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підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

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PMC. 5-35. UMKll lj>yHKl.IMOHMpOBaHMH G-6enKa. R, - pet.temop; r - ropMOH; AU - a,qeH111naTLt11JKfla3a.

6eJTKOM KJTeTO'IHOHMeM6paHbI H I13MeH5IeT ero aKTHBHOCTb. TaKHMH 6eJTKaMH MOfYT 6bITb <PepMeHTbl a,ueHl1JlaTUMKJTa3a, <Poc<Pomma3a C, <Poc<Pon113CTepa3a ufM<ll, Na+-KaHaJTbI, K+-KaHaJTbI.

CJieLJYIOIUHH 3Tarr u11KJia cPYHKUHOH11poBaHMH G- 6eJIKa - ne<Poc$op11JTl.fPOBaH11e IT<ll, CBH3aHHoro c a-cy6beLl,HHMUeH, rrpwreM <PepMeHT, KaTaJTH3MPYJOIUMH 3'fYpeaKu1110, - caMa a-cy6beLl,MHMUa.

Ll.e<Poc<PopMJTHPOBaHHe rrpHBOLl,MT K o6pa30BaHHIO KOMITJTeKca a-f)],¢1, KOTOpblM He KOMnJieMeHTapeH crreu:McPH'IeCKOMY 6eJTKY MeM6paHbl (HarrpMMep, a,ueHMJTaTUMKJia3e), HO MMeeT BbICOKOe cpOLl,CTBO K ~y-rrpornMepaM. G-6eJIOK B03BpamaeTcH K tteaKTHBHOH $opMe - a~y-f)],¢1. CTpH ITOCJTeLl,YJOIUett aKTHBaUMM peuenTopa H 3aMeHe MOJTeKYJTbl fLl.<ll Ha IT<D LIJfKJT rroBTOpHeTcH CHOBa. TaKHM o6pa30M, a-cy6beLl,MHHUhI G-6eJIKOB coBeprnaroT qenHO'lHoeLl,BIDKeHHe, rrepeHOC5I CTMMYJIMPYIOIUHH l1JTM 11HfM6MpyIOIUMH CMrHaJT OT peuemopa, KOTOpbIH aKTMBMPOBaH nepBM'lHhIMrrocpeLl,HMKOM (HarrpMMep, ropMOHOM), Ha <PepMeHT, KaTaJTM3HpyIOIUMM o6pa30BaHHe BTOpM'!HOfOrrocpeLl,HllKa.

HeKOTOpbre <PopMhI rrpoTeHHKMHa3 Moryr <Poc- <PopHJTllpoBaTb a-cy6beL1,MHHUhI G-6eJTKOB. <Doc<Pop11JTMpoBattHaH a-cy6beLl,MHMUa He KOMnJieMeHTapHa crreu11$H'lecKOMY6eJTKY MeM6paHhI, ttarrpMMep aneHMJTaTUMKJia3e MJTM <Poc<Po;rnrra3e C, no3TOMY He MO)f(eT yYaCTBOBaTh B nepeLl,a'IeCHfHaJia.

r. MEHMJIAT11,11K.JIA3A

<llepMeHT aLl,eHMJTaTUMKJTa3a, KaTaJTM3HPYIOIUMH rrpeBpa11.1ett11e AT<ll B uAM<ll (pMc. 5-36), - KJIIOYeBOH <PepMeHT a,ueHMJTaTUHKJia3HOM CHCTeMbl rrepe11a'IHc11rnana. ALi,emUiaTUMK.Jia3a o6Hap~eHa BO Bcex T11nax KJieToK.

<llepMeHT OTHOCHT K rpynne HHTerpaJTbHblX 6eJIKOB KJieTO'!HOH MeM6paHbl, OH HMeeT 12 TpaHCMeM6paHHblX Ll,OMeHOB. BtteKJieTO'IHhie $par-

MeHThI anemUiaTUHKJia3hI rn11K03HJI11poBaHhI. U11TOrrJTa3MaT11YecK11e Ll,OMeHbl aL1,eH11JJaTLI,11KJia3hl HMeIOT Ll,Ba KaTaJTHTM'IeCKMXueHTpa, OTBeTCTBeHHhlX 3a o6pa3oBaHtte uAM<D - BTOpwrnoro nocpentt11Ka, y'-1.aCTBYIOJ..UefOB perym1u1111 aKTMBHOCTH <PepMeHTa npoTel1HKMHa3bI A.

Ha aKTMBHOCTh a,uettHJiaTUMKJla3hI oKa3hIBaIOT BJIMHHHe KaK BHeKJieTO'iHhie, TaK M BHYTPHKJTeTO'!HbieperyJIHTOphI. BtteKJTeTO'IHhie peryJTHTOpbI (ropMOHbl, 3HK03aHOMLl,bI, 6MoreHHhie aMHHbI) ocymecTBJTHIOT peryJTHUHIO qepe3 cneuMcPHYeCKMe peuerrTOPhI, KOTOpble c ITOMOI..Uh!O a-cy6beLl,I1Hl1U G-6eJIKOB nepena10T cttrHaJihI Ha aneHHJiaTUHKJia3y. a,-Cy6beLl,MH11Ua (CTMMyn11py10maH) rrpM B3aMMOLl,ettcTBMl1 c aLl,eHMJTaTUMKJia30H aKTMBMpyeT <PepMeHT, ai-cy6beL1,MH11ua (Mttm611py10iuaH) 11ttm611pyeT <PepMeHT. B CBOIO oqepeLl,h, a,uemmaTUHKJTa3a CTMMyJT11pyeT rrpoHBJTeHMe IT<D-<Poc<Pa-

254

(<I>11<I>)

 

 

 

NH2

 

 

 

 

&:~N N

 

 

 

ex~~

 

 

 

I

I

I

 

o

 

 

 

 

S~0

o--~-o-~-o-~-o-cH2

 

 

AAeH11111aTL11111<J1a3a

O-CH2

 

o

o

o

~N

 

 

0-

0-

0-

H

 

H

 

 

 

 

 

 

""\

 

H

H

 

 

 

 

 

 

 

 

 

 

H

 

H

 

PPi

 

3

H

 

 

 

OH

OH

 

 

O=P--0

OH

 

 

 

 

 

 

 

 

 

I

 

 

 

A,QeH03111HTp111cpoccpar

 

 

 

o-

 

(AT<I>)

l...11111<J1w-1eCKlllH aAeH03111H-3',5'- MOHOcpoccpaT (l.lAM<I>)

P11c. 5-36. 06pa30BaH11e 1.111K1111o.iecKoro aAeH0311HMOHocj>occj>aTa (l..IAM<I>).

