;BY;:J;OJihHhie);

BHJihqaToe -

;:J;HXOTOMHqecKoe (rHHKro ;:J;BYJIOIIaCTHhIM).
<l>opMa JIHCTa

Ilpocmou Jtucm -

COCTOHT H3 0,ll;HOM JIHCTOBOM IIJiaCTHHKH H o;:i;uoro qepeIIJ-

Ka. Ilpu JIHCTorra.n;e oTna;:i;aeT u;eJIHKOM.

Jtucm -

CJtOJICHbtU

COCTOHT H3 HeCKOJihKHX JIHCTOBhIX IlJiaCTHHOK co CBOH-

MH qepeIIIKaMH. 06ru;uii qepeIIIOK -

paxuc. Ilpu JIMCTona;:i;e Kam;:i;aH JIHCTOBaH

UJiaCTMHKa OTrra.n;aeT He3aBMCMMO.
IIpocThle JIMCThH .n;eJIHTCH ua (puc.

76):

a) JIMCThH c u;eJihHOM JIMCTOBOM nJiaCTMHKOM no cpopMe MOryT 6hITh OKpyrJihle, npo;:i;oJiroBaThle, JIMHeiiHhie u ;:i;p.;

a

6

4
4

e

PMc. 75.

npOCTble nL'1CTbfl:

'iepewKa; I, A -

a -

B11aranl'1Ll.\Hble;

'tepewKy np1-1111-1CTHl'1KaM1'1; 3 OCHoeaH1-1e YepewKa;

3-

YepeWKOBbllii;

e -

6 -

C1'1Af!Yl'1i1; B -

C nOAYW8YKOi1 B OCHOBaHl'11'1

co CB060AHblMl'1 np1'1111'1CTHl'1KaM1'1;

)I( -

c np1-1pacTa10ll.\1'1Ml'1 K

c na3yWHblMl'1 cpaCTalOL1.\1'1Ml'1Cfl npl'1111'1CTHl'1KaMl'1. 1 -

a11ara111-1Ll!e;

4-

np1-1111-1CTHHKH;

5-

yepewoK;

6-

n11aCT1'1HKa; 2 -

na3ywHafl noYKa

BomanuKa

194

Tpoii'laTo-

na11b4aTO-

(rpex-)

0:
.0
f()

:s:

c::
Q)

Ji
f()

0

a
c

PMC.

76.

npoCTble lll1CTbfl C paC4118H8HHOVi lll1CTOBOVi n11aCT11HKOVi 11 CJ10)1(Hbl8 lll1CTbfl

6) JIHCTbH c pactIJieHeHHOM JIHCTOBOM IlJiaCTMHKOM MoryT 6bITh:
JIOIIaCTHbie (TpOii'IaTo-, rraJiht:IaTO-, rrepMCTO-);
paa,n;eJihHhie (pact:IJietteHHhie, 6oJiee 1/ 2 nmpHHbI);
paccet:IeHHh!e (pact:IJieHeHHhie .n;o cpe.n;Heii iKMJIKH).
BhI,n;eJIHIOT TPH THrra pacnoJioiKeHHH JIHCTbes:
oqepe,n;Hoe, HJIM CIIHpaJibHOe - OT KaiK,n;oro yaJia OTXO,ll;HT TOJihKO O,D;MH
JIHCT (.n;y6, 6epeaa, 3JiaKH, 30HTM'IHbie M ,n;p.);
cyrrpoTHBHoe - s KaiK,n;oM yaJie rrapa JIHCTbeB (KJieH, cHpeHh, ry6ou;seTHhle H ,n;p.);
MyTost:IaToe - B KaiK,n;oM yaJie TPH H 6oJiee JIHCTa (oJieatt.n;p, aJio.n;eH,
1rnatt-t:Iaii H .n;p.).
Ilo attaTOMH'IeCKOMY CTpoeHHIO BhI,n;eJIHIOT:
1. Pop3o6enmpaJLbHbte Jtucmb.R - accHMHJIHD;HOHHaH qacTh JIHCTa (cToJI6t:IaTa.H XJIOpocpHJIJIOHOCHa.H rrapeHXMMa) pacrroJIOiKeHa TOJlbKO Ha sepxHeH CTO-

fenepamu6Hble opzaubi (u,.6em01c, nnoiJ, ceMR)

poHe JIHCTOBOH IIJiaCTllHKII, ry6'-IaTaH -

195
Ha Hl:DKHeH (caxapnaH CBeKJia).

AHaTOMHt.feCKoe cTpoeHne .n;opaoseHTpaJthHoro JIHCTa

JI01Cpo61W.H mKaHb -

snII.n;epMa IIJIII KOiKIIlla. Rommi;a noKphITa KY-

TMKyJioH: II MMeeT yCThMlla, KaK npaBMJIO, Ha HIIiKHeH IlOBepXHOCTH JlllCTa. 3amnmaeT JIMCT CBepxy H CHH3y. Pa6oTa ycThHll 06ecneq1rnaeT nocTyrnreHne yrJieKHCJioro raaa H Bhrne.n;eHHe KHC.'lopo.n;a, a TaKme TpaHcnnpa11mo (HCilapCHMe BO,ll;hl). fa3006MCH B TKaHHX JIHCTa npoTeKaCT
IlOCTOHHHO, Tor.n;a KaK CBeTOBaH cpaaa cpoTOCHHTeaa -

Me3o</JuJIJI (cpoTOCMHTe3IIPYIOI11aH TKaHb) -

TOJihKO ,Zl;HCM.

CTOJI6'IaThIM n ry6t.IaThIH.

R'.JICTKH CTOJI6t.faToro MeaocpHJIJia 06hI'IHO co.n;epmaT 60JihllIOe KOJIII'ICCTBO XJioponJiaCTOB. fy6t.IaTbIH MC30cpHJIJI HMCCT MHOro MCiKKJICTHMKOB.

Cocy8ucmo-60J101CHucmbte ny'lKU (mMJIKH) BKJIIO'IaIOT KCHJieMy (.n;pesec1rny), cpJI03MY (ny6) H MCXaHII'ICCKYIO TKaHb (yroJIKOBYIO KOJIJICHXHMY
na.n; ny'IKOM H CKJiepeRXHMY KOJihllOM BOKpyr nyt.IKa).

2.

I130JLamepaJibTtbte JlUCmb.H -

CTOJI6qaTaH napeHXIIMa c .n;syx CTOPOH JIH-

CTa (r.11a.n;nonyc).

3. PaiJuaJibHbte JLucmb.H -

xJiopeHXMMa BOKpyr 11eHTpaJihHOH iKMJIKM

(XBOH COCHhl).
JincTona.n; -

ceaOHHaH a.n;anTallMH pacTeHMH K M3MCHCHMHM KJIMMaTa. Oc-

HOBHaH npM'IMHa JIMCTOna.n;a oceHblO H 3MMOH.

Heo6XO,ll;MMOCTh YMCHbIIIeHIIH ncnapeHMH BO,Zl;hl

c IlOHMiKCHMeM

TeMnepaTyphl BCaChIBaHMe BO,ll;hl KOPHHMH

cHIImaeTcH, pacTeHne MomeT o6easo.n;MThCH n norn6nyTb. C6pachrnaHne JIMCTheB npe.n;oTBpamaeT STOT npo11ecc n yMeHbIIIaeT ncnapenne BO.D;hI.
Bn.n;onaMeHeHMH nncTa: KOJIIO'IKM (KaKTyc H .n;p.); ycnKM (ropox n .n;p.); JIMCThH HaCCKOMOH,ll;HhIX paCTCHHH (pOCHHKa H .n;p.).

reHepaTHBHbie OpraHbl (I.J;BeTOR, IIJIO)l;, CeMH)
IJ;BeTOK

z.r6emo/C -

3TO opraH ceMeHHOro pa3MHO.iK8HMH IlOKpbITOCCMeHHhIX pacTe-

HMH. IJ;seToK npe.n;cTaBJIHeT co6oft yKopot.IeHHhIH cnopoHoCHhIH no6er, npncnoco6.11eHHhIM ,D;JIH o6paaosaHMH cnop n raMeT. B 11seTKe nponcxo.n;nT onhIJieHMe,
OII.110,lJ;OTBOpCHMC, pa3BMTMe aapO,Zl;hIIIIa, o6pa30BaHMe TIJIO.n;a c CCMCHCM.
CTpoeHHe u;senta (pl:lc.

77)

Il;BeTOK COCTOMT H3 cme6Jle60U 'lacmu (llBCTOHOmKa, llB8TOJIOiKC) H JLucmo60U 'tacmu ('IaUICJIMCTMKM, JierreCTKM, Thl'IMHKH, IICCTHKM).
I.{6emoJIOJ/Ce - paCUIMpCHHaH qacTb llB8TOHO.iKKM, K Heii npMKpenJIHIOTCH
Bee 'IaCTM llBCTKa.

l/ aute1t1Ca (Ca -

calyx) -

cocTOMT M3 t.IaUie.11ncTMKOB ( o6hI'IHO ae.11eHoro

D;BeTa). qaUICJIMCTMKM MoryT 6hITh CB06o.n;HhIMM, TOr.n;a qarnet.IKa Ha3bIBaCTCR

paaiJeJLbHOJiucmnou. EcJIM qarneJIMCTMKM cpacTaIOTCH, TO qarueqKa Ha3hIBaeTca cnaunoJLucmnou.

BomanuKa

196

Beu'luK (Co 7

6
8
9
.'3

9

JO
--2
-11
15

1.'3

HOCTh

corolla) -

JieneCTKOB.

coBoKyrr-

JienecTKM

MOryT

6hJTh CB060,ll;Hh1Mll IIJIM CpOClllMMIICR,
a seHqIIK, COOTBeTCTBeHIIO, pa38eJlbHOJtenecmHbtU IIJIM cnauttoJtenecmHbtil.
BettqIIK o6hlqtto RpKo oKpallleH, IIMeeT
pa3ttyro cpopMy,

npIIBJieKaeT ttaceiw-

MhIX-OIThIJIIITeJiett.

0KOJlOU,6emHUK, o6pa30BaHHhIH qa860UHbt.M. Ilpocmou OKOJlOU,6emHUI( MOJKeT
6b1Th 8eH'lUK06U8Hbl.M (TIOJihIIaH), T. e.
llleqKott II BeHqMKOM, Ha3bIBaIOT

COCTOHTh TOJihKO II3 JIIICTQqKOB BeHqII-

PMc. 77. CxeMa CTpoeH111si useTKa: 1 PblJlbL(e; 2- 3aBSl3b; 3 - CTOJ16111K; 4 - C8MSl3a<iaTOK; 5 - Tbl'lLllHO'lHaSl HLllTb; 6 - CBSl3HLllK;
7-

nbtllbHLllK;

8 -

nbtJ1bL(8Bbte 3epHa;
we11111crnK;
Ka;

nbtJlbHLllK B pa3pe3e;

9 -

10 - 11enecrnK; 11 - o.ia-

12- L(BeTOJlOlKe; 13- L(B8TOHOlK-

14- npLllL(B8THLllK; 15 -

npLllL(B8THLll'l8K

Ka, IIJIII

'lGUle'lK06UOHbl.M (KpaIIIIBa).

ll,BeTKII 6e3 OKOJIOD;BeTHIIKa Ha3hIBaIOT
rOJihlMM (IIBa).

I'a.Memo<Pum -

rarrJiott,n;ttoe rroJiosoe

IIOKOJieHI:Ie, rrpo,n;yu;HpyIOID;ee raMeThI.

Cnopo</Jum -

,n;ttrrJioII,n;ttoe rroKoJie-

HHe, KOTopoe IIPOH3BO,ll;HT rarrJIOM,ll;Hbie
crrophl (MHKpo- H Meracrropb1) y nseTKoBhIX pacTeHHH.

Tbt'lUHKa COCTOHT H3 mbl'lUIW't/WU
ttumu H nbtJlbHUKa. CosoKyIIHOCTh ThIqHHOK Ha3hIBaIOT aniJpou,ee.M (A).

10

IlbIJlhHHK o6pa30BaH ,D;BYMH IIOJIOBHHKa-

6

MII, coe,n;tttteHHhlMH CBH3HHKOM.

8

,n;oii: IIOJIOBHHKe IIhIJihHHKa

7

IIhIJihIJeBhlX rtte3,n;a -

B

Kam-

eCTh ,n;Ba

.MUKpocnopattzua.
llbl.Jf,bU,e-

rtte3,n;a IIhIJihHHKOB Ha3bIBaIOT

6bl.MU .Mew1ca.Mu. B HHX o6pa3yroTCH MHKpocrrophl II co3peBaeT IIhIJihIJa.

II btJlbU,a -

aTo pe,n;yu;ttposaHHhltt

MYJKCKOH raMeTOcpIIT'

PMc. 78.

CTpoeH111e ceMA3a<iaTKa 111 aapo-

Abtwesoro MeWKa (lKeHCKOro raMeTOCj>LllTa) noKpbnoceMeHHbtX pacTeH111iii: 1 -

no-

KPOBbt ceMsiaa<iaTKa, LllJlLll LllHTeryMeHTbl (a Hap~Hbtiii,

6-

BHyrpeHHLlliii);

2- ML11Kpon111-

11e; 3 - xanaaa; 4 - Q>yH111Ky11yc; 5 - HyL(e1111yc;

6 -

KTleTKa,

3BPOAblW8Bbliii MewoK;

8 -

CLllHepfLllAbl,

9 -

7-

SliiiL(e-

BHTL!lnOAbl;

10- srnp111o.iHoe SlApo; 11 - n11aL(eHTa; 12nposOASll.l.!Llliii ny<ioK

COCTOHID;HH H3

,n;syx KJieTOK.
06e KJieTKH MyiKcKoro raMeTocpttTa
( IIblJlhlJhI) BhIIIOJIHHJOT orrpe,n;eJieHHYIO

eezemamuettaa, 06pa3yeT IIhIJihIJeByIO Tpy6Ky. MaJieHhKaa KJieTKa, HJIH zettepamuetta.R, o6pa3yeT ,n;sa crrepMHH -

cpyHKIJHIO. BoJihlllaH KJieTKa, HJIH

My»CCKHe raMeThI.

IIecmwc MOiKeT 6hITh o,n;IIH HJIH aeCKOJihKO. CosoKyrraoCTh rrecTHKOB aa-

197

iuneu,eeM (G ). IleCTHK o6pa30BaH IlJIO,ll;OJIHCTHKaMH. IleCTHK COCTOHT
aB.H3U,
cmoJL6uKa w pbtJlbU,a. Ha pbIJiblle rrecTttKa rrorra.n;aeT IIbIJihIIa, w rrpo113 3
3bIBaIOT

11cxo,n;MT rrpopacTaHtte. EcJIH IIJIO,n;OJIHCTMK o6paayeT e,n;11HCTBeHHhitt rrecn:IK,

MOHOKapnllbtM. ECJIH ,n;Ba H 6oJiee IIJIO,ll;OJIHCTHKa o6pa3yIOT
STO anoKapnHbtU iuneu,eu. Ilptt cpacTaHHH HeCKOJlbKHX IIJIO,ll;OJIHCTHKOB B O,IJ;HH rreCTHK 06pa3yeTC.H u,eHOKapnHblU iuneu,eu.
Ce.Mano'lKa (pnc. 78) Haxo,n;ttTC.H B 3aB.H3H rrecTHKa. IlpnKperrJI.HeTc.H
TO OH Ha3bJBaeTC.H

caMOCTO.HTeJibHhJe rreCTHKH -

K cTeHKe 3aB.H3M c noMOIIJ;bJO IIJia11eHTbI Ka COCTOHT H3
H

B3.ZJ;YTH.H TKaHell: 3aB.H3H. CeM.HIIOq-

ce.M.RHOJICKU ( rpynuKyJLyca ), noKp0606 ( 06a UHmeiy.Me1tma)

aopa CeM.HIIOqKH, B KOTOpOM <l>OpMHpyeTC.H 3ap0,1J;bIIIIeBbIH MeIIIOK.
3apOObtUle6btU .MeUlOK (COCTOHT H3 BOCbMl:I KJieTOK) - 3TO pe.n;y11npoBaH-

Hblll .meHCKHH raMeTO<l>HT, KOTOpbIH pa3BHBaeTC.H H3 Meracrropbl. IloKpOBbl

nbtJlbU,eBXOO ( MUKponu.lle). BJin.me K IIbIJiblleBxo.n;y Haxo,n;.HTC.H Hll:11eKJieTKa H .n;Be KJieTKH - cunepiuobt ( n). Ha rrpoTHBorroJio.mHoM KOHI1e - TPH KJieTKH - anmunoiJbt
(n}, B 11eHTpe - ou_nJlOUOHOe BmOpU'lHOe .HOpO ( 2n }, KOTOpoe o6pa30BaJIOCb

Ha BepxyIIIKe CeM.HIIOqKH He cpacTaJOTC.H H o6pa3yJOT

H3 CJIH.HHH.H ,n;Byx IIOJI.HpHbIX .H,n;ep.

o6oenOJlbte. Il;BeTKH TOJibKO c TbloononOJlbte (pa3,n;eJibHOIIOJible), HaIIpH-

Il;BeTKl:I c TbiqHHKaMH H rreCTHKaMH qHHKaMH HJIH TOJibKO c rrecnrnaMH Mep KyKypy3a, 6epe3a H .n;p.

KJiacc11<l>HKa11m1 pacTenHii
PacTeHH.H, y KOTOpbIX MymcKne (Tb1qnHOqHb1e) H .meHCKHe (rrecTnqHbie)
IJ;BeTKH Haxo,n;.HTC.H Ha o,n;HoM pacTeHHH, Ha3hlBaJOTC.H

oonoooMHbt.Mu (KYKYPY-

aa). PacTeHH.H, y KOTOpbIX o,n;HOIIOJibJe IIBeTKH Haxo,n;.HTC.H Ha pa3HbIX pacTeHHsx, Ha3bIBaJOTC.H

06YOO.Mllbl.MU (KparrHBa, HBa H .n;p.).
Co11seTHJ1

Cou,Bemua -

aTo rro6er HJIH cncTeMa rro6eroB, HecyIIJ;HX IIBeTKH.

CTBHH c THIIaMH BeTBJieHH.H rro6eroB co11BeTH.H (pHC.

79)

B cooTBeT-

paa,n;eJI.HJOT Ha:

1) Mononooua.llbHbte ( 6ompuouiJHbte) cou,Bemua 6hlBaJOT

rrpocThle H cJiom-

IIhle. BoKOBbJe ocn rrpocTbIX co11BeTHtt He BeTB.HTC.H, CJIOJKHbIX -

BeTB.HTC.H.

IlpOCTLie MOHOIIO,ll;HaJILHLie co11seTHJI. PocrrycK IIBeTOB rrpoHCXO,!J;HT CHH3Y
BBepx, ec.rrn co11BeTHe BepTHKaJibHOe (KHCTb, KOJIOC), JIH6o OT rrepn<PepHH
K :u;eHTpy, ecJIH IIBeTKH B COIIBeTHH pacrroJio.meHbl ropH30HTaJibHO (30HTHK,
KOp3I:IHKa):
KHCTb -

BCe IIBeTirn Ha y,n;JIHHeHHOH OCH, HMeJOT IIBeTOHOJKKH (qepeMyxa);

Ill;HTOK -

cxo,n;eH c KHCTbJO, HO IIBeTOHOJKKH pa3HOH ,IJ;JIHHbl H IIBeTKH

pacrrOJIOJKeHhl B 0,IJ;HOH IIJIOCKOCTH (rpyIIIa);
30HTHK -

IIBeTKH Ha IIBeTOHOJKKaX 0,IJ;HHaKOBOH ,IJ;JIHHhl, BbIXO,!J;.HT H3 o.n;-

HOH TQqKH (BHIIIH.H);
KOJIOC -

IlpOH3BO,ll;HOe KHCTH, HO 6e3 IIBeTOHOJKeK, IIBeTKH CH,ll;.Hqne (no-

.n;opomHHK);
noqaTOK -

KOJIOC c TOJICTOH OCbIO (KyKypy3a);

198

6omanuKa

rOJIOBKa -

BH,Zl;OH3MeHeHHbIM 30UTMK; OCb pa3pacTaeTCJI, Ha Heii CM,Zl;H'!He

IJ,BeTKH (KJICBep);
KOP3HHKa -

CH,ll;Jl'!He IlBeTKH Ha pacurnpeHHOM KOHUe OCH cou,aenrn

( CJIOJKHOIJ,BeTHbie).
CJimKHLie MOHono,11;HaJILHL1e cou.BeTH.JI:

MeTeJIKa (CJIOarnaH KHCTb) CJIOJKHhIH KOJIOC -

o6pa3oBaHa H3 OT,ri;e.TCbHhlX KHCTeii (cttpeHb);

o6pa30B8H H3 OT,ri;eJihHbIX IlpOCTbIX KOJIOCKOB (muemm;a);

CJIOJKHhlll 30HTHK -

COCTOHT H3 IlpOCThIX 30HTHKOB OT Bepxyunrn rJiaBHOH
OCH (MOpKOBh);
CJIOJKHhIH ILl,HTOK

-

COCT8BJieH

H3

npOCThIX ILl,HTKOB (mrmMa);

2)

CUMnoauaJtbHblf' (U.UMOJHbte} co-

U,BemuR. I'JiaBHaJl OCb 38KaHqHaaeTCR:
I.J,BeTKOM. Iloc;re,11;y101I.1,He IJ,BeTKH B03HHKaIOT B na3yxax npHll,BeTHbIX JIHCTb-

•

eB H np0,!1;0JIJK8IOT POCT rJiaBHOH OCH •
PocnycK ll,BeTKOB npo11cxo,n;HT caepxy

4

BHH3 mrn H3 I.J,eHTpa K nepmpepHH.
Pa3JIH'laIOT TPH THIIa CHMno.n;HaJih-

a

8

9
6

\

HhIX COll,BeTHH:

~

a) MOHOX83HH (H3BHJIHHa H 3aBHTOK) -

TpeThero rrop.JI.l(Ka H T. ,ZI;., TO c O.!l;HOH,
TO c .l(pyroii CTOpOHbI (JIIOTHK e,n;KHH);
38BHTOK ro -

T1>1nb1

co4sernj;j: a -

HOM

ll\lllTOK;

6-

3 -

30HTlllK;

noYaTOK;

7-

4 -

6-

5 -

2 -

11 -

3111j;j);

14-

IIO,ZI; ll,BeTKOM rJiaBHOH

KOJlOC;

meHHbie BeTBH (OCH), 3aKaH'lHB8IOILl,HeCJI ll,BeTKOM. Kam,n;aH H3 3THX oceii TaK-

1113BlllJllllH8 {MOHOX83111M};

13 -

pa3BHJIHHa -

OCH o6pa3yIOTCH cynpOTHBHO pacrrOJIO-

CJlO>t<Hble 6oT-

8 - CJlOJKHblM KOJlOC; 9 - M8T8JlKa;
10- CllO>t<HblM 30HTlllK; B - l_\lllM03Hbl8 (ClllMnO-

3aBl1TOK (MOHOxaa111j;j) ;

JIOJKHO,!(HXOTOMH'leCKOe

pacrroJiomeuue oce:H:

PlllOWJ,Hble:

Al18JlbHb18):

38KPY'lHB8HCb

6) ,Zl;HX83HH (pa3BHJIHHa H ,Zl;BOHHOH
38BHTOK) -

KlllCTb;

rDJlOBKa;

KOP3111HKa;

HanpaBJieHHH,

B 38BHTOK (ue3a6y,Z1;Ka);

npocrb1e 6orp111-

I -

ocb TpeThero nopH,n;Ka H T • .n;.,

HO IlpH 3TOM OCH OTXO,Zl;HT TOJlhKO B 0,Zl;-

(J

PMc. 79.

OT rJiaBHOH OCH OTXO-

,Zl;HT OCh BTOporo rropH,n;Ka, OT BTopo-

~ Y,

0WJ.Hble (MOHOnOAlllMbHb1e):

OT rJiaBHOH OCH OTXO-

.l(HT OCh BTOporo IIOPH,ri;Ka, OT BTOporo -

~12

11

O'lepe,ri;Hoe pacnoJiomeHHe oceH::

H3BHJIHHa -

12-

pa3BlllJ111Ha (A111Xa-

MHoro11Y4eB11K (n11ej;joxaa111M)

JKe

o6pa3yeT .n;se cynponrnHhIX

ocH

(rB03.D;HKa);
.!l;BOHHOH 3aBHTOK -

ITO,ZI; IJ,BeTKOM

rJiaBHOH OCH cynpOTHBHO o6pa3yIOTCH
,Zl;Ba rrpoCTbIX 38BHTKa (KapTo¢leJib);

On""nenue. OnnoiJom6openue

B) nJieii:oxa3tt:ll -

199

MyToBqaToe pacnoJiometttte oce:ll -

no.n: IJ;BeTKOM rJiaB-

aoil: OCH pa3BIIBaeTCH MYTOBKa 60KOBbIX oceii: c IJ;BeTKaMH IIJIII COI1BeTHHMH
(l\WJIOqaii:).
OnbIJieHHe

OnbtJlettue -

aTo npoIIecc nepettoca TihIJihIJ;hI II3 nhIJihHIIKOB Ha phIJihIIe ne-

CTHKa. Pa3JIIIqaIOT caMOOIIbIJieHHe, nepeKpeCTHOe OilhJJieHtte, IICKYCCTBeHHOe
OilbIJJeHtte.

Ca.MoonbtJleHue MOJKeT npowcxo,n:HTh y pacTeHHH, HMeIOIIIHX ,n:ByTIOJihie
I1BeTKH. IlhIJihIJ;a nepeHOCHTCH Ha pbIJihIIe necTHKa BHYTPH o,n:ttoro IJ.BeTKa. CaMOOilhIJIHIDIIIHec.H pacTeHHH -

ropox, .fiqMeHb, pOJKh, OBeC II ,n:p.

Jlepe1Cpecmttoe onbtJlettue -

Ha phIJihIIe nona,n:aeT TihIJihIIa c ,n:pyrwx pacTe-

HHH Toro me BII,n:a.
IlepeKpeCTHOe OilbIJieHHe OCyIIIeCTBJIHeTCH no.n: B03,n:eil:CTBHeM pa3JIHqHbIX
BHeIIIHHX q:>aKTOpOB:
aaeMocpttJitt.a -

onhrnem1e c noMOIIIhID BeTpa. PacTeHH.H paao 3au;BeTa-

IDT, .n:o TIO.HBJieHII.H JIHCTbeB. ll,BeTKH MeJIKHe, HeB3paqab1e. IlbIJibIJ;a
MeJIKa.a, cyxaH. Phrnhu;a necTHKOB qacTo nepttCThie. 3To MHOrIIe 3JiaKH,
OCOKa, 6epe3a, ,n:y6 H ,n:p.;
3HTOMOcpHJIHH -

OTibIJieHHe c TIOMOIIIbID HaCeKOMhIX (nqeJI, IIIMeJieH, oc,

6a6oqeK). l.I;BeTKII KpynHhie, .HpKO OKpaIIIeHhI, eCTb HeKTap H 3anax.
PacTeHII.H, OilbIJI.HeMhie HaceKOMhIMH, -

IIIIITIOBHHK, H6JIOHH, qepeMy-

xa, BHIIIH$I, MaK, poMaUIKa;
rw,n:pocpttJIH.H -

OTihIJieHtte c noMOIIIhID BO,ll;hI. XapaKTepao ,ll;JI.H MHorwx

BO,ll;HbIX UBeTKOBbIX pacTeHHH.

HcJCyccmeeHHoe onbtJLettue -

aTo npway,n:wTeJibHoe onhrJieawe, npott3Bo,n:w-

Moe qeJIOBeKOM c TIOMOIIIbID co6CTBeHHOH IlhIJibUhI u;BeTKa HJIH IlhJJlhIIhl ,n:pyI'HX pacTeHHH.
Omrn,n:oTBOpeHHe

OnJLoiJomeopettue -

aTo npou;ecc CJIHHHHa meacKoil: H MymcKoli noJIOBhIX

KJieTOK c o6pa30BaHHeM 3HrOThl (o,n:HOKJieToqHoro 3apO,ll;hIIIIa).

y

IIBeTKOBbIX pacTeHHH STOT TIPOIIeCC Ha3hIBaeTC.H iJ60UHOe onJLoiJomeope-

nue. Oa 6b1JI OTKphIT B

1898

r. PYCCKHM 6oTaHHKOM

c. r. HaBaIIIHHbIM.

IlhIJihIJ.eBoe 3epao nona,n:aeT aa phIJih1Ie necTHKa. BeiemamueHaR JC.TLemJCa
npopacTaeT B BH,D;e Tpy6KH qepe3 TKaHII pbIJihIIa H CTOJI6IIKa no aanpaBJieHHIO K 3aB.H3II. Tpy6Ka npoxo,n:HT qepe3 MIIKponIIJie (rrhIJihIIeBxo.n:) w oKa3hIaaeTc.a BHYTPII 3apo,n:hnueBoro MeIIIKa. I'ettepamueHaR JCJLemJCa 06pa3yeT
.n:aa crrepMIIH. 0Htt ,n:BII?KyTc.a B IIhIJihUeBoli Tpy6Ke Ha ee KOHIJ.e. Ilh1Jih1IeBaa
Tpy6Ka pa3pbJBaeTCH, crrepMMH OKa3b1BaIOTC.H B 3apO,ll;hJIIIeBOM MeIIIKe. 0,n:HH
cnepMHH CJIIIBaeTC.H c .HHIJ.eKJieTKOH, II 06pa3yeTC.H ;wioma -

rrepBaH KJieT-

Ka 3apo)J;hIIIIa ceMeHH. BTopoil: crrepMIIH CJIHBaeTCH c ,n:HnJIOII.ll:HhIM H.n:poM
IJ.eaTpaJihHOH KJieTKH, 06pa3yeTc.a mpunJLouiJHaR JCJLemJCa. TpHIIJIOH,D;Ha.H
KJieTKa .n:aeT HaqaJio 3HiJocnep.My (nHTaTeJibHOH TKaHH ceMeHII).

BomanuKa

200

IIocJie onJio.a;oTBope1uu1 H3 ceMJrnaqaTKa 06pa3yeTCH ce,11.fl. CeMH nMeeT ceMeHtty10 1COJ1Cypy, aapoiJbtUJ, :Janae numameJtbllbtX Bew,ecmB. 0.nttoBpeMeHHo
pa3pacTaeTC.H 3aBH3b neCTHKa. lfa CTeHOK 3aBH3H 06pa3yeTCH OICOJLOnJIOOllUIC,
oKpymaIOID;Hlr ceMH. IlpOHCXO,ll;HT o6pa30BaHHe ITJIO,Lla.

flJIO.ZJ;

flJtoiJ -

opraH pa3MHOJKeHHH IlOKpbITOCeMeHHblX pacTeHHH. IlJIO,ll; co.nep-

JKHT ceMeHa 11pa3BHBaeTCH H3 O,LIHoro u;BeTKa (pHc.

80).

<l>yHKIJ;HH TIJIO,n;a: cpopMHpOBaHn:e, 3aID;HTa, pacnpocTpaHeHHe ceMHH.
CTeHKa nJio.a;a Ha3hIBaeTca nepuKapnue.M ( 0KoJtonJtoiJ1tuKoM) H HMeeT TPH
cJioH:3K3oKapnuu,Me301Capnuu,a1tiJ01Capnuu.
B COOTBeTCTBHH c THITOM rMHen;eH Bee ITJIO,Llbl .neJIHT Ha MOllOICapnuu, anoKapnuu, u,e1t0Kapnuu H nceBOOKapnuu.
flJLOObl MOHOICapnuu pa3BHBaIOTCH H3 u;BeTKOB, MMerouvrx MOHOKapnHhIH
rHtteu;eif:
a) O,ll;HOJIMCTOBKa -

MHOroceMeHHOH ITJIO,D;, BCKphIBaIOID;HHCH TOJibKO no

6pIOllIHOMY llIBy;
6) o,n;HoKocrnHKa -

o.nHoceMeHHoif, HeBcKpb1BaIOID;nifca ITJIO,LI. Me3oKap-

nnif O,ll;HOKOCTHHKH MOJKeT 6bITb coqHbIM (BHllIHH, CJIMBa H ,n;p.) HJIH cyXHM (M11H,1l;aJih H ,Lip.);
B) 606 -

O)IHO- HJIH MHOroceMeHHOH rnIO,n;, pacKphrnaeTCH no ,D;BYM JlIBaM

(cnnHHOMY n 6pIOllIHOMy). CeMeHa, pacnoJiomeHHbie Ha CTBOpKax (.z:IOHHHK, apaxn:c, aKaIJ;HH H ,n;p.).

flJtOObt anoKapnuu pa3BnBaIOTCH n3 anoKapnHoro rnHeu;eR:
a) MHOrOJIHCTOBKa -

MHoroceMeHHOH c6op-

HblH ITJIO,D;, COCTOHT H3 IlpOCTblX JIHCTOBOK
(KYilaJihHHIJ;a, KaJiyJKHMD;a H ,n;p.);
6) MHoroopellleK -

c6opHbIH MHoroceMeH-

HOH ITJIO,D;, COCTOHID;HH 113 OT,D;eJihHblX O,LIHOceMeHHblX HeBCKpbIBaIOID;HXCH IIJIO,ll;HKOB
-

2

opellIKOB (JIIOTHKH, JianqaTKa H ,n;p.);

B) MHOrOKOCT.HHKa -

c6opHblH ITJIO,ll;, COCTO-

HID;HH H3 ,n;ByX HJIH MHOrnx ITJIO,ll;HKOB -

3

KOCT.HHOK. Me30Kapn11ii Kam,n;oro nJio.nnKa coqHbIH, 3H,ll;OKap1rn:if -

TBep,n;blif (Ma-

JIHHa, emeBMKa H ,n;p.).

flJLOObl u,eliOKapnuu pa3BlfBaIOTCH H3 u;eHOKapnHoro rnHeu;eH. 3aBH3h o,n;Ha, HO o6paPMc.80.CTpOeHV1ennoAa(oAHOKO-

30BaHa ,D;BYMH H 6oJiee cpOCJlIHMHCH ITJIO,D;O-

CHIHKVI) nepcviKa o6blKHOBeHHOro:

JIHCTMKaMM:

1-3 -

a) Hro,n;a -

oKononnoAH~K. vinvi

nviiil ( 1 -

3K30Kapn~~.

2 -

nepviKapMe3oKap-

nvii1, 3 - 3HAOKapn~~); 4 - CeMft; 5 cneA nnoAOHQ)l(KVI;

6 - nnOAOHOJKKa

MHOroceMeHHOH HeBcKpbrnaro-

ID;Mifca ITJIO,ll; c
KOM -

coqHbIM OKOJIOITJIO,D;HH-

nepMKaprrn:eM (noMn,n;op, KapTo-

cpeJih, KPhliKOBHnK 11 ,n;p.);

201

flJ1,oiJ. CeMH

6) TbIKBHHa -

IIJIO,ZJ;, 6JIH3KHH K Hro.n;e, HO OTJIHqaroru;HHCH KOIBHCThIM 3K-

30KaprrtteM (ThIKBa, ap6y3, orypeu; H .n;p~);
B) Kopo6oqKa -

MHOroceMeHHOH IIJIO,ZJ;, HMeeT cyxo:i1: BCKphIBaIOru;HHCH

OKOJIOIIJIO,ZJ;HHK (MaK, .n;ypMaH, HarrepcTHHKa H .n;p.);
r) cTpyqoK -

o6pa3oBaH .ZJ;BYMH cpocIIIHMHCH IIJIO.ZJ;OJIHCTHKaMH, BHYTPH

HaXO,ZJ;HTCH rrpo,n;OJihHaH rreperopo,n;Ka, K KOTOpoii rrpHKpeIIJIHIOTCH CeMeHa (KarrycTa, ropqttu;a H ,n;p.). CTpyqKH KJiaccmpttu;ttpy10T Ha BCKph1Ba10ru;ttecH H HeBCKpbIBaIOru;HecH;

.n;)

CTpyqoqeK -

CTpyqOK, y KOTOporo ,!l;JIHHa IIOqTH paBHa IIIHPHHe (rracTy-

IIlhH CYMKa H ,n;p.);
e) H6JIOKO -

MHoroceMeHHOH HeBCKpbIBaIOIIJ;HHCH IIJIO,n;, 3aBH3h KOToporo

o6pa3yeTCH cpacTaHHeM OT Tpex

,n;o

IIHTH IIJIO,!l;OJIHCTHKOB (H6JIOHH, PH-

6ttHa, 6oHphllllHHK H ,n;p.).

ll;wabt nceBa01capnuu -

3aKJia,!l;hIBaeTCH ,n;Ba H 6oJiee IIJIO,ll;OJIHCTHKOB,

a pa3BHBaeTCH TOJihKO O,!l;HH, OCTaJihHhJe pe,n;yu;ttpyIOTCH:
a) ceMHHKa

O,!l;HOCeMeHHOH

IIJI0,!1;

c

KO:IKHCThIM

rrepttKaprrHeM,

He cpacTaIOIIJ;HMCH c ceMeHeM (rro,n;COJIHeqHHK, BaCHJieK, poMaIIIKa H .n;p.);
6) aepHOBKa -

O,ll;HOCeMeHHOH IIJI0,!1; c IIJieHqaThIM rrepHKaprrHeM, cpacTalO-

ru;ttMCH c ceMeHeM. XapaKTepeH ,n;JIH 3JiaKOB (rrmeHttu;a, pmKh, THMo<}JeeBKa H ,n;p.);
B) opex -

O,!l;HOCeMeHHOH, pe,n;Ko ,n;ByCeMeHHOH IIJIO,!I; c ,n;epeBHHHCThIM rre-

pttKaprrtteM (Jieru;ttHa, OJihxa, 6epeaa H ,n;p.).

Con;waua -

aTo coBoKyrrHocTh apeJihIX rrJio,n;oB, B03HHKa10ru;ttx IIYTeM cpacTa-

HHH 3aBH3eH u;BeTKOB B rrpe,n;eJiax o,n;Horo cou;BeTHH (aHrurnc, Hmirnp, IIIeJIKOBHu;a).
IIJio,n;hl, o6pa3oBaHHble H3 e,n;HHCTBeHHoro necTHKa B u;BeTKe,
BaIOT

nacmoaw,uMu.

EcJIH B

<}JopMttpoBaHHH rrJio,n;a

Ha3hl-

yqacTBYIOT .n;pyrtte

'laCTH u;BeTKa (u;BeTOJIO:iKe, OKOJIOll;BeTHHK, ThJqHHKH), 3TOT IIJIO,ZJ; cqHTaIOT
AO:JICltbtM. IIJio,n;, c<}JopMttpoBaBIIIHHCH H3 HeCKOJihKHX rrecTHKOB o,n;Horo u;BeT-

Ka, Ha3bIBaeTCH cJtOJICltbt.M. IIJio,n;hI 11 ceMeHa pacrrpocTpaHHIOTCH c rroMOIIJ;hIO:
BeTpa -

MeJIKHe, JierKHe ceMeHa, HMeIOru;He KphIJiaTKH, XOXOJIKH (ce-

Meii:cTBO OpxH,n;Hhle);
BO,!l;bl -

ceMeHa BO)l;HhIX pacTeHHH (KyBIIIHHKa, Ky6hlmKa H ,n;p.);

:lKHBOTHhlX IITHll; -

HMeIOru;He KpJOqKH, IIIHIIHKH (qepe.n;a, perreHHHK H .n;p.);

HMeIOru;He coqHhie OKOJIOIIJIO)l;HHKH (pH6HHa, KaJIHHa, BHIIIHH H ,n;p.);

IIyTeM caMOIIPOH3BOJihHOrO pa36paCh!BaHHH ceMHH (6emeHhlH orypeu;,
He,n;oTpora H ,n;p.);
qeJIOBe:KOM -

B X03HHCTBeHHhIX u;eJIHX HJIH HerrpOH3BOJihHO.

CeMa
CeMa -

opraH rroJioBoro pa3MHO:>KeHHH H pacceJieHHH. Pa3BHBaeTcH H3 orr-

Jlo.n;oTBopeHHoro ceMH3aqaTKa.
CeMH cocToHT H3

3apoabtut -

B

aapoabtUta, anaocnepMa, ceMennou 1wJ1Cypbt.

OCHOBHaH qacTh ceMeHH (MHHHaTIOpHbIH crropo<}JHT -

:e:eM pa3JIHqaroT

mpu aapoabtuteBbtX opzana:

2n).

3apo,n;h1II1eBhlH KopernoK;

Bomanu1w

202

3apO.D;bIUieBhIH CTe6eJreK c 38pO.D;bIUieBOH rroqKoii; 3apO.D;bIIIIeBhie JIHCTb.H M.H.D;OJIH (,n;Be -

y .D;BY.D;OJlbHbIX, o;:i;na -

ce-

y O.D;HO.D;OJibHbIX).

°YqacTOK CTe6eJibKa B 3apO.D;hIUie BbIUie ceM.H.D;OJie:H H83bIBaeTC.H anu1wmUJlb
(Ha,n;ceM.H.D;OJihHOe KOJieHo), Httme ceM.H.D;OJreil: -- zuno1wmu.1lb (rro.n;ceM.H.D;OJihttoe KOJieHO).

Ce.Me1t1la.fl KOJICypa MHOrOCJl:OHHa.H, 3aw;11w;aeT 3apO.D;hIIII OT BbICbIX8HI1.H
tt npem.D;eBpeMeHHoro rrpopacTaHtt.H.

Ce.Mella iJeyiJoJlblibtX 6hIB8IOT (pttc.

81 ):

6e3 3H.D;OcnepMa (CeMeHa ¢aCOJII1, 6o60BbIX, ThIKBeHHbIX, KpecT01IBeTHhIX, CJIOJKHOIIBeTHhIX tt ;:i;p.). BOJihIIIa.H qacTh rrttTaTeJihHhIX Bew;ecTs
38KJIJOqeHa B ceMH.D;OJl.HX;
c 3H.D;OCIIepMOM (ceMeHa 30HTI1qHbIX, rracJieHOBhIX, TOMaTa, ¢ttaJIKI1,
JIIOTttKa H .n;p.). 8H,n;ocnepM OKpymaeT ceMH.D;OJIH. Mem.n;y ceMa,n;oJI.HMtt
KOHyc ttapacTaHtt.H, rroqeqKa ew;e He c¢opMttpoBaHa;
c nepHcrrepMoM (ceMeHa rB03.ZJ;J1qHhrx 11 .n;p.). Ilepucnep.M -

3arracaIOIJI8.H

TKaHb, o6pa30B8HHa.H TKaHbIO 1tyU,e.1l.1lyca B o6JiaCTI1 IIOKpoBOB ceM.H3aqaTKa.

Ce.Mella oiJllOOO.!lbHbtX 6brna10T:
6e3 3H,D;ocnepMa ( ceMeHa qacTyxtt 6oJIOTHOH, p.n;ecTa 11 .n;p. ). CeMa ttMeeT <I:>opMy IlO.D;IWBbI, 3H.D;OCnepM yme «C'be,n;ett»;
c 3H.n;ocnepMOM (60JihlllI1HCTBO ceM.HH O.D;HO.D;OJibHbIX, HanpHMep ttpttc
H .n;p.). BoJihrnaa qacTh ceMeHH -

3H,n;ocnepM, B Hero norpymeH naJioq-

KOBH.D;HhIH 3apo.D;hIIII. CeM.H.D;OJI.H IIHJIMHJJ;pHqecKaa, ee HHiKH.H» qacTh
(BJiaraJrnw;e) OXB8ThIBaeT KOHYC HapacTaHH.H;
ceMeHa 3JI8KOB. 3apO.D;bIIII B 3epHOBKe conpHKacaeTC.H c 3H.D;OCnepMOM
o;:i;noH: cToponoii -

ceM.H.D;OJieii HJIH LQHTKOM, npHmaThIM K an;:i;ocnep-

MY. Ill,HTOK ( CeM.H.D;OJI.H) BhIIIOJIHHeT cpyHKIIHIO BC8ChIB8HH.H IIHTaTeJihHbIX Beru;eCTB M3 3H,D;OcnepMa.
8H,n;ocnepM co;:i;epiKHT opraHHqecKHe H IlHT8TeJibHbie MHHepaJibHbie Bew;ecTBa. OpranHqecKHe Bew;ecTBa: 6eJioK (KJieiiKOBHHa), yrJieBO.D;hI (KpaxMaJI), iKHphI
(paCTHTeJihHbie M8CJia). MHHepaJibHhie Bew;ecTBa: BO,D;a H MHHepaJibHbie COJIH.

!Ipopacmallue ce.M.flll -

3TO nepexo.n; H3 COCTO.HHH.H IIOKO.H K pa3BHTHIO

H pocTy aapo.D;hIIIIa.
,ll;JI.H npopacTaHHa aeo6xo;:i;ttMhI onpe,n;eJieHHhre ycJIOBH.H:

3

6

7

a
Pr.tc. 81. CTpoeH111e ceM5lH ABYAOllbHblX: a -

6
6e3 3HAOcnepMa;

B

6-

c 3HAOCnepMoM; B -

c nepi-1cnep-

1 - KopewoK; 2 - 30111KOTll11lb; 3 - C8M6HHa5l KO)l(ypa; 4 - rnnOKOTll!llb; 5 - C8MSlA0115l; 6 - 3HAOCnepM; 7 - nep111cnepM
MOM.

l{naccut/JuKau,uR noKpbtmoceMe1t1tbtX pacmenuii

npOHHKHOBeHHe BO)J;hI: IIpHBO)J;HT K

203

Ha6yxaHHIO, aKTHBH3aD;HH <Pep-

MeHTOB ll MCTa60JIH3Ma;
TeMnepaTypHhlll peJKMM: XOJIO)J;OCTOHKMe paCTeHHJI npopacTaIOT npu TeMnepaType

+ 1 ... + 5 C (pOJKh, MOpKOBh H ,a;p.); Tern10JII06HBh1e npopacrn+ 15 'C (orypil;hl, TOMaThI, KYKypy3a H .D;p.);

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H aiJae.~uwe npopac mm1ue -

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IIoiJ3eMIWe npopacmaHue -

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2-'IJieHHbre

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Bomanu1w

204

OcHOBHbie ceMeHCTBa KJiacca ,ll;By,n;oJibHLie (KpecTou;BeTHbie,
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MaJibBOBbie, MapeBbie, Buaorpa,n;Hbie)

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CeMeiiCTBO HacqMThIBaeT OKOJIO

lKU31teltHbte <jJop.Mbl -

3

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pacTeHMR o6oerrOJibie, TpaBHHMCThie, 0,IJ;HO- M .n;By-

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205

HO'IHa.H qmamrn H ,n;p.; copH.HKH

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( ) -

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poMallIKa JieKapcTBeHHaH, TbICJiqeJIMCTHIIK, KaJieH,n;yJia H .n;p.; .n;eKopa-

THBHhie pacTeHH.H -

reoprHHhl, aCTPbI, xptt3aHTeMbI ll .n;p.; copHHKM -

Bac:rr-

JieK, OCOT, aM6po3HH H .n;p.

OcHOBHLJe ceMeiicTBa KJiacca 0.ZJ;HO.ZJ;OJILHLie
( JIHJieHHLJe, 3JiaKOBLie)

CeMeMCTBO nMneMHble
CeMe:HCTBO Jl:rrJie:HHbie HaCtJHThrnaeT OKOJIO 1300 a11.n;oa pacTeHHH.
IKU3HeHHbte rpop.Mbl -

B OCHOBHOM MHOroJieTHHe JIYKOBll'lHbie HJilI KOpHe-

BHIJJ;Hhle TpaBbl, peme BCTpe'laIOTC5l JIMaHhl H .n;epeBhJI.
JiucmbJl -

rrpOCTbie, c .n;yroBbIM MJIM napaJIJieJibHblM JKMJIKOBaHfleM. JlMC-

TOpacrroJJOji(eHHe oqepe.n;Hoe, cyrrpoTMBHoe HJIH MyToatJaToe.

~6em1<u -

O,Zl;llHOtJHble MJIM co6paHhl B cou;BeTHe MeTeJIKy HJIM KHCTb.

Il;BeTKH o6oerrOJihie. 01COJl01{6emHUIC -

rrpoCTOH, BeHtJHKOBll,Zl;HhIH H3 IIJe-

CTll cpoCIIIHXC.H MJIH CB060,Zl;HhlX JIHCTOtJKOB, pacnOJIOJKeHHhIX B .n;aa Kpyra.

AniJpol{eU

COCTOHT ll3 IIIeCTH ThltJHHOK, ThitJHHOqHbie HHTH MoryT np:rrpac-

TaTb K CpOCTHOJillCTHOMY OKOJIOD;BeTHllKY.

I'uHel{eU -

O,Zl;HH, u;eHOKaprr-

HhIH M3 Tp.ex cpocllIMXC.f.! nJio,n;oJIMCTMKOB. HaceKoMoorrhlJIHeMhie pacTeHM.H.

IlJLoiJ -

Kopo6otJKa MJIH .f.!ro.n;a.

3HatJeHMe: IIMJl\eBhie pacTeHHH pacTeHMJI -

JiyK, qecHOK, crrapma. Jie1<apCTBeHHbie

JiaH,D;hIIII, KyrreHa lf .n;p. ,Il;eKopaTMBHhie -

TJOJihITaH M .n;p. H.n;oBMTbie pacTeHHJI -

JIMJIM.f.!, r1rnu;nHT,

BopoHnii: rna3, tJeMepnu;a H .n;p.

ct>opMYJia u;aeTKa JlllJleHHhlX:

*P3 +3A3 +3G(Q)

Haa~apcnrno /.(<Hiaepubie, unu llpo1wpuombl

209

CeMeHCTBO 3naKH, HnH MsiTmtKOBble

(p"1C. 83)
CeMeil:cTBO 3JiaKOBhIX HacqHThIBaeT OKoJIO

11

TblC.Hq Bll.ll;OB.

)/(u:meH1tbte <jJopMbt -

MHoroJieTHHe Tpa-

BHHHCTbie pacTeHH.H c MOqKoBaTOM KOpHeBOM
cncTeMoil:. ,Il,peBeCHhie cpopMbI CTe6eJih 3JiaKoB -

6aM6y1rn.

COJIOMHHa, HMeeT IIOJihie

Mem.n;oy3JIH.H H llHJIHH.n;pttqecKyIO cpopMy.

J]ucmbJl -

COCTO.HT H3 BJiaraJIIU.11a ll JillC-

TOBOM nJiaCTHHKH. J111cT0Boe B.'!ara.JIHIIIe 3aIIIHlllaeT MepHCTeMy B y3Jiax, 38 cqeT 3TOM MepHCTeMbl pacTeHH.H cnoco6Hhl K

MY ( uHmep1caJLRpHOMY)

6Cma60'lHO-

pocTy. BeTBJieHHe

5

3JiaKOB Ha.n; HJIH rro.n; IIOBepXHOCTbIO noqBbl
B

30He KyW,eHUJl -

y OCHOBaHirn no6eroB co

6

c6JinmeHHhIMH y3JiaM11.

I.{ BemKu -

o6oenoJibie,

pa3.n;eJibHOno-

J1hle BCTpeqaJOTC.H pe.n;Ko (KyKypy3a). ll;BeTKH co6paHbl B co11BeTH.H KOJIOCOK, CJ10jJ{HbIM
KOJIOC H MeTeJIKa. Ha OCH KOJIOCKa BbIXO,ll;.HT
OT o.n;Horo

.n;o

HeCKOJibKHX 11BeTKOB. B ocHo-

BaHHH OCH H8X0,!1;.HTC.H BepxH.H.H ll HHJl<H.H.H
KOJIOCKOBbie qernyH -

BH,ll;Oll3MeHeHHbie

JIHCTb.H. Bhlrne Ha OCH pacnoJiaraIOTC.H IJ;BeT-

6
P11c. 83.

CxeMa cTpoeH"1Sl KOnocKa

31laKOB (a), A"1arpaMMa u,seTKa
OCb KOllOCKa:
'18Wyfl;
wySl;

-

3 -

4 -

2 -

sepxH51Sl KOllOCKOBaSl Ye-

Hi.l)l(HSlSl U,B8TKOBaSl Yewyfl;

sepxHSlSl U,B8TKOBaSl Yewysi;

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(6). 1 -

H"1)1(HSlSl KOllOCKOBafl

6-

5

110-

7 _ Tbl'1"1HKa; 8 _ necrnK ·

KH. IJ;BeTKll np1-mpblTbl BepxHeil: H HHmHe:ii
llBeTKOBhIMH <Ierny.HMH. Ha.n; BepxHe:ii :a;BeTKOBO:ii qernyiiKoii pacnoJiaraIOTC.H
,ZJ;Be MaJieHbKHe 6ec:u;BeTHbie IIJieHKH, H83bIBaeMbie

AHiJpou,eu
I'utteu,eu -

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(ocTaTKH OKO-

JIOIJ;BeTHHKa).

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Mll HHT.HMH.

O,ll;HH, nceB,ll;OMOHOKapIIHbIM H3 ,ll;BYX cpOCIIIHXC.H IIJIO-

,Zl;OJIHCTHKOB (TpeTHH pe.n;y:u;npyeTca); pbIJibIJ;a neCTHKOB nepHCTbie.

II.1wiJ -

3epHoBKa. IlJieH'IaTbIM OKOJIOIIJIO.ll;HHK IIJIOTHO np11JieraeT K ceMeHH

H HHor.n;a cpacTaeTC.H c ceMeHHOH Komypoil::. OnbIJieHirn nepeKpecTHoe. BoJibIIIHHCTBo 3Jl8KOB BeTpOOIIbIJI.HeMbie.
3Ha'IeHHe: XJie6Hhie 3JiaKH CTHMK 11 .n;p. KopMOBhie -

nrneHn:u;a, pomh, oBec, KYKYPY3a, caxapHbIM Tpo-

M.HTJIHK, TMMO<)leeBKa 11 .n;p. CopH.HKH -

<I>opMyJia IIBeTKa 3JiaKoB:

nh1peil: 11 .n;p.

tP(2)+2A3G(2)

Ha;:i;:u;apcTBO ,lJ;oMepui.ie, HJIH IlpoKapHoTi.1

K

npoKap110TaM OTHOC.HT 6aKTepn11, apxe6aKTepnH (oT rpe'I.

archaios -

,Zl;peBHeHIIIHM), ay6aKTepHH, 8KTHHOMHIJ;eThl (JiyqllCTbie rpH6KH) H CHHe3eJieIibie Bo.n;opocJIH (:u;11aH06aK'rep11n).
OrpoeHne npoKapHOTH'IecKoil:: KJieTKH paccMaTPMBaeTc.H B pa3.n;eJ1e «IJ;HTOJiorHH» •

210

BomanuKa

Ba1nepHH
BaKTepntt 6hJJIH om1caHbI ro"1"1aH,n;CI<HM yqeHhIM AHTOHH BaH JieBeHryKOM
B

1676 r.

Bee orpoMHoe pa3Hoo6pa3ne 6aKTepttif KJiaccm!rn!IHPYIOT no <t>opMe, OTHOwettmo K Te:vinepaTypHOMY pemnMy, cnoco6y ITHTaHH.H H ITOJIY'IeHH.H 3HeprHH,
poJIH B )KH3HH qeJioBeKa (CM. pa3,n;e.1 «U:HTOJIOrmI» ).
<l>opMhl 1rneToK 6aKTepuii:
naJ10'l!C06UO/-lbte, H,'IH 6au,UJIJlbt: K HMM OTHOCJITC.H KHIIIeqHa.H IIaJIOqKa,
ceHHaJI ITa.noqKa ( O,L(HHOqHhie 6aKTepHH), 6aIJHJIJibI CH6npe.H3BeHHhie
( co6paHhI B IJeITO'IKH);
!COK/CU (IIIapHKH): Ofrhe,n;HHeHHbie no ,[(Be - ,n;HnJIOKOKKH (B036y,n;HTeJIH
nHeBMOHHH); uerroqKH - CTpCITTOKOKKH (B036y,n;HTeJIH aHrHHhI); CKJieeHHbie no qeTbipe (caplI,HHbI); CTaq>HJIOKOKKH o6pa3yIOT «rp03,L(b»;
eu6puO/-lbl: HMeIOT H30rHyTyIO ¢opMy (XOJiepHbIM BH6pHOH);
cnupu.n.nbt H cnupoxemot: nn:TeBH.L\HhI n: 3aRpyqeHbI ITO crrnpaJin, pa3Jrn:qaIOTC.H ITO IIO,D;BI-IJKHOCTH. BJie,n;naJI cnnpoxeTa - B036y,n;n:TeJib cn:¢HJIHCa.
Ilp11 HaCTYIIJieHHH He6JiaronpmITHhIX ycJIOBHM 6aKTep11n: o6pa3yIOT cnopbI c TOJICTOH o6oJioqKoii. CITopbI 6aKTepnM: cJiymaT )J;JI.H rrepeHeceHHJI
He6Jiarorrp11aTHhlx yc.noBn:ll cpe,n;br, cnoco6HhI BbI,n;epmnBaTb oqeHh BhIcoKne 11 oqeHh HH3Kne TeMnepaTyphr.
Ilo OTHOIIIeHHIO K TeMuepaTypHOMY pe'1CHMY 6aKTepnn ,n;eJIJITC.H Ha HeCKOJibKO rpyurr:
mepMorjJUJlbl (o6IITaIOT B KHII.HIIJ;HX KJIIOqax);
Me3orjJuJLbt (o6nTaIOT npn TeMrrepaType 36-37 C, 6oJihIIIHHCTBO BhI3hIBaIOT pa3JIHqHhie 3a6oJieBaHH.H);
ncuxporjJu.not (o6nTaIOT rrpn 0 ... -1 'C; B ropax «KPOB.HHhre rraTHa» Ha
CHery - KOJIOHHH 6aKTepnii-ncnxpo¢HJIOB).
Ilo cnoco6y nuTaHHH u noJiyqeunn auepruu 6aKTepnn oqeHh pa3Hoo6pa3HhI.
Tttrrhr o6MeHa Be:rn;ecTB y 6aKTepnii onpe,n;eJI.HIOT no rrpnpo.n;e HCToqHHIWB Tpex
Heo6xo,n;HMbIX KOMnOHeHTOB JKI13HH: ucmO'l/-lUI( ;mepiuu, ucmff'l/-lUI( yiJLepoaa,
ucmo'LHUK eoiJopoiJa (Bo,n;opo,n; HymeH RaK ,n;oHop aJieKTpOHOB: H 2 ~ 2Ht + 2e)
H MOJKeT 6hITb 3aMeHeH ,n;pyrHMH 3JieMeHTaMII, OT,n;aIO:rn;IIMl13JieKTPOHhI.
Ilo HCToqHIIKY 3Heprnn:
rjJomomporjJbl (IICIIOJib3YIOT COJIHeqHbIH CBeT);
xe;nompor/Jbt (ncnoJih3YIOT ;rneprmo xnMnqecRnx CB.H3ett B HeopraHnqeCKIIX Be:rn;ecTBax).
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Ilo ncToqHnKy yrJiepo,n;a:
aemompor/Jbt (IICIIOJib3YIOT yrJieKHCJibIM ra3 H3 B03,n;yxa);

pa 3 Mno:HCenue 6aKmepuii. 3na•tenu.e 6aKmepuii

211

iemepompoqJbl (11CIIOJib3YI-OT opraHWICCKOe BCll~CCTBO ).
feTepoTpOcpbI B 3aBHCI1MOCTI1 OT cy6cTpaw1, Ha KOTOPOM OHli pa3BI:IBaJOTCH,
no,n:pa3,n:eJIHJOT ua rpynnhr:
canpompOc/Jbt (carrpoqmThI) - IIliTaJOTCH MepTBhIM opraHWICCKliM BCIIJCCTBONI. 3To 6aKTepliH MOJlO'-IHOKHCJIOrO 6pomeHHH, rHHeHliJI H .n;p.;
6aKmepuu-napaaumbt - pa3BHBaJOTCH Ha JKliBhIX oprattl13Max. HanpttMep, MCHHHrOKOKKH, MHK06aKTepHH Ty6epKy.'le3a. 06JIHraTHhie aua3p06bI - B036y.n;HTe.'IH CTOJI6HHKa, raHrpeHhI;
.MUKcompotjJnbte 6aicmepuu - crroco6HhI HK rrapa3HTH'-IeCKOMy, HK canpOTpocpttoMy IIHTaHHJO (6a!IHJ1Jlhl CbIIIHOrO THcpa).
Ilo HCTO'-IHHKY BO,n:opo.n;a:
opianompotjJbt (rroTpe6JIHJOT opraHHKy);
Jtumompoc/Jbt (OT rpe'I. lithos - KaMeHb) - HCIIOJ1b3YJOT rrpOH3BO,ll;Hhie
JIHToccpepbr: H2' H 2S, S, Fe 2 ' H .n;p. ).

Pa3MHomeHne 6aKTepuH:
BecnoJLoe, HJIH eeiemamuenoe, pa3MHomeutte.
BaKTepHH pa3MHomaJOTCH nyTeM paBHOBeJimwro 6ttHapHoro rrorrepe'-IHoro
.n;eJieHHH KJieTKH. IJ;HKJI .n;eJieHHH MO/KHO pa3,n:eJIHTb Ha TPH 3Tarra:
1. PerrJIHKa!IHH KOJih!IeBou .IJ:HK (xpoMOCOMbI) 6aKTepHH. Ilpo!Iecc ua'-IHuaeTcH B TO'-IKe npHKperrJieHHH .IJ:HK K I1HTOHJia3MaTH'-IeCKOH MeM6paue
H 3.n;ecb me 3aBeprnaeTCH.
2. CHHTe3 MeM6paHhI B o6JiaCTH KOHTaKTa c .IJ:HK. 3TO IIPHBO,ll;HT K pa3.n;eJieHHJO ,L(O'-IepHHX MOJieKyJI .IJ:HK.
3. 06pa30Bam1e rrorrepe'-IHOH rreperopO,LIKH MeJK,LIY .D:BYMH ,ll;O'-ICpHHMH xpoMOCOMaMH.
Y.n;BoeHne 6aKTepnaJihHOii: KJieTKH rrpottcxo.n;HT B 6JiarorrpHHTHhIX ycJIOBHHx KaJK,!J;hie 15-20 MHH.
IloJLoeou rrpo!Iecc y 6aKTepHii: (HJIH ienemu'leCKa.fl peKo.M6unau,u.R).
y MHOrHX 6aKTepHii: H3BeCTeH IIOJIOBOH IIPO!ICCC, rrpH KOTOpOM npOHCXO,!J;HT
TOJibKO o6MeH reHeTH'-ICCKOH HHlpOpMa!IHeii: Mem.n;y KJieTKaMH, HO o6pa30BaHHe
HOBhIX KJieToK He rrpottcxo.n;nT. Bhrp;eJIHIOT TPH cpopMhI reHeTH'-IecKoii: peKoM6HHa!IHH:
1. TpaHccpopMa!IHH (nepeHoc .IJ:HK H3 paapyrneHHoii: KJieTKH B JKHBYJO
KYJihTYPY 6aKTepnii:).
2. KoH'hrora!IHH (rrepe.n;aqa HacJie.n;cTBeHHoii: HHcpopMa!IHH rrpn npHMOM
KOHTaKTe KJieTOK).
3. TpaHC.D:YK!IHH (rrepeHoc yqacTKOB .IJ:HK OT o.n;Hoii: 6aKTepHH K .n;pyroii:,
rrocpe.n;cTBOM cparoBoro BeKTopa - rrepeHOC'-IHKa).

3Ha'leHne 6aKTepuH:
BaKTepHH HMeJOT orpoMHoe 3Ha'-IeHne B rrpnpo.n;e H B JKH3HH qeJioBeKa.
MomHo Bhr.n;eJIHTh 3KOJIOrH'-IecKoe aHa'-IeHHe H rrpnMeHeH11e B qeJioBeqecKoii:
,Zl;e.HTeJibHOCTH H Me,!J;H!IHHe.

BomanuKa

212

31CoJLozu'leC1coe .ma'leHue:
carrpocpHThl HBJI.HIOTCJ! pe.l(yLI;CifTaMH, pa3pyrnaIOlll;llMH H MHHepaJIII3YIOll~HMH

MepTBYIO opraHI·my;

6aKTepHH yqaCTBYIOT B KpyroBopoTe XIIMH'IeCKHX aJieMeHTOB (meJie3o6aKTepmi:, cepHhie 6aKTepirn, a30TcpIIKCaTOpbI If p;p.);
yqacTBYIOT B npou;eccax rro 1rnoo6pa3oBaHHJI. Pa3JiomeHrre 6aKTepIIHMH
a30THCTbIX Ber.u;eCTB Ha3hJBaeTC.ff I'HIIeHHeM.

3ua'leHUl' 6 'leJL06e'leCICOU aeJ1.meJ1bHOcmu U Jtt.eauu,une:
B IIHlll;CBOH npOMhIUIJieHHOCTH 6aKTCPIIH IICTI0Jlh3YIOTC.ff ,ll;JIH IIOJiy<IeHII.ff
KHCJIOMOJIO'!HhIX rrpo,a;yKTOB, Chipa;
B Xl'IMH'!eCKOH npOMhllliJieHHOCTH -

IIOJJyqemi:e CIIHPTOB,

yKcyCHOM

KHCJIOThl, aIJ,eTOHa, caxapoB H p;p.;
B Ml'IKp06HOJIOI'H'!eCKOH npOMhllliJieHHOCTH -

rrpoH3BO)J;CTBO aHTH6u:o-

THKOB, BHTaMl'IHOB H ,a;pyrHX Ber.u;ecTB (aKTHHOMHIJ,CThl -

npop;yIJ,eHThl

aHTH6HOTHKa CTpeIITOMHIJ,HHa);
HCIIOJih3YIOTC.H B I'CHHO-HHJKeHepHhIX HCCJie,noBaHHHX H )J;JIH IIOJiyqeHH.H
rrpop;yu;eHTOB pa3JIH'!HhlX BClll;CCTB, HaIIpHMep ropMOHa HHCYJIHHa;
OTpm~aTeJihHa.H

pOJih: nopqa npo,a;yKTOB, KHHr, MaTepMaJIOB; Bhl3hIBa10T

3a6oJieBaHHH pacTeHMM, JKHBOTHhIX, qeJIOBeKa. BaKTepttM HBJIH10TC.H B036y,a;ttTeJI.HMH MHOI'HX 6oJie3HCM '!CJIOBeKa: TH(j>, tJyMa, XOJiepa, Ty6epKyJIC3
HJJ;p.

ll;apCTBO rpu61>1
I'pu6bt -

MHOJ'OtJHCJieHHa.H rpyrrna opraHH3MOB, HaC'lHThIBaIO~aa 6oJiee

120 ThIC.II'! BH,Zl:OB.

8TO reTepoTp0¢Hhie opraHH3Mhl, KOTOphIC HMeJOT IIpH3HaKH

CXOJJ;CTBa c JRHBOTHhIMH H c pacTeHHHMH.

JlpU3HalCU cxoacmea C pacmeHU.R.MU: HeIIO)J;BHJKHOCTh, HeorpaHM'!eHHhIM
KJleTO'lHhIH pocT, HaJIH'!He KJieTO<IHhIX o60JIO'!eK, pa3MHOJKCHHC cnopaMH.

Ilpu3nalCU cxoacmea c JICU6DmHblMU: OTCyTCTBHe XJIOpOIIJiaCTOB, reTepoTpo¢HOCTh, na.n:H<I:He 3anacHoro noJI:Hcaxap:H,na a30THCTOJ'O o6MeHa -

rJIMKoretta, npo,n:yI<T

MO'!CBHHa, B KJICTO'lHhIX o60JIOtJKax Ilp:HCYTCTBYCT

XHTHH.

CTpoettne
I'pn6hI O'!CHh pa3HOo6pa3HhI no pa3MepaM H BHCUIHCMY BH'AY. 06~ett tJepTOH

zurjJ. CoBOKYilHOCTh
zpu6nuu,y, mrn Muu,e.nuu. Ilpn o6pa3oBaHnn oprattoB pa3MHOmemu1 rpM6Hhie HHTH IIJIOTHO rreperrneTaIOTCH, o6pa3yH JIOJKHYIO TKaHh nJteKmeHXUMy.
Ila CTpOCHHIO Bhl,Zl;CJI.IIIOT ,n:Be 60JihllIHC rpyIIIlhl rptt6oB: 6blCW.Ue H HU3W.Ue.
HH3IIIHe rpn6hr HMCIOT HCI\JICTO'!HhIH MHOI'O.H,ZJ;epHhIH MHIJ,CJIHH ( 1CJlaCc 3uzoMUU,embt). y BhICIIIHX rpM60B MHIJ;CJIHH MHOI'OKJICTO'lHhIH ( lCJlllCC Ac1CO.M.Uu,embt, Ba3uauo~wu,embt, Hecoaepw.eHnbte zpu6bt}.
npaKTH'!CCKH BCCX rpn6oB HBJIHCTC.II TCJIO B BH,ZJ;e HHTCH -

rncpoB 06pa3yeT

Il,apcmBo I'pu6b£. CucmeMamuica zpu6o6

213

PaaMHO:iKenne
fptt6hl pa3MHOJKaIOTC.H BereTaTHBHbIM, 6ecrrOJibIM H IIOJIOBbIM rryT.HMH.
Beiemamu6ttoe pa3MHOJICettue rrpo1:1cxo.n;1:1T: qacT.HMH MHIJ;eJ11:1.a; pacrra,n;oM
:~rnu;eJll:l.H Ha OT,n;eJibHbJe KneTKH - xnaMHll;OCIIOpbl c TOnCTOH o6onoqKoii; pacUall;OM Ml:II.\eJIH.H Ha TOHKOCTeHHbie apTpocrropbI, OHll;HH. y .n;pommeBbJX rp1:160B
BereTaTHBHOe pa3MHOJKeHHe rrpOHCXOll;HT rryTeM rroqKoBaHH.H.
BecnoJLoe pa3MHOJ1Ce11ue rrpo1:1cxo,n;1:1T rrocpe,n;cTBOM crrop - BereTaTHBHhIX
KneTOK 6ecrronoro pa3MHomeHH.H:
3oocnopaMH (no.n;BH.iKHbJe, y BOll;HhIX rp1:160B);
aH,n;oreHHoe cnopoHOIIIeHHe (cnophI BHYTPH oco6hIX KneTOK - cnopaHrHeB);
aK3oreHHoe cnopoHOllleHHe (Ha cneu;m1nhHhIX zu<jJax Kouuauenocu,ax
o6pa3yIOTC.H KOHH.n;Hecnophl, HarrpHMep rpn6 IleHHIJ;HnnyM).
IloJLoBoe pa3MHOJICettue npo1:1cxo,n;1:1T rryTeM cnH.HHH.H raMeT.
31:1roraMH.H. IlpH BCTpeqe reTepoTann1:1qHhIX meHCKHX ( «+») H MY.iKCKHX
( «-») HHTeH rpH6a OTqneH.HIOTC.H KneTKH H 3aTeM CJIHBaIOTC.H, o6pa3y.H
.mzocnopbt.
faMeTaHrHoraMH.H. y BhICllIHX rpn6oB cnHBaThC.H MoryT He OT,n;enhHhie
KneTKH, a nonoBhJe opraHhl - zaMemanzuu.
CoMaToraMH.H. Cn1:1.HH1:1e KneTOK MHu;enH.H.
Bo Bcex cJiyqa.Hx cHaqa.Jia cn1:1BaeTc.H nHTonna3Ma, 3aTeM cn1:1Ba10Tc.a H,n;pa
(CTall;H.H /J;l'.IKapHOHa) . .D:1:1rrno1:1,n;Hoe .H.n;po IIOCJie Meiio3a 06pa3yeT crrophl (n).
Y BhICllIHX rp1:160B rrocne nonoBoro npou.ecca 06pa3y10Tc.H nJLo8o6bte meJLa:
3aKphIThJe (Kneil:cTOTeu.nii) - .n;om,n;eBHKH; npHOTKphIThie (nep1:1TeU.HH) B BH.n;e 6yThIJIKH; OTKphIThle (arroTeU.HH) - y lllJl.HITOqHhIX rp1:160B.
CncTeMaTHKa rpn6oB
Bee rp1:16hl .n;en.HT Ha rpyrrnh1: HH3nme (M1:1u.en1:1:ll 6e3 neperopo,n;oK) 1:1 BhICllIHe (M1:1u.en1:1ii c neperopo.n;KaMH).
Hnamne rpn6L1: omaeJl XumpuauoMUKOmbt, omaeJl 00.MUKOmbt, omiJeJl 3u20.MUKOmbt, omaeJl M Ul(CO.MUKOmbt.
Xumpuauo.MuKombt - Hatt6onee npHMHTHBHhJe rp1:16h1. He HMeIOT MHU.eJIHH. Teno - B BH.n;e KOMoqKa u;1:1Tonna3Mhl. KneToqHa.H o6onoqKa oTcyTCTBYeT. Hatt6onee H3BecTHhle npe,n;cTaBHTenn: onbnH,n;HyM - Bhl3bIBaeT 6one3Hh
KarryCThl «qepHa.H HO.iKKa», CHHXHTPHYM - Bhl3hIBaeT paK KapTocjlenH.
00.MUKOmbt - HMeIOT pa3BHThIH MHIJ;eJIHM. B KJieToqHoH o6oJioqKe OTCYTCTByeT
XHTHH, HO eCTb u;enn10no3a. 06HTaIOT BO Bna:arno:ll HJIH BOll;HOH cpe.n;e. BamHeHllIHH
rrpe,zi;cTaBHTeJib - cjlHTOcjlTopa - Boa6y,n;1:1Tenh 6oJie3HH KapTocjlenH, TOMaTa.
3uzo.MUKOmbt - o6pa3yIOT xopolllo pa3BHThlii, BeTBHin;HMC.H MHn;enttii. XapaKTepHo paaMHO.iKeHHe - anroraMHf!. ill1:1p0Ko pacnpocTpatteHbI: MyKop, xne6Ha.H
IIJieceHb. 3HrOMHU.eTbl 06paay10T :miJompo<Jmy10 .MUKOPU3Y c KOPH.HMH pacTeHHM.
MUKCO.MUKOmbl - HHOr,n;a BhI.n;en.HIOTCH B OT.n;enhHYIO rpyrrrry opraHH3MOB,
TaK KaK coqeTaIOT rrpHaHaKH mHBOTHhlX 1:1 rptt6oB. OT.n;enhHhJe aMe6oBHll;HhJe
MHKCOMHU.eThl cnoco6Hhl cnon3aThCH B e.n;HHoe cnnaeBH.n;Hoe Teno. CnttaeBHll;aoe Teno MomeT o6pa30BhJBaTh nno,n;oBble Tena H cnophl. K MHKCOMHKOTRM

Bomanu1w

214

P111c. 84. Cnopb1Hb51 (Claviceps purpurea): 1 -·Kanae p)l(111 co crnepm-1111RM111; 2- CTPOMbl, sb1pocw111e
Ha nepe3111MosasweM CKnepm-1111111; 3 - npo1:ionbHb1i1 pa3pe3 Yepe3 CTPOMy c nep111TeL\111RM111; 4 - npoAonbHblM cpe3 yepe3 nep111Tel\111111 c CYMK3M111; 5 -- CYMl<a c H111TeB111AHblMll1 acKOcnopaM111

OTHOC.HTC.H napa3HThI KYJibTYPHhIX pacTeHHM (KHJia KanycTHa.H, nopOIIIHCTa.H
nap III a KapTo<f>eJI.H ).
BbICWHe rpH6hI: omoeJt ACICO.MUKOma, omoeJt Ba3UOUO.MUKOma, omoeJL
JJ,eumepo.MUKoma, u.riu H ecoBepweHHbte ipu6bt.
,[(JI.H BhICIIIHX rp:rr6os xapaKTepuhr <IJieHHCThie rmphr c neperopo,n;KaM:rr ( cenma.Mu ).
AcKOMHKOTa. OtieHh pa3HOo6pa3HhI no cTpoeHHIO. ,[(JI.H 6oJihIIIHHCTBa xapaK-

TepHo o6pa30BaH:rre GCIC06 -

cyMOK c ranJIOH,ll;HbIMH cnopaMH, o6pa3YJOJ:nHMHCJI

B npou:ecce Meii:o3a. Pa3MHomett:rre noJiosoe (cyMKH acK:rr), 6ecnoJioe (Kott:rr,n;:rrecrropbr), BereTaTHBHOe (llaCTH MHU:eJIH.H HJIH TIOl{KOBaHHe).
,n;pOJKJKH -

K :3TOH rpyrrne OTHOC.HT:

o,n;HOKJieTOl!Hbie rpH6hI, cnoco6Hhie K BereTaTHBHOMY pa3-

MHOJKeHHJO nyTeM TIOl{KOBaHH.H (rreKapCKHe ,n;pomm:rr, BHHHhie ,n;pomm:rr,
KOPMOBbie µ;poJKJKH H ,n;p.);
crropbIHb.H -

rrapa3HT 3Jia:KOB (pHC.

84). JleKapCTBeHHhIM rpH6, co,n;ep-

mam:rrii: amrnJIOH,n; aproTaMHH (HCIIOJih3yeTC.H B po,n;OBOH rrpaKTHKe -

co-

KpamaeT MaTI<y), a TaKJKe npOH3BO,ll;Hbie JIH3eprHHOBOH KHCJIOTbl CHJihHoro raJIJIIOU:HHoreHa. YnoTpe6Jiemrn 3apamettttoro 3eptta onactto
,ll;JI.H qeJIOBeKa, TaK KaK rrpHBO,ll;HT K T.HJKeJihIM OTpaBJieHH.HM.
CMOpl{KH H CTpOqKH TPJO<f>eJIH -

Cl{HTaJOTC.H YCJIOBHO C'he,n;o6HblMH;

C'he,n;o6Hbie;

MHKOPH3006pa3yJOm:ri:e rp:rr6hI -

HeKOTOpbie rrpe,n;cTaBHTeJIH o6pa3yIOT

3KTOTpo<f>HYIO MHIWpH3Y.
IJ;HKJI pa3BHTHH CIIOphIHhH:

1)

OCeHbJO Ha 3JiaKOBhIX pacTeHH.HX o6pa3yIOTCJI CICJLepou,uu -

TeMHO-<f>HO-

JieTOBhie POJKKH, o6pa3oBaHHhie MHU:eJIIIeM rp:rr6a B CTa,n;HH IIOKOJI;

2) 3HMoii: CKJiepou:H:rr Bblrra,n;aIOT H3 3JiaKOB, 3HMYJOT B rroqBe. Bectto:H: c1rnePOU:HH rrpopacTaIOT, o6pa3y.H Bhrpo,CThI c roJIOBKaMH -

cmpo.Ma.Mu.

Il,apcm6o I'pu6bt. CucmeMamuKa zpu6o6

B cTpoMax

215

o6pa3yIOTCH nJiop;oBhIC TeJrn -

nepumeu,uu,

3anoJmeHHhre ac-

KaMw c aCKOCIIOpaMw;

3)

JieTOM cnopbl pa3HOCHTCH BCTpOM Ha pbIJibl.J;a 3JiaKOB I1 npopacTaIOT. fII-

1w1wiJuu -

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cnopbr 6ecnoJioro pa3MHOlf<eHIIH.

IlpII 3TOM rII<lJbl BhI;:J;eJIHIOT KaneJihKI1 CJia;:J;KOM JKII;:J;KOCTH -

pocy.

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CHT CIIOphI rpw6a Ha ;:i;pyrIIe pacTeHI1H;

4)

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85).

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nyTeM o6pa3oBaHI1JI KOHII;:J;IIM. XapaKTepHo rroJioBoe pa3MHOJKeHIIe, rrpII KOTopoM CJIHBaIOTCJI /];Be BereTaTI1BHhie O;:J;HOJI;:J;epHbie KJieTKII rarrJIOII;:J;HOrO M1II1eJIMJI (coMaToraMIIH).

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K

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II IIIJIHUKOM. MHorwe II3 HIIX c'be,n;o6HhI (6eJihIH rpII6, rro;:i;ocIIHOBIIK, rro;:i;6epe3oBIIK, orreHOK II ;:i;p.). Cpe,n;II IIIJIJIIIO'IHhIX rpII6oB eCTh JI,D;OBHThre
(6Jie;:i;HaH noraHKa), IICIIOJih3YeMbie B Me;:J;III!IIHe (MYXOMOp). fpII6bI Ha

98 %

COCTOJIT II3 BO;:J;bl • .IJ:oJIIO cyxoro Be~eCTBa COCTaBJIHeT 3Ha'IHTeJibHOe

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6) ipu6bt-mpymoeu1Cu.

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rrpIIBO;:J;JIT K pa3pyIIIeHIIIO Il rII6eJIII ;:i;epeBheB;
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216

pa3HOC.HTCJI HaCeKOMhIMII. Iloc;re IlOJIOBoro npou;ecca ,l\IUIJIOll,l\Hhie cno-

(2n) nopa:maJOT JillCTbH 3JiaKOB, CHIDKaH ypo:maiiHOCTh.
,r:t;eiiTepoMHlleTLI, mm HecoBepmeHHbie rpH6hl. ::ha HcKyccTBeHHaH rpyrrna
phl

rpH60B BKJIJOqaeT Bll,l\hl, y KOTOphIX He H3BecTeH IIOJIOBOM rrpou;ecc. Mttu;em1:H
qJieHHCThIH.

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c KOHH,l\llHMM (Hapyarnoe cnopOHOIIIeHHe). K oTOH rpynne OTHOCHT:
a) neuuu,uJtJt BeTBJieHHH -

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KOMITOHeHT -

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pocJIH noJiyqaroT OT rpH6a MHHepaJibHhle Beru;ecTBa.

M u1Co6uonmbt - rroqTH y Bcex JIHIIIaHHHKOB cyMqaTbre (AcKOMHKOThI Ascomycota) rpH6hI, TOJibKO y 20 TporrnqecKHX BH.D;OB BCTpeqaroTCH Ba3H.L\HaJibHbre rptt6hI (Ba3H,l\HOMHKOTbI).

B

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OmiJe.'l 3eJteHbte eoiJopoc.1tu -

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BH,11,0B. HaH6oJiee qacTo BCTpeqaJOTCH npe,n:cTaBHTeJIH oT.n;eJia UHCTeKOKK
H Tpe6oyKCHH.

OmiJeJt Cutte3eJtellbte 60iJopoc.1tu (u,uano6a1Cmepuu) -

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qaroru;HMHCH ¢lHK06HOHTaMH HBJIHIOTCH npe,n:cTaBHTeJIH po,n:a HocToK.
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PacTyT Ha KaMHHX, CKaJiax, Kope ,n:epeBheB, HeKOTOpbie B BO,n:e.

m'.H3HeHHbie <J>opMbl (pHC. 86)
IIo ¢>opMe TeJia, HJIH ma.1t.1t0Ma, BbI,11,eJIHIOT TPH THna JIHIIIaHHHKOB:

1C0p1C06ble HJIH HGICUllllbte -

HMeIOT BH,l\ KopoqeK, ITJIOTHO cpacTaIOTC.li

c cy6cTpaToM. HanpHMep, JleKattopa;

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217

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paBHOMepHoe pacrrpe,11;eJiemrn Bo,11;opocJieii H rptt6os.
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cpbl rptt6a); rDHH,IIHaJihHbIH CJIOH (OT Ha3BaHHf! KJieTOK B0,1IOpOcJieii -

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,IIHii); cep,IID;eBHHHhlH CJIOH (rHcpbI, Me:tK,IIY HHMH eCTb rryCTOTbI); HH:tKHHii KOpKOBhIH CJIOH (rH<pbl rpH6a c pH3HHaMH, IIpHKpeII.lIHIOIJJ;HMH K cy6cTpaTy).
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pa3MHOiKaThCf! OT,IIeJihHO: BO,IIOpOCJIH -

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(Cladonia), TaK Ha3b1saeMb1j;i «011eH111j;i MOX», 11111111 s:irenb; 4 - nonepe'lHb1j;i cpe3 c11oes111ll.la 11111cTosaToro 11111waj;iH111Ka napMe11111111 (Parmelia physodes), np111Mep rerepoMepHoro cTpoett111s:i; 5 - nonePe'lHb1j;i cpe3 c11oes111ll.la 11111crnsarnro 11111waj;iH111Ka 11emorn111 (Leptogium), np111Mep roMeoMepHoro
CTpoeH111s:i; 6 - Ha'lanbHas:i CTa.A111s:i pocrn copeA111j;i y napMe11111111; 7 - HaK111nttoj;i, TaK Ha3b1saeMb1j;i

a - sepxH111j;i KOPKOBb1j;i cnoj;i; 6 r - Hll1)1(Hll1j;i KOpKOBblj;i CJ1Qj;i

«n111cbMeHHb1j;i 11111waj;iH111K» (Graphis scripta), Ha KOpKe Aepesa:
roHll1A111a!1bHblj;i cnoj;i; B -

cepAL(eBll1HHblj;i cnoj;i 1113 rncj> rp1116a;

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pacTyT B MecTax c qIICThIM BoJ.n;yxoM, c ceBepHoii cTopoHhI CTBOJIOB. RpoMe
Toro, MHOrIIe II3 HIIX IICIIOJih3YIOTC.H B X03.HHCTBe. HanpHMep,

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HO HCIIOJih30BaTh KaK MYKY npII BhIIIeqKe XJie6a.
3TO KOPM ,ll;JIH OJICHeii;

MOp.H, IICIIOJih3yeTca ,ll;Jl.H npIIrOTOBJICHH.H JI3KMyca •

.QapcTBO PacTeHHH. Ilo.ri;u;apcTBO HacTomu;ue Bo,r:i;opocJIH
BoiJopoc.rw -

3TO aKonorHqecKa.H rpynrra aBTOTpo<}lHhIX pacTIITeJihHhIX op-

raHH3MOB, rrpttcrroco6JieHHhlX K o6IIT3HIIIO B BO,ZJ;HOM cpe,n;e. Bo.n;opoCJIII OTHOC.HTC.H K

HU3utUM pacmeHUR.M, IIJIII npomoKmucmaM.

,D;na npOTOKTIICTOB xapaKTepHhI cne,n;yIDIQIIe rrpH3H3KM:
HCT ,ll;II<}l<}lepeHI!MaI!IIH Tena Ha opraHhI ( OTCYTCTBYIOT CTe6mI' KOPHH,
JIHCTh.H);
TeJio -

TaJIJIOM MJIII cJioeBIIIQe. HeT BHyTpeHHeil .n;tt<}l<}lepeHI!Hal!IIII Te-

Jia, T. e. OTCYTCTBYIOT BHyTpeHHIIe TK3HII;
IIOJIOBbie opraHhl Bcer.n;a O,ZJ;HOKJICTOqHhie.
Ilo crroco6y IIIIT3HIIH Bee BO,D;OpOCJIII .HBJI.HIOTCH aBTOTpo<}laMII, IIMeIOT pa3JIIIqHhie MO,ll;HQlIIK3I!HH XJIOpOQlHJIJia, QlOTOCHHTC3HPYIOT ( CMHe3eJieHhIC MOryT
6h!Th reTepoTpo<}laMM).
B Bo.n;oeMax Bo,n;opocJIM 06pa3yroT

rjJumonJtau1Cmon mm rpumo6ettmoc. cpn-

TOrrJiaHKTOH ,Zl;BM:raeTC.H, rrapIIT B TOJIIQe; QlIIT06eHTOC npIIKpenJieH KO ,D;HY.

.n;o 22 ThICJP-1
,n;o ,n;ecHTII OT,D;eJioB. HaII6oJiee MHoroqtt:cJieHHhI omiJeJl 3eJteHbte 6000pOCJlU, omoe.rt Bypbte BOOOpOCJlU, omiJe.'l KpaCHbte 60oopocJlu.
EcTh npecHOBO,ZJ;Hhre II MopcKtte <}lopMhI. Bcero HacqJIThIBaIDT

BH,ll;OB Bo,n;opocJie:i1:. Bhl.n;eJiaIDT

0T.ZJ;eJI 3eJieHbie BO,z:i;OpOCJIH
OT,n;eJI HacqlfThIBaeT OKOJIO

1.

3eJieHa.H OKpacKa -

6 ThICJiq

XJIOPOQlHJIJI

HH:H). ,[(pyrtt:e TIHrMCHThl -

2.
3.
4.
5.

BII,ZJ;OB. ,[(Jia HIIX xapaKTepHhI:

a,

X.llopo<}ll:IJIJI

b

(KaK y BhICillHX pacTe-

KapOTHHhI, KC3HTOQlHJIJlhI.

KJieToqHhie cTeHKH II3 I!€JIJIIDJI03hI II neKTIIHa.
3anaCHhie BCIQ€CTBa: KpaXMaJI, MoryT 6hITh MaCJia.
PaaMephr OT MttKpocKOnIIqecKIIX, KOJIOHHaJihHhIX

,n;o

KPYIIHhIX <}lopM.

Pa3MHomeHIIe BereTaTIIBHOe, 6ecnoJioe, rroJIOBoe.

OT,n;eJI 3eneHhre Bo.n;opocJIM co,n;epmIIT:

1. KJtacc BoJtbBOICCOBbte ( o,n;HOKJieToqHhle

MJIH KOJIOHttaJihHhre,

IIMeIOT

mryTIIKII, HaJittqtte qarneBtt,n;Horo xpoMaTo<}lopa): xJiaMII,ZJ;OMOHa,n;a (o,n;HOKJieToqHhle), naH.n;opttHa, BOJihBOKC (MHOf'OKJieToqHhie).

n.apcm 6o PacmenuR. lloiJ11rapcm60 HacmoRw,,ue BoiJopoc.n,u

2. K.nacc IIpomoKOK06bLe
KOJIOHliaJibHhie,

219

(o.n;HOKJieToqHhre,

HeT mryTHKOB,

TOJICTa.H

060JioqKa):xJIOpOKOKK,XJIOpeJIJia.

3.

KJiacc KoH'hIOraTbI ( cu;errJI.HHKM) ( o.n;HmrneToqHhre, CIIMpaJibHhIH xpoMaTocpop,

eCTb

KOH'hIOra11,MH): crrnporMpa, 3MrHeMa.

4. K.nacc Y.nompuKC06bLe

(HMTqaThIC M rnrac-

5

TMHqaThIC cpopMhI, IIJiaCTMHqaThIH xpoMaTocpop; raMeThI c .ZJ;BYMH mryTMKaMM, 300crropbl -

c qeTbiphM.H mryTMKaMM): yJIOT-

7

pMKC.
CTpoeuHe KJieTKH
(Ha rrpMMepe XJiaMM.ZJ;OMOHa.ZJ;bI)

PHc. 87.

XJiaMM.ZJ;OMOHa.n;a o6MTaeT B rrpecHhIX

87).

eMax (pHc.

Bo.n;o-

n~nog~~);

aKTHBHO .ZJ;BHJKeTc.H. KJieTKa co.n;epJKHT THrrHqMMTOXOH.ZJ;plm:,

pH60COMhI,

MeJIKHe

xpo.MamorjJop -

(YHAY-

KJl8TOYHa~

3 - 6a3aJlbHbl8 T8Jlb-

u,a; 4 -

~APO C ~APblWKOM;

u,~Tonna3Ma;

6 - Yaweo-

6pa3Hb1C'i xpoMarncjJop; 7 -

o.n;MH oqeHb KpyrrHhIH xJio-

n~peHO~A;

porrJiacT B BH.n;e qarnH,

nupenoui) -

2 -·

CT8HKa;

5-

BaKyoJIH M oco6b1e CTPYKTyph1:

KneTK~

(Chlamydo-

>t<ryrnK~

monas): 1 -

Ee KJieTKa MMeeT .n;Ba mryTMKa,

Hhle .ZJ;JI.H ayKapHOT opraHeJIJihI: 3H.ZJ;OIIJia3MaTHqeCKYIO ceTh,

CTpoeH~e

xnaM~AOMOHaAbl

8 - 3epHa KpaxMa-

na; 9 - KpaCHblC'i rna30K

6eJIKOBoe TeJibIJ,e B xpoMaTo-

cpope, KOTOpbIH ocy~eCTBJI.HeT cpMKCaIJ,HIO
yrJieKHCJioro ra3a M yqacTByeT B 3arracaHMH KpaxMaJra,

cmui.Ma (KpaCHbIH «rJia30K» B BM.n;e JIHH3hI, co.n;epJK~HH IIMrMeHT KapOTHHOH.n;) -

BOCIIpMHMMaIO~a.H CTPYKTypa, 6Jiaro.n;ap.H KOTOpo:H KJieTKa orrpe.n;e-

JI.HeT HaH6oJiee OCBe~eHHhIH yqaCTOK H rrepeMe~aeTC.H Ty.n;a
B3pocJia.H oco6b xJiaMM.ZJ;OMOHa.ZJ;hI rarrJioM.n;Ha

(n),

(rjJomomaKCUC).
(2n) TOJihKO

.n;MrrJioH.n;Ha

3HrOTa rrpM IIOJIOBOM pa3MHOJKeHMH.
U:mtJI pa3BHTHH 3eJieHhIX BO,!l;OpOCJieH
(Ha rrpHMepe yJioTpHKca)
YJioTpHKC -

rrpe.n;cTaBHTeJih HMTqaThIX Bo.n;opocJie:H, o6MTaeT s rrpecHhIX

BO.ZJ;OeMax; HMTb rrpHKpeIIJI.HeTCH K cy6cTpaTy O.ZJ;HOH 6ec11,BeTHOH 6a3aJihHOH
KJieTKOH (p1I30H.ZJ;OM).
B '1<H3HeHHOM 11,HKJie yJioTpMKca rrpeo6Jia.n;aeT rarrJioM.n;Haa cpa3a (n), .ZJ;MIIJIOH.ZJ;aa

(2n) -

TOJihKO 3HroTa. XapaKTepHa cMeHa 6ecrroJioro M rroJioBoro rrpo11,ecca.

ocy~eCTBJIHeTC.H

300CIIOpaMH (IIO.ZJ;BH)l{Hbie KJieTKlI c qe-

TblpbMH »<rYTMKaMM). IloJIOBOH rrpo11,ecc -

M30raMHblH (raMeThl IIOflBMJKHhI, TaK

BecrroJioe pa3MHOJKeHHe

KaK HMeIOT ITO .ZJ;Ba mryTMKa).

Paa.MnoJ1Cenue 3e.nenbtX BooopocJteu
1. BeiemamuBnoe pa3.MHOJ/Cenue (HHor.n;a Ha3blBaeTca 6ecrroJihIM): .n;eJiemie
KJieTKH Ha.n;Boe (y O.ZJ;HOKJieToqHhIX), o6phIB HHTH (crrHpOrHpa).

Bomauu1ea

220

2. Becno;we pa3M1toJ1Ceuue -

nocpe.n;cTBOM cnop. CnophI 6hrnmoT no,n;smKHhie

(3oocnopb1) H Heno,n;BIDKHhie (arnrnHOCITOphl, nepeHOCRTCR BO,ll;OH).

3. lloJtoBoe paJMHOJJCeHue CBR3aHo co CJIHRHHeM

raMeT:

cf'

11

<?. Bhl,n;eJIRIOT

pa3JIHqHbie THilhl TIOJIOBOrO npou;ecca:

1)

KoH'hIOrau;11R -

caMhIH npl1MHTI1BHhIH. IlpoTeKaeT aHaJior11qHo nucpy.

30pI1RM.

llpuMep: KJiacc Cu;ernurnoK -

noJIOBOH npou;ecc KOH'h10rau;11R.

