Карлсон Б. Основы эмбриологии по Пэттену

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PATTEN'S

FOUNDATIONS

OF EMBRYOLOGY

Bruce M. CARLSON, M.

D.,

Ph. D.

Department of Anatomy

and Biological Sciences

University of Michigan

New York St. Louis San Francisoo Auckland Bogotii

Hamburg Johannesburg London Madrid Mexico Montreal

New Delhi Panama Paris Sao Paulo Singapore Sydney

Tokyo Toronto

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,UonOJIHHTeJIbHbIe OnbITbI rrOKaSaJIH, qTO rrOCJIe,lI.HeM:,lI.eTepMHHH-

pyeTcSI 11-,U OCb KOHeIJHOCTH.TaKHM oopaSOM, ObIJIa BbISIBJIeHa rrOCJIe-

,lI.OBaTeJIbHOCTb,lI.eTepMHHHpOBaHHSIoceM KOHeIJHOCTH:11-3-+,U-B-+

-+11-,U

[34].

Eha nOCJIe,lI.OBaTeJIbHOCTb,rrO-BH,lI.HMoMy, OTpa}KaeT 00-

w.YIO saKoHoMepHocTb ,lI.eTepMHHHpoBaHHSI OceM: OCHOBHbIX BHeWHHX

opraHoB, rrOCKOJIbKY aHaJIOrHIJHaSI rrOCJIe,lI.OBaTeJIbHOCTb,lI.eTepMHHH-

PYIOW.HXCOObITHMxapaKTepHa ,lI.JISIrrpOII,eCCOBpasBHTHSI yxa H CeTqaT-

KH rJIasa.

POCT rrOIJKH KOHeqHOCTH. KaK TOJIbKO SaIJaTOK KOHeIJHOCTHHaqH-

HaeT BOSBbIwaTbCSI Ha,lI. rrOBepXHOCTbIOSapO,lI.bIWa, ero MO}KHO C rrOJI-

HbIM OCHOBaHHeM HaSbIBaTb

nOtf,TWU TWfletf,flOCTU.

UeHTpaJIbHaSI qaCTb

rrOIJIUI KOHeIJHOCTHoopaSOBaHa MaCCOM O,lI.HOP0,lI.HbIX MeseHXHMHblX

KJIeTOK, rrpOHHsaHHbIX ceTblO KpOBeHOCHbIX KarrHJIJISIpOB. B paHHIOIC

rrOIJKY KOHeIJHOCTHHepBHbIe BOJIOIma ew.e He BpOCJIH. MeSO,lI.epMaJIb-

HaSI qaCTb rrOIJKH KOHeIJHOCTHcBepxy rrOKpbITa 3KTO,lI.epMOM,KOTopaSI

y OOJIbWHHCTBa rrOSBOHOIJHbIX }KHBOTHbIX yTOJIw.aeTCSI K ,BepWHHe

rrOqKH.

1948

r. CayH,lI.epC rrpOBeJI OJIeCTSIw.He HCCJIe,lI.OBaHHSI,KOTopbIe

HHHII,HHpOBaJIH MHO}KeCTBO3KcrrepHMeHToB, rrocBSIw.eHHbIX MexaHHSMy

pasBHTHSI KOHeIJHOCTH.Ero HHTepeCOBaJIa POJIb yTOJIw.eHHOM:rrOJIOCKH

3KTO,lI.epMbI-

anutWJlbflOeO 3KTOaePMaJlbflOeO epe6flR.,

pacrrOJIO}KeH-

HOro Ha BepwHHe BHeWHero KpaSI rrOqKH KpbIJIa KypHHoro Sap0,lI.bIwa.

CayH,lI.epC rrOKaSaJI, IJTO eCJIH y,lI.aJIHTb arrHKaJIbHbIM 3KTO,lI.epMaJIbHbIM

rpeoeHb, TO BCKope rrOCJIe 3Toro POCT KOHeqHOCTH rrpeKpaw.aeTCSI

(pHC.

10.5).

TaK ObIJIO,lI.OKaSaHO,IJTO B rrpoII,ecce pOCTa rrOIJKH KpblJIa

PHC,

10.5.

BJlHllHHe Y.l\aJleHHll anHKaJlbHOrO 3KTO.l\epMaJlbHOrO rpe6Hll Ha pasBHTHe nO'lKH

KpblJla.

CxeMa paCnOJlOlKeHHll nO'lKH KpblJla.

nO'lKa KpblJla H anHKaJlbHblA 3KTo.l\epMaJlbHblll rpe:

6 pHHoro sap0.l\blllIa

He.l\OCTaTO'lHOCTb CKeJleTa. BOSHHKalOlliall B pe

S~~~T;~eex;;;~:~~~ ~~HKaJlbHOrO 3KTO.l\e·pM~JlbHOrO rpe6Hll Y Tpex.l\HeBHOro KypHHoro sap0.l\bl-

(r)

/l,

He.l\OCTaTO'lHOCTb CKeJleTa. BOSHHKalOlliall B pesYJlbTaTe Y.l\aJleHHll anHKaJlbHOrO 3K-

TO.l\epM~Jlblioro rpe6Hll Y '1eTblpex.l\HeBlIOro KyplIHoro sap0.l\bIllia

(E) (B -

/l,

[24]).