Ta3Hoii aKTl1BHocT11 a-cy6neinrn11u. B pe3yJihTaTe ,n:ect>occpop1rn11pOBaHm1 IT<]) 06pa3y10TcH cy6ne,n:11-

Hl1UhI a -f,LJ,<]) 11 a.-Ul<]), He KOMITJieMeHTapHbie

S I

a,n:emrnaTUHKJia3e.

lf3 8 113yqeHHhIX 1130cpOpM a,n:eH11JiaTU11KJia3hT 4 - Ca2+ -3aB11c11Mb1e (aKT11B11PYIOTCH Ca2+). PeryJIHU11H a,n:eH11JiaTIJ,11KJia3bl BHyTp11KJTeTO'IHhIM KaJThIJ,11eM IT03BOJTHeT KJieTKe 11HTerp11poBaTb aKT11BHOCTb ,n:Byx OCHOBHhlX BTOp11'IHhlXnocpe.11HHKOB uAM<]) 11 Ca2+.

,II,. <l>OCCl>OJUII1A3hl

$occpOJrnna3hI - ct>epMeHThI KJiacca rn,n:poJia3, KaTaJTH311py10m11e Karn60JT113M n111uepoct>occpoJIHn11.1IoB. Pa3JI11qa10T ct>occpoJT11na3b1 ceKpernpHhie, BX0.115llll,11e B COCTaB rraHKpeanrqecKoro COKa, 11 . KJiernqHhie cpoccpoJI11rra3hI. K.n:eTOIJ:Hhie cpoccpoJT11rra3h1 AI' ~' D, C pa3JI11qa10TcH no crrem1$11qHocT11 K oTmerrJTHeMoH: rpyrrne. Bee ct>occt>oJT11na3h1 - KaJThll,11H3aB11Cl!Mbie cpepMeHThI (p11c. 5-37).

<l>oc<l>oJIHRaJa c - ct>epMeHT, f11.1IPOJT113YI01U11H <t>oc<t>o3q1w:pHYIO CBH3h B rrrmJ,epo<t>oc<t>oJT11n11.1Iax. B KJieTKax IJ:eJToBeKa 11)leHrn<t>11u11poBaHo 10 mocpopM ct>occpoJI11na3h1 C, pa3JI11qa10m11xcJI no MOJTeeyAApHotl: Macce, JIOKaJI113au1111, crroco6y peryAAU11M, cy6CTpaTHoi1: crreu11cpttqHOCT11. B CTpYKTYPe BCex 1130cPOPM ct>occt>omma3hl c OTCYTCTBYIOT f11,[(- pocpo6Hhie ,[(OMeHbI, KOTOphie MOfJIM 6bI o6ecrreqMTh MX B3aMMO,n:eilcrn11e c MeM6paHOH. 0,n:HaKO HeKOTOphre cPOPMhI cpoccpoJI11na3bl c CBJI3aHhl c MeM6paHoi1: c IIOMOlll,hlO rn,n:pocpo6Horo «JIKOpH» -

0

II

R2-C-0-CH

I J-o-~-o-x

/6H\

ct>occpo11111na3a C

ct>occpomma3a D

P11c. 5-37. JleliicTBite cj>occj>on11na3_

aUMJihHOfO OCTaTKa MHpHCTHHOBOH KMCJIOThI MJIH 3a cqfa B3aMMO,n:eilCTBM5l c IIOBepXHOCThlO 6MCJT05l. KaTaJIHTlfqeCKaH aKTMBHOCTh Bcex M3o<PopM cpocct>oJTttrra3hI -= 3aBHC11T OT HOHOB KaJihUM5l.

fioJThIIIHHCTBO ct>occpoJI11rra3 C crreu11$11qHo B OTHOIIIeHMH cpoccpaTM,[(MJTMH0311TOJIOB 11 rrpaKTHqecKM He rn,n:poJimyeT ,n:pyrne THilhI cpoc<t>omm11- ,n:oB. AKT11BHhIH cpepMeHT MO)l(eT rll,[(pOJI1130BaTb ,n:o 50% OT o6IIIero KOJilfqecrna <t>occpaTI1)lMJI11tto3MTOJTOB KJieTOqHoil MeM6paHhT. IlpM fH.1IPOJIH3e cpocQ:Jarn,n:11JI11Ho3HTOJI-4, 5-611cQ:JocQ:Jarn o6pa3YIQTCH npo,n:yKThI ,n:11au,HJirJIMuepoJT (Mf) 11

255

anchoring proteins).
)J,HHHU, R2C 2 -

l1H03HTOJI- l ,4,5-TPH<POCcPaT (11<1>3), CJIY)l(aru11e BTOp11qHbIMl1 nocpe,IJ,HHKaMH B TpaHCMeM6paHHOM nepe,IJ,aqe CHrttaJia no HH03HTOJicPOCcPaTHOMY nyTH.

E. IIPOTEl1HKMHA3hl

Bee nom1pHhre c11rHaJihHh1e MOJieKYJihI, ,IJ,ewcrnyIOIUHe Ha KIIeJKY-MMllleHh qepe3 MeM6paHHh1e peuemopb1, ocyruecTRJHIIOT CBOIO 6110JIOIWieCKYIO cPYHKll,H!O nYJeM ¢oc¢opI1J1MPOBaHIIB cneutt<PWieCKMX 6eJIKOB l1 ¢epMeHTOB, peryJIMpYJOIUHX MeTa60JIH3M B KJieTKe. <l>oc¢opwrnp0Bam1e H3MeIDieT (YBeJIWIHBaeT I1J111 YMeHblllae1) I1X aKTHBHOCTh. KaTaJIH3HpyJOT ¢oc¢opttJIMpOBaHMe 6eJIKOB (npOTeMHOB) rrpoTeHHKHHa3bl no aMHHOKl1CJIOTHhIM OCTaTKaM cepIDia, TPeOHHHa, 111p031rna. llpoTettHK11Ha3hI MOryT 6hITh cy6be,IJ,ttHttueil MeM6paHHoro peuemopa, HarrpHMep THP03HHOBasi rrp01e11H1<11Ha3a peuerrrnpa ttHcymrna, aKTHBHOCTh KOTopow peryJI11pyeTcH ropMoHoM. ):Wyra51 rpynrra - npoTettHKHHa3hI, perymrpyeMbie BTO-

pttqHhIMl1 BeCTHl1KaMl1 ropMOHaJibHOfO c11rHa.rra (uAM<l>, ufM<l>, Ca2+, )J,Af), Hanp11Mep rrpoTel1H-

Kl1Ha3a A, npo1e11HKl1Ha3a C, rrpo1e11HK11Ha3a G,

KaJibMO,IJ,YJI11H3aBl1CHMbie npoTeHHKl1Ha3bI H ,Il,p.

