підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

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ITOJI51PHhIM ClffHaJihHhIM MOJieKyJiaM OTHOe51T 6eJIKOBhie ropMOHhl (HarrpHMep, fJIIOKaroH, HHeyJIHH, napaTropMoH), tteifpoMe)l,ttaTOphI (ttarrpttMep, aueTHJIXOJIHH, fJIHUHH, y-aMHHOMaeJI51Ha51 KHCJIOTa), ¢aKTOPhl poem, UHTOKHHhl, 3ttK03aHOl1,l.l,hl.

B. PEQEIITOPbl

no JIOKaJIH3aUIIH pa3JIM'-laIOTMeM6paHHhie, UHTOITJia3MaTM'IeeKHe M 51)lepHbie peuerrTOpbl. no )l.pyroi1 KJiaeempttKaUHM see peuenTophI MO)!(HO pa3)],eJIHTh Ha 6b1eTpOOTBe'-1aIOw,He (B rrpe)l,eJiax MHJIJIHeeJ<YH)l) H Me)l,JieHHOOTBe'IalOIUHe, B npe- )l,eJiaX HecKOJibKMX MHHYT MJIH )l,a)!(e qacon, '-ITO xapaKTepHO )lJl51 ropMOHOB, nepe)l,aIOW,MX CHfHaJI Ha BHYTPHKJieTO'-IHhie peuenTOphI. PeuenTophl nepnoro Tttna - HHTerpaJihHhie OJittroMepHbie

6eJIKH, CO)l,ep)!(aU.lMe cy6be)l,HHHUY, HMeIOmYIO ueHTp )lJl51 eB513bIBaHM51 eHrHaJihHOH MOJieKyJihl l1 ueHTpaJibHblH MOHHhltt KaHaJI (pwe. 5-29).

PeuenTophI BToporo THna, JIOKaJIH30BaHHhie B MeM6paHax H He CB513aHHbie e KaHaJiaMH, noi:rpa3- )l,eJI51IOT Ha 2 60JihWHe rpynnhl: KaTaJJHTlf'leCKHe peu,enTOpbI, 06Jia)laIOWMe eo6eTBeHHOH THp03HHKHHa3HOH HJIM ryaHMJiaTUMKJia3HOtt aKTHBHOCThIO,

B3attMonei1eTByIOw,tte qepe3

JIOK e MeM6paHHhIM <I:>epMeHTOM. CB513hIBaHwe JIHraHlla (HanpwMep, ropMotta) c peuenTopoM Ha ttapy)!(Hott eTOpOHe KJieTO'-IHOHMeM6paHhl npHBO)l,HT K H3MeHeHHIO aKTHBHOeTH UHTOITJia3MaTH'-leeKoro <l:>epMeHTa, KOTOphIH, B CBOIO oqepe)lh, HHHUHHpyeT KJieTO'-IHhIH OTBeT, T.e. qepe3 MeM6paHy nepeHOeHTe51 HH¢opMaUH51, a He 3ap51)lhl MJIM KaKMeJIH6o paeTBopetttthJe MOJieKyJihL

B eJIY'-laeUHTOITJia3MaTtt'-leeKMXpeuenTopos 'Iepe3 MeM6patty npoxonwT ropMoH, a tttt¢opMaum1 0 npweyreTBHH ropMOHa B KJieTKe e ITOMOW,h!O peuerrTOpa nepe)l,aeTc51 B 51)],po.

Pa3JIH4HhJe KJieTKM opraHH3Ma B 3aswettMOeTH OT Bh!TlOJIHJleMblX HMM cPYHKUHH HMeIOT onpei:reJieHHhIH ·Ha6op peuemopos. B MeM6paHe onttotf KJieTKM MO)i(eT 6bITb 6oJiee )l,ee51TKa pa3HhIX THITOB peuenTopos. B3aHMO)l,etteTBy51 e peuenTOpoM, BHeKJieTO'IHhie XMMH'-leeKHe noepe)l,HHKH OKa3bIBaIOT BJIM51HHe Ha MeTa60JIH3M l1 cPYHKUlfOHaJihHOe eoeT051Htte (npOJittcl:>epaUH51, ceKpeUH51 H T.)l,.) KJie- TOK-MHWeHeH:.