Hepas-

~~~~~~~ '1~~:~JI~~O~lI~'1';~;~'::YHr~:~~~~~:JlHb~~:· 3KTO.l\epMaJlblIb1A rpe6eHb; 4 -6 cJloA KJleTOK

MaJIb~aH

qaCTb nOtIKH

KpbtJIa;

6 -

KpaeBoA

CHHyC;

7 -

aaaJIbHaH

nJIa~

~;~:::PM8b~ K';'T~~SO;;~JleIllIOrO anHKaJlblIOrO 3KTo.l\epMaJlbHOrO rpe6I1n; 9

i'i

nJle6'eBan KO~~~;

10 _

JI~naTKa;

11-

JlOKTeBall KOCTb;

12-

JlYQeBa5I KOCTb;

13 -

KopaKOH.l\;. - 0 JlaCTb M -

Kapnyca;

15 -

q,aJlaHrH naJlbueB.

PRC. 10.6. MHKpo<!JoTorpa<!JHHnO'leK KOHe

TpOHHoro MHKpocKona. 'IHOCTH,nOJlY'leHHble C nOMOXUblOcKaHHPYlOxuero

9neK-

I(YPHHblll sapoAblw Ha CTaAHH 26 (c

~gefa4Hl7alCTaAHH 15 (no [17]).

AKCOJlOT1i,a~~e~:::~ ~1.a (I¥o>KTa~kJl:t~f'>' 1~773

blW

'1eJlo-

• . ' •. xp,

Zool..

06paTHTe BHHMaHHeHa OTCYTcTBHeanHKa

CTH8KCOJlOTJlH. JlbHOrO 3KTOAepMaJlbHOrOrpe6HH Ha nOqKe

KORe'lBO-

PRC. 10.7. CxeMa, HJlJlIOCTpHpYlOmaHH3MeHeHHHcTpoeHHH anHKaJlbHoro 3KTOAepMaJlbHOrOrpe6-

HH nO'lKH KOHe'lHOCTHsapOAblwa 'IeJlOBeKa (no [16]).

CTaAHli 12.

CTaAHH 13-14.

CTaAIIH 15.

CTaAHH 17.

J{.

CTaAHli 19. Ha CTaAIIH 12 6a-

~aJlbHali nJl8CTHHKa 06pasoBaHa ABYMll CJlOllMH, coeAHHeHHblMH nOnepe'lHbIMH CBllSllMH. Ha

OJleen03AHHx CTaAHHXpasBHTHHOHa npeBpamaeTC51B OAHOCJlollHYIOCTpYKTYPY.

~ - MeseHxHMa nO'lKH KOHe'lHOCTH;2 _ 6asaJlbHB51nJlaCTHHKa; 3 - 3KToAepMa; 4 - AecMocoMbl;

.n;;;:;~~~~Bble KOHTaKTbI; 6 - AeJl51mHeC51KJleTKH; 7 - AereHepHpYlomali .KJleTKa;

MHKPOq,H-

Me)J{)J.yanHKaJIbHblM 3K 0

JJ.epMOH npOHCXO)J.HT T' )J.epMaJIbHblM rpe6HeM H- n0)J.J1e)J{ameHMe3

BHH. onpe)J.eJIeHHble H BeCbMa Ba)J{Hble BSaHMO)J.eHC~=

AnHKaJIbHblH 3KTO)J.epMaJIbHbIHrpeOeHb.

CTpOeHHe anHKaJIbHOro 3KTO e

H H. JJ.JIH nOlIeK KOHelIHOCTeH3a 0 b u)J. P M a JI b H 0 r 0 r p e

)J{HBOTHbIX xapaKTepHbl TepMHHa~b~b~~eH OOJIblI1HHCTBanOSBOHO'IHbIX

lIae TaKoe yTOJImeHHe HMeeT

cpo

M yTOJImeHHH.

THnHlIHOM CJIy-

pe)J.He-sa)J.Hero KpaH nOlIKH BHe p y rpe6HH, THHymerocH B)J.OJIbne~

lI1eH nOSBOHO'IH~X y)J.HB~TeJIbH~He anHKaJIbHNe rpeOHH y sapo)J.~~

cpesax nO'IKH KOHe'IHOCTH sa O)J.bIlI1~~HOOOpasHbl. Ha nonepelIHblX

HM~lOT BH)J.COCOlIKOB(pHC.

16.5

10~)

llTYII. H MJIeKOnHTalOmHx OHlf;

JIHH anHKaJIbHblH 3nH)J.epMHc saKpYlIe'H' Pbl0 H HeKoTop~x penTH-