1. llpomeuHKUHa3bt A

Tipo1e11HK11Ha3hI A ( uAM <l>-c111MyJI11pyeMh1e) yqaCTBYJOT B a,IJ,eHHJiaTUHKJia3HOM CHCTeMe nepe,IJ,a- qw c11rHaJia. ITpoTe11HK11Ha3a A cocTOHT 113 4 cy6ne- ,IJ,Byx peryJIITTOPHhIX cy6be)J,11HHU (R2)

11 ,IJ,Byx KaTaJI11rnqecK11X (C2) (CM. p11c. 5-41). KoMIIJieKc ~c2 He o6Jia,IJ,aeT ¢epMeHTaTHBHOM aKTHBHOCTb!O.

KoMTIJieKc ~C2 pa3HhIMH crroco6aM11 npttKpenJIHeTcH K MeM6paHe. HeK0Topb1e ¢opMbI npo1e11HK11Ha3hI A «3a51Kop1rna10TC5I» c noMOIUhIO aJI11- ¢aTWrecKoro OCTaTKa MHPl1CTHHOBOH KMCJIOTbI KaTaJIHTHqecKHX cy6be,IJ,HHI1II. Bo MHomx TKaH5IX npoTe11H1<11Ha3a A CB513aHa c «3asIKopeHHhIM» 6eJIKoM

AKAP, (oT aHr.rr. cAMP-dependent protein kinase

AKAP, MMeeT ueHTIJ CB513bIBamm )JJIH peryJIHTOPHbTX cy6be)J,HH11U npoTel1HKHHa3bl A C rroMOIUhJO 6eJIKa AK.AP, rrpoTeMHK11Ha3a A

CBH3bIBaeTCH c MeM6paHOH B o6JiaCTH JIOKaJIH3aU1111 cj)epMeHTOB, KaTaJIH3ttpYJOIUI1X o6pa30BaHMe uAM<l> ( <l)leHI1J1aTUHKJia3a) l1JIM ero rl1)],poJI113 (¢ocq>0,1J,113CTepa3a), a TaIOKe 6eJIKOB, B peryJIHU,HM aKTHBHOCTH KOTOpb!X ¢epMeHT rrpI1HMMaeT yqacTHe, HarrpMMep rroTeHUMaJI3aB11Cl1Mhie Ca2+-KaHaJihI.

Pery.rrmopHbie cy6be,IJ,11Hl1UhI rrpo1e11HK11Ha3hI A

11MeJOT crreu11¢11qecK11e ueHTpb1 )J,JI5I CB513bIBaHM

uAM<l>. llp11coe,IJ,11HeH11e uAM<t> K peryJIHTOpHblM cy6be,IJ,HH11UaM rrpHBO)J,MT K H3MeHeHl1IO KOHcPOPMaUMM I10CJie)J,HI1X I1 CHH)l(eHMJO cpO,Il,CTBa K KaTaJIHTHqecKHM cy6be,IJ,ttHMUaM C, rrpo11cxo,IJ,11T ;rnccou11auM no cxeMe:

uAM<l>4 + ~C2 ~ uAM<l>4 ~ + C + C

Cy6be,IJ,HHMUhI C npe,IJ,CTaBAAIOT co6o.H aKTl1BHYJO ¢opMy npoTettHKttHa3bI A, KOTopa5I KaTaJIH311pye1 peaKUMH ¢oc¢op11JI11poBaHHH 6eJIKOB no cep11Hy M TpeoHMHy. KaTaJI11T11qecK11e cy6be- ,IJ,I1Hl1UhI C y pa3HhIX T11noB np0Te11HK11Ha3 A He H)J,eHTJiqHbl' OHH pa3JI11qaJOTCH rrpe)l()J,e Bcero crreutt¢11qHOCTblO B OTHOllleHMH 6eJIKOB-cy6- CTpaTOB.

2. llpomeUHKUHa3bl c

llpoTel1HKl1Ha3bl c yqacTByIOT B MH0311TOJicPOC- <PaTHOM CHCTeMe rrepe,IJ,aq11 CHrHaJia. <l>epMeHT coCTOMT 113 ,IJ,Byx cPYHKUHOHaJibHO pa3JII1qHbTX )J,0- MeHOB - peryJI5ITOpHoro 11 KaTamnttqecKoro.

PeryJIHTOPHhIM ,IJ,OMeH co,IJ,ep)l(HT 2 c1pyKTYPhI ( «IUIHKOBble IlllJib~I»), o6pa30BaHHhie ¢parMeHTaMl1 nerrTl1)J,HOH uen11, 6oraThIM11 UHCTel1HOM, H CO,Il,ep)l(aIUHMH 2 HOHa UliHKa (CM. pa3,IJ,eJI 1).

«11,HHKOBbTe IIaJibUhl» yqaCTBYJOT B CB513bIBamm )J,H- aUHJifJIMUepoJia. )l,pyro.H ¢parMeHT peryJIHTOPHOro ,IJ,OMeHa HMeeT BhICOKOe cpo,IJ,CTBO K Ca2+. lloBhillleH11e KOHUeHTpau1111 KaJibUHH B UHT030Jie yBeJIJ1qHBaeT cpO,Il,CTBO npoTeHHKl1Ha3hl c K ¢oc- ¢aTl1)],I1JICep11Hy MeM6paHhI. TpaHCJIOKaUM rrpoTeHHKMHa3hT c K MeM6paHe I103BOAAeT ¢epMeHTy Cml3aTbC5I C ,ll,Af, KOTOphIH erue 60Jihllle ITOBbilllaeT cpO,Il,CTBO rrpoTeHHKJ1Ha3bI c K HOHaM KaJibUM (p11c. 5-38). Hatt6oJiee pacrrpocTpaHeHHhie 1130- <PopMhT rrpoTeMHKHHa3hI C aKTHBHPYJOTCH Ca2+, )J,Hall,HJifJIMUepoJIOM J1 ¢oc¢aTl1)],HJICep11HOM.