Bo,n;opocJih cn11por11pa He HMeeT 6ecnoJioro pa3MHomeumi. IlpH 6JiarorrpnRTHhIX ycJIOBHRX HhIX yCJIOBIIRX -

BereTaTHBHOe pa3MHOllieHHe, npH He6JiaronpIIRT-

KOH'hlOI'aII;HR.

llpou,ecc IW/lo/Oiau,uu: a) KJieTKH KOH'hIOI'aThl o6pa3yIOT BhIPOCThI;
6) neperopo,n;Ktt Mem,n;y KJieTKaMH Hcqe3a10T; B) 06pa3yeTCR Konymi:u;ttOHHhIH KaHaJI; r) Bee co,n;epJKHMOe O,ll;HOH KJieTKH nepeTeKaeT B ,n;pyryIO;
,n;) 06pa3yeTCR ITOKORIII;aRCR 3HI'OTa, IlOKphITaR ITJIOTHO:H o6oJioqKOH.
IlocJie KOH'hIOI'aII;IIH, oceHhIO, HHTH BO,n;OpOCJIH onycKaIOTCR Ha ,ll;HO,
B IIJI. CTaphie HIITII crHIIBaIOT B II;HKJie cnnporttphI. TaJIJIOM ranJiott.n;HhIH

(n),

3III'OTa ,n;nnJioH.n;HaR

(2n).

BecHoii HaqlfHaeTcR Meiio3 MB KJieT-

Ke BOCCTaHaBJIIIBaeTCR ranJIOH,ll;HhlH Ha6op xpoMOCOM

(n).

TpH KJieTKH

OTMIIpaIOT, O,ll,Ha npopacTaeT B HOBYIO oco6b.

2)

faMeToraMMR -

pa3MHO;KeHMe B pe3yJihTaTe CJIMRHHR TIOJIOBhIX KJie-

TOK (raMeT):

+ .. ~ 2n -

H30I'aMHR:

n

reTeporaMHR:

n

+'

~

n

o6e raMeThl O,ll;HHaKOBhle H IlO,ll;BHJKHhie (xJiaMH,ll;OMOHa.n;a),

2n -

raMeThl pa3HOI'O pa3Mepa, IlO,ll;BHlliHbie,

n
~

n

2n
n

JKeHCKaR raMeTa KpynHaR, HeTIO,ll;BH;KHaJI
(xapOBhie Bo,n;opoCJIH -

BhICOKOOpraHII30-

BaHHhle).

B JKH3HeHHOM u;ttKJie so,n;opocJieH: B03MOmHa CMeHa noKoJieHIIH. HanpnMep,
yJtbBa (HJIH MopcKoii caJiaT) HMeeT cnopocpHT (2n) If raMeTocpnT (n), HO MopcpoJIOrlfqecKH OHH He OTJIIfqaJOTCJI U30MOp</Hta5l CMeHa llOIWJteHUU (,n;JIR cpaBHeHnR: nanopOTHHK - iemepo.lftoprjJHaR CMeHa llOICOJteHuil: crropocpHT HMeeT BH,11; KYCTa, raMeTocpHT -

3apOCTOK B BII,n;e MaJieHhKOH IIJiaCTHHKH).

OT,z:i;eJI Byp1>ie Bo,z:i;opocJIH
1500 BH,n;oB.

OT,n;eJI HacqlfTbIBaeT

Bee 6yphie Bo,n;opocJIH -

o6IITaTeJIH Mop-

CKHX Bo,n;oeMoB. XapaKTepHhie oco6eHHOCTH 6yphIX Bo,n;opocJieH::

1.

BypaR oKpacKa, rrHrMeHT -

2.

ABTOTpocphl. Co,n;epmaT xJiopocpHJIJI
H,ll;hI -

3.

rjJy1Co1CcaHmUH (KcaHmorjJuJtJt).

a,

XJIOpoqmJIJI

c, a

TaKme KapoTI1HO-

MaHHHT (rneCTHaTOMHhIH cm1pT).

XJioporrJiaCThl 3epHHCTOH HJIH ,ll;HCKOBH,ll;HOH cpopMhl; B KJieTKe IIX MHOro, MX Ha3hIBaIOT rjJeonJtacmbt.

orniJell- BypbLe 6000poc.riu. lloiJu,,apcm6o KpaCltbLe fJOOO]JOCJl,U

221

4. 3anacHh1e Beru,ecTBa: r.r110KaH, JiaMHHapHH (rrom1caxapu,n;), mHph1.
B KJieTKax OTCYTCTBYIOT nupettou.n;hi.
5. Pa3MHomeHue: 6ecnoJioe tt noJioBoe. 06a Btt.n;a pa3MHO:>t\eHHJI CO'IeTaIOTCJI B )KH3HeHHOM l1HKJie.
reTepoMopq:rnaJI, c qepe.n;oBamrnM .n;ttnJI011.n;noii:
11 ranJI011.n;Hoii cpaa.

6. CMeHa noKoJieH11ii -

QttKJI paaBHTHH ByphIX Bo,11;opocJieii
(Ha npttMepe JiaMttHaptttt) (pttc. 88, 89)

B m11aHeHHOM I1HKJie npeo6Jia.n;aeT 6ecrroJioe noKoJiem1e -- crropocpttT,
npe,n;cTaBJieHHhlii TaJIJIOMOM. CrropocpttT - .n;nnJiott.n;Hoe noKoJieHue (2n).
BecnoJioe paaMHomeune rrponcxo,n;HT c rroMOW,hIO ,ll,Byx mryTHKOBhlX aoocnop (n). 3oocnophI o6paayIOTCJI B peayJihTaTe Meiioaa n paaB11BaIOTCJI B raMeTo<}>HThl: meHCKHii raMeTocpHT - oorottnii H MymcKoii - aHTepn,n;uii. faMeTo<l>HThI (n) 06paay10T Riil1eKJieTKy (n) 11 no.n;B11.1Rnh1ii cnepMaToao11.u (n), rrp11 orrJ10,n;oTBopen1111 cpopM11pyeTCJI anroTa (2n).
IloJIOBOll IlP01leCC - reTeporaMHJ!.
311r0Ta pa3BHBaeTcJI B crropocpHT (2n).
OT,!J;eJI KpacHLJe BO,!J;OpocJIH, HJIH Barp.RHKH

.,'

HacqnThIBaeT OKOJIO 4 ThICJiq B11,n;oB. KpacHhle Bo.n;opocJIH - 0611TaTeJI11
MOPCKHX BO,ll,OeMOB, JIHillb HeKOTOpbie o6nTaIOT B npeCHOH BO,ll,e 11 noqBe.
XapaKTepnbie oco6ennocTn:
1. CJioeBnru,e MnoroKJieToqnoe, peme o.n;noKJieToqnoe, paaMepb1 OT M11KpocKonnqecKHX ,ll,O 1-2 M, TaJIJIOM qaru,e B BH,D;e KyCTHKOB.
2. CaMhle r.11y60KOBO,D;Hhie Bo.n;opoc.11n, TaI< KaK HMeIOT oco6hie nnrMeHThI.
QBeT BO,n;opocJieii: KpaCHblll, cpnoJieTOBblll, 6op.n;OBhlll, :lKeJIThIH, roJiy6oBaT0-3eJieHbIH; 3aBHCHT OT coqeTaHHJ!
llHrMeHTOB. IlHrMeHThI: XJIOpoqmJIJibl
a, b, d; <jJUK06UJlU1tbl (<jJUKOU,UUftUftbl CHHHe, <jJUK03pumputtbl KpaCHbie),
KCaHTOcpHJIJihl H .n;pyr11e KapOTHHOH)l;hl.
XpoMaTocpophI aepHHCThie, .n;ncKoo6paanhJe, nnpenon.n;oB neT.
3. 3arracHhle Bew,ecTBa, 06paay10w,necJI
B npo11ecce cpoTocnnTeaa: «6azp.HHK06btu KpaxMaJt», no cTpoennro 6oJiee
cxo,n;en c rJIHKOrenoM, qeM c KpaxMa2
1
JIOM.
PMC. 88. 6ypb1e BOAOPOCJWI: 1 - ca4. PaaMnomenne: BereTaTHBHOe - rroqxapHaR
naMl'1Hapl'1R ( Laminaria sacchariKOBanne, IIOJIOBOe (nOJIOBOll rrpona); 2 - cesepHaR naMl'1Hapl'1R ( Laminaria
11ecc - ooraMHJI). BoJihIDHHCTBO Kpachyperborea)
Hhlx BO,D;OpOCJieii ,ll,By,n;OMHhie pacTeHHJ!.

Bomauu1ea

Cnopo4>HT ( 2n)
/(()ccU0,1iLOCTlllllf")

MoooJJ:oA cnopo<jiITT

~
3oocnopaimm ( 2n )

I

3wora ( 2n)

2f'I - J\MMOitJJ,HlJI ~HSI
\

n-fi\fl.~OU!ltlUCTt.,a,~R

\

P111c. 89. CxeMa lK1!13HeHHOro u.1111<J1a 6ypoi't so,a,opocn111 naM111Hap111111
Ha raMeTocpHTax (n) o6paayroTCH My:>i<e1rne H meHcKHe raMeTbI 6ea mryn1KOB. IlpH OilJIO,ll,OTBopeHHH meHCKHe raMeThl He BhlXO,lJ.HT B OKpyx<a10rn;y10 cpep,y, a ocTalOTCH Ha pacTeHHH. My:m.c1u-1e raMeThI naccHBHO nepeHOC51TC.H TOKOM
BO,!J.hl. Ilocne onno,lJ.OTBopemrn H o6pa3oBamrn 3HrOThl (2n) cpopMHpyeTc.a cnopoc}>HT (2n). Ha cnopocpHTax 06pa3y10Tc.a opraHbl 6ecnonoro pa3MHOJKeHH.H cnopaHrHH.
5. lfaoMopcpHaH CMeHa IlOKOJieHHH.
Ilpe,1.1;cTaBHTeJIH 6arpaHKOBbJX: a1ujJeJlb1"UJl (HCilOJih3yeTCH ,lJ.JISf noJiyqemrn
arap-arapa), poau.MeHUJl, noprj>upa.
3Ha"!em1e BOJJ;OPOCJieii::
1. Ilpo,!!.yU,eHTbl B BO,ll,HhlX coo6rn;ecTBaX (mnu,a ,II.JUI MHOrHX )KHBOTHhlX).
2. Bbl,ll,eJIHJOT I<Hcnopo,11., HCilOJihayeMhIR ,ll,JIH AhIXaHHH ,1I,pyrHMH opraHH3M8MH.
3. Bh13bIBaIOT u~BeTeHHe» BO,!J;hl.
4. MHKpocI<onH"leCKHe I<pacHhJe BOAOPOCJIH BhI,ll,eJIHIOT Bern;ecTBa, ry6HTeJihHhle ,ll,JIH )f<liBOTHhlX.
5. flBJIHIOTcn CHM6HoHTaMH c rpn6aMH rrpH o6pa3oBaHHH JIHIIJaHHHKOB.
6. YnoTpe6.rraIOTCH B nHrn;y 'leJioBeKOM (JJaMHHapna, nopcpnpa).
7. HcrroJJh3YIOTCH ,ll,JJH rrony-qemui: arap-arapa (HcnonbayeTca: B KOH,ll,HTepCKOM npOH3BO,ll,CTBe - mene, nacTHJia JI )J.p., ,!J.JIH npli:rOTOBJieHH.H MHKpo6HOJIOrH"leCKHX cpe,!I.).
8. licnonh3YIOTCK ,ll,JIK noJiy-qeHHH ii:o,11.co,11.epmarn;Hx Me)J;IH~HHCKHX npenapaToB.
9. licnOJJh3YIOTCH B O'lHCTKe CTO"t!HhlX BO,ll, (XJJaMHAOMOHa,!J.a).

223

noal.fapcm60 Bbicmue cnopo6bie pacmenuR

Ilo;::i;n;apcTBO BbICWHe crropOBbie pacTeHHH

K

BbICIIIHM crropoBbIM pacTeHHHM OTHOCHTC.fl:

6uiJHbte, nAay1to6UOflbte

11

.woxo6u01tbU'. nanopom 1tww-

xeow,e6UOflbte.

OT;::i;eJI MoxoBH.!J:Hbie
OTp;eJI MoxoBHP:Hbie HacqttTbIBaeT

24-35 Tbrcaq BH,ll;OB.

8TOT rrpttMHTHBHhI:ii

oTp;eJI BblCIIIHX pacTeHHM He BCer.n;a COOTBeTCTByeT xapaKTepHCTHKe BblCIIIHX.
,I(JIH BbICIIIHX pacTeHHM xapaKTepHbI:

1.
2.
3.

BereTaTHBHbie opraHhI -

KopeHb, cTe6eJib, JIHCT.

,I(mp¢>epeHii;ttpoBaHHbie TKaHH (.n;o

85

BH,ll;OB pa3JJHqHbIX TKaHe:ii).

IloJIOBbJe opraHbl MHOrOKJieToqHbie.

,I(mi: MOXOBH,ll;HhIX xapaKTepHo:

1.

BHeIIIHee CTpOeHtte: CTe6eJib eCTb Hey Bcex, eCTh JIHCTbH, HO HeT KOpHeiL
<t>yHKIJ;HIO KOpHel:i: BbIIIOJIHHIOT BbipOCTbl KJieTOK HHJKHe:ii qacTH CTe6-

pu30UObt.

JI.fl -

2.

B aHaToM11qecKOM cTpoeHHH o6HapymttBaeTCH oTcyTCTBHe BhICOKocrreri;ttaJIH3HpoBaHHoii cocy.n;ttCTO-IIPOBO,ll;aIIJ;el:i: CHCTeMbl.

y

Hatt60Jiee BbICO-

KOOpraHH30BaHHhIX JIHCTBeHHbIX MXOB IIOHBJIHIOTCH TpaxeH,ll;hl I1 CHTOBH,ll;HhJe Tpy6KH. Mxtt He MOryT IIO,ll;HHMaTb BO,ll;y Ha 60JihIIIHe paCCTOHHHH I1 BCer.n:a HI13KOpOCJibie.

3.

IloJIOBOH rrpori;ecc CBH3aH c BO,ll;OH, qTo yKa3hIBaeT Ha CBH3h MXOB I1 BO,ll;OpocJieH (rrpe.n;KaMH MOXOBH,ll;HblX HBJIHJIHCh 6ypb!e BO,ll;OpOCJIH). IloJIOBbie
opraHhI -

MHOrOKJieToqHbJe: MymcKHe -

aHTeptt.n;Htt, meHCKHe -

ap-

xeroHHH.

4.

Oco6eHHOCTh JKH3HeHHoro IJ;HKJia Boe -

raMeTo¢>ttT, 6ecrroJioe -

qepe.n;oBaHtte rroKoJieHttii:: rroJio-

cn;opo¢>HT (rrpoH3BO,ll;HT crrophr). IIo-

JIOBoe rroKoJieHHe 06ecrreq11BaeT H3MeHqHBOCTh, 6ecrroJioe -

pacceJie-

HHe.

QHKJI pa3BHTHH MOXOBHJJ;HLIX
(Ha rrpttMepe KYKYIIIKHHa JlbHa) (pttc.

90)

B JKH3HeHHOM IJ;HKJie KYKYIIIKHHa JihHa rrpeo6Jia.n;aeT rroJioBoe, rarrJioH.n;Hoe (n) IIOKOJieHHe. 3eJieHbie pacTeHHH rrpe.n;cTaBJIHIOT co6oii: IIOJIOBOe
llOKOJieHHe n.

IIpe.n;cTaBHTeJitt:

MapIIIaHIJ;HH, c¢>arHyM,

KYKYIIIKHH JieH

H ,n;p.
raMeThI o6pa3yJOTCH B rrpori;ecce MHT03a. Crrophl o6pa3yJOTCH B rrpori;ecce
Mel:i:o3a.
MxH -

Ha

,n;By.n;oMHhie pacTeHHH. faMeTo¢>HThI MXOB MHoroJieTHtte pacTeHHH.

MYJKCKOM pacTeHHH cpe.n;tt BepxHHX po3oBaThIX JIHCToqKoB pa3BHBa-

lOTCH MeIIIKOBH,ZJ;Hhre MymcKHe opraHhI -

aHTeptt.n;ttH. CrrepMareHHaH rarrJio-

R,ZJ;HaH TKaHh aHTepM.n;HeB .n;eJIHTCH MHT030M, o6pa3ya crrepMaTo3ott.n;hr. CrrepMaT030H,ZJ;bl rro,n;BttJKHhI, ttMeJOT .n;Ba :m:ryTHKa. Becttol:i: crrepMaT030H,ll;hI BhIXO,Zl;aT B Bo.n;y.

224

/'~d.:k~
3HnJTa (

2n )

MarepHHCXllC KnC'JJW

cnop (2n)

C,,<>P.4{n) ·..

5lRLteMCncii(n) . CriepMajoooHA(il) ...

~~;;~,~(~
~- ~~~~:~~~~~~~~~~}{

'

~~

·.

r.~~n.10~,1ttaH_<"rn,~~>1. l

Pttc. 90.

CxeMa lKV13HeHHoro 1..1,111Kna MOXOBlllAHblX

:iKeHCKHC paCTCHHR HMeIOT CBeTJI0-3eJieHhie BepXHHe JIHCTQqKH, cpe,n;H HHX
paasnBaIOTCH 1wn6oqKOBH,D;Hhle apxeroHHH, rrpo,n;yu;Hpyrom;n:e Riin;eKJieTKy.
Bo BpeMH .n;o.m.n;R crrepMaT030H,n; rra.n;aeT Ha .meHCKHH raMeTo<fJnT n rrpOHHKaeT
B Riiu;eRJieTKy: q>OpMnpyeTC.H 3llI'OTa (2n). qepe3 HeKOTopoe BpeMH 3HrOTa (2n)
HaqnHaeT .D;CJIHThC.H H o6paayeTC.H ,D;HIIJlOH,nHhIM crropoqmT (n) Ha .meHCKOM pacTeHHH-raMeTO$HTe (n). Crropo$nT HMeeT BH,n; Ropo6oqKH Ha Ho.mKe. BHyTpH
Kopo6oqRn: apxecnopuaJtbftaR mKaHb rro,n;BepraeTCH Meiioay n o6pa3yJOTCH rarrJIOH,D;Hhie crrophl (n). Crrophr rrorra.n;aroT B rroqBy n: rrpopacTaIOT B rrpoToHeMy MHOrOKJleToqHyIO aeJieHyIO BeTBHID;YIOCH HHTh. 3aTeM Ha Heii <PopMHPYIOTC.H
Jin6o My.mcKHe, Jin:6o .meHCKHe rroqeq1nr. IloqKn rrpopacTaIOT B raMeTo$HThI.
CnophI reTepoTannnqHh!e. ll3 rrpoToHeMhl <PopMnpyeTCH raMeTo$nT.

~eJI MoxoBMA~
K;rncc IIe,1eno'IHhle MXH
(Mapuranu;m1)

KJiacc JiucTocTe6e.JihHbie MXH

~

/

HeT cTe6Jieii: M JIMCTbeB,
IIOJIOBble opraHhI Ha
(< IIO,ll;CTaBKax ~

n 00/CJtacc

lloiJ1C.rtacc

II oiJ ICJtacc

3eJteHbte

Be.llbte

AniJpeeBbte

(KYKYIIIKHH
JieII)

(clf>arHyM)

(HeMHOrO'IHCJieHHbie)

225

omiJ eJi n anopomnu1wBuiJnbu'
J\.Jiacc .J1HCTOCTe6eJILHble MXH, rro.ri;KJiacc EeJILie MXH. CcparHyM

(pffC. 91)
XapaKTepHbJe np113HaK11:
1. He 11MeeT p113011;:i;os.
2. Bee BpeM.H HapacTaeT nepxyrnKOM cTe6JI.H, HMJKH.H.H qacTb oTMMpaeT. JKeHCKHe apxeroHHH H MYll<CKMe aHTepH,ll;HH o6pa3yIOTC.H Ha O,ll;HOM pacTeHHH - 0,ll;HO,ll;OMHbie.
3. 0TJIJ1qaeTC.H CTpOeHMeM JIHCTa.
JlMCTb.H O,ll;HOCJIOMHhie, COCTO.HT
113 KJieTOK ,u;Byx THIIOB: 3eJieHbie, MeJIKMe c XJIOpOIIJiaCTaMH; rttaJIHHOBbie, 6ecu;BeTHbie,

.

a

6

PMc. 91. Ccj:JarHyM:

a-

K!leTO'lHOe CTpoeH111e.

1-

06L1.(111Vi s111A:

6 -

<j>OTOClllHTe3111pyto-

MepTBhie, rryCTbJe, c KOJibqall1lile K!leTKlll; 2- nyCTble KlleTKlll
ThIM yTomu;eHtteM o6oJioqeK
(o6oJioqKM MMeIOT RpyrrHbJe rrophr).
3HaqeHHe MXOB: o6pa30BaHMe ccparHOBbIX 6oJIOT; TOpcpoo6pa30BaHMe
(Top<l> - TOIIJIMBO, y,u;o6peHHe). He,u;ocTaToK KMCJiopo,u;a Ha 6oJioTax rrpttBo,n;HT K HerroJIHOMY pa3JIO/KeHIHO OTMepIIIMX TKaHeii; 06pa3yeTC.H TOpcp (1 CM 33
10 JieT).
OT,ri;eJI IIarropOTHHIWBH.IJ;Hbie

IIarropoTHHKH, IIJiayHhI H XBOiu;H MMeIOT p.H,u; oco6eHHOCTett, no KOTOpblM HX
paHee o6'be,n;MH.HJIM B o,u;Hy rpyrrrry - rrarropoTHHKOo6pa3Hhre. OT;:i;e;:r HaC'IHThIBaeT 6oJiee 12 TbIC.H'I BH,ll;OB.
:IKtt3HeHHbie cpopMbI pa3Hoo6pa3HhI. EcTb pacTeH1u1 TpaB.HHHCTbre 11 ,u;peBoBH,ll;Hhle (B Tpomrnax).
Bueumee cmpoeuue (p11c. 92) .
.HBJI.HIOTC.H HaCTO.Hiu;HMH BbICIIIHMH pacTeHM.HMM: MMeIOT CTe6JIH, JIHCTb.H
(Balitt), KOpHeBttru;a c rrpu,u;aToqHhIMM KOPH.HMM. JlttCT rrarropoTHttKa (naii:.H)
KpynHbitt (MaKpoc}mJIJI), ttMeeT cTe6JieBoe npottcxom.n;eHHe. JlttcT o6hrquoro
pacTeHHa pacTeT 3a cqeT MepHCTeMbl y OCHOBaHH.H JIHCTOBOH nJiaCTllHKll. JlHCT
nanopoTmrna yJI11T1wo6pa3HO 3aKpyqeH H pacTeT Bepxyumoii, r.n;e ttaxo,n;MTC.H
MepttcTeMa. Jltt:CT rrarropOTHMKa (Batt.fl) MOmeT tt:MeTb pa3HYIO c}>OpMy, qTO HCllOJib3yeTc.H B cttcTeMaTMKe pacTeHttii. PacqJieHeHHOCTb JIMCTOBOM rrJiaCTMHKH
MomeT 6bITh BToporo, TpeThero nop.H,u;Ka.
B auamoMU'leCl(O.M cmpoeuuu o6HapymttBaeTc.H BHYTpeHH.H.H .n;11<lJcpepeHu;MPoBKa TKaHeii. Xoporno pa3BHTa rrpoBo,n;.Hru;a.H cttcTeMa cTe6JI.H M KopH.H. EcTh ·
aacToaru;tte KOpHtt. IIarropoTHMKM - rrepBbie cocy,u;ttCTbre pacTeH11a. 8To pacTeatta, y KOTOpbIX eCTb cocy,u;ttCTaa, HJIH rrpoBo,u;aru;aa, TK8Hb, T. e. TK8HH KCHJieMbI H cpJIOaMbI.

226

lloJt06bl(' opiattbl, KaK l1 y BCCX BbICllIMX, - MHOrOKJieToqHbie. I1MeeTca riepe.n;oBaHHe IIOKOJICHMH B fKI13HeHHOM r.vm:re. flpeoforn.n;aIOw;ee TIOKOJieHile ~
cnopoqmT (2n). 3eJieHhre pacTeHtta - 6ecrro:we rroKoneune, cnopocp11T (2n).
faMeToqrnT (n) B Btt.n;e cep.n;n,em1,n;HOH TIJiaCTIIHKll. Tipo.zwmKHTeJibHOCTh ero
:>KH3HH - o.n;ttH BereTa:u;110HHhrii nep110.n;. CrrepMaTo3ott,n:hI - 4-mrynmoBhie.
floqTH Bee rranopoTHIIKM - paeuocnopoebte pacmettu5l. Pa3HocnopoBhIMH
HBJIHIOTCH BOil;Hhie narropOTHHKH, HanpHMep caJihBHHHH TIJiaBaIOr.u;aa.
Mymcx11e JI meHcKHe raMeTocjJHThI, HJIH 3apoCTKH, pa3Birna10Tca H3 MHKpon Meracrrop, KOTOphle o6pa3yIOTCJI B MHKpO- H MeracrropaHnrnx Ha HH)KHCII 110BepXHOCTH JIHCTOBOM IlJiaCTHHKll. faMeTOcpllThI o6pa3yIOT cf' ll
raMeThl. I1oc.1e
OIIJIO,IJ;oTBOpCHHH pa3BirnaeTCH 3HI'OTa, ll3 3HrOThI - MHOI'OJieTHllll crropocpHT.

9

6

7

P111c. 92. W,11TOBHl1K MY)l(CKOiil: a - cnopocjl11T; 6 - YaCTb saiil11 c copycaM11: a - nonepeYHbliil cpe3
Yepe3 COpyc; r- CnopaHr11H; .0. - cnopa; e - MOllOAOiil raMeTOcjll1T; !K - 3p61lblH raMeTOcjll1T-3apOCTOK;
3 -

aHrnp11A11iil;

copyca): 3 aHrnp11A11iil;

111 -

apxeroH11iil;

cnopaHr11iil; 4 -

8 - p113011Abl

K --

MOllOAOiil cnopocjl11T. 1 - nnaueHTa; 2 -

HO>KKa cnopaHrnsi; 5 -

11H,1:iy311ii1 (n0Kpb1sa11bl..!e

KOllbU.O YTOlllL\eH11si; 6 -

apxeroH11iil; 7 -

....-au1'ft pa36umuH nanopomttUIW6

227

B Do,n;MOCKOBhe Hatt6o;:iee l.J.8CTO BCTpe<raIOTCH rrnrropoTHttKtt ru;ttTOBHIIK
»{CKOH - paccel.J.eHHOCTh .1l:ICTa ,LIO BTOporo rropH,n;Ka, JIHCT OBaJ1bHbIH; rra1111 Y
..
nopOTHHK JKeHCKHH paccel.J.CHHOCTh ,ll;O TpeThero rropa,11,Ka; op.1HK TpeXCJIOJKHh!M J1HCTOM, rrpon3pacTaeT B COCHOBhIX Jiecax; rrarropOTHIIK
c
roJib'-I8Thlll paccel.J.eHHOCTh ,11,0 TpeThero nopa.a;Ka, m1cT TpeyroJ1hHoii:
11
cpop.llllbI.
nanopOTHHK IIJ;HTOBHHK MymcKoli ucrroJih3yeTCH B Me.a;nu:nne (rm1cToronu

808 cpe,D;CTBO).
l~HKJI pa3BHTHH rrarropoTHHKOB
(na np11Mepe rrarropoTHHKa IIJ;HTOBHHK MymcKoi1:) (pnc. 93)

B3poc;roe pacTenne HBJIHeTCH crropoqmToM (2n) - 3TO TpaBHHHCToe pacTeane c xoporuo pa3BHThIM KOpOTKHM H TOJICThIM KOpHeBHIIJ;eM c npn.a;aTO'IHhIMH
KOPBHMH. OT KopneBttru;a oTxo.a;aT Kpyrrnhre 3eJieHhre Ba:i1:1:1. CcpopMHpoBaBrunncR JIHCT ,l\OCTHraeT ,/l,JUIHhI 1,5 M.
JleToM na n1:1mne:H: cTopone Bali1:1 06pa3y10TcH copycbt - co6pamrn crropanrneB. B crropanr1:1ax rrpo1:1cxo,n;1:1T Meiio3 1:1 o6pa3yIOTCH rarrJiou.a;nhre crrophr (n).
CnophI KJieTKH 6ecrroJioro pa3Mttomen1:1a. 1Ip1:1 co3peBan1:11:1 crropanru1:1
BCKpbIB8IOTCH H crrophr BhIChIIIaIOTCH. ll3 rarrJIOH,ll;HhIX (n) crrop o6pa3yIOTCH
ranJIOH,ll;Hbie 3apoCTKH (n), HJIH raMeTO$HThl (IIOJIOBOe IIOKOJieHne). 3apocmmc - cep,n;u:eBHHHaH rrJiaCTHHOl.J.Ka pa3MepoM c O,l\HOKOIIeeqnyro MOHeTy' 38JieHoro QBeTa. Ha nnmnen cTopone 3apoCTKa - pH30H.L\hI. Ha 3apocTKe o6paayIOTCH apxerOHHH (n) B o6JiaCTH BhICMKH H aHTepH,ll;HH (n) B 06J1aCTH PH30H,l\OB. OrrJIO,/l,OTBOpen1:1e npottcxo,n;HT np1:1 HaJIH'IHH KaneJihHO-JKH.a;Koi:i cpe.a;hr.
CrrepMaT030H.L\hI rro,n;rrJihIBaroT K apxeroHHHM H CJIHBaIOTCH c a:H:u:eKJieTKoii:.
CoopotJrr ( 2n)

/(6ecoarrax~~
3aP.<JJ(hllll ( 2n )

I

3HrOTa ( 2n )

Copyc1>1( 2n )

'\

CnopllHrHH ( 2n )

Ma-repHlitKHe JOleTIOI
coop (2n)

P111c. 93. CxeMa )Kl-13HeHHOro u,1-1Kna nanoporn1-1Ka w,1-1TOBH1-1Ka My)KcKoro

228

Boma11u1'a

113 311.rOTbl 06pa3yeTC.fl 3apO,LihllU (211). ~fapO,l(blfll CBJI3aH c raMCTO<jrnTOM, lIORa
He o6pa3yJOTCJI KOpHH, 3aTeM CTaHOBHTCJI caMOCT05lTCJ1bHbIM.

OT.z:i;eJ1 IIJiayHoBH,J;Hh1e
IInayHOBl1L\Hhie -- O.[\Ha l1a tta116o;ree ;lpestt11x rpynrr pacTeH11i1. HaC411ThIBaJOT OK0.'10

200

BH.!J:OB IlJiaYHOB. CospeMeHHbie mrnyHbl -- 3TO MHOI'OJ1eTHl1e Tpa-

BJIHHCTbie B€'lH03eJICHb1C pacTeHl1.fl, TPOIIIfl-IeCKHe BH.[\bl B OCHOBHOM 3IllIQJl1TbI.

Bnewuee cmpoe11ue:

no6eru nonayqu:e, neTBJieHHe .n;u:xoToM11qecKoe IlJll!

Bl1J1bqaToe (caMhlll )J,pCBHHi't THIT BCTBJJeHHJ!). JlHCTbJI qemyeBHL\Hhie M8JIKHe,
MHKpoctm:nn11a, cn11paJ1huo pacnoJroJKeHHhie na no6erax. CTe6;rn cTemoru,11eca
no 3€MJ1e n

npHIIO.!J:HHMa10ru;11ec.a. fIJJayHbI HMCIOT II0/~3eMHOe KOpHesmu;e

H rrpu.n;aToqHbIC KOPHH.
l106er11 3aitanqnBalOTCJI: crroponoCHbIMH KOJ"lOCKaMH (pnc.
JJOCIW ,n.nrr.i:ro11,1vrh1

(2n).

94),

see TKaHI1 KO-

Ha crropo¢iurax (n11cToqKax) Haxo.n;JI:TCJI: cnopaHnrn.

B HHX npoHcxo/~HT Meii:o3 H o6pa3yIOTCJI: rarrJJOHL\Hhre cnopbr (n). Crropb1 ona.n;afOT Ha 3eMJ1IO. IlpopacTaIOT cnopb1 qepe3 IIJ!Th neT, ,n.aBaJI: Haqano o6oenoJihIM raMeToqn1TaM. faMeToqm:T y rrJJayHOB pa3MepoM

0,5

CM, He HMeeT xnopoqnvma.

JJ.o rrepsoro OII.TIO,Ll,OTBOpcmrn 11o6pa30BaHHJI:3apOJ(blllia rrpOXOJ(llT OKOJIO 22 JieT.
faMeToc}mT cyru;ecTByeT TOJihKO B cnM6Ho3e c rpu6aMH. CrrepMaT030II,Ll,bI JlBYXmryTHKOBble, o6pa3y!OTCJ! B aHTepII.n;HHX; JI:MIJ;eKJieTKII -- B apxerOHIIJI:X, TaM me rrpOI1CX0AlIT OilJJO,Ll,OTBOpe-

lfa 3IIrOThI (2n) 06pa3yeTCJI: crropoc}JHT (2n).
B auamoMU'iPCKO.M cmpoeHuu Ha norrepeqHoM

HHe.

cpe-

3e CTe6"1H 06Hapym11Ba10TC.a CTPYKTyphr, HanoMHHarouvre KOHUeHTp11qecKI1H cocy,n.MCTO-BOJJOKHJICThIH rry'lOK Il KOpy. 11poBO,ll;HUJ;l1e 3JieMeHTbl KCHJJeMbl xeHJ(hl. 3:rnMeHThI c}JJI03Mhl -

Tpa-

CIITOBIIJ(Hble Tpy6KM.

IlnaynoBH,ll;Hbie HMelOT paBHOCITOpOBbie 11 pa3HOcrropOBbJe c}JopMhr.
U:mm pa3BHTH}{ IIJiaYHOBHJJ;HhIX
Ha rrpHMepe miaytta 6yJiaBOBH)J;HOrO

Cnopoc}JHT rrnayHa

2

3eMHoe

KOpHeBHII~e

(2n)
c

HMeeT seTBJI:ru;eecJI rroA-

rrpH.n;aTO'lHbIMM

KOpHHMlI'

OT Hero OTXO.[\HT Ha.[\3CMHhie no6ern. CTe6JJH IIOJI3YqIIe, c ,ll;HXOTOMIIqecKHM BeTBJICHIIeM, BhICOTOH ):\0

25

CM. JlHCThJI: Me.i'lKHe, qeruyii:qaThie, paCIIOJIOJKeHbl

no crrHpaJJ11. BepxyrrreqHbre no6er11 3aKaHqIIsaroTcH
PMC.

94. npoAOllbHb!Ci

BepxyrneqHblMU noqKaMH lIJIH cnopoHOCHhIMII KOJIO-

pa3pe3 cnopoHOCHoro

CKaMH -

KO/lOCKa nnayHa: 1 -

pacrr0Jrara10Tcn crreu;uanH3HponaHHhie JJHCThJI:

OCb KOllOCKa; 2 - MV1KpO-

crropoc}J11JJ.nhl, menToro u;BeTa, c rnIIpOKHM ocnosaHH-

cnopo<jl1111111bl; 3 -

eM II 3aOCTpeHHOH BepxyrnKOH.

paHrnCi co cnopaM111

cno-

cTpo6IIJiaMtt.

Ha crropoHOCHOM KonocKe

B

-

OCHOBaHIIH crropo-

c}J1rnna ecTh cnopaHr11M:, B KOTOpoM 06pa3yroTCJl cno-

229

bl o.zr.HHaKOBbie no cpop'.'.1e l:! BeJ1wrnHe. I1oc.1e npopacTaHHH cnopbr IIJiayirn

pa~T

HaqaJIO 06oeno.1bIM 3apOCTl<aM. faMeTOcpHT (n) -- HCcpOTOCHHTe3ttpyIO-

~Hil: MHOrOKJICTOqHblM

3apOCTOK, BCTynaeT B CHM61I03 c noqseHHbIMH rpH6a-

ld'.H· Ba 3 apocTIW o6pa3yIOTCH attTepH,LIHH (My:>KCKHe no.10Bb1e oprattb1) u apxeroHHH (mencKHe IIOJ10BhIC opratthr). B attTepII,zr,HHX o6pa3yIOTCH .zr.syxmryJIJCOBbie crrepMaT030II}J,bl, B apxerOHH.HX -

~JIJIO,LJ;OTBOpeHII.H

IIO,ll,BH:>KHble Hihi;eK.1eTKII. floc:1e

06pa3yeTC.H 3I1rOTa, II3 KOTOpoii: cpopMIIpyeTCH B3poc.1oe

pacTettHe-cnopocpIIT.
3Haqemrn rnrayHoB. CnophI rnrayHa 6yJiaBOBII.n;Horo -

mrnono,11:i1ii: -

MOJK-

110 HCII0Jlb30BaTb KaK JJ,eTCKYIO npllCbIIIKy; IIJiayn 6apaHeri: (cpapMaKoneii:ttoe
pacTeHHe) -

KaK aHTHaJIKoro.11bHOe cpe.n;cTBO. HaH6oJiee II3BeCTHhre npe.zr.cTa-

BHTeJUr OT)l;eJia I1J1ayHOBII)l;HbIX: n.1ayH 6yJiaBOBI1)l;HbIM (Ha sepxy BepTHKaJibsoro no6era 6era ra -

.n;sa KOJIOCKa), nJiayn ropeqHhIH (Ha sepxy sepTIIKaJibHoro no-

O)l;1IH KOJIOCOK), IIJiayH CIIJIIOCHYTbIM (Ha Bepxy BepTIIKaJibHOl'O no6eTj)I1 II 60.Jiee KOJIOCKOB).

0TAeJI Xuo:m;eBHAHLie
OT.n;e.JI npe.n;cTaBJieH e,zr,IIHCTBeHHhIM po.n;oM, BKJI10qa10ru;HM
XBOru;H -

15

sn.n;os.

Bee

MHOrOJieTHHe TpaBHHHCTbie pacTeHM.H, npOM3pacTaIOT BO B.lJa:>KHh!X

HJIM 3a60JioqeHHhrx Jiecax.

Bnewuee cmpoeuue:

IIMeIOT KopttesIIru;a M npM.n;aToqHbie KOPHII. CTe6eJih

qJieHHCTbIH c MyTosqaTO OTXO.n;Hru;IIMII OT Y3JIOB BeTB.HMH. JlIICTb.H Me.JIKHe
H qeruyeBMJJ.Hbie.

B

KJieTKax XBOru;eii OTKJiaJJ,hIBaIOTC.H KpMCTaJI.'IbI KpeMHC3C-

Ma, qTo npM.n;aeT pacTeHIIIO mecTKOCTh. Xsoru; 06pa3yeT .n;sa TMna no6erou (M3
noqeK KOpHeBMru;a): BCCHOH -

3mUOJlUp06llliHbte

(HecpOTOCIIHTC3Mpyroru;He)

no6erH, KOTOpbre .HBJIHIOTC.H cnopOHOCHbIMH; JICTOM -

accuMuJtupy10w,ue.
Auam0Mu 11ec1we cmpoeuue:

3eJieHbie cPOTOCI1HTe3H-

pyroru;He,

Ha nonepeqHoM cpe3e cTe6JI.H BII.n;Ha ri:eHTpa.rrh-

aaa IlOJIOCTb II HeCKOJibKO rrepHcpepHqecKHX B03JJ.YXOHOCHbIX IIOJIOCTCH. IlpoBO,ll;HID;Me 3JieMeHTbI KCHJICMbl -

TpaxeM,ll;bl, cocy.n;hI. 8JieMeHTbI cpJI03Mbl -

CH-

TOBH}J,Hble Tpy6KH, napeHXHMHbie KJICTKH.
Xsoru;esM.n;Hhre -

pasHocnoposbre pacTeHHH.

lI;HKJI pa3BHTHJI XBOiu;eii Ha rrpHMepe XBOiu;a IIOJieBoro
CnopocpHT xsoru;a

(2n)

HMeeT no.n;3eMHOe KOpHeBHru;e, OT KOToporo OTXO-

,ll;HT npM.zr.aToqHhie KOpHH. Ha Kopttes11ru;ax tteKOTOphrx xsoru;eii: o6pa3yIOTCH
KJiy6HII,

KOTOpbre npe.n;cTaBJI.HIOT co6oii: YTOJiru;eHHOe H

BH)l;OII3MeHeHHOC

Mem,n;oy3JIHe. KJieTKM KJiy6He:ii Kpyrrnee M 3anoJIHeHhI KpaxMa.JihHbIMH rpaHYJiaMM. OT ropM30HTaJibHhIX KOpHesttru; sepTHKaJibHO oTxo.n;JJT Ha.zr.3eMHbre
no6ern.

HaJJ,3eMHhre no6eru o.n;HoJieTHHe,

rro.n;3eMHhre

-

MHoroJieTHMe.

Ha no6erax BH)l;Hhr y3JibI H BhITJJHYThie Mem.n;oy3.JIIIH. OT y3JIOB OTXO.D;HT MYTOB'IaTo pacrroJiomeHHhie 6oKOBhie BeToqKH. JlHCThJI xsoru;eBM)l;HbIX pe.n;yri:HPOBaHbI ,zr,o qeIIIyeK. CrropoHOCHhre KOJIOCKH (crropocpnThr -

2n)

nMeIOT pac-

llIMpeHHyIO IIOJIYIO OCh, Ha Heii HaXO)l;JITCH crropOJII1CTI1KH IIICCTHyro.JihHOH

230

Bomanu1<.a

cpopMbI. Ilo.n HIIMII cilopMIIpyeTCH Meruo'!eK - cnopaHrui!:. 3,nech npoIIcxo,ll;IIT Metto3 II 06pa3y10Tca cnopbr (n). Cnopbr HMeIQT 3JiaTepbr (JieHTbr). 3JiaTepbr rrptt C03peBaHHH crrneTaIQTCfl rpynnaMU, CIIOpbI BblCbIIIaJOTCfl II rrpopacTaIQT B 3apoCTOK (raMCTOcPllT - n). 3apoCTKH XBOlI(eH 3eJieHbie, CB060,ll;HOJKIIBYliltte. faMeTocpttTbI XBOlI(ett O,ll;HO- HJIH o6oerrOJibie, HttTeBH,ll;HOli
cpopMbI. Ha Httx o6pa3yIDTCfl attTeptt,nttH tt apxeroHtttt. ,[(JI.a onJiop;oTBopeHttH
Heo6xo,nttMa BO.n;a; cnepMaT030II,ll; co mryTttKaMH ,ll;OIIJlbIBaeT K flHJJ;eKJICTKe,
rrpottCXO.[(llT OfIJIO,ll,OTBOpeHtte. lfa 3lll'OTbI (2n) 06pa3yeTCfl 3apO.[(bIIII, KOTOpbIH HeKOTopoe BpeMfl CBfl3aH c raMeTOc}:>llTOM. 3aTeM 3apO,ZJ,billI rrpopacTaeT
B crropocpHT (2n).
Hatt6oJiee pacrrpocTpaHeHHble Blf,ll;bI XBOlI(eBlf.[(HbIX: XBOlil IlOJieBOH, XBOII~
JIYI'OBOH, XBOlil JieCHOH.

CeMeHHLie pacTeHHH

OT,ZWJI roJIOCeMeHHbie

0T.[(eJI fOJIOCeMeHHbie HaCqlfTbIBaeT OKOJIO 700 Bll,l.l;OB pacTeHHH. roJIOCeMeHHbie B OCHOBHOM Be'IH03eJieHbJe p;epeBbH, peme - KYCTapHMKM; TpaBHHHCTbIX cpopM HeT. y roJIOCeMeHHbIX ceMHIIO'IKll pacrroJIO)KeHbI OTKpbITO. JlMcTbH - lfrJIOBll,l.l;HbJe, MHOI'OJieTHlle.
Bee npe.n;cTaBI:1TeJin: roJioceMeHHbIX - pa3HocrropoBbre, T. e. o6pa3yIOT Meracnophr If MllKpOCIIOpbI. jf\'.eHCKlle MeraCIIOpbI rrpopaCTaIOT B MHOI'OKJieTO'lHbie meHCKMe raMeTocpn:TbI. MymcKtte MMKpocrrophr rrpopacTaIDT B Mymc1<11e
raMeTOcpllTbl, COCTOfllillle ll3 ,ll;BYX KJieTOK, npe.ncTaBJIHIOIII,llX co6oll: nbtJlblJ,y.
CTa,ZJ;ttH raMeTocpuTa (n) oqeHh pe,ZJ;yn;ttpoBaHa.
UnKJI pa3BttTllfl roJioceMeHHbIX MomHo pa3,11;eJIHTb Ha qeTbrpe 3Tarra:
1. Mn:KpocrroporeHe3 - o6pa3oBaHtte MymcKoro raMeTocpttTa, MttKporaMeToreHe3.
2. MeracrroporeHe3 - o6pa30BaH11e mettcKoro raMeTocpnTa, MeraraMeToreHe3.
3. OrrbrJieHue n onJIO.[(OTBopettne.
4. <I>opMttpOBaHHe ceMeHH.
1. B3pocJioe pacTeHne cocHhI - O,ll;HO.[(OMHoe, flBJIHeTCfl cnopocpttTOM
(2n). B Mae y ocHOBaHHfl ee MOJIO.n;brx no6eroB 06pa3yroTCfl nyqKn meJITOBaThIX
MymCKHX llIHIIIeK. Kam.n;aa: illllllIKa HMeeT OCb, K KOTOpoa npnKperrmIIQTC5I
MUICpocnopofjJuJtJtbt (roMoJiortt TbJqMHOK). Ha MttKpocnopocp:ttJIJiax eCTb no .[(Ba
MMKpocrropaHrna. B HHx M3 ,ZJ;HIIJIOH,!l,HhIX MaTepn:HcKMX KJieToK 06pa3y10Tca
ranJIOM,ll;Hbre MMKpocnophr (n). M:ttKpocnopbr rroRpbITbI .n;ByMH o6oJioqRaMH:
MHTHHoii (BHyTpeHHeii) n 3K3MHOH (HapymHoii). Mem.n;y o6oJioqKaMtt B pe3yJihTaTe paccJioeHna o6pa3yeTCH .n;Ba B03.n;yxoHOCHhlX MernRa. MHRpocrropa (n)
MMeeT BereTaTMBHYIO n aHTeptt,lJ;HaJibHYJO KJieTKH. BereTaT:ttBHaa KJieTKa 06pa3yeT IlbIJiblJeBYIO Tpy6Ky, aHTepll,ll;llaJihHaH KJieTKa - ,ll;Ba crrepMMH. 3peJiaa:
MHKpocrropa - :no IIhIJibJJ;eBoe 3epHo, MJIM MyJ1Cc1Cou zaMemofjJum. My.mcRoii
raMeTocp:ttT npo).J;yn;.ttpyeT raMeTbI - p;Ba crrepMna.

231

CeJK.ennbte pacmenuR. OmiJeft I'oftoceMenuble

2.

iKeHCKMC llilllUKll KpaCHOBaToro 11,BeTa II co6pa!Ihl ITO 0,ll;HOH MJIM TPI1 Ha

:l(OHU8X M0.'10,ll;blX rro6el'OB. Kam,n;aJI llllllllKa MMeeT OCh, OT KOTOpoH: OTXO,ll;RT YKOpoqeHHhle BH,ll;OH3MeHeHHhie rro6er11 ,ll;BYX THITOB: 6ecn;10;:i,nh1e (KpOIOIJJ;He) H CeMeHHhle (c lJ.ByMR ceMR3a'!aTKaMH). CeMR3a'IaToK
v;eHTPaJlhHaR qacTh, MHTeryMeaTa -

(2n). B

(2n)

cocTOMT M3 Hyu;e;rnyca

rroKpoB. Hy11,e.'l.'lyc -

aeM ecTh o;:i,Ha apxecrropttaJihHaR KJieTKa

(2n),

(2n) _

3TO MeracnopaHrttli

KornpaR ):\eJIMTCR MelioaoM

o6pa3yeT 'IeTh1pe Meracrrophr (n). Tpn n3 Hnx OTMttpaIOT. O;:i,Ha Meracrropa (n)

8
:r.moroKpaTHO ):\eJIHTCR MI1T030M, H o6paayeTca HOBaJI TKaHh 3Hll,OCrrepM (n). B TKa-

11H 3H,lJ;OCIIepMa (n) o6pa3YIOTCR ll,Ba apxerOHMR (n), B HHX ct:>opMnpyeTCJI RllIJ;eKJieTK8 (n). 8a;:i,ocrrepM H ;:i,Ba apxeroBHR -

3.

3TO

Jl(f'ftc1wu za.Memocpum coCHhI.

IlhIJihll,a rrepeHOCI1TCR BeTpOM Ha ceMJI3a'IaTOK, r;:i,e yJiaBJIHBaeTCR ry-

CTOH mH,D;KOCThIO na IIhIJihll,eBXOL\a. IlpH rro,n;chrxaHnH mH,D;KOCTH IIhIJih11,a BTRruBaeTCR qepe3 MHKpOIIHJie (IIhlJihll,CBXO;:i,). Mem;:i,y OITblJICHHCM H OIIJIOll,OTBOpeHHeM npoxo;:i,HT o;:i,nH ro;:i,. B

Te'IeHne 3Toro ro;:i,a BereTaTHBHaR I<JieTKa

rrh1Jihll,hI npopacTaeT B IIhIJihll,eByIO Tpy6Ky .no apxeroHnR. B KOHn,e rr1>Uih1~eBoi1
Tpy6KH Henocpe,n;CTBeHHO rrepe;:i, OIIJIOll,OTBOpeHneM o6pa3yIOTCR ;:i,Ba crrepMHR
na aHTepn,n;naJihHOH KJieTKH. CrrepMnn 6e3 mryTHKOB. O,n;ttH crrepMHH cJIHBaeTcR c Jiii:11,eKJieTKOli, a ;:i,pyro:H OTMHpaeT.
lJ.hlIII

(2n).

3apOll,hllII -

lfa

(2n) o6paayeTCR 3apo(2n). 3apOll,hlill COCTOHT

3HrOTbI

3TO MHHHaTIOpHhlll cnopoqmT

H3 KopernKa, CTe6eJibKa H HeCKOJihKHX CCMR,ll;OJieil

(5-12).

4. IlocJie orrJio;:i,OTBopeHHR ceMR3a'IaTOK rrpeBpamaeTcR B ceMR.

CeMR co-

CTOHT H3 ceMeIIHOll Komyphl (IIOKpOBhl CeMJI3a'IaTKa), 3H,ll;OCIIepMa (n) H 3apoll,hIIIIa

(2n).

CeMeHa coapeBaIOT oceHhIO, Ha BTopo:H ro,n; nocJ1e orrhIJieHHR. IlocJie coapeBaHHR ceMJIH ceMeHHhie 'IelllyHKH IllHilleK pacKphIBaIOTCR, ceMeHa BhIChIIIaIOT-

CR. CeMeHa nMeIOT npo3pa'IHoe KpbIJihIIIIKO, nepeHOCRTCR BeTpoM.

UnKJI

paaBHTHH cocHhI o6hIKHOBeHHOH (pnc.

Cocna o6btKHOBenna.R -

3TO ,n;epeso BhICOToii ,11;0

95)

20-40 M.

BapocJioe pacTe-

HHe HMeeT MOIJJ;HYIO CTepmHeByIO KOpHeByIO CHCTeMy. IIo6erH HMCIOT MOHOrIOll,HaJihHOe BCTBJieHHe. CTe6eJib HMeeT CJIO)KHOe CTpOeHne 6Jiaro;:i,apJI ll,JIHTCJihHOMy <PYHK11,HOHHPOB8HHIO ll.BYX MepHCTeM: KaM6HR H ct:>eJIJIOrena. Ilo aHaJIOl'H'IHOMy CTpOeHHIO CXO,D;eH co CTe6JIJIMH .npeBeCHhIX IIOKpblTOCCMCHHhlX co
BToptttIHhIM yTOJIIJJ;eHneM. CymecTBeHHOe paaJIH'IHe aaKJIIO'IaeTcR B TOM, 'ITO
B KCHJieMe HCT cocy;:i,oB, eCTh TOJihKO TpaxeHll,hl, BO l}>JI03Me HeT KJieTOK-CilYTHH11,, ecTh TaK Ha3hIBaeMbie aJih6yMHHOBhie KJieTKH (BhIIIOJIHRIOIJJ;He cxo;:i,HyIO
<PYHKll,HIO, HO OTJIH'Ia10mnecR no rrponcxom,11;em110). Kopa tt ,n;peBeCHHa co.u,epataT MHOro CMOJIRHbIX XO.D;OB.
Ilo6erH y cocHhI ,n;Byx THIIOB: Y.U.JIHHeHHhie H yKopo'IeHHhie. Ha y.u,JinHeHHLix rro6erax paaBHBaIOTCR 6yphre qewyeBH,ll;Hhie JIHCThR, B rraayxax ROTOPLix paaBHBaIOTCR yRopo'IeHHhie rro6erH. Kam)J;hIH yRopo'IeHHhIH rro6er HeceT .U.Ba JIHCTa -

XBOHHhl. XBOHHhl )KHBYT IIOJITOpa-,D;Ba ro.u,a, aaTeM orra.u.aroT

Bl\fecTe c rro6eroM. XBOHHhl IIOKpblThl ann.u.epMOH c TOJICTbIM CJIOeM KynrnyJILI. CTeHKH KJieTOK antt.u,epMhr yTOJIIJJ;eHHhre n

o.u,peBecHeB11rn:e. YcThHll,a

232

rJiy6oKO norpy1:1<CHhI B TI<:aHb .r1I:IcTa.

B

uc1:1Tpe XBOHHhI pac110.'101Kellbl"

11:sa

K0.1JI8Tepa.'IhHhIX 3a1<p1>1T1>1x nylfita. Oco6eHHOCTlI CTpoeunH TI03BOo"'I.HIOT cocne o6ttTaTh B 3aCyl1JJ1!1BbIX MCCT8X 11 nepeHOCl1Th 3H8lfHTC.::lbHOC nepeox.11an:;:.n;eHHe.
Cocua 061>1KHOBeHnaH JIBJrneTCH o.n:no,n:oMHhIM pacTeHHeM. BecHoil: na o;lHo~
.n:epese o6pa3yfOTCH II MY.iKCKIIC, 11 mencKHC IIJHIIIKH.

MyJ1Cc1Cue wuw1Cu

meJrT0-3e-

J"Ienoro useTa, pacnoJiaraIOTCH y ocHosaHIIH MOJIO.LJ;hIX no6eros. OnH o6pa3yIOTCH
s

na3yxax qernyeBII.LJ;HhIX JIHCTbeB BMCCTO YKOPO'lCHHhIX no6eros. MymcKaH

llIHllIKa npe.ll,CT8BJIHCT OCb co crrn:paJibHO pacnoJIO/f\CHHbTMH MIIKpocnopoqHrn.11a-

B

MfL

MY.iKCKIIX Iffl1llIKax QlOpMHpyIOTCH IlbIJ!hll,CBbie 3epna, COCTOHII.VIC II3 .ll.BYX

K.1eToK: BereTaTHBHOH H renepaTHBnoi-i. fenepaTI1BHaH KJ"'ICTKa o6pa3yeT .n:se MY.iKc1rne raMCThl -

cnepMHH (n). IlbIJibll,a rrepeHOCl-ITCH BCTPOM na JKCUCKHe lllHlliKH.

)i{eHc1Cue wuw1Cu

KpacnoBaToro u,BeTa, o6pa3yIOTCH B rra3yxax qernyeBH.'J:-

HhIX JllICTbCB no 0)1.HO}i -

PHC.

95.

TPH Ha KOHIJ,aX MOJIO,llblX rro6eroB. JR'.eHCK8JI IllHIU-

Yepe/],OBaHll18 noKoneH111j;j B >Ki113H8HHOM L\111Kn8 ronOC8M8HHblX Ha np111Mepe COCHbl

06b1KHOBeHHoj;j ( Pinus

sylvestris): a -

r-

03pocnb1iii cnopocpm;

6-

>1<eHcKa5l w111wKa (3pena<i 111 He3penaR):

a-

M\f>KCKa5l w111wKa;

,11, -

Meracnopo¢>111nn (ceMeHHaR yewy5l) c MeracnopaHrlllHMlll (ceM>i3aYaTKaM111);

MlllKpocnopo¢>111nn c Mi11KpocnopaHrnRMlll, pacce111sa10IJ..(111M111 nblnbL\8Bble 3epHa;

pa3BlllBalOIJ..(111MC5l MY>KCK111M raMeTOcplllTOM, COCT05lli.(il1M Ha 3TOiii CTa,D.111111 1113
KOTOPOM Meracnopa HaY111HaeT pa3BlllBaTbCR B >KeHCKllliii raMeTocj:>111T; 3 YeWyR; K -

2

e-

flblnbuesoe 3epHo c

KneTOK; >K -

ceMR3aYaTOK, 8

ceMeHHaR YewyH; V1 -- KPOIOl.l.\afl

3penb1iii MY>KCKOL1 raMeTOcj:>lllT, npeACTasneHHbliii nblnbuesoj;j Tpy6KOiii c YeTblpbM5l 5lApaMlll. OA-

HO 1113 KOTOPblX BblnOnHR8T ponb raMeTbl;
lllAHbJj;j 3HAOcnepM); M TOnbKO OAlllH); H -

n-

>KeHCKllliii raMeTOcj:>lllT (npespalJ..(alOIJ..(llli1C5l n03AHee B ranno-

apxeroHllllll co ccpopM111posasweiiiC5l RiiiL(eKneTKoiii (1113 KOTOpblX pa3BlllBaeTCfl

CeM<l, CHa6>KeHHOe neTY<JKOiii, B pa3pe3e; 0 -

3aPOAblW;

n-

npopOCTOK

QrniJe11, flOl(/JbtmOCe.Mellltbte, UJiU J~6etnK06bte

233

}Ca npep,CT3BJl.H€T co6oii OCh, Ha KOTOpOM pacr10.1o:meHhI 'ICWYMKH l~BYX nm on:
6ecnJIO,lJ;Hbie (KpOIOUVIe) H ceMeHHbIC.
aaxo,n,.!ITC.!I ,naa ceMH33'IaTKa.

B

y

OCHOB3HIIH K3JK,ll,OH CCMeHHOH 'Iemyu

tiettTpe ceMH3aqan;a pa3a11naeTCH ::>np,ocrrepM

(n). 8Hp,ocrrepM o6pa3yeTcH H3 Meracrropr,1 (n), Bero TI"3HH o6pa3yeTCH ,nsa apxeroHHH (n).

B

Kam;wM apxeroH1111 cpopM11pyeTcn ailu;eKJleTKa. 8tt,n:ocnep:-v1

c ,n;BYM.!I apxerOHll5IMH -

JKCIICKllM raMCTOcpllT. I1pou;ecc OT OIIbIJieHH5I ,ll;O 011-

JIO,lJ;OTBOpeHllH (OT rrona,n;aHHH IIbIJibl~bI Ha JKeHCKlle IIIHIIIKH ,L\O rrponmmoaeall.!I BHYTPh ceM.H3a'laTKa) ,n;mITCH OKOJIO
Hihi;eKJieTKH -

311roThI

(2n) pa3s11aaeTCH

Mecm~eB. l1_3 onJIO,L\OTBOpCHJIOil

13

3apo,n:hilll, a 113 acero ceM.H3a'IaT1rn --

ceM.!I. 3peJIOe CCM.H coe,ZJ,llH.HeT B ce6e raMeTOcjJllT (meHCKHH 8H,ll,OCilepM) II crropocpllT

(2n).

(2n).

Komypa ceMeHll 11 nep11cnepM -- 'IaCTh MaTep11HcKoro cnopocp11Ta

CeMeHa cpopNrnpyroTcn B Te'IeHHe ceM11 --- ,n:eB.HTH Mecmi;ea 11 nocJie co3pe-

aaHll.H BblCbIIIalOTCH H3 IIIHIIIeK. CeMeHa HMelOT npo3pa'IHOe KpbI.1lbIIIIKO H MOryT pacnpOCTpaHHTbCH BeTpOM.
OTp,eJI foJioceMeHHhre nop,pa3,neJIHJOT Ha rnecTh K.1accoa:

1. KJtacc CeMeHl-tbte nanopomHUliU -

nepahre roJwceMeH111>re. Bee Bhr:>tep-

JIH.

2. KJtacc BeH1temumbt.

Ilpe,n,KII noKpbITOCeMeHHbrx. I1MeJI11 o6oenoJibIC

IIIHIIIKII, Bee BbIMepJIH.

3. KJtacc Ca2061tUli06bte.

PacTeH11.H noxomH tta naJlhMhr. Caroa11HKOB1>1e -

,n,By,llOMHhIC pacTCHHH. MymcK11e H meHcK11e IIIHWKII 06pa3y10TCH Ha
pa3HhIX oco6HX.

4. KJtacc I'uHliZOBbte.
CTHhIH.

E,iIHHCTBeHHhIH npe,n,cTaBHTeJih -

rHHKro ,llBy.11ona-

JlHCTOBaH nJiaCTHHKa BeepOBll.D;HOH cpopMbI, ,llByJionaCTHaH,

c ,n;peBHHM ,llHXOTOMH'ICCKHM THIIOM m11JIKOBaHHH. CnepMaTo30llP,hr no,n;attmnb:re.

5. KJtacc I'Hemo6bU'.

8cpe,llpa xa01ri;esa.H (noJiy'IaJOT cna3MOJIHTII'Iecirne ae-

w;ecTBO ::>qJep,p1n1), aeJihBH'IH.H y,llHBHTeJihHaH -

HMeeT acero ,llBa o'IeHh

,llJIHHHhIX JIHCTa, pacTyr:u;He BCIO )KH3Hb.

6. KJtacc X60UHbte -

naM6oJiee 06wnpu1>rii n pacnpocTpaHeHHhiii KJiacc.

B Hero BXO.D;HT:

CeMeucmeo Coc1to6b1e ( cocHa,
aasr -

eJih, nnxTa, JIHCTBeHHmi;a). CocHa o6hrKHOBeH-

XBOHHKH no nape, COCHa cn6HpCKaH -

XBOHHKH no IIHTb WTYK (B Hapo,n;e

Ha3bIBaeTC.H Ke,llpOM, ceMeHa COCHhI CH6MpCKOM -

CeMeucmeo Kunapuc06bte

KC,llPOBbIC opernKH).

(KHnapnc, MommeaeJihHHK).

Y

MO:>KJKeBeJihHH-

KOB IlOCJie OIJJIO,llOTBOpeHH.H qewyn MeracnopocpHJIJIOB CTaHOB.HTCJI M.HCHCThI-

MH, cpacTalOTC.H, o6pa3y.H «IIIHWKOHro,n;y».

Ce.11teucmeo Tucoebte

(THC aro,n;Hhrii:).

0TAeJI IloKpbITOCeMeHHLie, HJIH QBeTKOBbie
::hoT OT,llCJI O'ICHb urnpoKO pacnpocTpaHeH II HaC'IIITbIBaeT OKOJIO

250 TbICfl'I

BH,lJ;OB. Cyw;ecTBYIOT B 60JibIIIIIHCTBe THnOB 8KOCllCTeM, B TOM 'IHCJie B BO,llHbIX
(KYBillHHKH, Ky61>rrnKM, p,lleCT, aJio,n;e.H). C'IIITaeTca, 'ITO ri;BeTKOBhre pacTeHHfl
B03HHKJIII B MeJIOBOM rrepHop,e; IIMelOT IJ;BeTKH II nJIOP,bI.

I>omanu1w

234

l~BeTOK -- 3TO Bil/1,0113:.YieHeHllbIH, YKOpOYCHHbIH, orpaill1lJeH!lbIIi B pOCTe
cnopoHOCHOCHhiii no6er, BbIIlO.iIHRJ{)IIJ;Hii cpyHKIJHIO 6ecno.11oro H IIO.'!OBOro pa3MH0)f(eHHR. TI pH cpopMHpOBaHHH UBeTKa MHKpocnopOJIHCTHKH npenpaw;aIOTCR
B TbllJllHKH, MeracnopOJIHCTHKH - B neCTHK.
B IJHKJre pa3BHTIIH npeo6Jia;i;aeT 6ecno.1oe noKoJ1enne (cnopocpnT --- 2n); raMeToqrnT cmrhHO pe.n;yunponan: MY)f<CKorr raMeTocpHT - 3pe.'Ioe nhI.'IhIJ,eBOe
3epHO llJill Ilbr.'IhIJa, meHCKiifI raMCTOqrnT - BOChMHKJieTO'-IHhIH 3apOJJ;hIIIIeBhiii:
MeruoK. XapaKTepuo JJ;Boiinoe on.'IO;lOTBopen11e. CJ1JIBaIOTCR:
HiiIJeKJieTKa (n) + cnepMIIH (n) ··~ 3apOJJ;hIIU (2n);
uenTpaJibHaH KJieTKa (2n) + crrepMnii: (n) ~ :rn.uocnepM (3n).
U:mw pa3BHTHH IlOKpblTOCCMeHHhlX

(pnc. 96)

QHKJI pa3BI:ITHH TioKpbITOCeMeHHbIX MO)f(HO pa3.n;eJIHTb Ha qeTbipe 3Tana:
1. MnKpocnoporeHea - cpopMnpoBaHne :vrymciwro ra:vreToq:mTa, MJIKporaMeToreHe3.
2. Meracnoporene3 - cpop;vinpoBanue mencKoro raMeTocj_Jnrn., MeraraMeToreHe3.
3. OnhrneHIIe n on.10.n;oTBopenne.
4. 06pa3oBaHJie ceM.HH n n.'Io,n;oB.
Mu1Cpocnopoieue.; II cj_JopMIIPOBaHne MymcKoro raMeTocpIITa nponcxo,n;IIT
B IIhIJihIJeBhIX rHe3,n;ax IlhIJ1hHHKa. MnKpocnopanrI:III (2n) - 3TO rne3.n;a IIhrnhHIIKOB. B HI:IX ll3 Kam.n;oii: /l,llilJIOH.IJ:HOll apxecnopna.'IbHOH KJieTKll (2n) B pe3YJibTaTe Meil:o3a o6pa3yIOTC.H qeThrpe ranJion,n;HLie MIIKpocnophr (n). MIIKpocnopa HMeeT o60JIOlJKY JI .H,n;po. Tipu .n;e.'IeHJIJI :vIHT030M JI3 MJIKpocnopbI 06pa3yeTC.H ,n;Be KJieTKII: BereTaTJIBHa.H (n) - Kpynna.H JI renepaTHBHaJ:I (n) MeJIKa.H. IlbtJlbU,€'60(' .;epHO U.J aByX lCJlemolC - MYJICCICOU iaMemo</Jum. TioKpbIT ,n;ByM.H o6oJIOlJKaMII - 3K3HHOM JI JIHTH11oii:. Tipw rrpopacTaHnn nhr.'IhIJeBoro 3epHa (B xo.n:e OTIJIO,n;OTBOpeHI:IH) 06pa3yeTC.H ,n;Ba cnepMH.H -- MYIBCKHC raMeThI (n).
Meiacnopoieue3 npowcxo/l,HT B ceM.HnoqKax, a TaKme B n;eHTpaJihHOM qacTH - nyueJIJiyce. Hyue.'IJiyc (2n) co.n:ep~JIT MaTepnncKyIO apxecnopwaJihHYIO
KJieTKY (2n). TiocJie Meii:o3a H3 Hee o6pa3yIOTCH qe'fhrpe ranJioII,n;Hhre Meracrrophr (n). Tpu H3 HHX OTMHpaIOT, 0/l,Ha OCTaeTC.H . .H.n;po Meracnophr (n) TPH pa3a
,n;eJIHTC.H MHT030M, H cpopMJipyeTCJ:I BOChMJIKJieTOlJHbIH 3apO,n;hIIUeBhIH MeIUOK.
3apoabtute6btil MeutolC - 3TO J1Ceuc1Cuu iaMemo</Jum (n). CTpoeHI:Ie 3apo.n;blrueBoro MeruKa: B uenTpe - ueHTpaJibHaR KJieTKa (2n); y IIhIJihIJeBxo.n;a - .HHIJeKJieTKa H ,n;Be CHHeprH,n;hr; Ha npOTHBOilOJIOIBHOM TIOJIIOCe - TPH aHTHilO,n;hI.
ApxeroHHH noJIHOCThIO pe,n;yri;upoBaJIHCh.
OnbtJieuue MomeT 6bITh: caMOOIIhIJieHHe H nepeKpecTHoe. TibIJihIJ;a nona,n;aeT Ha pbIJihIJ;e necTHKa. BereTaTHBHa.H KJieTKa o6paayeT TihIJihIJ;eByIO Tpy6Ky,
pacTeT qepe3 CTOJI6HK, MllKPOTIHJie, HyIJ;eJIJIYC H .IJ:OCTHraeT 3apo,n;bIIIIeBoro
MeruKa. feHepaTHBHa.H KJieTKa Ha KOHIJ;e Tpy6KH ,n;eJIHTC.H Ha .n;Ba cnepMHH.
B onJio,n;oTBopeHHH yqacTBYIOT o6a cnepMw.H: o,n:HH CJIHBaeTcH c .HHIJ;eKJieTKOM
(n) H 06pa3yeT 3HrOTY (2n), H3 Hee 06pa3yeTcJI 3apo,n;hIIII ceMeHH; a .n;pyro:H

U.uiai pa3<mniun noKp1>1moceMennbLX

P111c. 96.

235

YepegosaHvie noKOneHviiii B >KVl3HeHHOM u,viKne noKpb1ToceMeHHblX pacrnHviiii:

e3pocnb1iii cnopocpviT: 6 Kpocnopb1: ' -

06oenonb1iii L(BeTOK; B -

33B513b (BVIAOVl3MeHeHHbliii Meracnopocpvinn -

OAHOiii Meracnopoiii; /1, MY>KCKOiii raMeTOQ:>VlT;
MY>KCKVlx raMeT -

e-

a -

nblnbHVIK (MviKpocnopocpVlnn). pacceVlBaJOLl.(viiii MVlnnOAOnVlCTVlK) c OAHVlM CeM513aYaTKOM Vl

nb1nbu,eeoe 3epHo, B KornpoM MVlKpocnopa Q:>opMVlpyeT 2-siAepHbliii Monor.1oiii
o6pa30B3HVle 3penoro MY>KCKOro raMeTOQ:>VlTa (nblnbu,eeoiii Tpy6KVI) Vl ABYX

cnepMVleB c napannenbHblM cpopMvipoeaHVleM 3penoro >KeHcKoro raMeToQ:>Vlrn (3a-

POAb1weeoro MewKa) c siiiil(eKneTKOiii Vl L(eHTpanbHblM AVlnnOVlAHblM 51APOM;

!K -

npOHVIKHOBeHVle cnep-

MVleB B 33POAb1WeBbliii MeWOK c nocneAyJOLl.(VIM CnVl51HVleM OAHOro Vl3 HVlX c siiiil(eKneTl<.Oiii, a BTOporo c L(eHTpanbHblM 51APOM (ABOiiiHOe onnOAOTBOpeHVle); 3 - nnOAC ceMeHaMVl; 11- 3penoe ceMsi; K - pa3pe3 ceMeHVl c 33POAb1WeM Vl OKpy>KaJOLl.(VlM era TpVlnnOVlAHblM 3HAOCnepMOM (pa3BVlBaJOL1.(VlMC51 Vl3 L(8HTpanbHOro 51APa);

n-

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crrepMHH CJIHBaeTCH c u;eHTpaJibHOM KJieTKOM

(2n)

H o6pa3yeT TPHIIJIOH,D;HYIO

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(3n).

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aoonorMSl

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500

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CoBpeMeHHaJI 300Jior11JI -

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HCCJie.n:oBaHHJI OHa no;i:pa3.n:eJIJieTCJI Ha:

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npom0300JlO?UIO (113yqaeT 0,Zl:HOKJieTO'IHhIX :amBOTHhIX);
300J10i?Uf0 6ecn0380HO'lllbtX:
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(u3yqaeT paKoo6pa3Hhix); aKapoJiorIIJI (1-13yqaeT K.'le!IJett); apaxHOJior1rn
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2

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237

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KJiaccmtrn1rn1vrn: mnBOTHoro MHpa
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qTO OTpamaeT o6w;ee HanpaBJieHHe lfCTOpI1'IeCK01'0 pa3BHTIIH :tKIIBOTHOro MHpa
Ha 3cMJie. CncTeMa mnBOTHoro M11pa 6h1Jia coa.n::rna B Konue XVII -

nepBoi1:

rro.10BMHe XVIII B. B pa6oTax a11rJIHHCKoro Y'IeHoro .LlJfWHa Pean mne;1c1wro
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HOMCHK.'laTyphl H83B8HBH BII,ll;OB JKHBOTHhlX H pacTeHHH.
OcHOBHhIC KJrncc11c}lnKai~noHHh1e e,n;nHHUhI (TaKCOHhI) :tKHBOTHh1x:
UAPCTBO ~ ITO.Llll;APCTBO-_, TMIT __., KJIACC ~ OTP.H.Il-_,

CEMEi1-

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OcHOBHaa emrnnua cncTeMaTnKn -

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rra OT,D;eJibHhIX )l.UfBOTHhIX, CXO,D;Hh!X MeJK.[\y co6oil no CTpoemno, H\I13He,n;eaTeJihHOCTII, 06nTa10w;nx Ha onpe.n:eJieHHOM TeppnTop11n 11 ,n;a10w;11x IIJIO.Z~OBnToe
IlOTOMCTBO.
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K.iaccbl, munbt, KOTOpbie BXO.ll;HT B COCTaB ll8PCTBa.
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,D;Ba noau,apcmea: 0,n;HOKJieTOqHhie JKIIBOTHhie, HJIH IIpocTeiirnne, H M1rnrorpyTIIIhl :tKHBOTHhIX -

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20 TIIIIOB H HeCKOJihKO CO-

TeH KJiaCCOB.

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CJIOBO «,ll;OMHUlHHR» -

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po,ll. 0HII OO'hC,llHH.ff-

IOTCH B ceMeiiCTBO Komaqh11 ~ oTpR,n; Xuw;HhIC ~ KJiacc MJieKorrnTarow;ue
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IlpOCTCHIIIIIe -

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CKOH KJieTKH, BhITIOJIHmow;eii c}lyHKuuu ueJiocTHoro oprattu3Ma (rrnTaHHe,
,ll;BHJ:KeHue, pa3MHOJKeHue, aaw;nTa OT He6JiaronpHHTHhlX yCJIOBHH cpe,llhl).
ITpocTeii:rnne -

sTo opraHH3MhI na KJieToqttoM ypoBHe oprattnaauHn. l1Me-

IOT MHKpOCKOITUqeCKHe paaMepbl. :IB'.HBYT BO Bcex BO,D;OeMaX, BO B.'laJKHOH rroqBe, B oprauax pacTeHIIH, JKHBOTHhIX H qeJIOBeKa.
Brrepsb1e rrpocTell:mne 6hIJII1 o6HapymeHhI I'OJ'IJ'IaH,n;cKuM yqeHhIM AHTOHII
BaH JieBeHryKOM

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r.), KOTOpbIM HBIIJIC.ff OCHOBOIIOJIOJKHHKOM HayKH Mll-

:Kpo6uoJiorHH. Oco6eHHO MHoro rrpocTeifrnHx 6hrno o6Hapymeno B HaCTOHX
TpaB (OT JiaT. infusum -

B rrepBoll:

HaCTOHKa), II03TOMy HX H83hIB8JIII HHc}ly3opnHMH.

KJiacc11c}luKauHH JKHBOTHhIX KapJia JIHHHea

(1759

r.) npocTell:I1Ine

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238
K;iaccm}>ttKa~1u1

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Ilo,n;TIIII

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npocTeiiumx
Ilpe;1cTaBl1TCJIH

AMC6b1 (aMe6a 06bIKIIOBe1rna.H, K11llle 1 rnaH aMe6a,
/UI3e!I'repHliHaH aMe6a)

cn1rocpopa

PaKOBIHlHhie aMe6b1 (api~eJIJia, .n;mj.Jtpmorvrn)
<PopaMHHHcpepb!
Pa,n;HOJTHpm1 (;ry'leBnKH)
C0Jrne 1 rnm{H
JKryTl1KOHOCUhl

3BrJICHOBb!e (3Bl'JICHa 3CJieHaH)
BOJibBOKCOBbie ( BOJibBOKC)
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TpttxoMoHa.n;oBbie (TplIXOMOHa,n;a Kl1llle11Han
11 yporeu11TaJihHaH)
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Ail11KOMil-

CnopoB11K11

KpoBJJHbie cuopoBHKH (Ma.J1Hpl1ttHb1i1: n.1a3Mo,n;11li)
Kom~H,n;HH (<1liMep1rn, TOKCOIIJiaaMa)

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rperapttHbl
Pecmi'-1Hb1e

HHcpyaopHH-TycpeJihKa, 11HcpyaopttH-Tpy6aq,
CTH,'IOHHXHH (6p10xopeCHHl'.!HaH HHcpyaop11.rr),
cyBOHKa, ,Zl;H}I:l1Hl1YM, 6ypcap11.H, 6aJI3HTlf,ZJ;HH,
ryceK

6hVIH OTHeceHhI K

KJiaccy '-Iepne:ii:. To.JihKO B

XIX

B. AJih6epT R'.eJIJIHKep

H KapJI Teo,n;op 3tt60Jih,n; BhI,n;eJIHJIH Hx B caMoCTOHTeJihHhIH THil

B 1980

Protozoa.

r. npocTeihnue 6hIJIH o6'he,n;HHeHhI B rro,n;u;apcTBO o,n;HoK.'leT011HhIX

H no,n;paa.n;e.JieHhI Ha ceMh THIIOB: cap1w.Macmuzo</Jopbt, anu1Co.MnJte1CCbt, .MuKpocnopuiJuu, .MuKcocnopuouu, uttf/Jy3opuu, Jta6uputtmyJtbt. acu,emocnopoBbU'.

B

OCHOBY KJiaccmt>HKaUHH Ha Tllllhl IIOJIO:>KeHbl rrpMHil;llIIhI CTpoeHH.H HX

a.n;epHoro annapaTa, opraHe.JIJI .n;Bu:meHH.H, TunoB pa3MHO:>KeHH.H H :>KH3HeHHhIX UHKJIOB. BoJiee rro,n;po6Ho B KHHre .n;aHa xapaKTepHCTHKa TOJihKO Tpex
TlillOB npocTe:HUJHX: capKOMaCTHro<lJop, anMKOMIIJieKCOB ll HH<lJyaopH:ii:.

06m;aJI xapaKTepHCTHKa rrpocTeHIIIHX
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SI.n;ep, II;HTOI1Jia3Mhl, orpaHHl'.!eHHOH CHapy:mn MeM6paHoii:.

HapyJ1Cttbte noKpOBbt 06ec11e'-!nBa10T <lJopMy TeJia.

f]oiJ1,fapcm6o OiJH01a!-emo'lllbte JtCU60mllbte, u.nu Ilpocmeiiuwe

239

1. <l>opMa TeJia HenocmoRH!laJt: KJieTKa orpanII'-ICHa TOJihKO llIITOIIJia3MaTIIqecKoli: MeM6panoii (aMe6a).
<l>op!V!a Te JI a nae momt1ta.fl:
I!JIOTHa.H :rnacTtttiecKa.H 060;10'IKa (neJIJrnKyna), o6pa3yIO:w;aaca 3a cqeT
yrrnoTHeHII.H rrepmpep:wreCKOI'O CJIOH 3KTOilJia3Mbl 11 Ha.rrnqIIH OIIOPHblX
qm6pmrn (JKI'YT11KOHOCIJ,bl, cnopOB11KII 11 IIHcpy3opII11);
KJieToqna.H CTeHKa II3 IIOJIHCaxap11;:i;oB - u,eJIJUOJ103bl, rreKTHHa H T. )];.
(paCTIITeJihHhie :mryT11KOHOCIJ,bI);
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JiyqeBllK11).
ll,11TOilJia3Ma rro.n;pa3.n;eJiaeTCH Ha ,n;Ba CJIOH: ::ncmon.na3MY - HapymHhIH,
rrpo3paqHbIH, 6oJiee IlJIOTHblll ll 3HOOnJta3MY - BHyTpeHHllll, 3epHl1CTblll, rro,l:i;BmKHhIH, co.n;ep:ma:w;11ii: Bee KJieToqnhie opraneJIJihI n a;:i;po.
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TOUJia3MhI, IlOKpbITbJC cnapymII U:IITOIIJia3MaTHqecIWH MeM6panoli, BHyTpH
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,Il;hIXaHlie IIpOHCXO,ll;HT: qepe3 ll;HTOIIJia3MaTHlleCKYIO MCM6pany Bce:H rroBepxnoCTblO TeJia 3a cqeT .n;ncpcpy3MH ra30B; qacT11qno npn yqacTHM coKpaTHTeJihHhIX BaKyoJieii; y napa3HTOB, 06IITa10:w;11x B cpe.n;e, JIIIIIIeHHOH KHC.TIOpo.n;a,
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rnx BH.UOB MJieKOIIHTaroru:nx, a TaKme IITHU:, KOTOpbIH nopamaeT pa3H006pa3Hhle TK8HH H KJICTKH CBOlIX X03.HeB. Jl\'.H3HeHHbIM U:HKJI COCTOHT H3 .LIBYX <Pa3:

Tun anuKOMnReKca. Tun U1ujJy.Jopuu (pecnU'tltb<e)

249

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aa6mo,n:aeTCH B KI:Illie'IHHKe rrpoMeiKyTO'IHOro X03JIMIIa (qeJIOBeKa IUII:I ,n:pyr11x
MJieKOilI:ITalOIIJ;HX, a TaKiKe IlTI111,).

Tm1 nucpyaopHH (pecunquhie)
H11rpy3opuu -

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TeJio no cpopMe HarroMIIHaeT mencKyro TycpeJihI<y. Hapymnhrli

c.;wif

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rrpO,LIOJibHhle TJIJKH MHKpOcpII6pIIJIJI (MIIOHeMbI). IlpII HX COKpani;eJHl:H H3MeHJIeTCJI cpopMa TeJia TycpeJihKH, 'ITO II03BOJIJieT eli npOHHKaTb B Y3KHe w;eJIII.
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6
PMc. 101. CTpoeH~e ~H¢y3op~~-ry¢enb1<~: a - nepeAHHCi KOHeu,; 6 - 3aAHHC1 KOHeu.. I - Manoe
2- 6onbwoe HAPO (MaKpOHyKneyc); 3 - 3HAOnna3Ma; 4 - 3Krnnna3Ma; 5 -

HAPO (MHKpoHyKneyc);
pecHw.irn; 6 -

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9- nHU18BapHT8nbHaH BaKyOnb; 10- COKparnTenbHaH BaKyOnb; 11 - o6onOYKa (nennHKyna)

250

300.!WZUR

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251

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( aMe6ou;MThJ) M
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9

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1.

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OTBepCTIIeM, KOTOpoe OKpymeHO

2.
3.
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IlepBH:qHopOThie. IfMeeTCH POTOBOe OTBepCTHe (e,n;IIHCTBCHHOe) .l(JIH 3arJiaThIBaHIIH IIMIIJ;II H BhIBeJJ;eHMH HerrepeBapeHHhIX OCTaTKOB. IlIIr.u;eBapeHIIe ,n;Boilttoe (noJIOCTHOe n BHYTPHKJieToqttoe).

5.

HepBHaH CMCTeMa JJ;tt<P<PY3Horo Tttrra, COCTOMT H3 HCPBHhIX IOieTOK B 3KTO,n;epMe, KOTOpbie coe.ri;IIHHIOTCH .l(.'IIIHHhIMII OTPOCTKaMII, o6pa3yH ceTb
(pacceHHHOe HepBHOe CIIJieTeHIIe).

6.

.I1:hIXaHIIe II BhI.l(eJieHHe rrpo.n;yKTOB o6MeHa rrpOMCXO.l(HT qepe3 BCIO rroBepXHOCTh Te JI a.

7.

Pa3MHomeHIIe 6ecrroJioe (uoqK0Ba1rne, cTpo6IIJIJIIJ;ttH) u rroJioBoe (KonyJIHIJ;tt.H).

y

HeKOTOPhIX B JKM3HeHHOM IJ;MKJIC Ha6JIIO.l(aeTCH qepe,n;oBaHMe

IIOJIOBOro II 6ecrroJioro IIOKOJieHHll.

8.

Pa3BMTIIe c MeTaMop<lJo3oM (.'I11qnHKa -

Tun 1wwe'l1-tono.!locm1-tbte

rrnaHyJia), peme -- rrpHMoe.

BKJIIoqaeT: KJiacc fn.n;ponJJ;Hhre, KJiacc Cu;II<Pon.n;-

Hhle, KJiacc KopaJIJIOBhie rroJIMIIhI (cM. Ta6JI. Ha c.

260).

Knacc n.1ApOMAHble
K.!lacc I'uapouiJHbte H.l(hl. If3BeCTHO OKOJIO

4

B OCHOBHOM MOPCKMe, peme ThIC. BII.l(OB.

rrpecHOBO.l(Hbie rn,n;po-

IlpeCH0600HaR iuapa -

O.l(lIHOqHhill IIO-

JIHIT, npHKperrJieHHhrii no.n;omBoii K cy6cTpaTy. TeJio IJ;HJIIIHJJ;pnqecKoii <PopMhl, pa3MepoM .ri;o
HhIH

1

r.u;yrraJihIJ;aMII

CM. Ha BepxtteM Konu;e TeJia pacnoJiomeH poT, oKpymen-

(5-12).

06nTaeT

B

rrpy,n;ax,

03epax,

peKaX.

Be,n;eT

MaJIOIIO.l(BII:IKHhIH o6pa3 iKII3HII. Ilepe.l(BIIraeTCH 3a cqeT COKpar.u;eHHH TeJia
H r.u;yrra;ieu; -

«KYBhipKaHCh». XIIIIJ;HHK, rrIITaeTCH MeJIKMMII 6ecrro3BOHoqHhI-

MH 'RHBOTHhlMII (.na<PHMHMM, IJ;IIKJIOIIaMn) M MaJihKaMII pb!6.

Bnympennee cmpoenue iuapbt (pnc. 103):
1. HapyJ1C1-tbtu C.!lou (:ncmoaepMa). KJieTKM
3IIIITeJIIIaJihHO-MYCKYJihHhie -

JJ;II<P<PepeHu;IIpoBaHhr:

IIOKpOBHhie KJieTKM, IIMeIOr.u;tte COKpaTM-

TeJihHhie oTpocTKM, pacrroJiara10r.u;nec.H B.l(OJih TeJia, rrpn coRpar.u;eHMM
KOTOphIX TeJIO ITOJIMirn M ero r.u;ynaJibIJ;a YKOpaqwBaIOTCH M YTOJIIIJ;aIOTCH,
a npH paccJia6JieHMM BhlTHI'IIBaIOTCH;

254

300/WlUR

HHTepCTHlJ;IIaJibHbie (npoMef!<YTOY:Hhie) KJJeTKH -- Me.'IKHe, He,n;mp<PepeHI.IHPOBaHHhie, pacnoJiara10mHec2 rpynnaMH Mem.n;y anHTeJIHaJihHoMycKyJihHhIMH KJreTKaMH. Co.n;epmaT Kpynnbre H,n;pa H MaJio r~HTOnJia3MhI, o6pa3yIOT .npyrHe KJieTKH :JKTO)lepMbl H o6ecne'IHBaIOT IIPOI.IeCC perenepaI.IHH;
CTpeKaTeJihHbie KJieTKH -

pa36pocaHhI no BCett IIOBepxHOCTH Te.'Ia H o6-

pa3yIOT CKOII.'IeHHR Ha mynaJibI.IaX (cTpeKaTeJibHhie «6aTapeH» ), KOTOpbre BhIIIOJIHRJOT <PYHKI.IHH Harra,n;eHHR H 3an~HThI. Co.n;epmaT cTpeKaTeJihHYIO KancyJiy c .f!)lOBHTOM )l{H)lKOCTbIO, BHYTPH KOTOpoi1: HaXO)lHTCR
CBepnyTa.H CUHpaJihHO CTpeKaTeJibHa.H HHTh. KJieTKa HMeeT OC.f!3aTeJihHbIM (qyBCTBHTeJihHhIM) BOJIOCOK. IlpH pa3,n;pameHHH 3TOl'O BOJIOCKa
CTpeKaTeJibHaJI HHTb c CHJIOM Bhinp.f!MJI.f!eTC.H H npOHHKaeT B mepTBY.
CTeKaIOilIHM no KaHaJiy HHTH 2,11; napaJIH3yeT MeJIKHX mHBOTHhrx. IlocJie
BhICTpeJIHBaHHR CTpeKaTeJihHhie KJieTKH OTMHpaIOT, a HOBhie o6pa3yIOTC.H H3 HHTepcTHI.IHaJihHhIX KJieToK;
HepBHhie KJieTKH -

3Be3)l'IaTOM cpopMhI, pacrroJiaraIOTCR no.n; 3nHTeJIH-

aJihHO-MYCKYJihHhlMH KJieTKaMH, coe)lHHRRCh ,n;pyr c .n;pyroM TOHKHMH
OTpOCTKaMH, o6pa3yIOT HepBHOe cnJieTeHHe. TaKaR HepBHaR CHCTeMa Ha3bIBaeTCR .n;n<P<PY3HOM H RBJIJieTC.f! caMOM npHMHTHBHOM cpe,n;H MHOrOKJieTO'IHhIX mHBOTHhIX.

B

OTBeT Ha pa3,n;pameHIIe, HattocnMoe no.i1nny,

HarrpHMep IIl'JIOM, TeJIO ero COKpamaeTCR.

2.

Bnympe1muu cJiou (:mmoiJepMa).

KJieTKH .n;n<PcpepeHI.IHPOBaHhI:
3IIIITeJIHaJihHO- MYCKy JibHhie MYCKYJihHhie OTPOCTKH,

HMeIOT

pacrroJiaraIO-

ilIHeCJI rrorrepeK, rrpH COKpameHHH KOTOpbIX TeJio Y.UJIHHReTCR u cymaeTCR. Crro-

2

co6HhI K cparOI.IHT03y, o6pa3y.H rrceB,n;orro.n;uu, 3aXBaTbIBaR IIHilleBbie 'IaCTHI.IhI
ll rrepeBapttBaR HX B I.IHTOIIJia3Me (BHYTpHKJieTO'IHOe IIHilleBapeHHe ).
mryTHKH

(1-3),

lfMeIOT

c IIOMOillhlO KOTOpbIX

rrpOHCXO)lHT rrepeMelJlHBaHHe co.n;ep:>KHMOl'O rrumeBapttTeJihHOM IIOJIOCTH.

Ta-

KHM o6pa30M, 3Tll KJieTKH BbIIIOJIHRIOT
IIOKpOBHyJO, cmcpaTHTeJihHYJO ll IIHilleBapHTeJihHYJO <PYHKI.IHH;
:>KeJie3HCThie KJieTKH -

PMc. 103. CTpoeHi-ie npecHOBOAHOC1 r1<1Apb1: 1 - pornsoe ornepcrne: 2- ~ynanbt-1a;

3-

pa3s111sa10~asicsi MonoAasi r1<1Apa;

4-

Bbipa6aTbIBaIOT

ll BhI)leJIRJOT IIHilleBapHTeJihHhie cpepMeH Thi B racTpaJihHYIO IIOJIOCTh, r.n;e rrponcxo.D;HT

BHYTPHIIOJIOCTHOe

rrumeBapettue.

cnoC1 KJ18TOK (3HTOA8PMa); 5 Hapyit<Hblei cnoC1 KJ18TOK (3KTOAepMa); 6- Klll-

06pa30BaBIIlHeC.H IIHTaTeJihHbie BemecT-

W6YHasi nonOCTb

KH IIHilIH BhI6paChIBaIOTCR qepe3 POT.

BHyrpeHHlllC1

Ba ycBaHBaIOTCR, HerrepeBapeHHhie OCTaT-

J{nacc I'uiJpouiJnbie

255

Pa3MHomem1e: 6ecnOJwe (JieToM) - npo11cxoµ;11T noqKonamrnM . Ha Te.TJe o6pa3yroTc.H noqK11 - Bhm.Hq11Bam1e .n;Byxc.1oiIHoii: cTeHKH. IIoqKa pacTeT, 11 .n;otJepHHli opraHII3M cyw;ecTByeT 3a cqeT MaTep11ttcKoro. 3aTeM ua rroqKe IIO.HB;rneTC.ff poT, OKpymeHHhlll w;ynaJibIJ;aMH, H IIO,ll;OIIIBa. IJoqKa OTphIBaeTC.H OT
MaTepHHCKOro opraHH3Ma, rra.n;aeT Ha ,D;HO H rrepexo,D;l1T K caMOCTO.HTCv1hHOH
'1{M3HH. 1Io.rto60e - Kony.1HIJ;MH Ha6mo.n;aeTCH oceHhIO, c IIOHM:tl{eHI1eM Te:vmepaTYPhI, oceMeHeHMe nepeKpeCTHOe. fII.n;pa o6hIKHOBeHHaH - repMacl:>po,n;IIT.
B aKTo.n;epMe cI:>opM11py10Tc.H roHa.D;hI, B KOTOphrx pa3BMBaIOTC.H raMeThI. MymcKne KJieTKH 06pa3y10TC.H B He6oJihIIIMX 6yropKax Ha BepxHeli qacT11 TeJia rn.n;pbI, a Kpyrma.H .HHIJ;eKJieTKa pacnoJiaraeTC.H B BhIIIYKJIOCTH y OCHOBaHM.H. CnepMaT030H,ll;hl qepe3 pa3ph1B TKaHM BhIXO,D;.HT B BO,n;y 11 npoHHKaIOT B .HHI~eKJieTKy
,n:pyroli oco611. 311r0Ta IIOKphIBaeTC.ff IIJIOTHOH o6oJioqKoii: II oce.n;aeT Ha ,ll;HO BO,n:oeMa (MaTepIIHCKHH opraHH3M rrorII6aeT). BecHoli HaqmraeTc.H pocT 11 pa3Bl1TMe MOJIO,D;OH rn.n;phl (pHC. 104).
Mopc1Cue iuiJpouiJHbte no.rtunbt oT.'IHqaIOTC.H HeKOTophIMM oco6eHHoCT.HMM oT
npecHoBo.D;HhIX r11.n;p H IIMeIOT 6oJiee CJiomHoe pa3BIITn:e. Be.n;yT rrp11KperrJieHHhr:ii
o6pa3 )KH3HH, KOJIOHHH o6pa3yIOTC.ff B pe3yJibTaTe rroqKoBaHH.ff, HO oco611 He OTpbIBaIOTC.H, a ocTaIOTC.H Ha MaTepn:HCKOM opraHM3Me. B CJIO:IKHO ycTpoeHHhIX KoJIOHM.HX IIOJIMIIhl cneu;MaJIM3HPYIOTC.ff Ha BhIIIOJIHeHHM onpe.n;eJieHHOM cl:>YHKIJ;HH,
a caMa KOJIOHM.ff CTaHOBMTC.ff e.n;n:HhIM opraHM3MOM. y IIOJIMIIa 06eJIIIH 60JihIIIa.H
qacTh oco6e:ii - r11.n;paHThI, noxom11e Ha rII.n;py. OT.'IHtJaeTc.H rn:.n;paHT rro pacnoJiomeHn:IO pTa Ha BhICTyna10w;eM CTe6eJihKe co MHO:>KeCTBOM w;yna.'IeI~ BOKpyr.
K11IIIetJHhie noJiocTM noJIHrroB coo6w;a10Tc.H Mem.n;y co6o:ii, n: n11w;a, 3axBatJeHHa.H

PMc. 104.
TOAePMb1;

BHyrpeHHee CTpoeHHe npecHOBOAHOC1 rnApb1: 1 -

2-

3KTOA8pMa;

3-

6a3anbHaR MeM6paHa;

3nHTenHanbHO-MyCKynbHble KneTKH 3HTOAepMbl;
HepeHasi KneTKa;
Hasi KneTKa;

9-

7-

4-

3nHTenHanbHO-MycKynbHble KneTKH 3K-

3HTOAepMa;

5-

>Kene3HCTble Kn8TKH;

68-

npoMe>KYTOYHble (HHTepCTHU.HanbHble) KneTKH;

npoMe>KyTOYHble (HHTePCTHU.HanbHble) KneTKH;

10- pa3BHBalOW.aS1CS1 CTpeKaTenb-

11 - CTpeKaTenbHaS1 KneTKa; 12- YYBCTBHTenbHaS1 (ceHCOpHaR) KneTKa

256

:/oo!lozun

0,ll;HMM H3 HMX, ycsaHBaCTCJI nceivrn 'lJ!euaMM KOJIOHMM. )1pyraJI rpyrrrra oco6ei1 B KOJIOHHM -

6JiaCTOCTHJIM. Ha poToBo:vr CTe-

6eJihKe y HMX HeT pTa M IIIYIIaJiel1. Ha 6J1acTOCTHJrnx

KOJIOHMH

BhIIIOqKQBhIBalOTCH

Me,n;y3hI, KOTOpbre 3aTeM OTpbrnaIOTCH H Be,ll;YT IIJiamuou~11i1 06pa3 )f01.1HH.