)J.aHHoe OTKJIOHeHae B MOPCPOJI B nJIOTHylO CKJIa)J.Ky. Heo)J{H-

)J.aeTCH y aMcpHoHH.

oecxBoc~~~ :~To)J.ep;vrbl KOHelIHoCTH Ha6JIlO~

MaJIbH~H rpeoeHb CXO)J.eHC rpeoH <pHOHH anHKaJIbHhIH 3KTo)J.ep-

mHX. O)J.HaKO y XBOCTaTblX aM;M

S~PO)J.bllI1eHnTHII. H MJIeKOnHTalO-

nOlIKe KOHelIHOCTHOTCyTcTByeT (PH~

~)H~JIbHOe yTOJImeHHe Ha,

y sapO~~lI1eH 3TOH cpaBHHTeJIbHO ~O . , u· onpOCbl 0 TOM, nOlIeMY

anHKaJIbHOro rpeOHH H KaKHM 00 3)J.HeH rpynn~ )J{HBOTHhIX HeT

HOCTH,OCTalOTCHnOKa oes OTBeTa.pa30M oes Hero pa3BHBalOTCH KOHe'lI-

<l>OPMHpoBaHHe anHKaJIbHOro 3KTO

cpa3y )J{e nOCJIe nOHBJIeHHH nOlIKH KO)J.epMaJIbHOro rpeOHH HalIHHaeTCfll

oeHb opHeHTHpOBaH B)J.OJIbnepe)J.He_3=elIH~CTH. 3KT0)J.epMaJIbH~H rpe~

npe- K nOCTaKCHaJIbHOHlIaCTH H He )J.HeHOCHnOlIKH KOHelIHOCTH01"

HanpHMep, y 3ap0)J.bllI1eH nTH OH CKOJIbKO aCHMMeTpH'IeH no cpopMe.

nOlIKH KOHelIHOCTH. OObIlIHO irrHKaYTOJIn;eH B nOCTaKCHaJIbHOHlIaCTIr

)J.OCTHraeTCBoero MaKCHMaJIbHOro ~bHb1H 3KTO)J.epMaJIbHhIH rpeOeHb.

KOHelIHOCTH,nOCJIe lIero C HalIaJIO p 3BHTHH K CTa)J.HHJIOnaTKOBH)J.HOK

HaeTCH ero perpeCCHH. M )J.HCPCPepeHII.HpOBKHnaJIbII.eB HalIH-

BblClI1HX n03BOHOlIRbIX )J{

JIOrHlIeCKOH opraHH3aII.HH anHBOTH~X pa3JIHlIalOT )J.Ba THna rHCTO-

3ap0)J.bllI1eH nmII. rpeoeHb 00 ~~~~bHoro 3KTO)J.epMaJIbHOro rpeOHH.

pHlIeCKHM 3nHTeJI1:leM (pHC

165

nceB)J.OMHOrOCJIOHHbIMII.HJIHH)J.-

mHX - MHOrOCJIOHHbIMKyo~lIec~' ,a y 3apO)J.bllI1eH MJIeKOnHTalO_

H <l>aJIJIOH

16]

)J.eTaJIbHO on HM HJIH nJIOCKHM 3nHTeJIHeM. KeJIJIIf

MaJIbHoro rpeoHH B ne HO e HCaJIH cTpoeHHe anHKaJIbHOro 3KTo)J.e _.

treJIOBeKa (pHC.

10.7).

PO)J.~O:~ CTaHOBJIeHHHH perpeccHH y 3apO)J.bluia

anHKaJIbHOro 3KTO;QepMaJIbHOro

rJI~BHbIX CTpyKTypHbIX ocooeHHOCTeH

lIHCJIa meJIeBblX KOHTaKTOB ( rpe HH HBJIHeTCH HaJIHlIHe OOJIb1I10ro>

[12],

lITO nocpe)J.CTBOM TaKH/:~~

1.10).

3TH aBTopbI npe)J.nOJIO)J{HJIII

CKaH H MeTaOOJIHlIeCKaH CBH3b Me~KTOB ocymeCTBJIHeTCH 3JIeKTpHlIe-

XHMHlIecKHX HCCJIe)J.OBaHHH

19]

)J.y KJIeTKaMH. Pe3YJIbTaTbl rHCTO-

MaJIbHOro rpeoHH cO)J.ep)J{aT 00 nOKa3bIBalOT, lITO KJIeTKH 3KTO)J.ep-

a TaK)J{e meJIOlIHOH H KHC

JIb1I10e KOJIHtrecTBO rJIHKOreHa

PHK

J1.JIHaKTHBHO pacTyr:u;HX H

