підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin
.pdfY'IE.l>HHK ~JIR BY30B
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113,lJ:ATEJJbCKHH IJ:OM
r30TAP-ME~
2004
Y)lK 577.1(075.8)
EEK 28.072H73
E63
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3aB. Kacpe.upoit 6MOXliMMM c KYIJCOM KJIMHMqecKoit na6oparnpttoit .u11amocTHKM AcrpaxattcKoit rocy.uapcrnettttoi' Me.UMUMHCKOM aKa.ueM11.11, KaH.Il. Me.a. ttayK, npocp. !{.M. HuKyJluHa
3aB. Kacpe.upoit o6rueti: 11 KJIMHMqecKoit 6110XMMMM NQ 1 PocTOBCKoro rocy.uapcrnettttoro Me.UMUliHCKOro yttMBepc11TeTa,
.uoKT. 6110JI. ttayK, npocp. 3.11. MuKautuHoeu11
3aB. Kacpe.upoit o6rueti: 11 KJIHHMqecKoit 611oxMMMM NQ2 PocTOBCKoro rocy.uapcrnettttoro Me.UHI.!MHCKoro yttMBepc11TeT2
JI.M. IlycmOBaJlOBa
ABTOPCKMM KOJIJieKTMB
AneiiHuKoBa T.JI., Aeaeeea JI.B., AHapuaHoea JJ.E., EeJ1yutKUHa H.H, BoJ1Koea HII., Bopo6beBa C.A., I'oJ1eH11eHKO B.A., I'y6apeea A.E., Kop.!lRKOBa O.B., Jluxa11eea H.B., IlaBJlOBa HA., Py614oea I'.B., Cwiaeea C.A., CwiyRHOBa C.H., Tumoea T.A
663 JiHOXHMHJI: Yqe6HMK I no.a pe.u. E.C. CeBepMHa. - 2-e 113,ll., MCTip. - M.: f30TAP-ME)l, 2004. - 784 c.: MJI. - (Ceprni «XXI BeK»).
ISBN 5-9231-0390-7
B yqe6tt11Ke paccMorpettbI octtOBHbre noJio)!(eHMH KJiacc11qecKoit 611ox11MMM. IlpMBe.uetthr cBe.uettMH o CTPYK· zype M CBOMCTBaX 6MOMOJieKYJI, 6M03HepreTMKe, MOJieKYJIHpHbIX OCHOBaX cpM3MOJIOmqecKMX <PYHKUMM qeJIOBe· Ka. PacCMOTpetthI 6MOXMMMqecKHe oco6eHHOCTM Ba)!(HeMIIIMX opraHOB H TKaHeM. l13JIO)!(eHbI COBpeMeHHbH npe.ucrnBJieHHH 0 MOJieKYJIHpHbIX OCHOBax ttapyweHHM npM pH,Ile naTOJiomqecKHX COCTOHHMM M 6oJie3HeM.
Yqe6ttMK npe,utta1ttaqett .IlJIH czy.uettTOB Me.umrMHCKMX BY30B, acn11pattTOB.
Y)],K 577.1(075.8)
EEK 28.072H73
Ilpaea Ha aaHHOe u3amme npuHaaJleJICam u3aameJlbCKOMY aoMy «I'30TAP-ME/f>. BocnpOU3BeaeHue u pacnpocmpaHeHU4 B KaKoM 6b1 mo HU 6b1Jw euae 11acmu wiu 14eJ1020 u3aaHUR He Mo2ym 6bJmb ocyU1,ecmBJ1eHbt 6e3 nucbMeHH020 pa3peuteHw U3aameJlbCK020 aoMG.
ISBN 5-9231-0390-7 |
© |
l11.uaTeJibCKMM .UOM «f30TAP-ME)l», 2004 |
© |
KonneKTMB aBTOpoB, 2004 |
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CesepHH EsreH11rr Cepreem1q, .UOKT. XHM. HayK, npo<J:>., q]J.-Kopp. PAH, 3as. Ka<J:>e.uporr 611oxHMHH MMA HM. 11.M. CeqeHoBa, reH. .UHpeKTop OAO «BcecoI03Hhitt HayqHblll u.eHTP MOJieKYJHIPHOll L(HarHOCTl1KH 11 JieqeHHH»' pe.uaKTOp lf3L(aHHH.
AileifHHKoBa TaThR:Ha JieoHttaOBHa, KaH)l. 6ttoJI. HaYK,
.uou.. Ka<l:>e.uphI 6ttoXHMl11f MMA 11M. l1.M. CeqeHoBa (pa3L(eJI 7), OTBeTCTBeHHblll aBTOP H3L(aHHH.
AB,11,eesa JlIO.UMttna BHKToposHa, KaH)l. 6110JI. HayK,
.uou.. Ka<J:>e,11,pb1 6HonornqecKorr XHMlfH MMA HM. 11.M. CeqeHosa (pa3.UeJihI 6, 11).
AJmpuaHoBa JlIO.UMlfJia EsrettbeBHa, KaH)l. 6HoJI. HayK, crnpumi1 rrperro.uasaTeJih Ka<J:>e.uphI 6ttonornqecKoi1 xttMlflf MMA ttM. 11.M. Ceqettosa (pa3.uenh1 12, 15).
lieJQ'IUKHHaHaTaJIMI HttKonaesHa, .uou.. Ka<J:>e.uph1 fo.roXHMHlf MMA HM. l1.M. Ceqettosa, yqeHbIH ceKpernph Hl1l1. MoneKYmrpmm Me.UHU.ttHa MMA lfM. Ceqettosa (pa3.uen 2).
BOJIKOBa HaTaJIMI neTPOBHa, KaH)l. MeL(. HaYK, L(Oll,. Ka<J:>e.uphI 6HonornqecKoi1 XHMHH MMA HM. 11.M. Ceqettosa (pa3.uen 1).
fOJiell'leHKoBepa AneKcaH)lpoBHa, Katt.LI. 611QJJ. HaYK, .uou.. Ka<l:>e.UPhI 611onorttqeCKott XllMHlf MMA ttM. l1.M. CeqeHOBa (Pa3.lleJihI 4, 5).
Bopo6beBa Csernatta AttaTOJibeBHa, KaH.u. 6tton. HayK, L(OI..(. KaQ:>e.uphI 6HonornqecKoli XHMlflf M MA HM. 11.M. Ceqettosa (pa3.uenh1 7, 11).
I)6apesa AneKcaHL(pa EsreHheBHa, KaH.LI. Me.u. HaYK, .uou.. Ka<l:>e.UPhI 6HoJiornqecKorr XlfMllH MMA 11M. 11.M. CeqeHOBa (pa3.UeJI 8).
KoPJUIKOBa Onbra BeHHaMHHOBHa, KaHL(. 6HoJI. HaYK, crnpwttrr rrpeno.uasaTeJih Ka<l:>e.uphI 6ttoJiornqecKoi1 XHMHH MMA ttM. 11.M. CeqeHosa (pa3.uen 9).