Y

:vre,n;y3

pa3BMBaIOTCH IIOJIOBbie :meJie3hl --- l'OHa,ll;hI,
KOTOph1e o6pa3yIOTCJI B 3KTO,n;epMe (Ha BOrHyTOM IIOBepXHOCTH rro,n; pa,n;HaJibllhIM11
KaHaJiaMM HJIH Ha poTOBOM CTe6eJ1hKe).
Pa3,n;eJihHOIIOJibie, rroJioBoii ,n;ttMopcptt3M ue
Bhipa:meH. IloJIOBOe pa3MHO;KeHHe MOpCKMX rn,n;pon,n;oB CBJI3aHO c IIOJIBJieHHeM rroJIOBOro IIOKOJieHHH -

rn,n;pott,IJ;HhIX Me,n;y3,

Cmpoenue zuiJpo.MeiJyJbL. TeJio Me,n;y31>r
CTy,n;eHHCTOe, HMeeT cpopMy 30HTHKa HJIH
PMC.

1-

105.

Ri1l.IO;

KOJIOKOJia. Ilo KpaIO 30HTHKa cBeIIIHBaIOT-

>K113H8HHbli1 l.111Kn 06en1111:

2-

nnaHyna;

3-

KOnOHl1R non11-

CJI lllYIIaJihl1a co cTpeKaTeJihHhIMH KJieTKa-

nos c 6naCTOCTl1n8M, Ha KOTOPOM Bl1AHbl pa3Bl1BalOW,118CR M8AY3bl;

4-

MH. Ha BOrHyTOH CTOpoHe TeJia, B IIeHTpe,
HaXO,ll;llTC.ff poT, KOTOphIM HHor,n;a paCIIOJla-

M8/J,y3a

raeTCJI Ha ,ll;JIHHHOM POTOBOM CTe6eJibKe.
lUYIIaJihl1aMH Me,n;y3a JIOBHT ,n;o6hJtfY' KOTOpaJI IIO,ll;XBaThJBaeTCJI POTOBhIM CTe6eJihKOM 11 3aTeM rrporJiaThIBaeTcJI. ll30 pTa IIHllla rrorra,n;aeT B meJiy,n;OI<, pacrroJiomeHHhIM B 1leHTpe TeJia rro,n; KyrroJioM. OT Hero oTxo,n;JIT rrpJIMhie, He BeTBHIIIHecJI
pa,n;HaJihHhie KaHaJihI, Brra,n;aIOIIIHe B KOJlbl1eBOM KaHaJI, OIIOJICbIBaIOlllHM Kpai1
30HTHKa Me,n;y3hl. IlepesapeHHaJI IIHIIIa IIOl'JIOlllaeTCJI KJieTKaMH 3HTO,n;epMhl.
CJiomHaJI racTpaJihHafl IIOJIOCTh Me,n;y3 Ha3hIBaeTCJI racTpOBacKyJIJipHoii c11cTeMoii • .[l;BH.iKYTCH Me,n;y3hl «peaKTHBHO»' qeMy cnoco6cTBYeT COKpaTHTeJibHaJI
KOJlhl1eBaJI CKJia,n;Ka 3KTO,n;epMbl no KpaIO 30HTI1Ka, Ha3bJBaeMaJI «napycoM».
Ilp11 paccJia6JieHHH napyca BO,n;a 3axo,n11T no,n; KyrroJI Me,ny3hI, a np11 ero cm<pan1eHHH BblTaJIKHBaeTCJI, IIPH 3TOM Me,n;y3a ,ll;BH.iKeTCJI Bnepe,n; TOJJqKaMH KYIIOJia.

JKuJHeHHblU 4u1CJL iuiJpouoa 06eJLuu. M3 orrJio,n;oTBopeHHo:H HM1leKJieTKH
06pa3yeTcH ,n;syxcJio:HHaJI JI11q11HKa -

IIJiaHyJia, c racTpaJihHoii noJIOCThIO BHY-

TpH. IlJianyJia IIJiaBaeT nptt IIOMOlllH peCHHqeK, 3aTeM oce,n;aeT Ha ,l.IHO H npeBpaIIIaeTCH B IIOJIHII, KOTOpbIH nyTeM JIOqKOBaHHJI 06pa3yeT KOJIOHHIO. OT KOJIOHHH IIOJIHilOB OTIIOqKOBhJBaIOTCJI He60JihIIIHe Me,n:y3hl, CIIOC06Hbie pa3MHOmaTbCfl rroJIOBhIM 11yTeM. TaKHM o6pa30M, y MopcK11x KnmeqHonoJIOCTHhIX
IIPOHCXO,ll;HT qepe,n;oBaHne IIOJIHIIJIOH,llHOro ( 6ecnoJioro) H Me,n;y30H,D;HOro ( IlOJIOBoro) IIOKOJieHHH (pHc.

105).

Knacc C4111<J>o111AH&1e, 111n111 C4111<J>oMeAy3&1
KJtacc C4u<PouiJHble, uJLu c4u<Po.MeiJyabt JIOCTHhie. ll3BecTHO oKoJio

200

KpyuHhie MopcK11e KttmeqHono-

s11,n;os. BoJibIDaH qacTh .iKH3HeHrrnro l1HKJia

KJ£acc KopaJ£J£06bte noJ£unbi

257

npoXO,lJ;HT B ~OpMe nJiaBalOm;HX Me,n;y3, ~a3a nomma KpaTKOBpeMeHHa HJIH OTCYTCTByeT. IloJIHilbI MeJIKHe H 06pa3yIOT

He60JihlllHe

KOJIOHHH.

Pa3MHomaroTCH 6ecnOJihIM nyTe.M -

norrepeqHhIM ,z:r;eJieHtteM

c o6pa30BaHHeM MOJI0,11;bIX Me,z:r;y3. 06m;ttif IIJiaH CTpoeHHH
TaKOM

me,

KaK H y

rtt,z:r;poMe-

,z:r;y3, HO HMeIOTCH OTJilfqHTeJihHble oco6eHHOCTH: KpynHbte
pa3.11tepbt, CUJlbHO pa3Buma.H
Me30tJle.H, omcymcmBue napyca, a6UJICeHUe COKpaut,eHUeM
cmeHOK 30HmuKa; pa3aeJienue
nuw,eBapumeJibHOU cucmeMbt
Ha a6a omaeJia r.JIOTKY
H meJiy,z:r;oK, HMeIOm;Hif qeTbipe
pa3,z:r;eJihHhJe KaMephI c

racT-

P111c. 106. >K113HeHHblH L\11Kfl MeAY3bl Aypen1111:
1 - HHL\O; 2 - nnaHyna; 3 - CL\l1¢>11crnMa; 4 - no'lKosaH11e; 5 -- CTpo611JJHL\l1H; 6 - 3¢>11pa; 7 - B3pocnaH
MeAy3a

pa.JibHbIMH HHTHMH ,11;JIH yBeJIHqeHHH nttm;eBapHTeJibHOM IIOBepXHOCTH. OT .JKeJiy,z:r;Ka OTXO,ll;HT BOCeMb BeTBHID;HXCH M BOCeMb HeBeTBHID;HXCH KaHa.JIOB, Brra,n;arom;ttx B KOJihll;eBOM KaHaJI
(racTpOBaCKYJIHpHaH CHCTeMa), T. e. mim;a rrepe,n;BttraeTCH no BCeM yqacTKaM
TeJia;

ycJlOJICHeHue nep6HOU cucmeMbl 3a cqeT CKOIIJieHHH HepBHbIX KJieTOK
optaHbt 'lYBCmB 06paay10m KO.JltnJieKCbt - pona-

(raHrJIHeB) no Kpa10 30HTHKa;
JIHH (y aypemu1 -

BOCeMb) c rJia3KaMH H CTaTOll;HCTaMH. I'Jia3a c:u;11~0Me,n;y3

npe,n;cTaBJieHhI rJia3HhIMH ny3hipbKaMH c poroBtt:u;eif n

xpycTaJIHKOM, a He

rJia3HblMH HMKaMH, KaK y r11,n;pOH,ll;HblX Me,z:r;y3. 0,z:r;HaKO rJia3a o6Jia,n;aIOT CBeTo'IYBCTBHTeJibHOM ~YHKD;Heif. Ha yKopo'IeHHhlx m;ynaJih:u;ax pacrroJiaraIOTCH
opraHbl XHMHqeCKOrO qyBCTBa -

o60HHTeJihHhJe HMKH.

Pa30eJlb1WnOJlbte. I'o-

Ha,ll;bI o6pa3yIOTCH B 3HTO,n;epMe Ha HH.JKHeif IIOBepXHOCTH KapMaHOB .JKeJiy,n;irn.
OnJio,n;oTBopeHHe Hapy:arnoe.

JKu3neHHbtu u,uKJI cu,u<Po.11teoy3bt AypeJiuu (pHc.
HHH H3 3HrOThl pa3BHBaeTCH JIHqHHKa -

106). IlocJie OIIJio,n;oTBope-

IIJiaHyJia. 0Ha OJiaBaeT, oce,z:r;aeT Ha

AHO H, npHKpeIIHBillHCb K cy6cTpaTy' IIpeBpam;aeTCH B O,ll;HHQqHblM IIOJIHII cn;mpHCTOMy, KOTOpbIM pa3MHO:lKaeTCH IIyTeM IIQqKQBaHHH. Ha TeJie Cll;H~HC
TOMbI IIOHBJIHIOTCH IIOIIepeqHbie rrepeT.H./KKH H ~opMHpyeTCH CTpo6ttJia. IlyTeM
UOIIepe'IHOro ,n;eJieHHH (cTpo6HJIHD;HH) o6pa3yIOTCH MOJIO,ll;hle Me,n;y3hl (3~Hpbl),
ltOTOpbJe IIJiaBaIOT H rrpeBpam;aIOTCH BO B3pOCJlblX Me,n;y3.

Knacc Kopannoeb1e nonMnb1
KJiacc KopaJIJIOBbte noJiunbt -

MOPCKHe KOJIOHHaJihHhle, peme o,n;HHOq-

Hhle IlOJIHIIhl, pa3BHBa10m;11eca 6e3 CMeHhl IIOKOJieHHM. lf3BeCTHO OKOJIO

258

6

3001tozun

Thrc. s11,n;os. Ha1160J1ee IIInpoKo pacrrpocTpaHeHbI o,n;11HoqHbre KopaJIJihr (aK-

TMHn11) M KOJIOHHaJibHbie (KpaCHbIM, •repHbIM).
0TJill'IMTe.'lbHhie oco6eHHOCTM (OT r11,n;pOl1,L(HbIX KllllleqHorroJIOCTHbIX):
TeJio u;nJmtt,n;pnqecKoe, MeIIIKOBH,L(Hoii ct>opMhI, CTa,L(HH Me,n;y3LI oTcyTCTByeT.
Pa3Mephr KopaJIJIOBhIX noJIMIIOB KpynHee, cH.'lhHO pa3BMTa Me3orJieH.

y

MHOI'HX BH,ll;OB cKeJreT H3 M3BeCTM H.'111 pororro,ll;06Horo Beru;ecTBa, pacrro-

JiaraeTCH BHYTPM TeJia mrn cHapyarn (y aKTHHHH cKeJieTa HeT). MLrIIIeqHhre
KJieTKH rrepeMeru;aIOTC.H IIO,L( 3KTO,D;epMy ll o6pa3yIOT MYCKYJibHbIH CJIOH.
JiyqeBaH CHMMeTpH.H HapyIIIeHa, Ha6JIIO,L(aeTC.H rrepexo,n; K 6nJiaTepaJibHOH
cHMMe'l'PHH. IIo rrpnttu;nny cnMMeTpHn ,n;eJIHTCH Ha IIIecTHJiyqeBLre (qncJio ru;ynaJieu; KpaTHO IIIeCTH JI€II -

aKTHHHH) H BOCbMHJiyqeBbie (Bcer,n;a BOCeMb ru;yna-

KpaCHbIM, 6eJibIH KOpaJI.'I) (pHC.

107).

CJio.JKHaa inuueqHaH rroJIOCTh. 3KTo,n;epMMhHM rJIOTKa Be,n;eT B racTpMhHyro
IIOJIOCTh, KOTOpaa paa,n;eJieHa pa,n;HMbHbIMH rreperopo,L(KaMM -

cenTaMH (60KOBbie

CKJia,L(KM 3HTO,D;epMbl). OHM rrpupacTaIOT K rJIOTKe, a HH:>Ke rJIOTKH He CMhIKaIOTC.H,
o6paayH meJiy,n;oK. HaJiuque cenT yBeJiuq11BaeT nuru;eBapuTeJihHYIO noBepxHOCTh.
HepBHaa cucTeMa o6paayeT rycToe cnJieTeHHe Ha poTOBOM ,n;ucKe. Paa,n;eJibttonoJibie. fotta,n;hr ct>opMHPYIOTCH B 3HTo,n;epMe. Pa3MHO:>Kettue 6ecnoJioe (y KOJIOHMaJihHhIX TBopemrn -

rroqKoBaHMe, y o,n;MHOqHhIX -

,n;eJieHHe) u rroJioBoe. OnJio,n;o-

B racTpaJihHOH IIOJIOCTH :>KeHCKOH oco6u.

PaaBHTHe c MeTaMOPcP030M (JiuqMHKa -

nJiaHyJia), qepe,n;oBaHMH noKoJie-

HMH HeT.

3na'lenue ICUUlC'lHOllO.!WCmHbtX: 3BeHO B u;en.ax IIHTaHH.H MOpCKllX 6uou;eH030B; rrorJioru;eHHe B3BeIIIeHHOH opraHMKH H oq11ru;eHue MopcKoii BO,L(hr; KOpaJIJIOBhre IIOJIHilhI yqacTBYIOT B KpyroBopoTe Ca2+ B 6uocct>epe H o6pa3oBaHHH
oca,n;oqHbIX nopo,n;, PHcPOB, 6apbepoB, OKpymaroru;ux OCTpOBa, a TaK:>Ke aTOJIJIOB
(KopaJIJIOBbie OCTPOBa B ct>opMe KOJibIIa c JiaryHOH rrocepe,L(HHe); ynoTpe6JieHHe
B rrnru;y (KHTaii:, .HrrOHHH), H3I'OTOBJieHue npe.ll;MeTOB HCKyCCTBa (KopaJIJibI),
HCIIOJib30BaHHe B KaqeCTBe CTpOHTeJihHbIX MaTepHaJIOB (ll3BeCTb), B Me,L(HI!HHe
,L(JIH noJiyqeHHH 6HOJIOrHqecKH aKTHBHbIX Beru;eCTB.

THn nJIOCKHe qepBH
Ocno6Hbte apoMopfjJo.Jbt: ,n;ByCTopoHHHH CHMMeTpHR TeJia, IlOHBJieHue Me3o,n;epMbI, pa3BHTHe CHCTeM opraHOB.
lfaBeCTHO OKOJIO

12

TbIC. BH,ll;OB. lIJIOCKHe qepBM o6Jia,L(aIOT 6oJiee BbICOKHM

ypoBHeM opraHuaau;uu, qeM HH3IIrne MHoroKJieToqHhle. CBo6o,n;Ho.arn:Byru;He o6HTaIOT B MOPHX, npeCHblX BO,L(OeMaX, BO BJia:>KHbIX MeCTax cyIIIH, HO 6oJibIIIHHCTBO BH,L(OB napa3HTbl :>KHBOTHbIX ll qeJioBeKa.
XapaKTepHble npu3HaKH:

1.

MHoroKJieToqHwe, TpexcJIOHHhle :>KHBOTHhle. T1rnHH H opraHhI ct>opMMpyroTCH H3 Tpex 3apo,n;wIIIeBbIX JIMCTKOB: Hapy:>KHOro BHyTpeHHero -

2.

3HTO,L(epMbl H cpe,n;Hero -

3KTO,n;epMbI,

Me3o,n;epMbI.

,Zl;ByCTOPOHHH.H (6HJiaTepaJihHaH) CMMMeTpM.H TeJia, B03HHKHOBemre KOTOpoll: CBJl3aHO c aKTHBHbIM rrepe,n;BH:>KeHHeM.

Tun n.riocKue 'tep6u

259

2

6

a
P111c. 107. a. CxeMa CTpoeHlll~ wecrnnyyesoro KOpannosoro non111na: 1 - w,ynanbL.1,a; 2 - pornsoe
oTBepcrne; 3 - rnoTKa; 4 - cemb1; 5 sn~Y111sa10w,111e M~rK111e

nOAOWBeHHa~

nnacrnHKa; 6 - YaweYKa; 7 - CKnepocemb1,

TKaH111 non111na (8) s racTpanbHYIO nonoCTb.

6. CxeMa CTpoeH111~ BOCbM111nyYesoro

Kopanna: 1 - w,ynanbLl,a; 2 - pornsoe oTBepcrne; 3 - rnoTKa: 4 - cema; 5 - Me3eHrnp111anbHb1e H111rn; 6

-~i:1L.1,a

3.

TeJio JIMCTOBM,n;Hoif MJIM JieHTOBM,n;Hoif ¢opMhI, yrrnoru;eHHoe B crrMHH06prornHOM (,n;opcoseHTpaJihHOM) HarrpasJieHMM, xoporno BhrpameHbI rrepe,n;HMH M 3a,n;HMH KOHIJ;hI.

4.

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y

JieHTOqHbIX qepse:H ,n;Ba CJIOH MhIIIlIJ;.

y

cocaJibIIl;HKOB H JieHTOq-

HhIX qepse:H HMeeTCH IIOBepXHOCTHhIH aKTMBHO rrorJioru;aroru;n::H KYTHKyJIHpHbIH CJIOH -

5. y

TeryMeHT.

rrapa3HTHqecKMX qepse:H KJieTKH 3IIHTeJIMH (Koma) BhI,ll;eJIHIOT seru;e-

CTBO, 06pa3yroru;ee IIJieHKY -

KYTHKyJiy.

y

CBo6o,n;HO:>KMBYIIl;MX BH,ll;OB

O,ll;HOCJIOHHbIH peCHHqHhIH 3IIHTeJIHH.

6.

IloJIOCTh TeJia OTCYTCTByeT, rrpoMemyTKH Me:m,n;y opraHaMH 3aIIOJIHeHhI
pbIXJIOH coe,n;HHHTeJihHOH TKaHhIO -

rrapeHXHMOH,

BbIIIOJIHHIOru;e:H

orropHyIO, aarracaroru;yro, BhI,n;eJIHTeJibHYIO <l>YHKIJ;HH.

7.

IInru;esapn:TeJibHaH CHCTeMa COCTOHT H3 ,n;syx OT,n;eJIOB: rrepe,n;Hero (3KTO,n;epMaJihHOro) M cJierro 3aMKHyToro cpe,n;Hero (aHTo,n;epMaJihHoro). KnrneqHHK MomeT 6hITh pa3BeTBJieH, qTo o6JierqaeT rrpou;ecc pacrrpe,n;eJieHHH rrpo,n;yKTOB rrnru;esapeHHH B rrapeHXHMe. HerrepesapeHHhre ocTaTKM rrn:ru;n Bhl6pachIBaroTcH qepea poT.

Y JieHToqHbIX qepse:H rrn:ru;esapn:TeJihHaH cn:cTeMa

OTCyTCTByeT.

8.

HepBHM CMCTeMa CTBOJIOBaH, JieCTHttqHOro THIIa (opToroH), COCTOMT H3 rrapHhIX M03rOBbIX raHrJIHeB (CKOIIJieHHH HepBHhlX KJieTOK) H OTXO,n;Hru;HX OT
HHX rrpo,n;OJibHhIX CTBOJIOB (KOHHeKTHBbI), coe,n;n:HeHHblX Mem,n;y co6o:H rrorrepeqHhIMH TH)KaMH -

KOMHCCypaMH.

3001wzua

260
CpaBHHTem,HaH xapaKTepucTuKa KJJaccoB Knme'IHOIIOJIOCTHhlX
fll)J.pOH)J.Hhle

C1\H<}lo11)J.Hh1e

fII)J.pa,rII)J.pOII)J.HaH

AypeJIIIH, 11;1-ianeH,

MC,ll,y3a -

KopnepoT

Oco6eHHOCTli
CTpoeIHIH
n o6paaa
JKH3HH
I Ipe)l.eTaJJIITeJI11

IIOJHl.II

06eJIIIH

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IIOJJJ11lbl

AKTIIHl1H, KpaeHbIH
KopaJIJI, Mopc1we rrepo,
aJibIJJl.OHap11H, porosoH:
I<OpaJIJI

Cpe)J.a o6IITa-

MopH, rrpeenh1e so-

HIIH

)l.bl

06paa .iKII3HII

IIp1mperrJieHHhrH:,

Mo pH

Mo pH

lJJiaB8lOII.\IIH

II pIIKpennennhlrr

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Pa)J.IIa.JihHaH II qaeTl1'l-

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Cl1MMCTPIIH

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<I>opMa TeJia

I1HT8Hl'le

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noJI11n II Me,uyaa)

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IlOJIIIII

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lIJI8HKTOHHblMII

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HbIMl1, peme KpyrrHhlM.H
:>K11BOTHblMH ( aKTl'IHlfH ),

Mil.

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faeTpaJibH8H IlOJIOeTb e

Ha.H elfCTeM8

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rreperopo,uKaM.H (cerr-

)J.OK, I' JIOTKl1 HeT, y

Mlf 11 e11eTeMa pa-

T8Ml1), I'JIOTKa

Me)J.ya -

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pa)J.IIaJib-

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TCJ18

TeJia

HepsHaH enc-

,l],mp<PyaHaR, y Me-

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HepBIIOe

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Y IIOJil1TIOB HeT, y

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Te Ma

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Paa,n;eJihHonoJibre

PaaJJ;eJihHOIIOJibie

KJ£acc PecnU'tl!bte 'tep6u

f11.n,p011 1111L1e
Oco6enuocT11
cTpoemrn
H o6pa3a
:am:3HH
06pa3osa1rne I B 3KTo.n,epMe
I
noJIOBhlX KJieTOK

261

Cr~mpott.n,Hhie

i

H'.opa.:-rJ1oahIC
IlOJIHilhl

I

CTa,!\tt:.II pa3nuTH.II

5Hi:~o

- JIH'IHHKa (rrnaHyJia) no,1un - Me11Y3a (y ru.n,pbr:
.!Ill~O -- IIOJIHII)

B 3HTo.n.epMe

B 3HTo.n,epMe

Hii~o - m1qmma
(1IJ1aHyJia) - IIOJIUII
(c~mflHCTOMa) - CTpO6u.1a - Me.n,y3a (3<lmpa)

Hii~o - Jiuqmma
(rIJiaHy,1a) -- rro.1ttu

9. OpraHhr qyscTB Han6oJiee paasnThI y cso6o,n;Homnsyrn;nx sn.n;os, y KoTophIX MMeIOTC.H CBeToqyBCTBMTeJibHhie rJia3Kll, opraHhl paBHOBeCM.H - CTaTOIIHCThI, qyBCTBMTeJibHbie OKoHqaHH.H - ceHCMJIJihI: OC.H3aTeJihHbie KJieTKll ll opraHhI XllMMqecKoro qyBCTBa. y rrapa3MTOB opraHhI qyBCTB pe.n;y1IHpOBaHhI H HMeIOTC.H TOJibKO Ha CTa,D;llll cso6o.n;uorrJiaBaIOrn;Hx JIHqllHOK.
10. Bh1,n;eJIHTeJihHa.H CHCTeMa rrpoToHecppn,n;HaJihHOro TMrra. CocTOMT H3 crreIIHaJIMaHposaHHhIX TepMHHaJihHbIX (asea,n;qaTbIX) KJieTOK, paa6pocaHHhIX no see My TeJiy' H CllCTeMhI OTXO,ll;.Hlll;MX OT HHX BeTBHCThIX KaHa.IIbIIeB (aKTo.n;epMaJihHOro rrpoHcxom.n;eHnH). IlpoToHecppH.n;HH HaqHHaIOTCH
B rrapeHXMMe asea.n;qaToii KJieTKOH c rryqKOM peCHHqeK, OT KOTOpoii OTXO,ll;HT TOHKM:ii KaHaJieI1. KaHaJihllhI CJIHBaIOTCH B 6oJiee KpyrrHhie H oTKphrnaroTcH Ha aa,n;He:ii rrosepxHOCTH TeJia BhI.n;eJIHTeJihHO:ii rropoii. IlpoTOHecppH.n;Ha.H CHCTeMa y.n;aJI.HeT rrpo.n;yKThI ,D;HCCHMllJIHIIMH ll peryJinpyeT OCMOTHqecKoe ,n;aBJieHHe.
11. ll:hIXaTeJihHaH CMCTeMa OTCYTCTByeT. y CB06o.n;HOiKMByrn;nx raaoo6MeH
ocyrn;ecTBJIHeTc.H rryTeM .n;Hcpcpyann qepea scro rrosepxHOCTh TeJia. Y rrapaanTOB .n;hrxattne aHaapo6Hoe.
12. KposettocHaH cHcTeMa oTCYTCTByeT.
13. IlOJIOBaH CllCTeMa. IlJIOCKHe qepBM - repMacppo.n;MTbI. IloJIOBbie meJie3hI
nMeIOT cJIOiKHhre swso.n;ttwe rrpoTOKll. PaaMHomeHHe rroJiosoe (y HeKoTophrx oco6e:ii qepe.n;yeTc.H c rrapTeHoreaeaoM) H 6ecrroJioe (rrorrepeqHbIM .n;eJieHHeM). OrrJio.n;oTsopeHne BHyTpeHHee (rrepeKpecTttoe).
14. PaasnTne rrp.HMoe (cso6o.n;ttomnsyrn;He BH.n;hl) MJIH c MeTaMopcpoaoM (rrapaanT11qecKne). :IB'.H3HeHHhie IIHKJihl co CMeHOH X03HeB H qepe.n;osaHMeM
IIOKOJieHMH.
Tun llJLocH:ue 'lep6u BKJiroqaeT: KJLacc PecHU'lHbte 'lepBu, uJLu myp6eJL.Rpuu.
KJtacc CocaJLbUJ,UKU, uJLu mpe.MamoiJbt, KJLacc JI enmo'lHbte 'lepBu, uJLu u,ecmoiJbt.

Knacc PecHH"IHble "1epe1r1
KJtacc Pecnu'lHbte 'lepBu - rrpellMYIIIeCTBeHHO cso6o.n;Homnsyrn;ne. BoJihIIIMHCTso BH,ll;OB BCTpeqaeTC.H B MOp.HX H rrpeCHbIX so.n;ax. lfaneCTHO 6oJiee 3 500 Bll.n;OB.

262

30011.ozun

Ilpe,n:cTaB11TeJ1h -

pu.R -

6eJta.R (.1110JLO'l1WR) n.rtaHa-

HeOOJibllIOH xm:IJ;HhIH IIJIOCKMM '-IepBb,

0611TaeT B rrpecHh!X BO,n:OeMax, rro.n: KOpHraM11,
JIMCThHMM.

TeJio ynJI01u;euttoe,

JIMCTOB11.n;no:i1

cpopMbI, IIOKpbITO peCHM'-IHhIM 311MTeJIMeM, KOTOpbIH co.n;epmMT MHOmecTBO meJie3,
IIJ;MX CJIM3h.

7
8
9
JO
11

18

PacnrnpeHHhIH

BhI,ll;eJI.HIO-

rrepe,n:HMH

OT,n:e.1

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OC.H3aTeJihHhie ru;y-

IIaJihIJ;a, opraHhI paBHOBeCM.H (cTaTOIJ;MCThI). Cneu;ttaJihHhIX opraHOB rrpttKpenJieHM.H HeT • .ll:BmI<eHMe

ocyru;eCTBJI.HeTC.H

3a

C'-IeT

COKparu;eHMH

MhlIIIU KO)KHO-MYCKYJihHOro Mern:Ka, a TaKme 3a
C'-IeT peCHM'-IHOrO 3IIMTeJIM.H (pttc.

108).

KomHO-MYCKYJihHhIH MellIOK TMIIM'-IHOro CTpoeHM.H. IlpocTpaHCTBO Mem.n;y MYCKyJiaTypoiI M BHYTpeHHMMM oprattaMM 3aIIOJIHeHO rrapeHXMMOH.

Iluw,eBapumeJLbHa.R cucmeMa Ha'-IMHaeTc.H
c pacIIOJIOllieHHOro Ha 6pIOIIIHOH CTOpOHe TeJia

Pitc. 108.
1-

Hap1.11.1:
POCTbl;

CxeMa CTpoeH1.1s:i nnaLl.(ynMbl(eBl.1AHbl8 Bbl-

2-

ma3a;

4-

raHm1.1Vi;

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5-

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3-

npOAOnbHblH HepB-

6-

nonepe4Hble Heps-

Hble nepeMbl'lKl.1;

7-

rJlOT04HblH KapMaH;
BOA;

10 -

M03rQBOH

nepeAH515l B8TBb Kl.1-

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9-

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pornsoe ornepcrne;

11 - ceMeHHHKl.1; 12- COBOKyn1.1TenbHblH opraH;
Kl.1We4HHKa;
CT1.1e;

15 -

KJlOaKa;

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s:iVil(eBOA;

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pOTOBOro OTBepCTM.H,

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y

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K1£acc CocallbllfUKU, u1£u mpe;namoi11>i

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263

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no 6oKaM TCJia_.

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S:IMIJa. qepe3 pa3pbIB B CTeHKe KOKO Ha IIJiattapHH BhIXO,ll;S:IT uapymy.

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B

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a CBHHhH TOJihKO npoMe)KyTO'IHhIM.
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qeJIO-

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6

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268

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P111c. 111.

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)J.OJI.iKHhI rrorracTh B BO)J.y. H3 o6oJIO'IKH Hii:u;a ocB06om)J.aeTCH oHxoc<)>epa (KopaD;H)J.HH) -

CB060)J.HOIIJiaBaIOID;3H JIHqHmrn, KOTopaH rrorra)J.aeT B opraHH3M rrep-

BOI'O npoMemyToqHoro X03HHHa .a;HKJiorra, r)J.e npespaIIJ;aeTcH B qepBeo6pa3HyID JIH'IHHKY -

npou,epKouiJ.

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u;epxon:.n; H3 KHille'IHHKa npOHHKaeT BO BHyTpeHHHe opraHhl H MycKyJiaTypy
pb16hI n: npeBpaIIJ;aeTc.H B <)>HHHY

nJtepou,epKouiJ. Y qeJioBeKa, C'heBrnero phr6y,

nJiepou;epKon:.n; pa3BlrnaeTC.H B noJioBo3peJioro JieHTeu;a.

JKu3neHHbtU U,UKJl 3XUHOKOKKa. 0KOH'IaTeJihHhIM X03.HHH JIHCHD;a; npoMeJKyTO'IHhIH -

co6aKa, BOJIK,

qeJIOBeK, KOPOBa, OB.a;a. IlOJIOB03peJia.H CTa,n;mr

napa3HTHpyeT B KHille'IHMKe OKOHqaTeJihHOI'O X03.HHHa H COCTOHT H3
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5-6

3-4 qJie-

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-

u,ecmoiJo3aMu.

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H3BecTHO oKoJio

100

ThIC. BH)J.OB, cpe,n;u KOTOphIX 60Jihrnoe 'IHCJIO xa:K

napa3HTH'leCKHX, TaK H CB060)J.HO.IRHBYID;HX BH)J.OB, KOTOphle 3aCeJISIIOT MOpSI,
npeCHhle BO)J.hl H rroqBy. IIapa3HTH'IeCKHe qepBH BCTpeqaIOTC.H y BCex MHOI'O-

Tun Kpyz1t11te, u:iu nep6u'inono1tocmn11te, 'iep6u

269

KJieTO'IHhIX mIIBOTHhIX, a TaKme y MHOrIIx pacTeHnii. KpyrJihre T.IepB11 Bhillle
nJIOCKIIX 11epBeii ITO ypOBHIO opraHII3aU:HII.
XapaKTepHhie rrpII3HaK11:

1.

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2.
3.

CTeirna TeJia BepeTenoBII~Hoii cpopMhI, B rronepe11HOM ce11eH11II oKpyr-

06Jia;:J;aIOT 6mrnTepaJibHOH CIIMMeTpIIeii TeJia.
Jioe, c 3aOCTpeHHbIMH

4.

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5.

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6.

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ll3

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mn~KOCTH, Haxo~.Hru:eH:c.H no~ BbICOKHM ~aBJieHJfeM (rH~pocKeJieT).
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~YKTOB o6MeHa.

7.

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B o~HOM nanpaBJieHHH If JiyqIIIe BcachrnaeTC.H. Pa3JIHqaIOT rrOJIOCTHoe
n npncTenoqnoe nnru:eBapenne.

8.

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9.

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10.

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napa3H-

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11.

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y

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y

na-

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12.
13.

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y

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14.

Pa3BHTHe np.aMoe. :IKn3HeHHhIH I!HKJI napa3HTH'leCKHX neMaTo~ 11acTO

270

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corrpoBoJK,n;aeTCH CMeHOH X03.HeB, cpe,n; o6HTaHIU:I, MMrparrII.HMM B opraHl13Me X03RHHa.

Tun

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400 TbIC.

3TO caMhIH o6rrmp-

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AclCapui}a
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112, 113).

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20-30 CM y

caMOK,

15-20 CM y

caMu;oB, oKpyrJioe, opraHbI np11-

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· 3

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(20-25 'C)

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6

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a TaKme c so,n;oii. B

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PMc. 112. CTpoeH1-1e acKapHAbl: a - caMKa,
6 - caMeU,. 1 - ry6w; 2 - HepBHOe KOnbU,O;
3 - rnoTKa; 4 - cjlarou,mapHble KJJeTKH; 5 -

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no opraHI:I3MY (B Te'leHMe ,ll;BYX He-

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a OTTyµ;a no Jiel'O'lHOH apTepHH -

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271

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H qe,'IOBCKa

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272
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75

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PMc. 113.

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274

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6.

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11.

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275

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15.

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JIOBCK, KpyllHblH poraTblH CKOT) -

("IeJIOBeK) · - IlOJIOB03peJiaH

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pa H cpHHHa. 0HKOC<l:>epa pa3BJ1Da-

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j(btxame/lbHa.R cucme.Ma.

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3

2

IIIO pa3BHTyro ceTh. KHcJiopo.n; pacTBop.aeTc.a B CJIH3M, IIOKphrnaroru;eli T8JIO

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48pBS1: 1 -

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115)

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repMacppo,n;HThI. MymCKI18 IlOJIOBhie KJI8TKH o6-

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(10-11

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c pa3BMBalOru;MMMC.ff .aH:u;aMM.

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(nonMxeTbl)

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8

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Cmpoenue.

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116). CerMeHThI,
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Knacc lluR61m

279

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Ten~~;

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noA~~.

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KOCaS'I Mblwua; 11 -

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P111c. 116.

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400

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OHa npoKyChIBaeT KO/KY.

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IIannBIIIaHC.H xpoBtt Me.n;nu;ttHCKa.f! nH.f!BKa yBeJIH'-IHBaeTCH B o6'heMe B HeCIWJihKO pa3. 3arraca KpOBH en XBaTaeT Ha HeCKOJihKO MeCHIJ;eB.

280

300JWlUJI.

P111c. 117.

BHyrpeHHee c1poeH1-1e Me-

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3-

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8-

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npHCOCKa; 10 -

9-

3aAHflfl

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M8TaHe¢pi-iAHH; 12-

13- C8M8HHbl8 M8WKH;
14- C8MflnpoBOA; 15- peCHHYHafl BOPOHKa MeTaHeQ:>pHAHfl; 16- BnaranHUje;
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6oKoBafl naKyHa

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,[(hrxaHue ocyiu;ecTBJIReTcR Been rroBepxHOCThIO TeJia.
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a Hecpporropoii: - Hapymy. BopoHKa He
coe,n;MHJieTCR c KaHaJIOM MeTaHecj)pH,n;HR
M ;KH.ZJ;KHe npo.n;yKThI rrpOHHKaIOT H3 Hee
B KaHaJI rryTeM OCMOCa.
IIoJloBa.H c uc me Ma. fepMacppo,n;HThI.
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(phIXJIHT' 06orarn;a10T opraHH'-leCKHMH
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u npoHHKHOBeHHIO BJiaru, yJiyqrua10T
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3Ha'-leHHe.

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281

Tun MOJl,IZIOCKU, unu MRZKomenble

CpaBHHTeJILHaH xapaKTepucTHKa KJiaCCOB KOJIL'laT1.1x qepBeii

Oco6eHHOCTll
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06pa3 JKl13IfII

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282

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fepMacppo,11;IITbI

fepMacppO,Ul!Tbl

c Jlll'fHHKOH Tpoxo-

Ilpll.MOC 6e3 Jlll'lllH-

IlpHMOe 6e3 Jlli'lllll-

cpopou

Kl!

KM

H.f!TCJlhllhIC .f!MKll
IloJioBa.fl
CHCTeMa
Pa3BIITHe

CpaBHHTC.'IhHaH xapaKTepHCTHKa THDOB nJIOCKHx, 11:pyrJih1x H KOJih'faThIX qepBeif

Ilp!!3113Kll
<t>opMa TeJia

IJJIOCKI!C '!CpBl!

Y IIJIO~CHHb!C

Kpyr"1b1e qepBII
KpyrJIMe n rrorrepe'!HliKC

KOJ1h'IaTh!C '!CpB!!
0KpyrJIMe n rrorrepe'IHHKC, c 6oJiee BbirryKJ'IOH
CIIHHHOH ITOBepxHOCTblO

CerMeHTaIJ;H.fl

0TCYTCTByeT (HeKJIIO'fCHHe -

0TCYTCTBYCT

rra-

Pa3B11Ta Hapy:>Kna.fl H
BHYTPCIIH.fl.fl cerMeHTaD;ll.fl

pa3HTli'fCCKHe
JICHTO'lHble 'fCpB11)
Ilptt,z:i;aTK11

HeT

HeT

Te JI a

IIaparro,ll1111 11Jm pR,ll;hI
rrapHbIX

~CTHHOI<,

y

IIHHBOK IIPH.U3TKH OTCYTCTBYIOT
HaJIH'ftte

HeT

KyTttKyJia TOJICTaJI

KyTHKyJia TOHKaJI

KomHo-

3rrttTCJ111H,

fttrrn.uepMa,

3rrIITCJil1H, KOJibD;CBbIC

MYCKY JlbHb!H

TeryMCHT,

rrpo,n;OJihHbIC Mblll!IJ;bl

H rrpo,n;OJihHbIC MbllIIIJ;bl

KYTHKYJibl

MCIIIOK

rrorrepe'!Hhie,
IIpO,ll;OJibHhIC
II .uwaroHaJihHhIC
MbIIIII~bI

IloJIOCTh TeJia

0TCYTCTByeT, 33H.f!Ta KJICTK3Ml1
rrapeHX11Mbl

IIepB11'1HaJJ.

BTopH'IHaJI,cerMeHTHpoBaHHa.fl

Tun

283

MOJt1uocKu, u.n,u MRZKome/lbie

,
II.Jioc1rne 'IepBH

fJpH3H8!01

.il:mpcj:Jy3HbIH

TpaHcrropT

Kpyr,1i,1c •icpBH

Ko,1b'!aThJe •1epB11

-

.ll:11cj:Jcpy3HbIH

KposeHOCHaH cncTeMa

seru;ecTB
IIttru;esapn-

Ilepe;i;Hl1H

IIepe;i;HHH, cpe;i;Htttt

Ilepe,!J;HHH, cpe.n;HHH

TeJibH3H

11 cpe,!1;Hl1H OT,Ue-

H 33,!l;HHH OT,!l;eJibI

H 3a,!l;IIHll OT,ll;CJibI

CHCTCMa

JihI, 3a.n;Heii

Komnoc

KomHoe

KomHoe, MoryT 6bITh

IlpOTOHCcj:Jptt,D;Hl1

KomHhie meJie3hI, Bl1-

MeTaHecj:JpM;i;1111

KHIIIKH neT

.il:hIX3Hl1e

ma6phr
Bhl.n;eJI11TeJ1h-

,!1;0113MeHeHHhie rrpoTO-

HaH Cl1CTeMa

Hecj:Jpll,!1;1111
CTBOJIOBaH, oKoJiorJio-

HepsHaH

CTBOJiosaH

CTBOJIOBaH

CHCTeMa

JieCTHWIHaJI

JieCTHH'!HaH, c ,!l;BYMH

TO'lHOe HepBHOe KOJib-

(opToron)

6oJiee pa3Bl1TbIMl1 TH-

u;o 11 6pIOurnaH Heps-

maM11: CllllHHhlM H

Ha.H u;eno<rKa

6p!OII1HbIM
IloJIOBaH

I'epMacppO,UllTbl

Pa3p;eJrhnorroJibie

Pa3B11T11e

Pa3.zi;eJibHommh1e 11.JIH
repMacj:JpO,D;HTbl

Cl1CTeMa

IlpHMOe 11Jll1

C Jll1'-ll1HKOH

IlpHMOe

llJil1 npHMOe

C Jil1'!11HKOH

XapaKTepHhle rrpH3HaKir:

1.

MHoroKJieToqHhle, TpexcJIOHHhie mHBOTHhre.

2.

,[(ByCTOpOHH.H.H (6HJiaTepaJibHa.H) CHMMeTpH.H TeJia, KpOMe 6pIOXOHOl'HX

3.
4.

IlepBHqHopoTble.

MOJIJIIOCKOB (aCHMMeTpH.H).
IJ;eJIOMHqecKa.H IIOJIOCTb coxpaH.HeTC.H OKOJIO cep,a;u;a H IIOJIOBbIX meJie3.
IlpocTpaHCTBO Mem,a;y opraHaMH 3aIIOJIHeHO coe,ZJ;HHHTeJibHOH TKaHbIO.

5.

TeJIO HecerMeHTHPOBaHHOe -

COCTOHT H3 l'OJIOBbl, TYJIOBHIIJ;a H HOl'H HJIH

TYJIOBHIIJ;a H HOrH. IloKpbITO MaHTHe:H (KOmHa.H CKJia,a;Ka, CBHcaroru;aa co
crrHHhI). CHapyam MaHTH.H BhI.ZJ;eJI.HeT paKOBHHy. IloJIOCTh Mem,a;y TeJioM
H MaHTHeii: Ha3bIBaeTC.H MaHTHHHOH IIOJIOCTbIO.

6.

liMeeTC.H H3BeCTKOBa.H paKOBHHa, COCTO.HIIJ;a.H ll3 Tpex CJIOeB: HapymHOro -

poroBoro, cpe,a;Hero -

q>apq>0p0Boro, BHyTpeHHero -

rrepJiaMyT-

poBoro. Pa3JIHqaroT u;eJibHYIO H ,a;ByCTBopqaTyro paKOBHHhI, pa3JIHqHbie
no <jlopMe. PaKOBIIHa BhIIIOJIH.HeT <j>yHKu;mo 3aru;HThI TeJia OT BparoB
II MexaHHqecKHX IIOBpem,a;eHn:H; nrpaeT· PO.Jib Hapy:>KHOl'O cKe.JieTa, K KOTOpOMY rrpHKperrJI.HIOTC.H MhIIIIIJ;hI.
meT pe,a;yu;HpOBaThC.H.

B HeKOTOpbIX cJiyqaax paKoBnHa MO-

284

300JWZUR

7.

MycKyJiaTypa rrpe,D.cTaBJJeHa OT,D.eJ1hHhll\HI Mbllllll;8Mll, KOTOpbie Kperr.HT-

8.
9.

OpraH rrepe,n;mimemrn -

C.H K paKOBHHe.
nora (MycKy.TihHhIH BhipocT).

IImn;eBapttTeJibHa.H CHCTeMa COCTOHT H3 Tpex OT,n;eJIOB: rrepe,n;Heli:, cpe.n.Heli: H 3a,n;Heli: KHIIIKH. llMeIOTC.H CJIIOHHbie JKeJie3bl, OTKpblB8IOID;Hec.H
B rJIOTKy.

y

6pIOXOHOrHX B rJIOTKe HaXO)l;HTC.H crreu;HaJibHbIH opraH ,n;.1,H

COCKafornBaHH.H H nepeTHpamrn IlHJD;H -

TepKa (pa,n;yJia). o6pa30BaHH85I

MHomecTBeHHhIMH p.H,n;aMH nonepe'!HbIX KYTHKyJ1.HpHhIX 3y6'lttKOB. IlttID;eBo,n;, 'laCTO paCIIIHp.HIOID;HMC.H B 306, Bila,D;aeT B MeIIIKOBH,ll;HhlM me;1y,n;oK, .HBJI.HIOJD;HMC.H '!aCThlO cpe,n;Heii: KHIIlKH, B KOTopy10 OTKpblBaIOTC.H
rrpoTOKH ne'!eHH. Co6cTBeHHO cpe,n;H.H.H KHmKa nepexo,n;HT B KopoTKy10
3a,n;HIOIO KHIIlKy, OTKpbIBaIOID;YIOC.H aHaJibHbIM OTBepCTHeM B MaHTHHHYIO IlOJIOCTh.

10.

HepBHa.H CHCTeMa CTBOJIOBa.H -

,n;mpcpy3HO-y3JIOBOro THna, COCTO.HID;aH

H3 HeCKOJ!hKHX nap raHrJIHeB,

y

coe,n;HHeHHhIX HepBHbIMH CTBOJiaMH.

HeKOTOpbIX MOJIJIIOCKOB rrpOHCXO,ll;HT KOHll;eHTpau;H.H HepBHbIX raHrJIH-

eB c o6pa3oBaHHeM cJIOJKHoro roJIOBHoro M03ra.

11. y

Be,n;yID;HX aKTHBHblH o6pa3 JKH3HH xopomo pa3BHTbI opraHbI 'IYBCTB:

CJIOJKHO ycTpOeHHbie rJia3a, opraHbI OC.H3aHH.H (ceHCOpHbie KJieTKH), XHMH'leCKOro 'lYBCTBa ( occppa,n;HH) H paBHOBeCH.H ( CTaTOll;HCTbl).

12.

OpraHhI BhI,n;eJieHH.H -

«ITO'IKH» (Hecpp11,n;11H), HMeIOT BOpOHKY B oKo.10-

cep,n;e'lHOli: cyMKe H OTKphrnaIOTC.H BbI,ZJ;eJIHTeJihHbIM OTBepCTHeM B MaHTHMHYIO IlOJIOCTb.

13.

KpoBeHOCHa.H CHCTeMa He3aMKHyTa.H. KpoBh H3 cocy,n;oB Te'leT B rrpoMemyTKax MeJK,n;y opraHaMH. IJ;eHTpaJihHbIH opraH KpOB006paJD;eHH.H
cep,n;u;e, cocTOHT H3 o,n;Horo meJiy,n;otIKa H o,n;Horo-,n;Byx rrpe,n;cep,n;ttii:.

14.
15.

AhixaHHe ma6epHoe HJIH Jiero'!Hoe.
IloJioBa.H cttcTeMa.

Pa3,n;eJihHOIIOJihie (roJIOBOHorHe, ,n;BycTBopqaThie)

HJIH repMacppo,n;HThI. OT roHa,n; oTxo,n;.HT rrpoTOKH -

roHo,n;yKThI (u;eJio-

Mo,n;yKThI). OrrJio,n;oTBopemrn Hapymttoe HJIH BHyTpeHHee. Pa3BHTHe rrp.HMoe HJIH c MeTaMopcp030M (JIHqHHKa -

Tpoxocpopa, BeJIHrep, rJIOXH-

,ll;H.H).
Tttrr MoJIJIIOCKH BKJI10qaeT: K.!lacc Bp10xonoiue, K.1tacc /(eycmeop•tambte,
u.1tu II.1tacmutt'lamoJ1Ca6epttbte, K.!lacc I'o.1toeonoiue.

Knacc 6plOXOHOrMe MOnnlOCKM

K.1tacc Bp10xonoiue .MO.!l.!llOCKU KOB, H3BeCTHO 6oJiee

90

caMbili: MHoroqttcJieHHhIM KJiacc MOJIJIIOC-

TbIC. Bll,ll;OB. CBo6o,n;HOJKHBYID;He, o6HTaIOT B MOp.HX,

rrpeCHhIX BO,n;OeMaX H Ha cyIIIe. Ilpe,n;cTaBHTeJIH: BHHOrpa,n;Ha.H YJIHTKa, rOJiblH
CJIH3eHh, rrpy,n;oBHKH. PaKOBHHa u;eJihHa.H, crrttpaJihHO 3aKpyqeHHa.H, JIH6o oTcyTCTByeT (rOJibie CJIH3HH). Ha6JIIO,n;aeTC.H HapyIIIeHHe CHMMeTpHH BO BHyTpeHHeM H BHeIIIHeM CTpOeHHH.

BunoipaiJnaR y.1tumKa

(p11c.

118). Cmpoenue.

TeJio pa3,n;eJieHo Ha roJioBy,

TYJIOBHID;e 11 Hory. Ha roJioBe Haxo,n;.HTC.H ,n;Ba ID;yrraJihD;a -

opraHhI oc.H3aHH.H.

Jeaacc Bp10xonozue Mo11,11,10cKu

285

2

PHC. 118. BHyi-peHHee CTpoeH111e BlllHOrPaAHOi'i yn111TK111: 1 - neYeHb; 2 - KlllWKa; 3 - lKenyAO'leK
cePALl<l: 4 - npeAcepA111e; 5 - nerKoe; 6- aHanbHOe ornepcrne; 7 - PaAYlla (rnpKa); 8 - poT; 9 rnoTKa; 10-

y

n111~eBOA;

11 - HepBHble Y3Jlbl; 12- lKellYAOK; 13- HepBbl

OCHOBaHJi.H w;yrra.rreu; pacrroJiomeHbl rJia3a.

c

6pIOIIIHOH CTOpOHbl OTKphlBaeT-

C.H poT. TeJio 3aKJIIO<JeHo B cmrpa.rrhHYIO paKOBMHY, KOTopa.H MMeeT Beprrn:rny
M OTBepcTMe -

YCTbe, qepe3 KOTOpoe BbICOBbIBaeTC.H rOJIOBa, Hora M HHiK.H.H.H

qaCTb TYJIOBMru;a. 3a C<JeT CMJibHOro MYCKyJia-peTpaKTOpa B CJiyqae orracHOCTM
Hora MomeT 6b1cTpo BT.HrttBaThC.H BHYTPh paKOBMHhI. Ha 3a,n;tteii qacTM HorM 06pa3yeTc.H IIJiaCTMHKa, rrpOIIMTaHHa.H M3BeCThIO, KOTOpa.H rrpH BT.HrIIBaHMM rrpMKPhIBaeT, KaK KphIIIIe<JKoii, Bxo,n; B ycThe paKOBMHhI.

IluU{e6apumeJtbttaJl cucmeMa.
paCIIOJIOiKeHHOM B

B

rJIOTKe,

PacTMTeJibHO.H,ZJ;Hbie. C rrOMOill;hIO TepKH,

YJIMTKa COCKa6mrnaeT H

rJioTKY OTKpbrnaeTCH rrapa CJIIOHHhlX meJie3.

Y

rrepen1paeT IIMill;Y.

XHill;HhIX yJittTOK ceKpeT

CJIIOHHblX me.Jie3 co,n;epiKMT cepHyIO KMCJIOTY. 3a rJIOTKOM M,ZJ;eT 1rnw;esop;,
paCIIIMp.HIOilJ;MMC.H B 306, 3aTeM me.Jiy,n;oK, B KOTOpbIM Brra,n;aIOT rrpOTOKH rre'leHM. Ile<JeHh Bhl,ZJ;eJI.HeT <lJepMeHThI, paCTBOp.HIOill;Me yrJieBO,ZJ;bI, M BbIIIOJIH.HeT BCaChIBaTeJibHYIO <lJYHKD;MIO. Cpe,n;HH.H KMIIIKa o6pa3yeT rreTJIM B TyJIOBH~e M rrepexo,n;MT B 3a,ZJ;HIOIO KHIIIKY' 3aKaH'IHBaIOw;y10c.H aHaJibHhIM OTBep-

CTMeM Ha,n; rOJIOBOH.

KpoBettocttaJl cucmeMa

He3aMKHyTa.H. Cep,n;u;e cocTOHT M3 o,n;Horo rrpe,n;cep-

AHH M o,n;Horo meJiy,n;o<JKa. OT meJiy,n;o<JKa 6epeT Ha<JaJio aopTa, KOTopaH BeTBHTC.H M pacrra,n;aeTC.H Ha KaIIMJIJI.HpbI.

lfa JiaKyH

KpOBb co6ttpaeTC.H B BeH03Hble

CHHYChI, M3 KOTOpbIX rrorra,n;aeT B KaIIMJIJI.HPHYIO CMCTeMy CTeHOK MaHTMHHOH
UOJIOCTM, r,n;e o6oraw;aeTCH KMCJIOpo,n;OM M ITO BhIHOC.Hill;MM cocy,n;aM B03Bpaw;aeTCH B cep,n;u;e.

286

300.TWZUJl

BbtiJeJlum<'JlbltaJl cue me.Ma

rrpe,n;cTaBJieHa Herrapnorr rroq1wii MeTaHecpp11,n;11-

aJibHOro TIIITa, KOTOpaa o,n;HIIM KOHIJ;OM OTKpbrnaeTC}[ B OKOJIOCep.n;eqHyIO IIOJIOCTb, a ,n;pyrIIM -

B MaHTl1iiHyIO.

f( btXameJlbHGJl cue me.Ma.

JierKoe ( oco6bIH OT,[(CJI MaHTIIHHOH ITOJIOCT11),

CTeHKl1 KOToporo OITJieTeHbl KpOBeHOCHbIMl1 KaITIIJIJI.HpaM11, r,n;e rrpoIICX0,[(11T
ra3oo6MeH.

HepBHaa cucme.Ma

CTBOJIOBa.H ,n;mpcpy3HO-Y3Jiosoro TIIrra. CocTOl1T 113 rraT11 nap

HepBHbIX y3;ros: 1lepe6paJibHbIX ry; ITJieBpaJibHhIX -

11HHeps11py10T opraHhI qyscTB; rrel);a.'lhHbIX -

MaHTHIO; rrap11eTaJibHbIX II BHCI1epaJihHbIX -

Ro-

opraHbI ,ZJ;bIXaH11H,

occppa,n;IIH H BHYTpeHHHe opraHhI.

OpiaHbt ttyBcmB.

OpraH 3peH11H -

rJia3a: y OCHOBaHIIH IIIYIIMe1l (rrpy,n;oBHK); Ha

sepurnHe ,n;Jil1HHhrx IIIYIIMe1l (yJI11TKa). OpraHhI BKyca, oca3aHHH H 060HaH11a KopOTK11e IIIYIIMb1la. Oprattbr oca3aH11H H XHM11qecKoro qyscTsa uo Kpa10 MaHTHH. OpratthI paBHOBecna -

JloJLoBaJl cucme.Ma.

occppa,n;nu,

cTaTOI1HCThI, B Hore.

fepMacppOlJ;l1TbI. IloJiosaa meJie3a HerrapHaa. OrrJio,n;o-

TBopeHne rrepeKpeCTHOe, BHyTpeHHee. 0TKJia,[(hIBaIOT HHI1a B CJil1311CTbIH KOKOH.

Pa3Bumue

rrpaMoe.

BoJihIIIl1HCTBO BO,ZJ;HbIX BH,D;OB pa3s11sa10TcH c MeTaMopcpo30M HOqHhre cTa,n;11u: Tpoxocpopa 11 seJinrep (rrapyca11K) (pnc.

,n;se JIHq11-

119).

Knacc ABYCTBOPLt3Tble, i.mM nnaCTMHLtaTO>Ka6epHble

KJLacc l(BycmBopttambte, uJLu I!JLacmuHttamoJ1Ca6epHbte -

so,n;Hhre, MaJio-

rro,n;B11jKHhre ,ll;OHHhie MOJIJIIOCKl1 c ,D;BYCTBopqaTOH paKOBl1HOH, IIOJIHOCTblO np11KPhIBa10IIIeH nx TeJio. H3secTHO OKOJIO

20

ThIC. s11,n;os. Ilpe,n;cTaB11Te.'111: 6e3-

3y6Ka, nepJIOBl1IJ;bI, Ml1,ll;1111, MOpCKOH rpe6eIIIOK, ycTpllIJ;bl, jKeMqyjKHl1IJ;hl.
B11ocp11JibTpaToph1 (rr11Ta10TCH B3BeIIIeHHhIMl1 B so,n;e qacT111laM11 opraH11qe-cK11x BeIIIeCTB, IIJiaHKTOHOM).
Hl1eM pe,n;yu;11poBaJiaCh roJIOBa.
(ycTp11D;bl).

y

y

B

CBH3Il c naCCl1BHhIM ,ll;Bl1jKeH11eM 11 IIHTa-

HeIIO,ll;BlljKHbIX cpopM pe,n;yu;npoBaHa Hora

HeKOTOPhIX, rrpnKperrJIHIOIII11XCH K cy6cTpaTy cpopM B Hore

11MeeTCH 611ccycHaa jKeJie3a, Bhr,n;eJIHIOIIIaH 6nccyCHhre HHTl1 ,n;JI.a np11KperrneHl1H K

TBep,n;oH: nosepxHOCTl1 (M11,D;ll11).

Be33y61w
eMoB,

0611TaeT Ha ,n;He rrpecHbIX so,n;o-

3apbIBIIIl1Cb B

rpyHT

rrepe,n;Hl1M

KOHIJ;OM TeJia.

Cmpoettue.
,D;JIHHOH

10

PaKOBMHa ,n;sycTsopqaTa.H,

CM, CKpeITJIHeTCH Ha CIIl1HHOH

CTOPOHe aJiaCTllqHoiI CB.H3KOH ( crroco6cTBYeT pacKphIBaHHIO paKOBl1HbI). CTsopK11
IIJIOTHO CjKaTbl napoiI 3aMbIKaTeJibHbIX
MycKyJioB. TeJio cocTOl1T 113 Hor11 11 TyJioB11IIIa. MycKyJI11CTa.H Hora nMeeT KJillHOs11,n;HyIO cpopMy 11 CJIYjKl1T ,D;JI.H rrepe,n;s11-

PMc. 119. Jlw·H1HKa paKOBL'1HHb1x Mon1110c-

jKeH11.H, aa.HKop11saH11.H B rpyHTe. MaH-

KOB -

Tl1ttHa.H IIOJIOCTb C006IIIaeTC.H c BHeIIIHett

napyCHL'1K (sen1-1rep}

287
cpe,n;o:ii c IIOMOID;bIO CHcpOHOB -

OTBepCTHM, o6pa30BaHHbIX uenOJIHbl'.\1 CMh!Ka-

aneM paKOBHHhr w MaHTHH B 3a,n;Heii: qacTH MOJIJIIOCKa.

B

HHiKHHH (sso,n;Hhr:ii)

cncpoH IIOCTynaeT so,n;a c paCTBOpeHHbIM KHCJIOpO,n;OM, OMhIBaeT ma6pbr, nrnu,y,
MY;KCKHe IIOJIOBbie KJieTKH. qepe3 sepxHHM (BbIBO,D;HOH) CHcpOH BMeCTe c BO,ll;OM
BbIP:eJI.liIOTCH TaKme npo,n;yKTbl BbI,D;eJieHHH H HenepesapeHHhie OCTaTKH IIHID;H.
ToK BO,D;hI co3,n;aeTca KOJre6aHHHMH Mepu;aTeJihHhIX pecHwqeK, n0Kphrna101u,ux
MaHTHIO H3HyTpH.

[Juw,eeapumeJtbHa.fl cucmeMa. Cnoco6 nwTaHHH -

6wocpHJibTpau;Ha

(B CBH3H c naCCHBHbIM o6pa30M JRH3H11). qaCTHll,bl ITHID;H c TOKOM BO,D;bl HanpaBJIHIOTCH K pOTOBOMY OTBepCTHIO, OKpymeHHOMY ,D;BYMH napaMH JIOIIacTeii:. IIHm;a IIOCTynaeT B KOpOTKHM nwm;eso,n;, ,n;aJiee B OKpyrJihlM iKeJiy,D;OK,
Ky,n;a sna,n;aIOT npOTOKH ,D;BYJIOIIaCTHOM neqeHH c nHm;esapHTeJihHblMH cpepMeHTaMH. OT meJiy,n;Ka OTXO,D;HT cpe,n;HHH KHIIIKa, nepexo,n;am;aa B 3a,D;HIOIO
KHillKY' 3aKaHqHsarom;yroca aHaJihHbIM OTBepCTHeM B MaHTHHHOM IlOJIOCTH
y BblBO,D;HOro CHcpOHa. fJIOTKa, pa,n;yJia, CJIIOHHbie meJie3hl B CBH3H c pe,n;yKll,HeM roJIOBbl OTCYTCTBYIOT.

KpoeeHOCHaR cucmeMa He3aMKHyTaa. Cep,n;u;e Ha CIIHHHoii: CTopoHe oxsaTbIBaeT 3a,D;HIOIO KHIIIKY H COCTOHT H3 O,D;HOro meJiy,n;oqKa, ,n;syx npe,n;cep,n;wii:.
ApTepHaJihHaa Kposb H3 cep,n;u;a ll,HPKYJIHpyeT no cocy,n;aM w JiaKyHaM TeJia,
oT,n;aeT KHCJiopo,n;, Hachrm;aeTca yrJieKHCJihIM raaoM. BeH03Haa Kposb co6wpaeTCH s Kpy1rny10 npo,n;oJihHYIO JiaKyHy no,n; cep,n;u;eM H B03Bpam;aeTca B ma6phr

,n;Jia ra3006MeHa. 0KHCJieHHaa apTepwaJihHaa Kposb H3 ma6ep no Bhrnocam;wM
cocy,n;aM B03Bpam;aeTCH B cep,n;r:r;e. qaCTH'IHO KPOBb, MHHYH ma6phl, npoxo,n;HT
qepe3 IlO'IKH, OCB06om,n;aacb OT npo,n;yKTOB o6MeHa, H BJIHBaeTCjJ B BbIHOCJJID;He
ma6epHhle cocy,n;hI, sna,n;arom;He s npe,n;cep,n;He.

BbtfJeJtumeJtbltaR cucmeMa -

napHhie nOTIKH (Hecppw,n;HH). ,n:onoJIHHTeJihHO

Bbl,D;eJIHTeJibHYIO cpyHKll,HIO BbIIIOJIHHIOT BblpOCTbl CTeHKH nepHKap,n;a, BH,IJ;OH3MeHeHHhre s BH,n;e rrapHhlX meJie3.

!{btXaHue ocym;eCTBJIHeTCH ma6paMH, rrpOHH3aHHbIMH KposeHOCHbIMH cocy,n;aMH. :IB:a6pbl cpyHKll,HOHHpyIOT TaK me, KaK opraHhl o6MeHa BO,D;bl B MaHTHMHOM IlOJIOCTH.

H epenaR cue meMa

pa3BHTa cJia6o, s cBH3H c MaJiono,n;BHJRHhIM o6pa30M

3tH3HH. CocTOHT H3 Tpex nap HepBHhIX Y3JIOB, pacnoJiararom;Hxca Ha,n; nHm;eso,n;oM (r:r;epe6porrJieBpaJibHbIH), B Hore (ne,n;aJibHbIH) H OKOJIO 3a,n;Heii: KHIIIKH
(BHCr:r;epo-napHeHTaJibHbIM).

Opzattbl 'lyecme paaBHThI cJia6o. B Hore HMeIOTCH cTaTOll,HCThI PaBHOBeCHH.

y

OCHOBaHHH ma6ep occppa,n;HH -

opraHhI

opraHhl XHMHTieCKOM qyBCT-

BHTeJibHOCTH. Per:r;enTopHhre KJieTKH pacnoJiaraIOTCH Ha ma6pax, poTOBhIX
JiorracTax, Kpaax MaHTHH, CHcpOHax. fOJIOBHhre m;ynaJibll,a H rJia3a OTCYTCTBYIOT.

lloJtoea.R cucmeMa. Pa3,n;eJibHOITOJihre. IloJIOBhle meJie3hI napHbre, pacnoJiararoTca B rrepe,n;Heii: qacTH TeJia H B OCHOBaHHH HOrH. IIpoTOKH OTKpbIBaIOTCH
B MaHTHMHYIO noJIOCTh. OnJio,n;oTBopeHHe npowcxo,n;HT s MaHTHMHOM noJioCTH,
Ky,n;a cnepMaT030H,D;hl npOHHKaIOT qepe3 BbIBO,D;HOM CHcpOH.

288

Pa3eumue

3ooJWZUR

c MeTaMopcpo3oM • .Hiiu;a pa3BirnaIOTC.H B MaHTMiiHoii noJIOCTM Ha

ma6pax. JiwqMHKM (rJIOXM,ll;MM), 3aKJIJOqeHHhie B ,ll;BYCTBOpqaTyIO paKOBMHy,
BhIXO,ll;.HT qepe3 BhlBO,ll;HOii CMcpOH B BO,ll;y M npMKpenJI.HIOTCH JIMilKOM HMThlO
K nJiaBHMKaM, KOa<e, ma6paM phI6, r.n;e rrapa3MTMpyIOT OKOJIO ,ll;BYX Mecau;eB,
3aTeM qepe3 pa3phIBhl KOa<M ph16 Bhrrra.n;aIOT Ha ,l(HO BO.n;oeMa M Be.n;yT CaMOCTO.HTeJihHhlll o6pa3 a<M3HM. BpeMeHHhlii rrapa3MTM3M JIMqMHKM crroco6cTByeT
pacce"'leHMIO nw.n;a.

Knacc ronOBOHOn>te MOnmOCKlll

KJtacc I'0Jtoeo11ozue .MOJlJllOCICU

-

aTo caMhie BhICOKoopraHM3oBaHHhie MOJI-

JIIOCKH ( «rrpwMaThl MOPH» ). 8To KpyrrHhie XMID;Hhie MopcKtte mMBOTHhie, aKTMBHo IlJiaBaIOID;Me B TOJIID;e BO,ll;hl. H3BeCTHO OKOJIO

650 BH,ll;OB.
120).

Ilpe.n;cTaBMTe-

JIH: KaJihMap, OChMMHOr, KapaKaTwu;a, HayTHJIYC (pwc.

Cmpoe11ue.

TeJio cocTOHT H3 TYJIOBHID;a H roJIOBhI. <t>opMa TeJia: Topne,n;OBH,ZJ;-

HaH (KaJihMaph1), MeIIIKOBH.n;Ha.H (ochMHHorw), yrrJioID;eHHa.H (Kapai<:aTHD;hI).
Hora rrpeo6pa3oBaHa B ID;yrra.n:hu;a, pacrroJiomeHHhre Ha roJioBe BOKpyr pTa
H ,ll;BHraTeJihHOM BOpOHKH Ha 6pIOIIIHOM CTOpoHe TeJia. 0TCIO,n;a IIOIIIJIO Ha3BaHHe «rOJIOBOHOrHe». Inyrra.n:hu;a CJiya<aT ,ll;JI.H 3axBaTa IIHID;H, rrepe.n;BMa<eHMH
no ,ll;HY H ,ll;JIH rrepeHOCa IIOJIOBhIX rrpo.n;yKTOB B MaHTMMHYIO IIOJIOCTh caMKli

(y

ochMMHoroB).

Y

BhICIIIMX roJIOBOHOrHx ID;yrra.n:hu;a ,ll;JIMHHhle, c MOID;HOii MY-

cKyJiaTypoii, c KPYIIHhIMH rrpwcocKaMH Ha BHyTpeHHeii rroBepxHOCTM. ,I:(BwmeHHe peaKTHBHOe, .n;o

50 KM/q.

HMeIOTCH 3aID;HTHhie rrpwcrroco6JieHHH:
(nHrMeHTHhie KJieTKH KOa<M -

14

rroKpOBHTeJihCTBeHHaH 0Kpac1<:a

xpoMaTocpophl, rrpH H3MeHeHHM cpopMhl KOTO-

11

JO
I

!

PMC. 120. BHyrpeHHee CTpOeH11e rDJlOBOHOroro MOJllllOCKa: 1- uwnanbl.¥l (Hor11) c np11cocKaM11;
2- TepKa; 3 - Jl0BYl1e HOr11; 4 - BOPOHKa; 5 - aHyc; 6 - )l(a6pb1; 7 - opraH BblAelleH11R; 8 - cepAue;
9- YepHl1JlbHblH MeWOK; 10- MaHT11R; 11 - noJlOBOH opraH; 12- opraH n11~esapeH11si; 13- paKOB11Ha (PYA11MeHT); 14 - maa; 15- M03r

run 'f,Jienucmonozue

289

pblX npOHCXO)J;HT CMeHa OKpaCKH), qepHHJihHhlH MeIIIOK (npoTOK OTKPhIBaeTCH
B 3a)J;HIOIO KHIIIKy), m;ynaJihlla H «KJIIOB» (BH)J;OH3MeHeHHhie qeJIIOCTH). llMeIOT
Hapy;irnyro JIH6o BHyTpeHHIOIO (py)J;HMeHTapHyIO) paKOBHHy.

y

HeKOTOphIX pa-

JWBHHa OTCYTCTByeT.

pbtxameJlbJ-ta.R cucmeMa

npe.n:cTaBJieHa ma6paMH (.n:BYM» -

qeThiphM»)

sHYTPH MaHTHHHOH noJIOCTH.

IIuw,eeapumeJlbHa.R cucmeMa. B

rJIOTKe HMeeTCH pa.n:yJia H poroBhie qeJIIO-

cTH ( «KJIIOB» ), Bna.n:aIOT npOTOKH CJIIOHHhIX H H)J;OBHThIX meJie3. JK11.n:KaH IlH-

~a nocTynaeT qepea y3KHH n11m;eBo.n:, npoxo.n:»m;11M: qepea Moar, B meJiy.n:oK
c npOTOKaMH napHOH neqeHH. JKeJiy)J;OK HMeeT CJienoM: MeIIIKOBH)J;HhlH OTpocTOK )J;JIH yBeJI11qeHHH o6'heMa. ToHKaH cpe)J;HHH KHIIIKa nepeXO)J;HT B 3a)J;HIOIO,
OTKPhIBaJOm;yrocH aHaJibHhIM OTBepCTHeM B MaHTHHHYIO IlOJIOCTh.

Hepena.R cucmeMa

npe.n:cTaBJieHa CKOIIJieHHeM raHrJIHeB OKOJiorJioToqHoro

KOJihD;a, 06paayrom;11x Moar, aaKJiroqeHHhIH B xp»m;eByro KancyJiy.

OpzaHbt 'lyecme

xopolllo pa3BHThI. llMeIOTCH cJiomHo ycTpoeHHhie rJiaaa,

opraHhl CBeqeHHH, opraHbl paBHOBeCHH (B KancyJie M03ra), opraHhl o60HHHHH
(no.n: rJia3aMH), opraHhl BKyca (Ha m;yrraJihD;ax), OCH3aTeJihHhie H CBeToqyBcTBHTeJihHhle KJieTKH (Ha Kome).

Kpoeenocna.R cucmeMa rroqTH aaMKHyTaH (JiaKyH MaJio, KopoTKHe).
BbtOeJlumeJlbHa.R cucmeMa npe,n:cTaBJieHa .n:ByMH - qeThiphMH rroqKaMH.
IloJWBll.R cucmeMa. Paa.n:eJihHOilOJihle. roHa)J;hl H IIOJIOBhle rrpOTOKH HenapHhle. OnJio.n:oTBopeHHe uapymHO-BHyTpeHuee, rrpo11cxo.n:11T B MaHTHHHOH rroJioCTH caMKH. 0TKJia)J;hIBaIOT »ifu;a Ha .n:He. Ha6Jiro.n:aeTCH aa6oTa o rroTOMCTBe.

Pa3eumue

rrp»Moe.

3HaqeHHe

MOJlJl/-OCKOB.

MOJIJIIOCKH HBJIHIOTCH COCTaBHOH qacTbIO 6eHTOCa

(o6pa3yIOT KpyIIHhie rroceJieHHH -

HeCKOJihKO ThICffq Ha O)J;HH M2) H 3BeHOM

B u;enHX IlHTaHHH; crroco6cTBYIOT 6HOJior11qecKOH oqHCTKe BO)J;OeMOB H .n:eCTPYKD;HH opraH11qecK11x Bem;ecTB. CJiy.acaT o6'beKTOM rrpoMhICJia (ynoTpe6JieHHe B IIHlll;y, IIOJiyqeHHe rrepJiaMyTpa, meMqyra H .n:p.); HBJIHIOTCH Bpe)J;HTeJIHMH ceJihCKOro X03HHCTBa (cyxonyTHhie BH/J;hl) H IIO)J;BO)J;HhIX .n:peBeCHhlX coopymeHHH (noBpem.n:eHHe )J;HHlll; cy.n:oB H CBaH npHqaJIOB Kopa6eJibHhIM qepBeM).
Me.n:11u;11HcKoe 3HaqeH11e: rrpoMemyToqHhie xoaHeBa qepBeif, rrapa3HTHpyrom;11x y qeJIOBeKa, H H)J;OBHTble .atHBOTHhle (KOHyc, Tepe6pa).

Tun 'IJieHHCTOHorue
0CH08Hbte apoMop</J03bt:

IIOHBJieHHe qJieHHCThIX KOHeqHOCTeH, xopOIIIO pa3-

B.HTaH rronepeqHo-rroJiocaTaH MYCKyJiaTypa, 06ecrreq11Barom;a» 6oJiee 6hICTphle

Ii CJIO.atHhle )J;BH.ateHHH, XHTHHOBhIH IIOKpOB, BhIIIOJIHHIOlll;HH cpyHKil;HIO HaPY.atHoro cKeJieTa; rroHBJieHHe OT)J;eJioB TeJia (reTepoHOMHaH cerMeHTau;H»),
rrporpeccHBHOe pa3BHTHe HepBHOH CHCTeMhl, IIOHBJieHHe cep.n:u;a.
T11rr qJieHHCTOHor11e -

aTo caMa» rrpou;BeTarom;a» rpynna mHBOTHhIX Ha

3eMJie, OTJIHqarom;aHCH 60JihIIIHM BH)J;OBhIM pa3H006pa3HeM, aKOJIOrHqeCKHM
MH:oroo6pa3HeM H BhICOKOH q11cJieHHOCTbIO. ll3BeCTHO 6oJiee

1 MJIH

BH)J;OB, cpe-

.D;H lWTOphIX BCTpeqaIOTCH KaK CB060)J;HO.atHBYlll;He, TaK H napa3HTHqecKHe

290

3ooJWZUR.

CpaeunTeJibHUI xapaKTepncTHKa KJiaccoe MOJIJIJOCKOB

BpIOxouon1e

Ll:BYCTBOp'laThie

f0.10BOHOfHC

IIpep;cTaBMTeJUi:

Ilpy,D;OBMK, BHHOrpap;HaSI yJIHTKa, paIIaHa, CJIH3eHb

Beaay6Ka, ycTpmi;a,
MH,!l;HH

KaJihMap, ocbMHHor,
KapaKaTHUa,HayTHJIYC

Cpep;a o6HTaHHSI

Mope, npecHhie sop;br, cyrna

Mope, upecHbte sop;hr

Mope

06pa3 JK.H3Hlf

IlOJI3alOlllHC MopCKHe Xlfll.\Hl1KH, !IO.II3al0ll.\HC rrpeCHOBO,D;Hbie H cyxorryTHbre
paCTHTCJlbHOSl,!l;Hble

Po10ll1lfe 11JI11 Herra,!l;BHJK.Hhle qmJibTpaTOpbI

IlJiaBalOllll1C
Xlfll.\HHKl1

<l>opMa paKoBHHhl

CrrnpaJibHaSI KoHyCOBH,D;Ha.!l (y HCKOTOpbIX pep;yu11posaua)

Ll:BYCTBOp'laTaSI

Pep;yunposaua (ncKJIIO'ICHHC - uayTHJIYC)

<l>opMa TeJia

I'OJIOBa, TyJIOBHll.\C,
Hora

TyJioBHll.\e, Hora

I'oJIOBa, TYJIOBHll.\e,
Hora BH,!l;OH3MeHeHa
(lllYllaJibUa, BOpOHKa)

IlOKpOBbl

3rrHTeJIHH (Mepu.aTeJihHhlH Ha ma6pax), 3IlHTeJIHH
M3HTHH

3rr11TeJIHH (Mepu.aTeJibHbIH Ha »<a6pax
H CTeHKax MaHTHHHOH IIO./IOCTH), 3IIHTeJIHH M3HTHH

3nlfTCJIHH (Mepu,aTeJibllhlH ua ma6pax), 3IIHTeJIHH
MaHTHH

IloJIOCTb Te.Ila

I..J;eJIOM BOKpyr cepttu.a H IIOJIOBblX meJie3

I..l;eJIOM BOKpyr cepp;u.a 11 noJIOBhIX me.Ilea

I..J;eJIOM BOKpyr cepp;u.a H IIOJIOBblX meJie3

IlHllleBapliTeJibHaSI
CHCTeMa

Ilepet(HHH (rJIOTKa c
TepKOH H CJIIOHHbIMH
meJie3aMH, IlHl.lleBO,!J;,
306), cpep;HHH (meJiy,!l;OK,neqeHb,TOHKaSI
KHillKa) H 33,!l;HHH OTp;eJibl

Ilepet1n11n (rrm.r~eBO,D;), cpe,D;HHH (meJiyp;oK,ne'leHb,TOHKaSI KHillKa) H 33,!l;HHH OT.ll.eJibI

Ilepep;HHH (rJIOTKa
C TepKOH H CJllOHHbIMlf
meJie3aMH, IlHl.lleBO,D;),
cpep;HHH (meJiyp;oK c
neqeHblO H noii;meJiy.ll.O'IHOH meJieaoii, TOHKaSI KlfillKa) H 33,!l;HHH
OTJ:(e./Ibl

Bhr,D;eJIHTeJibHaSI CHCTeMa

Ilo'IKH (op;ua-p;se)

Ilo'IKH (p;se)

IloqKH (,nse-qeTbrpe)

KpoBeuocua.11
CHCTeMa

He3aMKHyTa.R, c
.llBYX -TpexKaMepHbIM cep,nueM

HeaaMKHyTa.11, c
TpexKaMepHbIM cep,11U,eM

HeaaMKHYTaSI, c TpexKaMepHbIM cep,11u,eM
(y HayTHJiyca - Il.!ITHKaMepHblM) H ,ll;OIIOJIHHTeJibHblMH ma6epHbIMH cep,llUaMH

Oco6euuocTH
CTpOCHl1.R
H 06pa3a
JK!i3HH

-

Tun 1£aenucmonoiue

291

Bproxottor11e

Oco6eHHOCTH
cTpOeHHll
11 o6paaa
;Kl13lfl1

,J:(ByCTBOp'laTbie

I'OJ10BOHOI'l1e

,UbrxaTeJihHaJI

:>Ka6pbI ( Ot(Ha-,n;Be)

IIJiaCTHH'laThre ma6-

:>Ka6pbI (o,n;Ha-,n;Be

cn:cTeMa

HJIH JierKoe

pbl (o,n;Ha rrapa)

rrapbr)

HepsHaJI cue-

CTBOJioBa.H,paa6po-

CTBOJIOBa.JI,paa6pocaH-

CTBOJIOBaJI, KOHIJ;en-

TeMa

caHHO-Y3JIOBaJI

HO-Y3JIOBaH (TpH-1'.J.eTbI-

TpHpoBaHHO-y3JIOBaJI

(rrJITb nap raHrJIHeB)

pe rrapbI

fJia3KH, CTaTOIJ;HCTbI,

CTaTOIJ;HCTbI, pe,n;Ko

fJiaaa, CTaTOIJ;HCT,

rJia3KH Ha MaHTHH

opraHbl XHMH'IeCKOro

OpraHbI

•opraHbI XHMH'l:ecKoro

-qyBCTB

rattrJu:res)

-qyBCTBa H OCJI3aHHJl
IloJIOBaJI CHC-

fepMaQ:lpO,!J;HTbI HJIH

TeMa

pa3,n;eJibHOIIOJibJe

Pa3BHTHe

<l>opMhl.

B

'l:YBCTBa H OCJI3aHHJI
Pa3.n;eJibHOIIOJihie

Pa3,n;eJihHonoJibre

IlpJIMOe

C MeTaMOp¢I030M

c MeTaMopcp030M (JIM-

(JIH'l:HHKa-rrapyc-

'l:HHKa-napycHHK HJIH

HHK) HJIH rrpJIMOe

I'JIOXHt(HR)

rrpon;ecce SBOJIIOD;HH qJieHnCTOHorne ,n;ocTnrJin Bhicllleii: opraHH3a-

n;nn H no CTeneHH pa3BHTHH CTOHT Bblllle Bcex 6ecn03BOHoqHbIX JKHBOTHbIX.
XapaKTepHbie npH3HaKn:

1.
2.

,UByCTOpOHHHH ( 6HJiaTepaJihHaH) CHMMeTpllH.

3.

I'eTepOHOMHaH cerMeHTall,HH TeJia (cerMeHTbl pa3Hb!X yqaCTKOB TeJia OT-

MHoroKJieToqHble, TpexcJIOHHbie, nepBnqHopoTble mnBoTHbie.

JIH'IaIOTCH no BeJinq1rne n BHyTpeHHeMy CTpoeHHIO). Cxo,n;Hhie no cTpoeHHIO cerMeHTbl o6'be,!l;HHHIOTCH B OT,ZJ;eJibI: rOJIOBy, rpy,n;b H 6p10111Ko.

4.

qJieHl:ICTbie KOHeqHOCTH, <l>YHKU,l:Il:I KOTOpbIX MHOroo6pa3HbI: ,!l;BHraTeJibHaH, .n;o6hrnaHl:Ie nlllll,l:I, qyBCTBl:ITeJibHaa, ,!l;bIXaTeJihHaa, pa3MHOmeHHH H ,n;p.

5.

Ilonepe•rno-noJiocaTaa MycKyJiaTypa, o6pa3yIOlll,aa oT,n;eJihHbie nyqKn
MhlIIIU., o6ecneqnBaeT 6blcTpbie H paaaoo6pa3Hbre ,n;BHJKeHHH oT,n;eJibHhlMl:I qacTaM1:1 TeJia. Komao-MyCKYJihHhIH MellloK yTpa<IeH.

6.

X1:1T1:1H0Baa KYTHKyJia (Hapya<HhIH cKeJieT) aalll,Hlll,aeT OT BbICblXaHHH
1:1 Mexam1'!ecK1:1x noBpem,n;eH1:1:H. PocT Ha-aa aepaCTHJK:HMOCTH KYTHKYJihl rrpephlBHCTblH H corrpoBom,n;aeTCH

Jll:IH~KOH.

Ilo,n; KYTl:IKYJIOH -

rH-

no,n;epMa.

7.

IloJIOCTb TeJia -

MHKCOD;eJib (cMeIIIaHHaa) -

o6paayeTCH B peayJibTaTe

CJIHHHHH rrepBl:IqHOH H BTOpHqHOH IIOJIOCTeH, aaIIOJIHeHa reMOJil:IM<l>OH,
KOTOpaa BblIIOJIHHeT <l>YHKU.HIO KPOBl:I, IIOJIOCTHOH )Kl:I,!l;KOCTH l:I D;l:IPKYJillpyeT B KpoBeHocaoif CHCTeMe. OcTaTKH n;eJioMa coxpaHaIOTCH OKOJIO rOHa,ZJ; l:I BOKpyr He<t>pn,n;neB.
peHHHe opraHhl.

B

IIOJIOCTH TeJia pacnonaraIOTCH BHYT-

292

8.

3oOJWlUR

ITm:u;esapttTeJihHaH cttcTeMa cocTOHT tt3 Tpex oT,n;eJioB. B rrepe.n;HeM pOT, OKpymeHHhlll pOTOBhlM arrrrapaTOM, rJIOTKa, IIHIIJeBo.n;, 306, meJiy,ll;OK. B rJIOTKY OTKphIBaIOTCH rrpoTOKtt CJIIOHHhIX meJiea. B cpe.n;HeM cpe.n;mrn KttIIIKa, r.n;e rrpoucxo.n;ttT IIHIIJeBapeHtte, scachIBaHtte IIHIIJtt
tt srra.n;aIOT rrpoToKtt rreqeHtt. B aa.n;HeM oT,n;eJie - 3a.D;HHH KttIIIKa, aaKaHq11saI01IJaHCH aHaJihHhlM OTBepCTHeM.

9.

HepBHaH CHCTeMa -

rrapHhlll roJIOBHOH M03r, OKOJIOrJIOTOqHoe HepBHOe

KOJihU:O H 6pIOIIIHaH HepBHaH u:erroqKa. Ha6JIIO,n;aeTCH CJIHHHHe raHrJIHeB
(u:eq:>aJitt3aU:ttH). XapaKTepHo cJiomHoe rrose.n;eHtte tt optteHTaU:ttH B rrpocTpaHCTBe.

10.

OpraHhI qyscTB xopoII10 paasttThI. OpraHhI apeHtt.H (CJIWKHhle ttJitt rrpo-

11.

Bhl,n;eJIHTeJihHa.H cttcTeMa -

CThie rJia3a), OC.H3aHHH, CJiyxa, paBHOBeCHH, XllMttqecKoro qyBCTBa.
Btt.n;ottaMeHeHHhle MeTaHecpptt.n;tttt (r.i:eJio-

Mo,n;yKThI, o.n;Ha-.n;se rraphl). MaJihIIttrtteBhl cocy.n;hl -

cttcTeMa CJierro

3aMKHYThIX Tpy6oqeK Ha 0,ll;HOM KOHU:e, pacrroJiomeHHhIX Mem.n;y cpe.n;Heii tt aa.n;Hen KHIIIKOH, r.n;e rrpo.n;yKThI o6MeHa rrpespa1IJaIOTCH B KpttcTaJIJihI Moqeson KHCJIOThl HJIH ryaHttHa, KOTOPhie BbIBO,ll;HTC.H qepe3
3a,ll;HIOIO KllIIIKy. jKttpOBOe TeJIO (rroqKa HaKOIIJieHHH) -

crrer.i:ttcplfqe-

CKlfe KJieTKH IIOJIOCTH TeJia, B KOTOphIX HaKallJittBaIOTC.H KPHCTaJIJihl
MOqeKHCJihIX COJieii, HO ll3 opraHH3Ma He BhIBO,ll;.HTCH. 3eJieHhle ll KOKcaJihHble meJie3hl.

12.

KposeHocHa.H CHcTeMa HeaaMKHyTaH. Cep.n;u:e pacnoJiaraeTc.H Ha crrHHHOH CTOpOHe TeJia ll ttMeeT pa3JillqHoe CTpOeHtte.

13.

)l;hlxaTeJlhHa.H CHCTeMa y BO,ll;HhIX cpopM -

ma6pbl (BH,ll;OH3MeHeHHhie KO-

HeqHOCTH, B KOTOpbIX U:HPKYJIHpyeT reMOJIHMcpa), y Ha3eMHhlX cpopM JieroqHhie MeIIIKll (Bll,ll;Oll3MeHeHHble KOHeqHOCTH, rrorpymeHHble
BHYTPh TeJia) ll Tpaxe:n (c:ncTeMa paaBeTBJieHHhIX Tpy6oqeK c XHTHHOBhlMll KOJihU:aMtt, rrpOHHKaIOIIJaH BO see opraHhl TeJia).

14.

8H,ll;OKpttHHaH CHCTeMa -

meJie3bl, KOTOpble peryJI:npyIOT MeTaMopcp03

(HeiipoceKpeTOpHble KJieTKH M03ra).

15.

IloJioBaH cn:cTeMa. BoJihIIIttHCTBO su.n;os paa.n;eJihHOIIOJihle mn:BOTHhle,
xoporno Bh1pameH rroJiosoii .n;:nMopcpH3M. IloJIOBhle meJie3hI napHbie
:n HenapHbie. OrrJio,n;oTBopemre BHyTpeHHee H HapymHoe.

16.

Pa3BHTHe rrpaMoe H c MeTaMopcpoaoM: HerroJIHhIM H IIOJIHhIM.

Tun lfJtenucmonoiue BKJIJOqaeT: noiJmun JKa6poiJbtwaw,ue, KJlacc Pa1eoo6pa3Hbte; noiJmun XeJtuu,epoebte, KJlacc llay1Coo6pa3Hbte; noiJmun TpaxeunoiJbtwaw,ue, ICJlacc H ace1eOMbte.

Knacc PaKoo6pa3Hble
KJtacc Pa1eoo6pa3Hbte OKOJIO

20

rrpe:nMylIJeCTBeHHO so,n;Hhie m:nBOTHhie. lfanecTHO

ThIC. BH,ll;OB. Ilpe.n;cTaBHTeJih -

pe'lHOU palC, OTp.H.n; )l;eCHTHHOr:ne,

BhICIIrne paK:n. 06HTaeT B rrpecHhIX rrpoToqHhIX no.n;ax. Be.n;eT HoqHoii o6paa
'KH3HH.

Cmpoenue. TeJio cocTOHT :na Tpex OT,n;eJioB -

roJIOBhI, rpy.n;H H 6prorn1<a.

JCn,acc Pa1wo6paJu&te

293

foJIOBHOM OT,ZJ;eJI BKJIIO'IaeT: aKpOH (roJIOBHaH JIOnaCTb) c aHTeHHyJiaMH (KopoTKHe ycHKH) H

4

cerMeHTa c napHbl-

l'dlf KOHe'IHOCTHMH; aHTeHHbl (,n;JIHHHbie

ycHKH)

,Zl;JIH

OCS3aHHH

lf o6oHHHHH; pOTOBOM annapaT BepXHHX

qeJIIOCTett

napa

(MaH,n;H6yJibl)

lf ,n;Be napbl HHiKHHX (MaKCHJIJibI).
fpy,n;HOM OT,ZJ;eJI (BOCeMb cerMeHTOB)

4

aeceT: TPH napbI HoroqemocTett ,n;Jis
y,n;epmaHHH H npo,n;BHJKeHtm KO PTY
nm1~lf;

XO,ZJ;HJibHbIX

Hor,

lf3 KOTOpbIX nepBas napa HeCeT

nHTb nap

MO~

Hble KJieIIIHH ,Zl;JIH 3aXBaTa

nH~H,

Ha-

na,n;eHHH H 3a~HTbI OT BparoB. Bee
rpy,D;Hble KOHe'IHOCTH paKa HeCyT y ocHOBaHHH ma6pbl H BblnOJIHHIOT ,n;onoJIHHTeJibHO
(pHC.

il>YHKIJ;HIO

,Zl;bIXaHHH

121).

fOJIOBa H rpy,n;b CJIHBalOTCH B MacCHBHYIO roJIOBorpy,n;h. AKpOH, CJIHBmHCb c nepBbIM rOJIOBHblM cerMeHTOM, 06pa3yeT nepBH'IHYIO rOJIOBY npoTou;eit>aJIOH.

OcTaBrnHecs roJio-

BHble H rpy,n;Hble cerMeHTbl co CnHHbl

PHc. 121. CTpoeH11e pe'-IHOro paKa (co

H 60KOB noKpbITbl XHTHHOBbIM naH-

cnl1HHOiil CTOpOHbl): 1 - aHTeHHyna; 2 -

IJ;HpeM -

rOJIOBOrpy,ZJ;HblM

~HTOM

aHTeHHa; 3- pocTpyM; 4- 3aTb1noYHas:i

(KapanaKCOM). IIepe,n;HHM Kpali ~HTa

6op03ACl; 5 - lKa6epHocepAe'-IHb1e 6opo3-

06pa3yeT 3aOCTpeHHblM BbICTyn -

Abl; 6 - nocneAHs:is:i napa 6p10WHb1x HO-

POCTpyM, no 6oKaM KOTOporo Haxo-

lKeK - nnasaTenbHb1e nnacT11HK11; 7 -

,Zl;.HTC.H

TenbCOH; 8 - nnaBHl1K

CTe6eJib'IaTbie

qiaceTO'IHbie

rJia3a. Ha KapanaKce BH,ZJ;HbI co cnHHuo:H CTopoHbl 6opo3,n;hl: nonepe'IHa.H, HaMe'IaID~a.H rpaHHIJ;Y Mem,n;y roJioBHbIMH H rpy,n;HbIMH cerMeHTaMH (3aTblJIO'IHa.H 6opo3,n;a), H ,n;Be orpaHH'IHBaIO~He o6JiaCTb cep,n;u;a (ma6epHocep,n;e'IHble 6opo3,n;w). BoKOBbie qacTH Kapa-

naKca npHKpwBaIDT .ma6phl.
BplOIIIKO (a6,n;oMeH) COCTOHT H3 rnecTH IIO,ZJ;BHJKHblX cerMeHTOB H aHaJibHOH
JionacTH (TeJibCOHa) c aHaJibHblM OTBepcTHeM. Bee cerMeHTbl 6prornKa HecyT
KOHe'IHOCTH: y caMIJ;OB nepBble ,n;Be napbl BH,Zl;OH3MeHeHbl B KOIIYJI.HTHBHble oprattbl, a ocTaJibHbie a OCTaJibHble -

nJiaBaTeJibHhle; y caMOK -

nepBas napa pe,n;yu;lfpoBaHa,

IIJiaBaTeJibHble H y,n;epJKHB8IOT .Httu;a B nepH0,11; pa3MHOJKemm,

a TaKme BbIJIYIIHBIIIHXC.H pa'IKOB. IIocJie,n;HHH napa 6prornHbIX Hor HMeeT qiopMy c,n;BoeHHblx rnHpOKHX nJiaBaTeJibHblX nJiaCTHHOK (ypono,n;w). BMecTe
c YTOJI~eHHbIM TeJibCOHOM ypono,n;w o6pa3ylOT II.HTHJIOIIaCTHblM

«IIJI8BHHK ••

294

300/WZUJI,

Ilo.n;rtt6a.H M pacnpHMJI.ll.H MYCKyJIMCTOe 6pIOIIIKO, paK IIJibIBeT 3a,n;OM Hanepe.n,,
a

no ,!!;HY nOJI3aeT Ha n.HTM

(pttc.

XO,ll;MJihHhIX

Horax

B

JII060M HanpaBJieHHtt

122).

JI01cpo6.