KaTaJIMTI1qeCKHH ,IJ,OMeH HMeeT ueHTIJ, cBH3hTBa10ru11w AT<t> 11 6eJioK-cy6cJPaT. AKTttBHasi ¢opMa cj)epMeHTa npOTeHHKHHa3hI C <Poc¢opl1JII1pycr 6eJIKM no OCTaTKaM cepMHa H JpeOHttHa. CHJOKeHHe KOHueHTIJau1111 HOHOB KaJibUH5I B KJieTKe Hapyrnacr CB513h rrpOTettHKHHa3hI C c ¢occj)aTl1)J,I1JicepttttoM 11 ,IJ,HaUHJirJIMuepoJIOM, ¢epMeHT rrepeXO)J,MT B HeaKTHBHYJO ¢opMy J1 OT,IJ,eAAeTCH OT MeM6paHbl.

3. llpomeuHKUHa3bl G

8 01JI11q11e 01 rrpoTel1HKMHa3hI A, rrpoTeHHKIIHa3a G rrpMCYJCTByeT He BO Bcex TKaH5IX, ee 06Hap)')KHBaJOTB .rrefKMX, M03)1(eqKe, fJia)J,KJIX Mblilluax H TIJOM6ouMTax. l13o¢opMhI rrpOTeHHKIIHa3hI G

256

 

 

 

 

<t>C ,lJ,AI

 

 

nKC

Ca2+ \__.

nKC

~~

 

 

____.ca2

2

 

HeaKTlllBHafl

HeaKTll!BHafl

 

Ca

 

E-OH

 

/

 

 

E-OPO~-

 

~

 

np0Te111H 7

~

 

 

 

 

<t>occ:f:>onpore111H

 

AT<t>

A,D,<t>

 

 

PMC. 5·38. Peryn:111.1M:11 8KTMBHOCTM npoTeMHKMH83bl c (nKC). <PC - cpoccparnA"1nCep1o1H; ):!Ar -

 

A1<1a41o1nrn1o14epon.

MOryr 6bITb CBH3aHbl c MeM6paHOH IDIH HaxO,IJJ1TbCJ:l B UIITOIVIa3Me. PacTBopttMM rrpoTeHHKHHa3a G coCTOHT H3 ,LJ,Byx Jf,Il,eHTifqHblX cyfu,eattHHU, KIDK,ll,aH H3 KOTOpbIX HMeeT asa ueHTpa )J)UI CBJ:l3hIBaHHH ufMCl>. IlpttcoeattHeHtte ufMCl> K perymrropHhIM uempaM Bbl3bIBaeT KOHcpOpMaUHOHHbie H3MeHeHHH cy6i,eattHHU H IIOBblIIlaeT KaTaJIHTWieCK)'IOaKTHBHOCTb cpepMeHTa (p11c. 5-39). JlpOTettttKtttta3a G, rroao6tto rrpOTettHKHHa3e A 11 C, crreumpwma B OTHOlllemm orrpeaeneHHblX 6eJIKOBblX cy6CTPaTOB, KOTOpbre Otta <t>oc¢opIDI11pyeT rro ocTaTKaM cepttHa H TPeott11tta.

.IK. <l>oc<I>O,IJ;H3CTEPA3hI

Cl>occpoatt3CTepa3hI - cpepMettThI, Karnnm11pyIOW:tte rrpespamett11e uAM<D (pttc. 5-40) HJIH urM<D B tteaKTHBHhle MeTa60JIHTbl AM<D HJIH fM<D. Cl>occpoa113cTepa3bJ, CHmKaH KOHUeHTpaUHH BTOpl1'lHbIXrrocpeaHHKOB, pa3pbIBaIOT uerrb rrpespamett11tl, Bhl3BaHtthIX aKTttBaTopoM peuerrrnpa.

 

E-OH

 

npoTell!H

 

--- t ~ AT<t>

 

~ A,D,<t>

 

E-OPO~­

 

cl>occj:lonpore111H

nporell1HKll1Ha3a G

npoTe"1HKll1Ha3a G

HeaKTll!BHafl

8KTll1BH8fl

P11c. 5·39. Peryn:111.1M:11 aKTMBHOCTM npoTeMHKMH83bl G (nKG).

NH2

(x~ ):__N

~..J-9~

o~~:~O,~I "°""'°""30Tep83a _ W ~NN

0-~-0-CH2 o

oJ-...JQ OH

 

O-

 

H,O

H

H

 

 

 

I

!

 

HO

OH

ff

Mecro r1111=1pom13a

 

 

 

L1111KI1111'leCKll1Ha1=1eH03111H - 3', 5'-

AAeH03111HMOHoc:f:>occj:Jar

MOHO<:l>Occt>aT (L\AM<t>)

 

(AMcl>)

 

 

 

 

PMC. 5-40. npeepa11.1eHMe l.IAM4> B AM4>.

257

9-2128

<l>oc¢ontt3cTepa3bI ITpttcyrcrnyIOT B KJJeTKax TKaHeti B 2 ¢opMax: B ¢opMe pacrnopttMoro 6eJIKa H MeM6paHHOCBH3aHHOro. <l>opMbl ¢epMeHTa, CBH3aHHbTe c MeM6paHOH, B pa3HbTX TKaHHX COCTaBJIHIOT 5-40%. B O)lHOH 11 TOii )K:e TKaHH MOryr ITpwcyrCTBOBaTb pa3Hbie ¢opMbl <t>oc¢0JJH3CTepa3bI, pa3JI11':taJOll.l,l1eCHTIO cpO)lCTBY K cy6cTf)aTaM, MOJieKyMpHOMY Becy, 3apH)ly, perymrrnpttblM CBOHCTBaM H JIOKaJIH3auim B KJJeTKe.

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B 6oJiblilllHCTBe TKatte:i1 rrp11cyrcrnyeT ¢oc¢0.n11- 3CTepa3a-l, 6oJiee cneuw¢wqua51 K uAM<I>, aKTMBttpyeMaH Ca2+ , KOMTineKCOM 4 ea2+-KanbMo.nymm 11 ufM<I>.