~~~:pocCPaTa3.

3TH CBOHCTBa xapa~TepH~

.'¥.epeHII.HpylOmHxCH KJIeTOK 3apO)1.bllI1a.

B 0 H CT B a a rr HK a JI b H0 r 0 3 K T 0 )J.e p M a JI b H 0 r 0 r p e

II 51.XOTH Borrpoc 0 pOJIH arrHKaJIbHOrO 3KTO)J.epMaJIbHOrO rpeOHH B

t>lOpcporeHese KOHelIHOCTH OCTaeTCH )J.HCKyCCHOHHbIM, OOJIblI1HHCTBO

f1CCJIe)J.OBaTeJIeHClIHTalOT, lITO 3Ta CTpyKTypa CJIy)J{HT CTHMyJIHTOpOM

pOCTa nOlIKH KOHelIHOCTH.B COOTBeTCTBHHC )J.pyrHMH npe)J.nOJIO)J{eH~H·

Mfl arrHKaJIbHbIH 3KTO)J.epMaJIbHbIHrpe6eHb BbIITOJIHHeTperyJIHpylOmylO

f1JIfi MexaHHlIeCKYIO CPYHKII.HIOB npOII.eCce POCTa H MopcporeHesa KOHelI·

HOCTH

[1].

PH)J. 3KcrrepHMeHTaJIbHblX )J.aHHbIX cBH)J.eTeJIbCTByeTB

nOJIbSY rrepBoH TOlIKH 3peHHH. TaK, BCKope nOCJIe y)J.aJIeHHH anHKaJIb-

Horo 3KTo)J.epMaJIbHOrO rpeoHH POCTrrOlIKH KOHelIHOCTHnpeKpamaeTcH

[24].

Pa3BHTHe 6eCKpbIJIblX reHeTHlIeCKHX MyTaHToB Kyp

[36]

CJIy)J{HT

npHpO)J.HblM 3KcnepHMeHToM, TaK)J{e nO)J.TBep)J{)J.alOmHMnepBylO TOlIKY

3peHHH.

3THX MyTaHTHbIX sapo)J.bIlI1eH (wingless) paHHee pa3BHTHe

KpblJIbeB npoHcxo)J.HT HOpMaJIbHO, O)J.HaKO BCKope OCTaHaBJIHBaeTCH,

lITO CBHsaHO C perpeCCHeH H HClIe3HOBeHHeM anHKaJIbHOrO 3KTO)J.ep-

MaJIbHOrO rpeOHH. BMeCTe C TeM )J.onOJIHHTeJIbHble anHKaJIbHble 3KTO-

,lI.epMaJIbHbIe rpeOHH, nO)J.ca)J{eHHble B 3KcnepHMeHTe HJIH eCTeCTBeHHO

B03HHKalOmHe npH paSBHTHH y MyTaHTHoH JIHHHH Kyp (eudiplopodia).

BbISbIBalOT paSBHTHe )J.onOJIHHTeJIbHbIXCTpyKTyp KpbIJIa.

KaKOBa ObI HH ObIJIa HCTHHHaH POJIb anHKaJIbHOrO 3KTO,ll.epMaJIbHO-

ro rpeOHH B npOII.eCce paSBHTHH KOHelIHOCTH,ero BJIHHHHenpOHBJIHeTCH

llopa3HTeJIbHO HeCneII.HcpHlIHO. MeTO)J.OM rrepeca)J.OK nOKasaHO, lITO

anHKaJIbHblH 3KTO)J.epMaJIbHbIH rpeoeHb KypHHoro 3apo)J.bIlI1a cnocoo-

cTByeT pOCTy Me30,ll.epMbI KOHelIHOCTH 3apO)J.bIlI1eH HHbIX BH)J.OBnTHII.

H )J.a)J{e Me30)J.epMbI KOHelIHOCTH 3apo)J.bIlI1a MbIlI1H. BepOHTHee Bcero,

lITO 3KTO)J.epMaJIbHbIH rpeoeHb HeCrreII.HcpHlIeCKHCTHMyJIHpyeT POCT

KOHelIHOCTHH He onpe)J.eJIHeT nOCJIe)J.OBaTeJIbHOCTbMopcporeHe3a B)J.OJIb

ee npOKCHMO)J.HCTaJIbHOHOCH. PyOHH H CayH)J.epc

1[22]

oOHapy)J{HJIH,

lITO npH nepeca)J.Ke 3KTO)J.epMaJIbHOrO rpeOHH nOlIKH HorH KypHHoro

sapo)J.bIlI1a. Ha nOlIKY HorH KypHHoro sapO)J.bIIIIa, HaXO)J.HmerOCHHa

npe)J.IIIecTBylOmeH CTa)J.HHpasBHTHH, npo)J.OJI)J{aeTCHHOpMaJIbHOe pas-

BHTHe, HeCMOTpH Ha paSJIll-lIHH B BospaCTe H aHaTOMHlIeCKOM nOJIo)J{e-

HHH (<<CTeneHH)J.HCTaJIbHOCTH»)B)J.OJIbnpOKCHMO)J.HCTaJIbHOHOCH KO-

HelIHOCTHnepeca)J{eHHoro rpeOHH H nO)J.JIe)J{ameHMeSO)J.epMbI.110 Mepe

perpeCCHH anHKaJIbHbIH 3KTO)J.epMaJIbHbIH rpeoeHb yTpalIHBaeT cno-

COOHOCTbyCKOpHTb POCT nOlIKH KOHelIHOCTH. BbIBepHyTbIH 3KTO)J.ep-

MaJIbHbIH rpeoeHb (T. e. nO)J.ca)J{eHHbIHTaK, lITOObI HCXO)J.HO

HapyLK:-

Hble KJIeTKH 6bIJIH oopameHbI BHyTpb, a BHyTpeHHHe COOTBeTCTBeHHO

Hapy)J{y) H )J.HCCOII.HHpOBaHHbIHMaTepHaJI 3KTO)J.epMaJIbHOrO rpeOHH

MoryT peopraHHSOBaTbCH H npO)J.OJI)J{aTbOKaSbIBaTb OObIlIHoe )J.eHCT-

BHe Ha paSBHBaIOmYIOCH nOlIKY KOHelIHOCTH. 110JIylIeH pH)J. KOCBeHHbIX

)J.aHllbIX, CBH)J.eTeJIbCTBylomHX0 TOM, lITO KJIeTKH anHKaJIbHOrO 3KTO-

)J.epMaJIbHOrO rpeOHH BbI)J.eJIHIOTB nO)J.JIe)J{amyIO Me30)J.epMy onpe)J.e-

JIeHHbIe )J.HCP<pYH)J.HpYIOmHeBemeCTBa. O,ll.HaKO, npe)J{)J.e lIeM npHSHaTb

STy CPYHKII.HIOanHKaJIbHOrO 3KTO)J.epMaJIbHOrO rpeOHH, HeOOXO)J.HMbI

)J,onOJIHHTeJIbHbIeHCCJIe)J.OBaHH51.