Jluxa'lesaHHHa BttKToposHa, KaHL(. 6ttoJI. HaYK, .uou..
Ka<l:>e.uphI 6ttonornqecKoi1 XlfMlfH MMA HM. 11.M. Ceqettosa (pa3.uen 9).
IlaBJio&a HHHa AneKcaHL(poBHa, KaH)l. 6110JI. HaYK, .uou..
KaWe.UPhI 6HonornqecKott XlfMHH MMA lfM. 11.M. CeqeHOBa (pa3L(eJihl 3, 6).
Py6u,oBa ranttHa Bac11JiheBHa, KaH)l. 6ttOJI. HaYK, L(Oll,. Ka<l:>e.uphI 6ttonornqecKoi1 x11MttH MMA HM. 11.M. CeqeHoBa (pa3.ueJibI 3, 6).
Cm1aeBa CsernaHa AneKceesHa, .UOKT. 6110JI. HayK, npo<J:>. Ka<J:>e.uphr 611onornqecKorr Xl1Mlflf MMA HM. 11.M. CeqeHosa (pa3.uenbr 4, 10, 16).
CHJIYHHOBa CseTJiaHa HttKonaesHa, KaH.u. 6HoJI. HayK, .uou. Ka<l:>e.uphr 6HonornqecKorr XHMl111 MMA HM. 11.M. CeqeHOBa (pa3.UeJihI 12, 13, 15).
TuTo&a TaTMma AneKceesHa, KaHL(. 6110JI. HaYK, crnpwHtt rrperro.uasaTeJih Ka<l:>e.uphI 6HoJiornqecKoi1 XlfMHl1 MMA HM. 11.M. CeqeHosa (pa3.UeJihI 13, 14).
nPEJlVICllOBVIE
DHOXHMIDI - cpaBHITTeJibHO MOJIOL(ruI HaYKa, B03HHKwm1 Ha CThIKe 6ttonornH H mM11H B KOHU,e XIX seKa. Otta H3yqaeT rrpou.eccbI pa3BHTIDI H <l:>YHKU,HOHHPOBaHIDI opraHH3MOB Ha H3hIKe MOJieKYJI, CTPYKTYPY H XHM11qecKlfe rrpou.eccbI, KOTOphre o6ecneqHsaIOT )l(J13Hh O.UHO- H MHOroKJieToqHhIX cywecTB, HaceJIHIOIU,11x 3eMJIIO. BhmaIOm11ecH OTKpbITlfH B o6JiaCTH yqeHIDI 0 <J:>epMeHTax, 6HOXl1Ml1qecKOH reHeTl1Klf, MOJieKYJIHPHOH 6110JIOrttl1 11 61103HepreTHKlf rrpespaTlfJilf 611ox11MHIO B <l:>YHaaMeHTaJibttyIO
.u11cu.11nmrny' II03BOJIHIOIU,YIO peuraTh MHOr11e BCUKHbie rrpo6JieMbl 6HOJIOflfl1 11 MeL(l1ll,11Hhl.
HacTOHIU,11rr yqe6HHK HBJIHeTCH pe3ynhTaTOM MHOroJieTHei1 npeno.uasaTeJihCKOtt pa6oThI KOJIJieKTl1Ba Ka<l:>e.uphI 611ox11MH11 MocKOBCKOtt Me.u11u11HCKOtt aKa-
.ueMH11, 11 co.uep)l(11T ttH<J:>opMau.mo, Kaca10myIOCH, rnasHhIM o6pa30M, 611ox11M11tt qenoseKa. Yqe6H11K npenHa3HaqeH cry.ueHTaM 11 nperro.uasaTemrM Me.UHU.HHCK11x BY30B. no CTPYKType 11 conep)l(aHl1IO OH COOTBeTcTByeT rrporpaMMe no 611ox11M1111 .UJIH cry.ueHTOB Me.u11u.11HcK11x sy3os, yrnep)l(L(eHHOtt M11HttcTepcTBOM 3.Upasooxpattett11H P<l>.
Yqe6HttK 3HaKOMITT q11rnTenei1 co CTPYKTYPHO-<l:>YHK- u.110HaJihHhIMH KOMIIOHeHTaMH KJieTOK 11 rrpou.eccaMl1, Jie)l(aIUl1MH B OCHOBe )l(J13HeL(eHTeJibHOCTl1 3.llOPOBOfO opraHl13Ma, a TaK)l(e c HeKOTOpbIMl1 HapyweHIDIMl1, KOTOPbie np11BOL(ITT K B03Hl1KHOBeHl1IO 6oJie3Hett. ~ Hero xapaKTepHO aKU,eHT11pOBaHHe BH11MaHID1 qmaTeJIH Ha 3HaqeHl111 paccMaTP11BaeMhIX oorrpocos L(JIH Me.u11u11Hhl. Kirnra co-
.uep)l(JiT 16 pa3L(eJIOB 11 IIplfJIO)l(eH11e, B KOTOpoe BXOL(HT CIIl1COK COKpaIU,eHl1H, rrpeL(MeTHhIH YKaJaTeJih, CJIOBapb repMHHOB H Jia6opaTOpHhie IIOKa3aTeJil1. IlepBhie pa3)leJihl nOCBHIU,eHhI CTPYKTYPe H <l>YHKlll1HM 6eJIKOB, <J:>epMeHTOB, BttraMl1HOB 11 HyKJiel1HOBbIX Kl1CJIOT. CTo;:ipo6HO paCCMaTp11BaeTCH Cl1HTe3 11 peryJIHU,IDI lfH<l:>opMaU,110HHOfO IIOTO- Ka .UH.K-?PH.K-?6eJIOK, rrp11q11Hbl, Jie)l(all..ll1e B OCHOBe 6ttOXJ1MttqeCKOH 11HL(HBl1L(yaJibHOCTl1 opraHl13MOB 11 B03HHKHOBeHIDI HaCJie.UCTBeHHhIX 6oJie3Hett. B pa3L(enax no o6MeHy yrneBOL(OB, Jil11Itt)lOB H aMl1HOKHCJIOT BHHMaH11e qHTaTeJiett <J:>oKycwpyeTCH Ha npoueccax, o6ecneqmiaro- ll.(11X 06pa30BaH11e 11 rr0Tpe6nett11e 3Heprn11 B TKaHHX opraHl13Ma 11 yqacTHe 3Tl1X coe.u11HeH11tt B <J:>opMttpOBaHl111
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CT]JYJCTYPHbIX KOMnoHeHTOB KJieTOK. B KHJilre o6c)')K)J,aeTCH KJIK)'-leBaHpOJib ropMOHOB B Me)!(KJleT04Hh!X B3aHMO- ,netl:CTBJi!HX 11 peryJii:iu,1111 o6MeHa Bemecrn. Y4e6HHK co- ,nep)!(HT TaK)!(e pi:i,n cneuttaJIHJHPOBaHHhIX paJ,neJios: 6HOXJi!MHH Me)!(KJieT04HOrO MaTp11Kca, o6e3Bpe)!(ttBaHJile TOKCl14eCKl1X BemecTB B opraHH3Me, 6110XHMHH KpOB11, OHKOreHe3, npe,nCTaBJH!K)llll1X OC06bltt 11HTepec ,UlliI Me,UHKOB 11 Bpa4eti. Hmj:JopMaumr, npttBe,neHHaH BO Bcex rnaBax )"le6HHKa, ,natta B COOTBeTCTBHl1 c COBpeMeHHhJM YPOBHeM Ha)"lHblX 3HaHHH B 3Tl1X o6Jiacrnx.