CHapyarn TeJio noKphITO XMTMHM3ttpoBaHHOM KYTMKyJio:ii, nponn-

TaHHOM CJIO.llMM M3BeCTM. XMTMHOBbIH naHIJJ:lpb BhIIIOJIH.lleT 3arn;MTHhie <PYHKII;MM M Hapyarnoro CKeJieTa (MeCTO npttKpenJieHM.ll nyqKOB nonepeqHO-IlOJIOCaThIX MhIIIIn;). PocT conpoBom,n;aeTC.ll JIMHhKO:ii. B cocTaB XMTMHOBoii KYTMKYJihI
BXO,ll;HT nMrMeHThI,

nptt,n;arorn;tte noKpOBMTeJihCTBeHHYIO OKpacKy.

Oco6oM:

CTOMKOCThlO o6Jia,n;aIOT KpaCHhie IIMrMeHThI.

IloJWCmb meJta - CMelIJaHHaH
IIuw,eBapumeJtbl-WR cucme~w.

(MMKCOII;eJib), 3aIIOJIHeHHaH reMOJIMMcpo:ii.
PaKM Bcea,n;Hw (pa3Jiararorn;tteca JKMBOTHhie

M pacTMTeJihHhle OCTaTKM). KttIIIeqHhIH KaHaJI MMeeT BM,ll; npaMOH Tpy6KM M COCTOMT M3 nepe,n;Heli, cpe,n;Heli M 3a,n;Heii: KMIIIOK. l13MeJibqeHHaa rphI3YIIl;MM poTOBhIM annapaTOM nttrn;a qepe3 rJIOTKY M nttrn;eBO,ll; nona,n;aeT B J!<eJiy,n;oK, COCTOHIIl;MM M3 ,n;Byx OT,n;eJIOB: JKeBaTeJihHOro (nttrn;a nepeTttpaeTCH c nOMOIIl;hlO XMTMHOBhIX

3y6n;oB

Ha

BHyTpeHHMX

CTeHKax) M u;e,n;MJihHOro (nttrn;a (jJMJibTpyeTCH,

npon;eJKMBaeTCH). 113 me-

Jiy,n;Ka nwrn;a nona,n;aeT B KopoTKyro
cpe,n;HIOIO KMIIIKY c npoToKaMM ,n;ByJionacTHOli neqeHM ( cMeIIIaHHaa meJie3a, BhIIIOJIHHIOrn;a.H TaKJKe cpyHKII;MIO
no,n;meJiy,n;oqHoli meJie3w), r,n;e npottcXO,ll,MT nepeBapttBaHMe M BCaCbIBaHMe
nMTaTeJihHhIX Bern;eCTB. ,z:i:aJiee M,ll,eT
,ll,JIMHHaH 3a,ll;HHH KMIIIKa, 3aKaHqlfBa rorn;aaca aHaJihHhIM OTBepCTMeM.

,abtxameJtbl-WR cucme.Ma.

:IB:a6phI

npe,n;cTaBJIHIOT co6o:ii TOHKOCTeHHble
BhlpOCThI KOJKHhIX noKpOBOB Tpex nap
HoroqeJIIOCTeM M nHTM nap XO,ll,MJihHhIX KOHeqHOCTeli no 6oKaM rOJIOBOrpy,n;M. B ma6epttoM: noJIOCTM HenpephIBHO npOMCXO,ll,MT TOK BO,ll,hl 3a cqeT
,ll,BMJKeHMH OTpOCTKa BTOpo:ii napbI HMJKHeii: qeJIIOCTM. Bo,n;a Bxo,n;ttT B ma-

PMc. 122. BcKpb1Tb111i peYHOlli paK (caMKa):
1 - maa; 2 -

lKenyAOK; 3 - neYeHb; 4 -

6epHhle nOJIOCTM qepe3 rn;eJib Mem,n;y
rOJIOBOrpy,n;HhIM

Ill;MTOM

M

TeJIOM,

sepxHsisi aprnp1-1si 6pt0wKa; 5 - cepA1.1e; 6 -

a BhIXO,ll,MT qepe3 OTBepCTMe, pacnOJIO-

nepeAH1>1e aprnp1-11-1; 7 - lKa6pb1; 8 - si1>1YH1>1K;

JKeHHOe OKOJIO nepe,n;ttero Kottn;a no-

9- 6pt0wHasi HepsHasi 1.1enoYKa; 10- Mb1w-

nepeqHoli 6op03,ll,hl rOJIOBOrpy,n;M. I'a-

L1b1 6pt0wKa; 11 - aHTeHHYJlbl; 12 - aHTeH-

3006MeH

Hbl; 13 - 3MHS1Sl Ka.1WKa; 14 - MblWl.lbl MaH-

M B KJieTKax TeJia ,n;ttcpcpy3HO B reMO-

A1-16y11

JIMMcpy.

npOMCXO,ll,MT

B

ma6pax

K;iacc Pa1wo6pa3nwe

295

KpoBnwcuaR cucmeMa

He3aMKHyTaJL CocTOHT H3 cep.n;:a;a rrn:TttyrOJihHorr

cpopMhI, pacnoJIO./KeHHOro Ha CTIHHHOH CTOpOHe roJIOBOrpy,n;H, lI OTXO,n;.m:a;ux OT
aero cocy,n;oB. Cep.n;:a;e oKpymeHo oKonocep,n;eqHoii cyMKoii H HMeeT TPH naphI
OTBepCTHH, qepe3 KOTOpbre KPOBh nona.n;aeT B cep.n;:a;e. OT cep.n;:a;a OTXO,ll;.HT TIHTh
KPYTIHhIX cocy.n;oB: nepe.n;mur aopTa (HJIH HenapHaa rna3Haa apTep1rn); ,n;Be 6oKOBhie CHiKKOBhie apTepHH (K aHTeHHaM); 3a,n;HHH aopTa; HHCXO,n;an~aa apTepHH, KOTOpaa ,n;eJIHTCH Ha rpy,n;HyIO apTepHIO H 3a,ll;HIOIO 6pIOIIIHYIO apTep1110.
KpoBh (reMOJIHMcpa) TeqeT no cocy,n;aM H naKyHaM MHKco:a;ena, co.n;epmHT ,n;hrxaTeJihHhle TIHrMeHThl reMOD;HaHHH HJIH reMorno6HH, CBH3hlBaIO~He KHCJIOpo.n;. JfMeIOTCH HeiipoceKpeTOpHhle KJieTKH, BhI,ll;eJIHIO~He ropMOHhI, KOTOphie
nocTynaIOT B reMOJIHMC}:ly H BJIHHIOT Ha o6MeHHhie npo:a;eCChl B opraHH3Me,
JIHHhKY H pa3BHTHe.

HepBHaR cucmeMa

npe,n;cTaBJreHa napHhIMH Ha,n;rnoToqHhlMH raHrJIHHMH

H no,n;rJIOTOqHhIM raHrJIHeM (HHHepBHpyIOT rJia3a H YCHKH); OKOJIOrJIOTOqHhlM KOJihll;OM (HHHepBHpyeT pOTOBhie opraHbI); 6pIOITIHOH HepBHOii :a;enoqKOH (HHHepBttpyeT KOHeqHoCTH H BHyTpeHHHe opraHhI). BpIOrnHaa HepBHaa
:a;enoqKa COCTOHT H3

12

HepBHhIX Y3JIOB, CB.H3aHHhIX Mem.n;y co6oii npo.n;o.'Ih-

HhlMH H KOJihIJ;eBhlMH HepBaMH.

Opzattbt 'lYBCmB

xoporno paaBHThl. OpraHhI o6oHHHH.H -

KH (aHTeHHYJihl). OpraHhl ocaaaHHH 3peHHH -

KopoTKHe ycH-

,n;JIHHHhle ycHKH (aHTeHHhl). OpraHhI

napa CJIOiKHhlX cpaceToqHhlX rJia3 Ha TIO,ll;BHiKHhlX CTe6eJihKax.

<l>aceToqHhie rnaaa COCTOHT H3 npOCThIX rJia3KOB -

OMMaTH,ll;HeB, KOTOphre

npe,n;cTaBJIHIOT co6o:tt KOHYCOBH,ll;HhlH 6oKaJI, orpaHHqeHHhIH TIHrMeHTHhIMH
KJieTKaMH, BKJIIOqaIO~Hii npo3paqHyIO KYTHKYJIY (rrpeJIOMJIHeT CBeTOBOH
nyqoK), xpycTaJihHhlH KOHYC (npOBO,ll;HT CBeTOBOH nyqoK) H peTHHaJihHhie
KJieTKH (BOCIIpHHHMaIOT CBeT). 0MMaTH,ll;HH H30JIHpOBaHhl .n;pyr OT .n;pyra,
noaToMy H3o6pameHHe CKJia,n;hlBaeTca H3 pa3Hhlx qacTeii. TaKoe apeHHe Ha3hlBaeTca M03aHqHhlM. OpraH paBHOBeCHH -

CTaTOD;HCThI B BH,n;e ,ll;BYX HMOK

c qyBCTBHTeJihHhIMH BOJIOCKaMH H rrecqJIHKaMH y OCHOBaHHH aHTeHHyJI, KOTOphle coo6~a10Tca c BHernHeii cpe.n;oii.

BbtiJeJLumeJLbHaR cucmeMa

npe,n;cTaBJieHa napHhlMH aeJieHhIMH (camKo-

BhlMH HJIH aHTeHHaJihHhlMH) meJie3aMH, KOTOphle pacnoJiaraIOTCH B rOJIOBHOM OT,n;eJie. Kam.n;aa meJie3a COCTOHT H3 KOHIJ;eBoro MernoqKa, H3BHTOrO
Bhl,lJ;eJIHTeJihHOro KaHaJih:a;a, KOTOphIH MOiKeT pacrnHpHThCH, o6paaya «MOqeBo:ii: ny3hlph», H Bhl,ll;eJIHTeJihHOro OTBepcTHH y OCHOBaHHH BTOpOH naphl
YCHKOB (aHTeHH).

II OJLOBaR cucmeMa.

Paa,n;eJihHOTIOJihre. IloJIOBhle meneahl HenapHhie, pacno-

JiaraIOTCH B rpy,n;HOH o6JiaCTH. jKeHCKHe TIOJIOBhle OTBepCTHH HaXO,ll;HTCH Ha
mecTOM rpy,n;HOM cerMeHTe y

OCHOBaHHH TpeTheii naphI XO,ll;HJlhHhlX HOr,

a MYiKCKHe -

Ha BOChMOM rpy,n;HOM cerMeHTe y OCHOBaHHH TIHTOH naphl xo-

~HJihHhlX Hor.

y

caM:a;OB paKa o,n;Ha-,n;Be naphl 6pIOITIHhIX Hor npeBpa~eHhl

B KonyJIJITHBHhle Tpy6oqKH. Onno,n;oTBopeHHe HapymHoe. BecHOH caMKa BhlMeThlBaeT HKPY
RO.IKKaM.

(150-200

HKPHHOK), KOTopaJI rrpHKJieHBaeTca K 6p10rnHhlM

296

300/1,0lUR

Pa3Bumue

rrp.HMOe.

lfa

.HHI.J; Bhrxo.n;.HT cqJOpMIIPOBaHHbie pa'IKII, KOTOphre

HeKOTopoe BpeM.H ocTa101'c.H Ha 6prorrme y MaTepII, a 3aTeM rrepexo,n;.HT K caMoCTO.HTeJihHOMY o6pa3y :>KII3HII. PocT H pa3BIITIIe rrpoIIcxo,n;IIT BO BpeM.H rrepHo,n;H'IeCKIIX JIIIHeK, B O,ll;HO JieTO pacTyT O'IeHh 6hICTpO II JIHH.HIOT
BO BTOpOe JieTo -

.n;o

TeJihHOCTh :>KH3HII

15-20 JieT.

y

.n;o

.n;ec.HTH pa3,

II.HTII pa3. IlocJie II.HTII JieT paKH He pacTyT, rrpo,n;OJI:>KM-

MHOrHX paKoo6pa3HhIX 113 .a::iiu;a BhIXO,ll;HT IIJiaHKTOHHa.H JIII'IHHKa -

Ha-

yIIJIMYC II pa3BIITMe rrpOlICXO,lJ;HT c MeTaMopcp030M.
3Ha'IeHHe

paiwo6pa31fbtX:

yqacTIIe B KpyroBopoTe BO,ll;HhlX SKOCIICTeM;

KOPM ,ll;Jl.H pbI6, MOJIJIIOCKOB, BO,ll;HhIX MJieKOIIHTalO~HX (KHTOB); caHMTaphl BO.n;oeMa; o6'beKThl rrpOMbICJia (MCJIOJih3YIOTC.H B IIH~y); Me,ll;Hll;MHCKOe 3Ha'!eHHe -

rrpoMe:>KyTO'IHhJe X03.HeBa reJibMIIHTOB (u;IIKJIOIIhl,

paKlI, Kpa6bl);

y~ep6 X03.HHCTBeHHOH ,n;e.HTeJihHOCTM qeJIOBeKa (rrapa3MTM'IeCKIIe cpOpMhl, Bhl3hIBaIO~He 6oJie3HII II ra6eJih pb16).

Knacc nayK006pa3Hble
KJlacc Ilay1Coo6pa3Hbte OKOJIO

35

rrpeIIMy~eCTBeHHO o6aTaIOT Ha cyrne. lfaBeCTHo

ThIC. BH,ZJ;OB. Ilpe,n;cTaBIITeJih -

nay1C-1Cpecm06UIC,

IIIHpOKO pacrrpo-

CTpaHeH B cpe.n;ue:ii qacTII Harne:ii CTpaHhl. Ha CIIMHHOH CTOPOHe 6plOIIIKa IIMeeTC.H KpecToo6pa3HhJH pIICYHOK, 3a 'ITO rrayK II IIOJIY'IIIJI CBOe Ha3BaHIIe.

lfa rra-

YTHHhl rrayK CTpOIIT JIOB'!He ceTII B <}>opMe KpyroB c pa,n;IIaJihHhIMII ll KOHI.J;eHTPH'!eCKHMH HMT.HMH,

y6emn~e • .H:iiu;eBOH KOKOH,

a

TaKme c

IIOMO~hlO

rrayTMHHO:ii HIITII pacceJI.HIOTC.H MOJIO,ZJ;hre oco6II. XII~HHK, rrnTaeTC.H HaceKoMhIMM. Be,n;eT CKphIThIH o6pa3 :>KH3HII (pHc.

CmpoeHue.

123).

TeJio cocTOHT H3 ,n;Byx OT,n;eJioB: roJioBorpy,n;II II 6prornKa, coe,n;II-

HeHHhlX rro,n;BII:>KHO rrepeT.H:>KKO:ii (cTe6eJihKOM). CerMeHTau;II.H BhlpameHa cJia60. foJIOBOrpy,n;h HeCeT UieCTh nap O,lJ;HOBeTBIICThIX KOHe'!HOCTew: rrepBhie ,n;Be

PMc. 123.

BHyrpeHHee cTpoeHHe nayKa-KpecTOBHKa:

HepBHaS! ClllCTeMa;

1 - S1AOBHTSS1 )l<e11e3a; 2 - mmiesoA; 3 -

4 - J18rD'lHblH M8WOK; 5 - Tpaxe111; 6 - nayrlllHHbl8 )l<e11e3bl; 7 - no1100asi )1<811e3a;

8 - KHWKa; 9 - C8PAL\8; 10 - )1<811YAOK

297
naPhl -

poTOBhie opraHbI. HoroqeJIIOCTH (xeJittuepb1) cocTOJIT H3 ,n;syx '!JieHtt-

JWB, cHa6meHHbIX OCTpbIMH, 3arHyThIMH BHH3 KOrOTKaMH, Ky,n;a OTKPhIBaIOTCJI
opoTO:KH JI.ZJ;OBHThIX meJie3. CJiyJKaT .ZJ;JIJI Hana,n;eHHJI, 3axBaTa H yMePIIIBJieHHJI
p;o6hlqH; HOrOIIIYIIaJihUa (rre,n;HrraJibllbI) -

opraHhl OCJI3aHHJI. qCTbipe rrapbI

:x:o,D;HJlhHhlX Hor IIOKpbIThl qyBCTBHTeJihHbIMH BOJIOCKaMH, qTo KOMIIeHCHpyeT
oTCYTCTBHe y HHX ycHKOB. Ha KOHUaX HOr pacnonaraIOTCJI KOrOTKH, yqaCTByJOlll;He B nepe,n;Btt:lKeHHH H IIJieTeHHH nayTHHbI.
BpIOIIIKO KpynHee roJioBorpy,n;H, JIHUeBH,n;Hoe, Bee qJieHHKH CJIHThI BMecTe,
KOHeqHOCTH pe,n;yunpoBaHhI.

B

3a,n;Hell: qacTn HMeIOTCJI TPH naphl nayTHHHhIX

6opo,zi;aBO:K (BH,ZJ;OH3MeHeHHhie KOHeqHOCTH), Ha BeprrmHe KOTOpbIX OTKPhlBaJOTCJI rrpOTOKH rrayTHHHhIX JKeJie3.

!IoKp06 TOHKHH, XHTHHOBbIH, ,ZJ;JIJI o6JierqeHHJI Beca TeJia. HapyJKHbIH CJIOH
(anHKYTHKyJia) 3aII1HII1aeT Teno OT BhICbIXaHHJI.
Oco6eHHOCTbIO ,n;BnraTeJihHOH cncTeMbI JIBJIJieTCJI OTCYTCTBtte Mb1ruu-pa3rn6aTenell: (pa3rH6aHne ocyII1eCTBJIJieTcJI naccHBHO, 3a cqeT BhICOKoro BHyTpnnoJIOCTHoro ,n;aBJieHHJI).

IIuw,eeapumeJtbHa.H cucme.Ma

xoporuo ,n;m:)lcpepeHu;ttpoBaHa H cocTOHT H3 ne-

pe,ZJ;HeH, cpe,ZJ;HeH, 3a,ZJ;HeH KHIIIOK H aHaJibHOro OTBepCTHJI.
pa3yeT pacrunpeHHe -

II epeiht.H.H

KUUJKa o6-

MyCKYJIHCTYIO rJIOTKy, KOTOpaJI CJIYJKHT B Ka'leCTBe Haco-

ca, BTJirHBa10II1ero rroJiyJKH,n;KyIO IIHIIIY.

B

nepe,n;HIOIO KHIIIKY OTKphIBaeTCJI napa

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opraHhI.
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308

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CpaBHHTeJibHaH xapaKTepncTHKa KJiaccoB 'IJieuncTonornx
Oco6eH11ocT11
CTpOeHMH
11 o6pa3a m1131111

PaKoo6pa3Hh1e

Ilay1rno6pa3Hhle

HaccKOMhle

II pep;cTaB11TeJil1

Pe'IHoii paK, KpeBeTKa, MOKp11ua,
U11KJIOII, p;acpH11.H

C1eoprr11ou, rrayK,
KJieIU

Capa11qa,Ky3He'l11K11,
TapaKaHhI, myK11,
6a60'IK11, MYXH, II'leJ1h1

Cpep;a
0611TaHl1.fl

Mop.fl, rrpecHhie BOAhl, pep;Ko cyrna

Cyrna, pep;Ko
rrpeCHhie BO,ll;hl

Cyrna, rrpecHhre BOAhI

06pa3 ml13Hl1

I1JiaBaIDIUl1e cp11JihTpaTOphl (p;acpH11.H ),
l10JI3aIDIUl1e 11 IIJiaBa101u11e

Bera1011111e Xl1IUH11Kl1, rrapa311Thl 11JI11
paCTl1TeJibHOH,ll;Hhie
(qacTh KJie111eii)

Bera1011111e, Jia3aIOn~11e,
po1011111e, rrJiaBaIDIU11e,
MHOro JieTaIDIU11X; pacTl1TeJibHOH,ll;Hhie, Xl1IUHl1K11 11 rra pa311Thl

CTpoeu11e TeJia CerMeHTHpoBaHHhle CJIHTHaH roJioBorpyp;h, 6pIDIIIKO cerroJIOBa, rpyp;h,
6pIDIIIKO
MeHTHpOBaHHOe HJIH
CJil1THOe

CJIHTHaSI roJIOBa,
cerMeHTHpOBaHHhie
6pIDIIIKO H rpyp;h

Yc11K11 na
rOJIOBHOM
oTp;eJie

ABe rraphl

HeT

Op;ua rrapa

THII
KOHe'IHOCTeii

ABYBCTBliCThie

O;i;ttoseTBliCThle

Op;HOBeTBHCThre

IloKpOBhl H
CHCTeMa MhlIIIU

KyT11KyJia, Koma
(rHrrop;epMa)

KyTttKyJia, Koma (rtt- KyTttKyJia, Roma
(mrrop;epMa)
rrop;epMa)

IloJIOCTh TeJia

CMeIIIaHHaH,

CMernauHaSI

CMe111auHaH, c p;11acpparMaMH

IlHIUeBapHTeJihHaH CHCTeMa

Ilepep;u11ii (IIHIUeBop;, meBaTeJihHbIH
meJiyp;oR), cpep;HHH ( c rre'leHhID) H
3a,ll;HHH OT,lJ;eJihl

IIepep;HHii (rJIOTRa,
rr11111eBop;, meJiyp;oR),
cpep;HHH (c rre'leHO'IHh!Ml1 rrpHp;aTKaMH)
H 3ap;H11ii OTp;eJibl

Ilepep;HHll (rJIOTKa,
IIHIUeBop;, 306, meJiyp;oR), cpep;uHii (c meJie3HCThlMH BblpOCTaMll) 11 3a,ll;HHH OTp;eJihl

Bhrp;eJIHTeJihHaH Cl1CTeMa

MaJihITHr11eBhI cocyp;hI
AuTeHHaJihHhle me- Ilo'IRH (op;ua rrapa),
Jie3hl (op;ua rrapa)
MaJihITHrHeBhl cocyp;hl

KpoBeHocuaSI
CHCTeMa

He3aMRHyTaH c
He3aMRHyTaH c Tpy6Tpy6qaTh!M MHOro- 'laTh!M MHOPOKaMepRaMepHhIM cepp;ueM HhIM cepp;ueM

He3aMRHyTaH c Tpy6qaThIM MHOPOKaMepHh!M cepp;ueM

AhrxaTeJihHaH
CHCTeMa

:IB:a6phl (Ha KOHe'IHOCTHX)

JierRHe (B11,ll;OH3MeHeHHbie KOHe'IHOCTH),TpaxeH

Tpaxe11

HepBHaH
Cl1CTeMa

BpIDIIIHa.fl uepBHaH
uerro'IRa

BpIDIIIHaH uepBHa.fl
uerro'IRa

BpIDIIIHa.fl uepBHaH
uerroqRa

Tun xopiJo6bte

309
0/WH'lUllU ('

PaKoo6pa3HhIC

IlayKoo6pa3Hble

OpraHhI

<PaceTO'IHhre H rrpo-

IlpocThre rJiaaa, opra-

ct>aceTO'IHbie H rrpOCTbie

qyBCTB

CThie r.'Iaaa, cTaTo-

Hhl OC.!I38HHJI, XHMH-

rJiaaa,opraHhICJiyxa,

Il;HCTbl, OCJI38HHJI

'!eCKOI'O 'IYBCTBa

Oco6eHHOCTH
CTpOCHlUI
H o6paaa :1KH3HH

HaceKOMhie

XHMWieCKOl'O 'IYBCTBa,
OCJI3aHHJI H p;p.

IloJIOBaH

Paap;eJihHorroJiaa

Paap;eJihHorroJiaa

C .'IH'IHHKOM H•'IH

IIpa Moe

Paap;eJihHorroJiaa

CHCTeMa
PaaBHTHe

C JIH'IHHKaMH
(HeilOJIHOe H IlOJIHOe

rrpaMoe

rrpeBparu;eHHe)

CBeT HX 3al10JIHeH KpHCTaJIJiaMH MO'leBOll KHCJIOTbl -

11po,n;yKTOM ,Zl;HCCHMHJIH-

:a;HH. B Bhr,n;eJieHHH TaKme 11pHHHMaeT yqacTHe a<HpoBoe TeJio ( «110'!Ka» HaR011JieHHH), B KOTOPOM CKal1JIHBaeTCH H He y,n;aJiaeTCH MO'leBaH KHCJIOTa, OTKJI8,Zl;blBaJOTCH 11HTaTeJibHbie Ber:u;ecTBa, Heo6xo,n;HMbie 11pH ,ZJ;JIHTeJibHOM I'OJIO,ZJ;aHHH.

IloJloBaR cucmeMa.

Paa,n;eJibHOl10Jible, BblpameH 110JI0Boii: ,ZJ;HMop<t>HaM. IIo-

JIOBbie meJieabl 11apHbJe. 011JI0,n;OTBopeHHe BHyTpeHHee. PaaMHOa<eHHe 110JI0Boe.

Pa3Bumue

Maii:cRoro myKa 11p0Hcxo,n;HT c 110JIHbIM 11peBpar:u;eHHeM. Ha HH:a;

B 110'-IBe B Te'leHHe

3-4

JieT pa3BHBaJOTCH JIH'IHHKH, KOTOpble 11HTaJOTCJl KOp-

HJlMH ,n;epeBbeB H 11peBpar:u;aJOTCH B Hel10,ZJ;BH:IKHYJO KYKOJIKY. Ilo,n; o6oJIO'IKOll
KYKOJIKH 11pOHCXO,!J;HT 11epecTpOii:Ka opraHOB H TKaHeii: JIH'IHHKH, KOTopaa aaKaH'IHBaeTCH Bbrxo,n;oM BapocJioro HaceKOMoro. MaccoBbIH BbIJieT myROB 11poHcxo,n;HT K8:1K,ZJ;bJe
3Ha'!eHHe

3-5 JieT,

naceKOMbtX:

B3POCJibJe oco6H 11HTaJOTCH JIHCTbJlMH 6epe3bl.
yqaCTHe B 11HI:a;eBbIX :u;e11ax 11HTaHHH, 011bIJieHHe

:a;BeTKOBhlX pacTeHHM, yqacTHe B 110'1B006pa30BaTeJibHbIX 11po:u;eccax; HCl10Jib30BaHHe B OTpaCJIJlX X03JlllCTBa -

11qeJIOBO,Zl;CTBO, rneJIKOBO,ZJ;CTBO H ,n;p.' HC-

Tpe6JieHHe .Bpe,n;HTeJieH pacTeHHM. flBJIJllOTCJl Bpe,n;HTeJIJlMH ceJibCKOX03JlllCTBeHHbIX pacTeHHM H Jieca, 11apa3HT8MH H 11epeHOC'IHKaMH aa6oJieBaHHll qeJIOBeKa H :IKHBOTHbIX, a TaK:IKe HCilOJib3YJOTC.H B Me,ZJ;Hl.l;HHe.

Tun xopii;oBI.ie
XopiJOBbte -

aTO HaH6oJiee BhlCOKoopraHH30BaHHa.H H CJIO:IKHa.H rpy1111a :IKH-

BOTHhlX, :IKHBYI:a;HX B paaHoo6pa3HbIX ycJIOBH.HX. Pac11pocTpaHeHbI 110 BCeMy
aeMHOMY rnapy. 06r:u;ee 'IHCJIO BH,ZJ;OB OKOJIO

43

TbIC.

XapaRTepHbie 11pnaHaRn:

1.

,n;ByCTOpOHH.HJl (6HJiaTepaJibHa.H) CHMMeTpH.H TeJia.

2.
3.
4.

TpexcJioliaocTb.
BTopnqaopoToCTb.
BTopnqaa.H 110JIOCTb TeJia (:a;eJioM).

310

30011,0ZUR

5. BHyTpeHirn:ii ocesoii c1<eJieT s stt.n;e cnttHHOM CTPYHhI, MJIM xop.n;h1. Y HM3IUMX xop.n;OBbIX xop.n;a coxpaH5IeTC5I B TeqeHMe sceii iKM3HH, y BbICIUHX B :3M6pHOHaJibHOM neptto.n;e, a aaTeM aaMem;aeTC5I xpam;aMH If KOCTHbIM
IT03BOHOqHifKOM.
6. HepBHaa CHCTeMa pacnoJiaraeTC5I Ha CITHHHOH CTopoHe TeJia B Btt.n;e HepBHOH Tpy6KH, ITOJIOCTb KOTOpoii Ha3hIBaeTC5I HeBpou.eJih. y IT03BOHoqHbIX
.n;m:pcpepeHu.HpyeTC5I Ha roJIOBHOH If CITIIHHOll M03r.
7. IIHm;esapIITeJihHaa Tpy6Ka pacnoJiaraeTca no.n: xop.n;oii.
8. EcTh ma6epHhie neperopo.n;KH, JIII6o B Te'-IeHIIe sceii iKH3HII, JIH6o B sM6pHoHaJihHOM COCT05IHHH. Ilepe.n;HIIH OT.n;eJI (rJIOTKa) npOHH3aH ma6epHblMll m;eJIRMH. y BO.n;HhIX cpopM (pb16b1) OHM coxpaH5IIOTC5I B TeqeHHe Bceli
;KH3HH, y Ha3eMHhIX 3aKJia.ll;hlBaIOTC5I B 3M6pHoreHe3e If npeo6paayIOTC5I
B .n;pyrHe opraHbI.
9. KposeHOCHaa CHCTeMa aaMKHyTaa, cep.n:u.e (IIJIH aaMeHarom;Hi1: ero nyJihcIIpyrom;Hli KpOBeHOCHbIH cocy.n:} pacnoJiomeHO Ha 6pIOIUHOH CTOpOHe TeJia no.n: ITHm;esapHTeJihHOH Tpy6JwiL
10. MeTaMepHa, cerMeHTaU.HR TeJia.
11. .D:BYXCJIOHHaa Koma: HapymHaa ( :3ITH.n;epMHC ); BHyTpeHH5I5I (.n;epMa).
Tun XopiJ06ble no.n;paa,n:eJiaeTca Ha TPH no.n;THna: Jlu'lUHO'lHOxopiJoBble
(06o;wqnu11:u), Becqepennbte, Ilo360ffO'iffbte (I/epennbte).
JlM"'IMHO"'IHOXOPAOBble

Jluquno'lnoxopiJoBbte (06oJt01lffUKU) o6HTaIOT B Mopax, oKeaHax. Acu.H.n;HH
Be,n;yT CH,ll;5JqHli o6paa ;KH3HH, CaJibilbI H arrneH,n;HKYJI5IpHH - ITJiaBaIOID;IIH.
B npou.ecce :3BOJIIOU.HH nponaoIUJia pe.n;yKU.HR MHOrHx opraHOB.
TeJio 6oqoHKoo6paaHoe, noKphITO aam;ttTHOH o6oJioqKoi1: mynu11:ou, cocToam;eii H3 u.eJIJIIOJI030IIO,ll;06Horo MaTepHaJia. y acu.H.n;Hn CHJibHO pa3BHTa
rJIOTKa c ma6epHbIMM OTBepCTM5IMM, KOTOpaa 5IBJI5IeTC5I opraHOM .n;wxaHH5I
H IIHTaHH5I. Ilo crroco6y IlMTaHM5I - aKTMBHbie cpHJibTpaTOpbI. lfMeIOTC5I poTOBOll H KJioaKaJihHhlH cHcpOHhI. KposeHOCHaa cttcTeMa HeaaMKHyTaa. IIoJIOBaa c:ucTeMa repMacppo,n;HTHaa. PaaMHomeH:ue IIOJIOBhlM :u 6ecnOJihIM nyTeM. Paas:uT:ue c MeTaMopcpoaoM. Jlttqnmrn HMeeT xop.n;y, HepBHYIO Tpy6Ky,
ma6epHbie m;eJIH. y B3pOCJlblX oco6eii B CB5I3H c o6pa30M iKH3HH pe.n;yu.npoBaJiaCh xop.n;a, TYJIOBHID;Haa Myc:KyJiaTypa, HepBHa5I Tpy6:Ka, Hecppn.n;HH.
ToJibKO nayqeHHe JIHqHHOK noaBOJIHJIO OTHecTH acu.n.n;H:ii K xop,n;OBhIM .mHBOTHhIM.
Ilo.IJ;THil 6ecqepeilHhie. Jlanu,emHUKU - MOpC:KHe .IKHBOTHhie, KOTOpbie o6IITaIOT Ha ,n;He, 3aKailblBa5ICb B neco:K. Ilpe.n;KH Il03BOHOqHbIX, coqeTaIOm;He B ce6e KaK npH3Ha:KH xop.n;OBbIX, Ta:K H 6ecrr03BOHOqHblX iKHBOTHhIX.
Pw6oo6paaHaa cpopMa TeJia, HarroMttHarom;aa xttpyprnqec:Kttii HHCTpyMeHT - JiaHu.eT, ,ll;JIHHOH 6-8 CM. lfMeIOTC5I KOiKHbie CKJla.ll;KH. 0.n;Ha HerrapHaa CKJia.ll;Ka o6paayeT CilHHHOH, 6pIOIUHOH, XBOCTOBOH IIJiaBHH:KH. Ilo 6o:KaM
TeJia pacnoJiararoTca napHhie MeTarniespaJlhHhre CKJia.ll;KH. CpacTaach Mem.n;y
co6o:ii, OHH o6pa3yIOT O:KOJIO.ma6epHyIO (aTpHaJibHYIO) IIOJIOCTb, KOTopaa OT-

J[u~uno~noxopoo6~e

311

Kp&rnaeTCH OTBepcTtteM -

aTpttorropoM. I'OJIOBHOM OT,!J;eJI He o6oco6JieH, rrap-

HbIX KOHeqaocTeii HeT, MeTaMepua TeJia BhipameHa xoporno (puc.

fl 01Cp06

TeJia COCTOllT ll3 ,!J;BYX CJIOeB: HapymHOrO -

N!J1Ca II BHyTpeHHero -

126).

0,!l;HOCJIOMHOrO 3IIH,!l;ep-

.n;epMbl (KYTIIC).

CKeJieT rrpe.n;cTaBJieH xop.n;oif, pacrroJiomeHHOM no rrpo.n;oJI&Hoii OCH TeJia.
MbtUle'lHblU CJlOU COCTOIIT II3 ,!l;BYX rrpo.n;OJibHbIX JieHT, pa3.n;eJieHHhlX Ha cer-

MeHThl -

MIIOMep&1. MII0Mep&1 OTrpaHIIqeHbI .n;pyr OT .n;pyra coe.n;IIHIITeJihHOTKaH-

HhIMII rrpOCJIOMKaMII -

MIIOCeJJTaMII, C03,!l;aIOIIl;lIMII orropy MbIIIIeqHbIM BOJIOKHaM.

Hepe1ta.R cucme.Ma

rrpe.n;cTaBJieHa HepBHOM Tpy6Koii Ha CIIIIHHoii cTopoHe

TeJia. Kpaa Tpy6KII Ha crrIIHHoii cTopotte cpacTaIOTCH II IIMeIOT BII.n; TOHKoii:

5

::;;+;<:_-~;;;,."'"·
----> . .. . "... ·.
~~;~
1

2 ------ ----

4

5

12

13

6

-------

3

a
7

10

6

11
6

14

6

PMc. 126. CTpoeH1-1e 11aHu,eTH1-1Ka: a - BHewH1-1H B1-1A; 6- onopHaH, HepBHaH 1-1 n1-1L11eBapme11bHaH c1-1B - KpOBeHOCHaH Ci-lCTeMa. 1 - POTOBble L11ynanbu,a; 2- 60K0Bble KOlKHble CKJlMKl-1; 3- no110Bb1e lKe11e3b1; 4 - oTBepcrne, np1-1 nOMOLl.11-1 KOToporo 0Ko11olKa6epHa11 nonoCTb coo6Ll.13eTcH c oKpy)KalO-.

CTeMb1;

Ll.leH cpeAOH; 5 - n11aBHl-1K; 6 - aHanbHOe OTBepcrne; 7 - HepBHaH Tpy6Ka; 8 - xopAa; 9 - poTOBaH
nOJlOCTb; 10- rllOTKa, npOHl-13aHHaH lKa6epHblMl-1 Ll.lellHMl-1; 11 - neYeHb; 12- cpeAHflH Kl-lWKa; 133aAHflH Kl-lWKa; 14 -

Cnl-lHHaH aoprn; 15 - KOPHl-1 cn1-1HHOH aopTbl; 16- COHHble apTep1-11-1; 17- COCYAbl

Me)K)Ka6epHblX neperopOAOK; 18- 6plOWHaH aoprn; 19- Kan1-11111Hpbl neYeHl-1; 20- nOAKl-1WeYHaH BeHa; 21 - K"1WeYHble Kamrn11Hpb1

300J£ozu11,

312

~eJIM, ITO Kpa.HM KOTopoii: pacrroJiaraIOTC.H CBeTO'-lyBCTBMTeJibHhie o6pa30Ba-

mrn -

2Jta3/CU I'ecce (cBeTO'-IYBCTBMTeJihHa.H M IIMrMeHTHa.H KJieTKM). Ilepmpe-

pnqecKa.H HepBHa.H CMCTeMa rrpeµ;cTaBJieHa HepBaMM,

OT HepB-

OTXOµ;.H~MMM

HOM Tpy6KM M MHHepBwpyro~MMM MycKyJiaTypy. Ha oµ;nH MhIIIIe'-IHhIM cerMeHT rrpnxoµ;.HTC.H µ;Be rrapbl HepBOB -

CIIMHHhie M 6pIOIIIHhle, He CB.H38HHhie

Memµ;y co6oii:.

OpzaHbl 'lyBcme pa3BMThl CJia6o. fJia3KM recce -

CBeTO'-lyBCTBMTeJibHbie

opraHhI. Jlam.i;eTHMKM tt36eraroT cBeTa, aKTMBHhI HO'-lhIO. BoKpyr poTOBoro OTBepcTMR Haxoµ;.HTCR ocRaaTeJihHble

~yrraJihIJ;a,

coµ;epma~ne

peIJ;errTopHhie

KJieTKM opraHOB XMMM'-leCKOro '-IYBCTBa M OC.H38HMR. B IIOBepxHOC'rHOM CJIOe
KO)Klf paCIIOJIO.iKeHbl HepBHbie KJieTKM,

BOCIIPMHMMalO~Me

XMMM'-leCKMe paa-

µ;pa;KeHMR. Ha rrepeµ;HeM KOHIJ;e HepBHoii Tpy6KM Haxoµ;MTCR opraH o6oH.HHM.H.
OpraHa cJiyxa HeT.

lluw,eeapumeJlbHaJl cucmeJlta Ha'-IMHaeTCR rrpeµ;pOTOBOM BOpOHKOM, OKpymeHHOM

10-20 rrapaMM

~yrraJieIJ; (IJ;eµ;HJihHhIM arrrrapaT). B rJiy6nHe BopoH-

KM Haxoµ;MTC.H POTOBOe OTBepCTMe. Ilo TMIIY IIMTaHMSI JiaHIJ;eTHMK -

l}lMJihT-

paTop. Boµ;a c rrn~eBoii B3Bechro rrorraµ;aeT B rJIOTKy, rrpoxoµ;11T CKB03h ma6epHhre

~eJIM

(100-125)

M

rrorraµ;aeT

B

OKOJIO;Ka6epHyIO

IIOJIOCTh,

OTKphIBaro~ylOC.H Hapymy OTBepCTHeM (aTpMorropoM). BaBeIIIeHHhle IIM~e

Bhie qacTHJ.J;hI oceµ;aroT Ha µ;He rJIOTKM. Ilo cncTeMe 6opoaµ;oK, pacrroJiomeHHhIX Ha 6pIOIIIHOM M CIIMHHOM CTOpOHax, IIM~eBhle qacTMIJ;hI IIOCTyrraIOT B KMIIIe'-IHMK. B rrepeµ;Heii qacTM Kttrne'-IHMKa nMeeTC.H rreqeHoqHhIM BhrpocT. IlepeBapttBaHne H BCaCbIBaHMe

IIM~H

rrpoucxoµ;MT KaK B KMrneqHMKe, TaK

M B rreqeHM. HerrepeBapeHHhie OCTaTKM BbIBOµ;RTC.H M3 3~Heii: KMIIIKM qepea
aHaJihHOe OTBepCTMe,

OTKphrnaro~eecR

y OCHOBaHMJl XBOCTOBOro IIJiaBHMKa.

l{btxameJtbHaR cucme.+ta rrpeµ;cTaBJieHa .ma6epHhIMM rreperopoµ;KaMM.
Boµ;a, Bxoµ;R~a.H B rJIOTKy, 6Jiaroµ;apR MepIJ;aTeJibHOMY µ;Bu.meHMIO pecHMqeK, BblCTMJialO~MX BCIO IIOJIOCTh KMrneqHOM Tpy6KM, OMblBaeT .ma6epHble
rreperopoµ;KM, M coµ;ep.ma~MMC.H B BOµ;e KMCJIOpoµ; IIOCTyrraeT B KpOBh, µ;BM.my~yrocsi: no TOHKMM cocyµ;aM .ma6epHhIX rreperopoµ;oK. JKa6epHble ~eJIM

OTKpbIBaIOTCR B aTpMaJibHyro, MJIM OKOJio.ma6epHyIO IIOJIOCTb. ,D;hIX8HMe ocy~eCTBJIReTCR

oµ;HOBpeMeHHO c IIHTaHMeM.

BbtaeJtumeJtbnaR cucme.Ma rrpeµ;cTaBJieHa Hel}lpuµ;HRMH (100-150 nap),
pacrroJio.meHHhIMM H~ rJioTKOM B Me.m.ma6epHhIX rreperopoµ;Kax. Hel}lptt,D;MM
rrpeµ;cTaBJieHhI KOpOTKMMM Tpy6oqKaMH, OTKPhIBalO~MMMCR Ha O,D;HOM KOHIJ;e
HeCKOJihKMMM OTBepCTMSIMM -

Hel}lpoCTOMaMM B u,eJlO.M, a Ha µ;pyroM -

oµ;HMM

OTBepcTMeM B 01COJlOJ1Ca6epny10 noJtocmb. Hel}lpocTOMhI coµ;ep.maT 6yJiaBoBM,D;Hhle KJieTKM -

COJieHOIJ;MTbl. Ilpoµ;yKTbI Bblµ;eJieHMR BblBOµ;RTCR B OKOJIOJKa-

6epHyIO IIOJIOCTb Herrocpe,n;CTBeHHO H3 IJ;eJIOMa.

KpoeenocnaR cucme.Ma aaMKHyTaR. Oµ;ttH Kpyr KpOBoo6pa~eHM.H, cep,n;IJ;a
HeT. PoJib cep,n;IJ;a BhIIIOJIH.HeT rryJihCMpyro~aR 6prornHaR aopTa, OT KOTopoii: oTxo,n;si:T IIpMHOCR~Me .ma6epHble apTepMM (BeH03HaSI KpOBh), rrpoxo,n;SIIIJ;Me
B Me.m.ma6epHhIX rreperopoµ;Kax. JKa6epHhle apTepttM He o6paayroT Karr11JIJI.HpHhIX paaBeTBJieHMM. 06ora~eHHaSI KMCJIOpoµ;oM KpOBb no BhIHOCR~MM .ma6ep-

J[u~uno~noxopoo6we

313

HhlM apTepH.HM co611paeTCH B KOPHH aopThI, KOTOphie K nepe.n;HeMy KOHIIY CJIHBalOTCH B COHHhie apTepHH, a K 3a.n;HeMy KOHIJ;y TeJia CJIHBaIOTCH H o6pa3yIOT
CIIHHHYIO aopTy. 0Ha HaXO,ll;HTCH rro.n; xop.n;o:H H pa3BeTBJI.HeTCH Ha apTepHH,
lf.ll;YIIIHe KO BCeM opraHaM.
BeH03Ha.H KpOBh OT rrepe.n;He:H 'laCTH TeJia co611paeTCH B rrapHhie rrepe.n;HHe
Kap.n;HaJihHhie BeHhI; OT 3a.n;Hell 'laCTH -

B napHhie 3a,ll;HHe Kap.n;HaJihHhie Be-

Hhl. Ha.n; 3a.n;HHM KOHIIOM rJioTKH OHH CJIHBaIOTCH B KroBhepoBhI npoTOKH
l{

Brra.n;aroT B BeH03Hhlll CHHyc, a .n;aJiee BHOBh B 6prournyro aopTy. OT BHYT-

peHHHX opraHOB KpOBh co611paeTCH B IIO,ll;KHille'!HYIO BeHy' KOTopa.H B rreqeHO'lHOM BhlpOCTe pacrra.n;aeTCH Ha ceTb KaIIHJIJIHpOB, o6pa3y.H BOpOTHYIO CHCTeMy. Ilo KOPOTKOH ne'leHO'lHOH BeHe KpOBh BJIHBaeTCH B BeH03Hhlll CHHyc.
KpoBh JiaH11eTHHKa 6ec11BeTHa H co.n;epmHT He3Ha'!HTeJihHOe KOJIH'leCTBO
KpOBHHbIX KJieTOK.

lloJtoea.R cucmeMa. Pa3.n;eJihHOIIOJihie. IloJIOBhie meJie3hI uapHhre (npHMepHo

25

nap), pacnoJioiKeHhI B cTeHKax OKOJIOma6epHo:H uoJIOCTH. IIocTOHHHhIX rrpo-

TOKOB He HMeIOT' IIOJIOBhie rrpo.n;yKThl BhlBO,ll;.llTCH qepe3 BpeMeHHhie npoTOKH
B oKoJioma6epHyro noJIOCTh H .n;aJiee qepea aTp»onop BO BHenrn1010 cpe.n;y. OnJio.n;oTBopeHHe HapyiKHOe, pa3BHTJ{e c JIH'lHHO'lHOH CTa.n;11e:H.

lfa

3HrOThI pa3BHBa-

eTCH JIH'lHHKa (ma6epHhie 111eJIJ{ OTKphIBaIOTCH Hapymy), IIOKphITa.H peCHH'!KaMH, KOTOpa.H aKTHBHO IlHTaeTCH, pacTeT H qepe3 TPH Mecmi;a oce.n;aeT Ha ,ll;HO,
npeBpa111aHCh BO B3pOCJIOro JiaHIJ;eTHHKa.

Ilo)J;THrr rroauoao11ai.ie, HJIH 11eperrHLie, -

Ha1160Jiee BhICOKoopraHH30BaH-

Hhle xop.n;OBhie :lKHBOTHhle.

IIpoipeccueHbte 'lepmbt:

1.

2.

Xp.H111eBOH mm KOCTHhlll BHyTpeHHHH CKeJieT (Il03BOHO'!Hhlll CTOJI6).
fOJIOBHOH M03r COCTOHT H3 Il.HTH OT.n;eJIOB. Pa3BHBaIOTC.ll CJIOiKHhie opraHhl 'lYBCTB: 3peHHH, o6oHHHH.H, OC.H3aHHH, paBHOBeCH.ll, CJiyxa, BKyca,
opraHhl 6oKOBOH JIHHHH.

3.
4.
5.

KocTHo-xpH111eBo:H qepen, 3a111u111aro11111:H roJIOBHOH M03r.

6.
7.

OpraHhl Bhl.n;eJieHHH -

8.

KomHhle noKpOBhl ycJIO:lKHHIOTCH, H noHBJIHIOTCH aa111FITHhie o6pa3oBa-

YcJIOiKHeHHe rroBe.n;eHHH, 3a60Ta o noTOMCTBe.
3aMKHyTaH KPOBeHOCHa.ll CHCTeMa; cep.n;11e pacrrOJIOiKeHO Ha 6pIOilIHOH
CTOpOHe TeJia.
no'!KH.

AKTHBHOe rrHTaHHe npHBeJio K pa3BHTHIO .n;BHraTeJihHOH aKTHBHOCTH
H IlOHBJieHHIO opraHOB 33XBaTa IlHllIH -

qeJIIOCTeH.

HHH (qelllyH, neph.H, BOJIOChl), meJie3bl.

9.

Pa3BHBaIOTCH rrapHhle KOHe'!HOCTH H HX no'.Hca.

II 0360HO'lHbte rro.n;paa.n;eJIHIOTCH Ha .n;Be 6oJihllIHe rpyrrrrhl:
AHaMHUU, UJlU HU3utue ll0360HO'lHbte, KOTOphle o6'be,ll;HHHJOTCH B TpH KJiaCca ( KJtacc Kpyi.!Wpombte, KJtacc Pbt6bt, naoKJtacc 3eMH0600Hbte) Ha ocHoBaHHH CJie.n;y1011111x oco6eHHOCTe:H:
B Ka'leCTBe opraHOB ,IJ;hlXaHHH HMeJOT ma6phl HJIH MellIKOBH,ll;Hhie JierKHe;
npH pa3BHTHH HH1Ia He 06pa3yeTCH aapo.n;hIIlleBaH o6oJIO'!Ka.

314

AM1tuombt, UJlU Bbtcwue no:JBOltO'lHbte, o6'be.n;aH.HIOTC.H TaKJKe B Tp11 KJiacca
(KJiacc IlpecMhIKarorn;Hec.a, KJiacc IlTHIJ,hI, KJiacc MJieKOIIHTarorn;u:e) Ha ocHoBaHHH CJie.n;yrorn;HX oco6eHHOCTeil:
JKa6epHoro .n;blxaHH.H HeT;
rrpH pa3BHTHH .a:ih:i;a ct>opMu:pyeTc.a 3apo.n;hmieBa.H o6oJioqKa-aMHHoH .

Knacc Kpyrnop0Tb1e
OcHOBHhie npH3Haim:
1. ll;eHTpaJihHa.H HepBHa.fl CHCTeMa no.n;pa3.n;eJI.HeTC.H Ha rOJIOBHOM H ClIHHHOH M03r, pa3BHBaIOTC.H 6oJiee coBeprneHHhle opraHbI qyacTB, qeM y JiaHu,eTHHKa.
2. <l>opMHpyeTC.H xp.arn;eao:H cKeJieT B BH,n;e qeperra, BHCIJ,epaJihHoro CKeJieTa H 3aqaTKOB BepXHHX .n;yr II03BOHKOB.
3. HacTo.arn;wx qeJIIOCTeH HeT, poT OTKPhIBaeTc.H B rJiy6wHe cBoeo6pa3HOii:
npncaCbIBaTeJihHOH BOPOH.KH, KOTOpa.a no.n;.n;ep:m:irnaeTC.H xparn;oM, HMeIOII.\HM q,opMy KOJibU,a. POT BOOpy:meH porOBblMH 3y6aMH.
4. B KpOBeHocHOH CHCTeMe nmrnmi:eTca ueHTPaJihHhIH opraH - cep,n;u;e.
5. Bo3HHKaIOT 6oJiee coBeprneHHble opraHhl ,n;hrxamrn B BH,n;e .ma6epHhlX
MeIIIKOB.
6. Pa3BHTHe c JIHqHHOqHoii: cTa.n;Heii - rrecKopoHKH (!l>HJibTpaTopa), :>KHByrn;eii B necKe, KaK JI8HIJ;eTHHK.
K KJiaccy OTHOC.HTC.H .MUKCUHbl - MOpcKMe q,opMbI, napa3HTHpyrorn;we Ha pb16ax; MHHOrH - MOpCKHe H npeCHOBOAHhie ¢opMhI, T8Ka<e napa3HTHpyrorn;He Ha
pbI6ax (rroJiynapa3MThl).
Cmpoe1tue pe'lnou .MUHotu (pHc. 127). 3MeeBH.n;Ho y.n;mrHeaaoe TeJio, ,n;ocTHra10rn;ee 40 CM ,ll;JIHHhl. B TeJie pa3JIHqa10T TPH OT.n;eJra: roJioBy, TYJIOBlill.\e
H xBocT. IIapHhle KOHeqaocTH OTCYTCTBYIOT. Ha cnHHHOH cTopoHe xoporno 3a-

2

3

4

25 24

5

9

22 21 20

7

19 18

8

10

13 12 11

Piiie. 127. npOAOllbHbli:I pa3pe3 pe4HOi1 Mlt1HOn1: 1- xpsu~ltl poTOBOH BOPQHK1t1; 2- H03ApSI; 3ni-i~eBOA; 6 - XOPA3; 7 - 060110'-iKa XOPAbl; 8 cnHHHOi1 M03r; 9 - npeAcep,A1-1e; 10 -cni-iHHasi Myct<ynarypa; 11 - nonoaasi nopa; 12- MO'ienonosoe
OTBepcrne; 13- aHyc; 14 - M04enonosoi1 CHHyc; 15 - MOYeTOYHHK; 16- f104Ka; 17- flOJlOBaSI lKene33; 18- Kl-IWKa; 19- ne4eHb; 20- seH03H3SI na3yxa; 21 - OK0110CepAe4HblH xpsi~; 22- lKenyAOYeK;
23 - lK36epHble MeWKH ; 24 - MYCKYl13TYP3 Sl3b1Ka ; 25 - rJlOTKa; 26 - POTOB3SI nollOCTb
00oHSITellbHaSI Kancyna; 4 - ronOBHOi1 M03r; 5 -

315

CpaBHHTeJILHaH xapaKTepHCTHKa 6ecnoauouo11uL1x H noauouo11HL1x :IKHBOTHLix
Becno3BOHO'-!Hh1e

Jlp113HaKl1
C1<eJieT

HepsHa.II c11cTeMa

i HapymIIhIH
i

i BnyTpeHmi:ii:: xop-

Li:mpcpyaHoro 11JI11
Y3JIOBoro Tn:nos.
I HepBHble CTBOJihl

CIIHHHOH Moar B BHfoJioBHOH M03I' COCTOHT
113 Il.IITH OT,ll;CJIOB
1

1

OpraHhI .n;L1xa1111.1I:
MeCTOIIOJIO)KeH11e
OpraHhl ,ll;hrxaHn:H:
cPOPMhl

Ifpy.n;h, 6pIOIJlKO

cep.n;~a

OpraHbI 3axsaTa
IIHillH

i~

CrrHHHOH M03r
B BH,ll;e Tpy6x11

I .n;e Tpy61rn.

fOJIOBa,rpy.n;h

! l'oJIOBa (r JIOTKa)
I

BhrpoCThI KHIIIe'-!HH- : :IB:a6ph1
xa: IIJiaBaTeJihHhIH I
rryaLrph (y .n;peBHHX
1
1103BOHO'lHhlX), Jierx11e, ma6ph1, xoma

0TCYTCTByeT,
BTop11'-!Ha.1I(~eJioM) ; BTop1111Ha.1I (r~eJIOM)
11epB1111Ha.1I,
I
I
BTOp1111Ha.1I (~eJIOM)
I

0.n;HOKaMepHOe HJIH ' 2-, 3-, 4-KaMepHoe,
i MHOI'OKaMeyHoe,
i Ha 6pIOIIIHOH
Ha CIIHHHOH
CTOpoue TeJia
CTOpOHe TeJia

KposeHocua.H
CHCTeMa
PacrroJiomeHne
opraHOB 'lYBCTB

.n;a, KOCTHhlH HJIH
XP.IIIIIeBOH

JlaHIWTHl1K11
BnyTpemmii:
(xop.n;a)

I

TiosepxHoCTh TeJia,
,ll;bIXaTeJihHhJe
! Tpy6KH, Tpaxe11,
, ma6ph1, JierK11e

IIoJIOCTh TeJia

CTpoeHne n
UOJiomeHHC

Jl03BOHO'lllblC

He3aMKflYTa.H(KpoMe 3aMKHyTa.H
KOJih'laThlx 11epseil:)
1

PaaJIH'lHhle 11acTn

HeT

I
I

i 3aMKHyTa.II

ifoJioBa

PaaJIH'IHhle 11acTn
TeJia

i TeJia
KoHe11uocTn

qeJIIOCTH;
Y HCKOTOphlX KOHC'-IHOCTH

I

HeT; IIHTaIOTClI
naCCHBHO

I
I

MeTHhI ,n;Ba KO.mHCThIX HenapHhIX IIJiaBHHKa, IlOCJie,ll;HHH H3 KOTOphIX nepexo,ll;HT Henocpe,n;CTBeHHO B XBOCTOBOH ITJiaBHHK.
Crrepe,n;H Ha roJIOBe urn:poKa.H npncachIBaTeJihHa.H BopoHKa, no Kpa.HM KOTopo:i1: pacnoJio.meHhI KO.mHCTbie BhlpOCThI. BHyTpH BOpOHKH H Ha KOHUe
MOIIJ;Horo .H3hIKa Haxo,n;.HTC.H porOBhie 3y6h1. Koma IIJIOTHa.H, roJia.H, 3ITH,n;epMHc co,n;epmHT .meJie3HCThle KJieTKH, BhI,ll;eJI.HIOru;He Ha IIOBepXHOCTh TeJla
CJIH3h. HenapHoe H03,n;peBoe OTBepcTHe pacnoJio.meHo y MHHOr Ha Bepxy roJIOBhI, Me.m,n;y rJJa3aMH, y MHKCHH OHO Haxo,n;HTC.H Ha rrepe,n;HeM KOHUe roJIOBhI. Ilo3a,n;H H03,ll;PH HMeeTC.H CBeTO'IYBCTBHTeJihHOe II.HTHO. Ilo 6oKaM nepeµ;Hero KOHUa TeJia p.Hµ; OKpyrJihIX (a He ru;eJieBH,ll;HhIX, KaK y pbI6) ma6epHhlX
OTBepcTHH.

316

300ll0ZUJt

OceBoii cKeJieT TYJIOBMrn;a o6paaoBaH xop.z:i;oli, B coe,11;MHHTeJ1hHOTKaHttoii
o60JIO'!Ke KOTOpoii 3aKJia,ll;bIBaIOTCH 3JieMeHThI no3BOHKOB: 3a'IaTKM BepxHux
.z:i;yr n03BOHKOB • .IJ:nH aarn;HTbI rOJIOBHOro M03ra pa3BMBaeTCH qepenHa.H Kopo6Ka. Xp.Hrn;eBa.H ma6epHa.H perneTKa COCTOHT M3 He paC'IJieHeHHhIX Ha OT.z:i;e.'IhI
BepTMKaJlbHhIX ma6epHbIX .z:i;yr.
TyJIOBMrn;Ha.H MycKyJiaTypa cerMeHTHpoBaHa. IloJioCTh Tena -

LWJIOM, 110 _

KphlThIM TOHKOM 3nHTeJIMaJihHOM o6ono'IKOM (6pIOlIIMHoli).

IIuw,eeapumeJlbttaJl cucmeMa

Ha'IHHaeTCH wupoKoli npe.z:i;poToBoli Bopou-

Koli, BOOpymeHHOM porOBblMH 3y6aMM. PoTOBOe OTBepCTHe nemHT Ha ,ll;He BOpOHKH M Be,ll;eT B rJIOTKy.

y

MMHOr rJIOTKa pa3.z:i;eJieHa Ha ,ll;Be 'IaCTM: BepXHIOIO -

nMrn;eBapMTeJihHYIO (nMrn;eB0,11;), no KOTOpoli npOXO,ll;MT nHrn;a, M HMmHIOIO ,ll;bIXaTeJibHYIO Tpy6Ky. :/Keny.z:i;o.K He pa3BHT, M nMrn;eB0,11; nepeXO,ll;HT Henocpe.z:i;CTBeHHO B .KMlII.KY' .KOTopa.H HMeeT CnHpaJibHhIM .KJianaH B BH,ll;e M3rH6arorn;eikH
C.KJI~.KH

,ll;JIH yBeJIM'IeHHH noBepXHOCTM .KMlII.KM H 3aMe,11;J1eHMH npoxom.z:i;eHHH

nHIIW. 3~HHM .KOHeIJ; .KHlIIe'IHOM Tpy6.KM OTKPbIBaeTCH aHaJihHbIM OTBepcTMeM.
KpyrJiopoTwM CBOMCTBeHHO Bhl,ll;eJIHTh rmrn;eBapMTeJihHhre co.KM B TeJio mepTBhl, r.z:i;e H Ha'IMHaeTCH XMMM'IeC.Ka.H o6pa60T.Ka llMIIl;M (BHe.KHlIIe'IHOe nMrn;eBapeHHe ). Ile'IeHh pa3BMBaeTCH .Ka.K BhipocT 11epe,11;Hero OT,ll;eJia .KMlII.KH. Ilo.z:i;meny.z:i;oqHaH meJieaa .z:i;mpcpy3Ha.H. Ee .KJieT.KM pacce.HHhl 110 CTeH.Ke .KMrne'IHHKa.

/{btxameJlbHaJl cucmeMa

11pe,11;cTaBJ1eHa ma6epHhlMH Mern.KaMH (ceMh nap),

pac110JiomeHHhlMH no 60.KaM .KOpOTKOM, CJ1e110 3a.KaH'IMBaIOrn;eiic.H ,ll;hIXaTeJibHOM Tpy6.KM. CTeH.KM MelII.KOB 110.KphlThl MHorO'IMCJieHHhIMH nJiaCTHH.KaMM ma6epHbIMM JienecT.KaMM, npOHM3aHHhlMM KpOBeHOCHhlMM .KallMJIJI.HpaMM, r,11;e
npoHCXO,ll;HT raaoo6MeH. BHyTpeHHHe OTBepCTMH ma6epHbIX MelII.KOB OTKphlBaIOTCH B ,ll;bIXaTeJihHYIO Tpy6.Ky' a HapymHhie -

BO BHelIIHIOIO cpe.z:i;y. :/Ka6epHbie

Melli.KM M JieneCT.KM MMeIOT 3HT0,11;epMaJibHOe npoHcxom.z:i;eHMe.
B CBH3H c napa3MTH'IeCKHM THllOM IIHTaHHH (nyTeM llpHcacbIBaHHH K TeJiy
mepTBhl) TO.K BO,ll;hl npH ,ll;hlXaHHH H,ll;eT qepe3 HapymHhle ma6epHhie OTBepCTMH
B ma6epHbie MelIIKH H BhIXO,ll;HT o6paTHO 3THM

Kpoee11oc11aJl cucmeMa

me nyTeM.

aaM.KHyTa.H. Cep.z:i;ri;e cocTOHT M3 ABYX .KaMep -

npe.z:i;cep.z:i;MH H meny.z:i;oqKa. K npe.z:i;cep.z:i;HIO rrpHMhlKaeT TOH.KOCTeHHhlM BeHo3HhlM CHHyc (BeH03HaH naayxa), .Ky.z:i;a Bna.z:i;a10T Bee BeH03Hhle cocy.z:i;w. Bee OT.z:i;eJiw cep.z:i;ri;a co.z:i;epmaT BeH03HYIO .KpOBh. OT rrepe.z:i;Heli qacTH meJiy,ll;O'IKa OTXO,ll;HT 6pIOlIIHa.H aopTa, KOTOpa.H .z:i;eJIHTCH Ha rrpHHOCHIIl;He ma6epHbie apTePMM,

H.z:i;yrn;He K

Memma6epHhIM rreperopo,11;.KaM.

BwHoc.Hrn;He ma6epHhie

apTepHH Bn~aIOT B HerrapHbIM .KopeHh aopThl, OT KOToporo OTX0,11;.HT Bnepe,11;
COHHwe apTepHH, a Ha3a.z:i; -

CllHHHa.H aopTa. KpoBh OT cnHHHOM aopThI no-

cTyrraeT KO BCeM opraHaM Tena. OT roJIOBhl BeH03Ha.H KPOBh co6HpaeTCH
B napHhie rrepe.z:i;HHe .Kap,11;HaJihHhle BeHw, Bna.z:i;arorn;He B BeH03HYIO rra3yxy.
Cro.z:i;a me rrocTyrraeT KPOBh H3 aa.z:i;HHX .Kap,11;HaJihHhlX BeH, co6Hparorn;Hx
KpOBh OT TYJIOBHrn;a. OT .KHlIIe'IHHKa KpOBh co6HpaeTCH B no,11;.KHlIIe'IHYIO BeHy; nonaB B neqeHh, OHa o6paayeT BOpOTHYIO CHCTeMy KPOB006parn;eHHH.
Ha rre'IeHH KpOBh rro rre-•rnHO'IHOM BeHe H3JIHBaeTCH B BeH03Hyro rraayxy. BoPOTHOM cHcTeMhI noqe.K HeT.

317

HepBHaJl cucmeMa cocTOHT

H3 roJIOBHoro H cnHHHoro M03ra. Mo3meqoK noq-

Tfl He pa3BHT. Bee OT,ZJ;eJihl pacnoJiomeHhl B O,ZJ;HbH nJIOCKOCTH, TO eCTh He o6pa3yJOT THnHqHhIX .ZJ;JIH 6oJiee BhICOKoopraHH30BaHHhIX no3BOHoqHhIX H3rH6oB. KpbIuia M03ra He HMeeT HepBHoro Be~eCTBa H u;eJIHKOM 3nHTeJIHaJihHaa. OT roJIOBHo-

10

nap rOJIOBHhIX HepBOB. CnHHHOM03rOBbie HepBbl OTXO,ZJ;HT

,ZJ;BYMH KOpernKaMH -

cnHHHhIM H 6pIOIIIHhIM, KOTOphie, B OTJilfqHe OT .n;pyrHx

ro M03ra OTXO,ZJ;HT

rr03BOHOqHhJX, He coe.n;HHHIOTCH H He o6pa3yIOT o6~ero CMernaHHOro HepBa.

Opianbt 'lyBcmB
peHHHM YXOM -

ycTpoeHhl oqeHh npocTo. OpraH cJiyxa npe.n;cTaBJieH BHYT-

nepenoHqaThIM Jia6HpHHTOM c .ZJ;BYMH (y MHHor) noJiyKpymHbl-

MH KaHaJiaMH. I'Jia3a pa3BHThI cJia6o, pacnoJiomeHhI no 6oKaM roJIOBhI. HMeIOT
cTpOeHHe THnlfqHOe ,ZJ;JIH no3BOHOqHhIX mHBOTHhIX, HO noKpbIThl noJiynpo3paqHOH 1wme:H. MHHOrH MoryT BH,ZJ;eTh KOHTyphI no.n;BO.ZJ;HhIX npe.n;MeToB JIHIIIh Ha
6JIH3KOM paccTOHHHH. OpraH XHMHqecKoro qyBcTBa (o6oHHHHH) npe.n;cTaBJieH
uenapHOH H03,ZJ;pe:H, HOCOBhIM XO,ZJ;OM H CJienhIM BblpOCTOM, 3aKaHqHBaIO~HMCH
no.n; qepenoM. B

Rome c o6eHx CTopoH TeJia npoxo.n;HT 6oKoBaH JIHHHH. Ona

npe,ZJ;CTaBJieHa HerJiy60KHMH HMKaMH, Ha ,ZJ;He KOTOpbIX pacnoJiomeHbl OKOHqaHHH 6.Jiym.n;aIO~ero HepBa

(X

napa roJIOBHhIX HepBOB). BocnpHHHMaeT TOKH BO-

,ZJ;bl, perHCTPHPYH npH6JIHmeHHe .n;pyrHX mHBOTHhIX HJIH BCTpeqy c npenHTCTBHHMH. Ha roJioBe y MopcKo:H MHHOrH HMeeTCH cJia6hrH 3.JieKTpHqecKHH opraH.
Ilo H3MeHeHHIO 3.JieKTpHqecKoro nOJIH mHBOTHOe MOmeT qyBCTBOBaTb npH6JIHmeHHe .n;pyroro opraHH3Ma HJIH npenHTCTBHH. B

TOJI~e KomH paa6pocaHhl

HepBHhre oKoHqaHHH MexaHo-, xeMo- H TepMopeu;enTopoB.

BbtOeJtumeJtbHllJl cucmeMa

npe.n;cTaBJieHa TYJIOBH~HhIMH, HJIH nepBHqHhI-

MH, noqKaMH, HMeIO~HMH JieHTOBH.ZJ;HYIO <PopMy H Jiema~HMH no 6oKaM cnHHHOH aopThI B.ZJ;OJih Bce:H 6p10rnuo:ii noJIOCTH. MoqeToqHHKH (BOJih<POBhI KanaJihl), OTXO,ZJ;H~He OT napHOH noqKH, Bna.n;aIOT B MOqenoJIOBOH CHHyc, OTKpbIBa-

lO~HHCH Hapymy Ha KOHu;e MoqenoJioBoro cocoqKa.

IIoJtoBaR cucmeMa.

Pa3.n;eJihHOnOJihre. IloJIOBhre meJie3hI HenapHhre, cneu;H-

aJihHhIX npOTOKOB He HMeIOT. IloJIOBhie npo.n;yKThl qepe3 pa3pb1Bhl CTeHOK

me-

Jie3bl Bhlna,ZJ;aIOT B 6pIOIIIHYIO nOJIOCTh, OTKy.n;a BhIBO,ZJ;HTCH B MoqenoJIOBOH CHHYC H qepe3 Kana.JI MoqenoJIOBoro cocoqKa uapymy. OnJIO.ZJ;OTBopeHHe Hapymuoe. Pa3BHTHe c MeTaMop<}}o3oM. JIHqHHKa (necKopo:HKa) mHBeT B peKax, H JJ:O
ee npeBpa~eHHH BO B3pOCJIYIO MHHOry npoxo.n;HT HeCKOJihKO JieT.

HaA1<11acc

pb16b1

OcnoBHbte apoMop<jJoabt:

1.

3aMeHa xop,ZJ;bl XpH~eBhIM, aaTeM KOCTHhIM n03BOHQqHHKOM.

2.
3.
4.
5.

AKTHBHhIH nOHCK H 3axBaT nH~H (nOHBJieHHe qeJIIOCTeH).

6.

Bo BHyTpeHHeM yxe nOHBJIHIOTCH TpH noJiyKpymHhIX KaHaJia, Jiema~He

06pa3oBaHHe qepena, 3a~H~mo~ero M03r.
Bo3HHKHoBeHHe napHhIX KOHeqHOCTe:H -

nJiaBHHKOB.

Pa3BHTHe roJIOBHoro Mo3ra H opraHoB qyBCTB. Be.n;y~HH oT.n;eJI roJIOBHOro M03ra -

cpe.n;HHH M03r (HXTHOnCH,ZJ;HhIH THn M03ra).

BO B3aHMHO nepneH,ZJ;HKYJIHPHhIX nJIOCKOCTHX.

318

30011ozu11,

KpoBoo6pa~emrn:. Cep,n;u;e ,n;ByxKaMepHoe. B cep,n;u;e Bea 03 _
Ha.H KpOBb. y ,ll;BO.HKO,ll;hlIIIa~HX pb16 HaMeqaeTC.H 06paa0Bam1e BTOporo,
(neroqHoro) Kpyra KpOBoo6pa~emrn.
8. 06oHHTenhHhie MeruKH rrapHhie. KaJK,n;hlil: M3 HHX OTKphrnaeTc.a Hapy)l\y
caMOCTO.HTenhHhIM HOCOBhIM XO,ll;OM (H03,ll;pH napHbie).
9. OpraHhI ,D;hIXaHHH - JKa6phI.
10. II11~eBap11TenhHhiil: TpaKT ,n;mpcpepem111poBaH: o6oco6n.aeTc.a JKeJiy,n;oK,
KHilieqHHK no,n;pa3,n;en.HeTC.H Ha TOHKMM H TOnCThIM OT,n;enhI.
11. ,r:i:11cpcpepeHu;11poBKa rr11~eBapnTenhHoil: JKeJie3hI Ha no,n;JKeny,n;oquy10
11 rreqeHh.
12. 06paaoBaHHe TynoBH~HhIX rroqeK (nepBHqHhIX rroqeK Me3oHecppoca).
13. B KOJKe B03HHKaIOT aa~HTHhie, rrpettMy~ecTBeHHO KOCTHhie o6paaoBaHHH - qeruyM.
3TOT rro,n;Knacc BKJIIoqaeT ,n;Ba COBpeMeHHhIX Knacca pb16: XPRW,e6ble H 1(0cmHble. Cpe,n;a 0611TaHMH - MopCKMe 11 rrpeCHhie Bo,n;oeMhI. B HaCTO.H~ee BpeMH H3BeCTHO 6onee 20 TbIC. BH,ll;OB.
IIpe,n;cTaBHTenh KJtacca Kocmttbte pbt6bt - pe•uwu oKyttb. 0THOCHTCH K ompRiJy
0Kyneo6pa3Hbte ( naiJompJliJ Kocmucmbte ). PacrrpocTpaHeH B rrpeCHhIX Bo,n;oeMax.
Cmpoenue pe'lttozo OKYHR. 06TeKaeMa.H cpopMa Tena, yrrno~eHHaH c 6oKoB.
Teno COCTOMT 113 i:'onOBhI, TynOBH~a H XBOCTa. fpaHMIJ;eM MeJK,n;y ronOBOM M TYnOBH~eM cnymHT JKa6epHa.H ~enh, a MeJK,n;y TYJIOBH~eM H XBOCTOM - aHaJihHOe OTBepCTMe.
PoT y KOCTHCThIX phl6 pacrronomeH Ha rrepe,n;HeM KOHu;e ronOBhI, OTqero TaKHX phl6 Ha3hIBaIOT KOHeqHopoTbIMM (B 0Tn11qwe OT rrorrepeqHOpOThIX pb16 aKyn). IIo 6oKaM ronoBhI Haxo,n;HTCH KpyrrHhie nnocKHe rnaaa. Brrepe,n;H JiemaT
rrapHhle Hoa,n;pw - opraHhl o6oHHHHH. Hoa,n;p11 OTKphIBaIOTCH B 060H.HTeJihHy10
HMKY 11 He coo6~aIOTCH Mem,n;y co6oil:.
:IB:a6epHa.H KphlllIKa KOCTHa.H. B CB.H3H c 3TMM BMeCTO Il.HTH ma6epHblX OTBepCTHM (y xp.a~eBhIX pb16) o6paayeTCH o,n;Ha JKa6epHa.a ~enh.
IIapHbie rrnaBHMKH (rpy,n;Hhie, 6pIOilIHhle) pacrrono)R;ehI B BepTHKanhHOii:
rrnoCKOCTH. CrrttHHhie nnaBHMKH CHa6meHhl MHrKMMH BeTBMCThIMH 11 TBep,n;hIMH Kon10q11MH nyqaMH. Jlyqw CBH3aHbl Mem,n;y co6oil: TOHKOM KOJKHCTOM rrepeIlOHKOM. XBOCTOBOM nnaBHHK roMou;epKaJihHoro TMrra (paBHonorracTHoil:) ocHOBHOM opraH rrepe,n;BHmeHHH. Ha HHmHeil: cTopoHe Tena 6nwme K aa,n;HeMy
KOHIJ;Y HaXO,ll;HTC.H ITO,ll;XBOCTOBOM, HnH aHaJihHhlM, nnaBHHK. Ilepe,n; HHM B 06~eM yrny6neHHH nemaT TPH OTBepcTH.H: aHaJihHOe, rronoBoe H Bhl,n;enHTeJibHOe
(y xpn~eBhIX phl6 HMeeTC.H KnOaKa).
OpraHhl ,n;BHmeHHH, nJta6HUKu, ,n;ennTc.H Ha ,n;Be rpyrrrrb1:
rrapHhle (rpy,n;Hhle, 6pIOilIHhle) o6ecrreqwBaIOT rroBopoTbI, rro,n;,n;epJKaHHe
paBHOBeCMH, Me,n;neHHhie ,ll;BHmeHHH;
HerrapHhle (crrHHHOM, aHanbHbIM) 06ecrreq11BaIOT ycToil:qHBOCTh rrpH ,ll;BHmeHHH, xBoCTOBoil: BhlrronHneT cpyHKIJ;HH pyn.a.
B,n;onb Tena T.HHeTCH xopowo aaMeTHa.H 6oKoBa.H nMHHH - opraH optteHTaIJ;MH phl6.

7. O,n;1rn Kpyr

319
Te.!lO OICYHJl IIOKphITO KOCTHOH (raHOH,ll;HOH) qewyeli, COCTOHII~e:i1: H3 rrpo3pa'IHhIX KOCTHhIX IIJiaCTHHOK. Kam,n;aJI qerny:i1:Ka JJeJKHT B oco6oM KapMaHe. 1IewyH HaJJeraIOT ,n;pyr Ha .n;pyra B BH,n;e qeperrm~hI, 3a~11~a10~e:i1: Te;:ro
p.bI6hJ.
TeJJo p.b16b1 rroKp.bITO CJJH3.bIO, crroco6cTBYIO~e:i1: JJYTIWeMy rrepe,n;BHmeHHIO
OKYHJI B IIJJOTHOH BO,ll;HOH cpe.n;e, yMeH.blliaIO~eil TpeHHe rrpH IIJiaBaHHH H 06Jia-

.n;a10~eil 6aKTepm~H,ll;H.bIMH CBOHCTBaMH, rrperrJITCTBYIO~HMH rrpoHHKHOBeHHIO
B KO;KY 6aKTepHH.

CKe.Tlem KOCTHhIH. Pa3,n;eJJeH Ha OT,n;eJJhI: oceBoil cKeJJeT (rro3BOHO'IHHK),
cKeJieT roJJOBhl (qeperr) H CKeJJeT KOHe'IHOCTe:H (rrJJaBHHKH).

Ilo360HO'lHUIC ,n;eJJHTCJI Ha ,n;Ba OT,n;eJJa: TYJJOBH~HhIH H XBOCTOBOH. CocTOHT
H3 ,ll;BOJIKOBOrHYThIX II03BOHKOB, Mem.n;y KOTOphIMH coxpaHHJJHC.b OCTaTKH xop,n;.bl. TyJJOBH~HhJH II03BOHOK COCTOHT H3 TeJJa II03BOHKa, BepxHeH H HHmHe:H
,n;yrH. KoH~.bI BepxHHX .n;yr cpacTaIOTCJI, 06pa3y51 crrHHHOM03roBo:H KaHaJJ,
H 3aKaH'IHBalOTCJI ,ll;JJHHH.bJM HerrapHhIM OCTHCThIM OTpOCTKOM. HnmHHe ,n;yrH
paapacTaIOTCJI B CTOpOH.bl B BH,n;e ,n;Byx rrorrepe'IH.bIX OTpOCTKOB, K KOTOphIM
rrpH'IJJeHJIIOTCJI ,ll;JJHHHhJe H TOHKHe pe6pa.

y

XBOCTOBhIX II03BOHKOB xopowo

B.blpameH.bl HHmHHe .n;yrH, KOTOphle o6paayroT reMaJJhHhJH KaHaJJ H aaKaH'IHBaIOTCSI HH;KHHM OCTHCThIM OTPOCTKOM. IIo reMaJJ.bHOMY KaHaJiy rrpoxo,n;HT KpoBeHOCH.bJe cocy,n;hl. Pe6pa oTcyTCTBYIOT.
Ilo3BOHKH coe,n;HHSllOTCSI .n;pyr c .n;pyroM rrpH IIOMO~H cycTaBHhIX OTpOCTKOB, paCIIOJJO;KeHHhIX y OCHOBaHHSI BepXHHX .n;yr. TaKoe coe,n;HHeHHe o6ecrre'IHBaeT rrpO'IHOCTh CKeJieTa rrpH coxpaHeHHH ero IIO,ll;BHjf(HOCTH.

y

aKTliBHO

IIJJaBaIO~HX p.bI6 BCer.n;a HMeeTCSI pa3BHT.bIH II03BOHO'IHHK. Ilo3BOHO'IH.bIH
CTOJI6 y phI6 HeIIO,ll;BHmHO coe,n;HHeH c qeperroM.

tfepen COCTOHT H3 6oJihllIOro KOJJH'IeCTBa KOCTeli: M03rOBOH (qeperrHoii) KOpo6KH, qeJJIOCTHhIX KOCTeH, ma6epHhIX .n;yr H ma6epHhIX Kp.billieK.

CICe.Tlem ICOHe'lHO<;meu BKJJIOTiaeT IIOSIC H CKeJJeT CB060,n;H.bIX KOHe'IHOCTeH.
IIepBH'IH.bIH rrJJeTieBoH rroac rrpe.n;cTaBJJeH JJorraTKOH H KopaKOH,n;OM (BpaHoBo:H KOCThlO ). CBepxy K HeMy rrpHKperrJJHeTCSI 6oJJ.blliaSI ceprroBH,n;HaSI KOCTh c BhlTHHYThIM, aaocTpeHH.bIM OTpOCTKOM -

KJJeHTPYMOM.

c IIOMO~.bIO ,ll;BYX MaJJeHh-

KHX KOCTO'IeK BTOPH'IHOrO rroaca (Ha,n;KJJeHTPYM H 3a,ll;HeTeMeHHaSI KOCT.b) rrJie'IeBOH IIOSIC HeIIO,ll;BHjf(HO CO'IJJeHSieTCSI c qeperroM. Ta30BhIH IIOSIC rrpOCT H HMeeT
$opMy TpeyroJJhHOH KOCTH. Coe,n;HHeH c rrapH.bIMH 6pIOllIHhIMH rrJJaBHHKaMH.
JlemHT B

TOJJ~e 6pIOllIHOH MYCKyJJaTyp.bl H He coe,n;HHeH c II03BOHO'IHHKOM.

CKeJJeT CBo6o,n;HhIX KOHe'IHOCTeH (rrJJaBHHKOB) COCTOHT H3 xpa~eBhIX HJJH
KOCTHhIX JJyqeH, IIOKphIThIX Kome:H (pHC.

M blUle'lHllJl cucmeMa

128).

COCTOHT H3 OT,n;eJJ.bH.bIX cerMeHTOB, paa,n;eJJeHHhIX co-

MhlllI~ (maMhlllI~hI rrapH.bIX rrJJaBHHKOB). CoKpa~eHHH MhlllI~ o6ecrreTIHBaIOT H3-

e.n;HHHTeJihHOTKaHHhJMH rreperopo,n;KaMH. IlOHBJJSIIOTCSI rpyIIIIhl
6epH.bie,

I'H6aHHe II03BOHO'IHHKa H rrpHBO,ll;$IT B ,ll;BH;KeHHe XBOCTOBYIO JJOIIaCT.b.

H ep6Hll.R cucmeMa

COCTOHT H3 CIIHHHoro H roJIOBHOro Moara. I'OJJOBHOH

Moar rrpe,n;cTaBJJeH Il$lT.bIO OT,ll;eJJaMH: rrepe,n;HHM, rrpoMemyTO'IH.bIM, cpe,n;HHM,
npo,n;oJJroBaT.bIM H MoameTIKOM. Xopomo pa3BHThl o6oHHTeJI.bHhle ,n;oJJH rrepe.n;-

320

4
.5

128. CxeMa CTpoeH"15l KOCTHblX pb16: 1 - M03r; 2 - rna3a; 3 - 060H5lTellbHble S!MK"1; 4 - )l(a65 - cepAu.e; 6 - rpyAHOH n11aBH"1K; 7 - K"1WeYH"1K; 8 - 6p10WHOH n11aBH"1K; 9 - nOllOBaSl )l(eJle3a;
10- aHa!lbHOe OTBepcrne; 11 - aHa!lbHblH n11aBH"1K; 12- XBOCTOBOH n11aBH"1K; 13 - cn"1HHOH n11as-

PMC.
pb1;

H"1K;

14 -

n03BOHOYH"1K;

15 -

nOYKa;

16 -

n11aBaTellbHblH ny3bipb

Hero Moara. Ilepe,n;HMH Moar Marr rro paaMepaM, Kphlrrrn ::m11TeJIMaJibHa.H 11 He
co,n;ep:mnT MoaroBoro Bem;ecTBa.
TeJibHhle HepBbI. B

K

rrpoMe:>KyToqHoMy Moary rro,n;xo,n;.HT 3PH-

cpe,n;HeM Moare xoporno paaBMTbI ap11TeJibHhle u;eHTpbI.

Moa:meqoK OTBeqaeT 3a KOOp,ll;HHau;mo ,ll;BHJKeHHH H B CB.H3H co CJIOJKHhlMH ,ll;BHJKeHH.HMH B Bo,n;e ,n;ocT11raeT KPYIIHbIX pa3MepoB. B rrpo,n;oJiroBaToM M03re Haxo,n;.HTC.H u;eHTpbI peryJI.Hll;HH ,/J;bIXaTeJibHOH, KpoBeHOCHOH, IIHID;eBapHTeJibHOH
cncTeM, a TaK:me u;eHTpbI opraHoB qyBCTB (cJiyxa, 6oKoBoii JIMHHll). OT roJioBHoro M03ra OTXO,IJ;.HT ,n;ec.HTb nap qeperrHO-M03rOBbIX HepBOB.

OpzaHbt 'lyBcmB.

BKycoBbie peu;errTophl (opraHhl XHMlfqecKoro qyBCTBa)

pacrroJiaraIOTC.H ITO BCeii IIOBepxHOCTH TeJia -

Ha roJIOBe, IIJiaBHHKax, B rJIOT-

Ke, IIHID;eBo,n;e, pOTOBOH IIOJIOCTH. 8TH peu;errTOphl Ba:IB:Hhl ,/J;JI.H IIOHCKa H pacII03HaBaHll.H IIHID;H. 8Kcrrep11MeHTaJibHO ycTaHOBJieHO, qTo phl6bI MOryT pa3JIHqaTb ropbKOe, COJieHoe, KHCJIOe, CJia)J;KOe.

Opzanbt o6onRHUR

rrpe,n;cTaBJieHbI rrapHhlMH cJierrhlMH o6oH.HTeJibHhlMH

Meru:KaMn, Ha CIIHHHOH CTOpOHe rOJIOBbl, BbICTJiaHHbIMH o60H.HTeJibHhlM aIIHTeJIHeM. IIapHbie Hapy:IB:Hbie Hoa,n;pn coe,n;nH.HIOT rroJIOCTll aTMX MellIKOB c BHelllHeii cpe,n;oii. BHyTpeHHHX Hoa,n;peii (xoaH) HeT. OpraH o6oH.HHll.H rroaBOJI.HeT pbI6aM HaXO,/J;HTb rrapTHepa B rrep110,n; pa3MHOJKeHH.H, qyBCTBOBaTb rrpn6JIMJKeHMe
,D;06hlqll HJIH orraCHOCTb.

Opzanbt 3penuR (rJiaaa) pacrroJiaraIOTC.H rro 6o:KaM roJIOBbI.

3peH11e MOHOKY-

JI.HpHoe. PoroB11u;a nJioc:Ka.H, xpycTaJinK 11MeeT rnapoBM,n;HyIO (c¢>ep11qecKyIO)

321

HaiJ1aiacc P""6bl

<}>oPMY· PhI6hr MoryT BII,ll.eTh TOJihKO Ha 6JIH3KOM paccTO.HHIIH. KpyroBhie MhIIIIn;hl apaqKa II xpycTaJIIIKa OTCYTCTBYIOT. AKKOMo.rr.au;na (II3MeHeHIIe cpoKycHo-

ro

paCCTO.HHII.H) rrpOHCXO,ll.lfT aa cqeT rrepeMern;eHHH xpyCTaJIIIKa K rrepe.rr.Hett

KaMepe llJilf OT Hee c IIOMO:rn;hIO OTpOCTKa cocy.rr.ncTOH o6oJioqKif ( ceprrOBll,llHOro OTpOCTKa), KOTOpblH rrpnKperrJI.HeTC.H K aa,n;HeH CTeHKe xpyCTaJIIIKa. B CeTqaTKe co.rr.epmaTC.H IlaJIOqKn If KOJI6oqKif (u;BeTOBOe apeHIIe).

Opzatt cJtyxa u pa6tt06ecu.R

rrpe,n;cTaBJieH BHyTpeHHHM yxoM c TpeM.a rroJiy-

KPYiKHhIMII KaHaJiaMII (opraH paBHOBecna), co,n;epmarn;IIMH JKII,ll.KOCTh c B3BeroeHHhIMII TBep,ll.bIMll KpHCTaJIJiaMII. Pb16bI CIIOC06Hbl lf3,ll.aBaTb If BOCilpllHHMaTh pa3JIHqHhle 3BYKH B IIIHPOKOM ,n;IIarra30He. 3ByKOBa.H IIpOBO,lllfMOCTb TKaaeH TeJia pbI6hI 6JIH3Ka K

3BYKOilpOBO,llHMOCTlf BO,llbl, Il03TOMY 3ByKOBbie

BOJIHhl rrepe,n;aIOTC.H Herrocpe,n;cTBeHHO qepe3 TKaHH.
BoKoBaa JIHHH.H

(opzatt oc.R3attu.R) -

opraH, BocrrpttHHMaro:w;tt:B aarrpaBJie-

Htte, IIepeMern;eHHe BO,llbl, ee ,n;aBJieHtte H CHJIY IlOTOKa. :3TO rpyrrrra qyBCTBHTeJihHblX KJieTOK, rrorpymeHHbIX B KaHaJibl, rrpoxo,n;a:w;tte rro.rr. KOJKeH H OTKpbIBaIO:rn;HeC.H Ha IlOBepxHOCTb TeJia OTBepCTH.HMH.

Ilull-'te6apumeJtbtta.R cucme.Ma

HaqttHaeTCR poTOBhIM OTBepcTIIeM, orpa-

HHqeHHhIM qeJIIOCT.HMH c KOHHqecKIIMH ay6aMH (,n;JI.H aaxBaTa If y,n;epmaHH.H
rrtt:w;tt). HacToa:w;ero .H3b1Ka HeT. KphIIIIeH poTOBOH rroJIOCTH aBmrnTCR ocHoBaHtte qepeIIa (rrepBttqHoe TBep,n;oe He6o). I'JIOTKa, rrpoHtt3aHHaa ma6epHhIMH :w;eJI.HMH, rrepexo,n;IIT B KOPOTKHH IIH:rn;eBO,ll, a aaTeM B pacTRJKHMblH

me-

Jiy,n;oK. OT Hero OTXO,llHT TOHKa.H KHIIIKa, o6pa3yR xapaKTepHbIH H3I'H6, rroCJie qero KHIIIKa T.HHeTC.H rrp.HMO Haaa.rr. (TOJICTa.H KHIIIKa) H OTKpbrnaeTC.H
Hapymy caMOCTO.HTeJihHhIM aHaJihHhIM OTBepcTtteM.
TOHKOH KHIIIKII pacrrOJIOJKeHbl CJieIIbie BblpOCTbl -

B

HaqaJihHOH qacTH

IIHJIOpttqecKHe rrpH,n;aT-

KH, BbIIIOJIH.HIO:rn;He <l>YHKll;HIO yBeJittqeHH.H rrttrn;eBapHTeJibHOH IlOBepXHOCTH
KHIIIeqHHKa.

y

pb16 pa3BHTa rreqeHb, JKeJiqHblH rry3hlpb, B rreTJie KHIIIeqHH-

Ka pacIIoJiaraeTca rro,n;meJiy,n;oqHaa meJieaa B BH,n;e ,n;oJieK. OT rrepe,n;Hero oT,n;eJia KHIIIeqHHKa o6oco6JI.HeTC.H

nJta6ameJtbllblU ny3blpb

(BblpOCT CITHHHOH

CTeHKH rrttrn;eBo,n;a), HarroJIHeHHhIH cMecbIO raaoB. Ero cpyHKD;HH: rH,n;pocTaTttqecKaa, raaoo6MeHHaR (qacTttqHo), peaoHaHCHaR (yBeJIHqeHne ,n;ttarraaoHa BOCilpHHHMaeMhIX 3BYKOBblX BOJIH).

/(btxameJtblla.R cucme.Ma

rrpe,n;cTaBJieHa ma6paMH, KOTOpbie cocTo.HT H3

qeThlpex IIap ma6epHhIX ,n;yr. Ha ma6epHhIX ,n;yrax pacrroJiomeHhl ma6epHbie
TblqHHKH, BbIITOJIH.HIO:rn;He <l>YHKll;HIO u;e,n;HJibHOI'O arrIIapaTa, a TaKme ma6epHbie JieIIecTKH, cHa6meHHhie rycTo:H ceTbIO KaITHJIJIHpoB. Bo,n;a, rrorra,n;aro:w;a.a: B rJIOTKY qepe3 POT, OMhIBaeT ma6epHbie JieneCTKH, B KOTOpbIX rrpOHCXO,n;HT ra3006MeH (rrocTyrrJieHne KHCJIOpo,n;a B KpOBb H BbIBe,n;eHHe ll3 Hee yrJieKHCJIOI'O raaa).
AKT ,n;hIXaHHH y pb16 npoH3BO,llHTCH ,n;BHJKeHHeM ma6epHhIX KpbIIITeK: phl6a npttno,n;HttMaeT ma6epHyIO KPhIIIIKY, ma6epHaH nepenoHKa npIIJKHMaeTca K ma6epHoH :w;eJIH.

B

peayJihTaTe o6paayeTca npocTpaHCTBO c nom1meH-

BhlM ,n;aBJieHHeM, H BO,lla H3 pOTOrJIOTQqHoH ITOJIOCTH BCaCblBaeTCH B 60KOBy10 ma6epHyIO ITOJIOCTb. Ilptt orrycKaHHH ma6epHOH KpbIIITKH coa,n;aeTCH

322
tt36hITO'IHOe .n;aBJieHMC, H BOJ~a qepe3 Hapy:>KHhie :ma6epHbie OTBepCTMB: BbITaJIKMBaeTCB: Hapy:my.

y

pbr6 MoryT B03HMKaTh ITpHCITOC06JieHMB: ,ll;JIB: llCITOJih30BaHMB: aTMOccpep-

HOrO KHCJiopo.n;a: CTCHKH KMIIIe'IHMKa c KposeHOCHhIMll cocy.n;aMtt, BhIPOCTb!
.n;opcaJihHOM n seHTpaJihHOii: cTeHKM rJIOTKM, ITJiasaTeJihHhIM ny3Liph. OpraHhI
B03,ll;YIIIHOrO ,ll;hIXaHMB: o6ecITe'IMBaIOT Bhl:>KMBaHMe oco6eii: B CllJihHO 3arpB:3HeHHblX so.n;oeMax, r.n;e KOJIM'IeCTBO KllCJ'IOpo.n;a B so.n;e CHM:>KaeTCB:.

Kpoae1toc1w.fl

cucme.11ta

3aMKHyTaB:.

O.n;ttH

Kpyr

Kposoo6pa~emrn.

Ha 6pIOIIIHoii: cTopoHe pacIToJiaraeTCH .n;syxKaMepHoe cep.n;u;e, COCTOH~ee n 3
ITpe.n;cep.n;ttH tt meJiy.n;O'IKa, co.n;ep:ma~ee seH03HYIO KpOBh. BeH03HaB: KpOBh n3
meJiy,ll;O'!Ka cep.n;u;a ITOCTyITaeT B 6pIOIIIHYIO aopTy ll .n:a-'Iee ITO 'IeTh!peM ITPMHO-

CB:~llM :ma6epHhIM apTepttB:M B :ma6phr. B :ma6epHhIX JieITeCT.Kax .KpOBh Hacbr~aeTCH .KMCJIOpO,ll;OM ll CTaHOBMTC.H apTepttaJihHOll. 3aTeM BhIHOCH~Me ma6epHble apTepttll BITa.n;aIOT B .KOpHM aopTbl Ii .n;aJiee B CITMHHYIO aopTy'
KpOBh K roJioBe, MYC.KyJiaType n

Hecy~yIO

ITO BCeM BHyTpeHHHM opraHaM. BeH03HaH

.KpOBh B03Bpa~aeTCB: B cep.n;u;e ITO HeITapHOll XBOCTOBOll BeHe ll no ITapHbIM
3a.n;HttM M ITepe.n;HttM Kap.n;ttaJihHhIM seHaM. IIpasaB: 3a.n;H.H.H Kap.n;HaJihHa.H seHa ITPOXO,ll;MT qepe3 ITO'I.KM, He pa3BeTBJIHB:Ch, a JieBa.H .n;aeT ceTh KaITMJIJIB:pOB.
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324

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326

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KJtacc KocmHbte pbt6bt:
1. Kucmenepbte pbtObl.
Oco6eHHOCTil:

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coxpaHIIBWIIe

xop.n;y; napHbie IIJiaBHIIKH Il.MeIOT IIO,ll;BilJKHOe coqJieHeHile c IIOHCaMil.
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rJIOTKa IlMeeT rrapHbIM BblpOCT -

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HOro ,IJ;bDWHIIH.