Me30,ll.epMa p,aHHeH nO,lIKH KOHellHOCTH.-

C T pO e H H e. MeSO)J,epMa paHHeH nOllKH K0HellHOCTH: 06pa30BaH~

MeseHXHMHbIMH KJIeTKaMH, SaKJIIOlleHHbIMH -B PbIXJIbIH MaTpHKC H3

KOJIJIareHOBbIX BOJIOKOH H MyKonOJIHCaxapH)J,OB (rJIHKOSaMHHOrJIHKa-

HOB). KJIeTKH BblrJI5I)J,5ITHe)J,HepepepeHu.HpOBaHHbIMH,HMeIOT KpynHble

5I)J,pa C BbIpa:>KeHHblMH 5I)J,pbIIIIKaMH, u.HTOnJIaSMa HX CJIaOO 6a30<pHJIb~

Ha. MHTOTHlIeCKa5I aKTHBHOCTb 9THX KJIeTOK BbICOKa5I BO Bcex 30Hax

nOllKH KOHellHOCTH. Henocpe)J,CTBeHHO no)J, anHKaJIbHhIM 9KTO,ll.epMaJIb~

HbIM rpeOHeM HaXO)J,HTC5ISOHa TOJIIII.HHOHB HeCKOJIbKOC,?TeH MHKpOH

C OOJIbIIIHM KOJIHlIeCTBOMOCHOBHoro BeIII.eCTBa, B KOTOpOH OTMellaeTC5I

BbICOKHH MHTOTHlIeCKHH HH)J,eKC.2:ha OOJIaCTb, HaSBaHHa5I

SOliOU npo.-

epecca

[33],

B03MO:>KHO,HrpaeT Ba:>KHyIOPOJIb B pOCTe H MopeporeHese

KOHellHOCTH. [lollKa KOHellHOCTH OOHJIbHO CHaO:>KeHa KpOBeHOCHbIMH

cocy)J,aMH, a no,ll. anHKaJIbHbIM 9KTO)J,epMaJIbHblM rpe6HeM pacnOJIO:>KeH

)J,OBOJIbHOKpynHbIH KpaeBOH CHHyC. PaHH5I5I nOllKa KOHellHOCTH He

HHHepBHpOBaHa, HO no Mepe paSBHTH5I B Hee BpaCTaIOT IHepBHble BO-

JIOKHa. .LLHepepepeHu.HpOBKaJMeSO)J,epMbI 06cy:>K)J,aeTC5I B 3TOH rJIaBe

nOS)J,Hee.

Po JIb Me s O)J,e p M bI. B npou.ecce- paSBHTH5I rrpOHeXO)J,HTHe O)J,HO-

CTopOHHee, a BsaIIMHoe BJIII5IHHe 3KTO)J,epMbI H MeSO)J,epMbI, 06pasyIO-

III.IIX nOllKY KOHellHOCTII. P5I)J, )J,aHHblx cBH)J,eTeJIbCTByeT 0 TOM, liTO

MeSO)J,epMa paHHeH nOllKH KOHellHOCTH CTIIMyJIHpyeT 3KTO)J,epMaJIbHble

KJIeTKH K oopaSOBaHHIO anHKaJIbHOrO 3KTO)J,epMaJIbHOrO rpeOH5I, a

nOCJIe ero BOSHIIKHOBeHH5IBJIH5IHHe MeSO)J,epMbIHeOOXO)J,IIMO)J,JI5Ino,n;-

)J,ep:>KaHH5Iu.eJIOCTHOCTH3TOH CTpyKTypbI.

I1cCJIe)J,OBaHH5IHa «oeCKpbIJIbIX» H «MHorOnaJlbIX» MyTaHTHbIX

KypHHbIX sapo,n;bIIllax nOKaSaJIH Ba:>KHyIOPOJIb MeSO)J,epMbI.

oeCKpbI-

JIbIX MyTaHToB pafIHee paSBHTHe nOllKH KpbIJIa npOHCXO)1:HTHOpMaJIb-

HO, HO BCKope anHKaJIbHbIH 9KTO)J,epMaJIbHbIH rpeoeHb )J,ereHepHpyeT

H )J,aJIbHeHIIIIIH POCT nOllKH KpbIJIa npeKpaIII.aeTC5I. B OnbITaX C peKOM-

oHHau.H5IMH TKaHeBbIX SaKJIa,n;OKObIJIO nOKasaHo, liTO eCJIH MeSO,ll.epMy

nOllKH KOHellHOCTH «oeCKpbIJIOro» sapo)J,bIIlla nOKpbITb 9KTO)J,epMOH

nOllKII KOHellHOCTII HOpMaJIbHOrO sapo,n;bIIlla, TO «HopMaJIbHbIH» ana-

KaJIbHbIH 3KTO)J,epMaJIbHbIH rpeoeHb )J,ereHepIIpyeT II paSBIITHe KpbIJIa

OCTaHaBJIIIBaeTC5I

'[37].