Ha,neJOCh, 4TO )"le6tt11K JattHTepecyeT urnpoKHtl: Kpyr 4ITTaTeJietl: 11 OKIDKeTCH noJieJHhlM ,UlliI cry,neHTOB, acrn1paHTOB 11 H3)"lHhlX corpy,UHl1KOB, pa6oTaJOllll1X B o6JiaCTl1 o6metl: H KJIJilHJil'-leCKOll6ttOXHMHH, MOJieeyJIHpHOH 6HOJIO-
rn11 H Me,UHUHHhl. EoJibllJQe KOJill'-leCTBOcxeM 11 p11CyttKOB, KOTOpblM11 CH36)!(eHa KHlffil, ,UOJI)!(Hbl o6JieI"IJi!Tb YCBOeH11e Marep113Jla 11 MOryr 6b1Tb 11cnOJih30BaHbl B nperro,naBaHl111 6110XHMHH. B TO )!(e BpeMH KOJIJieKTHB aBTOpoB roroB pacCMOTpeTh rrpetvJO)!(eHHH no YJIY'lllleHHJOKHl1lli 11 Kpl1Tl14eCIG1e JaMe'-laHJi!H'-IHTaTeJiett,KOTOphie MO)!(HO BhICJI3Tb no 3JieKTpoHHotl: no4Te: info@geotar.ru.
oJiaro,napJO KOJIJieKTttB aBTopoB H oco6etttto rpynny B cocraBe T.Jl. A.JiettHHKOBotl:, Jl.B. ABlleeBott, H.fl. BoJIKOBotl:, B.A. foJieH'-leHKO,A.E. fy6apeBoi1 H C.A. CHJiaeootl: Ja nOMOlllb B pe,naKUHOHHOH pa6oTe, a TalOKe coTJJYllHl1KaM HJ)laTeJibCKOro llOMa «f30TAP-MELI,».
3aa. Ka¢. 611ox11M11H MMA HM. 11.M. Ce- '-lettoaa,lffi.-Kopp. PAR, rrpo¢.
Cnfl1COK COKPAIUEHfl1fll
*UJIU # - c nocJiellYIOlllHM KO,UOM H3 6 umj:Jp (CHMBOJihl
*uJ1u # yKaJbJBaJOT Ha H3J1114tte aJIJieJietl:, pa3HhIX <j:Je-
HoT11noB Ja6oJieBaHHH HJIH )!(e BKJIJO'-leHtte B COCTaB H030JIOfH4eCKOH e,UHHHUhl HeCKOJibK.11X H pa3HbIX nopa)!(eHHblX reHOB) - MeH,UeJieBCKOe H3CJie,noBaHHe (rro http://www.ncbi.nlm.nih.gov/Omim/).
¢o> - CHHOHl1M
9\ - ayrOCOMHOe llOMl1H3HTHOe HaCJie,UOBaHJile p - ayrOCOMHOe peueCCHBHOe HaCJie,UOBaH11e
N - CBHJattttoe c X-xpoMOCOMotl: ttacJie,noBaHtte
B-KJieTKH (B rrpo11JHOCHT KaK 63) - B-JI11M<j:Jou11ThI Cl, C2, C3 (rrpoHJHOCHT KaK cu) 11 T.,n. - KOMnotteHThI
CHCTeMbl KOMTIJieMeHTa 1, 2, 3 H T.)l.
Ca2+ - KaT11ott(b1) KaJihUHH, 110H(b1) KaJihUm1; 110HttJ11poBaHHh1tt (CB06ollHhltt) K3Jlhll11H
CD (oT cluster of differentiation [npottJHOCHT KaK cu au], KJiacTep ,n11<j:J<j:JepeHUHPOBKH), CM. MapKep
CJ- - aHHOH(bl) XJIOpa
Fab CM. «<l>parMeHT»
FAD - <j:JJiaBttHa,nett11HllHHyKJieOTHll
FMN - <j:JJiaBttHMOHOHyKJieOTHll
H+ - 110H(h1) BO,nopo,na, npoTOHhl
[H+j - KOHUeHT)JaUHJI HOHOB BO,Uopo,na Hb - reMorno6ttH
HbCO - Kaj:!60Kc11reMomo6ttH
Hb02 - reMorno6ttH oKcttrettttpoBaHHhitt
HLA (npottJHOCHT KaK 3U'I 3J/b 3ii, OT human leukocyte antigens), CM. «AHTttreH», CM. «MHC»
Jg - HMMYHOfJI06yJIHH, HMMyHOrno6yJil1Hhl
IRE - iron-responsive element, )!(eJieJ0-4YBCTBttTeJih- HhIH 3JieMeHT
K+ - K3THOH(bl) K3Jll1H
[K+] - KOHUeHTpaUHH HOHOB KaJIHH LT - leucotrienes, JiettKOTptteHhI MetHb - MeTreMornofarn
MHC (rrpo11JHOCHT KaK 3M 3U'I cu, OT major histocompatibility complex, rnaBHbIH KOMTIJieKC lliCTOCOBMeCTHMOCTH)
M, - Ka)!(yuiai:ici:i MOJieKYJIHpHaH Macca
Na+ - KaTttoH(hI) ttaTpIDI
[Na+j - KOHUeHTpauttH 110HOB HaTpHH
NAO - HHKOTHHaMH,Ua,nettttH,U11HyKJieOTHll
NADP - HHKOTttHaMH,Ua,nett11H,nttH)'KJleoTH,U<j:Joc<j:JaT NO - OKCHll aJora (NO), Bh1pa6aTb1BaeMhIH B 3HllOTe-
JIHH <j:JaKTop peJiaKcau1111 (BaJO,UHJiaTaUHH) HT<!> - ttyKJieOJHllTpH<j:Joc<j:JaThI
P.co2 - rraputtan:httoe ttarrpi:i:)!(etttte ,nByoKHctt yrJiepo- ,na B apTepttaJibHOH KpOBH
pC02 - rrapu11anhttoe ,naBJiett11e llBYOKHCH yrnepo,na PG - prostaglandins, npocrarnaH,UttHhI
Pi (H 3P04) - <j:Joccj:JaT HeopraHtt'-leCKHH p02 - naputtaJlhHoe ,naBJieHtte KHCJiopo,na
PPi (H4Pp1) - rrttpo<j:Joc<j:JaT tteopraHH'-leCKttti: SAr - S-a,nettoJHJiroMou11cTe11tt
SAM - S-a,nettoJ11JIMeT110HttH
T3 - TPHHO,UTl1POHl1H
T 4 - TeTpatio,nT11pott11H, TttpOKCl1H
T-KJieTK.11 - T-JIHM<j:JouttThI
TNF - tumor necrosis factor, <j:JaKTop HeKpoJa onyxoJiett
TX - TpoM6oKcattbI
VIP (OT Vasoactive Intestinal Polypeptide) - BaJoaKTHBHhlH 11HTeCTHH3JlbHbltt (K11llle4Hb!H) noJittnenTHll (He,norrycTttMo ttan11catt11e - Bl1fl)
Ar - attTttreH, attnffeHhI
7
All - |
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apTepHaJJbHoe ,llaBJieHwe |
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Allf - aHTH,UHypeTwqeCKHM ropMOH (Ba3orrpeCCHH) |
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A,llef> - |
a,lleH03HH.llH<Poc<t>opttaH KHCJIOTa, a,lleH03HH,llH- |
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<t>oc<t>aThI |
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AKTf - |
a,llpettoKopTHKOTponHbIM ropMoH |
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AJIT |
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aJJatt1rnaMHHOTpaHc<Pepa3a |
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AMCf> |
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a,lleH03HHMOHo<t>oc<t>aT(b1) |
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u.AMCf> |
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UHKJIHqecKHM a,lleH03Htt-3',5'-MoHo<t>oc<t>aT |
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anoJIIl |
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arroJiwnorrpoTeHH |
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Aflef> |
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attrwoTeH3Htt-npeBpaiua10ruwit <t>epMeHT |
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ACT - |
acnapTaTaMHHOTpattc<Pepa3a |
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AT - |
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attTHTeJio, attTHTeJia |
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ATCf> |
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a,lleH03HHTpw<tJoc<t>opttaH KHCJIOTa |