2. /{ 60Jl1WiJbtUlllW,Ue pbt6bt.
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HaJIII'IIIe ma6epHoro II JierO'IHOro ,ll;bIXaHIIH ( O,ll;IIH liJIII .n;sa IIY3blpH, OTKpbIBaIOlll,neca Ha 6pIOWHYIO CTopotty nmn;eBo.n;a); HaMeqaeTca pa3.n;eJieHne rrpe.n;cep.n;IIa II o6pa3oBaHIIe Jiero'!Horo Kpyra Kposoo6paw.eHIIH.

3. KocmHo-xp.HW,e6bte pbt6bt.
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CHa6meH ma6epHblMII KpblllIKaMII; BCTpe'IaIOTCR npOXO,ll;Hbie II npecHOBO,ll;Hbie pbI6bI.

4. Kocmucmbte pbt6bt.
Oco6eHHOCTII: CaMa.H MHOrO'IIICJieHHaH rpynna pb16; IIMeIOT KOCTHbIH
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Knacc 3eMHOBOAHble (aMcl>1116111111)
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3eMH060iJHbte -

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Knacc 3eAfn06oanbte ( aM<Pu6uu)

327

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200

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;I<:HBOTHbie aKT11BHhl B yTpeHHlfe H BeqepHHe qachI, KOr,n;a BJiaJKHOCTh B03.D;yxa
11 TeMnepaTypa ,n;oBoJihHO BhICOKH. IIpe,n;cTaBHTeJih oTpH,n;a BecxBOCThie mpa6R1WR Jt.Hiyw1ca. 0611TaeT BO BJiaiKHhIX MeCTax yMepeHHOll 30Hbl.

Cmpoettue mpa6RHOU JutiyUJKU. TeJio cocT011T H3 roJIOBhI, TyJIOB11m;a 11 napHhIX KOHeqHQCTeii. fOJIOBa IIIHPOKaH, ynJiom;eHHa.H, c 60JibIIIOll pOTOBOll m;eJiblO H BbIITYKJibIMH rJia3aMH, C3a,D;H KOTOpbIX 11MeIOTCH ,D;Be OKpyrJibie 6apa6aHHble nepenoHK11, 3aKphrna10m;11e caapym11 noJIOCTh cpe,n;Hero yxa (pn:c.

129).

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H o6JierqaeT ra3006MeH (KOiKHOe )J,hIXaHHe).

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roJIOBhI (qeperra) H CKeJieTa rrapHhIX KOHeqaocTeii.

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3BOHKH npoqHo coqJleHeHbl .n;pyr c .n;pyroM. CoCTOHT 113 qeTblpex OT,D;eJioB:

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HYIO KOCTOqKy -

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3aTblJIQqHhlMH KOCTHM11.

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(m1eqeBa.H KOCTh), rrpe.n;mieqh.H (cpocumeca JiyqeBa.H H JIOKTeBaH KOCTH) H KHCTH (KOCTH 3arracTh.H, II.HCTH 11 q>aJiaHrH rraJIL:u;eB).

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328

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8
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ycJiommi:eTca H crren;HaJIH3HpyeTCH B CBH3H c rrepe-

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rro,n;HHMaIOT H orryc1rn10T HHiRHIOIO qeJIIOCTh. MbIIIIIJ;hI ,n;Ha poToBoii: rroJIOCTH
yqacTBYIOT B rrpo:u;ecce JieroqHoro ,n;hIXaHHa. TyJIOBHIII;HaH MycKyJiaTypa
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HHHMH;

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K.nacc 3eMIW6oiJnbte (aMt/Ju6uu)

npoaoJLZOBambtit

329

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qepeIIHO-M03rOBblX Hep BOB.
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pa3BHTa CHMTiaTnqecKaH HepBHaH CHCTeMa, npep;cTaBJieHHa.H )J;BYMH HepBHhlMH CTBOJI8MH, pacnOJIOiReHHhlMH no 6oKaM II03BOHoqHHKa.
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CKJia,n;OK u BhICTJiaH rrpocThIM arrMTeJIHeM. OpraH o6oHHHHH cpyHKUHOH11pyeT
TOJibKO B B03,ll;YIIIHOH cpe,n;e, a B Bo,n;e HapyiRHhie H03,n;pu 3aKphIBaIOTCH KJianaHaMH.

B

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noJIOCTH.

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(rJia3a) uMeIOT cTpoeHne, xapaKTepHoe ,n;JIH Ha3eM-

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(B BO,n;e yrnrnruaeTCH),

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-

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130).

330

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KOTOpaa JIHIIIeHa ma6epHbIX rn;eJie:H
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OCHOBaHHe

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XBOCT' KOHe'IHOCTH H HX no.aca OTcyTCTBYIOT; py,n;ttMeHThI opraHOB
3peHHH ll cnyxa; HMCCTCH KOilyJIHTHBJihIH opraH (BbIIlH'IHBaHHC
cTeHKH KJIOaKH); ae,n;yT no,n;3eMHhIH o6pa3 iKH3HH

I HacTomu;a.a qeps.ara,

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i
i

OT aa,n;HeH 'IacTH TeJia KpOBh co6HpaeTC.H B HenapHy10 aa,n;H1010 noJiy10 BeHy,
npoxo,n;.H qepe3 BOPOTHYIO CHCTeMy ne'IeHH H no'IeK, Bna,n;aeT B BeH03HY10 naayxy H npaBoe npe,n;cep,n;He. OT nepe,n;Heif qacTH TeJia BeHoaHa.H KPOBh co6HpaeTC.H B npaBy10 H JieBy10 nepe,n;HHe noJihle BeHhl, B BeH03HYID naayxy H npaBoe
npe,n;cep,n;He.
M aJtbtu ( Jtezo•U-tbtu} Kpyz KpoBoo6paw,ettu.H Ha'IHHaeTc.H KOmHo-JieroqHhIMII
apTepH.HMH, Hecy~HMH BeH03HY10 KPOBb K opraHaM ,llhIXaHH.H, r,n;e npOHCXO,llHT raaoo6MeH. OT JierKHX o6ora~eHHa.H KHcJiopo,n;oM KPOBh no napHhIM Jiero'IHhIM BeHaM nona,n;aeT B JieBoe npe,n;cep,n;He.
BOJtblUOU Kpyz 1CpOB006paw,eHU.H Ha'IHHaeTC.H ,n;yraMH aopTbl H COHHbIMH apTepH.HMH, KOTOphle BeTB.HTC.H B opraHax H TKaH.Hx. BeH03Ha.H KPOBh no napHhIM
nepe,n;HHM nOJihIM BeHaM H HenapHoif aa,n;Heif nona,n;aeT B npaBoe npe,n;cep,n;He.
B nepe,n;HHe IIOJible BeHhl Bna,n;aIOT H KOmHhie BeHbl COOTBeTCTBy10~eif CTOpOHhl, KOTOpbie HecyT apTepHaJihHYlO KPOBb.
BbtiJeJtumeJtbHa.H cucme.Ma npe,11;cTaBJieHa napHhIMH npo,n;oJiroBaThIMII
TYJIOBH~HhIMH (Me3oHe<t>poc, nepBH'IHhIMH) nO'IKaMH, Jiema~HMH B IIOJIOCTM TeJia no 6oKaM OT n03BOHO'IHHKa; MO'IeTO'IHHKaMH H MO'IeBbIM ny3hipeM.
B TYJIOBH~HhIX noqKax He nponcxo,D;HT o6paTHoro BcachrnaHH.H BO,D;bI, noaTOMy MO'IeBoif nyahlpb .HBJI.HeTC.H peaepByapoM BO,llbl, B KOTOpOM npoHCXO,llHT ee o6paTHOe BCaChIBaHHe. IlpH HaIIOJIHeHHH IlY3blp.H MO'Ia Bb16pacb1Ba-

KJ1.acc llpecMbuca10utuecR (penmuJtuu)

333

eTC.H aapymy qepe3 OTBepCTlle KJIOaKH. ,r:(onOJIHHTeJibHbIMH oprattaMH BbI.n;eJieHH.H .HBJI.HIOTC.H Koman JierKne. OcHOBHhIM KOHe'lHhIM rrpo.n;yKTOM o6Meua
jlBJI.HeTC.H MO'leBHHa. BoJibllia.H IIOTep.H BO}J;bl opraHH3MOM qepe3 opraHbl BbIAeJieHll.H ll IIOBepxHOCTb J{Ojl{fi He II03BOJI.HeT JI.HI'YIIIKe H8}];0JII'O IIOKH.n;aTb
BJiaIBHbie MeCTa.

IloJLoBaJl cucmeMa. Paa.n;eJihHOIIOJihie. IloJIOBhre meJie3bI rrapHhre.

Y

caM-

11;os ceMeHHHKH He HMeroT caMOCTO.HTeJihHhIX BhIBO,D;HhIX rryTeli. CeM.HBhIHoc.am;ne KaHaJihUbI npOXO,D;.HT qepe3 rrepe,D;HllH OT,D;eJI IIO'IKH H Brra.n;aroT B MO'leTO'lHllKll, KOTOpbie O,D;HOBpeMeHHO CJiymaT ll CeM.HIIpOBO}J;8Mll. Ilepe.n; Brra.n;e1meM
B KJIOaKy o6paayeTC.H pacnrnpeHne -

ceMeHHOH rry3blpeK, B KOTOpOM BpeMeH-

HO peaepsnpyeTC.H ceM.H. Ha.n; ceMeHHHKaMn JiemaT mnpoBhie TeJia, cJiymaru;ne
AJI.H rrnTaHH.H ceMeHHHKOB n paaBnsaroru;nxc.H B HHX cnepMaToaoH.n;os. BeJIH'lHaa mnpOBbIX TeJI MeH.HeTC.H no ce30H8M. OceHhlO OHH BeJIHKH; BeCHOH, BO BpeMJI HHTeHCllBHOI'O cnepMaToreHeaa, seru;eCTBO HX ::rneprH'lHO pacxo.n;yeTC.H,
n paaMephI mHpOBhIX TeJI peaKo coKparu;aIOTC.H. KonyJI.HTHBHhIX opraHOB HeT •
.flnqHHKll caMOK BeCHOH yBeJinqeHhl ll 3aIIOJJH.HIOT BCIO 6pIOIIIHYIO IlOJJOCTb.
OHH co.n;epmaT apeJible ali::a;eKJieTKH (nKpHHKH). l.Jepea paaphIB TOHKoli CTeHKH
HllqHnKa HKPHHKH Bhma.n;aroT B IlOJIOCTb Te.Jia ll qepea BOpOHKY n:ii:ueso.n;a nona.n;aroT B ,D;JIHHHhIH H3BHTOH ali::u;eso.n;, KOTOpbIM OTKPhIBaeTC.H B KJIOaKy. Ornrn,D;OTBopeane HapymHoe, npOHCXO,D;llT B Bo,n;e.

Paaeumue (Henp.HMoe) nponcxo,n;nT c MeTaMopcpoaoM. l.Jepea He,n;e.Jiro noZOJl08aCmUKU.

c.Jie OITJIO,D;OTBOpeHHH ll3 HKPHHKH BbIJIYITJIHIOTC.H Jill'lllHKH -

0HH Be,n;yT BO,D;HbIH o6pa3 jl{ll3HH, HMelOT HapyJKHbie ma6pbl, .n;syxKaMepuoe
cep.n;:u;e, O,D;HH Kpyr KPOBoo6paru;eHH.H, opraHhl 60KOBOH JillHHH, napHhie KOHelJHOCTH oTcyTCTBYIOT. HeKOTOphre BH,D;hI aeMHOBO,D;HhIX npoHBJI.HIOT aa6oTy
0 IlOTOMCTBe.

Knacc npeCMblK310l14MeCS1 (penn'lnMM)
OcnoBttbte apoMop<jJoabt:

1.

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2.

IlporpeccnBHhre npeo6paaosaHHH CKe.JieTa n

aM6pnoHa B Ha3eMHhIX yc.JIOBH.HX.
cpopMnpoBaHne rpy,n;Holi

KJieTKll.

3.
4.

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5.

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CKy.JiaTyphr).
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6.
7.

PaaBHTHe Heno.JIHOH Memme.Jiy.n;o'IKOBOH neperopo.n;KH cep.n;ua.
<l>opMnpoBaHne T830BOH (BTOpll'IHOH) ITO'IKH c o6paTHbIM BCaChIBaHHeM
seru;ecTB.

lipeC.Mbt}(UIOUfUeCJl -

nepBhie Ha3eMHhie Il03BOHO'IHble, yTpaTHBIIIHe TeCHYIO

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HYIO

CTnxnro. CospeMeHHhie npecMhrnaroru;necn BKJIIO'laIOT 'leThrpe oTp.a:.n;a:

334

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KJllOBO?OJtOBbte, 'lepenaxu, KpoKoiJuJtbt u 'lewyu'lambte (3."deu, Rw,epuu,bt, xa;~wJte
Ollbt). B HaCTOflru;ee BpeMfl ll3BeCTHO OKOJIO 7 ThIC. Bll,[\OB.
Ilpe.n;cTaBHTe.nh ompRiJa 'lewyu'lambte, noiJompRiJa Rw,epuu,bt npbtm1CaJ1
Rw,epuu,a.
Cmpoenue npbtmKou Rw,epuu,bt. TeJio y,n;JIHHeHHoiI cpopMhr, cocTOMT M3 roJIOBbI, rneM, TYJIOBMIIJ;a, XBOCTa lI IlflTMIIaJihIX KOHe'IHOCTeH.

JloKpoe.

Koma cyxafl, rroKphrTa poroBhIMM qernyf!MM, JIMrneHa meJie3 (3a-

IIJ;MTa OT IIOTepM BO,[\hI). Ha naJihu;ax lIMelOTC.H porOBhie o6pa30BaHM.H -

KOPTM.

PocT conpoBom.n;aeTcH JIMHhKOH (rrepMo,n;MqecKa.H cMeHa KomHhIX rroKpOBOB).
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noJIHOCThIO OKOCTeHeBaeT. CocTOMT M3 oceBoro CKeJreTa

(Il03BOHO'IHlIK), CKeJieTa rOJIOBhI (qeperr) lI CKeJieTa KOHe'IHOCTeH. Ilo3BOHO'IHlIK .n;eJIMTC.H Ha 'IeThrpe OT,[\eJia:

1.

IIIeiIHhIH -

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ne'IMBaeTc.H .LIBYMfl rrepBhIMII rneHHhIMlI Il03BOHKaMM -

aTJiaHTOM lI oce-

BhIM (arrMCTpocpeeM), 06pa3yroru;MMI1 cycTaB. IIepBhIH IIMeeT BM.LI KOJihu;a
lI CO'IJieH.HeTC.H c HerrapHhIM MhIIIJ;eJIKOM qepena, BTOpoil: rrpe.n;cTaB.JJ.HeT
co6oii: 3y60BM.D;HhIH OTpOCTOK, BXO.D;.HIIJ;lIH B KOJihIJ;O nepBoro II03BOHKa
ll o6ecrre'IMBaIOru;IIii: IIOBOpOT I'OJIOBhI.

2.

fpy,n;Ho-rro.HCHH'IHhIH -

cocTOMT H3

22

II03BOHKOB. BKJIIO'IaeT rpy,n;-

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II.HTh -

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rpy,n;IIHe,

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3.

KpecTu;OBhIM -

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IIO.D;B3,[\0illHhie KOCTlI Ta3a.

4.

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B

cJiyqae orracHOCTH Hru;ep.Hu;a crracaeTC.fl OT r.H6eJIH rryTeM oT6pachrnaHMH

XBOCTa (ayTOTOMll.H -

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qeperr IIO'IT.ll IIOJIHOCThIO OKOCTeHeBaeT .ll COCTO.llT .ll3 60JihlliOI'O KOJI.ll'IeCTBa
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a TaKme 60Jihllia.fl .ll MaJia.H 6epu;OBhie He cpacTalOTC.fl; CTOIIa II K.llCTh Kopoqe,
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MbtUle'IHaJl cucmeMa

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(.D;B.llmeH.He qeJIIOCTe:ii:), rpyIIIIhI Mhlllill; rneII ( o6ecneqIIBalOT .D;B.ll)KeHII.H rOJIO-

K//,aCC

npeCMblKlllOUfUeCH ( penmU//,UU)

335

1

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Bbl), Mblllll.l;bl KOHeqHQCTeiL XopolllO pa3BHTbl Mempe6epHble Mbllllll;hl, KOTOphJe yqacTBYIOT B MexaHH3Me ,ll;bIXaHH.H.

HepBnaH cucmeMa

6oJiee paaBttTa, qeM y aeMHOBO,LJ;HhIX. foJIOBHOM Moar

3eMHOBO,LJ;HbIX COCTOHT H3 11.HTH OT,LJ;eJIOB.

II epeiJnuit

M03r 11pe,n;cTaBJieH 60Jib-

illHMH IIOJiyIIIapM.HMH, paa.n;eJieHHblMH Ha 11paByIO H JieByIO ,LJ;OJIH, 60JiblllHX
pa3MepOB aa cqeT CKOIIJieHH.H IIOJIOCaTbIX TeJI.

B

M03rOBOM C.TJOe pa3JIHqaIOT

11epBHqHbIM CBO,LJ; (apXMIIaJIJIHYM), 3aHHMaIOillHM 60JiblllYIO qaCTb KpbllllH
IIOJiyIIIapttii:, a TaKme 3aqaTOK HOBOM KOpbl (Heo11a.'1JIHyMa). IlporpeCCHBHOe
pa3BHTHe

11epe.n;Hero

<PopM 110Be,n;eHM.H.
a caa.n;tt n

M03ra

06ec11eqnBaeT

IIpoMeJ1Cymo'lHbtU

cpe.n;HttM MoaroM.

B

B03HHKHOBeHHe

CJIO:IKHhlX

Moar c11epe.n;tt 11pttKpb1T 11epe.n;HttM,

KphIIIIe xopoIIIo pa3BHTbI TeMeHHoii: opraH

3IItt<l>M3. TeMeHHOM opraH CBOHM cTpoeHtteM Ha110MMHaeT r.11aa.

B

HeM

paaJinqa10T y11JIOTHeHHhIM 11poapaqHb1ii: 11epe.n;Httii: OT,n;eJI (110.n;o6HhIM xpycTaJittKy) Ii 6oKaJioo6pa3HYIO aa,n;HIOIO qaCTh, COCTO.HillYIO H3 qyBCTBYIOillHX
H IIRrMeHTHbIX KJieTOK. OH a<Pc):leKTRBHO BOCIIPHHHMaeT CBeTOBbie paa.n;pam.emrn n cJiymnT peu;e11TopoM ceaoHHhIX H3MeHeHMM.

CpeiJHuii Moar yBeJIHM oaJ/Ce'lOIC cttJihHO

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Pa3BHT, .n;BttmeHtt.H rrpeCMhIKaIOillHXC.H 6oJiee HHTeHCHBHhie H paaHoo6paaHhle.

IlpodOJlZ06ambtU

M03r o6paayeT H3rH6 B BepTHKaJibHOH IIJIOCKOCTH,

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TeJibHhlMH MelllKaMH, paa.n;eJieHHblMH Ha o60H.HTeJibHbIM (BepXHHM) H ,!l;bIXaTeJibHhlM (Httmanii:) oT.n;eJihI. Ilo.HBJI.HeTC.H o6oH.HTeJibHa.H paKOBHHa. I1MeIOTc.H o6oH.HTeJibHbie XO)J;bl c 11apHbIMH H03,11;p.HMH, OTKpb1BaIOI11HMHC.H Hapymy,

336

300//,0ZUR

a B poTOBYIO IlOJIOCTb -

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aH 3a C'!eT pa3BHTH.H BTOpll'IHOro KOCTHoro He6a CMeru;aIOTCJI Ha33l\, 6JIJDKe
K rJIOTKe, 'ITO o6ecIIC'IHBaeT CB060,D,HOe ,ll;bIXaHile BO BpeMH Haxom,n;eHHH BO
PTY IIHIIJ;H. 06pa3yeTCH HOCOrJIOTO'!Hblll

xo,n;. ,lJ;JI.H BOCilpHHTllH 3anaXOB Illl-

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H3BHTaR ll CJieno 3aKaH'IHBaroru;a.HC.H IlOJIOCTb,

oTxo,n;am;aa BBepx OT KpbIIIIH poTOBOH IIOJIOCTtt. Opzan apeHUJl. I'Jia3a 3aru;tt u~eHhI IIO,ll;BH:>KHbIMll HapyJl\HbIMH BeKaMH H MuraTeJibHOH neperrOHKOH .
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HO cnoco6HhI K Bparu;aTeJihHbIM ,n;Btt:>KeHttaM.

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cnyxosoro rrpoxo,D,a. Ha rpaattue c BHyTpeHHHM yxoM B 3a,n;ae:ii cTeHKe noaBJI.HeTcH, KpOMe OB8JibHOI'O,

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yBeJia'lnBaeTC.H B ,n;JittHY H Ha'IMHaeT H3rM6aThC.H KaHan YJIMTKH, r,n;e cpopMHpyroTc.a qacTH BocnpttHHMaroru;ero annapaTa. CTpoeHHe cpe,n;Hero yxa He
H3MeH.aeTc.H.

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llu14e6apume.!lb/-laJl cucme.Ma Ha'IHHaeTcH pOTOBhIM oTsepcTHeM, orpaHH'leHHblM 'leJIIOCT5IMH c KOHR'leCKHMR, O,ll;HHaKOBhIMH, npO''IHO npnpacTa-

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meny,n;oK n KHIUe'IHHK. B ,n;BeHa,n;n;aTMnepcTHYJO KHIUKY OTKPhlBaIOTCH rrpoTOKH rre'IeHH If no,n;meny,u;o'IHOH :>KeJie3bl. Ha rpaHHn;e Mem,u;y TOHKHM H TOJICTbIM OT,u;enaMH KHIUe'IHHKa pacnonomeH 3a'IaTOK cnerroli KHillKH. 3aKaH'IHBaeTCH

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337

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TI pe;~cnt Bl!Temi

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(2700-3500

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338

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340

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CKeJLem c60600HbtX ICOHe'lnoemeu:

rrepe,z:i;tttte KoHeqttoCTH TpattccpopMttpo-

BaHhI B KphIJlh.ll H COCTOHT H3 TIJieqa, npe,n;nJie'IhSI H KHCTH. KocTH aarraCTha
H Il51CTH CJIHBaIOTCa, o6pa3yH npm:KKy.

ffa

IIaJihll;eB coxpaHaIOTCSI TOJihKO

3
2

3

\

\

-·-----·- 6
P111c. 132. T11mb1 nepbes: 1 - KOHTYPHOe nepo; 2 no6o•moe 6oKosoe nepo myxapR; 3 - nyxosoe nepo;
4 - H11rnm.1AHOe nepo; 5 - ll.!ernHKa; 6 - co6cTseHHO nyx

P111c. 133. CTpoeH11e nepa: 1 - onaxa110;
2 - CTBOJl; 3 - nyxosaR YaCTb onaxana;
4 - sepxHee ornepcrne; 5 - 041'1H; 6 Hl'1JKHee OTBepCTHe

342

Tpll -

300.!WZUJt

BTOpOtt, TpeTMH M 1IeTBepTblll, rrpu 3TOM TpeTHH rraJreu HMeeT /(Be <:J.:>a-

JiaHrH, a BTOpOtt H qeTBepTblH -

no 0,L{HOH. 3a,L{HHe KOHelJHOCTH CJiymaT /(J1.ll

rrepe,D;BmKeHH.H ITO 3eMJie H COCTO.HT H3 6e,D;pa, rOJ1eHM (Ma.1rna 6epn;oBa.H KOCTh
py,D;ttMeHTapHa H rrpttpacTaeT K 60JihIIIOH 6epn;osoi1) H n;eBKH (cpocrrrneca KOCTH rrpe,D;IIJIIOCHhI tt IIJIIOCHhI). IIaJihII;eB l.JeThipe, Tptt H3 HHX HarrpasJieHhI Brrepe,D;, O,D;HH -

Haaa,D;.

Mb1We1J,Ha.H cucmeMa

CHJihHO paaBttTa H ,D;HcjJcjJepeHII;HpoBaHa. HaH6oJiee

pa3BHThl MhllllII;bl rpy,D;H, o6ecrrel.JHBaIOIIJ;Jie ,L{BHJKeHHe KphIJibeB. IlapHhie
6oJihlllHe rpy,L{Hhie, rrpMKpe11JieHHhie K rpy,D;MHe M ee KHJIIO, CJiymaT ,L{Jl.H
orrycKaHH5l

KPhlJia,

IIO,L{KJIIO'IH'IHhie MhlIIIII;hI

-

,L{JI.H

IIO,ll;'beMa KpbIJia.

,D;JIHHHhie MhilllJJ;hI llleH o6ecrrequBaIOT CJIOlliHhie ,D;BMJKeHna roJIOBhI. CHJihHO pa3BMThJe MhlllllJhl 3a,L{HHX KOHel.JHOCTeii: rrpe.n;Ha3Ha'IeHbl ,ll;Jl.H rrepe,D;BMJK€HH.H IIO 3€MJie.

Hep6HG.fl cucmeMa

COCTOMT M3 CIIMHHOro H I'OJIOBHOro M03ra c OTXO,L{.HIIJMMH

OT HUX H€pBaMM. roJIOBHOH M03r rrpe,TJ;CTaBJI€H CJie,n;yIOIIJHMM OT,ll;eJiaMu:

/

/ 17

3

5
6

7

13

21

P111c. 134.
3BOHKH;

4-

CKeneT rony6s:1 (cxeMa):
n11e40;

5-

1-

2- HH)l(HHH 4e11t0cTb; 3- wei1Hble no7 - BOPOHbs:l KOCTb; 8 - rpyAHHa; g - KHJlb;

sepxHs:is:i Yent0CTb;

fPYAHb18 no3BOHKH;

6-

KJll04HLICI;

10 - roneHb; 11 - u,esKa; 12- 6eApo; 13- raa; 14 - KOn4HKOBas:i KOCTb; 15 - XBOCTOBble n03BOHKH;
16 - pe6pa; 17 - npeAnlle4be; 18- KHCTb; 19 - nonaTKa; 20 - 4epen; 21 - nanbu,b1 crnnb1

Knacc IImu!fbt

1. II epPiJuuu M03Z

IIMeeT pa3BIIThie 6oJihurne uoJiywapIIH. l l x noBEpx-

HOCTh co;::i;epJKIIT CKOnJieHIIe HepBHhIX KJieTOK, o6pa3yIOll(IIX apxIIraJIJIIIyM. O;::i;ttaKo 60JihIIIaH qacTh nepe;::i;ttero M03ra o6pa3oBaHa noJioci:rhiMII TeJiaMII. Bopo3;::i; II II3BIIJIMH HeT. IloJiyIIIapIIH peryJrnpyIOT CJI>mHhre cpopMbl noBe.n;eHII.H nTIIll, 3pIITeJibHhie .ll;OJIM pa3BHThl CIIJIIHO,
a o60H.HTe"1hHbie .ll;OJIII -

2. IJpoMPJ/Cymo'lllbtu M03t

CJia6o.
pa3BIIT cJia6o. Ha Bepxtteii: cTopotte Haxo.n;ITc.H

anIIcpII3, a Ha .n;tte, no3a.n;u nepeKpecTa 3pttTeJihHhIX ttepBoB, -

KPYITihIH

nmocpII3.

3. Cpeottuu M03t IIMeeT xopoIIIo
4. MoaJ/Ce'lOK xopomo pa3BIIT B

pa3BIIThie 3pIITeJihHhie 6yrpL1.
CBH3H c KOOp,D;IIHa11IIeii: )J;BIIJKeHHH II }aB-

HOBeCIIeM BO BpeMH noJieTa. CocTottT. II3 cpe.n;tteii: .n;oJIII -

qepBHma

II .n;Byx 6oKOBhIX BhICTynoB. C3a;::i;M npttKpwBaeT cpe.n;HMH M03r M qicTh
npo.n;oJiroBaToro M03ra.

5. IIpoiJoJtioBambtil M0.1t

nepexo.n;MT B cnMHHOH Mo3r.

OT roJIOBHoro M03ra OTXO.ll;MT

12

nap qepentto-M03rOBhIX ttepBoB.

CnuHHOU M03t MMeeT yToJirn;ettIIe B nJieqeBoii: M nO.HCHttqttoH: 06.JiaCT.Hx,r.n;e
OT Hero OTXO)J;.HT HepBbl K nepe,D;HHM II 3a.ll;HHM KOHeqHOCT.HM, o6pa3y.H nJI•qeBOe H Ta30BOe cn.JieTeHIIe.

Opiattbt 'lyBcmB. Be.n;yrn;IIMM .HBJI.HIOTC.H 3peHMe II cJiyx. Opiatt BK yea - IKYOpian o6oHJlHUJl pa3BIIT cJia6o. Opiatt apeHUJl o<eHh

COBhie JIYKOBMllhI B rJIOTKe.

xopoIIIo pa3BMT. f.Jia3Hhie H6JIOKII KpyrrHhie, pacnoJiaraIOTC.H no 6oKaM romBhI.
IloJie 3peHMH Kam.n;oro r Jia3a cocTaB.JI.HeT
HM.H -

30-50°.

150°,

a rroJie 6ttHOKY.JI.Hpttoro :pe-

OcTpoTa 3peHIIH oqeHh BhICOKa.H. lfao6pameHMe Ha ceTqfl'Ke

IlOJiyqaeTCH 60.JihIIIIIX pa3MepoB, qTo Il03BOJI.HeT pa3JIMqaTh .n;eTa.JIH o6'beITa.
CeTqaTKa MMeeT BhICOKYIO rrJIOTHOCTh cpoTope11errTopoB, ITTIIllhI pa3JIMqaroT ~Be
Ta H OTTeHKH. AKKOMO,D;a1III.H (,D;BOHHa.H) ocyrn;eCTBJI.HeTCH 3a cqeT H3MeHem.H
cpopMbl xpycTa.JIMKa (rro.n; .n;eii:CTBHeM peCHHqHOH MhlIIIllbl) II 0,ll;HOBpeMeHihlM
rrepe:vrern;eHIIeM OTHOCHTeJihHO ceTqaTKH. B 06.JiaCTII CJienoro Il.HTHa MMeeTC5COCY.ll;IICTOe o6pa30BaHMe -

rpe6eHh, KOTOphIH .HBJIHeTC.H HCToqHMKOM ITMTaT•.Jlh-

HhIX Bern;ecTB M KMCJiopo.n;a. MexaHIIqecKa.H npoqHOCTh rJia3 o6ecrreqMBarrc.H
YTOJill(eHIIeM CK.JiephI H IlO.HBJieHIIeM B HeH KOCTHhIX nJiaCTHHOK. BeKH xop•IIIO
paaBMThI. llMeeTC.H MH:raTeJibHa.H rreperroHKa (TpeThe BeKo).

OpiaH cJtyxa iaa-

BMT xopoIIIo. CocTOIIT II3 BHyTpeHHero M cpe.n;Hero yxa. Bo BHyTpeHHeM yxEYk
JlHH.HeTC.H BhIPOCT Kpynrnro MeIIIoqKa H yBeJIJiqIIBaeTC.H qHCJIO ceHCOpHblX meTOK. B cpe.n;HeM yxe B03pacTaIOT paaMepw 6apa6aHHOii: rroJIOCTII, yc.JIOJKH.HITC.H
cpopMa O)J;HOH CJIYXOBOH KOCTOqKII -

CTpeMeqKa, qTO yBeJmqMBaeT ee ITO)J;BIJK-

HOCTh. YBeJIMqIIBaIOTC.H pa3MephI 6apa6aHHOH rreperrOHKM. EBcTaxMeBhl Tpr6w
OTKphIBaIOTCH B r.JIOTKY O.ll;HHM o6rn;IIM OTBepCTHeM. Ilo.HBJI.HlOTC.H 3aqaTKHHapyarnoro yxa, H MMeeTC.H CJIYXOBOH rrpoxo.n;, aaKaHqMBaIOll(MHC.H 6apa6aHIOH
neperroHKOH.

Opiattbt ICOJICHOZO OCJl3UHUJl -

CKOIIJieHMe qyBcTBMTeJihHhIX me-

TOK, pearMpy10rn;IIx Ha II3MeHeHMe rro.JiomeHM.H rrepheB.

Tep.MoiJemeKmopb1 pe-

rMcTpMpyIOT H3MeHeHII.H TeMrrepaTyphl Te.Jia, COCTO.HT II3 CKOIIJieHH.H qyBC'BHT€JibHhIX KJieTOK, OIIJieTeHHhIX HepBHhIMH OKOHqaHH.HMII.

344

.100/WlUJl

IJuw,NwpumeJLb1-taJl cue me.Ma CBH3ana c oco6eHHOCTHMH npo.u;eccoB JKH3HeJJ.CHTeJibHOCTH. Tern10KpOBHOCTh H Bb1co1rnH no,n;BHJKHOCTh rrTHIJ. Tpe6yeT 3HaqnTeJihHhIX KOJIH'IeCTB TIHIIl.Il I1 ee 6bICTporo ycBOeHHH. IlHIIJ,a B JKeJiy,n;oquoKHillel.JHOM TpaKTe 6hICTpO rrepeBapHBaeTCH 3a Cl.JeT aKTHBHOCTH n:mr~eBapH
TeJihHbIX cpepMCHTOB lI ysem1qeJIHH BCaChIBaIOIIJ,e:H IIOBepXHOCTlf KHIUCl.JHHKa.
PorOBhre Kplrn qe,110cTe1i: 06pa3y10T 1rn10B, KOTOphr:H cJry.iKHT .n;.1a 33XBaTa rrnIIJ,H. 3y6on HeT. 5l3h!K MyCKyJIHCTbIH, KOHHl.JeCKHH, HMeeT oporoBeBalOIIJ,Re
IIIHIIHKH JJ,JIH y.n;epmaHHH TI11IIJ,H. IlpoTOKH CJIIOHHbIX .iKeJie3 c TIHIIJ,eBapHTeJibHhIMH cpepMeIITaMH. L10TKa HMeeT ropTaHHYIO IIJ,eJih c ,n;hIXaTeJihHOH cttcTeMOM. ITm.u;eBo,D, .n;.1HHHhIH 11 JierKo pacTmKHMhIH, HH.iKHaa t.JaCTh ero o6pa3yeT
BpeMeHIIOe BMeCTHJIHIIJ,e TIHIIJ,H -

306. JKeJiy,n;oK pa3,n;eJieH Ha ,n;Ba OT,ll;eJia: JKe-

Jie3HCTbIH, c TOJICThIMH CTeHKaMH, BhI,ll;eJIHIOIIJ,HMH IIHIIJ,eBapHTeJibHhJe COKH;
MYCKYJihHhIH, c BHyTpeHHeii: ITJIOTHOM porOBOM IIOBepxHOCThIO, r.n;e TIHIIJ,a,
CMOtJeHHaa IIHIIJ,eBapHTeJihHhIMH cpepMeHTaMH, MexaHHl.JeCKH o6pa6aThIBaeTca COKpau~eHHHMH MYCKyJiaTyphI H H3MeJibl.JaeTCH 3a Cl.JeT KaMeIUKOB (racTpoJIHTOB).
Knrnet.JHHK ,n;JIHHHhrii:, .n;ncpcpepen.u;11poBaH na ,n;Bena.n;uaTnrrepcTHYIO KnrnKY (oTKphrnaroTca rrpoTOKH rret.Jettn n rro.n;meny.n;oqHott meJie3hI), TOHKHH KHIUel.JHHK, napHhie CJienhie l1 KOpOTKHH TOJICTbIM KHiliel.JHHK, 3aKaHl.JHBaIOIll,HHCH B KJIOaKe. IIpaMaa KlfllIKa Ol.JeHh KOPOTKaa, H HenepeBapeHHhie OCTaTKH rrnIIJ,H,

He naKanJinBaacb,

6hICTpo Bhrno.n;aTCH Hapymy.

CTOpOHe KJIOaKII HMeeTCH CJieno:H Bh!pOCT -

Ha cnnHHOtt

cpa6pttl~HeBa cyMKa, BhIITOJIHHIO-

IIJ,aH cpyHKil.HIO meJie3hI BHyTpeHtteH: ceKpeum1 11 yqacTBYIOIIJ,aa B HMMYHHOM
3aIIJ,ttTe opran113Ma. ITet.JeHh 6oJihlliaH, ,n;ByJiorracTttaa. JKeJil.JHhIH rry3hiph
y roJiy6a OTCYTCTByeT.

JI. btxame.~ bliUJl

cucmeJl.ta COCTOHT 113 CJIO)KHOH Cl1CTeMhl B03,ll;YXOHOCHhIX

rryTeH: 11 .1erK.11x. Bo3.n;yxottoctthre rryTH BKJIIOl.JaIOT H03,n;pn, ttoc0By10 rroJioCTh,
HOCOrJIOTKy, ropTaHb l1 JJ.JIHHHYIO Tpaxero. B MCCTe pa3BeTBJieHMH Tpaxen Ha
6poHXH HMeeTCH H11JKHmI ropTaHb, CBOHCTBeHHaa TOJihKO ITTHuaM. B He:H pacrro.1ara10TCH rOJIOCOBhie CBJI3KH, KOTOpbre rrp11 rrpoxom.n;eHHH B03,n;yxa Btt6pttPYIOT 11 II3,n;a10T 3BYKH. Pa3Hoo6pa3tte II3,n;aBaeMbIX 3BYKOB KOHTpoJI11pyeTca
COKpaIIJ,eHneM CIICil.IIQJHl.JeCKHX rreBl.JHX Mblillil.. JierKHe -

ITJIOTHhie ry6t.JaTbie

Tena, ManopacTa.iKHMhre, c tte60JihIUHM o6'heMoM. CocTO.HT 113 pa3BeTBJieHHhIX
6poHXHaJibHhIX Tpy60K. I'JiaBHhIH 6pOHX BXO,ll;HT B JierKoe II .n;aeT

15-20 pa3-

BeTBJieHHH (BTOpH'IHhIX 6pOHXOB), coe.n;nHeHHhIX Mem.n;y co6o:H rrapa6pottxaMH c

MHOrOtJl1CJieHHbIMII BbipOCT3MH (6pOHXHOJI3MH),

orrJieTeHHhIX ceThIO

KpoBeHOCHhIX KarrnJinapoB (rrponcxo.n;11T ra3oo6MeH). qaCTh 6pottxnanbHhIX
OTBeTBJieHIIH ( 4-5 6poHXOB) BhIX0,11;.HT 33 rrpe.n;eJihI JierKHX 11 o6pa3yIOT TOHKOCTeHHhie pacrnttpeHH.H -

B03,ll;YIIIHhJe MelIIK11, pacIIOJlOJKeHHhie Mem.n;y BHyT-

peHHHMH opraHaMn. ITo o6'heMy ·oHII npttMepHo B

10

pa3 rrpeBhirnaIOT o6'heM

nerK11x. Pa3JIHl.JaIOT rrapHhJe B03,ll;YIIIHhie MeIUKH (rneii:Hh1e, rrepe.n;Herpy,n;Hhie,
3a.n;Herpy,n;Hhie H 6pIOIIlHhie) H HenapHhJe B03,ll;YIIlHhie MelIIKH (Me:»<KJIIOtJHl.JHhle). BhlpocThI B03JJ.YlllHhIX MernKOB npoHHKaIOT B rronocTH KPYIIHhIX KOCTeii:. 3ttat.JeH.11e Bo3,n;yIUHhIX MeIUKOB: oxnam.n;ea11e 11 06nert.JeH11e TeJia, «Haco-

KJLacc llmu!fbl

CbI»,

345

HaKa'lHBaIOru;He B03,ll,YX B

nJIOXO pacnDKHMhle JterI01P (raaoo6'.\1eH

B MernKax He rrpottcxo;:i;HT). MexaHH3M ,nhrxaHHH r·py/~Hoii. ITpu pacmwpemm
rpy,11,HOH KJieTKH npOHCXO,ll,HT B,11,0X, npH paCCJ:ta6JieHHH rpy,ll,HhIX MhIIIIIJ, H cymeHHH rpy,ll,HOH nOJIOCTH -

BhI,ll,OX. B OTJIIFIHe OT BCex Ha3eMHhlX II03BOHO'l-

Hh1X y IITHU rryTh B03,ll,yxa B JierKHe 0,ll,HOCTOpOHHHH, npH 3TOM npOHCXO,ll,HT
aachIUJ;eHHe KHCJIOpO,ll,OM qepe3 JierKHe K31( na B,ll,OXe, TaK H na Bhl,ll,Oxe (Tai{
H33hIB3eMoe ,ll,BOHHOe ,ll,hIXaHHe).

JCp06<'flOC/taJl cucme~ta

33MKHyTaH, HMeeT ,ll,Ba Kpyra KpOB006pau~eHH.H.

Cep,ll,ue qeThipexKaMepttoe (npaBan no.110B11Ha co,ll,epmHT BeH03HYIO 1<pos1>, JieBaJI -

apTepHaJlhHYIO), II03TOMY apTeptta.'IhHan H BCH03HaH KpOBh pa3,11,e.TieHhl

no.'IHOCThIO, 'ITO o6ecne•IHBaeT BhICOKYIO HHTeHCHBHOCTh o6MeHa BeU_\eCTB
(Ten.'IOKpOBHhie .IKHBOTHhie). OT cepm~a OTXO,/J;HT ;:i;Ba caMOCTOHTeJihHhIX cocy,ll,a: JlerO'lHbIH CTBOJI (HeCeT BeH03HYIO KpOBb) H npaBaH ,11,yra aopTbl (apTep11aJ1hHaH KpOBh).

Af aJtbtU l(pyz l(pOBoo6paw,e1wJl.

OT rrpaBoro me.'Iy,ll,O'lKa OTXO,ll,HT Jieroq-

HhIH CTBOJI, KOTOphlH nptt BhIXO,ll,e H3 cep,11,ua ,ll,CJIHTCH Ha npasyIO H JleByIO Jiero'lHhie apTep11H. B JierKHX npo11cxo,ll,HT ra3006MeH, H o6oraru;eHHaH KHC.'Iopo,ll,OM apTepHaJlhHaH KpOBh no npaBOH H JleBOH Jlel'O'lHhlM BeHaM BIIa,ll,aeT B JleBOe npe,ll,cep,ll,He.

BoJLbwou l(pyz l(p06oo6paw,eHUJl.

OT Jresoro meJiy,ll,o'lKa OTXO,ll,HT npaBaH

,ll,yra aopThI, KOTOpaH npH BhIXO,ll,e H3 cep,ll,ua ,ll,CJIHTCH Ha ,ll,Ba cocy,ll,a: npaByIO
H JleByIO 6e3hIMHHHble apTepHH, Kam,ll,aH H3 KOTOpb!X BCTBHTCH Ha o6ru;yro COHHYIO H no,ll,KJIIO'lH'lHYIO apTeptttt (nJie'leByIO H rpy;:i;HyIO). ITpasan ,ll,yra aopThI
ortt6aeT cepJJ;ue 11 cpopMHpyeT crrHHHYIO aopTy. Bee opraHhI Te.Tia cHa6maIOTCH
apTep11aJ1hHOH KpOBhIO. BeH03HaH KpOBh OT 3a,ll,HHX qacTeii TeJia co6ttpaeTCH no
HH./KHeii noJio:B BeHe H Bna.n;aeT B npaBoe npe.n;cep,ll,He, npe,ll,Bap11TeJ1hHO oq11ru;aHCh qepe3 BOpOTHYIO CHCTeMy ne'leHH H IIO'leK. BeH03HaH KpOBh OT nepe,ll,HeH
qacTH TeJia co6HpaeTCH B nepe,ll,HHe nOJihie BeHhI, KOTOphle BIIa,ll,aIOT B npaBoe
npe.n;cep,ll,He. CepJJ;ue KpynHoe no cpaBHeHHIO c pa3MepaMH TeJia. Bo.'IhrnaH cKopocTh UHPKYJIHUHH KpOBH no cocy.n;aM o6ecne'lHBaeTcH BhICOKOH 'IaCTOToii cep,ll,e'lHhIX COKparu;eHHii.

BbtdeJLume.l/bHaJl cucmeMa

npe,ll,cTaBJieHa Ta30BhIMH nO'lKaMH (MeTattecp-

poc, BTOpH'lHaH nO'lKa). 0THOCHTeJibHhlH pa3Mep no'leK y nTHU 6oJihlllOH H HaXO,ll,HTCH B npHMOH CBH3H c O'leHh HHTeHCHBHhIM o6n~HM o6MeHOM Beru;ecTB.
OT no'leK oTxo.n;HT MO'leTO'lHHKH, Bna,ll,aIOru;He B KJIOaKy. Kotte'lHhIH npo,ll,yKT
o6MeHa -

MoqeBaH KHCJioTa. Mo•rn rrpoxo,ll,HT qepe3 BhI,11,eJIHTeJihHbie rryTH 6hI-

CTpo B CBH3H c IIJIOXOH paCTBOpHMOCTbIO MO'leBOH KHCJIOThl, B03MOJi<HOCThIO 3aKyrropKH MO'leBbIMH COJIHMH rrpOBO,ll,H:UJ;HX nyTeH H BhIBO,ll,HTCH B BH,ll,e 6eJihIX
KpHCTaJIJIOB BMeCTe c IIOMeTOM.

c 3THM CBH3aHO OTCYTCTBHe y

ny3hipH. IToTepH BO,ll,hI npn MO'leHcnycKaHHH

y

nTHU MO'ICBOrO

IITMU HeBeJIHKa, TaK KaK B KJIO-

aKe npOHCXO,ll,HT o6paTHOe BCaChIBaHHe BO,ll,hI.

lloJLo6aJl cucme.Ma.

Pa3.n;eJihHOIIOJihie. IToJIOBhie meJie3hI napttbie.

Y caM-

UOB ceMeHHHKH 6o60BH,ll,HOH cpopMhl, nO,ll,BerneHHhie Ha 6pblmeiiKe, pacnoJiaraIOTCH Ha.n; BepxHeii ,11,oJie:B no'leK. BeJIH'IHHa ceMeHHHKOB MeHHeTca no

3oOJiOZUR

31wJioru•rncKue rpyrmbI ITTHU
IIpe,llcTaBHTeJm

Mccn1 0611ra11HH

IlTHilbI J1eca

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CHH!1IJ;3, IIHII\YXa, KOpOJJeK,

Oco6eHHocn1 CTpOCHH;;---t
H IlHTillllHI
, ToHKHtt xpcrrK11i1 I\JllOH.

flOIT0,13CHh

1

U:eu-

1ow 11 OCTpb!C KO!'OTKH, ,'\JillH' Hb!C IJao1bHh!. Jfl'.CCTK!1C llC)Jbf!
: XBOCTll. ncpe.'IeTHbie

-

I

:

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I

Bwibll!OH rrecTph1iI ,llJITCJI

1

,2J,0;10TOBH,l1Hhli'I IC1lOB, l\o1HHHbIH,

'TOHKHii H meCTKHH H3hl!C ,U:JlHH: Hhre, 1ie1m11e na.iihU,hI

(J~Ba Ha-

' npan,1cHhI snepell,, a ,nBa Haaa,l.().
'IlcpbJI XBOCTa meCTKHe n ynpy: r11e. fluraeTCH JIH'IHI!I<a:l-U! HllCC1

KOMblX, 3HMOll -

CCMeHaMl1

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PMC. 140.

roMOnonrn ronOBHOIO M03ra no3BOHOYHblX (BV1A csepxy): a -

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m111ua: ,i:i -· MneKon111rat0U1ee (Kpon111K).

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3-

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npOAOnrOB8Tbl~ M03r

Y

penmu.ll.uil -

aayponcuoHbtil nm

M03ra, Be,n;yrn;ttif oT.n;e.n -

nepeouuu

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HMeeT Me,ll;MaJibHbJH -

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376

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J(BH:>KCHllI1 H .UOCTIIraeT orp0"'1HhIX pa3MepoB, COCTOHT 113 Te.1a M03/KC'!Kn

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142).

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COCTOHT 113 BOPOHKII (HecppOCTO!\ia)

11 Bbl,ll;8JJHTe.1bHOrO KaHa,1bu;a, Bila,D,aIOruero B MO'I8TO'IH11K. Kam;ra.H BOpoHKa OTKphrnaeTCH B ueJIOM. He;racJCKO OT BOpOHKH pa3B11BUCTCH cocy;rnCTh!H K.rry6oCJeK. Ilpo;ryKTbl JJ;I1CCHMH.'1HUHH, co;repmau~11ecH B KPOBl1, IIOCTyrraIOT B ueJIOMH'IeCKY!O JKH,D;KOCTb ll3 cocy;r11cTb!X KJry6oCJKOB. 3aTeM
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K.1y60Y"eK,

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378

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emoputtH.a.R nott1w ( ma3oea.R, ;nemaue<fJpoc)

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30-34;

xapaKTepH3yeTca rroJIHhIM

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B

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pacceqeHtte, pac<u:reHeHtte) --

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opraHM3Ma, ero CMCTeM M opraHOB. AHaTOMM.H paccMaTpMBaeT CTpOeHMe TC.la
qeJIOBeKa, ero opraHOB B pa3JIMqHhie rrepMO,ll;hI JKM3HM, MCCJie,n;yeT oco6eHHOCTII
opraHM3Ma B YCJIOBMHX B:m.HHM.H BHeIIIHeM cpe,D;hI.

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381

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ocmeou (pttc. 143),

rrpe,ncTaBJI.HIOlll,MM co6o:i1 CMCTeMy KOCTHhIX
TIJiaCTHH, 3aMKHYThIX B IJ,IIJIIIH.JJ,PhI, KOTOphie
pacnoJiar8IOTCJI BOKpyr u;eHTPaJihHOI'O KaHaJia.

B KaHaJie npoxo,n.HT KpoBeHoCHhie cocy.n;hr

11 HepBHhie BOJIOKHa. Mem.JJ,y ocTeoHaMM ttMeIOTC.H BCTaBO'lHhie TIJiaCTMHhl, npe,n:cTaBJI.Hl:Olll,He co6oil OCTanrn paHee cylll,CCTBOB8BIIIMX
OCTeOHOB. Mem.n;y KOCTHhIMil IIJiaCTMHaMM
JieJKaT K.1eTKM OCTeOIJ,MThI. PocT OCTeoHa ocyIIJ,eCTBJI.HeTC.H M3HYTPM 38 ClJ:eT KJieTOK OCTe06JiaCTOB.

2

KoHcmpyKU,UH KocmHoit m1caHu o6ecne'lMBaeT npO<IHOCTh KOCTeH. Ila KOCTHOH TKaHM IIOCTpoeHhI:

P111c. 143. CTpoeH1-1e ocrnoHa s pa3pe- ICOMnaKmHoe Kocm1we ee14ecmeo - o6paayeT HapyJKHhIH CJIOH KOCTH, COCTOHT Il3 IIJIOT3e: 1 - nnacrnHKl-1 OCTeOHa; 2- KOCTHble KneTKl-1 (ocrno~1-1Tbi); 3- ~eHTpanb- HO np11Jiera1:0lll,HX .n;pyr K .n;pyry nepeK.'!aHb1i1 KaHan (KaHan OCTeOHa)

,LlMH.

B

.n;ttacpuae .JJ,JIMHHOH KOCTII KOCTHhle

Kocmu u ux coeiJuHeHUR

387

fIJJaCTHHbl pacno:IO:>EeHhl napaJJJIC"lhHO
,ZVIHHHOH ocn;

zy6'-lamoe 1wcmuoe eew,ecmeo -

cocTo-

HT ll.3 nepeKJia)J;ll.H 1l. 6aJIOK, Memll,y KOTOph!Mll. ll.MeeTCH 11pocTpaI1CTBO, 38110JIue1rnoe KOCTHhIM M03rOM. Pac110JiaraeTIIO)J;

CH

KOMnaKTHhIM

Bern,eCTBOM

B KOHu;ax Tpy6'-I8ThTX KOCTeiI -

311ttqrn-

38X, B TeJrnX ry6lJ8ThIX H c:vrewaHHhIX
KOCTeii,

B lVTOCKHX 1l. B03ll,YXOHOCHbIX

KOCTHX (pHC.

144).

3TO CTpOCHHe COOTBeTCTByeT HanpaBJieHll.lO OCHOBHhIX JIHHHH CJK8TIUI (p;aBJieHHH)

1l. paCTHJKeHHH, ;:i;eiICTBYIOllJ,HX Ha KOCTh.
Pac110.TiomeHtte KOCTHhIX rrepeK"1a;:i;IIH no;:i;
yrJIOM ;:i;pyr K ,n;pyry 06ec11e•rnBaeT paBHOMepHoe pacnpe;:i;eJieHtte ;:i;aBJieHIIH w cII.TihI
;:i;eiICTBHH MhIWU Ha KOCTH CKeJieTa.

K JI a cc II qrn Ka u II H
Me, CTpOeHIIIO H

Koc Te iI. Ilo cpop-

oco6eHHOCTHM pa3BIITIIH

KOCTII ;:i;emIT Ha (pIIC.

mpy6'-lambte:

145):

)J;JIHHHhie (11Jie'-Io, 11pe;:i;-

11Jie'-Ihe, 6e;:i;po II I'OJICHh) II KOpOTKHe
(cpaJiaHrH 118"1hUCB, KOCTH 11.T(lOCHhl

P111c. 144. CTpoeHi-ie Kocrn: I - rn-

II

'-I8CTh CKeJieTa KOHe'-IHOCTeii:, Bhll10JI-

nosKa Tpy64aTOC1 Kocm: 2 - ry64aTOe
Bell(ecrno; 3- KOMnaKTHOe Bell(ecrno:

HHH pOJih pbl'Ial'OB 11pII )J;BIIJKeHIIII;

4 - H3,ll,KOCTHl-1LJ,3; 5 - KOCTHblC1 M03r;

zy6'-lambte:

6- cpeAH5l5l 43CTb rpy64aTOC1 KOCm

11.HcT11).

CocTaB"lHIOT

11pell,l1JIIOCH8,

KOpOTKIIe

60Jihwy10

(3aIIHCThe,

1103BOHKII) II cecaMo-

BH)J;Hhie (Ha;:i;KoJ1eHHIIK). CocTOHT II3 ry6'-IaToro Bern,ecTBa, B H'-IeiIKax KOToporo co;:i;epJKllTCH KpaCHhIH KOCTHhlH M03r;

n.noc/Cue
CTH) -

(KOCTH KphllilII qepe11a, pe6pa, .TIOl18TKa, rpy;:i;ttHa, Ta30Bhle KO-

110 CTpOeHIIIO CXO,[(Hhl c Tpy6<raThlMII, yqacTBYlOT B o6pa30B8HIIII

110JIOCTeii: TCJia;
CMeUJGHHblP -

IIMeIOT CJIOJKHYIO cpopMy

ti pa3BIIBaIOTCH ll.3 HeCKOJihKIIX

'18CTeii:. Ha11pIIMep, TeJIO 1103BOHKa 110 cpopMe II 110 CTpOeHHIO OTHOCIITCH
K ry6<raTbIM KOCTHM, a ;:i;yra H OTPOCTKH -

K 11JIOCKIIM;

6030YXOHOC/lble (KOCTH <repena: JI06HaH, KJIIIHOBII)J;HaH, peweT'-IaTaH,
BepXHHH <reJIIOCTb) -

IIMeIOT B TeJie 110JIOCTh, BbICTJiaHHYIO CJIII3HCTOH

o60JIO'IKOH H 38110JIHeHHYIO B03)J;YXOM.
CTpoeHwe KOCTH (Ha 11pHMepe Tpy6<raTo:H).

B Tpy6<raThIX KOCTHX pa3Jitt<raroT:

Te.no, HJIH iJuarjJu.J - cpe;:i;HHH <raCTh, o6pa3oBaHo KOM11aKTHhIM Bern,ecTBOM.
3nurjJu.J - YTOJillJ,eHHhie KOHIJ;hl KOCTH c cycTaBHhIMH IIOBepxHOCTHMH,
IIOKpblThie cycTaBHhIM xp.HllJ,OM.

AuamoMUJl u cpu3uo:wzuJi 'le1w6e1w

388

2

P111c.145.

B~Abl KOCTeiil:

rne) KOCrn;

4 ·--

1 -AJl~HHa>l

(Tpy6'·JaTa51) KOCTb:

2-

nnocKa>l KOCTb:

3-

ry64aTble (KOpOT-

CMewaHHaS< KOCTb

MemarjJu3

(rrnaCTMHKa p0CT8) -

y<I8CTOK KOCTM, p8CIIOJIO)KeHHhlll Mem.ny

,ll;M8cpM30M M 3IIMcpM30M. LJ:o OKOCTeHeHM.H 3,ll;eCb p8CIIOJI8raJI8Cb xpmu,eB8.H
IIJiaCTMHK8, 38 cqeT KOTOpoH. KOCTh pOCJI8 B ,ll;JIMHY (.no

Ano</JU3bl -

20-25 JieT).

6yropKM, K KOTOpbIM rrpMKperrJI.HlOTC.H MhIIIIll,hI M MX cyxo-

)KMJIM.H, cp8Cll,MM, CB.H3KM.

H ai:hwcm Huu,a -

H8pymtt8.H, rrpoqtta.H coe,n:MHMTeJihHOTKaHH8.H IIJI8C-

TMHK8, 6or8T8.H KPOBeHOCHhIMM ll JIMMcp8TMqecKMMM cocy,n:aMM, HepBaMM, 38 MCKJIIOqeHMeM cycTaBHhIX IIOBepxHOCTeiI. CocTOMT M3 ,n:Byx CJIOeB:
Hapymttoro
cf>yHKll,MM:

(BOJIOKHMCToro)
38Ill,MTH8.H,

ll

BHyTpeHHero

TpOcpMqecK8.H

(KOCTeo6p83YIOill,ero).

(IIHT8TeJihH8.H),

HepBHO-peryJI.H-

TOpHa.H, KOCTeo6p830B8TeJihH8.H (pocT B TOJIID;MHY ll cp8CT8HMe rrpM rrepeJIOM8X).
BttyTpM KOCTM MMeIOTC.H KOCTHOM03rOBhie IIOJIOCTM ll Jiqeti:KM ry6q8Toro BeIll,eCTBa, B KOTOpbIX H8XO,ll;MTC.H KOCTHhIH M03r.

y

HOBOpom,n:eHHOro pe6eHKa

KOCTHOM03rOBhie IIOJIOCTM 38IIOJIHeHhI KpaCHhIM KOCTHhlM M03rOM, KOTOpbIH
BhlIIOJIH.HeT KpOB006pa3yIOill,YlO ll 3aill,MTHYIO cpyHKD;MM.

lfa CTBOJIOBhIX KJieTOK

Kp8CHOro KOCTHOrO M03ra o6pa3yIOTC.H KJieTKM KpOBM M KJieTKM MMMYHHOH CMCTeMbl (JIHMcpon;MTbI).

y

B3pOCJioro qeJIOBeKa KpaCHhIR KOCTHhIR M03r coxp8-

H.HeTc.H TOJihl{Q B JiqerrK8X ry6qaToro Beill,eCTBa IIJIOCKMX KOCTeli (KOCTM qepeII8, rpy,n:r·IH8, IIO,ll;B3/J;OIIIHhie KOCTM), 3{1McpM3aX Tpy6q8ThIX KOCTeli. B KOCTHO-

Kocmu u ux coe<lu1u'1WH

389

M03ro1mi:i II0.10CTH .n.naqrn30B Tpy6'IaThIX KOCTeii HaXO.[\HTCH me.'lTh!H KOCTHbJH
M03r, rrpe,ll,CTaB.1HJOlll,HH co6oii cKorr.1ettne mnpcrnoii TKattn. Macca KOCTHoro
M03I'a COCTaB.15IeT

4-5 'X,

OT MaCChI Teo"Ia.

CoeiJuHeHue 1wcmeu. KocT11 cKe:1eTa Mem,n:y co6oii coeti;trneHhI rrpw noMomu
pa3Jiwrnoro B11.n.a coe.n.nHen11ii:

1.

HerrpepbIBHLie COe,ll.HHCHHR ( CU!iapmpO.Jbl) ·-- Mem.n.y KOCTHMH HeT rrepepb!Ba, a TOJihKO npoCJIOMKa, COCT05Illl,a5I H3 pa3HhIX TKaHeii. Pa3Jlll'IaJOT:
cu liOPC.M.03 -

coetrHHeHHe KOCTe:ii npn ITOMOlll,H ITJIOTHO:ii coe,n:HHHTe.TJh-

HOH TKaHH (WBhl KOCTeii qepena);
CU/lXO!iOp03 -

coetiHHeHne KOCTe11: npn IIOMOlll,H xpRlll,a ( coe,n:HHeHHe

pe6ep c rpytiHHOH, MCiKIT03BOHO'IHhie xpRlll,eBbie ,ll;HCK11);
CUfWCmo.1 -

COC,ll.HHeHne KOCTeii rrpH IIOMOlll,H KOCTHOH TKaHn (B aM6-

pnoreHe3e JI06Ha5I KOCTh COCTOHT H3 .n.syx IIOJIOBHHOK, KOTOphie K pom.n.eHHIO cpaIIJ;HB8IOTC5I Mem.n.y co6oii).

2.

IloJiycycTaBbI ( CU.M</JU3bl) -

npe.n;cTaBJIRIOT co6oii coe.n.uneHHH ,ll;BYX KO-

CTeii rrpw IIOMOIIJ;H xpRlll,a (.106KOBhIM CHMcpH3).

3.

IlpephIBHhie coetiwnemrn ( cycmaBbL, uJLu iJuapmpoabt) -

Mem.n.y coe.n.11-

HRIOlll,HMHCR KOCTRMII eCTh nepephIB, KOCTII o6'be,ll;HHeHhl BMeCTe cycTaBHOM cyMKoii. 0TJIIIt.IaIOTCR 6oJILwoii no,n:BIImnocTLIO.

KJiacc11cp11Ka1~HR cycTaBOB. CTpoe1me cycTaBa (pIIc.

146).

B cycTaBe

pa3JIII'!aJOT OCHOBHh!e H BCITOMOraTeJihHhie aJieMeHThI.
OcttOBHhie aJieMeHTh1: cycmaB1tbte noBepxHocmu cot.I.'IeHRIOlll,HXCH KOCTeii,
noKphIThie cycTaBHhIM xpRlll,OM; cycmaB1ta.fl 1cancyJ1a ( cy.M1ca ), oKpy:maIOlll,aR
KOHUhl COt.I.'IeHRIOlll,HXCH KOCTeii; cycmaB/la.R llOJlOCmb, 3anO.'IHeHHaH CIIHOBllaJihHOM JKII,ll;KOCThlO, yMeHbIIIaIOlll,eii TpeHHe.
BcnoMoraTeJihHhie aJieMeHThI: CBR3KII, MeM6paHLI, xpRlll,eBhie .n.ttcKII, MeHIICKII, CHHOBII8JlhHhie CKJia,ll;KII II ,ll;p.
KJiaccIIqrnKaunR cycTaBoB:

1.

Ilo 'IllCJIY cycTaBHhIX IIOBepXHOCTeii:
npocmou -

HMeeT ,ll;Be cycTaBHbie IlOBepxHOCTH (Me:mcpaJiaHrOBhlH II

rr.'Ie'leBOM cycTaBhI); CJlO;HC/iblU -

HMeeT TPII II 6oJiee cycTaBHhie no-

BepXHOCTH (JIOKTeBOM cycTaB); /COM6UiiUp08aliHblU -

COCTOIIT II3 .n.syx

H 60.1ee aHaTOMII'-IeCKII caMOCTORTeJibHhIX cycTaBOB, cpyHKUHOHIIpyJOlll,lIX 0,ll;HOBpeMCHHO (BHCO'-IHO-HIIiKHet.IeJIIOCTHOM cycTaB); /CO.MllJlPICCllblU -

Mem,n:y CO'-IJieH5IIOlll,HMIIC5I ITOBepxHOCTRMH HMeeTCR tJIICK HJJII

Mhilll,eJIKH, KOTOpb!e .n.eJIRT cycTaB Ha ,n:Be KaMepbl (KOJieHHbIM cycTaB).

2.

Ilo cpopMe: wapOBUOHble (nJiet.IeBoii, Ta3o6e,n:peHHblii); .JJlJlUllC06UOllblP
(Jiy•re3anacTHhiii); ceiJJLoeuiJHbte (3anacTHO-nacTHbiii nepBoro naJ'Ihua);
6JlO/C06UOHbte (MemcpaJiaHrOBbie CO'!JieHeHHH naJibIJ;eB); n.rWC/CUP (3anaCTHO-ITHCTHbIH); u,uJ1uHiJpu'lec1Cue (JiyqeJIOKTeBoii,

I

II

II

3BOHK11); .MblUJ,PJl/COBblU (KOJieHHbIH).
<I>opMa cycTaBa onpe.n.emi:eTCR KOJIIIt.IeCTBOM oceii Bpan~euIIa.

3.

Ilo OCRM ,ll;BHiKeHMH:

rueiiubie no-

AnamoMull u rpu.1uo1wzull 'le.rioBeKa

390

2

3

3
4
5

5

4

6

Pvic. 146. a - cxeMa crpoeHWl cycrasa: 1 - HaAKOCTHVIL\a; 2 - KOCTb; 3 - cycrasHasi Kancyna; 4 cycrasHoi1 xpsil!1; 5 - cycrasHasi nonocTb; 6 -

pa3nVIYHble cj:>opMbl cycrasos (cxeMa): 1 - waposvig-

Hb1i1; 2 - 3nmmcOBVIAHbli1; 3 - CeAflOBVIAHbli1; 4 - nnOCKVli1; 5 - 6noKOBVIAHbli1; 6 - L\VIJlVIHAPVIY8CKVIC1

OOHOOCHbte (BOKpyr
i)eyocHble (BOKpyr

0,11,HOH OCH) ,11,BYX oceii:)

IJ;MJIMH,11,pMqecKMH M 6JIOKOBM,ll,Hhitt;
-

3JIJIMTICOBM,ll,HhIH,

ce,D,J10BM,ll,Hhlll

M MhIIIJ;eJIKOBhitt;

MHO?OOCllble (BOKpyr Tpex oceii:) - IUapOBH,ll,HblH M TIJIOCKMH.
CKeJtem (OT rpeq, skeleton - BhICyIUeHHhrii) (pttc. 14 7) qeJioBeKa npe,11,cTaBm:ieT co6oii: coBoKynHoCTh KOCTeii:, onpe,11,eJieHHhIM o6pa3oM coe,11,ttHeHHhIX ,11,pyr
c ,11,pyroM.

B

cKeJieTe BhI,D,eJ1HIOT TPM OT,11,e.'la: cKeJieT TYJIOBMIIJ;a, cKeJieT qepena

M CKeJieT KOHeqHOCTeii.
KocTM cKeJieTa BhITIOJIHHIOT onopttyIO, ,D,BttraTeJihHYIO M 3aIIJ;MTHYIO cpyHKU:MM. KocTM TaKme HBJIHIOTCH ,11,eno ,ll,J1H MMHepaJihHhIX coJ1eii:.

CICeJtem my.weuw,a

o6pa3oBaH no3BOHOqHhIM CTOJI60M (no3BOHoqHMKOM)

M rpy,11,Hoii KJieTKOH.

Il0360HO'lHblU cmoJt6

.HBJIHeTC.H OCHOBHhIM CTepmHeM, KOCTHOH OChIO TeJ1a

M ero onopoii. Ott 3aIIJ;ttIIJ;aeT cnHHHoii: M03r, cocTaBJIHeT qacTh cTeHoK rpy,11,Hoii:, 6pIOIUHOi1: M Ta30B0ll IlOJIOCTeii:, a TaKme yqacTByeT B ,11,BMmeHMM TYJIOBMIIJ;a M roJIOBhI. CocTOMT M3

33-34 II03BOHKOB,

TeJihHO o::i;ttH Ha ,11,pyroll:.

B

IUeHHbIH

(7

Il03BOHKOB); rpy,ll,HOH

KOB); KpeCTIJ;OBblll
KorrqMKOBhIH

(4-5

(5

HaK.'la,D,hIBaIOIIJ;MXCH nocJie,11,oBa-

no3BOHOqHMKe pa3JIMqaIOT cJie,11,yIDIIJ;Me OT,ll,eJihI:

(12

Il03BOHKOB); IIOHCHMqHhIH

TI03BOHKOB, cpOCIUMXCH B e,11,IIHYIO KOCTb -

i)yiy,

meJto,

o6paIIJ;ettttoe Bne-

KOTOpaH npHKpenJIHeTCH K Te.'ly Il03BOHKa C3a,D,H • .IJ:yra M TeJIO rro-

3BOHKa 06paay10T IUHpoKoe noaBoHoqHoe OTBepcTHe.
,11,HT

II03BOH-

KpecTeu;);

CpOClllMXCH Il03BOHKOB).

Kam,11,hrii no3BOHOK HMeeT yTOJIIIJ;eHHYIO qacTh pe,11,, M

(5

ompocm/Cu

OT

,11,yrH noaBOHKa oTxo-

pa3HOM BeJIMqHHhl Il HanpaBJieHHOCTM:

no cpe.n;Heii JIHHHH OT ,11,yrM Ha3a,D, OTXO,ll,MT HenapHblll
TOK ( CJiymHT ,11,JIH npMKpenJieHMH CBH30K M MbllllIJ;);

ocmucmblU

OTpOC-

391
Kocmu u ux coeiJunettuJl

Pr.tc. 147. CKener yenoseKa: a - cnepeA11: 6 - c3;m11. 1 - Yepen; 2 - KJ110Y1<1ua : 3 - nonarKa; 4 rpyAHall rneTKa ; 5 - nneYeBall KOCTb; 6 - pe6pa; 7 - n03BOHOYHl1K; 8 - KOCTl1 Ta3a; 9 - KOCTl1 npeAnne'lbSl ; 10- K1'1CTl1; 11 ·- 6eAP8HHaS1 KOCTb; 12- KOCTl1 roneH11; 13 - KOCTl1 cTOnbl

392

AnamoMUR u <fJuauonotuR 'leno6eKa

B CTOpOHLI OT ,IJ,yr11 OTXOJJ;5IT napHbie nonepP'IHbl<' OTPOCTKH

(;vrn:

COe,L\H-

HeHHH c pe6pa:vm);
BBepx I1 BHH3 HaupaBJJeHbl napHb!e nepXHMe Ii HM)l<HMe cycma6HblP OTPOCTKM

(.zvrn coe;:(MHeHIIH c

BhIIIJe M HM/Ke pacno.'IO:IKeHHbIMI1 II03BOHKa:vm).

Pa3MephI rro3BOHKOB Heo,zr11HaKOBhr, qeM 6oJibllle Harpy3Ka, TeM 6oJihWe
pa3Mepbl II03BOHKOB.

ill PUHblP

ll0.J60H/CU MMCIOT cpaBHMTe.'IbHO He60JibllIOe Te.'IO, TaK KaK Ol{J}

MCIIbITbIBaIOT MeHhIIIYIO Harpy3KY' qeM II03BOHKI1 ,zrpyrMX OT;:i;e.'IOB. IlorrepetrUhie OTpOCTKM IIIettHhIX II03BOHKOB HMeIOT OTBepcTHH, qepe3 KOTOphre rrpoxo/:(MT no3BOHoqHaH apTep11a. OcTMCThre oTpocTKH Ha KoHrr;e pa3,zrBoeHbI.
JiaHT) M

II (arrMcTpo<lJeil:) IIJettHhie rr03BOHKH

MMeIOT OTJIMqHyIO OT .n;pyrnx

I (aTIllett-

HhIX I103BOHKOB <lJopMy' IIOCKOJibKY OHM CO'IJieHHlOTCH c qeperroM' HecyT Ha
ce6e ero THllieCTb M yqacTBYIOT BO BPaII~eHHM.
I'pyihtbte ll0.360/ilCU (pMC.

148) KpyrrHee IIIettHhIX M

coqJieHHIOTCH c pe6paM11.

0CTI1CThie OTpOCTKM rpy.n;HbIX II03BOHKOB pe3KO HaKJIOHeHhl BHM3 H He pa3,QBOeHbl, qTO nperrFITCTByeT HaKJIOHY rpy;:\HOro OT,ll;eJia II03BOHOqHoro CTOJI6a Ha3a,zr.
IIORC/lU'IHblP

ll0360HKU

OTJIMqaIOTCF!

MaCCMBHOCThIO TeJI,

qTo CBH3aHo

c 60JibllIOll uarpy3KOll Ha HHX. OcTHCThie OTPOCTKH y HIIX HanpaBJieHhI Ha3a/:(.
Kpecmu,o6bte no.JBOHKU B IOHOCTM cpacTaIOTCF! B e;:\IIHyIO KOCTb -

KpecTerr;,

.D;JIH npMcnoco6JieHIIH K 60JihllIOll Harpy3Ke, KOTopyIO HeceT 3Ta qaCTh CKeJieTa.
Kon'lUIWBblP ll03601l/CU -

OCTaTKM Mcqe3HYBIIIero XBOCTa, OHM py.n;nMeH-

TapHbI M CJIMBaIOTCH B O,ll;HY KOCTb -

Konq11K.

Il03BOHOllHhie OTBepCTMF!, HaKJHl,QbIBaHCb ;:i;pyr Ha ;:i;pyra, o6pa3yIOT Il03BOHOqHblll KaHaJI, B KOTOpOM pacrro.'laraeTCH CIIMHHOH M03r BMeCTe c IIOKpbIBaIO-

~MMM ero o6oJioqKaMM.

1

2

5

60-65 CM, y Mymqm1 60- 70 CM, qTo COCTaBJIFieT rrp11MepHO

eT ,QJIMHY
2

I 3 /:\JIMHhI TeJia.
Y

4

Il03BOHOq-

Hblll CTOJI6 B3poc.'IOH llieH~MHbI MMe-

HOBOpOlli;:\eHHhIX Il03BOHOqHhIH

CTOJI6 npFIMOtt, no B npou;ecce pocTa pe6eHKa IIOF!BJIHIOTCF! ecTeCTBeHHhie <lJM3MOJIOrIIqeCKMe M3rM6b1: Ha3a.n -

cpo.J

f(U-

(rpy.n;HOH 11 KpeCTII;OBblH), nnepe;:\

-

Jl0p003 (mettHblH II IIOF!CmfqHblH).

B

JielliaqeM noJiollieHIIM TeJia M3rM6hr

pacnpF!MJIHIOTCH, II03BOHOqHMK y.z:vrnHHeTCH. Ilp11 BepTMKaJibHOM ITOJIOJKe-

P111c. 148.
(B~A

CTpoeH~e

rpyAHOro no3BOHKa

csepxy): 1 - Teno no3BOHKa; 2-

nonepeYHb1i'! OTpOCTOK; 3 - n03BOH04HOe
OTBepcrne; 4 -

sepxH~i'! cyCTaBHO~

OTPOCTOK;

HMII, oco6eHHO np11 ttarpy3Kax, ouu
BhipallieHbI 60.11ee qeTKO. BoKOBhre 11cKpIIBJieHIIH II03BOHOqHMKa IIOHBJIFIIOTCF! o6hrqHo M3-3a HenpaBIIJihHoro cu.zre-

5 - pe6epHa5l 5lMKa nonepeYHOro OTpOCTKa;

HIIH 3a CTOJIOM, 3a napTOll (y Il!KOJibHH-

6 - ocrnCTbli'I OTPOCTOK; 7 - 1:wra n03BOHKa

KOB) -

CKOJlUO;J -

M3rn6 B CTopoHy.

Kocmu u ux coedunenun

393

I'pyiJ1W5l ICJlem1w COCTOIIT II3

12

rrap pe6ep, rpy,[IIIIlbI II rpy,ilHblX rro-

3BOHKOB.

I'pyau1w -

HerrapHaH rr.'IOCKaH KOCTh, COCTOH~aH II3 PYKOHTKH, Te.1a II Me-

qeBII,[\HOro OTpOCT!rn.

Pe6pa rrpe.z:1cTaBJIHIOT co6oil ilJIIIHHhre II30rHyThie KOCTHhre rrJiaCTIIHKH,
nepeXOilH~He B rrepe.z:1HeM OT,[leJie B xpJI~eBhre. C3a,[ltt Ka)l<µ;oe pe6po 3aKau'lIIBaeTCH rOJIOBKOll, coe.n;IIHflIO~eiicR c TeJiaMU COOTBeTCTBYIOII~HX IT03BOHKOB, a 6yropoK weHKH pe6pa 06pa3yeT cycTaB c rrorrepe'lHhIM OTpOCTKOM norrepe.n;HHMH CBOHMH KOH1~8MH ilO-

3BOHKa. CeMb nap BepXHIIX pe6ep

(I-VII)

CTHraIOT rpy.n;ntthr, nx Ha3hIBaIOT

ucmuftftblMU pe6paMu.

VIII, IX, X

pe6pa

rpy,[IHHhl He ,ll;OCTHraIOT, OHH coe,n;HHHIOTCH c BhIIIIeJiema~HM pe6poM, II03TOMY

rroJiy'lHJIH Ha3BaHne

JlOJICflbtX pe6ep.

XI

H

XII

pe6pa aaKawmsaioTcH

B MhlIIIIJ;aX 6pIOI1IHOH CTeHKH, OTJIH'lalOTCJI 60JihllIOH IIO,ll;BH)l{HOCThlO.
3bIBalOT

KOJle6Jl10W,UMUCR pe6paMu.

K

11x

Ha-

pe6paM rrpnKpernreHhI .n;hlxaTeJlhHhre

MhIIIIIJ;hI II .n;nacpparMa.
rpy,n;Ha.H KJieTKa YIIJIO~eHa B nepe,n;He3a,n;HeM HarrpaBJieHHH, lIMeeT cpopMy
ttenpaBIIJlhHOrO KOHyca.

06Jia,n;aeT 60JihIIIOH npoqHOCThlO,

3JiaCTHqHOCThIO

II IIO,[\BH)l{HOCThlO. 3To o6ecrreqnBaeTC.H HaJIH'llleM rtt6KnX pe6epHblX .n;yr, HMe-

IO~llX xpH~eBbie ll KOCTHhie cerMeHThI. IlpII ilhIXaTeJibHhIX ilBH)l{eHHHX rpy.n;Ha.H KJieTKa II3MeHHeT CBOII ofrheM l1 cpopMy.
rpy,n;HaH KJieTKa MO)l{eT .n;ecpOpMllpOBaTbC.H npn HeUpaBHJihHOM ClI)'.(eHIIll
n HeKoTophrx aa6oJieBattnax. TaK, ecJin IIIKOJihHHK, Kor,n;a IIIIIIIeT, JIO)l{HTCH na
napTy, rpy,n;HaH KJieTKa MomeT ccpopMIIpoBaThCH yrrJ'IO~eHHOH. IlpII paxnTe
rpy,n;Ha.H KJieTKa rrpIIo6peTaeT cpopMy «KYPlIHOH rpy,n;KII», BhICTyrraH Brrepe.n;.
MyahIKaHThr, urpaIO~ue Ha Tpy6e, cTeKJio,n;yBhI nMeIOT IIIHpoKyIO u BhIIIYKJIYIO rpy,n;ttyro KJieTKY.

CKeJlem ZOJlOBbl ( ttepen) (p11c.

149) o6pa3oBaH

rraptthIMH II tterrapHhIMH Ko-

CT.HMII, rrpo<IHO coe.n;utteHHhIMH c rroMOill;hIO IllBOB. CocTOIIT H3 ,n;syx oT.n;eJioB:
JIHIJ;esoro (micu;epaJihHoro) u M03roBoro (qeperrttoii Kopo6KII).

Jiuu,eaou omiJeJl ttepena cocTaBJIHIOT:
1) aepxHetteJl/OCmHa.fl Kocmb - rrapttaH, yqacTByeT B o6pa30BaHHH HHmtteH:
CTeHKlI rJia3HlIIJ;hI, HOCOBOH lI pOTOBOH IIOJIOCTeH, IIO,[\BIICO'lHOll lI KphlJIOBH,ZJ;HO-He6HOH HMOK;

2) cKyJloBa.fl KOCmb - rrapHa.H, Hau6oJiee KperrKaH;
3) fte6Ha.fl /COCmb - rrapHaH, yqacTByeT B o6pa30BaHHH

IIOJIOCTH HOCa, pTa,

rJia3HHIJ;hl H KphlJIOBH,ll;HO-He6Hoii .HMKH;

4) flOC08llJl /COCmb 5) cJle3Ha.fl Kocmb -

rrapHaH, o6paayeT KOCTHYIO CIIlIHKY HOCa;
TOHKaa, napHa.H, pacrroJiaraeTCH B o6JiacTJI rrepeilHero

OT,n;eJia BHyTpeHHeH CTeHKH rJia3HHIJ;hl, npHHIIMaeT yqaCTHe B o6pa30BaHHH HOCOCJie3HOro KaHaJia;

6) ftUJICftRR ftOCOBaJl paK08Ufla -

rrapHaH KOCTh, pacrroJiaraeTCfi Ha 6oKO-

BOH cTeHKe HOCOBoH: noJIOCTII;

7) COUlftUI( -

HerrapHaa KOCTHaH IIJiaCTIIHKa, pacnoJiaraeTCH B HOCOBOH no-

JIOCTH ll o6paayeT neperopo,n;KH HOCOBOH IIOJIOCTH;

AnamoMUR u <jJu.1no;iozuR •w;i0He1rn

394

8

6
PMC.

149.

Yepen YenoseKa:

n04H35l KOCTb;

3-

a-

Bi-lA c6oKy;

4-

B"1C04H35l KOCTb;

sepXH5l5l 48nlOCTH35l KOCTb:

7-

6-

HOCOB35l KOCTb;

8) ftUJICHH.R 'le.'l/OCmb -

B"1A cnepeAi-1.

CKynosa5l KOCTb;

8-

5-

1-

T8M8HH35l KOCTb;

2-

H"1)1(H5J5l 48nlOCTH35l KOCTb;

33Tbl-

6-

no6Ha5l KOCTb

HenapHa.H, KpenKa.H, .HBJI.HCTC.H e,IJ,HHCTBeHHOll no-

,L\BH:iKHOll KOCTblO qepena, KOTOpa.a COC,L\HH.HCTC.H c BMCOqHbIM11 KOCT.HMH
cycTaBOM;

9) noiJoR:Ibt'lHaR Kocmb -

HenapHa.a, pacno;rnmeHa B o6;rncTM we11, Mem-

.n;y Hll:>KHeH qeJIIOCTblO 11 ropTaHhlO.
ct>yHKil,HH

JIHIJ,eBoro OT,L\eJia:

onopa Haqa.TJhHbIX OT,IJ,CJIOB /(bIXaTeJihHOH

11 IlHIIl,eBapttTeJibHOll Cl!CTeM; KOCTHa.H OCHOBa Jll!IJ,a.

Moazoaoil omiJeJt 'tepena cocT011T 113 KpbIIIIH 11 ocHoBaHH.H. KocTH nJIOCKHe,
BHyTpeHH.H.H IlOBepxHOCTb c yr;ry6.JieH.H.HM.H, COOTBeTCTBYJOIIl,MM.H 113B.HJI11HaM
11: 6opo3.n;aM M03ra. Pa3JI11:qaJOT HenapHhie 11: napHbie KOCT.H M03roBoro •iepena.

H enapubte Kocmu:
1) 3ambtJlO'lHa.R l(OCmb -

np.HH11MaeT yqacT.He B o6pa3oBaH.H.H OCHOBaHH.H

qepena .H 3a.n;Hero OT.n;eJia ero KpbIIII.H; .HMCCT 60JibIIIOe 3aThIJIOqHOC OTBepcT.He, qepe3 KOTopoe npoxo.n;11:T npo.n;oJiroBaTbitt M03r 11: no3BOHoqHbie
apTep11:11:;

2) Jl06Ha.H l(OCmb -

yqaCTByeT B o6pa30BaH.Hli KpbIIIIH H OCHOBaHH.H qepe-

na. lfMeeT JI06Hy10, rJla3Hl{qHbie .H HOCOBYIO qacT11;

3) pewem'lama.R ICOCmb -

JierKa.H .H xpynKa.H, COCTOHT H3 .L\BYX nJiaCTl!HOK:

peIIIeTqaToil nJiacT.HHK« 11: peIIIeTqaToro Jia6ttpHHTa. PemeTqaTa.a rvracT.HHKa pacnoJiaraeTc.a rop.H30HTaJihHO B peIIIeTqaTOH Bhrpe3Ke Jio6noH:
KOCTH. 0Ha HMeeT MHO:>KCCTBO OTBepCTHll, a B cpe.n;Hell IlJIOCKOCTM OT Hee
OTXO,L\.HT o6paIIl,eHHbIH KBepxy KOCTHbIM BbICTyn -

neTyIIIHHbIM rpe-

6eHb, K KOTOpOMY npMKpenmieTC.H OTpOCTOK TBep.n;oii M03rOBOM o60JIOq-

Kocmu u ux coe{}unenufl

395

Kll. qepe3 OTBepcnie peweT'IaTOH IIJ:IaCTllHKll rrpoxo,n.HT ll3 HOCOBOH IIOJJOCTH B IIOJIOCTh qeperra o6oH.HTeJJhHhie HepBhI;

4)

l(JlUfl06UOftaJI, KOCmb -

co.nepJKWT B031~YXOHOCHYIO rra3yxy, coo6w;a10-

w;y10ca crrepe.nu c Hocoso:H: rroJIOCThIO. Ha sepxHett rrosepxnoC'rw rra3yxu

Typer~Koe ce,n:JJo, r.ne pacrroJraraeTca rurroqrn3. qe-

ecTh yrJJy6JJeHwe -

pe3 MHOrO'IHCJJeHHbie OTBepCTll.fl rrpOXO,ll;JIT qeperrHhie HepBbI.
IIapHbte Kocmu 'lepena:

1)

meMeWtaJl KOCmb -

COCTaBJrneT ~efITpa.'lhHYIO 'IaCTh CBO,n;a qeperra.

Ha ee BhIIIYKJJOH IIOBepXHOCTll eCTh B03Bhllll8Hlle -

TeMeHHOM 6yrop.

BnyTpeHH.HJI (BOrHyTaa) IIOBepXHOCTh TeMeHHOM KOCTH llMeeT apTepttaJJbHble 1I CllHYCOII,ll;aJJhHbie 6op03,ll;hl, JIBJJJIIOill;MeC.H MeCTOM KpOBeHOCHOrO CMHyca TBep.noii M03rOBOH o60JJO'IKM;

2)

6UCO'lHaR KOCmb -

npMHMMaeT yqacTwe B o6pa3oBaHMM OCHOBaHM.fl qepe-

na M KphlllIM. CoCTOMT M3 Tpex qacTeii: KaMeHMCTOH (co.nepJKMT KOCTHhIH
JJa6MpttHT M cpop:vmpyeT BHyTpeHHee yxo), qernytt'IaToif tt 6apa6aHHott;

3)

CJtyX06bte KOCmO'll(U (MOJJOTO'IeK, HaKOBaJJhHJI 1I CTpeMe'IKO).

<I>yHK~MH M03rOBOrO OT,ll;eJJa: BM8CT1IJJ1Iill;e 1I 3aill;MTa rOJJOBHOro M03ra.

qeperr coe.nuHeH ,ll;BYM.H 3aThIJJO'IHhIMll Mhlru;eJJKaMM c II03BOHO'IH1IKOM. KoCTM qepenHOM KOpo6KH rrpOXO,ll;JIT ,ll;Be CTa,n:ttll pa3B1ITMJI: nepen01t'lamy10, ocTaTKOM KOTOpOH JIBJJJIIOTCJI pO,ll;HM'IKll: rrepe,ll;HllH (JJ06HblH) -

caMhIM 6oJib-

IlIOH, IlOJIHOC'fhIO 3apacTaeT Ha BTOpOM ro.ny JKM3HM; 3a,ll;H1IH (3aThIJJO'IHhIM) 3apacTaeT BCKOpe IlOCJJe pom.neHllJI, 1I KOCmHy/O -

IIOCJie 3apacTaHll.fl pO,IJ;HM'I-

KOB o6pa3yIOTC.H rnBhI, IWTOphre o6ecrre'IttBaIOT coe.nttHeHtte KOCTeif qepena.
CKPJtem Ko1te'lnocmPil. CKeJieT BepxHeif KOHe'IHOCTtt cocTOllT tt3:

1. II oJLca

eepxuux KOHe'lHOCmPu, o6pa3oBaHHoro rrapHhIMll JionaTKaMM

1I KJIIO'Ill~aMll. JionaTKa -

KOCTh TpeyrOJJhHOM cpopMbI,

rrpwJieJKllT

K 3a.nHeif noBepxHOCTM rpy.nHoii KJieTKll. PacrroJJaraeTca Ha ypoBHe OT
.no

VII

II

rpy.nHoro pe6pa. BoKoBoif yroJI JiorraTKll MMeeT yToJJru;eHHe,

Ha KOTOpOM HaXO,ll;llTCJI cycTaBHaa Bila,ll;ttHa ,ll;JIJI coe,ll;MHCHllJI c rOJIOBKOH
IlJJe'IeBOH KOCTll. I\JIIO'IH~a -

S-o6pa3HO ll30rHyTaa Tpy6qa'fa.H KOCTh,

pacrroJiomeHHaa Mem.ny rpy.nMnoil M JJonaTKoii. Y.nepmttBaeT rrJJe'IeBoH
cycTaB na HeKOTOpOM paCCTOJIHHll OT rpy,ll;HOH KJieTKll, o6ecrre'IIIBaJI
60JJhllIYIO CB06o.ny ,ll;B1IJK8H1IH BepxHeH KOHe'IHOCTll.

2.

CKe.TlPm
KOCTh -

ceo6ooHou

eepxHeil

Ko1w'l1tocmu

cocTaBJiaroT:

1rneqeBaa

,ll;JillHHaa Tpy6qaTaa, yqacTByeT B o6pa3oBaHttH rrne'!enoro cyc-

TaBa; KOCTll rrpe.nrrJIC'IhJI: JJOKTeBaa 1I Jiy'IeBaa -

,ll;J"IllHHhie Tpy6'IaThie

KOCTll, JIOKTeBa.H pacnoJJOJKeHa co CTOpOHhl MH3HH~a. a Jiy'IeBaJI -

co

cTopOHhI 6oJihWOro naJJh~a. OHM yqacTBYIOT B o6pa3oBaHllll JJOKTeBoro
1I ;ryqenoro cycTaBOB. KocTtt KttCTll BKJJIO'IaIOT:
noJiomeHHhIX B ,ll;Ba p.H.na (ry6'IaThie),

5

8 KocTeil 3arracThR, pac14 cpaJJaHr ITaJJh-

KOCTeM Il.HCTll II

~eB ('IeTbipe naJih~a MMeIOT ITO 3 cpaJiaHrM, 60JibilIOH rraJJe~ -

2) -

KO-

pOTKHe, Tpy6'IaTbie.
CKe.Tlem HUJICHeil KOHe'lHOCmu COCTOllT II3:

1.

I I ORCa HUJICHUX l(OHe'l1tOcmeu, o6pa30BaHHOrO rrapHOH Ta30BOH KOCTblO.

AnamoMuR u <Pu3uonoiuR •1e1w6eKa

396

Ta30BaH KOCTb .n:o

12-14

J1CT COCTOHT lI3 COC.[\HHel!HblX XpHmOM Tpex OT-

.[\eJibHhlX KOCTeli: uo.n;B3/l.OIIIHOH, JI06Kosoi1 lI ce.na.attmnoii:. Cpametttthre TeJia 3T.IX KOCTett o6pa3yIOT Bila,IJ;IIHY JJ;JIH ro.10BKH 6e.upeHHoii KOCTlI. Ta3
npe,ll;CTaBJIHCT co6oii: KOCTHOe KOJihl~O, BHYTPH KOToporo HaXOD;lITCH no.10CTh
(rroJIOCTh Ta3a), co,nepmamaH BHyTpeHHHe opranhr. Caa,n;H KOCTH Ta3a coq.1eHHIOTCH c KpecTl~OM, crrepe.n:u --- .n;pyr c ,npyroM, o6pa3yH KOCTHhlH Ta3.

2.

CKeJieT ceo6oiJnou 1-1u.HCneu KOIU''l1t0cmu cocTaBJIHIOT:
Be.n:pettHaH KOCTh, caMaH 60J1hmaH H .n;JIHHIIaH Tpy6liaTaH KOCTh. Ha ee
BepxHeM Kouue pacnoJiaraeTCH rOJIOBKa, KOTOpaH COlfJieHHeTCH c TR30BOH KOCThIO. HmKHHii KOHeU 6e.n:peuHoii KOCTH yToJimeH. Ott tteceT Ha
ce6e ,ll;Ba B03BbIIIIeHHH -

Mhime.JIKa, IlplI IIOMOmH KOTOphIX 6e.n;peHHaH

KOCTh COlfJieHHeTCH c 60JihIIIe6epu;oBOH KOCTbIO lI Ha.[\KOJieHHlIKOM. Ha.n:KOJieHHlIK -

cocTaBHaH lfaCTh KOJieHHoro cycTaBa. Ha.n:KoJieHHHK rrpe)l,-

cTaBJIHeT co6oii: KpyrrHyIO cecaMOBHAHYIO ynJiomeHHYIO KOCTh OKpyrJIOH
cpOpMhl.
KocTn roJieHn: 6oJihme6epu;oBaH H MaJio6epu;oBaH. 3To .ll;JIHHHhie Tpy6lJaThie KOCTH. BOJihIIIe6epu;oBaH KOCTh TOJime MaJio6epuoBOH. BepXHlIH
ee Kotteu MaccnBHhrii:, To.1cThIH. Ott cJiymnT .n;JIH coe.n:ttHeHHH c MhIII~eJI
KaMu 6e.n:peHHOH KOCTH H c roJIOBKOH MaJio6epu;oBOH KOCTH. Hnm1mii:
KOHeu rrpO.ll;OJIJKaeTCH B YIIJIOU~eHHbIH BblpOCT -

Me.n;ttaJibHYIO .10)1,bl)R-

Ky. MaJio6epu;oBaH KOCTb TOHKaH, )l,JIHHHaH, c yToJimeHHbIMH KOHUaMlI, JieJKHT c6oKy (JiaTepaJibHO) OT 60JihIIIe6epu;oBOH KOCTH. BepXHHH
KOHeu 06pa3yeT cycTaB BMecTe c 60JiblllOH KOCTbIO. Hn.muttii IWHeI( 3aKaHlfHBaeTCH JiaTepaJihHOH JIO,ll;bIJKKOH -

o6paayeTCH roJieHOCTOI!HbIH

cycTaB.
KocTH CTOilbl
pJI,na,

3

5

BKJIIOqaIOT:

KOCTeH IIJIIOCHbl 11

7 KOCTeH npe.n:rrJIIOCHbI, paCIIOJIO)KeHHh!X B .ll;Ba
14 cpaJiaHr IlaJihll;eB (lfeTbipe IlaJibUa lIMeIOT no

cpaJiaHrlI, 60JiblllOH naJieu -

2) -

KOPOTKHe, Tpy6qaThie.

KocTH IIO.f!Ca lI CB060.ll;HOH HlI)KHeii: KOHelJHOCTlI 6oJiee MaCClIBHbie H TOJICTbie, lfeM Bepxtteii KOHelfHOCTlI, TaK KaK OHH npncnoco6.1eHbl HeCTlI Ha ce6e
THJKeCTb BCero TeJia.