CXO)J,HbIM oopasoM ObIJIO nOKasaHO, liTO eCJIa

9KTO)J,epMy KOHellHOCTH TpaHCrrJIaHTIIpCmaTb Ha)J,Meso,n;epMOH, He Ylla-

cTByIOIII.eH B oopaSOBaHIIII KOHellHOCTH, TO 9KTO)J,epMaJIbHbIH rpeoeHb

TaK:>Ke)J,ereHepIIpyeT. B CJIyllae rrOJIII)J,aKTIIJIIIII rrpH KOMoHHau.HII HOP-

MaJIbHOH Meso)J,epMbI C .9KTo)J,epMoHsapo)J,bIIlleH-MyTaHToB Ha6JIIO)J,aeT-

C5I paSBIITHe HOpMaJIbHbIX KOHellHocTeH, Tor)J,a KaK npH COlleTaHFlH

MyTaHTHoH Meso)J,epMbI C «HopMaJIbHoH» 3KTo)J,epMoH rrOllKH KOHellHO-

CTH BOSHIIKae-r.MHOrOrraJIa5I KOHellHOCTb. 9TH orrbITbI CBII)J,eTeJIbCTBYIOT

o TOM, liTO MeSO)J,epMa CTIIMyJIIIpyeT .9KTO)J,epMyrrOllKH KOHellHOCTH K

oopaSOBaHIIIO anIIKaJIbHOrO 9KTO)J,epMaJIbHOrO rpeOH5I H no,n,nep:>KH-

BaeT ero B aKTHBHOM COCT05IHIIH; B CBOIO Ollepe,nb arrIIKaJIbHbIR 9KTO-

,nepMaJIbHbIH rpeoeHb CTIIMyJIHpyeT ,naJIbHeHIIIIIR POCT Meso,nepMbl

nOllKH KOHellHOCTH. UBIIJIJIHHr

{37]

IJpe,nrrOJIO:>KHJI,liTO BJIH5IHHe MeSO,.

.zr.epMbl ocyIII.eCTBJI5IeTC5I nocpe,nCTBOM MeSO)J,epMaJIbHOrO «nOMep:>KH-

BaIOIII.erO epaKTopa», o,nHaKO BbI,neJIHTb H H,neHTHepHu.HpOBa.Tb.9TOT

epaKTOP,no CHX rrop He y)J,aJIOCb.

MHorHe )J,aHHbIe cBII,neTeJIbCTBYIOT 0 TOM, liTO MopeporeHes KOHell-

HoCTH onpe)J,eJI5IeTC5ICKopee Meso,nepMOH, a He 3KTo,nepMOH rrOllKH KO-

HellHOCTH. ECJIH, HanpIIMep, Meso,nepMy rrOllKH KpbIJIa sapo,nbIIlla

coe,nHHHTb C .9KTo,nepMoH nOllKH HorH, TO paSBHBaeTC5I KpbIJIO, rrOKpbI-

Toe rrepb5IMH. B pesyJIbTaTe COlleTaHH5I ,nIICTaJIbHOH Meso,nepMbI HorR

C .9KTo,nepMo'H KpbIJIa oopasyeTC5I Hora, rrOKpbITa5I porOBbIMH rrJIaCTHH-

KaMH. BsaIIMHoe BJIH5IHHe Meso,n;epMbI H .9KTo,nepMbI rrOllKH KOHellHO-

CTeH HaOJIIO,naeTC5I,na:>KeB TOM CJIyllae, Kor)J,a ooa KOMrrOHeHTa rrpII-

Ha,nJIe:>KaTpaSJIHlIHbIM BH)J,aM. COlleTaHIIe Meso,n;epMbI rrOllKH KOHellHO-

CTH yTIIHoro sapo,nbIIlla C .9KTo,nepMoH KypIIHoro sapo,nbIIlla npHBo,nHT

K paSBHTHIO rreperrOHlIaTOH KOHellHOCTH, a HS Meso,nepMbI paHHeR nOll-

KH KOHellHOCTH sapo,nbIIlla MbIIllII, rrpIIBe)J,eHHOH B KOHTaKT C .9KTo,nep-

MOH rrOllKH KpbIJIa KypHHoro sap O)J,bIIIIa, pasBHBaeTC5I KOHellHOCTb c:

naJIbu.aMH

'[3].

fHOeJIb KJIeTOK B pa3BHBaIOIIJ.eHC5I KOHellHOCTH. Cpe,nH MHorHX

rrpou.eccoB, npoIIcxo)J,5IIII.HX BO BpeM5I paSBHTII5I KOHellHOCTH, HaHOOJIee

saMeTHbIM 5IBJI5IeTC5IrHOeJIb KJIeTOK (CM.

[36]

B cnIICKe JIHTepaTypbl

K rJI.

1).

,9TOT rrpou.ecc paHbIlle Bcero HaOJIIO,n;aeTC5IHa rrepe,nHHX KO-

HellHOCT5I: B OOJIaCTH oy,n;YIII.eH rro,nKpbIJIbU.OBOH5IMKII, rros,nHee SOHbl

KJIeTOllHOH rHOeJIH BOSHHKaIOT B OOJIaCTII JIOKTeBOro cycTaBa H Me:>K,ny

paSBHBaIOIII.IIMHC5I naJIbu.aMH. ,UJI5I paSBHTH5I sa,nHHX KOHellHOCTeH xa-

paKTepHa TaKa5I :>KenOCJIe)J,OBaTeJIbHOCTbrr05IBJIeHH5ISOHHeKposa KJIe-

TOK(pHC.