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AT<D-a3a - |
a,lleH03HHTpw<tJoc<Parn3a |
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AU - |
a,lleHHJiaTUHKJia3a |
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AXAT |
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aueTHJI-KoA-xoJiecTepoJiauHJITpattc<Pepa3a |
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BMK - |
BhICOKOMOJieKyJIHpHbre KHHHHoreHbl |
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r AMK - |
y-aMHHOMaCJIHHaH KHCJIOTa |
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fLl,Cf> |
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ryaH03HH.lltt<Poc<PaT |
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rM K - |
rJia,llKOMbillieqHaH KJieTKa |
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fMCf> |
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ryaH03HHMOHo<t>oc<t>aT |
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[fl3TE |
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rH,UporrepOKCH)l3MK03aTeTpoeHOaTbI |
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rT - |
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rnyTaTHOHTpattc<tJepa3a |
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IT<D - |
ryaH03HHTpw<Poc<t>aT |
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f3TE |
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fH.llPOKCH3MK03aTeTpOeHOaTbl |
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LI. - |
llaJ!bTOH (rroCJie qwcJIOBOro 3HaqeHHH) |
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Ll,A [ |
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,llHaUHJirJIHUepoJibl |
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n:e30KCHpH6ottyKJieHHOBaH KHCJIOTa |
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Ll,Oef>A |
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,llHOKCH<t>eHHJiaJiaHHH |
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Ll,Cf><D |
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)KKT - :JKeJiy,lloqHO-KHIIIeqHb!M TpaKT |
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HJI - |
HHTepJieMKHH, HHTepJieMKHHhI |
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HM<l> - |
HH03HHMOHo<Poc<PaT |
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11©3 |
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HH03HHTp11<tJoc<t>aT |
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11<DH - |
11HTep<t>epoH, HHTep<tJepOHbI |
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K,l( - |
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KHJIO,llaJibTOH |
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KK - |
KpeaTHHKHHa3a |
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KoA - |
Ko<t>epMeHT (Ko3H3HM) A |
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KoQ - |
Ko<PepMeHT (K03H3HM) Q |
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KWP - KHCJIOTHo-ureJioqttoe paBttosecwe |
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K<D |
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KJiacc11<PwKaUHH <DepMeHTOB (<http: //wwW. |
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expasy.ch/sprot/enzyme.html> ). K<l> rrpwse;:i:ettbI rro Enzyme Nomenclature (NC-lUBMB, KoMHTeT rro HoMeHKJiaTYPe Me)[(,Ify'Hapo;:i:ttomCo103a no EwoxHMHH H MoJieKyJIHpttoi1 EwoJiorn11)
Jff - JIIOTeHHH3HPYIOIUHH ropMOH, JIIOTpOIIHH JI,l(r - JiaKTaT.llem;:i:poreHaJa
Jin - Jil1fIOflpOTeHHbl
JIIlBTI - JIHnonpoTeHHbl BbICOKOM illIOTHOCTl1 JIIl-JiwrraJa - JIHnonpoTeHHJ111na3a
JIIlHil - JIHnonpoTeHHbl HH3KOM illIOTHOCTH JlITOHTI - JIHIIOnpoTeHHbI oqeHb HH3KOM ITJIOTHOCTl1 Jinnn - JIHnonpoTeHHbl npoMe)!()'ToqHoMITJIOTHOCTH JIXAT - JieuHTHHXOJiecTepoJiauHJITpaHc<t>epa3a
MAf - MOHoaueHJirnwuepotthr
MAO - MottoaMHHOOKcH,Ua3a OTIK - o6urwit nyTb KaTa6oJIH3Ma
MHPHTI - MaJibie H;:i:epttbre pw60HYKJieorrpoTeHHhI TITr - rrapaTwpeoH.Utthli1 ropMoH
CE - cy6be;:i:wttwua
ITKA - npoTeHHKHHa3a A ITKC - rrpOTeHHKHHa3a C
IlOJI - rrepeKHCHOe OKHCJieHHe JIHilH)lOB IlOMK - rrpoorrHOMeJiaHOKOpTHH
TI<D - rrHpH,UOKCaJih<Poc<PaT
ITUP - noJIHMepaJHaH uenttaH peaKUHH TIM - noJI11Mop<tJttoH,llepttb1e 11eitKOUHThI PHK - pw6oHYKJieHHOBM KHCJllOTa MPHK - MaTpwqHaH PHK
pPHK - pw6ocoMHM PHK TPHK - TpattcrroprnaH PHK PHP - pw60HYKJieOTH.Upen:YKTa3a
P3C - peTHKyJI03HllOTeJIHaJibHaH cwcTeMa CCC - cepn:eqHo-cocyn:wcrnH cwcTeMa CTr - coMaTOTporrtth1i1 ropMoH
TAr - TpHauHJirnwuepoJJb1
T,l(<D - THaMHH.llH<Poc<t>aT lTf - THpeOTpOITHbIH ropMOH YLl,<D - YPH.UHH.llH<Poc<PaT YM<l> - YPH.UHHMOHo<Poc<PaT Yr<D - YPMHHTpw<t>oc<PaT
Y<DO - yJJhTpa<PwoJieTOBoe 06Jiy'lett11e <l>A<l>C - 3-<Poc<Poa,lleH03HH-5-<Poc<PocyJJh<PaT <l>P,l(<D - <Poc<t>opw603wmw<Poc<PaT
<I>Cr - <PoMHKyJJOCTHMYJIHPYlOUIHH ropMOH, <PoMHTPOilHH
XIT - xopwottWieCKHi1 roHa,llOTporrHH XM - XHJIOMHKPOHbl
ll,ll<D - UHTH)J;HHllH<Poc<PaT UM<D - UHTH.UHHMono<Poc<t>aT
UHC - ueHTPaJJbHaH nepBHaH CHCTeMa Ull3 - uenh rrepettoca 3JieKTpOHOB
UTK - IIHKJI TpHKap6oHOBbIX KHCJIOT, IIHKJI Kpe6ca UT<D - UHTHlJ:HHTpw<t>oc<t>aT
3P - 3H,llOITJia3MaTHqecKHti peTHKyJIYM
CTPOEHlllE, CBOLilCTBA
Ill <l>YHKUllllll 6EJ1KOB
B )l(HBhIX KJieTKax rrpo11cxoJJ.HT CHHTe3 MHO)l(eCTBa opratt11qecK11x
MOJieKYJI, cpe)J,H KOTOpbIX rnaBH)'IOpOJih HrpaIOT IlOJIHMeptthle MaKpOMOJieJ<YJihl - 6eJIKH, HyKJieHHOBbie KHCJIOThl, rron11caxapH)J,hl.