AKTHBHa.fl lfaCTh onoptto-.n:BnraTeJihHoro annapaTa -

MhIIIIUhI, KOTOphre

npnKpeTIJIJIIOTC.fl K KOCTHM cKeJieTa.
<l>yHKUHH MbIIIIU:
y,nep.mHB8IOT IIOJIOJKeHHe TeJia lI ero lf8CTeH B rrpocTpaHCTBe; o6ecnelfHBalOT nepe.n:Bnmettne TeJia (6er, xo.n:b6a n .n:pyrne BH.ll;hI ABHJKeHnii:), nepeMemaIOT 1laCTlI TeJia .n:pyr OTHOCHTeJihHO .n:pyra;
onpe.n:emIIOT cpopMy n pa3Mephr TeJia; aamumaroT BHyTpeHHHe opraHbI;
ocymeCTBJI.f!IOT ,ll;bIXaTeJibHbie lI l'JIOTaTeJibHhie ,ll;BH:>KeHHH; yqacTBYIOT
B apTHKYJIHUlIH peqn H qmpMHpOBaHHlI MlIMHKH;

Mbtuu11,1-WH

cucmeMa

397

Bbipa6aThIBal<n Teruo; npeo6pa3yIOT XHMH'ICCKym ::mepr1110 B MPXaim1recKyIO.

B Te.1e l.J:Cj!QBeKa HaCl.J:HTbIBalOT OKOJIO 600 :vILIIIIU. 06u~a.H
17 -18 JieT ii,ocTttraeT 40 °;;., y rromrr:IhlX
crropTCl\teHOB 50 '1'1 MaCCbl TC.'!a.

MYCKyJ1wryph1 B
y

Macca CKe:JeTJroi:i

.1m;~ei'r --

30

'.'!,.,

OcHOBHhre c}lyHKUttOHaJ1hHhie CBOHCTBa :vrhrmu:
B036y!J;HMOCTh -

crroco6HOCTb 6hICTpO OTBC'1aTb Ha J(CHCTBHC pa3µ;pa-

:iKIITeJIR B036ym;:i:eHIIeM, B pe3yJihTaTe qero MbIIIIUa crroco6Ha COKpaIQaThCH;
rrpOBO/];HMOCTb -

crroco6HOCTL K rrpoBe.n;eHH!O B036ym11;eHHH OT HepBHblX

OKOHqaHHH 11;0 COKpaTIITCJ1bHblX CTPYKTYP MhlilleqHbIX BOJIOKOH;
COKpaTHMOCTb -

cnoco6HOCTb K COKpaIQeHII!O, K yKopoqeHHIO IIJIH II3-

MeHeHII!O HanpameHIIH;
:JJiaCTIIqHQCTh -

crroco6HOCTb rrpIIHHMaTh rrpemttee IlOJIOJKCHHe IIOCJie

pacT .HmeHII.H.
CTpoeHHe MhIIIID;hl

B Mhimue pa3JinqaIOT 6pIOIIIKO,

cocTO.HIQee II3 rrorrepeqHo-rroJiocaToi1 TKaHH

( COKpaIQaIOIQa.HC.ff qacTb ), I1 cyXO)KIIJibHbie KOH II bl ( cyxO:iKHJIII.ff ), o6ecrrequBa!OIQHe rrpHKpem:retttte K KOCT.HM CKeJieTa, KOme ll opraHaM. IlpOKCHMaJibHOe
cyxomttJIHe (HaXO/];IITCH 6-'rnme K cpe.n;IIHHOH OCH TeJia) HOCHT Ha3BaHne «r0.'10BKa»,

!J;IICTaJibHOe cyxomIIJ:rne

o6o3Ha'1aIOT

TepMIIHOM

«XBOCT».

ll HepBbl BXO,IJ;.HT B MbIIIIIIY c ee BHyTpeHHeH CTOpOHbl. 3.n;ech

ApTepHH

me H3 MbillIULI Bbl-

XO,ll;HT BeHhI I1 JIIIMc}JaTMqecKne cocy,ll;hl. MbIIIIIIa COCTOHT II3 nyqKOB norrepe'lHO-IlOJIOCaThlX BOJIOKOH, o6'helJ,HHeHHbIX coe/];HHHTeJILHOTKaHHLIMII o6oJIOqKaMl1, II csepxy IlOKpbITa rr.10THOH <PacuIIeH.
Pa3JittqaIOT 11;sa TIIrra Mhilllu:

1. I'JtaOKUe

MblUlU,bl -

o6pa30BaHbI rJia,ll;KOH MbIIIIeqHOH TKaHhID, COCTaB-

JI.HIDT CTeHKH BHyTpeHHIIX opraHOB II KpOBeHOCHblX cocy.n;on, IIHHepBIIpyl()TC.ff BOJIOKHaMII BereTaTIIBHOH HepBHoii cIIcTeMhI. CoKpauwHtt.ff HeJipoII3BOJ1LHhie, Me,n;JieHHbie, xapaKTepII3YIDTC.ff HII3KOH YTOMJUieMOCTh!O.

2.

llonepe'lno-no:wcambte MbtUJU,bt (cKeJieTHhie) -

o6pa30BaHhI nonepeq-

Ho-noJiocaToii Mbllllel.J:HOH TKaHbl() (cKeJieTHbie, MHMHqeCKIIe), UHHCpBIIpyIOTCH

BOJIOKHaMII COMaTII'1eCKOH

HepBHOH CHCTCMbl.

CoKpaIQeHH.ff

npOII3BOJibHbie' 6LICTphre.
Ilpu coKpaIQeHIIII MhrIIIua yTO.JIIQaeTc.H u yKopaqIIBaeTC.H. KamlJ,a}I MhIIIIua
orrpe11;eJI.HeT ,D;BttmeHIIe TOJibKO B OlJ,HOM HanpaBJieHHH.

MbIIIII~hr sepeTettoo6pa3HOM II JieHTOBII.D;HOM <PopMhI MoryT 6hITh 11;JittHHhIMtt
HJIH K.OpOTKIIMH, HMeTb 0.D;HY IIJIH HeCKOJihKO I'OJIOBOK. JleHTOBH,D;Hbie MhlIIIIIhl
yqacTBYIDT B o6pa3oBaHHII cTeHOK 6pIOIIIHOM II rpy,n;aoli rro.1ocTeiJ:. MbIIIIULI,
IlY'IKII KOTOpblX

npMKP~IIJ'I.ffIDTC.ff

K cyxomHJIIIl() c O,D;HOM CTOpOHbl, Ha3bIBaIDT-

C.ff O,D;HOilepIICTbIMH, c .D;Byx CTOPOH -

,n;ByrrepIICTbIMII, c HCCKOJihKIIX CTOPOH -

MHOrorrepIICTbIMII. IlepIICTble MblllIUhl pacnoJiaraIOTC.ff B OCHOBHOM Ha HII)R.HIIX KOHeqHOCT.ffX (pIIC.

150).

398

AnamOMUH u rpu3uo1ioiuH

'le.~.o6eKa

25

9
20

14

P111c. 150.

OcHOBHb1e MblWL\bl 4enoseKa:

cL1eB11AHafl;

4-

6onbWafl rpyAHafl;

MblWL(a nne4 (611L1enc);

5-

1-

2- MV1MV14eCK11e; 3 - rpyA11Ho-Krn04V14Ho-co6 - KOCafl MblWL(a )f(VIBOra; 7 -AByrnasafl
V1 nMbL\8B: 9- MblWL(bl K11crn; 10 - crpoe1Hafl; 11 - nopTHfl:-K-

>KesaTenbHb1e;

npflMafl MblWL(a )f(VlBOTa;

8- crn6aren11 3anf!CTbfl

Hafl; 12- npflMafl 6eApa; 13 - 4erb1pexrna0afl MblWL\a; 14 -

11KpOHO)l(Hafl; 15 -

nepeAHflfl 6onbwe6epL10Bafl:

16- KaM6anOBV1AHafl; 17 -- MblWL\bl cronb1; 18- nflT04HOe (ax11nnoso) cyxo)l(11n11e; 19-Asyrnasafl 6eApa;

20- nonycyxo)l(VlnbHafl; 21 MblWL(a nne4a (TPV1L1enc); 24 -

6onbwafl flfOAV14Hafl;

22-- pa3rn6aren113anflCTbfl11 nanbl\8B; 23 - rpexrnasafl
25 -A8nbTOBV1AH3fl; 26 - rpaneL(V18BV1AH3fi

WV1P04ae1waf! MblWL(a CnV1Hbl;

M b£tue<tna1l cue me Ma

399
KJiaccmtm1rnu,uH CKe;1eTuL1x MhillIU.

no CTpOeHHKl
Mbillle'!HbIX
BepeTeHoo6pa3HhH~

JlcHTOBH11Hble
KopoTKHe
,Il;.1HHHbie
IIIttpoKHe
PoM6oBH.UHhre
3y6'1aThie
Kpyr.1L1e
Ksa1waTHb1e
,TJ;Byr JJaBbJe
TpexrJiaBbJe
qeTb1pexrJiaBbIC
,Il;By6pIOIIIJ!b!C

: no l!0,10)KeHHIO
Ilo
: pacuo:10meHHKl
B Te:w

,O.u;norrepncThIC IlonepxHOCT.,Il;BynepncTbie
HbJe
·MHoro11e:LTJy6oKne
pnCTbie
!IIpH:v!b!C
KpyrOBbIC
!KocbIC
(IWJibUeo6pa3Hb!C)

Mb1mrlbI
Ty,10Bmqa
MbiillI(bI
rOJIOBbl
iMbiillUbI
IBepxuux n
lttnmunx
1
KOHC'IHOCTCH

flo CjlyHKI(HHM

Crn6aTe.1n
Pa3ru6aTc.1n
Ilp1rno.u;m1v1e
0TBOl(HllllfC
CmuMaTeJrn
(cc}>UHKTepbi)
BpanlaTeJrn
I 110,QHUMa!Oll(lle
lorrycKa1011ine

I

Ilo BbIIIOJIH.HeMbIM QJYHKI.1;11.HM pa3JII1'IalOT .ll:hIXaTeJibHbie, JKeBaTe.'lbHhie,
MllMM'leCKMe MbllllUbl. Ilo ,11;e:i1:CTBMIO Ha cycTaBhI CKeJieTa BhI,ll;eJI.HIOT MblllIUhI:
cru6a TeJIM u pa3ru6a Te,1u; OTB0,11;.HIIJ;Me M rrpttB0,11;.HIIJ;Me; BpaIIJ;aTe.1u.

OC1-t061tbte .MbtWU,bt myJ10Buw,a: TparreuueBM,ll;Ha.H MhIIllua, Illttpoqa:i1:ma.H
MhIIllua crrttHhI, 60.rrhllla.H rpy,11;Ha.H MhIIllua, MaJia.H rpy,11;Ha.H MhIIllua, HapymHhie ll BHyTpeHHMe Mempe6epHbie Mbllllllhl, ,ll;MatjJparMa, HapymHa.H KOCa.H
ll rrp.HMa.H MblllIUbI JKMBOTa, IIO,ll;KOJKHa.H MbIIllua meu, rpy,11;MHO-KJIIOlfl1'lHO-COcueBl1,1J;Ha.H MhIIlIUa.

M btWl{bt

io.1t06bt: M11M11tfecK11e (Ha,ll;'leperrHa.H MhIIlIIJ;a, MhIIllua rop.neuoB, Kpyro-

Ba.H MhIIllua rJia3a, KpyroBa.H MhIIllua pTa, MhIIllua c:vrexa) 11 meBaTe.1hHhie (meBaTe,1hHa.H ll Bl1CO'lHa.H MblllIUhl, :vre,11;11aJihHa.H 11 JiaTepaJibHa.H Kph!JIOB11,ll;Hhie MblllIUbl).

MbtWU,bt Bepx1tPi.1 1C01tel/.1tocmu: rrJielfeBoro rro.Hca (,11;eJibTOBM,11;Ha.H MhIIlIUa)
ll CB060,ll;HOH BepxHeH KOHe'lHOCT11 (,11;ByrJiaBa.H ll TpexrJiaBa.H MbIII.IUbl IIJielfa).

MbtWU,bL 1tUJ1C1teu 1C01te•mocmu: Ta3oBoro rro.Hca (6oJibllla.H, cpe,11;H.H.H 11 MaJiaa .HI'O,ll;lllfHbie MhlIIIUhl) ll CB060,ll;HOH HllJKHeH KOHe'lHOCT11 ('IeTbipexrJiaBa.H
MblllIUa 6e,11;pa, rropTH.HIBHa.H MbIIllua 11 TpexrJiaBa.H MbIIllua roJieHM: MKpOHOJKHa.H ll KaM6aJIOBll,ll;Ha.H).
BcrroMoraTeJihHhie arrrrapaThI MhIIllu:

<Pacu,uu -

coe,ll;MHMTeJibHOTKaHHhie lfeXJihI, KOTOpbie OKpymaIOT Kam,11;y10

llJlll rpyrrrry MblllIU, pa3,11;eJI.HIOT MbIIllelJHbie CJIOM. <l>yHKUllll: ycTpaH.HIOT TpeHlle MhIIllu .npyr o ,11;pyra, cJiymaT orropo:i1: rrpu coKpaIIJ;eHMM, orpaHMlfttBaIOT
pacrrpocTpaHeuue KpOBM u BocrraJittTeJibHoro rrpouecca.

Cu1to6uaJ1b1tb1e 6J1a?a.11uw,a o6pa3oBaHhI cttHoBua,1bHOH o6oJIO'lKOH, o,n;Ha
IIJiaCTMHKa KOTopo:i1: rrpo'lHO cpacTaeTC.H c cyxomttJitteM. Cu1toBuaJ1bHbte M CJIU-

3ucmbte cy.MICU ycTpaH.HIOT TpeHMe cyxomMJillll ll MblllIU 0 KOCTHbie BbICTYIIhl
MJIM ,11;pyrue o6pa3oBaHM.H.

AnamoMUJl u </JUJuo1wzuJ1

400

'lCJWeeKa

HepBHan CHCTeMa
Hep6HaJl cucme.wa -

rpynna oprattoB, o6pa3oBaHHhIX ttepBHOH TKaHhIO,

o6ecne'IHBaIOIIJ,HX peryJum;i1ro ue.TJOCTHoro opraHH3Ma.

3Ha1.Je1tue HepBttou cucmeJ111>1:
o6ecne'IeHIIe cor.rracoBaHHOH pa60TbI KJICTOK TKaHei1, opraHOB JI CHCTeM
opraHH3Ma B e.n;11noe IJ;cJ:roe;
pery.'IJII~lHI

,n;eJITeJibHOCTlI BCeX opraHOB lI ClICTeM;

ocyIIJ,eCTB.>eHtte CBJI3H oprattl13Ma c BHenrne:i1 cpeJJ,o:i1, np11cnoco6.TJem1e
K 6bICTPO MeHJIIOIIJ,lIMCJI yc.'IOBHJIM :IKII3HlI;
MaTep11aJibHaJI OCHOBa C03HaTe.'IhHOH ,11,eJITeJibHOCTlI 'IeJIOBeKa:

pe•Ih,

MblIIIJieHIIe' IIOBe;:i:eH11e.

RJiaccm}m1mu,H11 11epB11oii: CHCTe:11L1

I.

AttaT0Mtt'Iec1we ,11;eJiettne

[

11,eHTpa.'IbHaH HCpBHaH
CliCTeMa

IIepmjJepH'ICCKaH !!CpBHaH CHCTCMa

/

I'ommnoil Mo~

II.

Cmurnoil Moar

<l>ymm;ttottaJihHoe ,11;eJienne

CoMaT11'Iec1<aH

ABTOH0:11HaH (nereTaTHBIIaH)

IlapacHMrraTH'IecKaH

I

HepBHGR mKGHb COCTOMT ll3 HeHpoHOB 11 BCIIOMoraTeJihHhIX KJie'l'OK (r.'llIaJihHhJe KJieTKM, tte:ITporJI11JI; nrnaHHOBCKMe KJieTKM). HepBHhie KJieTKH crroco6tthI BOCilpHHlIMaTh pa3,n;pamen11e, IIplIXO,[(lITh B COCTOJIHlie B036ym,n;eHirn,

l/ep61tafl cucme.tia

401

BbJpa6aTbll33Tb H nepe.uaBa'fh HepBHbJe 11!VlllY-1hChl. llepBHbJ(' i-:.1en~11, cuocooHble CH!l'fe3IIpOBaTb 6tto:rorttt.JeCKI1 aKTI1BHbl(' BeW:eCTBa (Me,IJJia'ropbl), Ha3b!-

11eilpuCP1(pemop1tblMU.
Heupo2:1uJ1 ocyw:ecTB,'!aeT onopHy10 qlyHKu;ino, nttTaTe,1bHy10 11 cnerLmlm-

BaIOTCa

qeCKYIO 3arn;HTHYIO cpyHKU,Hio. HepBHhie K.1eTKn pa3HhIX oT.ne:roB Hep1rnoii CHcTeMhI 0Tm1qa10Tca no pa3:\lepaM II q>opMe.

B

Kam.uoi-i: ttepBttoiI rrneTKe pa3mI-

t.JaIOT ·re.10, OTPOCTKII II HepBHbie OKOWiaHl1 (pHC.

151).

Cyw;ecTsyeT ;:i:sa su.ua OTpocTKos:

1. AKco1t -

.UJJIIHHhIH oTpocToK, KOTOPhIH npono;i:MT ttepBHbie ttMnyJib -

CbI OT TeJia HCPBHbIX K:IeTOK H nepe.uaeT MX Ha .upyroii HettpOH 11,·rn Ha
KJieTKM pa6oTaroui;ero opraua -

MhIUJ1I,bI, me,'!e3bI. Bee HeiipoHbI ttMeroT

o;urn aKcott.

2. /{eHiJpumbt -

CMJJbHO BCTBHW:Meca, MOJKeT 6bITh OT O.UHOro

,D;O

15 .ueH-

.UPMTOB. 0Htt nposo.uaT ttepBttbie MMny,'IhChI K Te,'Iy ttepnttoii rrneTKII.
Ilo KOJJIIt.JeCTBY OTpOCTKOB HeiipOHbl pa3-

152)

yuunOJLJl,PflbU' (K,1enm
6unOJLRpflblf (KJieTKlI
.UBYMH 0TpOCTKaM11), MY.!lbmuno.u1pflbte (TpII
II 6onee OTpocTKa), nceaiJonoJLJlpflbte dByompoc'lambte (KJJeTKM qyBCTBMTe,'!bHblX y3,'!0B, y KO-

p,eJJRIOTC.ff (plIC.

c
c

0,11,HHM

Ha

OTpOCTKOM),

TOpblX 11,Ba OTPOCTKa OTXO,[\.ffT OT BblpOCTa TeJJa).

Tunbt ueupoHOB (no cpyHKI1MHM)
lfyBCTBIITeJibHbie (peu;enTOpHbie,
peHTHbie) -

acp<l>e-

BOCTIPIIHl1M3IOT pa3.upame-

4

HHe, npOBO,[\HT lIHcpOpMaIIMIO OT noBepxHOCTM TeJJa

lIJJlI

BHyTpeHHMX opraHOB

B M03r.

6

Accou;11aTIIBHhie (BcTaBoqHbre) -

ocyw;e-

7

CTBJJaIOT CBH3h MeJKIJ,y qyBCTBMTb'JhHbIMM
I1 .ll;BMraTeJJbHhIMH KJJCTKaMH,

HX TeJra

H OTpOCTKM He BhIXO.ll.HT 3a npe,ueJibl l~HC;
COCTaBJJ.ffIOT OCHOBHYIO Macey HelipOHOB
UHC.
,Il;BttraTeJibHbie (acpcpepeHTHbie) BbIHOCHru;ue -

npOBO,ZJ;JIT HepBHhIH HMTIYJJbC OT

l'OJJOBHOro M CillIHHOI'O M03ra K pa6oTa:IO-

P11c. 151. CxeMa cTpoeH1M1

W:I1M oprattaM.

ABll1raTellbHOro (MYllbT111nom1p-

HepBHaH KJieTKa oKpymetta n;taaJnamu'tecKou MeM6paflOU, KOTopaH peryJIIIpyeT TpaHc-

HOro) HeliipoHa: I -AeHAPVlf:

nopT Bew;ecTB, yqaCTByeT B o6pa30BaHIIlI KOH-

KlleTKVI; 4 -

TaKTOB (CIIHancoB) Mem.uy HelipoHaMM, RBJJH-

5 -- aKCOH; 6 - M8elli11HOBa>l 060-

eTCH

MeCTOM

oJICKTp:uqeCKOH

3KTMBHOCTl1

2 -- HApo: 3 -- TellO

nOYKa;

HepBHOVl

WBaHHOBCKaH KJleTKa;

7 - nepeXBaT PaHsbe: 8 --

H .ueiiCTBHa 6HOJJOJ'HqecKM aKTHBHhlX Bew;eCTB

HanpasneHvie HepsHoro viMnynb-

(Me,11,HaTopoB).

ca;

9-

HepBHOe OKOHYaH111e

4U2

Anamo.Mull u <}Ju3UOllOZUJl •u?.rtoBe/Ca

Hep6Hbte eo.1wn:1ta

-

&TO OTpOCTKH

HepBHhIX K.rreTOK, fIOKpbIThie o6oJIO'IKa-

.·

MH.
y11uno.1J1p11btu

1
.

B

3aBHCHMOCTH OT cTpoeHHH o6oJio-

<IeK pa3JIH'laIOT: MJllWmHblP (MHeJIHHOBhle) H 6e.JMJl1WmHbte (6e3MJie,'JHHOBbie)

.

HepBHhie BOo'IOKHa •

-

Ocesoi1 utt.'IMHJJ:P HepBHhIX BOJIOKOH
COCTOJ.IT H3 UHTOII,'la3MhI II Heiipoqrn.'la-

nceeiioy llU nol!.Rpnbtu

MeHTOB, CHapylKII IlOKpbIT MeM6paHOH,
06ecneq1rna10rueii npose.n;eHIIe ttepBHoro
KJ!l'nl!W

DypKUHbC
6unOJ!JlpHblU

HMITYJibCa.
MtteJIHHOBhJe HepBHhie BOJIOKHa TOJI-

(Ii .WOJJI(!'~ Ke)

ru;e, 'leM 6e3MHeJIJIHOBhie, TaK KaK B 060-

PMC. 152.

JIO'IKe JIMeeTC.H MIIeJIHHOBhIM CJIOM, co-

THnblH8MPOHOB

CTO.HII(JIH

(no CTpOeHHIO)

20-30

H3

CJIOeB MeM6paHbI

UIBaHHOBCKoli: KJieTKH. BoJihUIOe co,n;epma1rne cpoccpoJIHIIJI.UOB B o60JIO'IKe o6ecne'IHBaeT ;rneKTptttieCKYIO JI30JIHUHIO,
.ueJia.H B03MOIKHbIM npOXOiK,Zlemi:e HMilYJihCa.

y

MaJieHbKHX ,ZleTeli o6oJIO'IKH

eru;e He IIOJIHOCThIO ccpopMJipoBaHhI, Il03TOMY y HHX Ha6mo,n;aeTC.H 3a:vre,n;JieHHaH peaKUli.R JI IlJIOXa.H KOOp,ll;HHau1rn ,ll;BHiKeHHM. CKopOCTb npose.n;eHH.H HepBHoro JIMIIYJihCa B MJieJIJIHOBhIX 6hICTpee

(1-2

(5-120

M/C), <IeM B 6e3MlieJIJIHOBhIX

M/C).

B rrpo:vremyTKe MeiK,n;y JJ;BYM.H coce,n;HHMH urnaHHOBCKJIMH KJieTKaMJI MJieJinHOBa.H o6oJIO'-IKa HepBHoro BOJIOKHa rrpephrnaeTca, JI 06pa3y10Tca rrepexsaTbI PanBhe. B HHX npOJICXO,L(HT ,ll;BHiKeHne HOHOB, KOTopoe BhI3bIBaeT «CKa'IKH» IIOTeHuna.ii:a ,n;e:HcTBHH OT o,n;Horo nepexBaTa

.n;o .n;pyroro B

xo.n;e rrpose.n;eHJI.R JIMIIYJihCa.

B 3aBHCHMOCTH OT BhIITOJIHHeMOM cpyHKIVUI pa3mI<IaroT Tpn Tnna Hepsos:
<JBuiameJtbHbte (u,eHmpo6eJ1CHbte) ttep6bt -

co,n;epmaT oTpocTKH ,n;snra-

TeJihHhIX HettpOHOB;
'l.lJBCmeumeJlbHbte ( u,eHmpocmpe.lfwme.!lbHbte) HPp6bt -

co,n;epmaT oTpo-

CTKH 'IYBCTBHTCJihHbIX HeiipoHOB;
C.MPWGHHbte Hep6bt, B COCTaB KOTOpbIX BXO,ll;JIT ll qyBCTBHTeJihHbIM II ,L(BllraTeJibHhiii HepBHbie BOJIOKHa.
HepBHhie BOJIOKHa 3aKaH'IHBaIOTCJI HepBHhIMH OKOH'laHHJIMH B pa60'lHX opraHax H meJie3ax.
<Pu.JUOJtoZU'lPCICUe C60UCm6a HPp6HO<!O 60.!WKHa:

1.

Bo36YAHMOCTh -

cnoco6HocTh socrrpttHHMaTh pa3ApameHHJI

n:

OTBe'laTh

Ha HHX.

2.
3.

IlpoBOAHMOCTh -

cnoco6HOCTh nepeAaBaTh B036y1K,n;eHHe.

Il;eJIOCTHOCTb HepBHOro BOJIOKHa, T. e. B036YIKAeHHe pacrrpocTpaH.HeTCfl
no HepBHOMY BO.'IOKHY JIHlllh npII coxpaHeHIIlf ero aHaTOMH'leCKOM H

qrn-

3HOJIOI'H'leCKOH ueJIOCTHOCTH.

4•

.IJ:sycTopoHHee rrpose,[);ettne so36y1K,n;emrn HaeTc.H B o6e CTOpOHhl nepBHOI'O BOJIOKHa.

so36y1K,[J;eHne pacrrpocTpa-

[Jepeda 'la 6n3{JyJ1CiJe1tUR nep61w1i 1rnenuwii

5.

403

l1aO.'IHpOB3Hl-lOe uponc.11.e11ne HepBHOI'O HMIIy.:1bca -- - BO;i<)~l)l-(;wmH~ HC

HaXOJUlll~H('CH napa:1-

:vrOJ-1\eT nepcifTH W-l ;wyrue HepBHh!e B0.'10l{Ha,
JICJ!bHO B Oll;HOM H TOM me HCPBHOM CTBO.'Je.

6.

0THOCHTCJibHaR

11eyTo:vrnHe:vroCTb 11cps11oro no.10KHa

---

cnoco6HocTh

K npone.uemrro B036ym.zi;emur name np11 .11..1111C.'1hHOM HenpqJbIBHo:vr pa 3 .11.PaJKCHH11.

Ilepeµ:aqa B030Yll\'.11,CHHH uepBHOH K.'leTiwii
Tiepc.11.a 1 ra B036ym)l,CHirn c He]JBHOrO BOJIOKHa ocyn(CCTB.1HeTCH B crrnance -cneu;Ha.'1113IIpOBaHHOH

30He KOHTaKTa

Mem.uy Hettpo11aM11 (n:m

Hcl'ipoHOM)

II ii;pyrH:vrII B036y,n;H:vrhIMH o6pa30B3HHHMl1 (MbllliI(a, me.'Ie3a).
B

cmrance (pnc.

YTOollIJ;CHHH,

153)

pa3o111'-IaIOT npec11HaTITH'ICCKHiI OT;ie.1 (IWHIWBbie

11.1JI 6ml.11IKH

3KCOHa) H

IIOCTCl1H3IITIIqecl{IIH OT;J,eJI (y•rnCTOI{

MeM6paHhI .n;ctt,ll,pHTa .n:pyroro tteii:potta m160 MhillieqHott HJm

n-ce.1ea11cToii

K.1eTKII). I1MeIOTCH pa3HbJe BH,ll,hl CHHancos: c XHMII'ICCKH'.VI' :3c1CETpIJtieCI\HM
MeXaHH3MO:vr ncpe}(a'lII B036yJK,ll,CHH.H' a T3I<Jl\e CMemaHHbie CIIH31!CbI.

CuHancb1 c

:XUMU'U'CKU!ft Mexa1-tu.JN0.~1

qacTh cnnarrTIIqecKoro arruapaTa
Cl1H3IlTII'!eCK3H w;eJlb llIHPOI\a.H

UHC

nepeaa'iu

cocTaBJlHlOT 60.1hIIIYIO

Y

x11M11qecKHX c11Ha11coB

'ICJlOBCKa.

(10-20

HM) II Il03TOMY 06.1a,i:i:aeT 60J!hI1IIIM

corrpoTHBJleHIIeM, nperr}ITCTBYIOUJ;IIM Herrocpe.n:cTBeHHOMY uepexo.n:y llOTCH-

l~HaJia ,ll,CHCTBHH c npe- Ha IlOCTCHHaIITHqecKyIO Me:vr6patty. JJ,.1.H 3TOI'O HCfl0J1h3YCTCH XH:VlII'IeCI<lfH nocpCll;HHK --- Me,llHRTOp, Bb1pa6HThIBaCMhlH TC.'JOM Hei\pona I1 orry;:ia Tp~lHCIIOPTHpye:vrblll B OKOl!qaHIIH ero aKCOHa, r.11.e
OH HaKaIIJIHB3CTCH B ny3hiphKax B,ll,OJ!h BHYTPCHHCH IIOBepX!WCTl1 rrpec11HaIITIIqecKol'i ;vi:eM6panbr. Kor.n:a no.n:
.uel'iCTBJ1eM

npIIlliCl(lllCI'O

H3II1'IIqecKOH
HMny.'Ihca
u;Jia,

MeM6paHe

ttacTynaeT

MC,ll,U3TOphl

rry3hipbKOB
3K30IJ;IIT03a
ru;e.'Ih.

I{

I

rrpecn-

I

ttepnttoro
-[)

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CHHaIITIIqecKJIX

Bb16pacbrnaIOTCH nyTeM
B

MC:iKCHHaIITII'IeCKYIO

06H3aTeolhHhIM

yCJIOBIIeM

l(JIH nponecca Bbrcno6om.n:eHIIJI MeJ(IIaTopa
JIJICTCH

HepBHbIM

HMIIy.'IbCOM

npHCYTCTBHe

ca~

.HB-

BHYTPII

6JI.Hl1IKH aKCOIIOB.
BhrCB06om11enHhrii Me.11.11aTop csH3hIBaeTCH c peu;ef!TOpHblMH 6e.1KaMH

Pi-ic. 153.

IIOCTCHHaIITH'ICCKOH Me:vr6paHhI, BbI-

Cl'1Hanca: 1 -

3bIBaH noBhI11Ien11e ec rrpom1u;aeMoc-

2--

TH

,rwa

Na-'-. Bo3HIII<HOBettnc rroToKa

CxeMa cTpoeHWl M8)J(HeC'ipoHHOrD

6poc M8AV13TOpa;

5-

Na+ 113 CIIHanT11•1ecI<ol'i ru;eJm qepe3

ropa;

IIOCTCIIH3II1'IIqecKyIO

nornOLl\8HV18;

MCM6pauy

se-

.n:eT K ee .n:enOJ1HpII3au;1111 II BhI3bIBaeT

npecV1HanTV1YecKafl MeM6paHa;

nOCTCV1HanTV1YecKafl M8M6paHa:

4 -

CVIHanrnYecKafl Ll\ens:

7-

MVITaXOHAP11fl;

3- Bbl-

P8L\8nTOpb1 AJlfl M8AV1a-

6-

o6parnoe

CV1HanTV1Y8CKV1C1 ny3blpeK:

9- aKCOH

8-

(HepBH08 BOJIOKHO)

404

A11amoMuR u <Jluauu.iotuR 'le.1w6eica

reHepaI]J1!0 8(1;l6y.uneJihl!OfO JIOt:TCIIIIHIITHlJCCKOl'O IlOTeHI~l1aJra -- 11po11cxoJ~HT

K

nepe,n,a 1c1.n 11cpBHOI'O n:vrny :1 b<'H.

xu31u•1ec1Cuc11 .!fzec)uamopa.ll onuwcHmcR: aueT11.1xoJIHH, Hopaµ:petta.111H,

cepOTOHHH; a:vrHHOI\I:IC:JOTbI -

r.'IyTbMIIHOBaH, acnapar11HOBaH, rJII11Il1H, y-aMII-

HOMaC.'IJIHaH lf .upyr11e BeU~CCTBa.
J{eiiCTBJ1e HeI<OTOph!X H.L(OB OCHOBauo Ha OJIOKHPOBaHHII CHHaTITH<JeCKOII nepeµ:arrn B o6ac\CTl1 HepBHO-MhiillP'IHOro coeJJ;11Hemrn:. TaK, ,n,elicTBIIe H,n,a Kypape OCllOB8HO Ha CBH3bIBaHHJ1 cro c peuerITOpHhl'.\111 6e:maMI1 IIOCTCHH3IITH'-IeCKOH MeM6patthI, 'ITO npenHTCTByeT B3aHMOJJ;eilcTBHIO c HHM11 aueTHJixoJI1rna.

CuHancbt c Jo1e1Cmpu'lcc1Cu.>1 .'rtexaHUJMOM nepeiJa'lu illIIpOKO pacrrpocTpaneHhI B cep,n,e<rnoii Mblll11Ie, rJra,D,KOii MYCKyJiaType BHyTpeHHHX opraHOB, 3Til1TeJIHaJ!hHOH l1 me.rte3UCTOH TKHHJIX.
narrTIIqecKOII ~CJIH Ma.'Ia

(2--4

B

3.1eKTp11<1.eCKI1X c1rnancax nmpnHa CH-

HM). qepe3 Hee nepCKHHYTbI MOCTHKII l13 6eJI-

KOBbIX qacTm~, o6paayJO~ux csoeo6pa3Hhre KaHa:1b1, cnoco6Hhie nponycKaTh
113 KJICTKH B 1..:JieTKy HeopranIIqecKHe MOHhl H Me.rmue MOJieKyJ'Ih!. B.1aro,n,apH
3TOMY 3JieKTpuqCCKOe conpOTHBJICHIIe B THKOM Cl1Ha11ce o<JeHh Hl13KOe, qTO
II03BOJIHCT npec11nanTII'ICCKOMY IIOTeHUMaJiy pacnpocTpHHH'fhCH Ha TIOCTCl1H8IITHqecKylO KJJeTKy 6e3 yracaHIUI.
OT.1uquH: XIIMil'ICCK11e cuuanchI oT.'IH'IaIOTCH o,n,nocTopottHMM npoBe,n,eHneM OT rrpe- K IIOCTCMH<UICy; B 3J'IeKTpH'18CKOM c1-rnance qame ,D,BYCTOPOHHee
rrpose,n,eHue;

xuM11qecKue c11Hanch1 o6ecne<rnBalOT 1rnK Boa6ym,n,ettne, TaK

l1 TOPMOJKCHIIC IlOCTCUHaIITH<JeCKOro neiipotta, B TO BpeMJJ KaK 3.'ICKTpU'IeCKHe -

TOJibKO BOa6ymµ;eHue.

KoJiuqeCTBO CHHaIICOB Ha OJ];HOM neH:poHe MOIBeT 6bITh OT e,D,MHHU ,D,O neCKOJihKHX Th!CH'I.

CTpyKTyptto-<f>yttKu;HoHaJJLHLie qacTH HepBHoii cHcTeMLI
K ueHTpaJJhHOii: HepBHOH CHCTCMe (IJJIC) OTHOCHT i'0.706/lOU .!fl032, pacrroJIO)KCHI!bIH B IIO.'IOCTH 'Iepena, H cnUH/-lOU MO.Ji!, KOTOpblM JJeJKIIT B TJ03BOHOqHOM
OT,D,eJTe CKe.'IeTa. foJIOBHOH 1l ClllIHHOH M03r COCTOHT 113 ceporo l1 6eJioro Be~ec'f
Ba. Cepoe Be~eCTBO rrpe,D,CTaBJrneT co6oii CKOTIJieHMe Te.'I nepBHblX KJieTOK l1 MX
OTpOCTKOB, a 6e.'Joe -- HepBHh!X BO.TIOKOH, 06pa3y10mux rrpOBOl(HlIJ;He nyTM.
foJIOBHOH l1 CIIHHHOII M03r HMCIOT TPII M03rOBhle o6oJioqKH (p11c.

meepiJaR M03ZOBaR 060JIO'i1Ca (napymHaR) -

154):

rrpO<JHaJI, COCTOHT l13 IIJIOTHOH co-

e,D,MHHTCJibHOU TKaHu; naymuHHaR (cpe,D,HHH); MRZICaR, MJIH cocy,n,HCTaH, 060JIOqKa (BHyTpeHHHH) -

co,n,epIBHT 60JibllIOe KOJrnqecTBO KpOBeHOCHblX cocy-

,D,OB. IlpOHMKaeT BO Bee 6op03,D,bl H JKeJiy,D,O'IKH M03ra, o6pa3yfl B HUX cocy,n,11cTbie CIIJieTeH11H.

<PyHKU,UU Jlt0.J206blX 060JIO'le1C: Jat4um1taR (OT MexaHHqecKMX IIOBpemµ:eHIIH HepBHOH TKaHn); 6apbepHafl (OT rrpOHMKHOBeHHH Ml1Kpo6oB l1 .n;pyrux BC~eCTB B M031'), celCpemopHaR (ceKpeuuH KPOBCHOCHblMH cocy,n,aMII CIIMHHOM03-

rOBOH JKM,D,KOCTH).
Mem,n,y rrayTMHHoii: II MRrKoii o6oJioqKaMII uMeeTcH cy6apaxHou,n,aJihHOe
rrpocTpaHCTBO, 3aIIOJIHCHHOe cnUH/-lOMOJ2060U JICUOICOCmb/O ( JIU/(60pOM ). JlIIK-

Cnu1uwu M03Z

405

6
P111c. 154. CxeMan1YecKoe 111ao6pa)!(eH111e o6onoYeK:

a-

ronoaHoro M03ra, 6 - cn111HHOro Moara.

1 - KOCTb CBOAa Yepena; 2 - rnep,[\as:i 060110'-lKa; 3 - naynl\HHas:i 060110'-lKa; 4 - Ms:irKas:i 60110'-lKa:

5 - no,[\nayrnHHOe npoc1paHCTBO; 6 - cepoe B8li\8CTBO; 7 - 6enoe B8li\8CTBO: 8 -- )!(ll1POB851
TK8Hb C COCYA8Mll1

BOP Bb1pa6aTbIBaeTCH cocy,ll;HCTbIMII CIIJieTeHHHMII )lWJiy,ll;O'-IKOB M03ra.

Ilo

co-

CTaBy noxom Ha TIJia3MY. 06'heM ;noM: JKil,ll;KOCTil y B3pOCJIOro qeJIOBeKa OKOJIO

J50

MJI.

Bhmo.1m1eT cJie,11;y1011~11e c}:>yHKLI;II11:

· 1.
2.
3.

Y'IaCTByeT B o6Mette BeJ:QecTB M03ra.
Co3,11;aeT onpe,11;eJieHHOe .n;aBJieH11e BHYTPII 'Iepena H B no3BOHO'JHOM Katta;:re.
BhmOJIHaeT 3aIQHTHYIO poJih, npe,u;oxpattaJI M03r OT TOJI'IKOB H coTpRcettnM:.

CrrHHHoii M03r
8To ttan6oJiee ,11;peBH11M: OT)J,eJI UHC no3BOHO'IHhIX, noaTOMY ou B 60.11>llleil:
Mepe, 'IeM ,11;pyrne OT,ll;eJibI UHC, coxpaHIIJI qepThI ,11;peBtteM: opratt113au;1111, Bhlpama101Q11ecJI B MeTaMepHOM CTpoeHIIII (pttc.

155).

Cn11HHOH M03r pacnOJIOJKeH B II03BOHO'-IHOM KaHaJie H 11MeeT BH.L\ 6eJJoro T.Hma ,IJ;JIIIHOM OKOJIO

45 CM,

,ll;IIaMeTpOM OKOJIO

roBaTbIM M03r, a BHII3y, Ha ypoBHe

II

1 CM,

BBepxy nepexO,UIIT B npo,UOJ1-

noRCHII'IHoro II03BOHKa, 3aKaH'IIIBaeTcfl

M03I'OBblM KOHyCOM, nepeXO,ll;JIIQIIM B KOH8'IHYIO HHTb. l1MeeT ,ll;Ba YT0Jll.Q8HHR,
o6pa30BaHHbie CKOTIJ18HI18M HepBHhlX KJieTOK, IIHH8PBI1py101.u;Hx KOHe'IHOCTil:
lll8HHoe Il IlORCHil'IHOe; pa3,ll;8JI8H .L\BYMR 6op03,ll;aMII (rrepe.utteif Il aa,ll;Heii) na
rrpaByJO II JieBylO IIOJIOBHHbI.
Y3Killl u;eHTpaJibHbil':I. KaHaJI CIIIIHHOI'O M03ra OKpymeH

cepblJl,l 6ew,ecm60.M

(CKOIIJI8HIIe TeJI H8pBHblX KJ1eTOK) B BII,ll;e 6a6o<mH. Ilepe,ll;HHe «KPhl.'IbR» ceporo
B8IQ8CTBa H33bIBaIOTCJI nepe,n;HIIMil poraMII, 3a,D;Hile «KpbIJihfl» -- 3a,IJ;HHMil poraMII. B rrepe,IJ;HHX porax ceporo BeIQeCTBa ttaxo,11;aTca 'l'eJia .n;BttraTeJihHhIX (IIeHTpo6emHhIX) HeMpOHOB, BOJIOKHa KOTOpblX o6pa3yJOT

nepeiJuue lCOpeuuw

CIIIIH-

HOM031'0BbIX ttepBoB. B 3a,D;HHX porax pacnoJJomeHhI KJJeTKII rrpoMemyTO'lHhIX

AnamoMuR u cpu3uo1wzuR

406

•teJW6e1w

6

12

3

6
PMC.

155. CTpoeHi-ie cni-iHHoro M03ra: a - pacnonm1<eH1-1e cni-iHHOro M03ra s no3BOHOYHOM 1<aHane

(Ha npOAOnbHOM cpe3e); 6 CTOK n03BOHKa; 3 KaHan; 7 -

CTpOeHi-ie cerMeHTa. 1 - Teno n03BOHKa; 2 -

cni-iHHOi-1 M03r; 4 -

6enoe sew,ecrno;

3i!AHl-1i'i KOpewoK; 5 -

HenapHbli'i (OCTl-lCTbli'i) OTpo-

nepeAHl-li'i Kopewot<;

6 - l_\8HTPMbHbli'i

8 - 3aAHi-ie pora; 9 --- 601<osb1e pora: 10- nepeAHi-ie para; 11 - cni-iHHO-

M03roBoi'i y3en; 12- cnl-1HHOM03fOBOi'i Heps

(BcTaBOqHbIX) HeHpOHOB, ocyn.~eCTBJIHIOUJ;HX CB.H3h MeJK.D,Y u;eHTp06eJKHbIMH
ll uenTpOCTpeMHTeJihHhJMH neii:pottaMH.

3aiJHUU 1wpewo1C o6pa3oBaH BOJIOKHa-

Mll qyBCTBHTeJihHbIX (1~eHTpOCTpeMHTeJibHhIX) HettpOHOB, Te.'Ia KOTOpbIX pacnoJiaraIOTC.H B CTIHHHOM03rOBbIX
HJIH qyBCTBHTCJibHbie,

(MeJKTI03BOHKOBbIX) Y3Jiax. qepe3 3a.D,Hlle,

KopeIIIKH

B036ym.n;ettn:e nepe.D,aeTCJI c

B CTIHHHOH M03r, a qepe3 nepe.D,HHe, HJIH ,ll,BHraTeJibHhie, -

nepmpepn:n:

OT CTIHHHOro M03ra

K MbIIIII.J;aM ll .D,pyrn:M opraHaM.

B 601C06blX pozax ceporo Beru;ecTBa cnn:ttttoro M03ra ( OHH pacnoJiaraIOTC.H na
I rpy.D,Horo .D,O II-III no.Hcttn:qttoro cerMeHTa cnn:nnoro M03ra) pac-

ypoBHe OT

eezemamu6Hble RiJpa CHMTI8THqeCKOH HepBHOH CHCTCMbI.
6e.1wzo Be~ecmea cnnnnoro M03ra, Jiemaru;ero no nepnllJepnn: ceporo Beru;ecTBa, o6pa3yIOT npoeoiJa~ue nymu. Ilo oTHM nyT.HM ycTanoJiaraIOTC.H

OcnoBHyIO Macey

HaBJIHBaeTC.H CB.H3b Me:lK.D,Y pa3Jill'IHbIMH qacT.HMH :UHC ll npOXO.D,.HT HMTIYJibCbI B BOCXO.D,HID;eM ll HHCXO.D,HID;eM nanpaBJICHll.HX.
IOT

31

cerMeHT.

I-13

Kam.D,oro

cerMeHTa

B

CTIHHHOM M03re BbI.D,eJIH-

BhIXO.D,.HT

nepe.D,nn:e

n:

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KOpeIIIKH, KOTOpbre CJIHB8IOTC.H ll o6pa3yIOT CMelllaHHbIH CTIHHHOM03rOBOH
nepB (Bcero

31

napa).

<l>yHKl~llll CTIHHHOrO M03ra:

1.

Ocy~ecmeJtettue cnuHHO.M03ZOBblX ( epo)l("iJeHHblX) pe</JJte1Ccoe.

B

cepoM

Beru;ecTBe CTIHHHOro M03ra 3aMbIK8IOTCH peilJJieKTOpHbJe nyTH MHOrHX
.D,BttraTeJibHhIX peaKUHH (KoJiennhr:H peq>JieKc). Cnunnoii: M03r unttepBH-

/'oJWHIWU MO.Jl

407

pyeT BC!O CKeJieTHYIO :vrycKyJrnTypy, I<po:vic MbIIIllI rOJIOBhl, EOTOPLIC l1llIICpBHPYKlTCH 'ICpCIIllhlMII HepBRMH. 3.11,ecb :me paCI!OJ10)KCHhl MHOr11c I~Cll
Tpbl BCI'CTRT!IBHOii HepBHoi1 CHCTeMhI. TaK, B I<pCCTI\OBOM OT;'l;C.'Ie 11axo,ll,HTCH I.LCHTpbI MO'ICHCITYCK3HlUI, /\CWCK3I.LHH II IIO.'IOBOH .11,CHTCJihHOCTH.

2.

Ilpoeor)HU/W(WJI r/Jy1t1W,UJI. IlcttTpOCTpeMHTC.'IbHhIC HMTiy'.'lbChl, fIOC'ryIT310ll\HC B CTIIIllIIOH M03r !IO 33.11,HIIM J<OpCllll:'8M, nepc.11,aKiTCH no npcrno,llHlll;HM rryTHM CHHHHOro M031'3 B BbillleJ1emaw;IIe OT/\e.ilbI I'OJIOBHOI'O
(BOCXO,'],Hlll;IIC IIYTl1 ).

B

CBOIO (Jt-Iepe,ll,h, II3 I'OJIOBllOI'O M03ra IIOCTYTIRKJT

Hl'llIIYJlhCbl B CIIIIHHOll :vI03r (HHCXO,!l,Hll.l,Ue IIYTII), KOTOph1e MoryT ll3MeHHTb /\CHTeJlbHOCTb CKe.1eTHOH MYCKYJIRTYPLI n

BIIYTPCHHHX Opl'3llOB.

Y.e:vr Bbillle opraHH30BaHa llCpBH3H CHCTe:vra, TC:YI 60.'Ibllle 3aBHCHMOCTb
cl.JyuKu;IIii: cmrnuoro :vro31•3 OT BhIUICJ1CiK3ll.LHX OT,ll,CJIOB

11,HC.

roJIOBHOH M03r
l'OJ10BIIOH M031' pacrroJraraeTCH B ITOJIOCTH M03I'OBOI'O qepen3 (M3CCa ()1{0J10

1500

r, ofrbeM OKO.'IO

1500

CM"). AHaTOMnqeCKH Bbl.11,CJI.HIDT c;1e;:i;y1orr~ne OT,Ue-

.'IhI: npO,llO.'II'OBRThIH, 33,llHHH (MOCT II M03JKeqoK), cpe.11,HHM, IIPOMCJKyToqnhrii
H I<OHC'-IHbIH (pIIC.

156).

0T,ll,CJibI I'OJIOBHOI'O M03r3, paCIIOJIOJKCHHhIC 6.111me K CIIIIHHOMy M03ry, 061,C,ll,HHHIOT TIO.II, o6ru;HM HR3BRHHCM cm60Jl .iVLOi/2a -

rrpo.11,0.'II'OBHTbIIi, 3R/1HIIH

(MOCT), cpc.11,ttnii: H rrpoMeJtcyToqHLIM M03r. Caa.uII K MOCTY np11.'H'JKIIT M03Jt<eqoK. CTBOJI COCTRBJIHCT C3MYIO .upeBHIOIO qaCTb I'OJIOBHOI'O M03!'a. 3;:i;ecb rrpoxo,L\HT HllCX0.11,Hlll;llC rryTH, CBH3bIB3IDII~HC KOHeqHbIH M03r co CI1llHHbl:YI MoaroM.
Oco6enuocTn:

01'CyTCTBHe cerMe11TapHOCTH,

cepoe nen~CCTBO pacrrpe.;:i;eJ1CHO

B BHJJ,C H,Uep, H3XOJ~HTC.H pCTIIKYJI.HpH3H QJOpMaHHH.

9
7

8

5
8

6

P111c. 156. CTpoeHvie ronosHoro M03ra: a - npoAOJ1bHb1Vi pa3pe3; 6 - ocHosaHvie ronosHoro M03ra.
1 -·- crnon: 2- npoAonroBarb1Vi M03r; 3 - Moci; 4 ·-· M03>K8YOK; 5- cpeAHVIVi M03r; 6 - YernepoxonMvie; 7- npoMe>KyTOYHb1Vi M03r; 8- Kapa 6onbWVIX nonywapviVi; 9- Mo3onvicrne 1eno: 10- nepeKpecT 3pV1TeJ1bHblX Hepsos; 11 - 060HmenbHb1e nyKOBVIL\bl

ANamoMUJt u tJ>u.1uo.rwzuJt •ie.rw6eKa

408

KoHeY:HhI}i Moar, oco6emw r10J1ymap11H, cocTaBJIHIOT c}rn:10reHeT11Y:ecK11 Hau60J1ee MOJ!OilOe o6paaouamie IJ;HC (no Becy 3aHHMaIOT OKOJIO

80 '/,,

OT MaCChI

ro.'JOBHOI'O M03ra). Ilo.11ymap11H IIOKpbIThI CJ10CM ceporo neu~eCTBa, KOTOpbIH Ha3bJBaeTCH KOpOH rOJ10BHOro M03ra H HBJUleTCH BhICIIIHM OT,[\eJIOM IIeHTpaJihHOH
HepBHOH CHCTeMbI. 0Tµ;eJ1bHhie CKOIIJieHHH ceporo BeIIWCTBa Ha3h!BaIOTCH a,n;paMH 11 o6pa3yl<Yf HepBHhre 11,ettTphr.

B

cooTBeTCTBHH c BhIIIOJIHHeMhIMH <PYHKIJ,II-

HMH BhI,[\eJIH!OT pa3JIH'IHble 'IYBCTBHTeJlhHhie 11,eHTphI, IIeHTphI BereTaTIIBIIhIX
c!>YHKIJMH, .L\S.IraTe"lhHhle 11,C!!Tphl, IWHTph! IICHXHlJ:CCKHX <PYHKil,HH HT.µ;.

<l>ymrn;HH OT,!J;eJIOB rOJIOBHOro M03ra

IIpodoJt206ambtU MO.J?

HB.'lHeTca npo.n:oJT>KeH11eM cnuunoro M03ra. CnttHHO-

M03rOBOH KaI-IaJI pacIIIttpaeTca u 06pa3yeT

IV

meJiy.n;oY:eK M03ra.

B

cepoM Be-

IIJeCTBe HaXO,il;HTCH ,!l;BHraTeJlhHhie H Y:YBCTBHTeJihHhie aµ;pa HeKOTOpb!X qepenHO-M03rOBhIX HepBOB.
<t>yHKIJHH:

1.

IlpoBO/\HIIJa.H -- BOJIOKHa 6eJioro BeIIJeCTBa o6pa3y}(Yf BOCXO,[\J:lIIJHe (<IyBCTBHTeJibHhie) II HHCXO,!l;HIIJHe (µ;BnraTeJihHhie) npOBO,[\.HIIJHe IIYTH.

2.

Petj)Jieinopttaa -

3.n;ech pacnoJiometthr JKH3HeHHO Bamtthre IIeHTphr pe-

ryJIHI1HH ,[\hIXaHHH, KpOB006paIIJeHHJ:l (TOHYC cocyµ;oB, µ;eaTeJlhHOCTh
cep.u;I1a), <PYHKIJHH IIHIIJeBapeHHH, o6MeHa BeIIJeCTB. IJ;eHTpbI 6e3yCJIOBHhIX pecj>JieKCOB -

JKeBaHHH, cocaHHH, !'J10TaHHH, pBOThI, 'IHXaHHJ:l, Ka-

IIIJIJ:l, MOpraHHJ:l. IJ;eHTpbI BeCTH6yJIHp!IbIX pecp;reKCOB -

petj)JieKTOpHoe

no.u;,n;epmaHHe HOpMa.'lbHOro IIOJIOJKeHHH TeJia B npocTpaHCTBe.
IloBpem,n;eHne npOl(OJII'OBaToro M03ra Be,n;eT K CMepTH BCJie,n;CTBHe npeKpaIIJeHHJi ,ll;hIXaHHJi HJ1H OCTaHOBKH cep.n;rw.

3adl-tUU .Jt/0.32 COCTOHT H3 ,n;Byx OTl(eJIOB:
Bapo.11ue6 Mocm pacno.rraraeTCH Bnepeµ;u

rrpo,n;oJiroBaToro M03ra B BH,n;e

yTomn:eunoro BaJ1HKa. BeJioe BeIIJCCTBO MOCTa COCTaBJI.HeT BOCXO,ll;J:lIIJHe
H HIICXO,ll;HII~He npOBO,ll;.HIIJHe nyTH. Cepoe BeIIJeCTBO MOCTa o6pa3yeT .H,ll;pa qepCIIHO-M03I'OBhIX HepBOB H co6CTBeHHhie H,n;pa.
<t>yHKIJHH:

1.

IlpoBO/IHIIJa.H -

BOJIOKHa 6eJ10rO BeIIJeCTBa CBH3hIBa}()T BhIIIIeJieJKaIIJHC

OT,n;eJib! c npo.n;oJirOBaThIM H CIIHHHhIM M03I'OM.

2.

Petj)JieKTOpttaa -

3,n;ech pacnoJiometthr IIeHTphr 6e3yCJIOBHhIX petj)JieK-

coB l'JIOTaHHH H MHraHHH. IJ;eHTphl, peryJ1HPYIOIIJHe pa6oTy MHMH'IeCKHX H JKeBaTeJihHhIX MhlllIIJ, ,ll;BHJKeHHJ:l rJia3 H .n:p.

M oaJ1Ce'lOK pacnoJiaraeTcJi no3a,n;H MOCTa H

npuKphrnaeT npo.n;oJiroBaThIH

M03r. CocTOHT H3 ,n;Byx noJiyIIIapMH, coe.n:ttneHHhIX

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Cepoe BeIIJeCT-

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418

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' 110.1yY!8HHe n:mpopMaUHH, llOCTYllHBUleii

KOil!{pCTHblX cn:rttaJ10B, rrpeJlMCTOB H

J

HBJienn::H 1rnewnero MHpa, H.UYII!HX OT

K qeJioBeKy B BttiJ:e CH:Y1Bo,10B (cJ1on,
3HaKOB, cj:JopMy,1)

peaKUHH opraHOB qyBCTB
06ecneqHBaeT IWHKpeTJIOe MhIIlIJieH11e

i 06ecrre'-urnaeT a6cTpaKTHOe MhilllJienne

Tuni.1 BblCIIIeii HepBHOH ,i:i;enTeJILHOCTH

y

LJ:eJIOBeI<a HMeIOTCH HH,L(HBH.ZJ;yaJihHbie oco6eHHOCTH BbICIIIeii HepBHOH .n;e-

HTeJibHOCTH. Pa3JIH"l!HH onpe,n;eJIHIOTCH B3anMOOTHOIUeHHHMH rrpol!eccoB B036ym,n;eHHH H TOpMOJKeHHfl, KOTOpbie

ll.

Il. IlaBJIOB B3HJI 3a OCHOBhl .ZJ;Jlfl l\.J'IaC-

CHcpHKaUIIH THIIOB BhICIUe:H HepBHOM ,n;eHTeJibHOCTH. Bb:r,n;e.'IHIOT LJ:eThrpe OCHOBHhlX Tttna BhlCIIIeM HepBHOH ,n;e5ITCJibHOCTH:

1. CUJlbHbtu 1teypa61-l06eUJe1i1tbtU mun -

npou;ecchI B036ym,n;eHHH rrpeo6Jia,n;a-

lOT Ha,[( rrpo:r:reccaMH TOpMoJKeHHfl (BbICOKHM ypoBeHh pa60TOCIIOC06HOCTM).

2. CUJlbHbtzt ypa61toeeweH1tbtu mun -

ypaBHOBerneHHOCTh B036ym.n;eHHH

H TOpMOIKeHHH, BblCOKaH no,n;BHJKHOCTb.

3. Cu.7blibtU ypa6H06eUJe1mbtU u1tepm1tb1u mun -

rrpeo6.'Ia,n;aHtte npouec-

cos B036ym,n;eHHH, HH3Kafl IIO.ZJ;BHJKHOCTh (3aMKHYTblM nepexo,n; H3 COCTO.HHHH B036ym,n;eHHfl B TOpMo:>KeHne HJIH Hao6opoT).

AnamoMUJl u </JU3uo;wzuR 'te.!loBeKa

420

4. C.1ta6b1it mun -

11peo6Jia,n:a10T npouecc1>1 Top:vi01:1<eHMH Ha;~ B036ym,n;errn-

eM (6bICTpa.H yTOMJ:rneMOCTb M MaJra.H pa60TOCTIOC06HOCTb).
TttrrhI ttepBtto:iI cHCTeMbI,

Bh1,n:e.1e1u1b1e

H.

IT. IIaBJIOBbIM, cooTBeTCTBYIOT

KJlaccur/JuKau,uu me.lftnepaN<'H moB 'le.,106eKa, rrpe,n:"i:omeHHOH BpatJOM L(peBttero
Milpa fmmoKpaTOM

2500 JICT H333.[\.

ITo THTIY TeMrrepaMeHTa pa3,n:eJI.HIOT mo,n:eH::

1.
2.
3.
4.

XoAepuK - tteypaBHOBemeHHhIH, "1erKo B036y,n:HMhIH.
Ca11ieuHUK - ypaBHOBCIIIenttbr:iI, c 110,n:BHmHoii ttepBHOH cttcTeMo:iI.
<P.r1ei.11wmuK -- ypaBHOBeIIICHHbIH, CITOl<OHHbIH, ImepTHhIH.
M eJlaHXOJLUK - MpaqHbitt, rro,n:aBJieHHhrtt.
Co.JHaHue - ::>To BhICilla.H cpopMa o6o6mettttoro oTpamemrn qeJioBeKOM 0Kpyma10.w;ei1: ;:i;e:i-icTBHTCJlbHOCTII II CBOero co6CTBeHHOro Mttpa ( OC03H3HIIe co6cTBeHHOro «.H» ). Otto 011pe,n:eJI.HeT ueJieHarrpaBJiettttoe rroBe;:i;eHIIe qe"10BeKa c yqeTOM H3KOIIJieHHoro OIIbITa, a T3K)l{C rrpIICIIOC06JieHI'.!e opraHM3Ma K OKpymaIO.w;eil cpe;:i;e.

MaTepttaJihHOH cpopMOH

B03HMKHOBeHII.H C03HaHH.H .HB.1.HeTc.H

.H3hIK. Co3ttattue orrpe;:i;e.1.HeT BHIIMatttte, crroco6ttocTh cocpe,n:oToqHThC.H, oueHIITh M rrporH03MpOB3Tb CBoe 110Be,n:ettMe. Co3H3HIIe cpopMMpyeTCJI B pe3yJihTaTe B33HMO,n:e:i1CTBM.H Memt1;y CTPYKTypaMM KOpbl II IIO.[\KOpKII.
fJiaBttoe OTJittqIIe BH,U qeJioBeKa -

MbtUJJleHue u pe'lb, KOTOpb1e rro.HBH-

JIMCh B pe3yJihTaTe Tpy,n:OBOH ,n:e.HTeJihHOCTH.

MbtWJleHue -

KOM11otteHT 1103ttaBaTe.1btto:i1 ,n:e.HTeJihHOCTM, xapaKTepH3YIO-

JUIIHC.H o6o6meHHbIM OTpameHtteM ,n;eHCTBIITeJibHOCTH. 3a CIJeT MblIIIJieHM.H 011pe,n:eJI.HeTC.H cymttocTh 11pe;:i;MeTOB tt .HBJieHHH. B

octtoBe MhIIIIJieHII.H JiemHT

rrpouecc o6pa3oBaHH.H accou:HaUIIH pa3HOH CJiomuocTM. Pa3JIIIIJaIOT 3JieMeHTapHoe (KOHKpeTHOe) MbIIIJJICHlle M a6cTpaKTHOe. 3JieMeHTapttoe npIIcyme )J{IIBOTHblM II qeJIOBeKy; a6cTpaKTHOe 11pucyme TOJlbKO qeJIOBeKy' pa3BMBaeTC.H
B CBJI3M c pe4hl0 M B03MO)l{HOCTblO o6o6meHM.H MaTepttaJia.
BJiaro;:i;ap.H

l'Jl06Y B03HMKJIM o6o6meHHbie TIOHHTM.H II npe,n:cTaBJieHMH, Cl10-

co6HOCTb K JIOruqecKOMY MbllllJieHHIO.

CJloeo -

pa3;:i;pamIITeJih, BhI3hrnaeT y qeJioBeKa 6oJihIIIOe KOJIIIqecTBO yc-

JIOBHhIX pecpJieKCOB. Ha HHX 6a3HpyIOTC.H 06yqem1e, BOCl1IIT3HIIe, Bblpa60TKa
rrpttBbJqeK, Tpy,n:OBbie HaBbIKM.
C.110Bo -

CIIMBOJI 11pe;:i;MeTa, JIBJieHtt.H; MomeT 6hITh 06061n;a101UIIM M a6cT-

paKTHhIM; OT06pamaeT HHcpOpMaUHIO M3 BHeIIIHero Mttpa, TO eCTb .HBJIJieTC.H
«CIIrHaJIOM cMrttaJioB» (110

H.

IT. ITaBJIOBy). Pe6eHOK yqIITC.H o6maThC.H c 110-

MOJUhIO CJIOB B B03pacTe OT O)J;HOro ;:i;o II.HTH JieT. «MayrJIH»,

pe6eHOK, BOCl1M-

T3HHblH )J{HBOTHbIMH II IIOl13BIIIIIH B qeJioseqecKoe o6mecTBO rrocJie II.HTM JieT,
KaK npaBMJIO, He OBJia.[\eBaeT qeJioBeqecKOH peqbIO.

Pe'lb -

3TO BhICIIIa.H cpyHKUII.H UHC, Bamtthiii MexaHH3M MHTeJIJieKTyaJih-

HOH ,n;e.HTeJihHOCTH, cpopMa 06mett11.H JIIO,n;e:H.
B11,n:h1 peqII:
BII,ll;HM3.H (nIICbMeHHa.H) -

o6yCJIOBJieHa pa60TOH BMC04HbIX II TeMeHHOH

,n:oJieii;
CJibIIIIHMa.H (yCTHa.H) JiyIIIaptt.H;

o6yCJIOBJieHa pa60TOH BHCOqHOH .[\OJIH JieBoro rro-

flpo11eccbL TWMRlnU

421

npoH3HOcHMaH (Rc.1yx, npo cc6H, apnmy.1mvrn) --- 06yc"10B.1etta pa6oToii: !I06HOH ll,OJIH H BHCOY:HOH ll,OJIH .1eBoro rro.1yIIIapH.H.

II a.11-lRrnb ---

::no 3ano:vrttttattHe, coxpatteHHe H noc.1e.n;y10r.u;ee Bocrrpon3Be.ue-

HHe Y:eJIOBCKOM paHee nepeJKHThIX HM Y:YBCTB, Mh!C.'IeH lI o6pa30B pattee BOCnpHHHTh!X npell,MeTOB n .HBJICHHii.

Ilpou;ecchI rraMHTH

K rrpou;eccaM

rraMHTH OTHOC.HTc.a:

3a nw11u 1w H UP --

rrponcxo.n;HT 3aKperr.1eHHe HoBoro MaTepHa.1a rryTeM

CBH3hIBaHH.H ero c rrptto6peTCHHh!M pattee.

CoxpaHeHue -- orrpell,eJI.aeTC.H cTerreHhJO yY:aCTH.H MaTepnaJia B ll,eHTe.1hHOCTH JIJJLIHOCTII, TO eCTh BaJKHOCThlO ,ll;JI.H qeJIOBeKa.

Bocnpou.:1BeiJe1tue -

rrpoucxo,n;nT aKTyaJIH3au;n.a 3aKperrJieHHoro MaTe-

pHa.1a rryTeM H3BJieY:eHHH ero H3 ll,OJirOBpeMeHHOH rraM.HTH II rrepc,n;aY:a
B rraM.HTh orrepanrnttyJO.
HaII60JihIIIero pa3BIITII.H rraM.HTh ,n;ocTnraeT K

50

JieT. 3aTeM cnoco6HOCTh 3aIIOMHIITh H

25

ro,n;aM H coxpatt.aeTCH ll.O

BCIIOMHIITh IIOCTerreHHO IIll,eT Ha

y6h!Jlb.

II oBmopeHue -

o,n;no H3 BaJKHhIX ycJioBnii: 3arrOMIIHaHII.H. ,[(JI.a yJiyY:IIIeHH.H

3aIIOMIIHaHH.H Heo6xo,n;IIMO Bhl,[l,CJICHIIe OCHOBHhlX MhICJiei1, IICIIOJih30BaHIIe
IIJIJIJOCTpau;IIii, COCTaBJieHIIe rr.1aHOB, cxeM, Ta6JIHIJ;.

BH,iJ,hl naMHTH

1.

Ilo xapaKTepy IICHXII'-!eCKOH aKTIIBHOCTII:

Jieuiame.'lb/ia.fl -

3aIIOMHHaHIIe H BOCIIpOH3Bejl,eHHe l(BHJKeHIIH (o6y-

'-IeHne crropTHBHhIM HaBhIKaM: IIrpa B TeHHIIC, IIJiaBairne H ,n;p.).

3.MOU,UOHaAbHa.R -

coxpaHeHIIe rrepeJKHThIX Y:e.10BeKOM '-IYBCTB (co-

'-IYBCTBIIe ,n;pyrIIM JIJOll,.HM).

C.MblCAOBaR -

3arroMIIHaHIIe, coxpattettne II BocrrpoH3Be,n;ettIIe npoqH-

TaHHhrx, yc.'IhIIIIaHHhIX IIJIII rrpOII3HeceHHhIX CJIOB (3aIIOMIIHaHIIe CTHXOTBOpeHnii:).

06pa.JH<l.fl -

3aIIOMHHaHIIe 3pHTeJihHhIX H 3BYKOBhIX o6pa30B (KapTH-

Hhl npHpo,n;hr, MY3bIKaJihHhle MeJlO,ll;HH).

2.

Ilo xapaKTepy u;eJieii 11e.aTeJibHOCTII:

Henpou360AbHaR -

3arroMIIHaHne n BocrrpoH3Be,n;ettne 6e3 crreu;IIaJih-

ttoii u;eJm (JKII3HeHHhlll OIIhIT qeJIOBeKa).

1Ipou.160AbHaR -

3arroMIIHaHIIe n BocrrpoH3Be.n;eHIIe, B KOTopoii npH-

cyTCTByeT crreu;naJihHa.a u;eJih (rrpocpeccnoHaJihHbre 3HaHH.H).

3.

Ilo rrpo,n;OJIJKIITCJ!bHOCTH coxpaHeHH.H H 3aKpenJieHH.H MaTepHaJia:

KpamKoBpe.MeHHaR -

KopoTKoe coxpaHeHIIe n 6brcTpoe BocrrpoH3Be.n,e-

HHe MaTepHaJia IIOCJie O,ll;HOKpaTHOro BOCilpH.HTH.H (MexaHII3M HaKOil·
JICHH.H orrhrTa). BIIll. KpaTKOBpeMeHHoii: rraM.HTH -

onepaTHBHa.H -

OCyIIJeCTBJieHHe BaJKHhIX .n,ettCTBHH B .n,aHHhIH MOMCHT (qTeHHC, CITH·
ChIBaHIIe).

Anamo ..uun u <jJu.Juo;wiun

422

ll,o.11?06pe,we111w11 -

•teJWHeKa

,LVIIITe.!IbHOC coxpaHeHHe MaTepna:rn rroc.1e :vit1oro-

KpaT110ro ero JIOBTOpemrn: H BOCIIPOH3Be.n:emHI (HaKOIUleHHC npoclJeCCHOHaJibHbIX 3HaHHH).

3.izou,uu -

3TO CjJOpMa IICIIXH'IeCKOro nepeiKHBaHHH, lWTOpaH OTpamacT

cy6'heKT1IBHOe OTHOIIIe1111e. Pa3Jll1lialOT ,ll;Ba THIIa 3MOUHJI: rro.nmKHTe.'lhHbie
(pa.n;OCTb) H OTpHI~aTe:IhHbie (rueB, CTpax, TpeBora). 8MOll,1101!8JlbHYIO HaCTpOeHHOCTb H MOTHBaJ~HH (rro6ym.n;eHHe) KOHTPOJIHPYIOT J1I1M611qecKaH CHCTeMa,
rHITOTa.naMyC 11 BhICIIIHe I(ef!Tpbl pery.'lm(HH 3MOUHH (.1106HaH H BHCOlJHa.H KOpa 6oJihlllHX noJryrnapnii: M03ra).

Cou -

3TO nepno.n;nqecKoe cpyHKUHOHaJihHOe cocTO.HHHe qe.'TOBCKa, xapaK-

Tep113y10w;eec.H OTCYTCTBHeM ue.'leHarrpaB.'leHHOH .n;e.HTeJibIIOCTH H aKTHBHhIX
cB.H3eH c 0Kpymaro1qei1 cpe.n;oii:. Bo BpeMH ctta aKTHBHOCTh M03ra He yMeHhlllaeTC.H, a nepecTpa1rnaeTC.H.
Ilpo.n;OJI:>Kl!TeJihHOCTh CHa 38BHC1IT OT B03pacTa 'leJIOBeKa 1I ero cp113110.nor11qec1wro COCTOHHll.H (y HOBOpom.n;eHHbIX -

21

7-8

q, y B3pOCJihIX -

TblO qacTb iKll3HlI tie.'TOBeK rrpOBO,Zl;lIT BO CHe: CITHT

25

113

75

lJ). Tpe-

JieT.

Cyw;ecTBYIOT pa3Hhie CT8,ll;llll CHa. 3a Bpe!\1.H HOlJHOro cua y qe.'IOBeKa Ha6JIIO.n;aeTC.H OT Tpex .no ITHTll

rrep110.n;HqecKHX CMCH Me,ll;JieHHOI'O H 6bICTporo

CHa, BO BpeMH KOTOpbIX H3MeHHIOTC.H cpH3HOJIOrHqecKHe HOpMbl opraHH3Ma
H rrcHxHqecKHe rrpouecchI M03ra.

CJI

M eo.rteHHblU

COil ( «OpTOJWKCaJihHbIM ») nacTyrraeT ITOCJie 38CbIIT8HHH, ,IJ;JI1IT-

60-90 MHH.

IlpH 3TOM CHHmaeTCH 06!\teH Bew;ecTB H aKTHBHOCTb cep,ne'IHO-

cocy.n;11cTOH,

,Zl;bIXaTe.1bHOH,

rrnw;eBapHTCJihHOii: II

BbIJJ:eJIHTeJihHOH CHCTeM,

Mbilllll,hI pacc.na6.'IHIOTCH, TeMrrepaTypa TIOHH:>KaeTCH.
Cuou11.n;eu11.a 0Tpa:ma10T npou.ecchr MhilllJieHH.H 11 rrepecKa3 co6hITHii: npollle,nmero .n;H.H, OHM a6cTpaKTHhI 11 no3HaBaTe.'IhHhI. Mo:meT rrponcxo.n;HTh pa3rouop BO CHe, B03HHK8JOT HOqHbie CTpaxu y .n;eTeii: II CHOXOJK.n;eHHe (JiyHaTII3M).

BbtcmpblU COii ( «rrapa.n;OKCaJihHbIH»)
,l(JIHTC.H

10-15

HaCTyrraeT IIOCJie Me.n;JieHHOro CHa,

MHH. IlpH 3TOM aKTIIBH3HpyeTC.H ,neHTeJibHOCTh BHyTpeHHHX

opraHOB: yqaw;aeTC.H IIYJihC, ,ll;hIXaHMe, TIOBhIIIJaCTC.H TeMrrepaTypa, COKpaw;aIOTC.H r.na30,ll;BHraTeJibHhie (r.TJa3a 6bICTp0 ,ll;BnraJOTC.H), MIIM!1lieCKHe Mbllllll,bI
H MbllllUhl KOHe'IHOCTeii:.

B

M03re np0Hcxo.n;11T B036y:m,neHne ueii:pouoB 38ThIJioquhrx .n:o.neii:, rroJIBJIJI-

IOTC.H 3MOUHOH8JibHhie CHOBH,IJ;eHHH co 3fJHTeJibHhIMH, 3BYKOBbIMH H o6oH.HTeJibHblMH o6pa3aMH. Ilpoucxo.n;HT K.naccucp11Ka11,u.H 11 yrrop.a.n;oqeaue rrocTynuBlllett
38 .n;eHb HHcpOpMaUHH. JlullleHHe qeJIOBeKa «rrapa.n;oKCaJihHOI'O» CHa npHBO,ll;HT
K paccTpoii:cTBaM rraMHTH H rrc11xHqecKHM 3a60.neBaHH.HM.

M.

M. CetJeHOB ua-

3bIBa.n CHOBH,ll;eHHJI «He6bIB8J'IbIMH KOM6HH811,H.HMH 6hIB8JihlX BrreqaTJieHHll».

OpraHbI

qyBCTB

Opzanbt 'lyBcmB -

3TO opraHhI uocrrpH.HTHH pa3,npameu11ii:, 11.n;yw;ux H3 or<-

pyma10w;eii: H BHyTpeuuett: cpe)Ihl.

K

HHM OTHOC.H:TC.H opraHhl OC.H38HH.H, 3pe-

HH.H, c.nyxa, BKyca H o60H.HHH.H. Ilo.nyqeune OT opraHOB qyBCTB HHcpopMaIJ,Hll

Opzan bl

'lYfH'tnH

423

0 COCTOHllHI! H H3Me!IeHHH BHernHeii: H BllyTpc1111eii: cpe;~bl 06ecne'111Ba10T auaJIII33TOpbI. Yqe1rne 06 attaJrn3aTopax 6hrJro co3.n;atto

AHa.1u.wmopb1 -

11. 11.

IIan.10Bhr:vr .

KOMTIJICKC CTPYKTYP HepBHoii: cttcTe:vrbr, ocyw;ecTB.'IH-

10ru;nx BOCllPHHTHe H aHaJIH3 IIHCpop:viall,l!H B COOTBeTCTBYKHll,HX OT,11,e.::rax
M03ra.
0T,ll,CJihl aHaJIH33TOpfl: nepurjJepul/.eCICUU umiJe.1 (peuenTOphl), npoeorh1u1w6blU omiJeJl ('-IYBCTBIITC.nbHbie HepBbI), U,Pllmpa.1bHblU omiJe.I[ ('-IyBCTBHTC.nbHb!C
30Hhl KOphl 60.'Ihllll:IX Il0J'IYlll3pIItt).

Peu,enmopbl -

3TO OKOH'-l8HIIH <ryBCTBIITeJihHhIX HCPBOB HJIII K.'ICTO'-lllhle

o6pa30BaHI·IH' BOCIIPHHIIMaIOw;tte BHCUIHI1e B03.!J:CMCTBIIH ( <l>II3H'-leCKIIe, Xll:v!HqecKIIC, MexattnlieCKIIe) IIJIII H3MeHeHIIH BHyTpen11ei1 cpe,ll,bI opraHII3Ma.
CBottCTBa peuerrTopos: BhICoKa.H eoa6yauMocmb; BhrcoKa.H 1tyecmeume.1bHOCmb; cneu,uqJU'lHOCmb (pell,CIITOP pearIIpyeT TO.'IhKO Ha COOTBCTCTBY!Olll;He
,ll,.'IH nero B031\CHCTBIIH); nopoz pa.:1iJpaJ1CCHUJl, T. e. orrpe.n.eJieHHa.H CIIJ'Ia H ,ll,.'IIITe.1bHOCTh B031\CHCTBIIH pa3.n;paJKHTeJia; <l>YHKll,IIOHaJibHa.H JlGOUJlbHOCmb, '!'. e.
B036y.n;IIMOCTh pell,CilTOpOB ue IIOCTOHHHa; aiJanmaU,UJl (rrpHBbIK8Hlle) pell,errTOpOB, rrpO.HB.'IHCTCH B CHliJKCHIIII lIYBCTBIITCJihHOCTII K IIOCTOHHHO .n:ei1CTBYIOU~IIM

pa3.n.pamIITCJIHM.

Bu,n:hr peuerrTopoB

31ccmepopeu,enmopbl -

opraHbl lIYBCTB, BOCilpIIHIIMaIOru;IIe B03.!J:CHCTBIIH

BHCIIIHeii: cpCJ1bI.
KonTaKTHhre: BocrrpIIHIIMaIOru;IIe TCIIJIO, xoJio.n; (TepMopeuerrTOphr); 6oJih,
IlpIIKOCHOBCHIIe II J18BJICHIIe (TaKTIIJihHbie); CJiaJIKOe, KIICJIOe, COJICHOe
(BKYCOBhie) • .IJ:IIcTaIITHhie: pearIIpyIOru;IIe Ha CBCT (<PoTopeu;errTOpbl), 3BYKII ( <PouopeIWTITOpbI ), 3arraxII ( o60H.HTeJibHhIC ).

Ilponpuopeu,enmopbl -

yJiaBJIIIBaIOT II3MCHCHIIH B COCTOHHIIII onopHO-

,ll,BIIf'aTeJ'IhHOf'O annapaTa II BOCIIPIIHIIMaIOT COKpaw;eHIIe MhillIC'-IHbIX BOJIOKOH, HaT.HJKCHIIe cyxomII.'IIIM, CBH30K II cycTaBHhIX KarrcyJI.

Humepopeu,enmopbl -

pacrro"1araIOTCH Bo BHyTpeHHIIX oprauax, cTeH-

Kax KpOBCHOCIIbIX, .JJIIM<PaTII'ICCKIIX cocy.n.oB; BOCIIpIIHIIMaIOT XII:vllI'ICCKIIH COCTaB, ,ll,8BJ1CHIIe TKaHeH II JKIIJIKOCTeii: B opraHII3Me.
ct>yHKll,IIII 8H8J'III3aTopoB: 06Hapymen11e Ii BOCilplIHTIIe CIIrHaJIOB peu;eIITOpaMII; npeo6pa30BaHIIe II KO,ll,lipOBaHIIe CIIrHaJIOB B HepBHhlll IIMIIYJihC; nepe.n.aqa CIIrHaJIOB no rrpOBO,ll,Hlll;IIM nyTHM; aHaJIII3 II OII03H8HIIC CIIrHa.'la c IICIIO.nh30B8HIICM

rrpe,n;hI,ll;yu~ero

«JKII3HCHHOro OIIhITa ».

OpraH 3peHHH
3TO BaJKHCMllIIIH II3 opraHOB 'IYBCTB, 06ecrre'IMBaIOill;IIM L[CJIOBCKY ,D,0

90 %

IIH<l:>OpMaUIIII. Ilpe,n;cTaBJIHCT co6o:H rrepII<PepML!CCKYIO liaCTh 3pIITCJih-

HOro atta.rm3aTopa, uaxo,n;IITCH B rJia3HIIu;ax qeperra. CocTOMT tt3 rJia3ttoro
.H6JIOKa (rJia3a) JI BCilOMOraTCJihHOro annapaTa (rJia3HhIC Mhlllill,hl, BCKII,
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10 %

70 'Yo

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30 %

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448

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450-500

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200 r

-

1000 r.

r, a npn noBhillleHHhrx yMCTBeHHhIX Ha-

YrJieno.nhr 3anacaIOTCJI B opraHl13Me B B11.ne rJI11KoreHa (oKo.10

B neqeH11), a np11 l130hITOqHOM yrJieBO,ll;HOM Ill1TaHl1H -

YrJieB0,11,hI (rJIIOK03a) -

B BH,11,e ;1mpoB.

ocHoBHOtt ncTO'IHHK 3Hepr11n.

Ilo,11, .n:ei1:CTBl1eM cpepMeHTa nmuaJlUlta, MUJlbmO:Jbl B pOTOBOH TIOJIOCTl1 yrJieB0,11,hl
pacw,eTIJIJIIOTCJI .n:o C0.10,11,0BOI'O caxapa l1 l'JIIOK03bI, ,11,aJiee B TOHKOM l{HillC'-IHl1Ke TI0,11,
,n:ettCTBHeM qJepMeHTa UMUJlG.Jbl pacw,enJIJIIOTCJI ,11,0 rJIJOK03hl. f.TJ:IOK03a, TIOCTYTil1Blll3Jl B KpOBh 113 Kl1llle'-1Hl1Ka, TpaHCTIOPTMpyeTCJI B neqeHh, r.ne 113 Hee CHHTe311pyeTCJI l'Jil1KOreH -

pe3epBHhIH 3anac yrJieB0,11,0B. I'Jil1KOreH OTKJia,ll;bIBaeTCJI TaKme

B MhIIIIU.ax. Ilp11 He,11,0CTaTKe yrJieB0,11,0B B n11w,e r.1HKOreH pacw,enJIJieTCH .no
rJIIOK03hI, l1 LTJ:IOK03a no Mepe Ha,ll;06HOCTl1 TIOCTYTiaeT B KJICTKl1 TKaHett, r,n:e OHa HCTIOJlh3yeTCJI KaK l1CToqHl1K ::meprnl1. qaCTh l'JIIOK03bl no,n:BepraeTC.H pacna.ny c OCB06om,11,eH11eM ::mepr1111

17,6 K.D:m

Ha

1r

yrJieB0,11,0B ,11,JIH MCXaH11qecKOH pa6oThI, KaI<

Ha Cl1HTe3 MOJieKyJihI AT<I> ,11,JIH 3anacaH11H B opraHl13-

l1CT0'-1Hl1K TenJia, a ,11,pyraH -

Me. KoHeqnhre npo,11,yKThr pacna.na yrJieBo,n:oB -

ao.n:a l1 yrJieKnCJihitt ra3 -

BhIB0-

,11,.HTCH 113 opraHl13Ma c TIOTOM, Moqou, BhI,11,bIXaeMbIM B03,ll,yxOM.
Ilp11 ynoTpe6Jiemrn c n11w,ei16oJihrnoro KOJIH'-leCTBa o6hrqnoro caxapa (.no

200

150-

r) co.n:epmmrn:e l'JIIOK03hl B Kp0Bl1 pe3KO B03pacTaeT. 3To HBJieHl1e Ha3hIBaeTCH

n11w,eBoi1: r11neprJI11KeM11ei1:.

B

3TOM cJiyqae tt36hITOK caxapa BhIBO,ll,l1TC.H 113 opraHn3-

Ma c Moqou. IlOBblllleHHYIO KOHU,eHTpau.1110 caxapa B MO'le Ha3bIB3IOT l'JIIOK03ypl1eH.
IlOHl1fKeH11e ypOBHH l'JIIOK03bl B Kp0Bl1 -

r11norJIHKeMHH -

MOJKeT BhI3BaTh HepB-

Hhie paccTpoiicTBa (noTepIO C03HaHl1JI, cy.n:opor11) 11 .name np11BeCTH K CMepTl1.

B CBH3H c

HapyllleHl1eM yrJieB0,11,HOrO o6MeHa pe3KO B03pacTaeT HCTIOJib30Ba-

HHe 6eJIKOB 11 iKHPOB B KaqeCTBe HCTOqHl1KOB 3HeprHH, qTo conpOBOiK,11,aeTCJI
o6pa30BaHHeM H HaKOTIJieHl1eM npoMeiKyTO'-IHbIX npo,11,yKTOB pacna,11,a l1 Be,11,eT
K OTpaBJieHHIO opraH H3Ma ( caxa PH hill ,11,11a6eT).

06Mett mupo11. BumaMuttbt

449

Pe;:y.,7Jimopa.~1u ofi.MJ'lla y<:.1eeoao6 HBJUIIOTCH ropMOHhl (r:1I01<0reu, HJ\PeHaJII1H, Jll1HOTpOIIHH, r.'lIOKOKOpTHKOH,ll,bl) II CIIM!1aTII'leCKaJI HeIJB!ICUI CIICTe'.Vla.

06MeH ;irnpon
CyT0 11HaH IIOTpe6HOCTb opraHII3Ma B iKIIpax COCTaB.'lJieT

20-25 % -

80--100

I', H3 BUX

paCTIITe.'lbHb!e MaCJia. JKIIpbl BXOJUIT B COCTaB paCTHTe.'lhHOll

H iKIIBOTHOii nm:.ri;II, HCU0Jlb3YIOTCH KaK HCTO'lHllK ;rneprllII. lfaBeCTHO, lJTO IIpII

pacu~en.'leHuII

1r

mupa o6pa3yeTCH

38,9

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IlocTynHBlllHe c rmui;eii mHpbl pacui;ernrnIOTCH UO,ll, .neiiCTBHCM q)ep'.\ieHTa .1U-

na3bl B

TOHIWM KIIlllelJHUKe ,ll,O rJim:.repuHa II mupHblX KHCJIOT II pacxo,nyIOTCH Ha

cuHTe3 co6cTBeHHbIX mupoB (accuMUJIHIIH.H). BoJihllla.H 1IaCTh BHOBh o6pa3oBaBmuxcrr mupoB nocTynaeT B JIIIM¢y, MeHbllla.H -

Henocpe,'l,cTBemrn B KpOBh

11 pa3HOCHTC.H no BceM opraHaM u TKaHJIM. MHoro m11pos nocTynaeT B mupoByIO
TKaHh, KOTopa.H IIMeeT 3HalleHIIe mIIpOBOrO .neno ,ll,JI.H opraHU3Ma (nO,ll,KOmHa.H
mIIpOBa.H KJieTllaTKa) u COCTaBJI.HeT

10-20 %

MaCCbl Te.rra qe.'lOBeKa.

Ilpo,nyKThI pacna.na JKupoB -

C0 2 II H 2 0 BhIBO,ll,HTCH u3 opraHII3Ma c noTOM, MO'lOH, Bbl,lJ,b!XaeMhIM B03,ll,YXOM. JKupbl MoryT CUHTe3upoBaThC.H B opraHU3Me 11e.'lOBeKa U3 yr.rreBO,ZJ;OB H 6eJIKOB.

Pec:y.iamopaMu o6MeHa J1Cupoe

JIBJU!IOTCH ropMOHhl (;1un0Tpom1H,

TII-

poKCUH, rJIIOKOKOpTIIKOH,IJ,bl, IlOJIOBhie ropMOHbl).

BuTaMHHhI
Buma.'du1tbl

(.'laT.

vita -

mu3Hb) -

6uoJ1oruqecKH aKTHBHhie seu~ecTBa,

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BuTaMHHhI

-

opraHtttiecKue coe,nuHeHHH pa3Hoo6pa3Hoi1

xuMHtiecKoii

npttpO,'],bl, Heo6xo,nuMbie ,ll,JI.H HOpMaJibHOro pOCTa u pa3RHTH.H opraHU3Ma.
3Ha'leHne BIITaMHHOB 6bIJIO ,n;oKa3aHo B pa6oTax pyccKoro Bpaqa H.
HuHa B onh1Tax Ha,n muBOTHhIMU
npo11yKTaX UMeIOTCH

(1880

Hel13BeCTHbie ¢aKTOpbl

nuTaHH.H,

mu3HH. TepMIIH «BH:TaMIIH» 6bIJI npe.n.rromeH B

IC

11.

Jiy-

r.). OH ycTaHOBUJI, l!TO B mrni;eBhIX

1912

Heo6xon;uMbJe

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3BaJI ero «Vit» (Beni;ecTBO, Heo6xo,nnMoe ,n;JIH mII3HII).
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1.

He HBJIHIOTC.H TIJiaCTIIlleCKIIM MaTepIIaJIOM H He c.rrymaT IICTOlIHIIKOM

2.

.HBJIHIOTCH 6IIOJIOrIIl!eCKU aKTIIBHhIMH Beni;eCTBaMII yme B MaJiblX IWJIII-

3.

BbITIOJIH.HIOT pa3HOo6pa3Hble ¢YHKIIIIII: BXO,ll;.HT B COCTaB ¢epMeHTOB,

3HeprIIII.

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4.

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MY qe.rroBeK no.rryqaeT IIX c nIIni;e-ii.

450

AnamoMUJl u rpu.1uo.noiu.n •te.no6eKa

XapaKTepttcTHKa BttTaMHHOB

Boµ,opacTsopnMhre BHTaMHHbI

llittpopaCTilO[lH'.\lhie BHTaMHHbl

A

c (ae1wp6HHOBaH KlfCJIOTa)

(peTHH0.'1) -- ;K11p 11e•ieH11 Tpec1<11,

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I pyeTCH

neTpywI<e, IIOMH,l\Opax, MQ,10 1 1HbIC npo-

(Ka,'lhIJ;H$epo.i:) -

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.HH'IHhltt /KCJITOK, C.ll!BO'IIIOC MaColO, MOJIOKO, Cl1HTC3I1pyeTC.H B KOJKe IIOJI ,ll,CMCTBI1eM

HOK; f!J!Oil;hl: UBIIIOBHHK; cppyl<TbI: IJ;HT-

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B 12 (u;uaHoKo6aJiaMHH) -- rrpo.z:1y1<ThI
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B

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Pau,uonaJlbnoe numanue -

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451

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( 50-100

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Kpoeoo6paw,eHue -

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RpoBh MOJKeT BhrnoJIH.HTh JKM3HeHHO BaJKHhie <PYHKUHM, TOJihKO Haxo.D:RCh B HenpepbIBHOM ,ll;BMJl{€HMM • .IJ;eJIT€JlbHOCTh Bcex opraHOB M opraHM3Ma
B ueJIOM Tectto CB.H3aHa c cpyHKUMeii: opraHoB Kposoo6paiu;eHtt.a. R cttcTeMe
KPOB006pameH:n.H OTHOC.HTCR:

cep,11;u;e,

BhlIIOJIHRIOiu;ee cpyHKUilIO Hacoca,

M nepttcpeptttreCKHe KpOBeHOCHhie cocy,D;hl -

apTepllll, B€Hhl :n Kantt:JIJI.HpbI.

Kpo6oo6paUfenue

453

CTpoeHtte KpoBeHOCHLIX cocyAOB
Ilo BbIIlO,lHHeMOH ¢YHKU,M.H pa3JHP-IaIOT TPH BH,Afl KpOBeHOCHhlX cocy.nos:
apTepHM, KailHJIJIHpbl H BeHbl.
He3aBHeHMO OT COCTaBa KpOBH Bee eocy,[\hl, npHXO.QHlll,He K eep.nu,y, rrp11Ha-

6efta.MU, OTXO,l\Hlll,He - apmepuR.MU (ptte. 166).
Apmepuu - eocy,n;hI, no KOTopbIM KpoBb Te'IeT OT cepAua K KarrnJ1.1IapaM

TO C'IHT8Tb
B opraHbI.

CaMaa Kpynaaa apTeptta -

aopma. 0Ha Bbixo;:i;ttT tt3 Jlesoro )f<e,'ly;:i;oq1rn,
apmepuo-

BeTBHTea Ha KPYTIHhie apTepHU H Ha eoey,l\hl MCHbllJero ,l\HaMeTpa J1bl (eaMhre MaJreHhKIIe).
B e Hbl -

eoey,[\hI, no KOTOphIM KpOBh Te'IeT K eepAU.Y OT TKaHei1: H opraHOB,

npoi1:.na eeTb KalIHJIJIHpOB, MeJib'IaHUIHe BeHhl -

KanuJt.:LJl,Pbt cy,Ahl,

6eHyJibl.

:vie.11h'l.aliurn.e co-

tl

KOTOpble rrpOHH3bIBaIOT see

opraH1.1 11 TKaHII.

B

HIIX npottexo.nHT

06MeH JKH,AKOeTaMII, IlIITaTeJihHhlMII
semeeTnaMII II ra3aMII Mem.ny KpOBbIO II TKaHHMII. CTeHKH Kanu.11JI.Hpos eoeTOHT II3 O,L\HOro eJIOH IlJIOeKHX
KJieTOK :3H,AOTeJIIIa, Jiemamero Ha 6a3aJihHOil MeM6paHe. .LJ:IIaMeTp KaIlHJIJI.HpOB

0,2-0,4

poe eemnw;
qepe3

7

-

MKM;

,AJIIIHa

-

MM . KoJinqeeTBO Kamvvrn-

1

MM" KpOBH npOXO,AHT

KaJK,AhIH

KanIImrnp

3a

He-

eKO.'lhKO qaeos. He see KanuJ1Jiapb1
¢YHKU,HOHupyIOT

B

O,L\HOBpeMeHHO.

opraHe, Haxo.uameMea B eocTo.H-

III

Hun TIOKOH, qepe3 60JihlllYIO 'IaCTh

91, III

KanuJIJIHPOB KpOBb He Te'IeT (OTKphlTO ,l\JIH KPOBOTOKa
JI.HpOB).

20-30 '%

Bo speMH pa6oThl

JO\
J

KanuJI-

opraaa KO-

JiuqeeTBO KaTIIl.lIJIHpOB yBeJIII'IUBaeTca. Ma.11eHbKaa BeJIII'IHHa KPOBOTOKa
qepe3 KaJK,AblH KanHJIJiap KOMTieHeHpyeTC.H ux orpOMHhIM KOJIH'IeCTBOM

(3000

ua lMM" TKaHn). KamrnJiaphI

Ha'IMHaIOTCH

OT

npeKaTIHJIJiapoB,

KaiK,ll,hlH 113 KOTOpbIX Bna,ll,aeT B .npyrne npeKaTIHJIJIHphl HJIH B BeHyJihl.
TaK OHH 06pa3y10T neTJIH, coe.nnHaIOlll,IIe apTepIIaJihHYIO cneTeMy e BeH03HOM. CKopocTb KposoToKa B Kamurn.Hpax -

0,5

MM / c.

PMc.