10.8). Y

KypHHoro sapo,nbIIlla HaIIOOJIee XOpOIIIO Bblpa}KeHa

oOJIaCTb MaCCOBOHrIIOeJIH KJIeTOK Ha MeCTe oy,n;YIII.eH no,nKpbIJIbU.OBOH

5IMKH; .9Ty OOJIaCTb HaSbIBaIOT

saolieu lieKpOTUl1.eC1WU 30liOU.

BOJIee

orpaHHlIeI;!Hble OOJIaCTH KJIeTOIIHOH ruoeJIII HaOJIIo,naIOTC5I npII pasBH-

THH CKeJIeTa, B lIaCTHOCTH cycTaBoB, H rHCToreHese onpe)J,eJIeHHbIX

rpyrrrr MbIIIIU..

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rrepBoM naJIbu.e B OTJIHlIHe OT OCTaJIbHbIX C TpeM5I epaJIaHraMII, nO-BH-

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u.eBOH IJaCTH SaIJaTKa 9Toro naJIbu.a

[19].

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KOHe1JHOCTH,6y,n;b TO HOpMaJIbHOe 5IBJIeHHe y HeKOTopbIX BH,nOB (Ha-

rrpHMep, y yTOK) HJIH pesyJIbTaT aHOMaJIpHOrO rrpouecca

(SU200aK-

TUJ/.UJl),

paSBHTHe neperrOHOK Me:>K,nynaJIbu.aMH OO'b5ICH5IIOTCHII:>KeHH-

eM HHTeHCHBHOCTHrrpou.eCCOB rIIOeJIII KJIeTOK B Me:>KrraJIbu.eBbIXyIJaCT-

Kax.

Bblpa:>KeHHble SOHbI KJIeTOIJHOHrHOeJIH, B IJaCTHOCTII B Me:>KrraJIb-

u.eBhIX y1JaCTKaX, npOCJIe:>KHBaIOTC5IB npolJ,ecce paSBHTH5I Bcex sapo-

,nhIllieH OCHOBHhIXrpyrrrr aMHHOT (penTHJIHH, nTHu. H MJIeKOrrHTaIOIII.HX,

BKJIIOIJa5IIJeJIOBeKa), o.zr.HaKOy sapo.zr.bIllieH aMepHOHHrHOeJIb KJIeTOK B

Me}KnaJIbu.eBbIX yIJaCTKaX OTCyTcTByeT. KaMepOH H <PaJIJIOH

[3]

npe,nnOJIO:>KHJIH,IJTO HeKpos KJIeTOK, KaK O,nHH HS MexaHHSMOB pasBH-

l'H5I KOHeIJHOCTH,BOSHHK B epHJIOreHeSe nOSBOH01JHbIXnOCJIe OT)J,eJIeHH51

aMRHOTOT aHaMHHH. .

B. KaJlJICOH

PYlC. 10.8. 06J1aCTH rH6e<IH KJleTOK (saUITpHXOBaHhI) B nOqKaX HH1KHHX KOHeqHOcTelt KypHHO-

(A)

H }'THHoro

(B)

sapoAblwell (no Saunders, Fallon, 1966, In: Major Problems in Deve.

lopmental Biology, ed. M. Locke, Academic Press, New York), a TaK1Ke B nOqKe pyKH qeJlO-

'BeqeCKoro sapoAhIwa

(B)

(no Menkes et aI., 1965, Rev. Roumaine Embryol. Cytol., 2, 161).

PYlC. 10.9, Cxe ••a OnbITa no nepecaAKe yqaCTKa SOHbl nOJl>lpHsYloIl.\ell aKTHBHOCTH HS aaAHeill

qaCTH nOqKH KpblJla KypHHoro sapOAblllla

(A)

B nepeAHlO1O qaCTb noqKH KpblJla APyroro KypH-

Horo sapOAblllla (5). (TIo Saunders, 1972, Ann. N. Y. Acad. Sci., 193, 29.)

,n:HcTaJlbHee

TpaHCnJlaHTaTa 06pasyeTc>I BTOPHqHbllt anHKaJlbHblll 3KTOAepMaJlbHbIll rpe6eHb

(I),

B peaYJlbTaTe.

qero paSBHBaeTC>I KpblJlO C ABollHblM Ha60poM TepMHHaJlbHblX CTpyKTyp

(F).

nepeAH>I>I no-

BepXHOCTb;

aa,l\H>I>I nOBepxHocTb; 2 - anHKaJlbHblll 3KTOAepMaJlbHbIll rpe6eHb aapOAblllla-

penHnHeHTa.

rHOeJIb KJIeTOK HeKOTOpbIX SOH nOtIKH KOHetIHOCTH, nO-BH,n:HMOMy,

TeHeTHtIeCKH npe,n:onpe,n:eJIeHa Ha KOHKpeTHbIX CTa,n:HHX paSBHTHH,

onpe)J.eJIeHHblH nepHO)J., no oopa3HoMy BbIpa:iKeHHIO CayH)J.epca,

BKJIIOtIaIOTCH «tIaCbI CMepTH» H «npHrOBopeHHbIe» KJIeTKH, OCTaBaHCb

B TetIeHHe ern.e HeKOToporo BpeMeHH BHeUIHe HOpMaJIbHbIMH, OOpeqeHbI

norHoHyTb B KpHTHtIeCKOH CTa)J.HH pasBHTHH

[25].