Oco6asr ponh B )l(J13HeJJ,eHTeJihHOCTH )l(HBhix opraHH3MOB rrp11tta)J,Jle- )l(HT 6enKaM. OT poJJ,HTenetl: JJ,ernM rrepe)lafacH retteT11qecKaH 11tt¢opMau11sr o crreu11¢11qecKotl: CTPYKType 11 ¢YHKU:HHx Bcex 6eJIKOB )lattttoro oprattH3Ma. C11ttTe311poBattttb1e 6eJIKH BhIIlOJIHHIOT Mttoroo6pa3tthre ¢YHKu1111: ycKopHIOT XMM11qecK11e peaKU:HH, BblilOJIHHIOT TpaHcnopTttyIO, CTPYKTYPHYIO, 3aw,11THYIO ¢YHKU11H, yqacrnyIOT B rrepeJJ,aqe c11rHaJIOB OT 0)J,Hl1X KJieTOK JJ.PYD1M H TaKHM o6pa30M peaJIH3)'IOTHaCJie)l- CTBeHH)'IOHtt¢opMaU:HIO. Il03TOMY 6eJIKH Ha3blBaIOT TaK)l(e rrpoTeMHaMH (oT rpeq. proteas - rrepBhitt).
Ha llOJTIO 6enKOB BttyTpM KJieTKH np11xo)J,11TcH 6onee nonoB11Hb1 11x cyxoro Bemecrna. B oprattM3Me qeJioBeKa ttacY.11Tb1BaIOT OKOJIO 50 000
MHJJ.HBH)JyaJihHhIX 6eJIKOB. BMOBaH n MHllHBH)lYaJThHaH crreu11¢11qttocTh tta6opa 6eJIKOB B )laHHOM opraHH3Me onpe)leJIHeT oco6ettttocT11 ero CTpoettHH 11 ¢yttKu110H11poBaH11H. Ha6op 6eJIKOB B JJ.11¢¢epettu:11pyiomnxcH KJieTKax O)lttoro oprattM3Ma orrpe)lensreT Mop¢onorw1ecKHe 11 ¢YHKUHOHaJTbHbie oco6eHHOCTH Ka)l()lOfO THrra KJieTOK.
KaK H mo6oi1 IlOJIMMep, 6eJIOK COCTOHT H3 MOHOMepHbIX e)ll1H11U:, HJil1 «CTPOMTeJihHbIX 6JIOKOB». B 6enKax oprattH3Ma qenoBeKa TaKHM11 MOHOMepaMH CJI~aT 20 113 HeCKOJTbKHX COTeH H3BeCTHhIX B rrpnpo)le aMHHOKHCJIOT. AMMHOKHCJIOThl, HaXO)lHW,HeCH B 6eJIKax, CBH3aHhl ,llpyr c ,llpyroM rreITTH)lHbIMH CB51351Ml1. JlHHettHaH TIOCJTe)lOBaTeJibHOCTb aMl1HOKHCJIOT B 6eJTKe YHl1KaJihHa )J,JIH Ka)l()loro HH)ll1BH)lYaJTbHOfO 6eJIKa; Mtt¢opMaQ11H 0 Hett COJJ,ep)l(HTCH B yqaCTKe MOJieKyJThI )],HK, Ha3hIBaeMOH rettoM.
IlommerrT11)ltth1e u:err11 3a cY.eT BHYTPHMoneKynHptthIX B3a11Mo)lei1crn11i1 o6pa3yIOT rrpocTPattcrnetttthre CTPYKTYPhI - Kott¢opMau1111 6eJIKOB. Ha orrpe)leJiettttoM yqacTKe 6eJIKOBoi1 MOJieKYJihI M3 pa,llHKaJIOB aMHHOKHCJIOT ¢opMnpyeTCH aKTHBHhlll ueHTp, KOTOPhill MO)l(eT crreun¢nqtto (KOMilJleMeHTapHO) CB513bIBaTbCH C MOJTeKyJTaMH-Jil1faH)J,aMH. B3a11Mo)leticrn11e 6eJIKOB c JIHraHJJ.aMM ne)l(HT B octtoBe HX ¢YHKU:HOttnpoBaHHH. l13MeHeHHH IlOCJTe)J,OBaTeJibHOCTH aMMHOKHCJTOT B 6eJIKax MOfYT rrpHBOJJ.HTh K M3Mettett1110 rrpocTpattcTBettttoti CTPYKTYPhI 11 ¢YHKU11ti )J,aHHbIX 6eJIKOB 11 pa3BHTHIO 3a6oJieBaHHH.
I. CTPOEHVIE VI CBOlllCTBA
AMVIHOKVICflOT,BXO.QSUUVIX
B COCTAB 6EflKOB. nEnTVlllHblE
CBSl3VI, COEllVIHSllOUlVIE
AMVIHOKVICflOTbl B UEnVI
BeJIKtt - rroJittMepHbte MOJieKyJihI, B KOTOPhIX MOHOMepaMH CJIJ')KaT aMttHOKHCJIOThI. B cocrnBe 6eJIKOB B opraHl13Me Y.eJIOBeKa BCTpeY.aIOT TOJibKO
20 a-aMHHOKllCJIOT. 0)lHH H Te )Ke aMHHOKHCJIOTbl rrpttCYTCTBYIOT B paJJIWIHhIX rro CTPYKTYPe H cPYHKUIDIM 6eJIKax. 11H)ll1BH)lyaJibHOCTb 6eJIKOBblX MOJie KyJI orrpe)leJIHeTCH rropH)lKOM qepe)lOBaHHH aMttHOKttCJIOT B 6eJIKe. AMHHOKHCJIOThI MO)!(HO pacCMaTpHBaTh KaK 6yKBbI aJI¢aBl1Ta, rrpH ITOMOlilH KOTOpbTX, KaK B CJIOBe, JarrttChIBaeTCH HHcPOpMallHH. CJioBo tteceT HH¢opMaQ1110, Harrp11Mep o rrpe)lMeTe HJIH )leHCTBHH, a ITOCJie)lOBaTeJibHOCTb aMHHOKHCJIOT B 6eJIKe HeceT HHcj)opMau:mo 0 rroCTPOeHMH rrpocTpaHcTBeHHOH CTpYKTYPbf l1 cPYHKlll1l1 )laHHOfO 6eJIKa.