3. MEHRJIATU:MKJIA3HAH CMCTEMA

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THBallHH a,L1,eHHJiaT11HKJia3hl:

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• 3To Bhl3bIBaeT JlHCcouttauw10 KOMrrneKca. OT-

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[f][R][G-IT<I>] ~ [f][R] + a-IT<I> + py;

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KOH¢0pMaUHOHHbie ll3MeHeHHH B KOMITJieKce

[a-IT<I>] [AU] CT11MYJI11PYJOT TIOBbIWeHHe rr<1>- ¢oc¢arn3uoi1 aKTHBHOCTH a-cy6be)l,HHl1Ubl. IlpOTeKaeT peaKUHH Jle¢oc¢opHJIHPOBaHl1H IT<I>, H OJJHH H3 ITPOJlYKTOB peaKu1111 - ueop-

rauHqecK11i1 cpoc¢aT (P.) OT)lem1eTCH OT a- cy6beJlHHHUhI, a KOMITJie~c [a-f)J,<I>] coxpaHH-

eTcH; CKOpOCTb rHJlPOJIH3a OITpe)l,eJIHeT BpeMH npoBe,ll,eHHH cwruana;

o6pa30Bauwe B aKTl1BHOM ueHTpe a-cy6be)ll1- Hl1Ubl MOJieKYJihI f)J,<I> CHH)K:aeT ero CPOJlCTBO K a).l,eHMnaTUHKJia3e, HO yBeJil1'lHBaeTcpOJlCTBO K py-cy6beAHHHUaM. G,-6enoK B03BpamaeTCH K HeaKTl1BHOH ¢opMe;

ecJIH peuernop CBH3aH c aKTHBaTopoM, HaITpHMep ropMOHOM, UHKJI <i>YHKUHOHttpoBamrn G, 6eJIKa IIOBTOpHeTCH.

AlcrHBallHH npoTeUHKHHa3hI A (TIKA)

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ITJieKCa C2~ Ha KOMTIJieKC uAM<l>4 ~MC + C.

• AKTHBHaH ITpOTel1HK11Ha3a A ¢oc¢opHJT11pyeT cITeUHWM'leCK11e6eJIKM ITO cepwuy 11 TpeOHHHY, B pe3ynhTaTe H3MeHHIOTCH KottcpopMaum.1. H aKTMBHOCTb ¢oc¢opHJTHPOBaHHhIX 6eJIKOB, a 3TO npHBO,[J,11T K H3MeHeHl1IO CKOpOCTM 11 HarrpaBJieHMH peryJittpyeMbIX HMH ITpoueccoB B KJJeTKe.

• KouueHTpauHH uAM<I> B KJ1eTKe MO)K:eT pefYJIHPOBaTbCH, Otta 3aBl1CHT OT COOTHOWeHl1H aKTl1BHOCTeii ¢epMeHTOB a).l,eH11JiaTIJ,MKJia3bl 11 ¢oc- ¢oJll13CTepa3hl.

oOJibWYIO pOJib B peryJIHUllll BHYTPl1KJieToqHOH c11ruanbHOM CHCTeMhI ttrpaeT 6eJioK AKAP . <<3m:1KopetttthIM» 6eJioK AKAPs yqacrnyeT B c6opK~ ¢epMeHTHbIX KOMITJieKCOB, BKJIJOqaJOIUl1X He TOJib-

258

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JleKyJI a.ueHHJiaTU!1KJia3bl c o6pa30BaHl1CM TblC51lJ MOJICKYJI uAM<P. Ha 3TOM 3Tane c11rna.JI ycl1J111BaeTcs1 B 102-103 pa3. 06pa3y10w11i1c51 uAM cD «BKJIIOtJaIOT» .upyroi1 ¢epMeHT - npoTeHHKHHa3y A, yc11mrna51 Cl1rHa'lewe B 1000 pa3. <PoccflOpl1JJ11pOBaHl1e ¢epMeHTOB npoTeHHKHHaJoti A eiue 6011bwe yc111111BaeT cJ..ffHari, B pe3yJihTaTe cyMMapHoe ycl1JleH11e paRHO 106-107 pa3. TaKHM o6pa30M, no MexaHH3MY KacK<lllHOro yc1111eH11H o.uHa MOJieKYJJa perym1rnpa

259

cnoco6Ha H3MeHl1Th aKTl1BHOCTh MHJIJIHOHOB LlPYfHX MOJieKYJI.

Ho MH mo6oti H3 cHcTeM TpattcMeM6pattttoti nepe,ll,al!l1 CHfHaJia KJieTKa HMeeT LlPYryIO Cl1CTeMy, no,ll,aBJIHIOJUYIO 3TOT c11rnan. K(l)K)J,hIH 113 3TaTIOB B <PepMeHTHOM KaCKa)l.e HaXO)l,11TCH no)], KOHTPOJleM cneIJ,11aJihHhIX TIO)l.aBJIHIOIIl,HX 3TOT Cl1fHaJI MexaHH3MOB. Hanp11Mep, )].JIHTeJihHoe )l.eticrn11e ropMoHa npHBO)l.HT K )l,eCeHCl16llJUf3aIJ,Hl1 MeM6paHHhIX peuenTopoB: OHll Jil160 ttHaKTHBttpyIOTCH, Jlll60 BMeCTe c ropMOHOM norp)')KaIOTCH B KJieTKY nocpe,ll,CTBOM 3H,ll,OUMT03a. B pe3yJihTaTe )l.ecettc11611n113au,u11 peuenrnpoB cTenetth aKTttBau1rn a)l.eHttJiaTUMKJia3HOH CHCTeMhl CHIDKaeTCH. Ecm1 B KJieTKe )l,Jll1TeJibHOe BpeMH TIOBhIIIIeHa KOHIJ,eHTpaU,ttH uAM<I> (nOBhIWeHa aKTllBHOCTh npoTellHKllHa3hI A), MO)KeT npoucxo)l.ttTh <Poc<PopllJlttpoBaHtte KaJihIJ,tteBhIX KaHaJIOB, qTO npHBO,ll,llT K IIOBhIIIIeHl1IO KOHu,ettTpau11u Ca2+ B KJieTKe. Kanhu,11ti aKTttBttpyeT Ca2+-3aBttCl1MYIO <Poc<Po)l.113cTepa3y, KaTM113npy10my10 rrpeBpamettne uAM<I> B AM <I>.