166. CxeMa CTpoeH111si cTeHOK aprnpvM (a)

BeHbl

(6):

2-

I-

BHyrpeHHSlSl o6on04Ka:

6a3MbH3Sl MeM6paHa;

cnoi1;

4-

5-

Ill -

7-

Hapyit<HaR o6ono4Ka:

CT1114ecKaR MBM6paHa;
TenbH8R TK8Hb;

II -

rna.t\KIA9 MblW94Hbl9 Kn9TKIA;

3n3CTl1'19CKl19 BOnOKHa:

noKHa:

nOA3HAOT8niAMbHbli;i

BHyrpeHHSlSl 3naCTL<149CK3R MeM6paHa;

cpep.HRR o6on04Ka:

6-

3-

111

1 -- 3HAOTen111i1:

9-

KOnnareHOBble BO·

8- Hapyit<HaR 3na-

BOnOKHl1CTaR coeg11Hl1-

10- KpoBeHOCHblB COCYAb'

AuamoMUJl u <Jm.mu;wzuJl '!elW6eKa

454

XapaKTepHCTHKH Beu u apn~puif
Bc11b1

ApTep1rn

1.

··-·--·

CTeHKll aopTbl II apTepuii COCTORT ll3

BHyTpeHHllll -

1.

CTCHKH COCTOHT H3 Tpex CJ!OeB, HO

MeIICC !IJIOTHbie (TOHKHe), TaK KaK

Tpex c.1oes (060.10'-lcK):
llHTHMa, fJ!a.J,Kllll, COCTO-

MCHblllC Mblll!C'-!Hb!X 11 3,1aCTl1'!HhIX BO -

UT ll3 u,10CKllX KJH'TOK 3!!/lOTCJ!llH;

JIOKOH, B CBJI311 c 3TlfM OHII :\iCHee yrrpy-

cpe!lHHll -- Me!lHa, COCTOllT 113 l'Jla.z\KOl\lhl-

ru:e, HO 60,1ee paCT.fDf\:IIMbie; CJKHMaIOT-

IIIe'-IHblX K.1eTOK ll 3JJaCTl1'-!HbIX B0.10KOII;

CH COC'el\Hl1Ml1 Mhlll!U:aMII. BHyTpCHHR.fl

uapymHb!H

o60JIO'-!Ka o6pa3yCT IIOJiy.TiyHHOJl cpopMbl

-

aj(BCHTHIVW, COCTOIIT 113

qm6p03HOll COel\llHIITCo1hHOH TIWHH. f'JI(>

CK,1al\KII -

IOJaTiaHbl

rrpOXO;:\JIT HepBhl.
Mem,n;y rrepBbIM u: BTOpbIM c.1oeM II Me)!{llY BTOpb!M II TpeTLIIM eCTL 3JlaCTI1'HihlC
MeM6paHLI. BJiaro,n;apa TaKoMy CTpoeHIIIO
Beru:ecTBa II3 apTCPIIll He MOryT rrpoca'-lllB3TbCR B MeJKTKaUCBble ru:eJ111

2. Ilo

xo,n;y ,ZJ;Bll/KCHllfl KpOBH KJJaII3Hbl

3.

R'.pOBh TC'leT ITOl\ 6oJJLlllllM ll3BJ1CHllCM

(rrpu: ITOBpe:ll<,ll,CHHll KPOBh 6LCT ¢oHT3HOM
BbICOTOtt

1,5 M),

'2.

IC1arraHhI 11MeIOTCJI II OTKpbrna10Tca

! ITO):{

OTCYTCTBYIOT

13.

Ha110po:-.1 KpOBH B CTOPOHY

cep111~a

KpOBb Te'!eT HOA ue60JihlllllM ,[\3B.1e-

i HHeM (rrpu IIOBpC:ll{,[\CHHH KPOBb Bb!TeKa-

I103TOMY OHII pacrrOJIOJKC- : eT MeJIJICHHOll CTpyci1), !103TOMY OHH pac-

Hbl rJiy6oKO ITOl\ MbllllC'lHbIM CJIOeM

uo.1omeHbl IIerJiy6oKO, IIO,[\ KOJKCH (elln11c, Me.1Kue)

4.

CiwpocTL KpOBOTOKa

0,5

M/c

4.

C1wpocTb KpOBOTOKa

0,25 M/c

B cocy.n;McToii cMcTeMe pa3Jrn'-laIOT ,n:Ba Kpyra KpoBoo6paru;eHMH (pMc. 167).
::no CMCTeMa cocy.n;oB, no KOTOpbIM KpOBb
Te'-leT M3 JieBoro Jf<:eJiy,n;oqKa B opraHhI, a M3 HMX B npaBoe npe,n;cep.n;Me. Ha<IMHaeTCH aopToii, BblXO.[l;Hru;eii ll3 JieBoro JKeJiy,n:O'IKa, Hecyru;eii apTepHaJibHYIO
KpOBb. Pa3JIH'-laIOT BOCXO.[l;Hru;yIO ll HHCXO,n:Hru;ylO 'laCTH aopTbl. OT BOCXO,D;.Hru;eii
'laCTH OTXO,D;.HT KOpottapHbie apTepHH. OT ,n:yrH OTXO,D;.HT TPM cocy,n:a: n.rte'leZOJt061WU cmBOJt, KOTOpblH ,n:eJIHTC.H Ha npaByIO 06ru;y10 COHHYIO ll npaByIO no,n;KJllO'IH'-IHYIO apTepMH; JieBaJI o6w,aR COHHllJl apmepuR; JieBa.H !WOh'..'l/O'lU'lHUJl
BoJtblllOU Kpyi Kp06006paw,ellUJl -

apmepuR •

.Uo ,n;uacpparMhI aopTa - ipyattaR, HHJKe -- 6p101111taR. IIo nyTu aopTa ,n;eJIHTCH Ha apTepnH, KOTOpbie BXO,D;.HT B opraHbI ll ,n;e.'l.HTC.H Ha MeJIKHe apTepHH --+ apTepHOJibI --+ KanHJIJI.HpbI, r.n:e nHTaTeJibHbie Beru;ecTBa ll KHCJIOPOll
nepexo,n;.HT M3 KpOBM B TKaHeByIO JKH,n:KOCTb, a npo.n;yKTbI MeTa60JIIf3M8
B KPOBh. KpoBh cTaHOBHTC.H BeH03Hoii, co,n:epmHT npo,n:yKThI MeTa6oJIH3M8
ll yrJieKHCJlbIH ra3, OTTeKaeT no CHCTeMe BeHyJI B BeHhl --+ ,D;Be nOJible BeHbI
(sepxH.H.H H HHJKHHH), KOTOpbre Bna,n;a10T B npaBoe npe.n;cep,n;ue; c10,n;a me
Bna,n;aeT KOPOHapHhIH CHHYC (HeCeT BeH03HYIO KpOBb OT cTeHOK cep,n:u;a).
Kpyroo6opoT KpOBH npoucxo,n;nT 3a 23 ceKyH,D;hI.

lipo61uwe <JatJJtenue

455

1VfU.'lbLil Kpyi 11:poeoo6paw,Pn11.H (,irerotIHhIH) na<nmaeTcH B rrpaBOM ;.Ke:ry;:i;otJKe cep,!(ua, OTKy.ua BhIXO.UHT CTB0,1 Jlero•moii apTepHH, necyU\Hll BeH03HYIO
KpOBb ~ .11.e.TIHTCJl Ha npaByJO II Jiesyro JierOtIHbie apTepHII ~ BXO,l\HT B JierKIIe
11 .neJ1HTCJl TaM Ha apTepHOJihl H KallHJ'IJUipb!, KOTOphre orr.1eTaiOT a,1bBeOJibI,
B Kamt.1: 1.HpHbIX ceTHX ,1erKHX npo11cxo.u11T 06orarn.em1e Kposu KHC,'Iopo11,oM,
OHa CTaHOBHTCH apTepHa,lhHOM II no .D;BYM BeHaM OT KaJKD,Oro J'Ie!'KOI'O Bna.naeT
B JreBoe npe.ucep.utte. Kpyroo6opoT KpoBH npo11cxo.11.11T 3a 1 ceKyH,nh1.
llpaBbIM lI .'ICBbIH me.'ly,!(OlJKH Hal'TieTa.IOT KPOBh B cocy/l;hl O,UHOBpeMeH2
HO, II OHa ,[l,BHJf<eTCJl cpa3y no OOOHM
1,pyraM KpoBoo6parn.eHIIH.
KpoBOCHa6J1CeHue cepiJu,a. B .uo3
nonHeHHe K 60JibllIOMY Kpyry HMeeTCJl TpeTHH (cep,n;eqHhIM) Kpyr KPOBOo6parn.eHHa, CHa6ma10rn.11ii KPOBbIO
cTeHKH cep.n,u;a. KposocHa6meHMe
cep.n,u:a HatJMHaeTca .n;ByMH BeHe'lHhIMH apTepHHMH, KOTOpbie OTXO.ll;JIT OT
BOCXO,n;arn.eii '!8CTH (JiyKOBHUbl) aopTbI. 3aKaH<urnaeTCH BeHaMH, CJillB8IOlll.HMIJCH B BeHeqHblll ClIHyc, BII8,[l,8!0lll.11ll n rrpaBoe rrpe.ucep.n,11e. qacTb BeH
OTKph!BBIOTCJl Herrocpe,D,CTBeHHO B IIOJ'IOCTh rrpe.ucep.zvrn.
IlpH'IHHhI .D;BHJKeHHH KPOBH no cocy.u.aM:
1. PHTMHqttoe coKpau~eHtte cep.uua.
2. HaupmI<eHHe cTeHoK KpoBettocHhIX cocy.uoB.
3. CoKparn.ettne cKeJieTHhrx MbIUII~.
4. Ha.TIHtIHe cep.n;etIHhIX KJranaHoB.
5. HaJIHtIHe BeH03Hhlx KJiarraHon.
6. Pa3H11ua Mem.n;y .uaBJieH11eM KpoBH B HaqaJihHblX II KOHetIHb!X llaCTHX COCY.UHCTOH CHCTCMbI.
7. IlpHcaChIBa10rn.an CHJia rpy,n;HOM
KJI€TKH.
Bee 3TO o6ecne-qHBaeT 6bICTpoe u o.uP111c. 167. 5onbwoC1 111 Ma1lbll1 Kpyr111 KpoeoHottarrpaBJieHttoe .UBHJKemrn KpoBH no
06pa~eH111R: 1 - aoprn; 2 -- 11ero'IHb1e aprecocy.uaM. Ilo Mepe BeTBJieHHa cocy.u11cp1-11-1; 3 - nero'!Hble eeHbl; 4 - aprep111111 sHyrToro pycJia ~aBJieHHe H CKOpOCTb B copeHH111x opraHOB; 5 - Kan1111111Rpb1; 6 - BOcy.uax CHHJKaIOTCH, KPOBb ycTpeMJIHeTPOTHaR seHa neYeHLll; 7 - sepXHRR Lil HLlllKHRR
CJI B o6JiaCTh HH3KOrO .n;aBJieHHH, T. e.

OT apTepHH K BeHaM.

no11b1e aeHbl

456

AnamoMuR u <jJu.1uonozu1t 'te.noeeKa

Kpomrnoe ;::i;aBJieHue
,UBIDKeHMe KPOBH 3aBHCHT OT .n:aBJieHM.H, C03.n;asaeMoro cep.n:u;eM BO BpeMH
ero COKparu:eHII.H II corrpoTHBJieHH.H TOKy KpOBII CTeHOK cocy.n;oB. CaMoe BhICOKOe .n;aB.'leHHe B aopTe (OKOJIO

100

MM pT. er.) I1 KPYIIHhIX apTep1rnx

(90

MM

pT. CT.). Ilo xo.n:y cocy.n;ttcToro pyc.w .n:asJieHIIe rrocTerreHHO na.n:aeT, oco6eHHO HM3KOe B KarrII.'IJIJ!pax (.n:o
yMeHblliaThC.H (OT

40

.n:o

10

40

:'vfM pT. CT.).

B

BeHax OHO rrpo.n:omf<aeT

MM pT. er.) H B IIO.'IhIX Bettax paBHO

(0

MM pT. CT.)

MJIM HM:>Ke aTMoccpepttoro .n;aB.'leHH5L To eCTh qeM .n;aJihIIIe OT cep.n;u;a, TeM Hume .n;aBJiemie.
,UJI.H Me,!l;HU:HHCKHX u;eJiett apTepu:aJibHOe .n;aBJieHHe H3MepHIOT B IIJieqeBOH
apTepnn

H.

c

IIOMOIIJ:h!O

cneu11aJibHOrO

rrptt6opa

-

TOHOMeTpa

(MeTO,!l;OM

C. KopoTKOBa: Hao'IO:>Kett11e pe3MHOBOH MattmeThI Ha rrJieqo).
Bo BpeM.H CMCTOJibI :>KeJiy.n;oqKoB .n;aBJieHHe B aopTe IIO,!l;HI'.IMaeTC.H ,Zl;O MaKCH-

MaJibHOro, HJIH cucmoJLu'leCKOi?O, i}a6Jle/iUJl, KOTOpoe IIO.HBJIJleTC.H B BM,ZJ;e rrepBOro TOHa. HopMa
(cTapIIIe

55

120-125

MM pT. CT. (B

18-50

JieT),

135-140

MM pT. CT.

JieT). Bo BpeM.H .n:uacTO.'IhI .n:aBJiettue rra.n:aeT .n:o MHHMMaJihHoro,

HJIH i}uacmoJLu'leCKOW (cocy,ZJ;HCToro), KOTOpoe COOTBeTCTByeT MOMeHTY MCqe3HOBeHH.H TOHOB. B HOpMe 3TO

60-80 MM

pT. CT.

Pa3HHU:Y Mem.n:y MaKCHMaJihHhIM H MHHHMaJibHhIM .n;aBJieHMeM Ha3bIBaIOT nyJLb-

C06bl.M 0G6JlC/iUe.M (B HOpMe

40-50 MM pT.

Ha3bIBaeTCSI rurrepTOHMeil:, a IIOHM:>KeHHe -

ApmepuaJtblibtU ny.7bC -

CT.). ilOBbIIIIeHMe KPOBJ!Horo ,ZJ;aBo1eHM.H
rurroTOHHeH.

:JTO pnn,mqtthre KOJie6aHM.H CTeHoK apTepuil: npn

npoxom.n;eHMH no HMM KPOBH. IlyJihC COOTBeTCTByeT COKparu:eHMIO cep.n:ua.
B HOpMe y qeJIOBeKa B CITOKOHHOM COCTO.HHIIH nyJihC

60-70

y.n;apoB B MMHyTy.

IlyJihC onpe.n;eJI.HIOT qaru:e BCero Ha o'lyqeBOH apTepHH, 6.'IH:>Ke K KHCTH.
Perym1u;uH ,11;BH:1KeHHH KpOBH no cocyi:i;aM
PeryJI.HUirn ocyru:ecTBJIJieTc.H M3MeHettueM ruttpMHhI npocBeTa cocy.uos, qTo
npHBO,!l;HT K H3MeHeHMIO o6'heMa KpOBOTOKa H .n;aBo'leHM.H KPOBIL

1. H epBliaR peiyJLRU,UJl ocyru:ecTB:rneTCJI peq)JieKTOpHo BereTaTMBHoil: HepBHoil: CHCTeMOH.

Cu.Mnamu'lecKuu omi}eJL BhI3hIBaeT cymemrn cocy.n;oB n nOBhIIIIeHne
napacu.Mnamu'lecKuil pacIIIupemrn cocy.n:oB

.n:aBJieHuJI KPOBH,

ll CHH:>KeHue ,ll;aBJieHH.H KPOBH.

Cocyi}oi)BuiameJtbHbtu u,eHmp Haxo.n:nrcJI B rrpo.n;o.'lroBaToM M03re IIO.ZJ;
KOHTpoJieM rnrroTaJiaMyca u KOpbI 60.1bIIIMX rroJiyIIIapnil:. CocTOM'l' n3
.!l:BYX OT,ll;eJIOB: cocyiJocyJ1CU6G/Out,eio, HepBHhie HMIIYJihChl KOToporo 3aCTaBJIJ!IOT COKparu:aThCJ! rJia.ZJ;KYIO MYCKyJiaTypy Il yMeHhIIIa!OT rrpocBeT
apTepnil:, n cocyiJopacwupR10w,eio, rrpnBO,ll;JIIIJ:ero K yBeJinqeHMIO rrpocBeTa apTepnil:, ycnJinBaroru:ero npnTOK KpoBn K opraHy.

2. I'y.MopaJtbHGJl peiyJLRU,UJl

ocy~eCTBJIJleTCH rro.n: BJIH.HHMeM Beru:ecTB, rre-

peHOCHMbIX KpOBb!O.

Ai}peHaJLuH n nopaopeHaJtuH, 6a3onpecculi, muuomeH3UH BhI3hIBaIOT
cymeHMe KpoBeHOCHhIX cocy,n:oB, ITOBhIIIIaSI TOHYC HX rJia.n;KMX MhlIIIU:.

CmpoenuecepiJita

457

[ucmaMUH pacumpHeT apTepHOJihI H settyJ11>1, ysc.1w111saeT npoHnu;aeMOCTh Kamuvrnpon.
c oomHOWPH.U(' 02 u C02 - CHHmeHHe co;:i;epmamrn 02 l-L'IH flOBbillleHHe
co11epmatt11H C0 2 BhI3hJBaeT pacnrnpett11e KposeHOCHhIX cocyJJ,os.

CTpoeu11e cepti;u;a
CepiJ~e - no u;eHTpaJihHhIH opraH Kposoo6pa~eHMH, pacno.TJomettHhIM
B rpy,n;ttoir nonocTM. IIpe,n;cTaB.1HeT co6oii rro.'IhIH MhIIIIe'IllhIH opratt, no q)Op"'le
ttenpami.1hHoro ynnoIQeHHoro KOHyca (p11c. 168). Pacrro,1araeTCH s rpy,n;ttoi1
rronocTH tte CMMMeTpHqHo ( 2I 3 cnesa, 1I 3 crrpasa). PacnmpeHHaH '-18CTh cep,n;u;a
_ aTo octtoBaHHe, o6paIQeHHoe ssepx 11 snpaso; Y3Ka.H qacTh - sepm11tta, ttarrpasJieH!Ia.fi BHH3 H BJieBO. )l,1tttta B cpe,n;HeM 10-15 CM, Macca 300 r (np116:maMTeJihHO cooTBeTCTsyeT senrrqHHe Ky,•aKa qe;rnse1rn); qeThipexKaMepnoe.

CTettI\lf cep,n;u;a:
sm'TpeHH.H.H -

;miJ01wpiJ, COCTOHT 113 IlJIOCKHX 3IIHTeJIHa)JhHbIX K,1eTOK

aH;lOTeJIH.H;
cpeilHlUi - MUOICapiJ, COCTOHT H3 rrorrepeqHQ-IIOJIOC8TOH MhIIIIe'-IHOM TK8HH;
ttapyJKH8H - anulCapiJ, COCT011T 113
IIJ!OTHOH coeJJ,HHHTeJihHOM TKaHH.
IIoil 3IlMKapJJ;OM pacnonaraJOTCH
Kop0HapHhie apTepH11 H HepBhl.
Cep,n;ue pacrroJiaraeTCH s oKonocepJJ;eqHoH: C)'MKe, HapyiKHhIH Jil1CTOK KOTOpoi1 _ nepu1CapiJ - cpaIQeH c rpy,D;11ttoH:
11 pe6paM!f; a BHyTpeHHHH ,111CTOK anuicapiJ- IIOKPhIBaeT cep,JJ;u;e 11 cpacTa11
4
eTCH c ero MhIIIIu;eH:. Mem.rzy arrmtap,JJ;OM
11 rrep11KaPl\OM Haxo,n;11TC.H rrep11Kap,n;11JO
aJibHaH nOJ10CTh, aano,1HeHHaH cep03HOH
:>KH,JJ;KOCTbiO, KOTOpaH yMeHhlllaeT TpeHHe cep,n;ua npH ero coKp~eHHH. Ha Ha9
pymaoii noBepxHOCTH cep,n;u;a HMeroTC.H
rrorrepeq11a.fl BeH03Ha.H 6opoa,n;a, OT,n;eJIH8
JOIQ8H upe.n;cep,n;H.H OT :>KeJiy,n;oqKoB,
H ,n;Be rrpo;.J;OJlhHhlX Memmeny,n;O'-IKOBhlX
PMc. 168. CxeMa BHyrpeHHero cTpoeH1-1s:i
6opo3.JJ;bI - nepe.n;HH.H H 3a.JJ;HHH, pacrrocepAll<l: I - aoprn ; 2 - sepxHl'ls:l nonas:i eeHa;

JiomeHHbie aa rpaa11u;e Mem,n;y meny,JJ;oqKaMH. B 3'fllX 6opoa,JJ;ax ne:>KaT setteqHbie

3 - KJlanaH 11eroo.iHoiil apTep1-11-1; 4 - npaeoe

apTepirn 11 BeHhI cepJJ;u;a.
Mhrllleq!Ihie cTeHKl1 meny.n;o'-IKOB 60-

npaBbliil lKBllYAO'leK; 7 -- HHlKHflfl no11afl eeHa;

Jiee TOJICThie, oco6eHHO nesoro, TaK KaK
oH cosepJJiaeT HaH60JihIIIYJO pa6oTy
(rrpasoro - 4-6 MM, nesoro 9-11 MM).
Ilpe,n;cep,llll.fl HMelOT TOHKHe CTeHKH -

npeAcepAHe; 5 - Tpexcrnopo.iaTbliil KJlanaH; 6 -

8 - neperOPQAKa; 9 -118Sbliil lK811YA0'l8K: 10aopTMbHblH KJlanaH ; 11 - M'1Tp311bHblH K11anaH
(ABYCTBOP'l3Tbliil); 12 - 11eeoe npeACBPA'18;
13 - 118rQ'lHbl8 88Hbl: 14 - 11ero'lHblB apTep1-1"1

458

AnamoMun u pu.1uo1wzun '!ellOfffKa

ll,HK;I pa60T1>1 cepAu,a
' TJpO,l\OJ!-

I

,L(BH;IWl!Hl' KpOilH

cJ)a3bI cep11e•rnoro l\HK.<a

m11Te..:1h-

l!O('Th <tJa3
I
--+--

C0Kpall\CH11e ( c11cTo.w) npe11cep.uttft

I tfa

·--0,1 c

npe.ucep/:\llll B )l<CJIY .uo•m11

CoKpall\CHMC (cttCTO.'Ta) nw;ry.uoqKoB' lfa me.'lyl\O'IIWB B ;1eroqnyIO
apTepttlO n aopTy
i lfa BCH B npel:(CCp,L\II.H If B
PaccJra6JicH11c (l:(rracTo;ra)
npel:(cep,nttft n :>KeJiyl:(o•mon
I :>KCJIYl:\Oq!{lf

0,3 c
\

'

0,4 c

I

2-3

MM. Ilpe,n;cep,n;mr l1 meJiy,Zl;O'IKH coe,n;irneHbI OTBepCTI1.HMM, ITO Kpa.HM KOTO-

pbIX pacIToJiaraIOTC.H CTBOp'IaThJe KJiaITaHhI cep,n;I1a. Mem,n;y JieBhIM ITpe,n;cep,n;tteM

iJeycm6op'l.a
mblM (MumpaJLbHbl.M-). Mem,11,y ITpaBhIM ITpe,n;cep,n;11eM n. ITpaBhIM meJiy,11,o•moM -mpexcmBop'lambtii K.wnaH. K cB060,11,HhIM Kpa.HM CTBopoK KJiaITaHOB ITpnKpeITJie-

H JieBbIM meJiy,IJ;O'IKOM I<JiaITaH HMeeT ,IJ;Be CTBOPKM H H83hIBaeTC.H

HhI TOHKMe cyxo:>KMJibHbie HMTM -

CTPYHhI, ,n;pyrMM KOH!IOM ITPMKpeITJI.HIOIIIHeC.H

K COCOlJKaM MbIIllelJHhIX CTeHOK :>KeJiy,Zl;O'IKOB.
<PyHK!IMM KJiaITaHOB -

ITpeIT.HTCTBOBaTb o6paTHOMY TOKy KpOBH ITPM COKpa-

IIIeHH.HX meJiy,n;o'IKOB.
B aopTe H JierO'IHOH apTepttll MMeIOTC.H

llOJlYJLYHHbte KJLanaHbl,

COCTO.HIIIMe

113 Tpex 38CJIOHOB. Ilpn COKpaIIIeHMM meJiy,Zl;O'IKa OHM ITPMJieraIOT K CTeHKaM
cocy,11,oB l1. ITponycKaIOT KpOBb. Ilpn: paccJia6JieHMH JKeJiy,ll,O'IKa (OH HaITOJIH.HeTC.H KPOBbIO) OHM ITpeIT.HTCTBYIOT o6paTHOMY TOKY KPOBH.
<PyHK!IMH :3TMX KJiallaHOB: o6ecITelJMBaIOT TOK KpOBM TOJihKO B 0,Zl;HOM HarrpaBJieHHM -

113 JKeJiy,D;O'IKOB B aopTy M JierO'IHYIO apTepMIO.

<PyHI-C!IH.H cep,n;I1a: pHTMH'IHOe HarHeTaHMe ITO apTepM.HM KpOBM, ITOCTyITaIOIIICH K HeMy n:a BeH. BhIITOJIH.HeTc.H 3a C'IeT norrepeMeHHhIX coKpaIIIeHMH (

moJL)

H paccJia6JieHHH (

iJuacmo.1)

cuc-

MMOKap,n;a.

CHCTOJia H ,n;naCTOJia B HOpMaJibHbIX ycJIOBH.HX corJiaCOB8HbI M COCTaBJI.HIOT
!IHKJI pa60Thl cep,11,I1a. Ilpo,n;OJIJKMTeJihHOCTh cep,11,e'IHOrO IU1KJia 3aBHCMT OT qaCTOTbI COKP8IIIeHH.H cep,11,I1a (y 3,11,0pOBOro qeJIOBeKa
BpeM.H o,n;Horo !IMKJia aaHMMaeT

0,8

60-80

y,n;apoB B MttHyTy).

c.

0, 1 c, a oT,11,hrxaIOT 0, 7 c. JKeJiy,n;oq0,5 c. Bo BpeM.H Kam,11,oro coKpaIIIeHM.H meJiy,n;oqKOB B cocy,D;hI BbIT8JIKHBaeTC.H 70-80 MJI KPOBM. 3a 1 MMH cep,11,!le B3pOCJIOro
qeJIOBeKa (B ITOKOe) ITpOK81IMBaeT 5-5,5 JI KPOBI1. Ilpn cpH3MlJCCKOH Harpy3Ke
KOJIH'IeCTBO KpOBM yBeJIHlJHBaeTC.H ,n;o 15-20 JI/MMH (y CITOPTCMeHOB
30-40 JI/MMH).
B Te'IeHMe !IMKJia rrpe,11,cep,11,M.H pa6oTaIOT

Kn pa6oTaIOT

3 c,

a OT,ll,hIXaIOT

Tonbt cepiJu,a -

aTo 3BYKM, B03HHKaIOIIIHe BO BpeM.H pa6oThI cep,n;I1a.

Pa3Jilf'IaIOT ,n;Ba TOHa:

cucmOJ1U'l.PCKUU -

Hl13KMH, ITpo,n;omKHTeJihHhlM, ITO.HBJI.H-

eTC.H B H81IaJie CHCTOJibI meJiy,IJ;O'IKOB, CB.H3aH c COKP8IIIeHHeM MYCKyJiaTypbI CTeHOK

Cmpoenul' cepiJLfa

JKeJiy)J;O'lKOB l1 3aKpblT!1eM CTBOpOK
rrpe11cep.n;1rn-JKeJIYAO'lKOBhIX KJiarraHOB,
II iJuacmoJLU'leCKUU - Bh!COKHH, KOpOTKHH, IIOHBJ1HeTCH B HR'laJie .n;uaCTOJ1hl,
Kor11a 3aKpbIBa10TCH 3aCJ10HKH rro.nyJIYHHhIX KJiarraHOB aopTbI H .'JerO'lHOro
CTBOJia.
CepiJe'lllbLU mo.ll'lOK 06pa3yeTCJI
npu tt3Menentttt rroJiomenirn: cep.n;n.a
rrptt CHCTOJie, KOr,n;a .1eBbIH )IWJIY.l(O'leK
rrpnmttMaeTC:H K rrepe.n;neH: rpy .n;noH:
cTenKe tt «y.n;apaeTCH» 06 Hee. Orrpe.n;emieTcH pyKott na ypoBne rraToro Mempe6ephH CJieBa OT rpy.n;HHhl.
Aemo.Mamu3.M cepiJu,a - STO crroco6nocTh cep.n;ua pHTMH'lecm1 coKpa rn;aThCH 6e3 BHemnux pa3;:i;pamenni1:
rro.n; BJIHHHHeM HMIIYJihCOB, B03HHKaIOrn;ttx B HeM caMOM.
llcTOt:JHHKOM aBTOMaTII3Ma HBJIHIOTCH THIIIIt:JHhie MhIIIIet:JHhie KJieTKII npoBO,D;Hrn;eil CIICTeMhI (puc. 169). IlpOBO.n;arn;aH cttcTeMa cep.n;ua BKJIIOt:JaeT:

Cu nyc no-npeiJcepiJ n bLU y:Je.ll

459

3
P11c. 169.
1-

8

npoBOAHW,aH C~1CT8Ma C8PALJ,a:

sepXH5!5l no11a51 BeHa; 2 -

KYll5lpHbli:1 y3e11;
rnanaHb1;

3-

aTpVIOB8HH1-

aTpvtoseHTV1KYll5lpHb1e

4 - HO>KKa ny4Ka lvica; 5 -11eBb1i:1
6 - neperopOAKVI M8>KAY >K81ly7 - ceTb BOllOKOH nypKV!Hbe; 8 -

>K81lYA048K;
A04KaMvi;

npasb1i:i >K81lYA04eK;

9 - HVl>KH5!5l no11a5l seHa;

10- ny40K lvica; 11 - 88H8'1Hbti'I CV!Hyc;

(CHHOaTpttaJihHbIM «BO.l(lI12- cvtHOaTpvianbHbti:i y3e11; 13- nesoe
TeJih cep,Aet:Jnoro pttTMa» ), pacnpeAC8PAvte; 14 - npasoe npeAcepAvte
noJiomennhIH B cTenKe npaBoro
rrpe.n;cepmrn Mem.n;y ycThHMH
IIOJlhIX Ben, r.n;e B03HIIKaeT B036ym.n;enIIe (70-80 HMIIYJihCOB B MHHYTy), KOTOpoe pacrrpocTpaHHCTCH B MlIOKap.n; npe.n;cep.n;ttH:.
JI peiJcepiJno-J1Ce.TLyiJo'l1C06bLU y:JeJt ( ampuoeenmpuKy.llHpttbtil ), pacnoJiomeHHhIM TaKme B cTenKe npaBoro npe.n;cep,n;IIH, y ero rpanIIUhI c meJiy.n;o•IKaMII, 'Iepe3 KOTOpbIH B036ym.n;emrn npOBO.ll;IITCH.

K.llemKu npeiJcepiJno-J1CeJtyiJ0'1-1C06oio ny'l1W ( ny'l-OIC I'uca) u eio pa:Jeeme.lle/WH ( iJee HOJIC/CU 60.llOICHa JI yplCUHbe) - ,D;JIH pacnpocTpaneHIIH B036ym.n;emrn K Kap.n;IIOMIIOUIITaM meJiy,D;O'lKOB, r.n;e B03HIIKaeT coKparn;enIIe.
Ilpoxom.n;emrn HMIIYJ1hCOB t:Iepe3 npe.n;cep.n;no-meJiy,n;ot:JKOBhJH y3eJ1 3aMe.n;JIHeTCH, II03TOMY B036y)l(,l(eHIIe .ll;OCTHraeT MHOKap,Aa meJiy,D;Ot:JKOB Me,UJieHnee,
'ICM MHOKap,Aa npe,Acep,Atttt. B CBH3H c STHM COKpa11~aIOTCH BHa'laJie npe.z:i;cep,AHH, a IIOCJie meJiy,AO'IKII.
Ilpn B036ym.n;emrn cep,Au;a MeJK)J;Y pa3JIH'lHhIMH TO'lKaMH TeJia C03.n;aeTCH
pa3nocTh noTenuHMOB. KpttBaH H3MenenHH sToil pa3HOCTH noTenuttaJIOB BO
BpeMeHH - SJICKTPOKap.n;ttorpaMMa (SKI'). 8Kr OTpamaeT COCTOHHIIe Mhillll~hI
cep.n;u;a II CJIY)KHT IIOKa3aTeJieM ee ,n;eHTeJihHOCTH.

AnanwMUJl u <fm.1uo1wzuR 'te:wHe1w

460

Pery.ru1.u;m1 .i:i;eHTeJILHOCTH cep.i:i;u;a

1. H eplrnaR pe2y.'1.H4UR ocym;ecTB.'IHeTC.fl pecp.'IeKToptto BereTaTIIBHoi1

ttepB-

HOII CHCTeMOH' KOTOpaJI HaXO!l;HTCH TI0,11; KOIITpO.'IeM npO,ll;OJ'IrtrnaTOI'O
Moara, r11noTa.'IaMyca II KOphI 60.JihIIIIIX no.'IymapIIf1. IlapacuMnamuI£ec1cuiL omoe.'l (6JiyJK,n;aIOill;Hll HepB) IIHHepBIIpyeT CIIHYCHO-IIpen;cep,D,HhIH
tt aTp110BeHTpttKy.;'IHpHLIH Y3JihI, BhI3LIBa.fl cmimeH11e qacTOTbI tt c11.1b1

cep,n:e'IHhIX coKpam;eHm1. CIIJihHoe pa3n;pameHIIe 6Jiy)K,n;arom;ero uepBa
MOJKeT npIIBeCTII K OCTaHOBKe cepn;u,a. Cwnnamu'leCICUU omue.11 IIHHepBIIpyeT MIIOKapn; npen;cep,n;IIH II meJiyn;oqKoB, Bbl3hIBa.fl TIOBhillieHHe qaCTOTbl II CIIJihl cepn;eqHbIX COKpam;eHIIH.

2.

I'yMopaAblWJl peiyAR4U.fl ocym;ecTBJ'I.fleTc.fl qepe3 Kponh. AcJpe1taAUH,
Ca2+ BhI3hIBaIOT ycm1enIIe coKpam;eHIIH cep,n;u;a. A4emuAXOAUH, J{- TopM03HT pa6oTy cep,n;u,a.

BHyTpeHHHH cpe;::i;a opraHH3Ma

B1tympe1tHJlR cpeoa opiaHU.JMa -

3TO COBOKYTIHOCTb JKII,ll;KOCTeH (KpOBb,

JIIIMcpa, TKaHeBaJI JKII,ll;KOCTb), npIIHIIMaIOm;IIx yqacTIIe B npoll,eccax o6MeHa
Bem;eCTB II rron;,n:epmaHII.fl roMeOCTa3a (TIOCTOJIHCTBa) opraHII3Ma.

T1Ca1te6aR J1CU01Cocmb 06pa3yeTCH 6Jiaro,n:ap.H nepexo,n;y (cpIIJihTpau;IIII) m11,11;KOH qacTII KpOBII (nJia3Mbl) II3 KaTIIIJ'IJI.flpOB B TKaHII.
MecTonaxomn;eHIIe -

npoMemyTKII Mem,n;y KJieTKaM11 Bcex TKaHe11:.

HcTO".fHIIK o6pa3oBaHIIH -

nJia3Ma KpOBII 11 npo,n;yKThI m113ne,n:eHTeJILHoc-

TII KJieTOK. 06'heM y B3pOCJIOro qeJIOBeKa COCTaBJIHeT

20

.'I.

CocTaB: BO,D,a, paCTBOpeHHhle B Hett TIIITaTeJihHhie Bem;ecTBa II HeopraHIIqecKue Bem;ecTBa, KIICJiopo,n;, C0 2 , npon;yKThI pacna,n;a, Bh1,neJI.HIOm;11ec.H 113
KJieTOK.
<l>yHKIJ;HH:

npoMemyToqHa.fl

cpe,n:a

Mem,n:y

KpoBeHOCHhIMII

cocy,n;aMlI

II KJ'IeTKaMII opraHlI3Ma; nepeHOC 113 KPOBH B KJieTKlI KlICJIOpo,n:a, 113 KJieTKH
B KPOBh -

yrJieK11cJioro ra3a.

BoJILIIIaH qacTh TKaHeBoil m11,n:KOCTII B03Bpam;aeTcH B KpOBh, npoH11KaJI qepe3 3H,ll;OTeJI11i1: KpoBeHOCHhIX Kamrnmi:poB • .D:pyraJI qacTh, He ycnenaJI nepHYThC.fl B KpOBh, co611paeTCJI Mem,n:y KJieTKaMH TKaHeH, r,n:e 6epyT Ha•rnJio JIIIMcpaTIIqecKIIe cocy,n;hI.

JluM<fJa -

3TO )KlI,ll;Ka.fl coe,n:IIHHTeJihH8H TK8Hb, u;MpKyJI11pyrom;a.fl B cocy-

,n:ax JIHMcpaT11qecKOH CIICTeMbl.
11CTOqHMK o6pa30BaHIIJI: B Me)KKJieTOqHhIX npocTpaHCTBax 38 cqeT TKaHeBOH m11,n:KocT11 6epyT HaqaJio JIMMcpaTMqecKHe cocy,n:hr H npoHH3hIBaIOT noqT11
Bee opraHbl, 3a IICKJIIOqeHIIeM KOCTeil, BOJIOC, KO)KII, porOBH:ll;bl. 3a CYTKII y qeJIOBeKa 06pa3yeTCJI

2-4

JI JIIIMcpbI. JlHMcpa, OTTeKaIOm;aJI OT pa3HblX qacTeH

TeJia, IIMeeT Heo,n:1rnaKOBblll COCTaB, KOTOpblll onpe.a;e.irneTC.fl cneu;HcpJiqecKOH
,n:eJITeJibHOCThlO pa3HblX opraHOB H TKaHeH.

BnympeHHJlR cpeiJa opzanUJMa

461

CocTas: so.na c pacTsopeHHhLVrn B He:ii rrpo,nyKTaM11 )Kl131ie.neHTe.1LttocTtt (pacrra.n;a opraHIIqecKIIX sew;eCTB), 6e.'!Klf -- 1-2 '1~.' J!l1:vnpo1~HTI.I, .'ICHKO~HThl.
Ilo COCTaBy JIHMCpa OTJ!HqaeTC.H OT TKaHeBOM )KM,UIWCTH 60J1ee Bh!C01{11M COJrepmamieM 6eJIKOB (2 r'X, ). XuMuqeCKHM COCTaB JIHMcpbI 6.TIH30K TaKme K COCTaBy
flJia3MhI Kp0Bl1, HO B Hett co,nepJKHTC.H MeHhllle (B 3-4 pa3a) 6e.'IKOB, 1103TOMy
JII1Mcpa o6Jia.n;aeT He6o.'IhllIOM B.H3KOCThIO. B J'IttMcpe co,nepm11TC.H clm6p1rnoreu,
oHa crroco6Ha csepThIBa'fhCH, HO ropa3.n;o Me,nJJemree, qe:vr KpOBh. Ilpu 110speJKi\eHl1II KpOBeHOCHhIX KaflHJL'l.HpOB qllC.'IO JIIIMcpOUJITOB B .'IHMCpe yBec1Hq11saeTc.H.
cflyHKUHH
Bo3spaw;eH11e B KpOB.HHoe pycJio TKaHesoH: m11,nKOCTu.
cfltt.'IbTpauHa 11 o6e33apam11BaH11e TKaHeBOM )KH,ll;KOCTH, KOTOpbIC ocyw;eCTBJI.HIOTCH B J111Mcpan1qecK11x y3.1ax, r.ne Bhrpa6aThIBaIOTC.H B-.111Mcpou;ttThI.
Yqacnie B o6MeHe semecTB - m11pos.
°YqacTtte B TpaHcnopTe n11TaTeJihHhIX sew;ecTB (.n;o 80 ';{, JKMPOB, BC3Chl·
saeMhIX B KIUneqH11Ke, nona,naeT qepe3 JIIIMcpaT11qecKyIO CHCTeMy).
JI11McpaTttqecKaa CHCTeMa Tecuo CB.H33Ha no CBOeMy CTpoeHHK> II cpyHKUH.HM
c KPOBeHOCHOM CIICTeMOM.

CTpoenue JIHM<l>aTuqecKoD: CHCTeMLI

1. 3a.MKttymaR cemb JtU.MrjJamu'leCKUX KanuJtJtRpo6 (p11c. 170), Ha Kouuax

2.

3.

KOTOpbIX B JIHMcpaTttqecKHX MellloqKax npOIICXO,ll;HT cpHJih'fpau11.H TKaHeBOH :>KH,UKOCTII.
Jlu.MrjJamu'leCKue cocyiJbt - o6pa3yIOT B03BpaTIIYIO CHCTe:vi:y c HH3KHM
,naBJieHIIeM, CJJymaw;ym ,ll;JIH oqttw;eHHH :vi:eJKKJie'foqHOM )KH,ll;KOCTH.
B cTeHKax J'IHMcpaTIIqecKnx cocy.n;os Haxo.n;nTca MHomecTso no.1y.1yuHhIX K.'JanaHOB, cnoco6CTBYIOlll;HX ,UBHJKeHHIO JlIIMcpbI no HanpaBJieHIIIO K cep.n;uy.
Hatt60J1ee KpynHLIMII J1HMcpaTnqecK11MH cocy.n;aMII .HBJI.HIOTC.H npaentit
JtU.MrjJamu'lecKuit npomoK, co6npa10w;ttH: JIIIMcpy n3 npasoii: cTopoHhI roJIOBhl, npasoH: pyKn II npasoH:
BepxHeM qacTn TeJia H H3JIHBaIOru;IIM ee B 60JlhllIOM Kpyr KPOBO·
o6paw;eHH.H B MecTe coe.n;nHeHH.H
npaBOM IlO,ll;KJIIOqIIqHOM H .HpeM HOM seH, II ipyottoit npomoK,
npIIHIIM3IOlll;HM JIHMcpy H3 Bcex
.n;pyrnx qacTeH: TeJia H B03Bpaw;a10w;nli ee B KpOBeHOCHYIO CIICTeMY B MecTe coe,nttHeHna JiesoH:
Pi.tc. 170. Ha4ano m1Mcj>arn4ecK1.1x
no.n;KJIIOqHqHOM H apeMHOM BeH.
Kan1.1nm1pos: 1 - Kan1.111m1p: 2 - nna3Ma:
Jlu.M<jJamu'lecKue y3Jtbt pacnoJ1a3 - KPOB5!Hb1e KneTKl.1; 4 - 111.1Mcjla; 5 - Ha4ara10Tca no xo.n;y JJHMcpaTttqecKHX
no 111.1Mcparn4ecrnx Kan1.1111151pos; 6 - TKaHeBafl
cocy.n;os, KaK npasnJio rpynnaMH.
0.n;Hn, 6JtuaKo om noBepxttocmu

AnamoMUH u ljJuauo;wzuH 'teJwae1w

462

(rneii:ttbJe, no.n;Mhrrne'lttbre, naxOBhre), Jier1w npo~ynhrna10TCH BO npe:vrn
6oJie3HH, a .n;pyrtte pacnoJiometthI

iJ1y601w 6 rneJte

(neepoBhI 6m-1wK11) -

JIHMcpaTII'leCKHe Y3JibI B CTCHKax TOHKOrO KHIJlC'lHHKa, 3arr~mu;aro~He OT
Bpe,D,HhIX MllKpoopraHH3MOB H norJIOIIJ;aIO~He

TOKCH'IHhie

JlnMcpa,

Be~eCTBa.

npOXO,ll,H

qepe3

8Tll

ya:Ihl,

cpHJibTpyeTCH OT np11Mece:ll (pnc.

171).

B ;rnMcl>aTnqecKHX y3;1ax cpHJih'l'paMH c.'Iy/I<aT CC'l'OlJKII 113 BO.'IOKOH. 3.necb me npOHCXO,ll,llT cparOIJ;llT03, ocy~eCTB.'IJieMb!H MaKpocparaMH, llJill YHll'lTOJKeHHe Ilpll IIOMO-

~ll

T-KJieTOK.

J111Mcpou,HThI

o6pa3yIOT

aHTHTeJ'Ia II.'Ill TIOKll,ll,aIOT y3eJI ll nepeMe-

~aIOTCJI BMeCTe c JllIMcpoH: no TeJiy.
06'beM KpOBII, BbITaJIKHBaeMhlll cep.nIJ,eM B 60Jibll10ll Kpyr KpOB006pa~eHirn,
paBeH

4,5

c:vr3 /MHH.

B

JIHMcpaTn'lecKoM

y3Jie cparou,HThI 11 JIHMcpOIJ.HThI y,ll;aJIHI01'
TOKCHHhI ll naToreHHbie opraHll3Mhl.
ToK JIIIMcphr ot.IeHh Me,n;JieHHhitt.

J1CeHue

,Tl,6u-

JlUM,(jJbl o6ecneqHBaCTCH pllTMll-

qecKHMll

COKpa~eHHHMII

CTeHOK Kpyrr -

HbIX JllIMcpaTH'leCKllX cocy,ll,OB, COKpa~e
HlleM

CKeJieTHbIX

Mh!WIJ,,

HaJilllJHeM

KJianaHOB B CTeHKaX JII1McpaTH'leCKllX cocy.n;oB,

npncachIBaIO~HM

.n;eli:cTBHeM

rpy,ll,HOll KJieTKll B CB.H31I c OTpHIJ,aTe.'IbHbIM BHyTpHrpy,ll,HblM .n;aBJieHHeM.

KpoBeTBOpHhie opraHhl
RpoBeTBOpHbre opraHhI:

a
P111c. 171.
nHMcPbl: a -

KpaCHblU 1wcrnHbtU MO.Ji! -

CxeMa HanpasneHHS'I TOKa
pacnono>KeH1-1e rpynn n1-1M¢arn-

Y8CKHX y3nos;

6-

pacnono>KeHHe o6nacreV!,

KJieTKll, KOTOpbie yqaCTBYIOT B pa3Bl1THH
BCeX cpopMeHHbIX 3JieMeHTOB KPOBH.

JI u~uparnu'lec1Cue y3JZbt -

OTKYAa co61-1paeTCS'I IlHMct:>a B rpyAHOCi H npaBb!Ci n1-1M¢>aTHY8CKHe npoTOKH. 1 n1-1Mct:>arnYeCKHe y3nbt;
Y3Ilb1;
Ilbl;

2-

LU8VJHbi8

nOAKPblflbl.l,OBbi8

npaBbl~ nOS'ICHHYHblVJ CTBOn;

BblVJ nOS'ICHHYHbiCi CTBOn;

7-

6 - ne-

KHW8YHblH

CTBon; 8 - rpyAHOCi npOTOK: 9 - MeCTO BnaA8HHS'I rpyAHOro npOTOKa;

10 -

MeCTO BnaAe-

HHS'I npasoro nHM¢>aTHY8CKOro npoTOKa

opraHhI, B KO-

TOpbIX npOHCXO,ll,llT aHTMreH3aBMCHMaH
.n;mp<)lepeHIJ,llpOBKa JIHMQlOIJ,llTOB KpOBll.

CeJZe3eJtlCa -

3 - nax0Bb1e y3nb1; 4 - nOKTeBble y3-

5-

pacnoc10-

meH B KOCT.HX, r.n;e HaXO,ll,.HTC.H CTBOJIOBhJe

caMoe 6oJihIIIOe B TeJie

JillMcpanI'leCKOe o6pa30BaHMe,

CJIYJKMT

,ll,JI.H cpllJih'l'paIJ,Hll KpOBH (a He JillMcph!,
KaK

B

.11;pyr11e

JIHM<j)aTtt'lecK11e

Y3JihI).

ceJie3eHKe B036y,n;ttTeJin 3a60JieBan11i\:,

naxo.n;a~HeC.H

B KpOBll,

TIO,ll,BepralOTCH

Kpo6em6opnbte opzau bl

463

B03.n:ei1cTBIIKl JIHMCj)Oll,HTOB, KOTOpLie Bbipa6aThIBaIOT COOTBCTCTBYKlilJ,He
aHTHTCJia.

B

ceJIC3eHKe ocyruecTBJIHeTCH cparOIJ,HT03 6aKTep11:i1 n OTIKHB-

mnx 3pttTpou;IITOB. Otta .n;ei1cTByeT KaK .n;erro Kpontt, a BO spe:\rn pa 3 snTHH IIJIO.n,a npo.n:yunpyeT KpaCHLie KPOBJIHL!e Te.1Lna.

Kpoeb -

3TO mtt.n;KaJI coe.n,HHHTeJILHaR TKaHL, ll,HpKy.•npy10ru;aH B cocy.n:ax

3aMKHYTOM KpOBCIIOCHOH CIICTeMhI.
<l>yHKll,HII
TpattcnopT rrnTaTeJihHhIX seu~eCTB (Tpoq:rnqecKaR cpyHKll,lrn).
TpattcrropT KOHeqHhIX npo.n:yKTOB MeTa6oJitt3Ma (3KCKpeTopttaH q)yHKI!HH).
TpaHcrropT ra3oB -

KHCJiopo.n;a H .n;ttoKctt.n:a yrJiepo,n;a (,n;h1xaTe.1LttaH

cpyHKUlrn).
TpattcrropT ropMOHOB (ryMopaJILHaJI pery.•RUHR).
3amIITHaH cpyHKll,HJI -

cparoll,IITapHaJI aKTIIBHOCTh JiettKOll,HTOB (K.leTO•I-

HhIH HMMYHIITeT) H Bbipa60TKa JIIIMcpO!JIITaMII aHTHTeJI, o6e3BpemnnmoIUHX reHeTnqeCKII qymepOJJ;Hhie BemeCTBa (ryMopaJihHhlll IIMMYHIITeT),
CBepThIBaHHe KPOBH, rrpeIIHTCTBYIOmee KpOBOIIOTepe.
foMeOCTaTI1qecKaJI cpyHKll,lI.H -

rro.n:.n:epmami.e IIOCTOHHCTBa BHyTpeH-

Heli cpe,n;LI opraHlI3Ma, rrpnro.n:Holi ,n;.'I.H KJieTOK B OTHOWeHHII HOHHoro
COCTaBa, KOHUeHTpau;IIH no.n:opo,n;HhIX HOHOB H ,n;p.

Cocmae Kpoeu: mtt.n;Koe MemKJieToqttoe seruecTBO - nJta3.Ma n B3BeweHHLie B Heil: cpopMeHHhie 3.TJeMeHThl: Jpumpou,umbt (KpacHbie KpOB.HHbie TeJihIJ,a),
JleUJCOU,Umbt (6eJihie KPOBJIHbie Te.lbll,a) H mpo.M6ou,umbt (KpOBHHhie IIJiaCTHHKH) • .D:OJI.H IIJia3MhI COCTaBJI.HeT OKOJIO

55 'Yo,

cpopMeHHhIX 3JieMeHTOB -

06ruee KOJmqecTBO KpOBH B opraHH3Me B3pOCJioro qeJIOBeKa MaCCbI TeJia, TO eCTh rrpnMepHO

4,5-6

JI.

B IIOKOe 40-50 'X,

OKOJIO

45 °f<,.
6-8 %

Bceil: MaCChl KpO-

BH IJHPKYJIIIpyeT ITO cocy.n:aM, a OCTaJihHaJI KPOBh CKaIIJIIIBaeTC.H B KaIIIIJIJIJipax
(.n:enoHIIpyeTCR). OcHOBHhre .n:erro Rposn: ceJie3eHKa, rreqeHh, Koma. IIoTep.a
1/8 ofrheMa KpOBII MomeT npnBeCTII K rII6eJIH opraHH3Ma.

Cocmae nJta3Mbt: BO,n;a

(90-92 % )

I1 cyxoe BeII~eCTBO

(8-10 % ),

COCTOJI-

ruee I13 opraHIIqecKHX coe.n:HHCHIIH II MIIHepaJihHbIX BemeCTB.

BeJtKU

(7-8 % )

nJia3MhI KpOBII. HecROJihKO .n:ecRTKOB pa3.1IIqHbIX 6e.TIKOB

o6'he,ZJ;HHeHhI B TPH OCHOBHbie rpynrrhr: aJib6yMHHhI (OKOJIO

(233 % ) H

4,5 % ),

rJio6yJilIHhl

(0,2-0,4 % ).

AJih6yMIIHhI H cpn6pIIHOreH cnHTe3npy-

IOTC.H B KJieTKax rreqeHH, rJio6yJIHHhI -

He TOJIJ,KO B neqeHII, HO I1 B ceJie3eHKe,

qm6pttHoreH

KOCTHOM M03re, JIHMcpaTnqecKHX Y3Jiax.
<l>yHKI~Hll 6eJIKOB:

yqacTBYIOT B no.n:.n:epmaHIIII pH KpOBH Ha IIOCTO.HHHOM ypoBHe;
npn,n;aIOT B.H3KOCTh KPOBII, qTo HMeeT BamHoe 3HaqeHne B rro.n;.n:epmaHHH
apTepttaJihHOro .n:asJieHrr.a;
o6ycJiaBJIHBaIOT OHKOTttqecRoe .n:asJieHIIe, KOTopoe onpe,n;eJI.aeT o6MeH
BO,ll;bl Mem,n;y KpOBbIO I1 TKaH.HMH;
yqacTBYIOT B CBepTbIBaHIIII KpOBH;
cpaKTOpbI IIMMYHIITeTa;
CJiymaT pe3epBOM ,ll;JI.H IIOCTpoemrn 6eJIKOB TKaHeli.

464

AnamOMUH u <Pu3uo1wzuH 'le/W6eKa

CTJI33Ma, .111mettHaH qm6p11ttoreHa, Ha3hIBaeTCH CbtBopomKou llpOBU. B OTJnrqne OT nJia3MhI ona He cBepThrnaeTcH.
Yi.rteBOUbt rwa3MhI Kpos11 rrpe.n;cTaBJieHhI rJiroK03ott B KOH11,eHTpm~11n 80-120 MI' (0,12 'X, ), .WnUUbl COCTaB.JlfllOT 0,5 % •
Mu11epaJ1bllbte BeUJ,eCmBa rnia3MhI (0,9 %): KaT11onhr Na 1 , K', Ca2+, Mg2,_
n aH11onhr Cl-, HC0 3-, HP0 4 -. l-IcKyccTBeHHhie pacTBOphr, 06.'la.n;aron~ne o.n:ntta1<0BhI:vI c KpOBhfO OCMOT11qecKMM .n;aBJI€H11€M, TO eCTh co.n:epman~ne paBHYIO
e:H: KOHf~eHTpa11,nro CO.'leM, Ha3hIBaIOT U300CMOmU'leC/CU.MU MJ111 U30mOHU'leCf(UJl1U. ll30TOH11qecKMM ,D;.'l.H TeITJIOKpOBHhIX )KMBOTHbIX M qeJIOBeKa .HBJI.HeTCfl
0,9%-Hhitt pacTBOp NaCl. TaKoil: pacTBop Ha3hIBaroT rpu3uo.r10iu'leC1CUM. PacTBOphr, 11Merow;ne 60Jibillee OCMOTnqecKoe .n;aBJieHne, qeM KpOBh, Ha3hIBaIOT
zunepmOllU'lPC/CU.MU, MeHhillee zunomOllU'leCKU.MU. 3pnTp011,HThI B M30TOH11qecKOM pacTBOpe coxpaH.HIOT CBOIO cpopMy' B r11rrepToH11qeCKOM paCTBope
CMOPW:11BaIOTC.H, a B r11ITOTOHMqecKOM - Ha6yxaroT M JIOITaIOTC.H. 0Tcro.n:a CJie.n;yeT HC06xo.n;MMOCTh rro.n:.n;epmaHM.H COJieBoro COCTaBa IIJia3Mbl KpOBH Ha IIOCTO.HHHOM ypostte.
KpoBh qeJioBeKa nMeeT cJia6ow;eJioqttyro peaK11,1110 (pH apTepnaJ1hHoi1: KpoBM - 7,4; pH BCH03HOM KpOB11 - 7,35).
«l>opMeHHLie 3JieMeHTbl KpOBH

3pumpou,umbt KpacHhie KpOB.HHhie TeJ1b11,a, nMeroT cpopMy 6e3'hH.n;epttoro
,D;BO.HKOBOrHyToro .UMCKa (.n;11aMeTpOM 7 MKM, TOJ1W:11HOH - 1-2 MKM), qTQ
yBeJinq11BaeT ITOBepXHOCTh .n;ucpcpy3HM ra30B II o6ecrreqnBaeT rrpOHHKHOBeHMe
B KarrnJIJIHphr. B 1 MM" KPOBH qe"10BeKa co.n;epiKHTC.H 4-5 MJIH ap11Tpo11,nToB.
06pa3yIOTC.H apMTpOll,HThl B KpaCHOM KOCTHOM M03re, paapyillaIOTC.H B rreqeHM
u ceJie3eHKe. IIpo.n;oJim11TeJihHOCTh mtt3H11 - OKOJIO 120 cyToK.
<l>yHK11,11li 3p1ITp011,HTOB: ocyw;ecTBJiflIOT rrepeHOC K11C.'lopo.n:a li ,D;HOKCH,D;a yrJiepo.n;a. LJ:brXaTCJibHhlll rr11rMeHT apuTpOll,HTOB - ze.11WZJ106Ull - JierKO rrpHCOC,D;11H.HeT 11 oT.n;aeT KHCJiopo.n: 6e3 113MeHeHH.H BaJieHTHOCTH meJie3a. Co.n;epmattue
reMorJio6uHa B KpOBH 3,D;OpOBOro qeJIOBeKa KOJie6JieTC.H OT 14 .n:o 16 r% (140160 r/JI). O.n;11H rpaMM reMorJio6Irna crroco6ett CB.H3aTh 1,3 M.1 KHCJIOpo.n;a.
feMOrJI0611H B KpOBeHOCHhIX KarrunJiapax JierKHX HaChiw;aeTC.H K11CJiopo.n;oM
li rrpeBpaw;aeTC.H B OICCUZeMOZJL06u11 (Hb02), rrpn.n;arow;uil: KpOB11 .HpKO-aJihIM
11,BeT. B TKaH.HX 11 opraHaX KHCJiopo.n; JierKO OTw;errJI.HeTC.H; reMorJio6HH BOCCTaHaBJIHBaeTC.H 11 rrp11coe1.1:uttaeT ,n;110Kc11.n; yrJiepo.n;a, rrpenpaw;aach B 1Cap6ozeMOZJL06u11 (HbC0 2). ll;BeT KpOB11 (BeH03HOH) TeMHO-KpacHhitt. B nerK11x .n:noKc11.n; yrJiepo.n:a on1~errJiaeTca OT reMorJI0611Ha, BoccTaHaBJIHBaeTC.H 11 rrp11coe.nuH.HeT KllCJiopo,n. feMor.'lo611H crroco6eH coe.n;11H.HThC.H c yrapHhIM ra3oM (CO),
o6paaya rrpoqHoe coe.n;11Hett11e - 1Cap601Ccuze.MOZJL06u11 (HbCO). B pe3yJihTaTe
reMOrJI06HH Tep.HeT crroco6HOCTh CB.H3hIBaThC.H c 02, 11 HacTyrraeT y.n;yllihe.
"YMeHhllleHue KOJIIfqeCTBa ap11Tp01I,HTOB B KPOBH H CHHJKeHHe co.n;epmaHtt.H
reMorJI0611Ha tta3hIBaeTc.H aHeM11eil: ( «MaJioKpoBHe» ). B HOpMe reMorJI0611H
y mettw;11tt 120,0-140,0 r/JI (12,0-14,0 r% ), y MymqnH 130,0160,0 r/JI (13,0-16,0 rcYo ).

465
,UJrn /~ttaruocTHKH IIaTOJIOrttLieCIOl:X l1BJieHHW HCllOJih3Y!OT lW.'lli'lllHy CKOpOCTH occ,riaumi 3p1npounTOB (C03) Kpostt.

3-10

paBua

MM/LI, y memu,11u -

7-12

B

uopMe BeJrnLittHa C03

MM/LI. YBb111L1eu11e

C03

y

MYJKLIHH

JIBmleTcn

np113uaKoM Bocrra.1eu11n.

Jiei:iKou,umbt -

a.11;epuh1e 6e.1h1e KpOBl1Hhie Te.1Ihl1,a, ue HMeIOmne nocToan-

noi1: ctiopMhI, crroco6Hhie rrpOHHKaTb MCJK/l,Y K.lJCTKaMH 3H.ll;OTC.1!1}f Karrn.1.1Hp<m
11 aRTHBHO rrepeMem,aTbca so Bee <IaCTH TeJia.

9

B 1

MM" KPOBl1 co,riepmttTCH

4- _

ThIC. ;1eii:KOl1,HTOB, o,riuaKO HX LIHCJIO MOJKeT H3MeHaThCH rroc.1e np11e:vrn IIH-

Ill,H' MhlllICLIHOH pa60ThI' BO BpeMa CHJlbHhlX 3MOI1HM ( r/JU.WO.'l02U '{('(;ICU ii .:1eil1wu,u moa). Ilpl1 HHQJCKI1HOHHhIX 3a60JieBaHHHX HX LIHCJIO pe:mo YBC.1HLIHBaeTca ( namO.'l02U'leCKUU JleUICOU,UnW3). Ilp11 JJyqeBOR 60J1e3HH, orryxo.1JJX
ua6mo,riaeTca 3Ha<IHTCJihHOe yMeHhllleu11e q11c;ra JieiiKOUHTOB ( JLPu1wneHu5l}.
JieiiKOUHThI o6pa3yJOTCH B KpacHOM KOCTHOM M03re, ce.1Ie3eHI-ce, JIHM<!mTHLieCKHX Y3JiaX, a pa3pylllaJOTCl1 B ceJie3eHKe H oqarax BOCITaJICHHH. Ilp0.11;0J1JKl11'CJihHOCTh JKH3HH

5-9 cyTOK,

HCKOTOPhie BH,ll;hl JICHKOUHTOB cyID,eCTByJOT ro.n;aMH.

11eaepuucm1>1e
tteumpo<jJU.rta.MU (50-79 •y., BCCX JlCHKOUHTOB), 303UllOr/JUJlll.MU (0,5-5 % ), 6a30<jJU.'ta.wu (0-1 % ).
B rpynny ne3epu11cThIX JiettKOI1HTOB Bxo,riaT .Mo1wu,umbt (3-11 'X,) H .rw.Mr/Jou,umbt (19-37 %, ). y 3,ll;OpOBhIX mo,n;e:H COOTHOllleHHe Mem,riy BCCMH BH.ll;aMn .1ei1:Jiei1:KOI1HThI ,n;e.1aT na ,n;Ba Tttna:

3epnucmbte

(rpauy.1ou11ThI) u

(arpanyJIOUHThI). 3epHHCThle JieHKOUHThI npe,n;cTaBJICHhl

KOI1HTOB .ll;OBOo'lhHO IIOCTOHHHO, H l13MeHeHHe ero Co'IYJKHT npl13HaKOM pa3JIH'lHhIX 3a60o'ICBaHHH. IlpOl1,CHTHOe COOTHOllleHHe pa3HhIX Bll.ll;OB o1CHROI~HTOB Ha3hIBaCTCl1

.r1euKou,umapHOU <jJop.My.'lOU.

cDyuKunn JieliKOUHTOB -

3aID,HTa OT M11Kpooprau113MOB, Liymcpo/l,HhIX 6e.1KoB,

HHOpO.ll;Hh!X Te.1, KOTOpaa BhlIIOJIHHeTca 3a CLieT cpa!'OIJHT03a l1 Bbipa6oTKH aHTUTe.'l.

<Pa2ou,umo3 TPHK.'ICTOLIHOe

II. IL

TIOI'JlOID,eHHe JlCRKOI1HTaMH qymepo,n;Hh!X qacTHI1 H HX BHYnepeBap11BaHHe.

Me<IHHKOBhIM.

5IBJICHHe

<)JaI'OIJHT03a

Ha1160Jibllle:li cnoco6uocThJO K

6brno

OTKph!TO

qiarou;HT03Y o6Jia,riaIOT

nei1:Tpocp11Jib1, MOHOUHThI 11 303HHOQJHJihI. OuII o6ecneLittBaIOT

K.'lemo'lHhtU u.M-

.11iyttumem.
,Upyro:li QJOpMOH 3aID,HThl l1BJIJICTCl1

2y.MopaJlbHbtU U.M.Myttumem,

JIHCMhiiI .1IHMQJOI.l,HTaMH. JlHMQJOIJ;HThl -

OCYII1CCTB-

ueuTpaJihHOe 3BCHO HMMYHHOW CHC-

TCMhl oprau113Ma. Ou11 06pa3y10T 3aID,HTHhie 6eJIKH -

anmume.rw,

HJIH IIMMY-

uorJio6yJIHHhI, KOTOphre pa3pylllaJOT 'ly)Kepo,n;Hhie 6eJIKH, 06ecne<I11BaIOT peaK11,1110 OTTOpJKCHHa TpaHcrrnaHTaTa l1 YHH'lTO)KCHtte eCTeCTBCHHbIX MyTaHTHh!X
KJICTOK. Jl11M<}Jou11Th1 06Jia.11;a10T HMMYHHOH naMaThIO, TO ecTh cnoco6HocThIO
OTBe'laTh YCHJICHHOH pemnvi:e:li Ha TIOBTOPHYIO BCTpeqy c qymepo,riHhIM TCJIOM.
3Ty QJYHKI.l,HJO OHH BhlllOJIHaJOT 6Jiaro,n;apa TOMy, LITO, B OTJIH'IHC OT .11;pyr11x
JieHKOIJHTOB, )KHBYT He HCCKOJlhKO ,n;He:li, a

20

H 6oJiee JieT (HeKOTOpbie Ha rrpo-

TaJKeHHH BCe:li JKH3HH 'lCJIOBeKa).
Ilo QJYHKUHH 11 MecTy co3peBaHna JIHMQJOUHThI pa3,n;eJIHIOTCa Ha

mbt

(THMyc3aBHCHMhre) 11

B-J1u.11t<jJou,umbt

T-JLu.M<jJou,u-

(6ypco3aBHCHMhie). 06a Btt.n;a JIHMQJO-

UHTOB o6pa3yIOTCH B KOCTHOM M03re 113 CTBOJIOBhIX KJieTOK-npe,rirueCTBCHHHU.
,U11<)J<)JepeHI1HPOBKa T-JIHMQJOI1HTOB nponcxo,n;11T B BHJIO'IKOBOH me.11e3e (TttMyce).

466

AnamoMuJl u <jJu3uo1wzuJl •te1w6eKa

Pa3JHI<JaIOT tiecKOJihKO pa3HOBH/lHOCTei1.

T-KuJt.rtepbt (y6:nli:u;hr) crroco6HhI y611BaTh qymepoii;tti,re K.IeTKH, narrp:nMep 011yxoJieBbie,

KJieTKH-MyTaHThI,

KJieTKH

qymepo,n:HbIX

TKaHeH

TP8HCllJI8HT8HTOB.

T-cynpeccopbt (yrneTaTeJu:1) 6J10K11pyIOT qpe3MeptthJe peam.i:m:1 B-JIHMc}Jou;HTOB, rro.n:.n:epm11Ba.H rapMOHH'Iecrwc pa3BHTHe HMMyrn:1TeTa.

T-xeJtnepbt (rroMOID;HHKH) peryJmpyIOT peaKD;HH HMMYHHTeTa. Ilptt 6oJre3HH CIIMJJ: nopamaIOTC.H T-xe"·:mephr, <JTO noii;aBJI.HeT HMMYHHTeT.
B-JIIIMQJOD;IIThI npoxo.11;.HT ,n:IIc}Jc}JepeHii;ttpoBKY B JmM¢011;i:uoli: TKaHH KIIIIIe<JHHKa, anrreH.D;HKCa, MI:IH.ll;aJIIIH, He6HO-rJIOTO'IHOrO KO.llhIJ;a. 0cHOBH8.H QJYHKD;H.H B-JIIIMQJOD;IITOB -

C03.D;8HIIe ryMopaJihHoro IIMMYHHTeTa nyTeM Bhrpa6oT-

KH aHTIITeJI. IIocJie BcTpe'III c aHTIIreHOM B-JIIIMQJOD;HThI MIIrpIIpyIOT B ceJie3eHKY,

JIHMcpaT11'lecKne

y3JihI,

r.ll;e

pa3MHo:arnIOTC.H

n

npeBparu;aIOTC.H

B IIJI83M8Tll'IeCKIIe KJJeTKll, 06pa3yroru;11e aHTHTeJia.

Ji.M.MyHumem -

CIIOC06HOCTh opraHH3M8 3aID;IIID;aThC.H OT JKIIBbIX TeJI, Be-

ru;eCTB, KOTOpbre HeCyT reHeTII'leCKH 'lymepoii;nyIO IIHQJOpMaD;IIIO (6aKTepIIH,
BIIpychI, 6eJIKII, KJieTKH-MyTaHThI H ii;p.).
Pa3JIII'IaIOT cJie)J;yIOru;IIe 3aID;IITHhie MexaHII3MhI, 06ecrre'l11Baroru;11e IIMMyHnTeT.
BocnaJI1:1TeJihHa.H peaKIJ;ll.H, ocHoBaHHa.H Ha c}Jarou;IITapHoli: aKTIIBHocTII JieHKOD;IITOB. IlpH 3TOM JieHKOD;IITbI rII6HyT, OCB06om.n;a.H JIII30COMHhie c}JepMeHThI, pa3pyIIIaroni:IIe OKpymaIOru;lle TKaHll. B O'Iare BOCIIaJieHW.H B03HllKaeT IIOBbIIIIeHlle TeMnepaTyphI, IIOKpacHeHIIe 11 c}JopMIIpyeTC.H

rHOHHllK.

fHOH

-

3TO CICOOJl.eHIIe rrorII6IIIHX

JieHKOD;IITOB

11 rrpo.n;yKTOB pacrraii;a TKaHeli:.
Crreu;IIQJH'leCKIIe MexaHH3MhI pa3BHBaIOTC.H B OTBeT na rrpoHIIKHOBenIIe IIHQJeKD;IIOHHbIX areHTOB, B BH,n:e KJieTO'IHoro 11 ryMopaJibHOro IIMMYHIITeTa.
B 3aBIICllMOCTII OT rrpoIIcxomii;eHII.H pa3JIII'IaIOT c.11e11:yroru;tte 6Udbt U.M.MyHumema:
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aTOM

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200-

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467

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11 TPOM60ul1TOB BhICB06om.n;aeTC.H mpo.M60n.llacmu1-t, KOTOpbIM COBMCCTHO conpe.n;eJieHHhIMH rrna3MeHHhIMl1 cpaKTopaM11 (VII, V, X) n 110HaM11 Ca2- cnoco6cTByeT o6pa3oBattmo cpepMeHTa npompo.M6u1-ta:Jbt (cpa3a I). Ilp0Tpovr611tta3a rrp11
y 1rncTn11 cpaKTopoB V, X n Ca 2 + rrpespamaeT tteaKTl1BHhitt cpepMeHT npompo.w6uH, 06pa3yeMhIH Ko'IeTKaM11 rre'IeH11 rrp11 yqacT1111 BHTaM11Ha K, B aKTl1BHhitt
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13 rrJia3MeHHhIX cpaKTopos. feHeT11'!ecKn:li .n;ecp11u11T cpaKTopos VIII, IX
11 XI npttno.n;nT K rrecsepThrnaeMOCTl1 Kpos11 reMocp11JittlI cpopMbI A,
B 11 C COOTBCTCTBeHHO.
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B Kpos11 HMeeTCH TaKme aHTHCBepThrna10maa cttcTeMa. O.n;ttnM 113 MOII~HhIX
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AnamoMu.n u <Pu.1uo1wzu.n "Le1w6e1w

468

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l1 TJIJl<eJihIX oc.1omHeH11ii

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1903

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aiz.1110muHoieHbl ( a1tmuieubt)

A 11 B. B ::ip11Tpou11-

Tax pa3HhIX .1IO,ZJ;eii OHH MOI'YT 6hITh no O;:\HOMY' BMCCTC J1tt6o OTCYTCTBOBaTh.
B

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aci.:110mU1-tUHbt (au mu me.Ila) -

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W) Ha3hIBaIOTCJI OdliOUMeHlibtMU.

ArrJIIOTHHHH a CKJieIIBaeT apttTPOUMThI c arrJIIOTMHOreHOM A, a arrJIIOTMHMH l~ CKJICHBaeT 3pl1TPOUMThI c arrJIIOTMHOreHOM B. Ilo3TOMY B KpOBH Ka:>K,ZJ;Oro 'ICJIOBeKa naXO;:\JITCJI pa3HOHMCHHhIC arrJIIOTMHOreH l1 arrJIIOTHHMH.
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(AB) rpyrrna -- B apHTPOUMTax HMeIOTCfl arrJIIOTHHoreHhI AM B, arrJIIO-

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15 %

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469

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470

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(BepXHHH,

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BepxHHM

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125

MJIH pe11,errTOpOB

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cpe.n;H11e 11 H11:>KHl1e - - ,ll;bIXaTeJibHYIO IIOJIOCTb.
CJI11311cTaH o6oJioqKa Hoca BhICTJ"IaHa
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ceKpeT CJ111311CTblX KJieTOK, 06BOJiaK11Ba10ru;11x •rncT11qKII IIbIJI11 11 yBJiammi:roru;nx

PHc. 172. CxeMa CTpoeHvisi Ab1xarnnbHb1x
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B03,n;yx, 60JibIIIOe q11CJIO KpoBeHOCHbIX

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11 oq11ru;eH11e Bo3.n;yxa • .IJ:b1xaH11e rrpo11c-

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B ropTaHII

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cpopMIIPOBaHIIII 'lJieHopa3.11,eJ1bHOH pc<rII yqacTBYIOT HOCOBaJI noo1ocTh, ry6r,i,

.H3hIK, MBTKOe He6o, Ml1MII'leCKHe :vlbIIlIUbI.

3. llonPpP'IHo-no.1tocambu'

.wbtWU,bt, 06ec11clJHBaIOw,uc ,ilBl1)I\eHne xpxuieii:

H l'OJIOCOBbIX CBJI30K.

4. C;rn.1ucmaJt 060J10'llW

(MepuaTeJibHhri1 sn:nTc.rrnii:), BhICT1u1a101uaJI rop-

TaHh 113HYTPH' 6oraTaH me.rre3aMH II JIHMQlOH.11,HbIMH y3eJIK3Ml1.

TpaXPJl -

Tpy6Irn

(10----15

CM), COCTOJII.l(UJI II3

16-20

xp.HI.l(eBh!X IIOJIYIW-

.1IeU, He33MKHYThlX, TaK KaK C3al(H pacnoJio:men TIHI.l(eBO,il. IlpH npOX0)1{.ll,CHl111
nmu:enoro KOMKa IIIII.l(eB0.11, MO:>KCT paCllIHp.HTbC.H l1 BHe,11,p.HTbCJI B upocBeT TpaxeII. CBepxy Tpaxe.a rrpttKpen.'l.HeTC.H K nepcTHeBn.ll,HOMY xpm11y 3a c 1 ICT CBJI3Kn Me:m.ll,y

VI-VII

meii:HhIM TI03BOHKOM. CHH3y, Ha ypom1e

IV -V

rpy,nnoro

no3BOHKa, Tpaxe.H rrepcxo,nttT B 6poHxn. CJm311CTa.H o6oJIO'lKa co.uep:m11T KJieTKH JlHMcpOII.ll,HOH TK3HH l1 COCTOl1T 113 MepuaTeJibHOro 3IlHTeJIHJI.
Bhipa6aTbIBaIOT CJIH3h, 3a,nep:m1rnaIOI.l(YIO rrocTopoHHHe <raCTHUhI.

B

Ee

KJieTH:tt

Tpaxee BhI -

,neJI.HeTC.H ,nsa OT.ll,eJia: IIIeHHbIH II rpy,nHOH.

B

rpy.z:i;tto:i1 rroJiocTH Tpaxe.H pacrroJiaraeTcJI B cpe,nocTettn11, r;i;c crrepeJ(II

rrpttJieraeT BHJI0 1 1K0Ba.H :meJie3a, aopTa n JieBa.H rrJie'leroJIOBHaH ncna.

Bpo1txu.

Tpaxe.H ,neJIHTC.H Ha ,nBa rJiaBHhIX 6pottxa -

JleBhIH 11 npaBhrii.

IlpaBhIH 6ponx OTXOJ(HT rro.z:i; MeHhllIHM yrJIOM, lJeM JieBbIH, IIO:=JTOMY HHOpo,nHhie TeJ13 1Iall\C IIOIIaJ(aIOT B npaBLIH 6ponx.
BpoHXH BXO,l\.HT B BOpOTa JlerKoro l1 pa3,ll,CJI.HIOTC.H no 'lIICJlY OCl!OBHhlX .ll,0Jieii: JierKoro Ha Tptt l\OJieBLIX 6ponxa: 0.11,tto -

n npaBOM JierKOM u ,11,sa -

B Jie-

BOM. KpynHhIC 6pottxn no cTpoen1110 HanoMttHaIOT TpaxeIO. IIo Mcpe yMeHhllICHH.H .ll,IIaMeTpOB 6pOHXOB BMeCTO xp.HI.l(eBbIX IlOJIYKOJieu; B HX CTCHKe IlO.HBJl.HIOTC.H

II

xpml\eBbie I1JI3CTl1HKH,

KOTOpbie

nopJI,l\Ka nepexo.z:i;.HT B 6pottx11

III

HC'le3aIOT

B

6pOHXHOJiaX.

BpOHXII

nop.H,l\Ka (cerMeHTapHbIC 6p0Hx11), .r\aJiec

rrepCXOJ(.HT B KOHI(eBblC 6pOHXHOJibl, o6pa3y.H 6poux11aJibHOC ,11,epeno.

Jieucue -

napnhitt oprau B cpopMe KOHyca, pacnoJiaraIOI.l(HHCJI B rpy,nttoii

KJieTKe. IIpaBoe JterKoc 1rnpo11e tt TOHhllle, <reM JieBoe. JierK11e J~eJI.HTC.H I.l(eJIHMH Ha ,l\OJIH: rrpaBoe -

Ha Tpll .ll,OJIII, JieBoe -

6poHxoJiero<rnL1e cerMeHThI (npaBoe -

ua .ll,Be. ,[(OJIH ,neJIJITC.H Ha

10 cerMeHTOB,

JieBoe --

9 cerMeHTOB).

CerMeHThI cocTOHT II3 rrepBl111HhIX ,l\OJieK. ,ll,OJihKII JierKIIX OT,neJieHhI ,npyr OT
,npyra rrpocJ1oi'maMII coe.11,11H11TeJILHoii: TKaHII. Ha BHyTpeHHei1 110BepxttocTH
HaXO.ll,HTC.H BOPOTa JierKoro, 11epe3 KOTOpbre rrpoXO.ll,HT 6pOHXII, KpOBeHOCHhie
COCY.l\hl,

JIIIMcpaTIIIIeCKIIe

COCY.l\hl,

HepBbI,

COCTaBJIH.H

BMCCTe

KopeHh

JierKoro.
TKaHL JierI<HX o6pa3oBaHa BeTB.HI.l(IIMIICH 6pottxaMH H aJihBCOJiaMH. CTeHKH MCJib'IallllIIIX 6pOHXHOJI 11 aJibBeOJIHpHbIX XO.ll,OB o6pa3yIOT IIY3hlpbKII (J\HaMeTpOM

0,2-0,35

MM) -

aJibBeOJihI, KOTOpbie O'leHh IlJIOTHO npirneraIOT .z:i;pyr

K ,11,pyry II rycTo orrJieTeHhI ceThIO KpoBeHOCHhIX KarrIIJIJI.HpoB. Kam,nhlli KarrHJIJI.Hp conpIIKacaeTCJI c O.ll,HOH IIJIH HeCKOJihKIIMH aJihBeOJiaMII, tITO o6ecrre'lHBaeT orrTIIMaJihHhie ycJIOBII.H ,nJI.H ra3006MeHa. BHyTpeHHHJI rroBepxttoCTh cTeHKH

Anamo.MuK u <Pu3uo;ioiuK 'tell06el(,a

472

aJibBeOJihl BhICTJiaHa IIJIOCKHM O,D;HOCJIOHHbIM 3IlHTe.111eM H npoHH3aHa 3JiaCTHqecKHMH BOJIOKHaMH. KJieTKH an11TeJIHH BhI,n;eJI.HIOT B noJIOCTb a;IbBeoJI cypcpaKTaHT (cpoccpo.111n11,n;HOH npHpO,D;bl), npen.HTCTBYIOlil;HH CJifJTiaHHIO CTeHOK
11 06Jia,n;arou~11:H 6aKTepHI1H,D;HhIM .n;e:HcTBHeM. B BbICTHJIKe aJihBeOJI npttcyTCTBYIOT aJihBeo.1.apHbie MaKpocparn, KOTOpbre cparOI1HTHPYIOT HHOpO.D;Hbie qacTHqKH H

oTpa6oTaHHhIH cyp{paKTaHT.

B

o.n;HoM JierKoM Haxo,n;11Tc.H

350 MJIH aJibBeOJI, o6w;a.a IlOBepXHOCTb o6ottx JierKHX COCTaBJI.HeT OT 40
Bhr,n;oxe ,n;o 120 M' nptt B.n;oxe.

300M

2

npH

Mopcpo.1orw1ecKoii 11 cpyHKI1HOHaJihHOH e,n;HHHI1eil: JierKoro HBJI.HeTc.H a1IHHYC -

CHCTeMa pa3BeTB.'IeHmI O,D;HOH KOHI1eBOH 6pOHXHOJibI. AI1HHYC Haq1rna-

eTC.H 6pOHXHOJIOH, nepexo.n;.aw;e:H B pa3BeTBJieHH.H aJibBeOJI.HpHbIX XO,D;OB, KOTOpbie 3aKaHqHBaIOTC.H a.1bBeOJI.HpHbIMH MeruoqKaMH.

12-18

aIIHHYCOB o6pa3y-

IOT JieroqHyIO ,D;OJibKY.
Coe,n;HHHTeJibHOTKaHHa.a o6oJioqKa JierKoro
CTKOB. IlepBbIH, BHyTpeHHHH

(6ucu,epaJlbHbtU),

(nJteBpa)

cocTOHT H3 ,n;Byx JIH-

TIJIOTHO cpacTaeTC.H c JieroqHoil:

TKaHhIO, np11KphrnaeT JierKoe H 3axo.n;ttT B 6opo3.n;hr. Hapy;KHhIH

HblU, npucmeHO'lHbLU)

(napuemaJtb·

cpacTaeTC.H co CTeHKaMH rpy.n;HOH ITOJIOCTH H ,n;eJIHTC.H Ha

pe6epHyro (rroKpbrnaeT pe6pa), .n;ttacpparMaJihHYIO (rroKphrnaeT .n;ttacpparMy),
cpe,n;ocTeHHYIO Me,n;11acnrnaJihHYIO (rroKphrnaeT cpe,n;ocTeHtte ).

CpeaocmeHue -

no KOMrr.1eKc opraHOB, pacrroJiomeHHhrx Me;K,n;y rrpaBbIM

H JieBbIM JIHCTKaMH Me.n;ttaCTHHaJibHOH ITJieBpbl (cep.n;I1e, rrttru;eBo.n;, ,n;yra aopTbI, BHJIOqKOBa.H )KeJie3a, HepBbI HT. ,n;.).
Me;K,n;y JIHCTKaMH nJieBpbI HMeeTCH rroJioCTh, co,n;ep;Karu;a.a cepoaHyro ;KH,z:t;KOCTh, KOTOpa.H yMeHhlliaeT TpeHHe Me;K,z:t;y JIHCTKaMH ITJieBpbI rrpH ,ll;bIXaHHH,
coa.n;aeT OTPHIIaTe.'IbHOe .n;aBJieHHe, I103TOMY JierKHe ITOCTO.HHHO paCTHHYTbI,
TaK KaK .n;aBJieHHe B Me;KITJieBpaJihHOH ITOJIOCTH Bcer,n;a qyTb HH;Ke aTMOccpepHoro (aTMOccpepttoe .n;aBJieHtte rrptt aToM rrpHHHMaIOT 3a HyJieBoe). JieroqHaH
TKaHh o6Jia.n;aeT 60JibllIOH 3JiaCTHqHOCTbIO, rrpOTHBO,z:t;eHCTBY..H paCT..H;KeHHIO, H
yMeHblliaCT HeJIHqHHY .n;aBJieHH.H B03,z:t;yxa

(756-751

(760

MM pT. CT.) Ha

4-9

MM pT. CT.

MM pT. CT.). qeM CHJibHee paCT.HrHBaIOTC..H JierKHe, TeM Bhlille CTaHO-

BHTC..H HX 3J1aCTMqecKa.H T..Hra H TeM MeHbllie MeH.HeTC.H .n;aBJieHHe B ITJieBpaJibHOH rroJIOCTH. Bo BpeM.a B.n;oxa OHO paBHO

4

9

MM pT. CT., BO speM.H Bhr.n;oxa -

MM pT. CT.

MexanH3M s;:i;oxa H BhI;:i;oxa
B JierKHX HeT MhIIIIeqHOH TKaHH, I103TOMY OHM He MOryT CaMOCTO.HTeJihHO COKparu;aTbC..H. 06MeH raaaMM Mem.n;y aTMOCcpepHbIM H aJibBeOJI.HpHbIM B03,z:t;yXOM rrpoucxo,l:t;HT 6Jiaro.n;ap.a H3MeHeHMIO o6'heMa rpy,n;Hoil: rroJIOCTH BO BpeM.H B,z:t;oxa H Bhr.n;oxa.

AK-

THBHa.H pOJib B 3TMX IIPOIIeccax rrpHHa,ZJ;JiemHT CKeJieTHOH MYCKyJiaTYPe.

AKm 68oxa

o6ecrreqttBaeTC..H o,n;HOBpeMeHHbIM coKparu;etttteM Hapy;KHbIX

Me;Kpe6epHbIX MbIIIIII, rrpHITO,ZJ;HHMaIOru;HX pe6pa H CJierKa OTBO,l:t;.Hill;HX HX
B CTOpoHy' a TaK;Ke COKparu;eHHeM ,n;:irncpparMbl (yrrJioru;aeTC..H, orrycKaeTCH),
qTo rrpttBO,ZJ;HT K pacruttpeHHIO rpy,n;Hoil: KJieTKH. BJ!aro,n;ap..H aJiacn1qHoCTH
JierKHX, 3aMKHyTOCTH Me)KITJiespaJibHOH IlOJIOCTH H HaJIHqMIO B Heii OTpttn;a-

Mexauu3M 6iJoxa u 6MiJoxa

473

TeJibHOro ,D,aBJieHIUI J1erKIIe c.1e.n.yIOT 3a ,ll,BIIJKYIIIIIMIIC.H CTeHKllMII rpy,ll,HOM
K.1eTIUI II rraCCIIBHO paCT.HrHBaIOTC.H. ,UaBJieHIIe B03.n,yxa B RJihBeo.1ax JierKIIX BCJie,ll,CTBIIe HX paCT.HJKeHI1.H CT8HOBHTC.H HHJKe aTMOCcpepttoro, LfTO IIpIIBO,ll,IIT K IIOCTYIIJ1eHIIIO B03.n,yxa H3 OKpymaIOrne:iI cpe,iJ,bl B .'lerim:e -- IIPOIICXO.D.IIT B,ll,OX.

Cm~,n:yeT OTMeTHTb, 'ITO paCIIIHpeHIIe rpy.n;HOH KJieTKII KaK IIpII CIIOHT8HHOM, TaK II npII ycirneHHOM ,ll;hIX8HIIII Tpe6yeT yqaCTH.H ,ll,hIXaTC.'lhHbIX MhilllIJ.
8To 3Ha'IHT, 'ITO B.D.OX -

AKm Bb1ooxa

Bcer.n;a rrpo1Iecc aKTHBHhitt.

tta'IHHaeTC.H c

pacc.1a6JieHII.H

ttapya<HhIX

Mempe6epnhrx

MblllIII H .D.IIacpparMhI. BcJie,n;CTBIIe <!Toro IIO.D. ,n;eHCTBIIeM SJiaCTII'leCKHX CMJI
JierKIIX, ,D,aBJieHH.H BHyTpeHHHX opraHOB, ClIJibI T.HJKeCTH rpy,n:Hoii KJieTKII 06'beM ee yMeHhIIIaeTC.H, ,D,aBJieHHe B03,n;yxa B Jiel'KHX CTaHOBIITC.H BhIIIIe aTMOCcpepHOro Il B03.D.YX BhIXO,ll,HT B OKpymaIOJ:HYIO cpe.n:y. BbI.D.OX BO BpeM.H CIIOKOMHOro
,ll;hIXaHII.H rrpOMCXO,ll,HT rrpM paccJia6JieHHH ,ll,bIXaTeJihHbIX MblllIIJ, T. e. rraCCHBHO. IlpHIIO.D.H.HTa.H II pacrnHpeHHa.H rpy,n:Ha.H KJieTKa B CHJIY CBOeH T.HJKeCTII
orrycKaeTc.H.
IlpH ycHJieHHOM ,ll;hIXaHIIH, Kor.n.a ttymno yBeJIHqMTh o6·beM Bhr,n:oxa, Oll;Horo rraCCHBHOro MexaHII3Ma He,n;ocTaTO'IHO. ,UorroJIHIITeJibHO ,ll,OJIJKHhI .n,eMCTBOBaTb BHyTpeHmre Mempe6epHbie KOChie MbIIIII~bI If MbIIII1Ihl 6pIOIIIHOH CTeHKII.
IlpII KamJie, qHxaHIIH 38,ll,eHCTBOBaHhI MblllIIIhl 6pIOIIIHOrO rrpecca: ,ll,IIacpparMa
pe3KO IIO,ll,HMMaeTC.H, rpy.n;Ha.H KJieTKa orrycKaeTC.H.
l1TaK, Bbl,ll;OX MOJKeT 6bITb rraCCHBHhIM (6e3 yqaCTH.H .D.hIXaTeJibHbIX MhillIII)
HJIH aKTHBHbIM, B 38BHCHMOCTH OT ypOBH.H MeTa6oJIM3Ma B opraHII3Me. ECJ1II
IIOCJie BhI,ll,OXa ,ll,bIXaTeJibHbie MhIIII1IhI OCTaIOTC.H paccJia6JieHHhIMII, TO HaCTyrraeT ,ll;hIXaTeJihHa.H rray3a. ,UaBJieHIIe B03,D,yxa B JierKIIx B rrpoMeJKyTKax Mem-

.D.Y

Bbl,ll;OXOM H B,ll;OXOM paBHO aTMOCcpepHOMy, TaK KaK HapyiKHbill B03,n;yx

II B03.n;yx B JierKHx cBo6o.n.tto coo6rnaIOTC.H Mem.n.y co6on.

BeHmuJtJW,UJl ;ieiicux -

:no ra3oo6MeH Mem.n;y aTMoccpepHhIM M aJihBeo-

JI.HpHhIM B03.D.YXOM. IloKa3a TeJI.HMH BeHTHJI.H1IHH JierKIIX .HBJI.HIOTC.H MHHYTHbIH o6'beM ,ll,bIXaHM.H (MO,U), KOTOphIH orrpe.n;eJI.HeTC.H KaK rrpOM3Be,l(eHHe
.D.bIXaTeJihHOro 06'beMa Ha 'IHCJIO ,l(hIXaHMH B MMHyTy. B Te'JeHIIe 0,l(HOH MHHYTbI qeJIOBeK B,ll;bIXaeT H BhI,ll;bIXaeT

5-8 JI

B03,n;yxa. Ilpn HHTeHCHBHOll cpH-

3M'JeCKOM ttarpy3Ke MO.U MOJKeT .n;ocTnraTh
TeJihHhIX ,ll;BMJKeHMH y

20-80

JI/MHH. qacToTa ,l(brxa-

B3POCJIOI'O qeJioBeKa B IlOKOe COCTaBJI.HeT

12-16

B MHHyTy.
l1HTeHCHBHOCTh BeHTMJIRIIHH JierKMX xapaKTepH3YIOT:

1. I'Jty6uHa U 'laCmoma dbtXaHUJl,

KOTOphie 3aBHCRT OT HeifporyMopaJihHOH

peryJIRIJHH, rrap1IM8JihHOI'O .n;aBJieHHR KHCJiopo.n.a B aTMOccpepe; B03pacTa
H IIOJia (y HOBOpOJK,ll,eHHhlX
y rro.n;poCTKa

18-20,

40-50 ,l(hIXaTeJihHhIX ,l(BHiKeHHH B MHHYTY,
15-18); cpll3H'leCKOH narpy3KH (rrpH

y B3pOCJIOro

yBeJinqeHMH MhIIIIe'JHOM pa60TbI rJiy6HHa H 'laCTOTa ,l(hIXaHHR yBeJIM'IHBaIOTCR, BO CHe -

yMeHhIIIaIOTCR) I1 cpH3HL!eCKOro COCTORHIIR qeJIOBeKa

(y TpeHHpOBaHHhIX JIIO,n;e:H

JKEJI

6oJree pe,n;KHM H r.11y60KHM).

yBeJIH'IHBaeTCR; ,l(hIXaHIIe CTaHOBHTCR

A namOMUR.

474

u tjJu.JuO.'WZUR. '!e11c111e1w

/KHaHeHHatt eMKOCTh .11ermtx

Boa pa CT

JKE.!l

~--------------------

.UeTlI

1,2 JI
---------------

5-6 JieT

~------------

·1,0-5,()

Mym<JinlbI

--

2,5 3,5 :1

iH:cmqHHbI

r-----------------

JI

--------------------

1-----

5,5--7

CrropTC:vteHbl

~:-I

2. Coom1wweuue ObtxameJtb1t0zo o6oe.Ma u 061,eJlW BeHmu;wpyeJ1wzo npocmpaucnwa.

3. JKu.JllC/l/WJl e.MKOCmb Jle?KUX ( JKEJJ ),

M3Mepem1e KOTOpOH OCYIUCCTB-

Jrn:eTCH MeTO,ll,OM CIIMpoMeTpMM c IIOMOIIJ;bIO crreu;MaJibHOro IlpM6opa crrnpoMeTpa.

Jit°u.JltellHUJl e.MKOCmb Jte?KUX ( JKEJI) -

3TO HaM60JibllleC

KOJIM'-ICCTBO B03,ll,yxa, KOTopoe qe.'lOBeK MO/KeT Bbl,ll,OXHYTh rroc.'le caMoro
rJiy6oKoro B)l,OXa. 3TOT IIOKa3aTCJih xapaKTCpM;~yeT COCTOHHMe arrnapaTa
BHCIIIHero ,ll,hIXaHHH qe.'lOBeI<a (JICl'KHX M rpy,IJ,HOH KJieTKH c ,IJ,bIXaTC.'lbHbIMM MbilllUaMM).

JKEJI

CK.'la,ll,bIBaeTCH M3 Tpex JICJ'OqHbIX o6'heMOB:

JKuJ1te/lHll5l eMKOCmb = )~btxame.1bHblU ome.M
3ep61tbtit o6oe.w 6Uoxa
,UbJXaTeJibllhlH 06'beM

(1500 CM")

-

+ Pe.wpBllblU o6oe.M BblOOXa T

Pe-

3TO 06'bCM B,n;oxa IIOCJIC CllOKOHHOl'O Bbl-

,n;oxa.
Pe3epBHblH o6'heM Bbl)l,OXa

(1500

CM") -

o6'be:'vl B03,ll,yxa, KOTOpbltt MO/KHO

,ll,OIIO.'lHHTe.'lhHO BLI,n;OXHYTh rroc.1e CIIOKOHHOrO BhI,n;oxa.
Pe3epBHblH 061,eM B,ll,OXa

(1500

CM") -

o6'beM B03,ll,yXa, KOTOpblH :'vlOiKHO ,ll,0-

IIOJIHHTCJibHO BJ~OXHYTb IIOCJie CIIOKOMHOro B,ll,OXa.
pacTa, COCTOHHM.H TpeHMpOBaHHOCTM M pocTa.

y

JKEJI

3aBMCHT OT IIOJia, B03-

cnopTCMeHOB

JKEJI

YBC.'lH'IM-

BaCTC.H 11 C03,ll,aIOTCH YCJIOBII.H ,n;JI.H 6oJiee r Jiy6oKoro ,ll,bIXaHHH' 'ITO CB.H3aHO
c pa3BHTMCM ,ll,bIXaTeJibHOM Mycxy JiaTyphI.