I. Peu,enmopbt aiJpentl.!luna - aiJpenope11ienmopb1

A):J:pettopeuenTOphI pa3m1qa10T no pacnpe)J,e- Jiem110 B opraHH3Me - ueHTpMhHhie 11 rrep11- <t>epwqecKwe. UeHTpMhtthie a)J,pettopeuenTOphI, JIOKMH30Batttthie B pa3_1rnqHhIX o6Jiacrn.x M03ra, yqacrnyIOT B perym1um1 <l>YttKUMti UHC, nepw- <t>epwqecKwe - KOHTPOJIHPYIOT pa60Ty BHyTpeHHHX opraHOB.

Bee a)J,pettopeuenTOphI KJJaccw<t>wu,wpyIOT Ha )J,Ba TMTia - a- lI f3-, HO Ka)K)J,hlll THTI MMeeT HeCKOJibKO nO)J,THnOB, ttaH60Jiee pacnpocTpaHeHHhie H3 ttMx - a,-, a 2-, f3 1- w f32-peuenTophI. B 3aBHCHMOCTH OT CBOero aHaTOMH'leCKoropacrrOJIO.lKeHH51 KJieTKM O)J,HOfO TMTia, HanpHMep rna)J,KOMblllleYHhie KJieTKM cocy)J,OB HJIH a)J,HnOUHThI, co- )J,ep.lKaT pa3Hbie THnhI peuenTOPOB.

HecMoTp51 Ha 3HaqwTeJibHOe no)J,o6we Me)K)J,y a- H f3-peuenTopaMH H HX nO)J,TttnaMM, OHH KO- )J,HPYIOTCH pa3HblMH rettaMH. A):J:pettopeuerrTOPhl npMHa)J,Jie.lKaT K ceMeticrny 6eJIKOB, HMeIOiuwx 7 TpattcMeM6paHHhIX a-cnwpaneti (KoTOpbie npttHHTO tta3bIBaTb )J,OMettaMH). Ll,Jitttta N- H C- KOHUOB, a TaK.lKe )J,JIHHa 1-4 )J,OMeHOB pa3JTHqaeTC51 y pa3HhIX THnOB H nO)J,THnOB peuenTOpOB (pwc. 5-30).

A):J:pettopeuenTOpbl - rJIHKOnpoTeHHbI, BKJIIOqa10iune B CBOH COCTaB pa3JIHqHbie yrneBO)J,Hbie <t>parMeHTbl. fJIHK03HJTHpOBaHHIO nO)J,BepraIOTC51

pacnoJio)l(eHHbie B o6nacTH N-Kottua ocTaTKH acnaparttHOBotl KMCJTOTbl.

f3-A,ri,peuopeu,eUTOpbI BCTpeYaIOTCH rrpaKTH'leCKM BO Bcex TKaHHX opratttt3Ma. Kom111ecrno f3-Mpe- ttopeuenTopoB, npwxo)J,5IIUeec51 Ha KJieTKY, BaphHpyeT OT 300 )J,0 4000.

UeHTp CB513bIBaHHH a)J,pettanwtta o6pa3oBaH aMMHOKMCJIOTHhIMH OCTaTKaMH TpeThero, n51TOro H wecrnro )J,OMeHOB. Ll,pyrotl: <l>YHKUMOHaJibHO Ba)l(- HblM uettTp - o6JiaCTh B3aMMO)J,etl:cTBH51 c G-6eJT- KaMH, yqaCTBYIOIUHMH B <t>opMHPOBaHMH KJieTO'l- HOro ornern. OcrnTKH cepwtta H TPeOHttHa B o6nacTH TpeThero BHYTPeHHero )J,OMetta M C-Kottua a)J,pettopeuemopa MOrYT <t>oc<t>opmmpoBaTbC51 no)J, )J,eticrntteM npOTeMHKMHa3bl A MJIH cneutt<l:>HYecKotl KHtta3oti f3-Mpettopeuemopa. <t>oc¢opwnwpOBaHHe npMBO)J,HT K H3MeHeHMIO KOH<l:>opMaUHH peuemopa 11 cttM)l(eHHIO cpo)J,cTBa K G-6eJTKY MJIM npenITTcrnyeT CB513hIBaHHIO c G-6eJTKOM.

a-A,ri,peuopeu,eUTOpbI pa3JTHYaIOT no JTOKaJTH3aLl,HM (ttanpMMep, renaTOUHThI ttMeIOT a 1-peuenTopbI, a)J,HIIOJJ,HTbl - a2-a)J,pettopeuenTOPbl) l1 MexaHH3MY Tpattc<t>opMauMM 611onorttqecKoro cttrttMa. 3<t><t>eKTOpHbie CHCTeMbI, CBH3aHHbie c a,- H a2- a)J,pettopeuemopaMH, BKmoqaJOT G-6eJIKH pa3HOro TMna - Gplc-6eJTKH (G-6eJIOK CTMMYJTHpyIOIUHH) H Gi-6eJTKH (G-6eJIOK HHrtt6ttpyIOIUMti) H COOTBeTCTBeHHO <t>epMeHTbI - <t>oc<t>onwna3y c HJIH a)J,eHHJiaTUHKJTa3y.