B pe3yJihTaTe ttHaKTMBau,u11 rrpoTel1HKMHa3hI A (R2C2) CHtt)KaeTCH CKOPOCTh <Poc<PopllJlttpOBaHl1H crreun<P11qecKHX <PepMeHTOB. 3aBepIIIaeT «BhIKJIIOlfeHtte» cttcTeMhI <Poc<PonpOTentt<Poc<Parn3a, ,ll,e- <Poc<PopllJlttPYIOmaH <Poc<PonpoTettHhI.

BJmsrnue 6aKTepuaJibHblX TOKCHHOB ua aKTHBuocTb a,l],eHHJiaTIJ,HK.JUl3bl (A,Il;<l>-pu6o3HRHpOBa- HHe G-6eRKOB)

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XoJiepHhIH TOKCl1H - on11roMepHhIH 6enoK. 0)].Ha 113 cy6be)l.11H11U - <PepMeHT NJ,<1>-pll603llJITPaHC- <Pepa3a; npOHl1KaH B KJieTKy, OHa KaTaJil1311pyeT rrp11COe)l,l1HeH11e ALl,<l>-p11603hl K a,-cy6oe)l.llHl1IJ,e KOMilJleKca [a,-IT<l>][AUJ (3Tan aKT11BaIJ,ttl1 a)l.eHttJiaTIJ,llKJia3hl).

NAO++ [a s-IT<l>][AU] ~ [ALl,<l>-p11603l1Jl-a,- IT<l>][AU] + HllKOT11HaMl1)]. + H+.

NJ,<l>-p11603l1JlttpoBaH11e ttttrn6ttpyeT rrpoHBJieHtte IT<l>-<Poc<Parn3HOH aKT11BHOCTtt a,-cy6'he)l.tt- H11UhI, He npo11CXO)l,HT )l,e<Poc<PopttJil1POBaHtte IT<I>. UnKJI cPYHKUHOH11poBaH11H G,-6eJIKa ocrnHaBJinBaeTcH Ha 3Talle aKTHBaIJ,1111 <PepMeHTa a)l.e- Hl1JlaTIJ,llKJla3hI, OTBetiaIOmero 3a o6pa3oBaHtte uAM <I> tt3 AT<I>. <l>epMeHT a,ll,eH11naTUHKJia3a coxpattHeT nOBhIIIIeHHYIO aKTllBHOCTh B TelJeHtte rurnTeJihHOfO BpeMeH11.

Cy6'he)l.11Httu,a KOKJilOIIIHoro TOKC11Ha, npoH11KaH B KJieTKY, KaTaJil1311pyeT ALl,<l>-p1160311Jil1PO- BaH11e CX;-Cy6'he)l.11HttUhIaKT11BHpoBaHHoro G;-6enKa (a;py-fT<I>).

NAD++ [a;py-IT<I>] ~ [ALl,<l>-p11603ttJI- a;PYIT<I>] + Hl1KOTttHaMl1)]. + H+.

M o)l.11<P11uupoBaHHaH a;-cy6'he)l.11H11uacoxpaHH- eT BhICOKOe cpO,ll,CTBO K py-cy6'he)l,HHHUaM, T.e. G;-6eJioK TepHeT crroco6HocTh )l.11ccou1111poBaTh Ha a;-IT<I> 11 py-cy6'he)l.llHttUhI.TaKttM o6pa3oM, HHrn6ttpy10m11ti Cl1fHaJI ( ai-IT<l>) He )l.OCT11raeT a,l],e- HHJlaTIJ,llKJia3hI, 3HalJl1T B 3TOM CJiyqae B03MO)!(Ha TOJlhKO ee aKTllBaIJ,ttH rrp11 CBH3hIBaHl111 c a,-IT<I>. Ll,ettCTBtte KOKJllOWHOfO TOKCttHa Ha KJieTKH TKaHeH Bcema np11BO)l,11T K IIOBhIWeHl110 ypoBHH uAM<l>.

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If. lfH03HTOJI<I>OC<I>ATHASI CHCTEMA

<l>yHKUHOHHPOB<lHtte HH03HTOJlcPOCcPaTHOH CHCTeMhI TpaHCMeM6paHHOH nepe,ll,alJl1 Cl1fHaJia (p11c.

5-42) o6ecnel!HMIOT: R (peuemop), <Poc<Ponttna3a C, Gplc - 6eJIOK, aKT11BttpYIQIIl,11H <Poc<PoJIHTia3Y C, 6eJIKll 11 <PepMeHThI MeM6paH 11 UHT030J1H.

IloCRe)l.OBaTMbHOCTb co6blTHH, DpHBO,ll,HUUIX K aKTHBaIJ,HH <1>oc<1>0Runa1h1 C:

• CBH3bIBaHHe Cl1fHaJihHOH MOJieKYJihI, Hanp11-

Mep ropMOHa c peuenTOpOM (R), Bhl3hIBaeT

l13MetteH11e KOHcPOPMaIJ,1111 H yBem1lJeHtte cpo,ll,-

CTBa K Gp1c-6eJIKY.

o6pa30Batt11e KOMnJieKca [f][R][Gp1c-fLl,<l>] rrp11BO)l,HT K CHIDKeHl1IO cpO,ll,CTBa a-npOTOMe-

pa Gplc-6eJIKa K fLl,<l> tt YBeJil1lJett1110 cponcrna K IT<l>. f.ll,<I> 3aMem1eTCH Ha IT<l>.

3TO Bhl3bIBaeT )l,11CCOIJ,11aIJ,1110 KOMilJleKca; OT- )l,eJ111BlllaHCH a-cy6'be)].l1H11Ua,CBH3aHHaH c MO:-

JieKYJIOH IT<I>, nptto6perneT cpo,ll,cTBo K <Poc- <Ponttna3e C.

260

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a-IT<l>

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B3al1MO)J.eilcTByeT c <Poc<PommaJoi1: C 11 aKTIIBl1pyer ee. no)]. )J.eHCTBl1eM ¢oc<}lomma- 3bI-C rrpol1CXO)J.HT fH)lpom13 .JIID1l1,Ila MeM6paHbI <Poc<Parn.ru1mrno311TOJI-4,5-611c¢oc¢aTa (<l>M<l>2).