06ut,a5l e.M!WCmb Jle2KUX = JKEJI
OcTaToqHLrii ofrheM

+ Ocmamo•lllblU o6oe.M

(1000-1500

CM") -

eoaoyxa

3TO o6'heM B03,ll,yxa, KOTOphrtt oc-

TaeTca IIOCJie HHTCHCIIBHOI'O Bbl,ll,OXa. Y,11;aJJl1Tb ero npaKTHTJeCKH HCE03MO}IOIO,
II03TOMY J'lCl'O'IHaH TKaHh B BO)l,e He TOHCT. Bo3HMKaeT IIOCJie rrepBoro B,ll,OXa HOBopmK,ll,CHHOI'O.

Mepmeoe npocmpaHcmBo -

3To o6'heM ,ll,hIXaTeJihHhIX

rryTeil:

(TpaxeII,

6pOHXOB, 6pOHXMOJ'1), B KOTOpOM He rrpOMCXO,ll,IlT rrpouecc ra3o06MeHa Mem,n;y
B03,ll,yxoM 11 I{pOBhIO. CocTaBJIHeT

140-150

cM", MJIM I

f3 ,ll,hIXaTeJibHoro o6'be-

Ma rrptt crroKOHHOM ,n;h1XaHMH.

f

a3006MeH B JierKHX H TKaHHX

fa3006MeH ocyIUecTBJI.HeTc.a no MexaHM3MY ,11;11¢cpy3HM. EcJIM Ha,n; mtt.n;xocThIO Haxo,n;MTCH CMeCb ra30B, TO Ka}l{,IJ;blll ra3 paCTBOp.HeTCH B }l{M,ll;KOCTM COOTBeT-

I'a3006Men

6

.riezi,:ux u m1rnn1lx

475

CTBeHHO eI'O napu;irnJihHOMY ,ll;3BjleHl1!0. Ilapu,IIaJih!IOC ,z:i;aBJJeHHe -

dTO TO /IaB.1e-

HIIe ra3a, KOTOpoe rrpl1XO,ll;IITCH Ha ero ,ll;OJIIO OT o6ui;ero ;i;aBJieHIU! CMec11 I'R30B.
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meny.n;o'IHO-Kttwequoro TpaKTa.
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0T,L(eJieHMe CJIIOHhI, B03HHKIUee IIPII pa3,npameHMII MexaHIIqecKIIX, XIIMIIqecKIIX II TeMrrepaTypHhIX peu;errTOpOB pOTOBOM IIOJIOCTII, HOCIIT Ha3BaHHe 6e3YCJIOBHOI'O CJIIOI-IOOT,L(eJIIITeJibHOI'O peQ:>JieKca.
HapH,ny c 6e3ycJIOBHhIM cJJIOHOOT,neJIIITeJihHhIM peQ:>JieKcoM cymecTsyIOT
YCJIOBHhie CJIIOHOOT,L(eJIIITeJibHble peQ:>JieKChI B OTBeT Ha 3pMTCJihHhie, o6oHHTCJihHbie, CJIYXOBhie H ,npyrIIe pa3,npaJKHTeJIII (BH,L(, 3arrax rmmH II T. ,[(.).

r JlOmaHUe -

pecpJieKTOPHhIM aKT. IIepemeBaHHaH, CMoqeHHaH CJIIOHOM IIH-

ma rrpeBpamaeTCH B rr:11meso:i1: KOMOK, KOTOphlll ,L(BIIJKeHHHMH H3hIKa, ry6

480

AnamoMUR u <Pu3UOIW?UR 'le1w6e1rn

H rneK JIOITaJJ;aeT Ha Kope!lb H3hIKa. KopeHb H3bIKa H MHI'KOe He6o HMelOT 60:1bmoe KOJIH'IeCTBO Y.YBCTBttTeJihHhIX nepBHhIX OKOH'rnHnii. Pa3JJ;pamenue, BbIJhrnae:vrne nnwei1, rrepeJJ;aeTCH B 1~enTpb1 r"10TaHHH npo,L1,oJ1roBaToro MOJra. OTCIOJJ;a nepBHhie HMIIYJlbCbI no JJ;BHraTe.JihHbIM (scpcpepeHTHbIM) HepBHhlM BO.lOKHaM B COCTaBC H3hIKOrJIOTO'IHOI'O H 6JiyJKJJ;a10wero Hep BOB lI.lJ.YT K MhIIIJI1aM
r.10TKH 11 ropTauu, BhI3hIBaH aKT rJIOTaHHH. B

STOT MOMeHT JaJJ;epmttBaeTCH

JJ;hIXaUHe, BXOJJ; B HOCOBYIO IIOJIOCTb 3aKpbIBaeTCH MHI'KHM ue6oM, Ha.n;ropT31lJIHl{ JaKphrnaeT axo.n; B ropTaHh. EcJitt Y.eJioBeit BO BpeMH e,nhI pa3roBap1rnaeT,
TO BXO,ll; 113 I'J10TKII B ropTaHb He 3aKphrnaeTCH, H IIIIrna MO)KeT rrorraCTh B rrpoCBeT ropTaHH, B ,ll;bIXaTe.1hHbie nyTII.

I-fa

POTOBOH IlOJIOCTH IIHrneBOH KO MOK norra.n;aeT B HatiaJibHYIO 'IaCTh r JIOTKH

II CHJihHhIM COKparneHHeM ee Mhlllll1 rrpoTaJIKHBaeTCH B IIIIrneBo.n;. BOJrnoo6pa3HOe c0Kpan~ern1e MhIWI1 rr11weBo.n;a rrepeMewaeT rrnwy B 'KeJiy.n;oK. Bech rryTh
OT pOTOBOH rro.10CTH
3a

2-3

.n;o

meJiyJJ;Ka TBep.n;aH IIIIII13 rrpoxo.n;IIT 3a

6-8,

)Klf,D;KaH -

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I'.!lOml(a

paCIIOJlaraeTCH II033,D;II IIOJIOCTII pTa H IlOJIOCTH HOCa, npe.n;CTaBJIH-

eT co6oii nopmrnoo6pa3uh1ii KaHaJI, cnJIIOrneHHhIH B nepe.n;ueJa.n;HeM HarrpaBJieHHII, JJ;JIHHOH

12-14

CM, B KOTOpOM rrepeKpernIIBaIOTCH JJ;hIXaTeJihHhIH II fIH-

rneBapIITeJ'IhHblll nyTtt. Ha rpaHIII1e Mem.n;y

VI

II

VII

IllettHhIM no3BOHKOM ona

nepexo.n;ttT B rrIIrneBo.n;.
B rJIOTKe Bh1.n;eJIHIOT TPH OTJJ;eJia:

nocoBou (Hocoz;toml(a) - c10.n;a OTKphrnapomOBOU (pomoz.nom1ca) - coo6zopmaHHbLU - coo6rnaIO~IIHCH c BXO,lJ,O!\i

IOTCH xoaHhI II OTBepCTHH eBCTaXIIeBhIX Tpy6;
rnaeTCH c fIOJIOCTbIO pTa qepe3 3eB;

B rop'l'aHh II nepexo.n;HrnIIii B rrmqeBo.n;.
IloJIOCTh I'JIOTKII BbICTJlaHa 3IlIITeJIIIeM: B o6JiaCTII HOCOI'JIOTKII TeJibHhIM, OCTaJibHaH qaCTb -

CTOH o6oJIO'!Ke I'JIOTKII pacno.11araeTCH meCTh MHH,lJ,8.JIIIH HIIH J'IIIMcpOH,ll;HOH TKaHII, IlnporoBa -

Mepl13-

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Ba.rih.n;eiiepa (napHhie He6nhie II Tpy6nb1e MIIn.n;a.11nHhI, OJJ;Ha rJIO-

To•rnaH MIIH.Lt;aJIIIHa). 3TO opraHbI IIMMYHHOH CHCTeMhI.
B CTenKax r.TIOTKH nMeeTCH xoporno Bhipa'KeHnaH MhIIIle'IHaH o6oJIO'IKa, coCTOHIIl3H II3 nonepetIHO-lIOJ10CaTbIX MbIIIIetIHhIX BOJ10KOH. IlpII ee COKparneHHH
IlOCTYIIIIBIIlIIH II3 IlOJIOCTH pTa ITIIrneBOH KOMOK 6hICTpo npoTaJIKIIBaeTCH B fIIIweno.n; II .n;aJiee B 'KeJiy.n;oK.

ITuw,eeoi} -

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CH Ha rpaHHI1e Mem.n;y

VI, VII

25-30

CM, KOTOpaH naqnnaeT-

IllettHhIM rroaBoHROM n aaKaHtIIIBaeTCH Bna-

JJ;enIIeM B meJiy,noK. IlIIrneBo.n; IIMeeT TPII cymeHIIH: y cBoero HatiaJia, Mem.n;y

IV

H

V

rpy.n;HhIM no3BOHKOM II Ha ypoBHe nnweBoro OTBepcTHH .n;IIa-

cpparMhI.
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rJia.n;KOMhIIIle'IHhlMH

481
K:reTI<a:vrH. llnpyiKHHH 000JHJ11Ka Illlll\eBO,TJ;a

--- a,UBCHTIH\HH ·-- COCT011T 113

pLlX.:10.ii HO.TIOKHHC:Toii coe.u111111TeJJbHOH TK3HH.
<PyHKUH.H mrn\eBO,':(a -

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pa3Hbl:vil1 CO!tparueHHHMH CTeHOK nHw;eno.ua B iKCJIY.UOK.

}ff P.'L,1Jr70K -

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OT

1,5 .uo 2,5

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35

MJIH), Bbl-

)J,eJ1H:!OI1VIC IBeJJy.UO'!Hb!H COK.
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B

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482

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B Tet.1e11IIe

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1,5--2,5

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me.11y;:i;ot.1HbIH COK yme qepe3 IIHTb -

;:i;eCHTb MIIHYT rroc.11e Hat.JaJia e.ll,hl. CeKpe-

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lfa

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14-16

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12

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K

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8

176).

0lJifaKO OCHOBHaJI pOJih B BbI,[(€JIHTCJ!hHbIX

npoueccax

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75 %

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Ma npo.uyKTOB o6MeHa B€Il.\€CTB.
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KIIC.'IOTHO-r.u;eJioqnoro paBHOBCCII.H;

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Pitc. 176.

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TenbHOi1 C~CT8Mbl: 1 -- Hat(n0484H~K; 2 -

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3 - aoprn; 4 - H~)l(HSlsi nonasi seHa; 5

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11 lIPYI'IIX

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8-

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Bbllll€HH!O KpOB.HHOI'O l(aBJICHH.H, CTII-

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noxaHKa; 12 -

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KpoBeTBopettHJI;

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488

AHanu1Mu11 u <f>u.1110.101un •1e:1C11wKa

11011;1.cpmaHHe ElIC.lOTHO-II\e.:w'f!!Oro
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no;ui:ep»WHHe

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II O'l l(ll 120-200 r,

napHhII1
1WHHOIJ

opnrn

10-12

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C'.\1,

6050-

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no 6oKaM

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XII

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P111c. 177.
pa3pe3):
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9-

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6 -

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Betta,

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2060(' 6l'UJ,l'C/7160

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1500-1800
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1 .1
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a TaKme rrpo,llyKThI o6MeHa 11 ,D,pyrtte

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(0,03 ';!,, ),

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(0,004 % )

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490

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11 ryMopa.JihHhIMH MeX3HH3M3M:H.

Cu.Mnamul/ecKoe

B.JIMRHHe 11epBHoii cncTeMLI rrpHBOL\HT K cy*eHHIO noqe'-I-

Hhrx cocy.l(oB n yc11JieHMJO pea6cop6I(1111.

II apacu.Mnamul/ecKoe

n.rr11Jittne, Ha-

rrpoTHB, YCllJIHBaeT KpOBOTOI{ B TIOlfKax 11 rrp11BOJ\11T K yMeHhllleHHIO pea6cop6ll,HH.

Cu.Mnamul/ecKaR Hep61taJl cue me.Ma

a rrapaCHMII3TH'IeCK3R -

PeHun -

cTnMy.rr11pyeT pea6cop6ll,HIO ttaTpMJI,

pea6cop6ll,11IO r.JIIOK03hl.

aHzuome1t.m1weaJ1. cucme.J1>w:

B K.rreTKax rro'IeK o6pa3yJOTCfl <P11-

3uo.rroruqecKue aKTHBHhre Bew;ecTBa, pery.rr11py10ill,He pa6oTy rroqeK u L\pyrux

A namo.HUR u <Jnt.1uo1wzu11 'H'ILnfieKa

492

opraHOB. Ilo.D, BJll15IHHeM peHHI!a 06pa0yeTC'5l ropMOH C\HI'llOTeH:lllH, KOTOpbiii
cyJKRCT cocy.D,bl, HOBbllIIC\}! ,IUlB.'1PHl1e Kj)OBl1 BO BllYTIJCHHHX opra1rax, a TRKffie
ycivnrnacT pca6cop6umo H8'rp11H 113 rrepnw111oii :vro•rn B 1-.:poBh.
fopMOH

:Jpumpono::mzuu

CTIIMyJ111pyeT rrp011ecc 06pa3onattHH .:Jp1rrp01u1TOB

B KpaCHOM KOCTHOM M03l'C.
fop:vioH 3ai1ueif JIOJIH r11rro¢113a -

zopMOH -

A/I.I') -

Ba.Jonpeccuf/ ( all!nwluypemu'lec1n1i1

ycn.111saeT o6paTHOe BCaCbIBaTrne B021bl B IIJBHTbIX IW-

naJibU:aX, y:vieHblllaf! 06'beM BTOplPIHOii: MOLIII, H, CJIC)!OBaTCJ!bHfJ, IIlJf'llilTCTBYeT ITOTepe BO,'J;hl. Ilpu Hapyweurm ,UCHTC.1hl!OCTH r11rrocjrnaa Bbl,i(P:IE:HHC
A,Uf MO:>KeT rrpeKparu;aThCJI, Bo.n;a rrepecTaeT pea6cop6upoBaThCB, 11 iruype3
B03pacTaCT (.n:o

20-25

Jl B cyTKH). 3To 3a60.1eBaHHe Ha3bIBaCTCR 11ecaxap11hIM

MOLICI13HypeHl1eM.
fopMOH ll\HT0Bl1,ll;HOH me.1e3hl

mupOICCUH

ycIImrnaeT .n;11ypeJ, OC.'1RUJUJ,f! pe-

a6cop6u;1110 BO,ll;hl B KaHaJihI(aX. fopMOH KOpbl Ha)lllO'IeLIHHKOB

Q.'lbiJocmepOll
aiJpe11a-

yseJIMLIHBaeT pea6cop6umo HaTpHH H y.n;anettne KaJIH.H H3 opranHJMa;

JlUfl B 60JihIIIHX .n;o3ax Bhl3bIBaeT cymeHHe rrpHHOCl'lll\HX cocy,1(0B-KJiy6o•rnoB

H TeM CUMb!M CHHmaeT .n:11ype3.

PaaMHomeHHe

Bee iKIIBhre opraHH3MbI o6na.n:aroT crroco6ttocTh!O 1\ caM0Bocnpon3se;ren1110
IIJIII pa3MHomeHIIIO. Cyru;ec'!'BYIOT rroJioBaH II 6ecrro.;ra.H l}:>0p:vi:b1 pa3:vi:1-1omettna.
BecrroJroe pa3Mttomettne xapaKTepIIayeTcH OTCYTCTBHeM rro.rrosoro rrpoIIecca
II ocymeCTBJIHeTCf! 6e3 yLiaCTIIH ITOJIOBhIX KJieTOK.
CBOHCTBeHHO BCeM )KIIBh!M opraHH3MaM, y
KJieTKII (raMeThI) -

flojIOBOe pa3MHO:>KeHIIe

KOTOpb!X o6paJyJOTCH no.TIOBhie

f!HUCKJICTKII (B :>KCHCKIIX no.'10BhIX :>KeJICaax) l1 cnepMRT0-

3011,ll;hI (B MYiKCKHX ITOJIOBh!X :>KeJie3ax).
HoBhiii opraHII3M paaBIIBaeTcH II3 31IrOTbI, 06pa3yroru;e11cR B pc3yJih'l'aTe orrno.n;oTBopeHIIH f!HUeKJieTKlI cnepMaT03011,ll;0'.\1:.

MymcKaH rroJioBaH CHCTeMa
BKJIIO•rneT B ce6H BHyTpetttttte H ttapymuh1e rroJIOBhie opratthI (pHc.

179).

BttyTpeHHIIe rroJIOBLie opraHhr:

1.

J1oJW6bte J/CeJle.3bt

(ce.JtteHHU/CU

UJlU .HU 1l1CU)

-

rrapHbIC,

ITOKpb!TbIC

IIJIOTHOH coe.n;IIHIITeJihHOTKaHHOH o6oJIOtIKOH, B KOTOpoii HaX0,11;.HTC.H 113BIIThie ceMeHHhre KaHaJihUhI. CTeHKH Ka11aJihIIeB BhICTJiaHhI cnepMaToreHHhIM an11Te.1neM, II3 KOTOporo o6pa3yIOTCf! cnepMaT030H,ll;hI, II KJICTKaMII CepTOJIH, KOToph:re crroco6cTBYIOT co3peBaHHIO crrepMaT030II.n;OB.
Mem.n;y ceMeHHhIMII KaHa.'1hU:aMII Haxo,llHTCH KJieTKII Jleii.n;IIxa, Bhrpa6aThrnaromIIe anirporeHhI, HBJIHIOll\IICCf! CTepon,r~aMH XOJieCTepIIHa.

mocmepoH

90 '%

npOII3BO,ll;HhIMII

att.n;poreHOB COCTaB.'1.HeT ropMOII TeCTOCTepoH.

Tec-

pery;rnpyeT 6eJIKOBhin o6MeH, CTIIMYJIHpyeT pocT II pa3BII-

Pa.JMllOJKenue

493

14
P111c. 179.

CxeMa c1 poeHvi>i My>KcK011 nonosoV! CV1CTeMb1:

cTarennHasi )!(811e::;a,
MG•!Cf3~·fi ny3blpb:

K3Hilff

10 --

3 -

7-

nCHV1C:

r1psiMa11 KV1WKa:

li00k0B351 KOCTb:

11 -

4 -

8 --

1-

ceMeHHof1 ny3b1peK;

aHanbHOe OTBepcn1e;

5-

C8MS1BblBOAS1LUV1fi npOTOK;

r1P1'11\3TOK C8M8HHV1Ka:

12

CeMeHHV1K;

2 -

npeA-

Kyneposa >1<ene3a;

9-

6-

MOY8V1CnycKaTenbHbl1'1

13 -- IOIIOBKa 11eHV1Ca; 14

MOWOHKd

THC

IIOJIOBOro arrnapaTa,

MY:ll<C101C BTOpl!Lllibie IIOJIOBbie IIPH311aKn.

Ilponecc o6pa3oBamrn crrepMaTo3011;~0B Ha3hIBaeTcH

cnep.1tamoze11e.w.w

(CM. pa3,LJ,CJI «06w;aH 6uo:IOrHH» ).

2. Ce,nJ16bt1t0cJ1w,ui1 npomo/C -

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IIO.JIOBh!e KJieTI(H CTaHOBHTCH crroco6Hh!MII K OIIJIO,ll;OTBUpeHHfO.

3. IlpeiJcmamPJlb/WJl .m:P.7P;JQ - Bhipa6aT1,maeT fKH,'l;KHII ceKpCT.
4. B.u.11b6oypempu.;1b11a.H (Kynepoeu) J1Ce.H'Jll - Bbr,n;eJI.HeT BH3KHii

ceKpeT,

rrpe.n;oxpaHHIOru;Hii C.'IH3HCTYJ:O 060.1o<my MO'leIIcnycKaTeJibHOrO 1rnnaJia
OT pa3,n;pamaJ:Ow;ero ,n;ei1CTBHH MO'IH.

5.

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Hapymuh1e Mymnrn:e no.'lOBhJe opratthi:

JloJW60U

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M OUJOHIW

-

(neuuc) -

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co,n;ep:>KIIT BHYTPH HH'IKH c HX np11,n;aTKa:vrII, 110,n;,n;epmrrna-

eT TeMIIepaTypy HIIme TeMnepaTypbI Te.1a.

i\1o'leucnyc1Came:tbllbti11Ca1-ta ..1 (ypempa) -

c:1y)KIIT J~.1.H n1>rne,n;eHHH MO-

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BKJIJOlfaeT B ce6H BHyTpeHHHe, Haxo;~Hll~HCCH B IIOJIOCTH Ma,1oro Ta3a, H HapymHhie 110.'IOBb!e opraHhl, pacnoJiaraJ:Ow;HeCH Ha IIOBepxHOCTH Te.1a (pHC.

180).

494

A.namUMUR u <J>U.IU0.102.UR '/, (' /lOH<'IW

BH y Tpe1-rnu e oprnHhl:

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B

KOpKOBOM Be111ecTB e p ac rro .' ro n <eHhI

n e pBH'IHhie cpo.1.11myJILI, OI<py/t\eHHhie OJ(HOc.10i'IHhl'.\t CjlO JL'Il1KY:InpHbIM 3 IlHTe.1HeM. 3.z:i;ecb o6p a 3yI-OTC.H II0.10Bh!C KJI8TKH .11 BLrpa6aThIBaI-OTC.H rrOJIOBhIC rop:vroHhr. Mo3rosoe seu1ecTBO o6pa3oB a Ho coe;1.11H.11Te"i.hHOii TKaHh!-0, B KOTOpoii upOXOLJ;.HT cocy,1,hl l1 HCPBhI. Ilpouecc pa3BHTHH

siiueK.::TeTOK

(OBoreHe 3 )

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rrepno.z:i;e. 8TOT npouccc cxo )~eH co cuepMaTorette3o:vr, on;ttaKo e c Th pa3.'IH'IHJI, CB.H3aHHhie co crreuna.'ln3au.11eii raMeT (cM. pa 3 .z:i;eJI « 06n~as
6nonorHJ1» ).

400-500

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MOMeHTY rroJioBoro c o3pesaHHJ1 B nn'IHHKe naxOj (HTC.H

nepBl14HLIX <}lOJrnHK Y JIOB, IWTOpbie rrpeo6pa3ylO T CH B 3pe-

Jibie c}lOJIJIHKYJihI (rpaacpoBbl rry3biphK11). PocT <}10.n JrnKyJIOB npoHcxo-

28
(cIJCf),

)IHT U:HK.lHl!eCKH KaJK)lble

)!Heii 110)1 BJIHHHJ1€M CjlO J1 JlHKy.i.oCTHMY-

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Bbipa6aTbIBaeMoro I'HIIOC}.>H 3 0M. ¢lO.lJ1H-

KYJIJ!PHhJe KJieTKH na'IHHaIOT BL1pa6aTLIBaTL JKH)II<OCTh cpo.rr;1.11KyJia,
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3cmpo ie Hbl.

5Ii111e1<,1eT-

Ka, OKpymeHHaJI CJIOCM <}lOJIJIHKYJI.HpHLIX !{J1CTOK, OTTeCH.HeTCH K 0)(HOMY H3 IIOJII-OCOB cpo.1JI.11Ky.'la, o6pa3y .H BTOpll'!Hblll qJO:I.::THKyJI, 3a'rCM

C xeMa CTpoeH~S1 )1(8HCKO~ nonoao~ c~CT8Mb1: I - 11i11..(esoA; 2- MarKa ; 3- MOYesoi'i
- no6KOsa11 KOCTb; 5 - no6oK: 6 - MOYe~cnycKarenbHbli'i KaHan : 7 - Kn~rop: 8 - Mana11
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P111c. 180.

ny3blpb ; 4

Pa.JM1toJ1Ceu ue

495

TpeTWIHb!H, KOTOphiii lIOC:Ie coapeBamrn. pa3pbIBaCTCH, OBO!U!T BLIXO.UHT B 6pHHIIHYIO IIOJJOCTh H nonanaeT B :vraTOl.J HYJO Tpy6y. Ilporrecc Bh!XOJJ:a OBOll;l:!Ta HOCHT naaBaH1:1e

06.IJ.'lRl(UJI. Otta npo11cxo.n:HT pery.151 ptto

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Bhrpa6aTLIBaeMoro nrrroqrn3o:vi. Ha MecTe JTOIIHYBIIIero cpo.irJlHKy:ia 113
J/CPJ!moe me.rw, KOTOJ)Oe c:1v-

ero cpoJI;IHKy.1HpHblX Ko'1E'TOK o6pa3yCTCH

JKHT Bpe:vreHHOH JKCJIC30ii: BHyTpe1rnei1 cexpeurrn. fopMOH JKPJIToro

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T~

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orr,10.n:oTBopeHHOH HMUCKJlCTI<H. EcJin OIIJIOD;OTBOpettne .aH:r1ex:1eTKH
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12-14

:IHer'I rop:vrottoo6pa30BaTeJihHa.a cpyHK-

I!HH JKCJIToro TCJ1a rrpeKparuaeTCH II Ha ero MeCTe OCTaCTC.H coe,n:HHHTCJibHOTKaHHbIH py6eu. Ilpn 3TOM CJ1H3HCTaH o60.10'1Ka MaTKH OTTOpraeTCH, nacTyrraeT KpOBOTe'ICHHC, KOTOpoe rrpHHHTO Ha3hIBaTb MeHCTpyaJihHbIM.

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rrepBOI'O

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45--50

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4. BJ1azaJ1u14e -

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Hapy:>KIILre mencKtte noJIOBhre opratthr:

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496

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3) rrpe.uoTBpau,vteT coeJJ,HHeHHe H-PHK c ptt60COMOH; 4) npeKpa1uaeT C'IHTblBaHMP H3CJ1 e,ll;CTBeHHOH HHcj>OpMaIJ;HH.

A 2. II Pn mua11a.R C6.R.Jb 6 .MO.TteKyJle nenmuaa o6pa.Jyemc.R .MeJ/fOy:
1) aMHHO- H Kap6oKCMJ1hHOH rpynnaMM; 2) Kap6oKCMJ1bHOH rpynrroH M pa3) pa.n;nKaJIOM n s-s-rpynnaMn; 4) s-s- H aMHHorpyrrna:vrn.
A 3. Henpa6uJ1b1toe cyJl<'iJeuue:
1) ryaHHH - rrypHHOBOe OCHOBaHMe; 2) IJ;HT03MH - IlHpHMH.UHHOBOe OCHOBaHHe; 3) KapOTHHOH,ll;hI - mnporro.n;o6Hhle Beu~eCTBa; 4) a.n;eHHH H ypal~HJI

.llHKaJ10M;

IlllpMMH,ll;l1HOBhie OCHOBaHMH.

A 4. r Jl/OKOZO/l OmliOC.Rm K:
1)

.n;ttcaxaptt.n;aM;

2)

IIOJIHIIeIITH,ZJ;aM;

3) IIOJmcaxaptt.n;aM; 4) CJIOIBHhIM 6eJJ-

KaM.

A 5. B cocma6 1-ty1uieomuiJa exoiJ.nm:
1)
Ta; 3)

MeH03MH n cj>occj>aTHaa rpynna;

2)

cpocq>aTttaH rpyrrna H aMttHOKMCJIO-

3MMHOKMCJIOTa H a30TMCTOe OCHOBaHne;

4) a30TMCTOe OCHOBaHIIe H uyK-

JieonpoTeH.n;.

A 6. HyKJteomuabt e K.:temKax opzauu:J.Ma 6btnOJlH.R/Om rjJyliKu,uu:
1)
3)

CTPYKTyptty10 H

peu;enTopHyIO;

TpattcnopTHYIO tt attepreTMqecKyIO;

A 7.
1)

2) pe1~enTopttyIO
4) attepreTM'leCKYIO

n

TpattcnopTtty10;

tt 3arraca10I11;y10.

I'aJtaKmoaa uwpoKo pacnpocmpaueua 6 npupoae u .fl6.iJU' mcR:

.'IMilH,ZJ;OM;

2)

rrerrTH,ZJ;OM;

3)

HYKJieOTH,ZJ;OM;

4)

MOHOCaxapM;10M.

Tecmbt

498

A 8. Ko;vi.n.7eJVLCllmapuocmb --- .mw npuHu,un rJono.HHC/lUH o'VlCJ/NlY 11yK,,7COmuaaMU - zyaHU.70Bb/M u:
1) TlIMIUlllJJOBbIM; 2) 8J(eHmTOBbIM; 3) lllITlIJ(lIJ10BbIM; 4) yplI,lllIJIOBbIM.
A 9. 3yxpo.wamuH - .Jmo:
1) cy6cTpaT ,ll.1.H ClIHTe3a 6eJIKOB; 2) 8KTlIBH8.H qaCTb MOJieKyJihI .IlHR'.;
3) cy6cTpaT J(Jrn: ClIHTe3a yrJieBO,iIOB; 4) HeaKTHBHaa qacTb M0.1eKy.1bI
JI:HK.

A 10. Tpemu'luy10 cmpy1Cmypy )JHK cocmae.Miem:
1)

caxapocpoccpaTHhitt ocToa;

2)

ocToB, o6pa3oBaHHhitt caxapaM11;

Ha.H CIIllp8J1h 113 IIOJIHHYKJieOTlI.ll;OB;

4)

3)

.n;aoi1-

KOMn.1eKC J(BOHHOH crr11pa.1111 JI:HR'.

C 6eJIK8MlI.

B.

fetteTHKa

B 1. XapaKmepucmuKou 1Capuomuna HBJtHemc.R:
1)

COBOKYIIHOCTb reHOB

B .Httl{eKJieTKe;

3)

B

opraHlI3Me;

2)

COBOKYIIHOCTb ayxpoManrna

MOpcpoJiorttqeCKlie rrptt3H8Kll opraHH3Ma;

4)

COBOKYIIHOCTh

xpOMOCOM B nepBHOll KJICTKe.

B 2. 3nume;waJtbltaJl KJtem/Ca 'leJtoee1Ca coiJepJ1Cum ... aJtJteJtJl (aJtJteJtb)
oiJHOZO zeHa:
1)

.n;Ba;

2)

TplI;

3)

0.ll;HH;

4)

qeTh!pe.

B 3. Y'lacmoK m-PH K, 1CoiJupy10w,uu aMUftOKUCJtomy. Ha3bl6a10m:
1)

reHOM;

2)

JIOKycoM;

3)

KOJlOHOM;

4)

aHTlIKOJ(OHOM.

B 4. I'e1t0M cocmae.'lJlem Ha6op:
1)
KlI;

4)

KO.ll;OB 11-PHK;

2)

HYKJICOTH.ll;OB T-PHR'.;

B 5. AJtJtCJlbHbte zeHbl pacnoJWJ/CeHbt
1)

3)

reHOB 6aKTeplI8JlbHOll KJieT-

reHOB 3IIlITeJIH8JlbHOll KJieTKlI KlIIIIeqHHKa.

rpyrrrrhI cuerrJiemrn;

2)

6 oiJuHaKOBblX Jto1Cycax:

.n;ayx rpyrrrr cuerrJiettm:1;

3)

roMOJior11qJih1x, no-

4) HerOMOJIOrliqHbIX ayTOCOM.
B 6. B.'lU.JHeu,oebtM MemoiJoM e .weiJuu,uHcKou zeHemuKe onpeiJeJtR10m:
1) reHOM H KapttOTlIII; 2) KaplIOTlIII lI MO.ll;HcpttKau1rn; 3) MOJ(licplIK8ll;HH II

JIOBhIX xpoMOCOM;

HaC.'le.n;oBaHH.H nptt3H8KOB;

4)

THII

TlIIl HaCJie,llOB8HHJI nplI3H8KOB lI THII MYT8l{lIH.

B 7. I { umo2euemu'lec1CuM MemoiJoM e Jl1.eiJuu,uHcKou 2euemu1Ce onpeiJeJtJl10m:
1)

peuecCHBHhie limI J(0MlIH8HTHhie MyTau1111;

3HaIWB;

3)

2)

TlIII HaCJIC.ll;OB8HlI.H nplI-

KOJI11qecTBO xpoMOCOM B HHTepcjJa3HOM .HJ(pe KJieTKH;

4)

KOmiqecT-

BO xpoMOCOM B KJieTKe B npou;ecce MlIT03a.

B 8. Y 'leJtoee1Ca zeH 1Cape2Jta.Jocmu (K) iJoMuHupyem Haa zeHoM 20Jty602.:ta.Jocmu (K), a zeH wecmunaJtocmu ( wecmb naJtbU,ee) (M) Hao zeHoJVL nJtmunaJtocmu
( M ). B ceJVLbe poiJuACJl 20Ay602.rn.Jbtu, nJtmuna.llbtil pe6eHOIC c nepeou zpynnou 1Cpoeu. TaKou JtCe <jJeHomun y Mamepu amozo pe6eHKa. Omeu, pe6eH1Ca - KapezAa3btu,
wecmunaAbtu, c mpembeu zpynnou !Cpoeu. I'eHomun omu,a amozo pe6eH1Ca:
1) KR
JAJA.

MM

JAJO; 2)

KK MM

JBJO; 3)

KK MM

JOJO; 4)

R'.K MM

JOJO; 5)

KK MM

B 9. Y 'leJwee1Ca wecmunaAocmb onpeiJe.ruwmcJl aymocoJVLHbtM iJoJVLuHaumHblM Ze/lOJVl (K), zeMO<jJUJlUJl (h) - cu,en.'leHHblU c llOJlOJVl peu,eccUBHblU npu-

Bon11ocb1

499

B ce.11ue podu:1cJ1 1u1nzuna:tbtil .11a.'Lb'lU1C c 11em8Ppmoil <pynnou KpOfHl,
60.!lhliOU ?e.lto(jJU.'lUCU. Ew poiJumeJlU /[(' 60.rtblibl ?(JJllOljJUJlU('U. re1t0munb1
.>fiat(.

.iza.'tb'IU/Ca -- a) u om4a -

6):

1) a) KK ,J A,JBXhy 11 6) KK JBJOXhy; 2) a) Kl{JAJBXhy H 6) rrnJRJBXhy;
3) a) K1{ ,J 1\JBXhy n 6) KK JA,JBXliy: 4) a) 1mJAJAXHy n 6) 1{ 1{ JO,JOXIIy;
5) a) KK JAJBXhy 11 6) KK ,JBJBXHy.

B 10 . .Y llYP zeli zupoxo8uiJ1wio 2pe61tJ1. ( P) ao.~ufiupyPm 1taiJ 2e11wvt npocmozo ipe6w1. ( p ·),a no ze!laJll 'le pH.oil ( B) u 6e.wil ( 6) Ol(pac1Cu fla611oiJaemc 51 itetlO.'lliOP iJo.wufiupotwHue . I'emPpo.m2om1tb1e oco6u UJvte/Om 20.11y6y10 oKpaci;;y.
C1Cpecmu.7u 'lep1tbtX u 6e.'tbtX 1Cyp c iopoxo8uiJflbtMU 2pe6flJUvtu. B nomo.Mcm 8 e
nO.'lY'lllJlll 20.!ty6b!X Kyp l' 2opOX06U01lbl.'VlU 2pe6flJ1..~U u 20.!ly6btX Kyp c npocmbtJvtll 2pe6flJUvtu 8 om1wutefiuu 8: 1. I'ei-wmun poiJume.!lbCKux oco6eil:
1) BBPP n BBpp; 2) BsPp n sspp; 3) BBPp H ssPp; 4) BBpp n BBPP; 5) B 8 pp
H Bnpp.
B 11. .Y apo.w<jJu.rtbt ie1t iJAUfifibtX 1tpbl.1tbe1~ (V) iJoMufiupyem 11arJ 2e1w;vt ( v)
y1copo'le1111111x Kp1>1Jtbe6 u 11ac.r1eiJyemcJ1 aymoco.MfiO, a 2e1t Kpac11utx i.rza.:1 (A)
iJoJvtll liupyem 1tar) zefio.M nypnyp1tbtX ?.!la.J (a) u fiac.rteiJyemc.R c4en.ie111w
c X-xp0Moco.111ou. Cr;;pecmu.w .JvteJ1CiJy co6oi.J iJJ1u1uwKpbt.!lbtX ca.MOK c KpaCHbtMu ?.!la.Ja,wu u 1copomK0Kpbt.7btX caM406 c nypnyp1t1>t.Mu 2.7a.JaMu. B nomOJHCm8e no.!ly'lUoW rJ.7UfiH01Cpbl.7blX CG.MO/( ll CGM/.,\08 c l(paC/lblMU 2Jta.:laJllU (:i om110ll/('//Ull 1 : 1. I'euomun poiJume:11>c1wx oco6eu:
1) VVXAXA 11 VvXAy; 2) VvXAXA H VvXAy; 3) VvXAXa 11 vvXay;
4) vvxAxa H vvxay; 5) VVXAXA 11 vvxay.

I'.

asomOI~HOHHOe yqeHHC

I' l. Jlo npeiJ.rfO.}l("('fiUIO .lJ'lel-lblX CGMblJll mpyiJllblM JmanO.llt Xll.HU'lel'KOU .380.!l/O/.,\UU .}l("U6btX cucme.111 c'tumaemcR 803fiUKfi06PllUe:
1) MOJJeKyJJLI ATct>; 2) MOJJeKyJJ Jnrnn;~os; 3) Koa1~epBaTHbIX KaneJJL;
4) MeM6pamrhIX CTPYKTYP·
r 2. )[{. B. JlaJnapK 06051.CfiU.7 npU'lU/lbl ll,:i.Jvt('fi'lU60Cmu opzaHU3Jll08."

1) 6opb60tt 3a cyu~eCTBOBaHHe H CTpeMJieHHCM opraHH3MOB I{ cosepmeHCTBOBaHJno; 2) CTpeMJJCHHeM opraHH3MOB K coseprueHCTBOBaHHIO H HaCJie~CTBCH
HOCTblO; 3) HaCJie~CTBeHHOCThlO H ecTeCTBeHHbIM OT6opoM; 4) ecTeCTBeHHbIM
OT6opo:vi 11 6oph6oii 3a cy~eCTBOBaHHC.
r 3. /J.1151. K.rncculjJuKat."1UU pacmeuuil K. Jluuueu ucno.,ib.J06a.!l oco6e11Hocmu
cmpoe11w1:
1) KOpHeii H n,10JIOB; 2) nJJOJIOB n no6eros; 3) no6eroB II necTHKOB; 4) necTIIKOB II TblqHHOK.
f 4. 0Cll06HGJ/ npU'lU/la npo4ecca iJu6ep2<'H/.,\llU."
1) YMCHbllICHIIe qIIcJJeHHOCTH BHJia; 2) cymeHne apeaJJa pacnpocTpaHeHlUI
BIIJia; 3) llIIIPOKOe pacceJJeHHe oco6ei1 BHJia B apeaJJe; 4) B03HHl{HOBeHIIC OJIIIllaKOBbIX npttcnoco6;ieuni:i y oco6ei1 BttJia.
f 5. fl.AH K.llUCCU<jJUKa/.,\Ull JICU80mfiblX K. J/UfifieU llCf10.!lb308U.!l oco6eHIWCmu cmpoe1wJ1.:

Tecmht

500

1) K.illOJ;d y IITHll II 3y6on y :VI.!leKOllHTaKllll.JIX; 2) :iy6oB II T:OH<"lllOCTeii
i.5) KOIW'!H()("l'Cli y ;vrnPKOili!Trl!Oilil!X H i{pbIJJl,eB y I!Tlll(;

y :vL'ICKOJIIJ'Lt!Oll~I1X;

4) Kpbl.'IbeB y IITHll 111;1nLte'IHlll;<1 y :VJ:lPl\OnltTaIOll(HX .
r 6. B pu.1uumue llPpU11/X ,)(j(),Ut!i/UIJ/ltf/;1X ll()pu cyu~ecmr-3('/UlblU (j/{.!IUO HlleCJIU i)ocmUJl(('ILUJl:
1) re11eT1;K11; 2) i~I1TO.'iOfHI: ; ;{) O!C:'J'O.lO!'lrn; 1) ITH . 'lCOHTO.'JVf'l!H.
r 7. JBO.'/IOUUO!lllp,IJ!Oll~eil ('()ll/tlll(<'U //{, S . .lla.1taph· c•wma.rt:
1) BH,11;; 2) ocofo,; 3) rJi<lJ\<lIUP\l; 4) ce:v1eiiCTBO.
r 8. Buobt ecmecnwe11no20 om(;opu:
1) onpe,.'l,e:reHHbiii H npHMOii: 2) ll]i5I:V10ii ll )l.Bll/Ky:1mii; 3) ,i(BIDKYIIlllii
l1 CTa6HJII13HPYIOII~Hi-i;

·1)

CTa6J.I.1113HPYEOllll1H H nenp.~:vioil.

f 9. I'Jl.UB!l'blC npU4llllbl 6op1>61JL .W cyU(N'llWIJRU/lUC. no lf. ,[f.ap6Ull,IJ:
1)

Me,IJ;.'IellHOe paa:vmomeIIH(' OJlI'ilHl13MOB If

BOCTh;

2)

J'CHOT!IJIHlJec1rnH H3M81Jlll1-

f'CHOTH!IWieeKHH l1 <j)EHOTll!IWI('CKHi-i BH;lhl Jl;JM8H'IHBOrT11;

Tl1IIl1qecKaR. rraMeH'-IHBOCTh 11 f:•bICTpoe paa:vrHOffWHHC opraHM3MOB;

4)

3)

<llCHO-

6i,1cTpoe

pa3MHOJKeHHe opraHH3MOB 11 11a:vrettemre yCJIOBHH cpCJ(hl.

r

10. Pe.JyJ1bmamu.i1u :UW.'llOl!UU. no r1. ,7,ap8UH.IJ. }/(i//>l!OmCJl:
µ;rrBepreHUHR. rrpn3HaIWB lI eCTCCTBeHHLrii. crr6op; 2) ecTeCTBCHHhiii

1)

6op 11 MHoroo6pa31re BHJJ.OB;
rattn3MOB K cpe.11.e

r

rrp11cnocoo.1eHHOcTh opraHH.JMOB I{ cpe.ne 0611-

ll3MeH4l1 ROCTh.

11. AflZ.'lUUCKUi' 6a6o4KU

1) c1a6H.'ll13IIpyIOu~eii
3)

OT-

'.V1Horoo6paa11e BHi(OB II npncrroco6.wnttocTh op-

06!1'rattm-1; 4)

HaCJ!eJ~CTBellH<l5I

TaHHR. 11

3)

«,W('JW/l ucm bl»

-- .-JfilO npu.11ep:
2) .7l;I10j)YlITHBHOi'i <l>opMbl OT6opa;

<J>Oj)Mhl OT6opa;

Heonpeµ;eJ1eunoii H3MCH'IHBOCTH;

1)

KOMfoIHaTHBHOH H3MCH'!HBOCT!1.

JI:. AttTponoreHe3
JI: 1. Ila npeiJnoJIOJl(e1w10 y•1e111>1x ar$cmpa.rwnume1w:
1)

0611TaJIII 6oJiee

5

MJ1H :1eT Ha3a,L( II JKHJl!1 CT<l/:(<-l:VIH;

rr xo.nrr.1H ua 'leTbrpex Horax;
Jia ll;O

70

KI';

"1)

3)

2)

JKH.'IH CTa;(aMH

xo;:(11.:u1 Ha 'ICTL1pex uorax H HMe.'Iu Macey Te-

HMeJ111 Macey TCJia /:(0

70

l(f' II pOCT ,no

150 CM.

JI: 2. Cou,ua.,1b1tbtil q1aicmop a1tmponoze11e.w:
1)

ecTeCTBeHHhrii oT6op;

BOBaH11e;

4)

2)

TPYNlBMI ,11;ewre:rhHOCTh;

JI: 3. Mopqio.wzu<tecKoe cxocJcmlW
1)
4)

3)

6opb6a aa cymecT-

nenacJie,LlcTBennaR. H3Mtrn•nrnocTb.

IIJIOCKHe HOrTH;

2)

npu~wmoB c CJe.weeKoJn:

MOHOKYJI.Hpnoe 3pe1rne;

3)

.n:Be CJIYXOBhie IWCTO'l.KH;

rrepBH'IHafl no.1ocTh Te.'Ia.

JI: 4. llpuuu,unuaJtb1we om.1w•we pace
1)

CHOCOOHOCTh K Tpy.ZJ;y;

T.J:eHmo;

4)

2)

te.1w6e•1ec1\"0.M o6u~ecmee:

1

06b€;\1 ro::ro1rnoro M03ra;

3)

crroco6HOCTb I{ 06y-

<t)eHOTHIIHqecKHe 11 Hac.rreJJ.CTBCHHhie upnaHaKH.

JI: 5. Buo.rwiu'lecKuu rjJaKmop aumponozeHe,w:
1)
4)

3)

HCKYCCTBeHHhIH OT6op;

rrpH3HaKH, CBOJICTBeHHLre JlaJINtHM rrpe.n:KaM;

2) opraHhI, yTpa'l'HBIIIHe

pa3BHTl1e peq11;

2)

ecTeCTBCHHhTJI

OT6op;

TPYJJ.OBaH .ZJ;eHTe.'IhHOCTh.

JI: 6. AmaBU3.M.bl
1)

-

3mo:

Bonpocbt

501

CBOe 3H8lJCHtte;

3)

npoueCChI yctt.'1eHIUI cpyHKill1H 0,ll,HOI'O oprana;

4)

npOI(CCCbI

yc11;1cHttH cliyHKUHH cttcTe:vi opraHoB.

)1, 7. /l,.1tJt aBcmpa.rnnu mP1W6 6bt.1to xapaKmep1w:
1)
4)

pa3Bl1THe pe•n1;

2)

CTa,nHhlH o6pa3 :>Kl13Hl1;

3)

061,e::.1 M03ra

1100

CM';

11c110.1b30BaHtte cJroflrnhrx opy.l\HH Tpy,ll,a.

)1, 8. B .JJ116puoua:tblWJ1t nrpuo()e 'iP.Jto6eKa xapaKmepllbte iJ.tJI xopauubtX
npu3HaKu -- amo ua.qu'iue:
1) JlerOlJHb!X MelllKOB H II03BOHO'-IHOro CTOJI6a; 2) II03BOHO'!H01'0 CT0.'16a
Il cep.n;ria Ha CIIIIHHOH CTOpOHe; 3) cep.n:ua Ha CIIHHHOII CTOpoHe lI xop.n;hr; 4) xopJl,bl H ma6epHbIX IQeJreii.

)1, 9. B 360Jt10u,uu pewa10UJ,UM waioM ua nymu om o6e.ibJUtbt K 'ie.1106P1fy J16U.rtocb:

1)

npHMoxom.n;eHtte;

m1111eii;

JR.

4)

2)

BHyTpttyTpo6Hoe pa3BIITtte;

3)

IIHTatttte flrnBoTHoii

KOpM.'1eHHe .n;eTeHhIIIIeii MOJIOKOM.

:J1W.'IOrHH

JR 1. l'pynna peoyu,eHmo6:
1)

xeMOTpOCpbr;

2)

cpOTOTpocphl;

3)

3eJieHhie pacTelIHH;

4)

6aKT€plll1 caIIpo-

cpHTbI.

JR 2. B JK0.1to2uu npuMP1t.R1omcJl Jvtemoobt ucc.1te006aHuJl:
1)

r116pH,ll,OJ'10rH'-IeCKHH ll llllTOreHeTHlJeCKIIH;

rrepIIMel!TaJJbHbIH;
Hlle;

3)

2) Ll,HTOreHeTH'IeCKHH

ll 3KC-

3KCIIepHMeHTaJihHbIH l1 MaTeMaTttlJeCKOe MO.n;e.TJHpOBa-

4) MaTCMaTH'-IeCKOe MO,neJIHpOBaHne ll rn6pllll,OJIOI'HlJCCKHH.

JR 3. Buou,euo3 - amo co6oKynHocmb:
1)

HeopraHtt'-IeCKOH cpe.n;hI ll paCTHTeJibHbIX opraHII3MOB;

l1 :>KllBOTHblX opraHH3MOB;

,i~hI;

4)

3)

2) paCTHTe.'1hHbIX

)KllBOTHbIX opraHll3MOB l1 HeopraHll'IeCKOH cpe-

HeopraHtt'-IeCKoii cpe.n;hr H MttKpoopraHtt3MOB.

JR 4. K npooyu,e1tmaM omHOCJlm:
1)
Bei1;

3eJieHbie pacTeHnH;

4)

2)

6aKTepIItt carrpocpttThr;

3)

rrapa3HTHlJec1rnx '-Iep-

napa311Ttt'-IeCKHe 6aKTepnH.

JR 5. :J1wcucmeMou Jia3bl6a/Om coBoKynuocmb:
1)

)KllBOTHb!X opraHH3MOB;

c HaCeJIHIOll~llMll ero Bll,ll,8Mll;

4)

2)

paCTHTeJibHb!X opraHH3MOB;

3)

6HOTOI1a

paCTHTCJlbHbIX ll )KllBOTHbIX opraHH3MOB.

JR 6. KoucyMeHmbt - ::Jmo op2aHU3Mbt-nompe6ume.r1u:
1)
4)

OKCH.n;a yrJiepo.n;a;

2)

,nHOKCH,ll,a yrJ'lepo.n;a;

3)

opraHWieCKHX BeIQeCTB;

3HeprIIH CBeTa B BH,ll,e cpOTOHOB.

JR 7. H au60.1tbmee K0.1t1t'lecm60 6uoMaccbt 06pa3yemcJt 6:
1)
4)

OKeaHe u TYH,ll,pe;

2)

TyH.n;pe H JieCOCTerrtt;

3)

JieCOCTeIIH 11 TpOIIHKax;

TPOIIIIK8X H yMepeHHOM 30HC.

JR 8. no muny accuMUJtJtu,uu JICU6bte opiaHu3Mbt 06aeouHe1tbt 6 zpynnbt:
1)

reTepoTpOcpbI ll cpOTOCIIHTeTIIKH;

n aHa3po6hl;

4)

2) cpOTOCHHTeTIIKll

II a3p06br;

3)

a3p06hI

aHa3po6hr tt reTepoTpocpbr.

JR 9. Buo.Macca a6mompo</JHbtX opiaHU3.M06 Ha 3eM.1te 06pa3yemCJl 6 pe 3y.1tbmame UCllO.llb306aflUJl UMU:

Tecmbt

502

1) KHCJTopo;ui; 2) co:rne'-IHOH :J11cprm1; 3) aTMocc}iepttoro a3oTa; 4) opraim'feCKHX

BeU~CCTB.

10. Ilpooyu,<'1tn1b1 - .Jmo opiaHu.u1n1-nompe6umP:1u:
1) KHCJJOPOlla; 2) 3Hepr:im CBeTa; 3) OKCHJla yrJ1Cpo.ua; 4) opraHH'feCKliX BC-

)I{

u~eCTB.

60TAH~KA

1. KoJutamepa.'tbllblU cocyoucmo-60.1zoK1tucmn1il ny'lOK xapa1CmPpuayPmcJl
meM, 'lmo:
1) MCJK/IY .11y'-IaMH .upeBeCHHbl paCIIOJIOJKeH :1y6; 2) Bero I~eHTpe eCTh .11y6,
OKpymeHHhIH .11;peBeCHHOH; 3) llPeBeCHHa H J1y6 COIIPHKaCalOTC.H TIO 0,llHOH CTOpotte; 4) Bero ueHTpe ecTh ;weBccutta, 01<pymeHHa.H JTy6oM.
2. lf ('l(('6U 'll(U cmP6JlR pacmPllUU Jl6Jl.HIOmcR y'laCmKaMu:
1) KOpb1; 2) KOPKH; 3) 3IIHllePMhl; 4) 3H/IOllepMhl.
3. 3a C'lPm ecmaeo'l1wu 06pa3oeamP.1bHOU mKaHu y u,6em1coen1x pacmeHuu:
1) Y.UJIHIUJ!OTC.H cTe6Jrn n pa3pacTa10Tc.H J1HCTh.H; 2) pa3pacTa!OTCH JIHCThH
H Y.UJIHH.HIOTC.H KOpHH; 3) Y,ll,J1HHHIOTC.H KOPHH H pacTyT B TOJIIIIHHY CTe6m1;
4) pacTyT B TOJIIIIHHY TO.TJhKO CTC6JIH.
4. ff,PHmpaJlbHblU OCe60U lf,UJ1UHOp 6 MllOi!O.~PmHPM OpP6PCHOM CmP6JlP JlUllbl 06pa.Joea1t:
1) ;1y60M H llPCBCCHHOH; 2) .upeBeCHHOH H cep.11;11:eBHHOH; 3) cepllneBHHOH
H Jiy6oM; 4) Jiy6oM H BTOPH'IHOH KOpoli.
5. KJty61tPJtyKoeuu,a - 3mo 6uooua.Me1tPHHbtu:
1) JIHCT; 2)no6er; 3)KopeHh;4)CTOJIOH.
6. BblOWiUilcR cmP6PJlb y:
1) XBOIIIa; 2) XMCJI.H; 3) IIJiaytta; 4) BHHOrpana.
7. I'oiJu'lHbte KOJtbU,a - 3mo Ko.inu,e6bte cJtou Ha nonepe'lHO.M cpe.Je:
1) CTe6m1 - B Jiy6e; 2) CTe6JI.H - B l1PeBec1111e; 3) KOPHH - B nepBH'fHOH
KOpe; 4) KOpHH - BO BTOpH'IHOH KOpe.
8. Bmopu'lHaJl Kapa mpexJtemneu eemKu .1zunb1 o6pa3oeaHa:
1) llPeBeCHHOM H ;iy6oM; 2) Jiy6oM H napeHXHMOH KOphI; 3) napeHXHMOH KOpbl H KaM6HeM; 4) KaM6HeM H llPeBeCHIJOH.
9. Bmopu'l~taR opeeecuHa pa36U6aemcR 6:
1) CTe6Jie O/IHO/IOJThHOro pacTeHHH; 2) KOpH.HX H JIHCTb.HX O,ll,HO.ll;OJihHOrO
pacTeHH.H; 3) CTe6Jie ,ll,peBeCHOrO llBY,ll,OJlbHOro pacTeHH.H; 4) KOPHHX TpaB.HHHCToro O,ll,HO/IOJlbHOrO pacTeHH.H.
10. CmoJLOHbt Kapmo<j)('JlR - 3mo euiJou.J..111eHeHHbte:
1) no6eru; 2) KOpHeKJiy611u; 3) KOpHeBHIIW; 4) KOpHeIIJIO/lbl.
11. B ceJtbCKOM xoaHucm6e ucno.11bayemcR 'lepeHKoea1tue npu pa3MHOJ1CeHuu:
l)KaKTYCOB;2)BHHOrpa.11;a;3)T10JlhIIaHOB;4)3eMJI.HHHKH.

Bonpoc1>1

12.

503

Ha r>epxywKe Kop11er3t1U<a n1>1pe.H .110)1(110 0611apy.1!(11nu,:

1) !WpHeBoii 'iCXma;:; 2) IIPHARTO'!llhJe Kopnn; 3) BcpxymetmyIO TIO'IKy;
1) BTOPl1'IHYIO Mepucn~:viy.
13. JJepBU'lHbte 1u1emKu llb1JlbllU1>a 1wKpb1moce.!lU'H111>1x pacme11ui1 rJe.'tJLm
CJ/ .11eiw.Woll c 06pa.J06a11ue31;
1) cnep:vrneB; 2) MHKpocnop; 3) nihLeICieTKH; 1) aHTepHL(HPB.
14. CfJ06o<huuenecm111>1il Be!l'lUK UJ11eem u,Bemo1c:
l)BHKH;2)p03hl;3)ITOMUL\Opa;4)KapToct>enH.
15. IIepeu'lJtafl 1uemKa ceJ115t.Ja 1tam1rn no1cpb1nwce,11eH1tbLX pacme11uil iJe.'lU mc.H .11eilo.JO.'li c o6pa .Joe a 1tue JW:
l)cuop; 2)crrepMHCB;3)HHUeKneTOK;4)aHTepnti;HCB.
16. Cou,eemue c.WJl(HblU .w11muK Uol!eem:
l)pomh;2)pH6HHa;3)MOpKOBh;4)qepeMyxa.
17. B pe.>y.rtbma me .!Jieilomu •tecKozo de.11e1tu5t nep6u 'lHbLX K ..qemo/C nbt.rlbH uKa zopoxa nocee1wzo 06pa.Jy10mcJ1 'lf'mb1pe:
1) raMeThI; 2) MeraCIIOphI; 3) MHKpocnophr; 4) BereTaTHBHh!C Ko"IeTKll.
18. ll,.rtJL npou,ecca aeou11ow on.rwaomeopeHUR no1Cpbtn10ce.l!e1111b1x pacme11uil xapaKmepHo 06pa.Joea1tue:
1) JLBYX apxcro1rneB; 2) Meracrropoq>HJwa; 3) TpHrrJIOH,nHoi1 I<JICTKn;
1) J~HIIJJOHtlHOI'O 3HJLOCrrepMa.
19. Ha u,eemo. 1t0.l!(e U,6em1ca euw11u pacno.rwza10mc51:
1) rrpnuBeTHlIK H IlpHJIIICTHlll{; 2) npH.'11ICTHlIK H rnHelLett; 3) rHHeI{eiI
H 8HL(poueiI; 4) RHL\POuciI II apxerOHHH.
20. Bo epeJ11R no.ruwzo BblCbtXaHUR y .rwwai11w1Coe:
1) .[\hIX3HHe 3KTHBH3HpycTCH; 2) cpOTOClIHTe3 rrpeKpamaeTCH; 3) ct>OTOCHHTe3 38MC.[\J'1HeTCH Ha 50 'X,; 4) o6MellllblC 11POILeCChI He H3MCHHIOTCH.
21. Y n.rwy11a no6ez r3emeumcJL:
1) CHMITOL(H8JihHO; 2) MOHOITO.[\H8JihHO; 3) JLHXOTOMlllJ:CCKH; 4) J'10)KII0.[\HXOTOMH'IeCKll.
22. 1111ou1camopaJ11u 'lucmomb1 eo3ayxa Re.11.R10mcR:
1) rpH6hr; 2) XBOllLH; 3) BOL\OPOCJIH; 4) JIH1II8HHHKH.
23. J'aJ11emo<jJUm n.rwyua 6y.1weoeua1wio - pacmeuue:
1) tLBYL\OMHoe; 2) reTepoTpoqrnoe; 3) 06pa3yromec crrophr; 4) qJOTOCHHTe3npyromee.
24. B.Jpoc.rwe pacme11ue 1C!J1CYW1CU1ta JtbllG uMeem:
1) rrpOCThie JIHCThH; 2) KOpHH c rrpHCOCKaMH; 3) pa3BCTBJ'1CHl!h!H CTe6eJih;
4) nerrap110rrepHCThre JIHCTMI.
25. B euae (J(';JUH<Puu,upyJOu~ux no6Jl.JOI( npu ueo6xoowvwcmu MOJICHO ucll0Ab306Gm b."
1) JI8MHHapHIO; 2) OJICHHH Mox; 3) MOX ccparHyM; 4) KYKYillKMH MOX.
26. I'aMemo<jJUm xeo~a noAeeoio - pacme11ue:
1) OJLH0.'1CTHee; 2) TpaBJIHHCTOC MHOrOJICTIIee; 3) ue HMeIOmee H8CTOJIIILHX
KOpHeii:; 4) HMCIOmee nHCThJI c pacqJieHeHHOH JIHCTOBOH ITJI8CTHHIWH.
27. Cnopo<jJUm MXa c<jJamyMa:

Tecml>l

504

1)

HMeeT .~.pesec1rny; 2) HMl'C'l' p!L1011Jlbl; ~~) O;J;HO,IJ;O'.\nIOe paCTCIIHC;

'1)

HMe-

eT KOJHICBYIO CHCTeMy.

28. B
1)
3)

u,wotf' pa.wunzuJl nanopom1w1;1u3;

cnopL!

i\11 HT030~•1;

cjl0p!V111pyi0TCH

3apO,D;hlHI pa3BHBaeTCH ll3 ;mrOThl:

29.
1)

2)

-1)

r11:v1eTbl

<llOpM upy10TC5I MeH030M;

cnopmjmT p<:3Bl1B3C'l'CH 113 cnopbl.

JlfUUl/W-CnzpOOU.•1bl OfipG.3.IJIOl/1('51 .IJ."

2)

XBOma;

3)

IlJlayHa;

mrnopOTHIIKa;

4)

,;rncTRCi:!H111Lhl.

p<'.f,lj/Lbmamc .Meuomu 1U'Cl(OVJ cJe.'l('lfUR nep6ll 1lllblX
1tam1Ca 6 u,BemKP R6.w1w ol5pa.;y1omc;1:

30. B
l)

raMeTJ,;;

2)

;3)

MCracnOphI;

apxeromii1;

4)

/(,'/('!71()1(

ceoltR.JU-

K.1CTKl1 3llLl,OCIICpMa.

aoonorl1S1
1. 1\lfri)y.m yiJa.11J1Pm 11enPpPeupe1111b1e ocmam1w nuw,u '!.epe.J:
1)

attyc;

2)

rropornm1y;

3)

1) BhlLl,CJ1HTeJ1hHhie

4)

aHa.'IhHYIO rropy;

2. Y u1upy.wpuu-my<jJe.r1b1CU

poTOsoe OTBepcTtte.

rzpou,eccbt oc,iwpczwrnu,uu BblnOJ!flJ110m:

BaKy0.'111;

2)

nopom1111a H }\jJeT04HaH rJIOTKa;

3) J~jWT0'-1-

HaJI rJIOTKR 11 TpHXOIIHCThI; 4) TPIIXOlUICThl.

3. Y Jlto.J11F11wu n .•w11apuu, e om.rw'lue om zuiJpbl, ecmb:
1)
Hlrn;
11

3HTO,[\epMa H n.1aMCHHbie K.'leTF\H;

3)

2)

II.'IaMeHHh!C !CleTI\11 11 opraHbl 3pe-

opr::tHhI 3peHIUI l1 ,L\hIXaTeJih!HHI CHCTCMa;

Bbl/~eJIHTCJlbHbie

3)

f!C'-IC!lll

CllCTe:vra

coca.1bu~u1Ca npoxoiJum e:

4. T!o.rtHu1u 1iunJ1 pa.mumuH fl<"fflW'llW?O
1)

4) l~b!X3TCJ!bH3H

rrpOTOKH.

qeJ10Beirn

11

BO,L\e;

2)

so;~e H

TIC'!CHH

ne'ICHH BOLl,HOro MOJ1·-IIOCIW 11 rreqeirn: 'ICJIOIWKa;

4)

BOll,HOro MOJIJIIOCKa;

IIE''-ICHH '-IeJIOBeKa, BOll,-

Horo MOJIJIIOCKa l1 BO,L\e.

5.

IIep£3U'IHUJl no.wcmb me.rw:

1)

eCTh

y

IIJIOCKlIX qepseii: 11 HMeeT ;)f!HTe.1n3J1bHYlO BhICTllJIKy;

3fllITCJIH3.'1hlIYIO BhICTH.iIKY l1 eCTh y EpyrJiblX '-ICpBeii;
Bett 11 He HMeeT 3IJHTeJIHaJihHOH Bh!CTHJIKH;
CTHJIKH H CCTh

y

4)

3)

CCTh

y

2)

HMCCT

Kpyr.JibIX qep-

He HMeeT 3IIHTeJ1Ha.'1hHOH Bhl-

KOJih4aThIX qepBCH.

6. B JICPJLy<loK KOJtb'lamoio '!epBR onll(pbwa10mcJ1 npomoKu .HCPJteJ:
l)

3eJieHhIX;

2)

CJ1H3HCThIX;

3)

l<OKCaJlhHhIX;

4)

n;.rneCTKOBhIX.

7.

061«u.nu .111op</Jo.w2u'tec1Cu111u npu.nw1Ca.11u .tW.'lO'lHOil n.ia11apuu, ne'leHO'l11020 coca.rtblll,UKa u C6UH020 u,enH.R JlB.'lJllOmCJl."

l)

Tptt C."IOH Mhlllill H TeryMeHT;

Macpp0,L\HTH3:vl l1 MeTaHecppH,L\HH;

4)

2)

TeryMCllT 11 repMacppO,L\HTH3M;

3)

rep-

MeTaHecppHi~HH 11 CTBOJIOBaH HepBHaH cnc-

TeMa.

8. JI eio'I Hoe Obtxa11ue
l)

pe4HOI'O paKa;

2)

xapaKmepuo iJ.~R:

6eJIOH IIJiaHaptt11;

3)

KOMH3THOH :vryxtt;

4)

60JihIJlOro

npy,L\OBlIKa.

9.
1)
Ke; 4)

Oco6eH1wcmb nuu~eBapumeJLbHou cucmP:Wbl npyiJoeu1<a:
OTCYTCTBHe rreqeHtt;

2)

3aH:aH'IHBaCTCH c.ierro;

pa3BlITHe ITO,L\JRe.'ly,L\O'IHOH me.Tie3hl.

3)

pa3BHTIIe TepKII B r.10T-

Ronpo1.'1Jt

505

10. Ep1ow1w pr 111w10 pa"a u ..weern KOll("llfocmu, <lh1no·11rn1ouu.e (/J.l JNKl<uu:
1) 33XBa"ra 11 TJ:taBilHIIH; 2) ll.13B3Hlll1 11 no.:IUBhlX opra HOB CaM !\OB; ;3) TIOJIORhIX opraHOB CBMHOB a ncpCTIIp3Hl15I IIllln11; 4) rr0peT11pan11rr lll!Il(H H IIO.'IOBh!X
opraHOB CRMO!{.

.w

11. Rcac1>[(W/OU(aJ1 nonepx1wc1111> f(Utue 1111u1w nayK006pa.J111>1x y 6 e:iu•irita
Cl/Pm R1>1pocmof3:
1) CTeHKH LTJOTKH; 2) 3RJiHei1 KIIIIiI·:11; :i) cpe;rneii Kl!IIIKII; 4) :Hn1em1.
12. 06UJ,tOIU llpUJHUKllJitll llU!JIW I.I 6a6o·um 5W.UU0/l1('Ji,'
1) TeryMeHT H CMCW3HHRH II0.10CTh TCJHJ; 2) CMemaHllR5I llOJ!OCTh TC.1a

33l\1KHyTa.H KpOBCHOCHRH CHCTeMa;
II 3ec1CHbie fKCJie3hI;

1:3.
1)

4)

ae.TJCHh!P meJIC3bl H 33MKHyTa5I KpoBeHOCHl'iH CHCTeMa.

CH.ll(K06W1 .11(<'.:te.w

Rll'IHIU( CaMKH;

I! HC-

3) He3aMl{HYTRH KPOBCl!OCHR.H CHCTeMa

2)

y pP'l1w20

pa Ka -·- amo 1zapH111u:

ceMeHHIIK Cil.MlL3;

3)

4)

opraH BhI.D;e:remrn:

opraH XH-

MII<JPCKOro qyBCTBa.

14.
1)

I{P11mpa.1b1rn51 flepe11aH cucmeJwa .WHW'm11u1rn npeucmau:1ena:

2)

.'l;By:vui: OT.ll;CJiaMH ro.TJOBHOJ'O :vioara H CIJHHHhIM M03l'OM;

3)

M03f0M H TpCM.H OTn;e.T!ilMH rOJJOBHOro :\103ra:

1"1031'H II HepBHOH Tpy61wii;

15.
1)

4)

CIIIIHHhI'.\1

TpCM.H OTJleJia:Y!H rOJJOBHOl'O

HepBHOii Tpyfowii c 11enpou:eJ1CM.

XapaKm<'pllbtil npu.utaK !itM6011o'l11b1x:
OTCYTCTBHe 1 1emocTei1:;

xaHHH

-

2) TPII OT.D;eJ1a
4) H8.TIIIqne

.1erO'lHhie '.\ICillKH;

l'OJIOBHOrO M03ra;

3)

opraHhl .ll;hl-

;i,mI)ciJepeHIUIPOBam10ii HepBllOH

Tpy6Jrn.

16. 06w,a5l
1)

CTOpOHe;

17 .
1)

4)

2)

CTBOJIOBa}I HepB!I8}1 CHCTeMa;

3)

ccp.11;ue Ha Cl1I!HHOii

xpmqenoil: BHyTpeHHHi1 CKeJ1eT.

.;7Ulll{Pll1HUI( u 6PCll03BOH0'/Nhte cxoihtbl.'

1rn;mqueM xop.D;hI B 3Mfipttoretteae;

2)

ttaJ1HqneM cep.11;ua B 3M6p110rette-

OTCYTCTBIICM paJ~H8.!IbHOii CHM:\1CTpHH;

3)

3e;

'iepma opi'aJfU,JUU,UU fiPC'U!penHblX u no.UWHO'lllb/X,'

Ha;rnqne ueJro:vrn;

4)

pa3BHTHeM Tpex 3apO.D;hlllle-

BhIX J111CTKOB.

18. I7paezt.TlbflbtJll J1B.rrnemcH cy.wiJeHue:
1) BIIHOrpamiaH y.TIHTh:a - l'CpMac}JpOLJ;IIT; 2)
J1el'O'IHhlC MCIIIKH;
JLhI;

4)

opraHhl JLhIXRHHR KJieru:ci'1 -

y ma6hI B I'CMl'l.TJhHOM rrnHaJie .1emaT KpOBC'HOCHhIC cocy-

y ll\YKH cerMeHTIIpOBRHLI BhI.D;eJIHTCJ'IhHa.H H IJ0J10B8H CHCTCMhl.

19.
1)

3)

IlpaBU.1bllhl.'.t .flB.'lJlPmCR cy.ll({}('f!U(','

.1eBaH ,n;yra aopThI y IlTHI( llCCeT apTepHIL1hHYIO Kf>OBh;

3TO COBOKyTIHOCTb yc.lIOBHhIX pec]lJICKCOH fHlCCKO'.\llbIX;
BhIHOCMT BeH03HYID KpOBh H3 cep,nia )tW6hr;

4)

3)

2)

HHCTHHKT -

apTepnaJihHhIH KOHYC

T330Bh!H IIOHC mnepIIU:hI o6pa-

aosaH np01wpaKOHl{OM, KOpaKOH.ll;OM II .nofoWBhl'.\111 KOCT.HMII.

20. Henpatill.'thHhtM J1a.'1J1emcJ1 cy.11coeui1e:
1) 1rncTenepb1e - · rJ1y6mmr,1e phr6h:; 2) ayTOTOM:IIH
pnn;

3)

pa3BIITHe CBepqKQB IIPOXO.ll;HT

c

xapaKTep1rn .n.1.H aru:e-

ITOJlHhIM MeTaMopq)030M;

4)

ueJ'IOMO-

JLYKThl paKa coe.11;nH.HIOT l(CJ10M c BHPII!Hett cpe.D;Oll.

21. llpGBU/lbHblJll f/6.'7.<lCTnCJl C.lJ.)/(0('/IU(','
1) acj:>pHKRHCI<He CTpayChl -- caMhJe KpynHhie

TITIIU:hI;

2)

y KOCTHCTblX pbr6 tteT

506

3) y 6hI'Ihero OlWJ(a :IH'll1HKH oKyK.1HBaIOT4) ceBCpHbIH OJICHb 11 JIOCh - nenapHOKOIIblTHbie IBIIBOTHbie.
22. II 06l'px1wcmb ml'.'l no.wo1uwe y p1J16:
1) IL'"IOCK<1H; 2) /(BOHKOBOrHyTaH; 3) l(BO}I!WBhlIIYKJJa}[; 4) TIJIOCKaH c o;rnoii:

OCTaTKOB xop;(bI Me):l<JJ;Y II03BOHKa:vrn;

CH IIO/( KO}!{Cl! 1wpoB;

11 Bh!IIYKJiaIT c JJ;pyroii: CTOPOHhl.

23.

y

Apmepua.11b11aJ1 1tpo8b

1)

2)

npeJJ;CCpl(ttH;

cyJJ;aX;

4)

24. y
1)

01(_1JHJ1 me'lem H:

KI{)BbCpOBbIX npoTOKax;

3)

BhIHOCHll(HX ma6epHbIX co-

npHHOCHll(HX ma6epHb!X cocyl(aX.

.JCJWIW600HblX nOR6LIJlUCb npo2peCCLl6Hbll' 'lepmhl op2UllU.JaU,LILI."

2)

cepJJ;ll,C 11 0/(HH mettHhlH fl03BOHOK;

Baa JI1111115I;

3)

O,ll.I1H meiiHhIH 1103BOH01{ H 601w-

6oKOBaH JJHHHH l1 ,l.l,Ba I<pyra KpOB006paII(eHm1;

1)

)];Ba Kpyra

KPOB006paIIWHI15I H TpexKaMepHoe CCp.L\I(C.

25.
1)

JiuMrjJamU'l€CKLll' JWKlJHbl .•1.fl2.tJUleK HQXOOJllnCR."

ITO)]; KO/Keii;

2)

3)

110/( ,l.l,I1a<llpar'.'.10ii;

B o6.18CT11 rrJieBpbI;

4)

B o6;iacTH rre-

p1rnap,na.

26.
1)

4)

Mo 11Peu1w RBJrnemcR 1to1te 1rn1J1.M npoay1cmoM o6MeHa y:

aKyj!bl 11 JIJirynrnn;

2)

HlI(epHll,bI 11 CTpayca;

ApoMoprjJo.J .MbtUle'l1t0il cucme.Mbl npec.1111J11ta1ou~uxc.H -

27.
1)

KO}!{Hhie;

29.
1)

4)

MeTaMep1111 weiittOI'O OT)J;eJia;

2)

ITOTOBhIC;

H aHyca;

1)

31.
1)

3)

4)

MhllIII(-cr116a-

Jmo 6UOOU.J.MeHCHlllJI(' ... Jl((' ..1('.Jbl."

CJIIOHJlhie;

4)

I'JIOTOl.JHb!C.

06U{lle npu.maKU .JC.'lt/W800/-l1JlX LI npeC.Mbl/{Q/O~LIXCR -

Ty.TIOBllll(HbIX IIOl.JCK;

30.

Jnw IWR6.'ll'llUl':

3)

JJ:llctlqlepeIII(llpOBaHHbIX MhJIIII(, KOHCl.JHOCTCH.

5laoeum1Jte JIN'.~('.Jbl 2QOIOKLI -

28.
1)

2)

Mempe6epHhIX Mbrrnu;

TeJrett 11 pa3rn6aTeJiett;

2)

3)

l(ByxKaMepnoro cepJJ;u,a;

;mw

/lQ ..qu'IUe:

MO'ICBOI'O ny3blp5I

BeH03HOH, apTCpttaJibHOH, CMCIJI8Hf!Oii KpOBH B cep)J;ue.

PyJteGbte nepbR nmuu, 06pa3y10m n.wc1tocmb:
rnc11;

2)

KpbIJia;

3)

XBOCTa;

4)

TYJIOBHll(a.

C1te.11em coJty6R 06.rtn'leH ;JQ C'lem:
rrOJJ;BIDKHOCTll

MhillIU

11 ITHeBM8THlJCCI01X KOCTe:i1:;

4)

3)

JIHryrnKll 11 Hll(CpHl(hI;

cTpayca 11 :vrne1wn11TaIOll(HX.

3)

11 I<pyrrHhIX

r.1a3HHI(;

2)

Kpyrrnhrx

rJia3HHU

ITHeBMaT11qeCKHX KOCTeii 11 061,eMHOll rpyJJ;MHhI;

06'heMHOll I'py,n11Hhl 11 peJJ;KllX 3y60B.

32.
1)

Apo.111oprjJ030M a.HR .'Vl.R€KOnuma10~ux CJl.IJJl(Um nORG.1enue:
l(Ha<tlparMbr;

nocTci1;

4)

2)

'ICThtpexKaMepttoro cep.n;ua;

3)

IIJIT11rra.1oro T11na 1wnel.J-

CTpeMe'!Ka 11 M0.10TO'IK8 BO BHyTpeHHeM yxc.

AHATOM"1SI

1.

K coeiJu numeJtbllblM mKGHR.!fl omuoc;im:

1) arr11l(ep:v111c 11 peT11Ky.1apHyIO; 2) peT11Ky:rnp11yIO 11
BYIO 11 3II11TCJ!Htt;

2.

4)

xpHll(eByIO;

3I111TCJI11H 11 CKeJieTHyIO.

Kocmepa3pywa10~ue KJ1em1tu:

1) OCTCOHhI; 2)

OCTeOI(llThI;

3)

OCTeOKJiaCThI;

4)

OCTeo6JiaCTbJ.

3)

xpmu,e-

BonpocbL

507

3. Bo.wcb1 y 'l('Jt06P1ca «6cma10m db160.lli» 6 pP.>y.ibmamP:
1)

ycmremrn cpyHKIJ;l1l1 ca.1bHbIX IBeJie3;

3)

2)

yc1rneHI1H cj_)yHh'.11.Hll IIOTOBLIX

4)

paccJrn6.1emrn

4. CPpor erw,Pcnuw .1wo.J2a 'lPJWBrh·a o6pa.wrwuo:
l) peu.enTopaMH ll Te.1aMH Heii:ponos; 2) Te.'laMn HeHpOHOB

11 ,Ll;CH.11,pHTaMII;

me.TJe3;

COKpalll,eHHH Mb!IIIU., IIO,Ll;HllM8IOlll,llX B0.10CbI;

Mbllllll., IIO,Ll;HHMaIOlll,llX BOJIOCbl.

3)

,n;ett,n;pHTaMH H ueiipoI1aM11;

4)

HeiipottaMH H aKcOHaM11.

5. CMbLKGHUP BPK y 'lPJWBPIW BO 8peMJt cu.ibHblX nopbtBOB Bempa -- ..nno pe.JYAbmam:
1) 6e3yC.'IOBHOI'O pecj_)JieKca, Tpe6yIOIIJ;CI'O o6pa30BaHIIH BpeMCHHhIX HCpBHhIX CB.H3eii:; 2) 6e3yCJIOBHOI'O pecj_).'IeKca, ocyui;ecTBJI.HIOlll,erocH no I'OTOBbIM
HepBHbIM CBH3.HM; 3) yCJIOBHOI'O pecj_)J]:eKca, Tpe6yIOlll,CI'O o6pa30BaHIIH BpeMeHHblX HepBHhlX CB.H3eii:; 4) ycJIOBHOI'O peq)JleKca, ocylll,eCTB.'IHIOlll,erocH no l'OTOBblM HepBHbIM CB.H3.HM.

6. JI uca nouMa.rta .Jauu,a, yc.ibtwae 6b1cmpP.1t oxomHU1ca, 6blllycmu.1w pzo
u 6pocu.iacb naymeK. Y .11ucbt 80.JHUKAO mopMOJ/CeHue:
1) yCJIOBHoe;

2) 6e3yCJIOBHoe; 3) 3anpe.n;eJibHOe;

4)

llH.ll:llBH.n;yaJibHOe.

7. Xpycma.iuK ZJta.Ja 6btnoAHJlem <jJyHKU,UIO:
1)

38lll,HTHYIO; 2) peu.enTopJiyIO;

3)

npeJIOM.TJ.HeT JIYLIH; 4) onpe/],eJIHCT U.BeT

rJiaaa.

8. C.iyxoeou aHa.iu.Jamop - anw:
1)

yrnHa.H paKOBHna; 2) yrntta.H paKOBirna ll CJIYXOBOH I!CpB;

HepB ll KOPKOBblH 11.eHTP BHCO'IHOH .ll:OJIH;

4)

3)

c.1yxOBOH

KOpKOBhlH 11.eHTP BHCO'IHOH /l,OJIH

ll TpH C.'lyXOBbie KOCTO'IKH.

9. y 'l('A06eKa 6/(YC06blP COCO'lKU, pacno.1t0JIC('HfiblP no Kpa!O Jl,JblKG, 'lyBcm6UmeAbHbl K.'
l)KHCJIOMy;2)cJia.n;KoMy;3)coJieHOMy;4)ropbKOMy.

10. llymb nocmyn.ielluJl zopMOH06 u.J K.iemoK w,umoeuihwu J1Ce.ie.Jht:
1) Henocpe.n;CTBeJIHO B KpOBh; 2) B JlllMcj_}y ll .n;aJiee B KPOBb; 3) B TKaHeByIO
)Kll.ll:KOCTh ll .n;aJiee B KpOBh; 4) B TKaHeByIO JKH.ll:KOCTh H .n;aJiee B JIHMcj_)y.

11. CaxapnbtU iJua6em y iJemeu pa.JBueaemcJl npu:
1)

rwnocj_)yHKll.llll lll,llTOBll.ll:HOH :>KeJie3LI; 2) rHnepcpyHKU.llll lll,HTOBll.ll:HOH

meJie3hI;

3)

rwnocj_)yHKlI.HH no.n;meJ1y.n;oLIHOH meJre3LI; 4) rm1epcpyHKlI.Hll no.n:-

meJiy.n;oLIHOH IBeJie3bI.

12. 3a Cl/em HGiJKOCnlHUU,bl KOCnlb.'
1)

paCTCT B .ll:JlllHY ll rnnp1rny; 2) pacTeT B urnpm1y 11 IIIITaCTCH; 3) IIHTaeT-

C.H ll pacTeT B .n;JIHHy; 4) pac.TeT B .ll:JlllHY ll HMeeT 3JIRCTll'IHOCTh.

13. I'pyihwJl KAemKa 'le.ioeeKa no rjJopMe:
1)

He MeHJICTC.H BCIO :>Kll3Hh; 2)

KHH3y;

3)

y

cnopTCMeHOB cmaTa c 6oKOB ll Bh!T.HHyTa

y cnopTCMCHOB cmaTa cnepe.n;H Ha3a.n; ll np116JrnJKeHa K II03BOHO'!Hll-

Ky; 4) o.n;nHaKOBa y cnopTCMeHOB ll HeTpCHHpOBaHHhIX JJIO.n;eiL

14. <Pu6puu - amo:
1)

IIHI'MCHT B 3pllTPOlI.HTax; 2) paCTBOpHMbIH 6eJIOK KPOBII; 3) uepacTBOpll-

MLIH 6eJioK KpOBH; 4) cj_)epMeHT no.n:me.TJy.n;oLIHOH meJie3LI.

508

15. B npo6upKy c 0,9'X.-J11 pacmeopo.w noeape111wil co.w rJof5amu1u 11e.;1111ozo
Kpoeu. C-Jepe.:111ecKo.·1bKO JWUHym u,Bem J/CUOKocmu 6 npo6upKe:
1)

HC l:I3:vIPHl:ITCH;

B Hl:I'.iKHCH -

2)

KpaCHbIM;

CT3HCT a.1br:vr;

3)

B BepxHeH 1 IaCTl:I CTaHeT rrpoapat.JHbIM,

4) B Bepxnerr qaCTll CTaHeT KpaCHbIM, B Hl:I)f\HCH -

npo-

3paqHbIM.

16. Coe.Hecmu.lf!OCmb zpynn Kpoeu npu nepe;1u6a11uu eo.J.lf!OJlflW, ec.r1u rJo11op:

1)

BTOpOH rpymihl, perrHill:ICHT -

t.JCTBCPTOH;

3)

perrI:IrrHeHT -

17.

rrepBoi1;

2) TpCTbeH

qeTBepTOH rpynrrhI, per~HTIHeHT -

rpynnhl, peU,l:ITIHCllT -

TpeTbett;

4)

qeTBCpTOH rpymrhl,

nepBott.

J(8yxcmeop 1tambtil KJtanaH 6 cepiJu,e 'le.wBeica 11axoiJumCR .11teJ1Cay:

1)

2)

npaBhIMH rrpe.ncep.z:i;HeM l:I meJiy.z:i;oqKoM;

roqHhIM CTBOJIOM;

3)

npaBhIM meJiy,noqKoM 1:1 JIC-

.WBb!Ml:I npe.ncep.z:i;HeM l:I JKeJiy)J;Ol.JKOM;

4)

JlCBhl;\11 JKeJiy-

.z:i;oqKOM H .nyroii aopTbI.

18. Y 'le.'to6e1ca om cepiJu,a no 60.1bw.o.;ny 1Cpyzy icpo6oo6paw,euu.R Kp06b
iJeuJ1Cemc.R no:
1)

Jiero1 rnoiI BCHe;

2)

JieroqHOH apTepHH;

3)

JieBOH .nyre aopTbI;

4)

rrpaBOH

.nyre aopTbI.

19.
1)
3)

Kypnwe npu6oaum ic:
cymemno npocseTa apTepMiI;

2)

BOCCTaHOB.1CHHIO KJICTOK ~m.noTemrn;

BOCCTaHOBJ1eHHIO MeM6paH MbIIlleqHbIX K.1eTOK;

.nmpcpy3tt1:1

02

4)

yBeJil:It.JCHl:IIO CKOpOCTH

B aJ1hBeoJiax.

20. 01Co11 'lameJtb1t0e qJOp.11tupo6anue .rny1Co6 6 npou,ecce neuu.R u.rw pa.:1zoeopa y 'le.rweeica npoucxoiJum 6.llazoaap.R:
1)
3)
4)

COKpan~eHI:IIO l'OJIOCOBbIX CBH30K;

2)

paccJia6J1eHHIO l'OJIOCOBbIX CBH30K;

KOJie6aHHHM l'OJIOCOBbIX CBH30K rro.n .nei'ICTBHeM BbI)J;bIXaeMoro B03,llyxa;
H3MeHeHHIO TIOJIOJfWHHH H3hIKa, Hl:I)KHeii qeJIIOCTl:I H MHI'KOI'O He6a.

21. B cno1Coil110.11t cocmo.R1wu 'leJtoaeK aeJtaem BiJox .Ja C'lem:
1) coKpall1eHHH MhIIlll~ .z:i;HacpparMhr; 2) paccJia6.JJemrn Mempe6epHhIX
MbIIllII;

3)

onycKaHI:IH pe6ep no.n co6CTBCHHOll THJKeCTbIO;

4)

paCC.lJafoieHHH

MblllIII )J;HatpparMhl.

22.
1)

llpaeuJtbHbLM .R6JLJlemc.R cyJ1CiJenue:
)Kl:I3HCHHaH eMKOCTb JICI'Kl:IX qeJIOBeKa -

xa IlOCJie rJiy6oKOI'O BbI)J;OXa;
ro B03JJ;yxa;

3)

2)

:3TO 06'heM B)J;bIXae'.VIOI'O B03.llY-

a30T COCTaBJIHCT OCHOBHYIO qacTh aTMOCcpepHO-

npH KCCCOHHOH 6oJie3Hll rry3blpbKH KHCJIOpo.na 3aKynopttBaIOT

MCJIKHe KpOBCHOCHhie cocy.z:i;hr;

4) CTpax,

rHeB ll CHJibHbie :3MOIIMH conpOBOJK)J;a-

IOTCH 3aMe)J;JieHHCM JJ;b!XaTeJibHbIX npoueccos.

23.
1)

CfeM 0.7UHHee ZOJlOC06bte C6Jl3/CU
xyme .llllKIIHH;

n. n

y 'le.rw6e1Ca, meM:

2) Jiyqllle )J;HK1IHH; 3) HlDKe TOH ro.JJoca; 4) Bblllie TOH I'OJIOCa.

24. H.
a6.706 iJ.rn u.Jy'leHUJl <PYHICU,UU nuw,e6apume.llbllOU cucme.'l'lbl
pa.1pa6omaJt .11temoiJbt:
1)
3)

peHTreHorpacpHM ll 30HJJ;HpOBaHHH;

:3Hj\OCKOilllll ll HaJIOJKCHHH cpl:ICTYJihI;

COCTaBa IIHll1CBapHTCJibHbIX COKOB.

4)

2)

30H.L\HPOBaHHH ll :3HJJ;OCKOTIHH;

HaJIOJKCHHH cpHCTYJibI ll H3Y1 IeHHH

Bonpoc1>1

509

25. I for} .J.1ut:11J10 .Jy6a :1eJl(wn:
l)

XprIU\;

2)

h:OC'Th;

26. ()()U(UU .HC('.'l
1) ne•ieHh;

3)

;\CllTIIH;

·1)

lly.1hf1<1.

npomoK '/('Jl(Jlff'h"U O/Jllifibl(-iQ('fllC.H n:
2) Jl(e.lYi\01(; 3) iIO,il;B3j\OlllHYID KUmI<y; 4) J\BCHR,1JUlTHnepemy10
1l//l,/U

El!Jl[J{y.

27. Ilenpaau.ru,H1,1.11 JW.c151emca cy.w:iJe111Le, 'Imo y 1e.wunra:
1

1)

no11.me.1y,n:o•rnhd! coK HMeeT c.1a6ou1e11o•myIO fJcar..:1uuo;

l'KHT (l)CpMeHTbI, paCIIJ;CTlJlHIDIIUW JKHphI;
IIOCTOHHllhre I\

10-12

ro,n:aM;

3)

4) OCHOHHYID

2)

n.:eJl'Ib conep-

"10JIO'!Hbie 3y6h! 3a:'vl\~H5HOTCH Ha

Macey ay6a COCTaBJ15ICT i\eHTlill.

28. !lepm.i.e npu.ma1w wn0Buma ..wu1w.Ja C:
1)

JIOMKOCTb KOCTeii H yxy;unem1e 3pcmrn;

meHHC 11:vr:vrymITeTa;

4)

3)

2)

yxy,TJ,rne1rne 3pemrn n CHH·

CHl:!JKeI\HC H!Vl.MYHIITeTa 11 KpOBOTO'll1BOCTb ACCeH;

KpOBOTOIJHBOCTb ,a;eceH I1 reMOQJHJifUL

29. Henpa6u.?11>11bui J1B.rnemc11 cy.J1CiJeH1u':
1)

1

MOIJa o6pa3yeTcH 113 1wa3MbI KpOBH;

MJIH IIO'Ie'IHhlX KaHa.'Ibl\CB;

2)

B Kam,n:oil: no•rne

y

<JeJJOBeKa

3) MO'!eTO'IHHI\ OTI{phlBaeTCH B MOlJCHCIIyc1rn-

Te.1hHhlH KaHa.il; ,1) IlO'!e'Il!aH .'IOXaliKR COOHptie'I' BTOpH'IHYlO '.\10'IY.

30 . .Y

'l<'JW6PKa npou,ecc .MO'leucnyc1rnHuJ1

1wxoaumc;1

norJ 1wHmpo.?1e.'n

;110.:1ia:

1)

Kotteirnoro;

2) cpc,n:m~ro; 3)

npo,n:oo1ronaToro;

4)

11poMemyTo•moro.

Tecmb1

510

OTBETbl
061.u;aH 6ttO.'IOrHH

A. ll1lT0:10nrn
A 1. 2; A 2. 1; A 3. 4; A 4. 2; A 5. 1; A 6. 3; A 7. 4; A 8. 3; A 9. 2; A 10. 4.
B. rcneTHKa
B 1. 4; B 2. 1; B 3. 4; B 4. 3; B 5. 3; B 6. 3; B 7. 4; B 8. 2; B 9. 5; B 10. 3; B 11. 5.
f. 8BOjlI<)[(llOHHOe yt1en:u:e
r 1. 4; r 2. 2; r 3. 4; r 4. 3; r 5. 1; r 6. 4; r 7. 2; r 8. 3; r 9. 4; r 10. 3; r 11. 3.

,n. AttTporrorcne3
,II, 1. 1; ,II, 2. 2; ,II, 3. 1; ,II, 4. 4; ,II, 5. 2; ,II, 6. 1; ,II, 7. 2; ,II; 8. 4; 11; 9. 1.
JK.

3KOJIOrMH

JK 1. 4;
JK 10. 2.

jl{

2. 3; JK 3. 2; JK 4. 1; JK 5. 3; JK 6. 3; JK 7. 4; JK 8. 1; JK 9. 2;

1. 3; 2. 2; 3. 1; 4. 2; 5. 2; 6. 2; 7. 2; 8. 2; 9. 3; 10. 1; 11. 2; 12. 3; rn. 2; 14. 2;
15. 1; 16. 3; 17. 3; 18. 3; 19. 3; 20. 2; 21. 3; 22. 4; 23. 2; 24. 1; 25. 3; 26. 3;
27.3;28.3;29.4;30.2.
3ooJiorttn

1.4;2.1;3.2;4.4;5.3;6.4;7.2;8.4;9.3; 10.2; 11.3; 12.2; 13.3; 14.4;
15. 4; 16. l; 17. 4; 18. 1; 19. 3; 20. 3; 21. l; 22. 2; 23. 3; 24. 4; 25. 1; 26. 1;
27. 1;28.3;29.4;30.3;31.2;32. 1.
AuaTOMHH
1. 2; 2. 3; 3. 3; 4. 2; 5. 2; 6. 2; 7. 3; 8. 3; 9. 1; 10. 1; 11. 3; 12. 2; 13. 3; 14. 2;
15. 1; 16. 2; 17. 3; 18. 3; 19. l; 20. 4; 21. 1; 22. 2; 23. 3; 24. 4; 25. 3;
26.4;27.2;28.3;29.3;30.1.

1113AATEnbCTBO ccMAXAOH» PEKOMEHAYET:
CepHSI cc.QnSI WKOnbHHKOB H a6HTypHeHTOB>> - Kmo~
KycnexyHa3K3aMeHax

Jlyl.fw1.1e cnpaeol.fHl1Kl1 1.1 noco61.1S1 aeTop1.1TeTHel:iw1.1x cne'-'1.1an1.1cToe

,IJ,.3 . Po3eHranb, /11.5. rony6
PycCKlltH Sl3b1K Ha OTnllt'-tHO. npas111na 111 ynpa>1<HeH111s:1
,IJ,.3. Po3eHranb, /11.5.rony6
PyccKMM Sl3blK. Cnpaeo1.tH11tK WKOnbH11tKa. Op¢orpa¢111s:1 111 nyHKTyau,111s:1
,IJ,.3.Po3eHranb,/11.5. rony6
PyCCKlltH Sl3blK. roTOBlltMCSI 6e3 peneTlltTOpa. Ynpa>1<HeH111s:1111 KOMMeHrnp~
,IJ,.3.Po3eHranb, /11.5. rony6
PyccK11tM Sl3blK. Co1.t1.1HeH11tS1 11t 3K3aMeHbl Ha OTnlltl.fHO. Crnn111crnKa 111 Kynbryp.

/11.5 . KO>i<YXOB, A.A. npoKOcjJbeB
MaTeMaT11tKa. nonHblH cnpaeo1.tH11tK

A.A. npoKOcjJbeB, /11. 5. Ko>KyXOB
MaTeMaT11tKa. roTOBlltMCSI 6e3 peneTlltTOpa. 3aAa4111111 peweHll1s:!

B.A. ryceB, /11. 5. Ko>KyXOB, A.A. npoKOcjJbeB
reoMeTp11tsi. nonHblH cnpaso1.tH11tK

E.E. BH3eMCKMlli, !7.B. >KyKOaa, B.A. WecraKoB
~cTop1.1si

Pocc1.11.1 c ApesHeMwMx epeMeH AO Haw1.1x AHeM

C. C. 5ep11.0Hocoa, E.A. MeH11.eneeaa
X11tM11tS1. HoeeMWlltM cnpaeol.fHlltK