2. Peu,enmopbt c mupo3U1tKUIUl311oii aKmU61tOCmb10

T11po311HOBh1e npoTe11HK11Ha3hI - cpepMeHThI,

¢>occpopwrnpyiom11e cneu,11¢>w-1ecKHe 6eJIKH no T11-

p03111zy, no,IJ,pa3,IJ,eJUIIOT Ha 2 nma - MeM6paHHhie

(peu,enTOpHhie) 11 u,11ToTIJia3MaTw-1ecK1-1e. BttyrpHKJieTO<JHhie T11p0311HOBhie npoTel1HKHHa3hl np11Hl1MaIOT yqacT11e B npou,eccax nepe,IJ,a<JH cHrHana B Mpo. Peu,enTOpHhie THP0311HOBhie npoTe11HKHHa3hI yqacTByIOT B TpaHCMeM6paHHOM nepe,IJ,a<Je c11rHaJIOB.

Ilp11MepoM peu,enTOpHOM T11p0311HOBOM npoTe-

11HKl1Ha3hI MO)l(eT CJIY)l(l1Th peu,enTop 11HCYJTl1Ha

(p11c. 5-31). Peu,emop 11Hcyn11Ha - THP0311HOBaH

npoTel1HKl1Ha3a, ¢>occpop11JTHPYIOIIIaH 6eJTKl1 no

0 H-rpynnaM T11po311Ha.

Peu,enTOp cocT011T 113 ,IJ,Byx a- 11 ,IJ,Byx P-cy6be-

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KaTanHTH<JecKHtl: ueHTP THP0311Hoootl: npoTe11HKHHa3hI HaxO,IJ,HTCH Ha BHYTPl1KJieTO<JHblX ,Il,OMeHax P-cy6be,IJ,11H11u,. B OTCYTCTB11e ropMoHa 11HcyJ111HoBh1e peu,emopbI He npoHBJTHIOT THP0311HKl1Ha3Hotl: aKTHBHOCTl1. Ilp11coe,IJ,11HeH11e 11HcyJT11Ha K u,eHTpy CBH3hIBaHIDI Ha a-cy6be,IJ,11Hl1IJ,ax aKT11B11pyeT cpepMeHT, ITpH<JeM cy6cTpaTOM CJT~T caMa THp0311HOBaH ITpoTe11HKHHa3a (p-cy6be,IJ,11H11IJ,b1), T.e. npo11cxo,IJ,11T ¢>occpop11n11poBaH11e P-cy6be,IJ,11H11IJ,hI ITO HeCKOJThKHM THP0311HOBhIM ocrnTKaM. <l>occpop11J111poBaH11e P-cy6be,IJ,11H11u, npo11cxo,IJ,11T ITO MexaHl13MY Me)l(MOJieKYAAPHOro TpaHccpoccpop11J111poBaHHH, T.e. O,IJ,Ha P-ueITb ¢>occpop11J111pyeT ,Il,pyry10 p-u,enh TOM )Ke MOJTeKYJThI peu,eITTopa. 3To, B cBoIO oqepe,IJ,h, ITPl1BO,Il,HT K 113MeHeHl1lO cy6cTpaTHOM CITeu,11¢>11qHOCT11 THP0311HOBOM npoTel1HKl1Ha3hI; TeITepb OHa CIToco6Ha ¢>occpop11JI11pOBaTh ,IJ,pyme BHyTpHKJieTO<J- Hhie 6eJTKH. AKTHBau,1151 H H3MeHeH11e cITeu,11cpmHoCTH o6ycJIOBJTeHbI KOHcpOpMaIJ,110HHbIM11 113MeHeIDIHMl1 peu,enTopa 11HCYJTl1Ha nocJie CBH3hIBaHl1H ropMoHa 11 aYT0¢>occpopIDIHpoBaH11H.

Kmo<JeBotl: 6eJTOK, ¢>occpop11JTHpyeMb1tl: THpo311HOBotl: npoTel1HKHHa3otl:, - cy6cTpaT 11HCYJil1HOBOro peu,enTOpa-1 (oT aHrn.