• B XO)J.e f11)J.pOJil13a 06pa3yeTC5I 11 Bb!XO)J.l1T B Ul1T030Jlb fH)J.POcPHJibHOe Bemecrno l1H0311TOJI- 1,4,5-Tp11<}loc¢aT (M<l>). )1,pyroH: rrpo)J.)'KTpeaKu1111 )J.11aUl1JifJil1UepoJI ()1,Af) OCTaeTC51 B MeM6patte 11 yqacrnyeT B aKT11Bau1111 <PepMeH-

Ta rrpoTel1HKHHa3hI c

• l1H0311TOJI-1,4,5-Tpl1<}loc<}laT (H<1>3) CB513hIBaeTC51 cneuJI¢wrecKHMH uempaMJI Cai+-Katta-

4>oc¢on111naaa C

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AKTWBHblH cpepMeHT

Jia MeM6paHbl 3P, 3TO npJIBO)J.JIT K l13MeHeHl1IO KOH<}lOpMaUJIJI 6eJIKa JI OTKpbITJIIO KaHaJia - Ca2+ nocTynaeT B UJIT030Jih. B oTcyrCTBJie B UHT030Jie 11<1>3 KaHaJI 3aKpbIT.

AKTuBal.UUI npoTeHHirnHa3hl C

• noBhillleHJie KOHileHTpauJIJI Ca2+ B Ul1T030Jie KJieTKH YBeJIWI11BaeT CKopoCTh B3aHMO)J.ettcTBmr Ca2+ c tteaKTJIBHhIM UJIT030JihHhIM <PepMeHTOM npoTel1HKHHa30H c (IlKC) JI 6eJIKOM KaJibMOJl)'Jil1HOM, TaKHM o6pa30M CJifHaJI, rrpMH51TbIH peuerrTopoM KJieTKH, pa3,llBal1Baerc51.

261

Cmt3hIBam1e npoTeHHKHHa3hI C c HOHaMH Kllilh-

11,IDI n03BOJUieT ¢epMeHTY BC'rynaTbB KaJlhll,HMonocpe.n:oBaHHOe B3aHMOL(eHCTBHe c MOJieK)'- JiaMII «KIICJIOro» ¢oc¢0JI1In1I.n:a MeM6paHhI, ¢oc¢aTttnHJicepHHa ( <l>C). Ll,1Ia11,HJirn1I11,epoJI, 3aHHMaH cneuttQ:>wrecKJie ueHTphI B npOTeHHKIIHa3e C, eme 6oJiee yileJilllIIIBaeT ee cpo.n:CTBO K HOHaM Kllilhll,IDI.

Ha BHyrpeHHei1 crnpoHe MeM6paHhI o6pa3yeTCH

¢epMeHTaTHBHbIM KOMilJieKC - [ nKC] [Ca2+l

[Ll,Af][<l>C] - aKTIIBHaH npoTe1IHK1IHa3a C, ¢oc¢opHJIIIPYJOmaH crreu1I¢wrecK1Ie ¢epMeHThI ITO cepHHY H TpeOHHtty.

Y11acme 6eJIKa KaJibMO,rzyJIHHa B HH03HTOJ](l>occt>aTHoii nepe,l1,a11e cumaJia

B KJieTKax MHOrHX TKaHei1 rrplICYTCTByeT 6eJIOK KllilhMOnyJIHH, KOTOpbIM cPYHKll,HOHHpyeT KaK BttyTPIIKJieTOlJHhlH peuerrTop Ca2+, OH HMeeT 4 ueHTpa nJUI CB513hrnamrn Ca2+. KoMITJieKc [KllilhMOD:YJIIIH]- [4 Ca2+] He o6JianaeT ¢epMeHTaTIIBHoi1 aKTIIBHOCThIO, HO B3aHMOL(eHCTBHe KOMilJieKca c pa3JUi'l- HhIMII 6eJIKaMH II ¢epMeHTaMII rrplIBOL(HT K HX aKTHBaUHII.

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<l>epMeHThI, KaTaJIII31Ipy10m1Ie BOCCTaHOBJieHIIe ¢oc¢oJIHTIH.LJ.a, aKTIIBHPYJOTCJI ¢oc¢op11JIIIPOBaHHeM npOTeHHKHHa30ll c.

KoHUeHTpauHJI Ca2+ B KJieTKe CHIDKaeTCJI .n:o HCxonttoro ypoBH51 nplI ,nei1cTB11H Ca2+-AT<l>-m UJITOTIJia3MaTH'lecKoi1MeM6paHb1H3P,a TaIOKe Na+/Ca2+ - H H+/CaH-TpaHCJIOKa3 (aKTHBHhIH aHTHIIOPT) KJie-

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AT<l>-a3 MO)!(eT aKTIIBHpoBaThcH:

KOMTIJieKcoM [KarthMOnyJIHH][4 Ca2+];

rrpoTeHHKIIHa3oi1 A (¢oc¢opHJI1IpoBaHHeM);

• npOTeHHKHHa30M c (¢oc¢opHJIHPOBaHHeM). I10HH)!(ett11e KOHUeHTpau,HH Ca2+ B KJieTKe H .n:1IauHJirn1IuepoJia B MeM6paHe nplIBO.IllIT K II3MeHeHHIO KoH¢opMau11H rrpoTeHHKMHa3hI C, CHII)!(e-

HHIO ee CpOL(CTBa K Q:>ocQ:>aTHL(HJICepHHy, Q:>epMeHT ,IlHCCOll,HHpyeT B 11,HT030Jlh (HeaKTHBHM ¢opMa).

<l>oc¢opHJIHpOBaHHbie rrpOTeHHKHHa30H c ¢epMeHThl II 6eJIKM rro.n: .n:ei1cTB1IeM ¢oc¢orrpoTeHH- ¢oc¢arn3h1 rrepexo,nHT B .ne¢oc¢op11JIHpoBaHHy10 Q:>opMy.

K. IIEPE,lJ,AqA ClffHAJIA c IIOMOIIIJ>IO

BHYTPHK.JIETQqffhIX PEUEIITOPOB

nepe.n:aqa CHrHarta JIHTIIIL(OpaCTBOplIMbIX CTepo- 1-0lHhIX ropMOHOB H THPOKCHHa B03MO)Klia TOJihKO rrpH rrpoxo~eHHH 3THX ropMOHOB qepe3 ITJia3MaTH'leCK)'IOMeM6paHy KJieTOK-MHWeHeM (p11c. 5-43).