A. CTPOEHHE H CBOHCTBA AMHHOKHCJIOT
I. 06111,ue cmpyKmypnbte ocooe1111ocmu a.MunoKuCAom, exoiJaw,ux 6 cocmae OeAK06
06maH CTPYKTYPHaH oco6eHHOCTb aMHHOKl1CJIOT - HaJIHY.He aMMHO- H Kap60KCl1JibHOH rpyrrrr, coe)lHHeHHhIX c O)lHl1M tt TeM )Ke a-yrnepo)lHbIM arnMOM. R - pa,uttKaJI aMMHOKHCJIOT - B rrpocTetirneM CJiy'laerrpe)lCTaBJieH aTOMOM BO)lOpO)la (rnHU:HH), HO MO)l(eT IIMeTb H 6oJiee CJIO)l(HOe CTpOeHHe.
a
+ |
- |
I |
- |
NH3 |
CH - COO |
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B BO!lHhIX pacrnopax rrptt HeilTpaJibHOM JHaqeHHtt pH a-aMMHOKMCJIOTbI cyw:ecTBYIOT B BH)le 611rroJIHpHbIX HOHOB.
B OTJI11q11e OT 19 ocTaJibHbIX a-aMHHOKMCJIOT, ITpOJil1H - HMHHOKMCJIOTa, pa,lll1KaJI KOTOpOH CBH3aH KaK c a-yrnepO)lHblM aTOMOM, TaK l1 c aMl1Horpynrroti, B peJyJihTaTe qero MOJieKYJia rrp1106perneT I1ttKJI11Y.eCKYIO CTPYKTYPY·
a
HN-CH-COOH
I I
H2C, ,.CH2 CH2
npOJH1H
19 l13 20 aMttHOKMCJIOT CO)lep)!(aT B a-ITOJIO)l(e- HHH aCHMMeTpHY.HbIH aTOM yrnepo)la, c KOTOpbIM CBH3aHbI 4 paJHhre JaMema10mtte rpyrrrrbI. B peJYJihTaTe 3Tl1 aMl1HOKl1CJIOTbl B rrptt:po)le MOryT Ha-
XO)lHTbCH B !lByx paJHhIX tt30MepHbIX ¢opMax - L 11 D. 11CKJIIOY.eHHe COCTaBJIHeT rJI11U:l1H, KOTOpblH He HMeeT acHMMeTpHY.HOro a-yrnepO)lHOfO aTOMa, TaK KaK era PMHKaJI rrpe)lCTaBJieH TOJihKO aTOMOM BO)lopo)la. B cocrnBe 6eJIKOB rrpHCYTCTBYIOT TOJihKO L-1130Mepb1 aM11HoK11cnoT.
COOH |
COOH |
COOH |
I |
I |
I |
H2N-C-H |
H-C-NH2 |
H2N-C-H |
I |
I |
I |
CH3 |
CH3 |
H |
L-AnaHlllH |
D-AnaHlllH |
rnllll..llllH (He lllM99T |
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11130MepHblX cpopM) |
qHCThie L- HJIH D-cTepeomoMepb1 Moryr 3a )lJIHTeJibHbIB cpoK CaMorrpoH3BOJibHO 11 He¢epMeHTaTHBHO npeBpamaTuCH B 3KB11MOJIHptty10 cMeCh L- H D- 1130MepoB. 3To1' rrpoQecc HaJbIBaIOT paueMttJamreti. Pau:eMH3alIIDI. KmK,11,oti L-aMHHOKHCJIOThT rrpH )lattHoti TeMITepaTYPe Hlli!T c orrpe)leJieHHOH cKopoCTbIO. 3ro o6cTOHTeJibCTBO MO)l(HO HCJIOJII30BaTb )lJlH yCTaHOBJieHHH B03pacra JIIO)leH H )l(HBOTHblX. TaK, B rnep)lOH 3MaJil1 3y6oB HMeercH 6enoK )le:HTMH, B KOTOpoM L- acnapnn nepexo)ll1T B D-H3oMep rrp11 TeMrrepaTYPe rena qeJIOBeKa co CKOpOCTbIO 0,01 % B ro.u. B nep110.u ¢opMl1p0BaHHH 3y60B B )leHTHHe CO)lep)!(J1TCH TOJibKO L-H30Mep, rro3TOMY rro co.uep:>I<aHHIO D-acrraprara MO)J(HO paccqJ1TaTh BOJpacT o6c:Jie)lyeMoro_
Bee 20 aMHHOKMCJIOT B opraHH3Me qenoBeKa pa3JI11qa10TcH rro CTPOett1110, pa3MepaM 11 ¢11311Ko-x11- M11Y.eCKMM CBOHCTBaM pa,llttKaJTOB, rrp11coe)l11tteHHbIX K a-yrnepO,UHOMy aTOMY.
2. KAaccu<}JuKaU,UR a.MUllOKuCAom
no XUMU'leCKOMY cmpoe11u10 paiJUKll.il06
CTo XHMH'leCKOMy CTpOeHHIO aMHHOKHCJIOTbT MO)!(HO pa3)leJil1Tb Ha aJI11¢an1qecK11e, apoMan1- Y.ecKMe 11 reTepoQHKJIHqecKHe (rn6JI. 1-1).
B cocrnBe aJI11cj)aTtt'lecK11xpa,urrKaJIOB MOfYT ttaxo.urrThCH cj)yHKUHOHaJibHhie rpyrrnhI, np11)la10w:11e HM crreQHcPHY.eCKHe CBOHCT1Ba: Kap6oKCHJibHaH (-COOH), aMrrHo (-NH2), TttOJTbttaH (-SH), aMH)l- HaH (-CO-NH2), rH)lpOKCHJibHaH (-OH) l1 ryaHH-
)lHHOBaH (-NH-C=NH) rpyrrITbl.
I
NH 2
Ha3BaHIDI aMl1HOKHCJIOT MO)l(HO rrocTpOHTb no 3aMeCTl1TeJibHOH HOMeHK.IIaTYpe, HO o6hil.fHO HCIlOJib3YIOT TPHBHaJibHbie Ha3BaHHH (Ta6JI. 1-2).