<l>occpopHJTttpOBaHHbitl: aKT11B11pyeT cpepMeHTbI, HaITp11Mep T11p0311HOBYIO cpoccponpoTe-

PMC. 5-31. AKTMBal..IMSI pe1..1enTopa MHCynMHa - TMp03MHOBOM

npoTeMHKMHa3bl.

HHcpoccpaTa3y, 11 6eJIKH, yqacrnyiom11e B peryAAIJ,HH KJieTO'-IHbIXnpou,eccoB.

)],ecpoccpop11J111poBaH11e peu,emopa ITO,IJ, ,IJ,etl:cTBHeM THP0311HOBotl: ¢>occpoITpoTe11HcpoccpaTa3bI B03BpamaeT era B HeaKTl1BHOe COCTOHHl1e. CpO,Il,CTBO peu,eITTOpa K MHCYJil1HY CHH)l(aeTCH np11 era ¢>oc- ¢>op11JT11poBaH1111 np0Te11HK11Ha3otl: A no aMHHOKl1CJIOTHhIM OCTaTKaM cep11Ha 11 TpeOHl1Ha.

3. Peu,enmopbt c zya11U11amu,uK.1u1311oii aKmlUJllOCmblO

fyaHHJTaTIJ,HKJia3a KaTaJIH3HpyeT o6pa30BaHHe u,fM<I> 113 IT<l>, O,IJ,Horo 113 Ba)l(HhIX nocpe,IJ,H11KOB BttyrpHKJieTO<JHOM ITepe,IJ,a'-111 CHrHaJia (pttc. 5-32, 5-33). fyaHHJTaTIJ,HKJia3a HaxO,IJ,HTCH B KJieTKe, KaK B MeM6paHHOCBH3aHHOM COCTOHHl111, TaK H B IJ,11T030JlbHOM.

CooTHOIIIeHHH 3THX ,IJ,Byx cpopM cpepMeHTa B pa3- Jil1'-IHhIXTKam1x pa3Hoe. Hanp11Mep, B KJieTKax TOH-

OH OH

Pitc. 5-32. 06pa30BaHHe 3',5'-l..lltKlllt"l9CKororMci> (1..1rMci>).

nenrnAHbllil

/ropMOH

OKC~ aaorn (NO)

+--NO

j

AKrnsa1.11-1r1

np0Te1-1HK1-1Haab1 G

Pitc. 5-33. Perynsi1..11tsi aKTl4BHOCTH MeM6paHHolii (1) 1t L11tTo30nbHOlii (2) ryaH1tnaTL11tKna3bl.

Koro KMllle'!Hl1Ka90% ryamrnan.1,11KJia3bI Haxo,L1,11TCH B MeM6paHaX, a B JierKl1X l1 nelJeHH - JIHUJb

20%. UttT030JihHaH l1 MeM6paHHOCBH3aHHaH ryammam11KJia3hI pa3JI11qa10TCH He TOJlbKO no JIOKaJll13aUHH, HO l1 no MOJieK)'JIHPHOH Macce, aKTl1BHOCTH, cnoco6y peryJIHUl111.

U11T030JihHaH <PopMa ryaHJ1JlaTUHKJia3bI cocTOl1T 113 ,ll,Byx cy6be,ll,HHl1U (a l1 ~) l1 CO,ll,ep:)lGiT B CBOeM

ryaH1maTL1111KJ1a3a

O=P - O

I_

0

cocrnre rrpocTent'!eceyJOrpynrry - reM. B o6JJacTH reMa CBH3bJBaeTCH aKTHBaTOP 3TOH <PopMbJ ryaHHJJaTJ.J,HKJla3bl - OKCH,ll, a30Ta (NO), o6pa3YJOIUHHCH 113 aprnHHHa JIO,ll, ,ll,eHCTBHeM <PepMeHTa CHHTa3bI OKCH,ll,a a30Ta (CM. pa3,ll,eJJ 9).

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B KJJeTKax TKaHert BbIHBJieHbI 3 OCHOBHbIX THna BHYTPl1KJJeToqHbIX peuenTopHbIX 6e11KOB, c KOTOpb1M11 B3aHMO,ll,ettcTByeT ufM<l>: ufM<l>-3aBl1Cl1MaH npOTe11HKHHa3a (npoTel1HKHHa3a G), ufM<l>-pery- n11pyeMbie 110HHbie KaHaJibl 11 ufM<J>-pery1111pye- MaH <Poc<Po,L1,113cTepa3a, cneu11<Ptt'-IHaH K uAM<l> (KaTan11311pyeT npeBpameH11e uAM<l> B AM<l>).