Peuemoph1 ropMOHOB MOfYT Haxo.n:HThCJI B UHT030Jie HJIH B Mpe. ll,HT030JihHhie peuemophr CB513aHhI c 6eJIKOM-warrepOHOM (qacTO 3TO rpyrrrra 6eJIKOBwarrepoHOB). 51.n:epHhie II UIIT030JibHhre peuemophr CTepOH.LJ.HhIX H THpeOIIL(HbIX ropMOHOB co.n:ep)!(aT Ll,HK-CBJI3hIBaIOI11,HM L(OMeH, xapaKTeplI3YIOI11,HMC51 HaJIH'IHeM.n:Byx crpyKzyp «UHHKOBbIX Illlilbll,eB».

IloCJie,lJ,OBaTeJILHOCTh co6LITHil, npuso)UlllUIX K aKTHBal.UIH TpaHCKpHOl.UIH:

ropMOH rrpoxo.n:HT qepe3 L(BOMHOM JIHII.H.llHhlM CJIOM KJieTO'IHOMMeM6paHhI.

B3aHMOL(eMCTBHe ropMOHa c peuenTOpOM (R)

IIPIIBO,Ll,IIT K H3MeHeHIIIO KOHQ:>opMaumt peuenTOpa w CHH)!(eHHIO cpo.ncrna K 6eJIKaM-warre- poHaM, OT,Ll,eJIJIIOill,HMCJI OT KOMITJieKca ropMoHpeuemop.

• KOMTIJieKC ropMOH-peuerrTOP npoxo.n:HT B MPO, B3aHMO,Ilei1crnyeT c peryJIJITOPHOM HYKJieOTH.LJ.- HOM IIOCJie.n:oBaTeJibHOCTbIO B Ll,HK - 3HXaHcepoM HJIH cai1JieHcepoM.

yBeJUtlJIIBaeTC51 (rrpH B3alIMO,Ll,eMCTBHH c 3HxaHcepoM) IIJIH YMeHhwaeTCH (rrplI B3aHMo.n:etiCTBIIII c cai1JieHcepoM) ,Ll,OCTynHOCTh rrpOMOTOpa .LJ.JIJI PHK-rroJIHMepa3hI.

cooTBeTCTBeHHO YBeJIIIlJHBaeTCJI HJIH yMeHhwaeTCH cKopocTh TpaHcKpHrruHII CTPYKTYPHhIX reHOB.

yBeJIH'IHBaeTCJI IIJIH yMeHbwaeTCJI CKOpOCTh TpaHCJIJIUHH.

II3MeHHeTC51 KOJIII'leCTBO 6eJIKOB, KOTOPhie MOfYT BJIHJITh Ha MeTa60JIH3M H cPYHKIJHOH3JlbHOe COCT051HIIe KJieTKII.

3¢¢eKThl ropMOHOB, KOTOpbie rrepe.n:a10T CIIfHaJI qepe3 BHYTPIIKJieTOlJHhie peuerrrnphI, HeJih3JI Ha6JI10.n:aTh cpa3y, TaK KaK Ha rrpoTeKaH1Ie MaTplI'IHhIXrrpoueccoB (TpaHCKplIIIUHIO II TpaHCJI51UHIO) Tpe6yioTCJI 'laCbl.

262

P111c. 5-43. nepeAa•ta CIArHana ~la BHyTpmmeT04Hble pe1..1enTOpbl.

JI. CrrEU,M(()lflfHOCTh CHrHAJUI3AUHH

Ll;rn !1CCJJe).lOUaTCJ1eti, 11MelOW,11X npe).lCTaBJJeH11e 0 KOJJMl.Jecrne C11fHaJJbllbIX MOJJeKyJI, 0 COOTBeTCTBYIOW,eM KOJ1111.Jecrne peuenrnpou, o TpaHcMeM6paHHhIX Cl1CTel\1aX nepe,lla1.J11 Cl1rHaJIOB, BTOpH'IHbJXno-

cpe,llH!1KaX, OCTaeTC51 33ral!KOH, K3K npOTeHHKl1Ha3bl Bb!6Hp3!0T COOTBCTCTBYIOlUH~i cpepMCHT MeTa60JIHlJeCKOfO nyT11 .IL151 cj:>occj:Jopvtm1ponaH11s:i. f!lccJieao-

BaTen11 !JJJ5l o6b51CHCHl151 3TOfO 51BJICHl151 npeJJ.naraIOT «rnnOTe3y M11weH11» (OT aHrn. targeting hypothesis).

no 3TOH rnnoTe3e CilCIJl1cP114HOCTb 11pOTel1HKl1Ha3 11 cj:Joccj:>onp0Te11Hcj:>occj:>aTa3 ,r1,ocrnranc51 nyreM 06-

pa30BaHl151 KOMnapnvteHTOB Ha MeM6paHe, u COCTaU KOTOpblX BXO,ll51T He TOJJbKO C3Ml1 npoTel1HKl1Ha3bl 11 cj:Jocqmnp0Te11Hcj:Joccj:>arn3h1, HO 11 cneu11cj:Jw1ecK11e 6eJIK11-cy6crpaTb1. Ha;111411e ocrnTKa M11p11cT11Houol1 11JIH naJ1bMMT11Houotf Kl1CJIOThJ n cTpyKType 6eJIKOBcy6cTpmon -- ycJIOBl1e 11X «3a51KOpi1B3Hl151» B COOTBCTCTBYIOW,CM MeM6paHHOM KOMnapTMCllTC.

0)lH3KO B 60JibW11HCTBe CJiytJaeB npouecc aKnrnawrn KaKoro-m160 MeTa60JI111.JecKoro npouecca H3X0)111TC51 non Kot!TpOJieM He O)lHOH, a HCCKOJ!b- Kl1X Cl1CTeM BHyTp11KJICTOlJHOH c11rHaJI113au1111, nO'.:ffOMY B3)1(Hb!M cj:JaKTOpOM OTBCTa KJICTOK CJIY- )1(!1T LlJal1MOCB513b 3Tl1X Cl1CTCM.

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