10
Ta611M48 1-1. KnaccM<IJMK8~MA OCHOBHl>IX 8MMHOKMCllOT 6e11KOB no MX XMMM'leCKOMYCTpoeHMIO
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TpueuaJibHbie uaJeauuH |
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CoKpaII1i'HHbieuaJeauuH |
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CTpoeuuepaAHKaJIOB |
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aMHHOKHCJIOT |
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pyccKue |
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JiaTHHCKHe |
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I. AMHHOKHCJIOTbI c aJiu<t>aTuqecKHMH pa,l1,uKaJiaMH |
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1. fJIHIJ.HH |
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frrtt |
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Gly |
G |
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-H |
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2. ArraHHH |
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Arra |
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Ala |
A |
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-CH3 |
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3. BarrttH |
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Barr |
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Val |
V |
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-CH<CH3 |
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CH3 |
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4. JleHIJ.HH |
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Jleli |
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Leu |
L |
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-CH,CH <CH3 |
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CH3 |
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5. Ifaorreliu.HH |
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Hrre |
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Ile |
I |
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-CH-Cf-1:1-CH3 |
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I |
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CH3 |
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II. AMuuoKuCJIOTbI, co)J,ep3rnII1ne e aJimt>aTuqeCKOM paAHKaJie )1,0DOJIHHTeJibHYIO <l>YHKUHOHaJILHyIO rpynny
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fH,11;pOKCHJibHYIO rpyrmy |
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6. CepHH |
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Cep |
I |
Ser |
S |
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-CHrOH |
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7. TpeoHHH |
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Tpe |
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Thr |
T |
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-CHOH-CH3 |
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Kap6oKCHJihHYIO rpyrrrry |
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8. AcrraparnHosrui |
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Aerr |
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Asp |
D |
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-CHi-COOH |
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KHCJIOTa |
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9. frryTaMHHOBml KHCJIOTa |
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frry |
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Glu |
E |
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-CH2-CHrCOOH |
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AMHAHYIO rpyrrrry |
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10. AcrraparnH |
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AcH |
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Asn |
N |
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-CH2-CO-NH2 |
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11 . f JIYTaMHH |
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fJIH |
I |
Gln |
Q |
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-CHi-CH2-CO-NH2 |
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AMttHorpyrrrry |
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12. J1H3HH |
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JIH3 |
I |
Lys |
K |
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fyaHH)J,HHOBYIO rpyrrrry |
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13. AprttHHH |
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Apr |
I |
Arg |
R |
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-(CH2k NH-C-NH2 |
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II |
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Cepy |
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c |
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NH |
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14. lJ;HCTeHH |
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LI;ttc |
I |
Cys |
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-CHrSH |
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15. MeTHOHHH |
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MeT |
Met |
M |
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-CHrCHrS-CH3 |
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III. AMHHOKHCJIOTbI, co,l1,ep1KaII1ue apoMaTuqecKun pa,l1,HKaJI |
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16. cDeHHJiaJiaHHH |
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cDeH |
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Phe |
F |
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-CH2-0 |
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17. Tttp03HH |
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Tn:p |
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Tyr |
Y |
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-CH2-00H |
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IV. AMHHOKHCJIOThl c reTepouuKJiuqecKHMH pa)J,HKaJiaMu |
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18. TpttmoQiaH |
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Tptt |
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Trp |
W |
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-CHn;----;t'""" |
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ll..~ |
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N |
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H |
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19. fHCTH)J,HH |
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fttc |
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His |
H |
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-CHi-1 I |
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HN"-""N |
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V. liMHHOKHCJIOTa |
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20. Tiporrn:H |
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Tipo |
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Pro |
P |
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V-COOH |
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N |
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H ,QaHa nonHaR |
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¢opMyna |
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11
Ta6m11.1a 1-2. npMMepbl H83B8HMM aMMHOKMCnoT no 38MecTMTenbHOM HOMeHKna-rype .., COOTBeTCTBYIOll.lllle TplllBManbHble H83BaHMH
HaJeauue aMHHOKHCJIOTbI no |
cI>opMyJia aMHHOKHCJIOTbl |
TpnBHaJibttoe HaJBauue |
3aMeCTHTeJibHOH HOMeHKJiaType |
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2-aMHH0-3-r11)J,pOKCHI1pouaHOBa}l KHCJIOTa |
H2 N-CH-COOH |
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Ccp11tt |