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peuenTOp3aBHCHMOro nOCTYJ1JleHlUJ 3TOro ttOHa B UHTOnJia3MY KJJeTKH. 3TH ::i¢¢eKThl onocpe,UOBaHbI LJ:ettCTBHeM npoTettHKHHa3hI G Ha MeM6paHHhie 6eJIKH, Y<JaCTBYJOlllHe B 06MeHe Ca2+.

B. CTPYKTYPHO-«I>YHK11;1fOHAJihHMI

oPrAHM3AU,IUI G-BF.JIKOB

G-6emrn (IT<l>-cBH3hrna10mue 6eJIKtt) - ytmBepcaJihHhie nocpe,UHHKH npH nepe,Uaqe cttrHMOB OT peuenTOpOB K ¢epMeHTaM KJieTOlJHOM MeM6paHhl, KaTaJIM3HPYIOIUHM o6pa30BaHHe BTOpH'-IHhIX nocpe,UHHKOB ropMOHaJihttoro cttrHaJia. G-6en- KH - OJIHroMepbl, COCTOHIUHe 113 a, p H y-cy6be- LJ:MHMU. CocrnB LJ:MMepoB py He3HalfwTeJihHO pa3JIH'IaIOTCHB pa3HhIX TKaHHX, HO B rrpe,UeJiax OLJ:HOM KJieTKtt Bee G-6enKM, KaK npaBMJIO, wMeIOT OLJ:M- HaKOBhitt KOMilJleKT py-cy6beLJ:MHHll. no::iToMy G- 6eJIKH npHHHTO pa3JIMlfaTh no MX a-cy6be,UMHM- uaM. BhrnBJieHo 16 rettoB, KOLJ:MPYIOIUHX pa31IttlfHhie a-cy6beLJ:HHHllhI G-6eJIKOB. HeKOTOpb1e M3 reHOB MMeIOT 6onee O,UHOfO 6eJIKa, BCJie,UCTBMe aJihTepHaTHBHOro cnJiai1c11Hra PHK.

Ka)((LJ:aH a-cy6beLJ:HHMUa B cocTaBe G-6eJIKa ttMeeT crreump11qecKMe uettTpbr:

•CBH3hIBaHH5I IT<l> HJIH fLl<l>;

•B3aHMO,UeMCTBH5I c peuenrnpoM;

•CBH3bIBaHH5J c py-cy6be,UMHMUaM11;

•¢oc¢opHJittpOBaHHH no,U ,UeMCTBtteM npoTeHHKHHa3hl C;

•B3aMMO,UettCTBHH c ¢epMeHTOM a,UeHMJiaTUMK-

Jia30tt HJIM ¢oc¢0Jiwna3oi1 C.

B cTpyKrype G-6eJIKOB OTCYTCTBYIOT a-cnupaJih- Hhie, rrpotttt3hIBaIOIUMe MeM6paHy LJ:OMeHhI. G-6eJI- KH OTHOCHT K rpyrrne «3aHKOpeHHbIX» 6eJIKOB (pttc.

Pery.JlllUHJI aKTueuocru G-6eJIKoe

Pa3JittlfaIOT HeaKTMBHYIO ¢opMy G-6eJIKa - KOMITJieKC apy- f)],<J> M aKTHBHpOBaHHYIO ¢opMy apy-fT<l>. AKTMBauua G-6eJIKa rrpowcxo,UMT rrptt B3aMMO,UettCTBHM c KOMnJieKCOM aKTHBaTOp-peuen- TOp, M3MeHeHwe Kott¢opMauww G-6enKa CHtt:x<aeT cpO,UCTBO a-cy6be,UMHMUhI K MOJieKyJie fLl<l> H yseJIHqlfBaeT K IT<l>. 3aMeHa fLl<l> Ha IT<l> B aKTHBHOM uettTpe G-6enKa HapywaeT KOMnJieMettrnpHOCTh Me)!(LJ:Y a-fT<l> 11 py-cy6be,UMHHUaMtt. PeuerrTop, CBH3aHHhii1 c cwrttaJihHOi1 MOJieK)'11oi1, MO)!(eT aKTHBHPOBaTh 60JibIUOe KOJIH'-IeCTBOMOJie- KYJT G-6eJIKa, TaKMM o6pa3oM o6ecrre'-fttBaHycu- JTeHtte BtteKJieToqttoro cwrttana Ha 3TOM ::iTane (pwc. 5-35).

AKTHBHpoaaHHaB a-cy6beLJ:HHHUa G-6eJIKa (a- fT<l>) B3aMMOLJ:ei1crnyeT co crreuu<Pw'!ecKHM

I

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