rHOeJIH KJIeTKH

npe)J.llIecTByeT CHH:iKeHHeypOBHH CHHTe3a OeJIKOBH HyKJIeUHOBbIX KHC-

.lIOT

H nOBbIllIeHHe aKTHBHOCTHrH)J.pOJIHTHlIeCKux <pepMeHToB. 3KcnepH-

MeHTbl C nepeca)J.KoH OOJIaCTHsa)J.HeH HeKpoTHlIecKoH SOHbl KypHHoro

3ap0)J.bIllia Ha )J.opCaJIbHyIO nOBepxHocTb nOllKH KpbIJIa nOKaSaJIH, liTO,

XOTH «tIaCbl CMepTH» sanycKaIOTcH npHOJIHSHTeJIbHO Ha CTa)J.HH

17,

oopaTHMocTb 3Toro npouecca HJIH «OTMeHa cMepTHoro npHroBopa»

BOSMO:iKHaBnJIOTb )J.OCTa)J.HH

22.

ITocJIe CTa)J.HH

rHOeJIb KJIeTOK He·

<oopaTHMa, HeCMOTpH Ha TO liTO HBHbIX npHSHaKOB HeKposa He HaOJIIO-

.n:aeTCHBnJIOTb )J.OCTa)J.HH

24.

308a nOJHlpH3YIOllIeH aKTlfBHOCTIf If Mop!f>oreHeTlfllecKlfH KOHTpOJIb

pa3BlfBaIOw.eHcSJ KOHeqHOCTlf, BOJUISH sa)J.Hero KpaH nOtIKH KOHetIHO-

eTH B MeCTe ee coe)J.HHeHHH co CTeHKOH TeJIa HaXO)J.HTCH)J.pyraH 00-

.lIaCTb MeseHXHMbI, OOJIa)J.aIOrn.a51OCOObIMHCBoHcTBaMH. I\JIeTKH 3TOl1:

OOJIaCTH, HaSBaHHbIe sOlioa nOJlflPUSYIOf4ea aKTUBliOCTU, CTHMyJIHpyIOT

!f>opMHpoBaHue )J.ooaBollHblx KOHellHOCTel1:npH nepeca)J.Ke Ha nepe)J.-

HIOIO npeaKCUaJIbHyIO tIaCTb nOtIKH KOHellHOCTU (puc.

10.9).

30Ha no-

JIHpH3yIOrn.eH aKTHBHOCTH BnepBbIe ObIJIa oOHapy:iKeHa B Henocpe,n:cT-

BeHl-IOHOJIH30CTH OT 3a)J.HeH HeKpoTHlIeCKOI1:SOHbI nOtIKH KpbIJIa KypH-

Horo sap0)J.bIllia

[28].

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KOHellHOCTeH nTHll, oeCXBOCTbIX H XBOCTaTbIX aM<pHoHH, penTHJIHH H'

MJIeKOnHTaIOrn.Hx, BKJIl(jqaH lIeJIOBeKa.

PesYJIbTaTbl onblTOB C nepeca)J.KoH TKaHeH nOllKH KpblJIa KypHHoro

sapo)J.bIllia. (oobeKT, Ha KOTOpOM ObIJIO npoHsBe)J.eHo nO)J.aB.nHIOrn.ee

60JIbllII1HCTBO HCCJIe)J.oBaHHH) nOKaSaJIH, tITO nOJIHpHsyIOrn.aH aKTHB-

HOCTbOlleHb CJIaOO Bblpa:iKeHa Ha paHHHX CTa)J.HHXpOCTa KOHellHOCTII:

(cTa)J.HH

16),

HO SHallIITeJIbHO YCHJUlBaeTCH Ha oOJIee nOS)J.HHX

cTa)J.HHX, Kor)J.a HallUHaeTCH )J.u<p<pepeHllHpoBKa naJIblleB (pHC.

10.10).

MaK- I\eH6 H )J.p.

[17]

B OnbITaX in vitro yCTaHOBHJIH, liTO SOHa nOJIH'

PHSYIOrn.eH aKTI1BHOCTH HBJIHeTCH HCTOllHHKOM )J.H<p<pYH)J.HpYIOrn.ero

?raKTopa (OlleBU)J.HO, HeU)J.eHTUlIHoro «no)J.)J.ep:iKUBaIOrn.eMY<paKTopy»

••••BUJIJUlHra), KOTOpblH MO:iKeT BbISbIBaTb oopaSOBaHue anHKaJIbHoro .

SKTO)J.epMaJIbHOrOrpeOHH,

DOJIHpHSYIOrn.UMU CBOHCTBaMU oOJIa)J.aeT He TOJIbKO SOHa nOJIHpU-

SYIOIlI,eH aKTHBHOCTH. B O)J.HOHHs paOOT ObIJIO nOKasaHO liTO KJIeTKH

COM '

HTOB, MeSO)J.epMaJIbHaH TKaHb, JIe:iKarn.aH Me)J.HaJIbHO no OTHOllIe-

Iililo K nOllKe KpbIJIa, H )J.a:iKe MeSOHe<ppoc KypHHoro sapo)J.bIllia oOJIa-