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I |
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CH2 |
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l |
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OH |
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2-aMl1H0-4-MCTHJITl10MaCm!Ha}l KHCJIOTa |
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H2 N-CH-COOH |
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McTHOHHH |
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I |
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CH2 |
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I |
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CH2 |
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I |
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s |
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I |
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CH3 |
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LI,.1rn 3am1cw aMttHOKttc.110THhrx ocTaTKOB B MO- |
HbIM MOJieKYJiaM, B pe3yJihTaTe 'leronoBepXHOCTb |
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JieKyJiax nenTl1,L.l,OB 11 6eJIKOB l1Cfl0Jlh3YIOT Tpex6yK- |
conpl1KOCHOBeHl15l |
11X c BOnoH yMeHbWaeTCH. |
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BeHHbie COKpamemrn l1X TPl1Bl1a.JibHbIX Ha3BaH11ti, |
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a B HeKOTOpbIX c.11yqa51x 11 O,flH06yKBeHHble Cl1MBO- |
AMunoKHCJIOThI c |
n0Ji11pub1Mu ue1apIDKeHHbIMH |
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Jlbl (CM. Ta6JI. 1-1). |
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pa,Ll,HKaJiaMH |
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Tp11Bl1a.JlbHhie Ha3BaHl1H 'lacTo npowcxonHT OT |
Pan11Ka.11b1 3T11x aM11HoK11c.110T .11y11we, 'leMmn- |
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Ha3Bamrn: 11CTO'!Hl1Ka, 113 KOTOporo OHM BnepBbie |
po<:Po6tth1e pan11Ka.11h1, pacrnopHJOTCH B Bo,rie, TaK |
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6bJJ111 BbI,lle.JleHbl, |
11.1111 OT CBOHCTB ,riaHHOti aMl1HO- |
KaK B 11X COCTaB BXOA5IT IIOJIHPHbie cPYHKU110Ha.Jlh- |
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Kl1CJ10Tbl. TaK, cepHH BnepBhie 6hrn |
BhIAeJieH 113 |
Hbie rpynnhI, o6pa3yIOIU11e BO,llOpOAHhle CBH311 c |
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(OT JiaT. serieum |
- |
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qm6po11Ha rne.11Ka |
we.11KOB11c- |
Bo,rioti. K HMM OTHOCHT cep11H, TpeoHttH 11 Tttpo- |
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Tb1ti)' a rJI11U11H |
IIOJ1Y1111Jl CBOe Ha3BaHl1e 113-3a |
311H, 11Me10m11e r11,L.1,pOKC11JibH1>1e rpynn1>1, acnapa- |
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CJia,llKOro BKYCa (OT rpe11. glykos - |
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mH 11 rnyrnM11H, conep)!(amm~ aMl1JlHhie rpynnhI, |
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3. KAaccu</JuKaU.UR aMUHOKUCAom |
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11 U11CTel1H c ero Tl10JibHOH rpynnoti. |
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U11cTe11H 11 TMP0311H co,riep:KaT coorneTcrneHHO |
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no pacm«opUMocmu ux paiJu1Cll.ll06 « «ode |
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THOJibHYIO 11 rnnpOKCl1JlbHYIO rpynnhI, crroco6Hble |
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Bee 20 aM11HoK11CJIOT B 6e.11Kax opratt113Ma qe- |
K ,ri11ccou11au1111 c o6pa30Bam1eM H+, HO rrpw pH |
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JIOBeKa MO)[(HO crpynn11poBaTb TIO CIIOC06HOCTl1 11X |
OK0.110 7,0, nOMep)l<HBaeMOro B KJleTKax, 3T11 rpyn- |
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pan11Ka.JlOB pacTBOpHTbCH B Bo,rie. Pan11Ka.11b1 MO)!(- |
Ilbl npaKTH11ecK11 |
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HO BblCTPOl1Tb B HerrpepbJBHblll PM, Haq11Ha10ru11ti- |
AMHHOKHCJIOTbl c |
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CH nOJlHOCTblO m,ripo<:Po6HbIMl1 11 3aKaWIHBaJOIUHH- |
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3apIDKeHHblMH |
pa,Ll,HKaJiaMH |
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Pacrnop11MOCTb pa,ri11Ka.110B aMHHOK11c.110T on- |
K 3TOti rpynne OTHOC5IT acnapar11HOBYJO 11 rny- |
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pe,rie.11J1eTCH ITOJlHPHOCThJO cPYHKUHOHaJibHbIX |
TaMl1HOBYJO aMl1HOK11CJIOTbl, 11MeIOIU11e B pan11Ka.11e |
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rpynn, BXO)UIIUHX B COCTaB MOJieKyJihl (IIOJI5IpHbie |
AOITOJIHHTeJibttylO Kap60KCHJThHYJO rpynny, nptt pH |
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rpynnhI np11rnr11BaJOT BOAY, HenomipHbie ee OT- |
oKoJTo 7,0 n11ccou1111pyIOlllYJ10 c o6pa3oBatttteM |
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AMuuoKHCJIOTbI |
c ueno.m:1pubIMH pa,ri,uKa.ri:aMu |
MbI rnyTaMHHOBOH 11 acnaparnHOBOH KMCJIOT Ha3bI- |
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K HeIIOJI5IpHblM (m,ripo<:Po6HblM) OTHOCHT pa- |
BaIOT COOTBeTCTBeHHO rnyraMaTOM H acnapTaTOM. |
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AMunoKHCJIOTbI c |
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Hbie uen11 (pa,n,11Ka.11b1 a.11att11Ha, Ba.Jll1Ha, .11etiu11tta, |
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pa,Ll,HKaJiaMH |
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To<:Patta). Pan11Ka.11b1 TaK11x aM11HoK11c.110T B Bone |
rpynny B pan11Ka.11e 11MeIOT Jill1311H 11 aprnHl1H. y |
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cTpeMJ1TC5l npyr K ,ripyry 11n11 K npyr11M rnnpo<:Po6- |
.11113HHa BTopa51 aM11Horpynna,. cnoco6HaH rrp11co- |
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12
e,1urnHTb H+, pacnoJiaraeTCH B £-noJIO)Kemu1 am1- <PaT11'-leCKOtiuen11, a y aprnmrna noJIO)!(l1TeJibHbIH 3apSI,ll npno6perneT ryattJI,IJ,ttHOBa51 rpynna. KpoMe Toro, nICTH)lHH CO)lep)KJIT CJia6o HOHH3HpOBaHHYIO HMJI,IJ,a30JibHYIO rpynny' n03TOMY rrpH cPH3HOJIOrWieCKHX KOJie6attH51X 3Hal.feHHH pH (OT 6,9 )10 7,4) fHCTJI,IJ,HH 3ap51)Kett JIH60 HetiTpaJibHO, JIH6o IIOJIO- )KJITeJibHO. CTp11 YBeJI11qettHH KOJIH'-lecrna npoTottoB B cpeLJ,e HMHLJ,a30JibHa51 rpynna fHCTH)lHHa crroco6tta rrpttcoeLJ,HH51Tb rrpoTOH, rrpno6pern51 rroJIO)l(HTeJibHbitt 3apHL\, a rrptt YBeJIHl.feHHH KOH11,ettTpa11,im nI.llPOKCHJibHblX rpynrr - OTLJ,aBaTb rrpoTOH, Tep51SI IIOJIO)KJITeJibHbIH 3ap51LI, paLl,HKaJia. CTOJIO)KHTeJibHO 3ap51)KeHHbJe paL!,HKaJibl - KaTHOHbl (CM. cxeMy HH)!(e).
Hatt60Jiblllei1 pacTBOPHMOCTbIO B BOLJ,e 06Jia11a10T IIOJIHPHbJe 3ap51%eHHbie paLJ,HKaJibl aMHHOKl1CJIOT.
4. HJ.Menenue cyM.Mapnow 1apa<Ja QMUHOICUCJlOm 6 3Q6UCUMOcmu om pH cpeObt
CTpn ttethpaJibHbIX 3tta'-!eHH51X pH Bee KMCJIOTHbie (crroco6ttbte OTLJ,aBaTb H+) 11 oce octtOBHhre (cnoco6ttbie rrpttcoeLJ,HH51Tb H+) cPYHKIIHOHaJibHbie rpyrrIIbI HaXOMTC51 B ).UfCCOUHHpOBaHHOM COCT051HHH.
CT03TOMY B ttetiTpaJibHOH cpe11e aMHHOKHCJIOTbl, COLl,ep)Kalll,He HeLJ,HCCOUHHPYIOIIIHH paL!,HKaJI, HMeIOT CYMMapttbiti ttyJieBoti 3ap5IL1,. AM11ttoKHCJIOThI, COLl,ep)l(alll,He KHCJIOTHble cPYHKil,HOHaJibHbie rpynnbI, HMeIOT CYMMapHbIH OTPHil,aTeJibHbltt 3aPHL\, a aMHHOKHCJIOTbl, COLJ,ep%aW,He OCHOBHble cPYHKUHOHaJibHbie rpynIIbl, - IIOJIO)KJITeJibHbIH 3a-
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5. Mo<Juqmu,upo6aHHbte aMUHOICUCAOmbt, npucymcm6y10w,ue 6 Oe.tllCax
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·ooc / "coo· |
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y-Kap60KcHmyra- |
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r1'1.QPOKCl'1ni1131'1H |
Ml'1HOB8SlKl'1CJ10T8 |
MOAMct>M4MPOB3HHble KMCJ10Tbl, H3iilAeHHble B COCTaBe 6enKOB
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13