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228
HapY?l(Ha>i nosepXHOCTb MeM6paHbl
BHYTpeHH>iH noaepXHOCTb MeM6paHbl
P111c. 5-1. nonepe'IHblMpaspes nnasMaT1111.1ecKoiii MeM6paHbl.
1. Cmpy«mypa u C6oucm6a AunuiJo6 MeMfipan
MeM6patttthre Jil1DH./lhI -- aMcpmj>HJibHbie (aM<tmnaTw1ecK11e) MOJieKYJibl, T.e. B MOJieKyJie eCTb KaK rn.upoQ:>11JihHbie rpynnhI (nom1pttb1e «roJIOBKH» ), TaK 11 aJIMQ:>aTw1ecK11e pa,/ll1KaJibl (rn.upo¢>o6ttble «XBOCTbl» ), caMonp0113BOJibHO Q:>opM11pyJOw,Me 611cJIOH. B 60Jihw11ttcrne 3yKap110THYecKHx KJieTOK OHM COCTaBJI51JOT OKOJIO 30-70% MaCCbl MeM6paHbl (p11c. 5-2). B MeM6pattax np11CYTCTBYJOT JI11n11nh1 Tpex rnaBHhIX Tl1CTOB - Q:>ocQ:>ommM.llhI, rJIMKOJTMCTM./lbl M XOJieCTepOJI (xoJiecTep11H).
JlHDMJlHblH COCTaB MeM6paH pa3JIJ1lieH, co.uep- )1\aHMe Toro HJil1 ,n:pyroro JIMCTl1./la, no-BH./lHMOMy, onpe.ueJI51eTC51 pa3H006pa311eM ¢>YHKU11H, BhinOJIH51eMblX 3TMM11 mm11,n:aM11 B MeM6pattax.
<l>occpoJinnll)J,hI. Bee ¢>oc¢>0JI111111.ub1 MO.lKHO pa3-
.ueJI11Tb tta 2 rpynnb1 - rnMuepoQ:>ocQ:>oJI11n11.uh1 11 cQ:>11H roQ:>ocQ:>OJIM n 11./lbl. rJI 11uepoQ:>ocQ:>OJIHn 11./lbl OTHOC51T K npOH3BO./lHblM Q:>ocQ:>aTH./lHOH KMCJIOTbl. HaM6oJiee pacnpocTpaHeHHbre rmm:epoQ:>oc- Q:>oJI11n11.ub1 MeM6paH - Q:>ocQ:>aTl1./lHJIXOJIHHbl 11 ¢>oc¢>arn.u11JI3TattoJiaM11ttb1 (pMc. 5-3). B MeM6pattax 3yKapMOTl1YeCKl1X KJieTOK o6Hapy.lKetto orpOMHOe KOJIWieCTBO pa3Hb!X ¢>ocQ:>oJIMill1./lOB, np11YeM OHM pacnpe.uenettw ttepaBHOMeptto no pa3HblM KJieTOYHblM MeM6pattaM. 3Ta HepaBHOMepHOCTb OTHOCMTC51 K pacnpe.ueJieHl1!0 KaK noJI51pHblX «roJIOBOK» (Ta6JI. 5-1 ), TaK M aU11JihHbIX OCTaTKOB (Ta6JI. 5-2).
Ka.lK./lblH rn11uepoQ:>oc¢>0JI11nM.ll, ttanp11Mep Q:>ocQ:>aTM./ll1JIXOJIMH, npeµ,cTaBJieH HeCKOJlhK11Ml1 )leC51TKaMM ¢>ocQ:>aTH./lHJIXOJil1HOB, OTJIJ1lfa!OLI.l,MXC51 ,rrpyr OT .upyra CTpoettMeM .lKl1PHO-Kl1CJIOTHhIX OCTaTKOB.
llhnerpanbHb!ill 601101<
J111m1Abl |
D |
6enKvi |
MeM6paHb1: |
CoCTaa, % |
|
0 |
|
100 |
Ml18Dl4HOB |
|
|
nna3ManNecKaH
annapaT fonbA>Klll
3HAonna3Ma-
Tlll4eCKl-IVI pen11cynyM
Hapy)l{HBR
Ml1TOXOHAP11anbH8H
BHYTpeHHRR
MlllTOXOHAP1118flbH8H
P111c. 5-2. CoAep>KaH111e 11111n111AOB 111 6enKoe e pasn1111.1Hb1x KneTO'IHblXMeM6paHax (%).
Ha µ,omo rn11uepo¢>oc¢>0JI11nn.uos (nOJI51pHa51 rpynna - 11H0311TOJI) np11XO./lMTC5! JIHWb 2-8% scex ¢>ocQ:>oJil1IlH./lOB, co.uep.lKaW11XC51 B KJieTO'-lHOHMeM- 6patte 3yKap110TOB. 11H0311TOJI B COCTaBe ¢>ocQ:>aT11-
./ll1JIMH0311TOJIOB MO.lKeT 6bITb ¢>ocQ:>opHJI11posaH no
C4 (Q:>ocQ:>aTH.llMJil1H0311TOJI-4-MOHOQ:>ocQ:>aT) HJil1 C4
11 C 5 ( ¢>ocQ:>aTM,ITMJIHH03HTOJI-4, 5-6McQ:>ocQ:>aT).
229
0 - |
nom1pHa5! |
1 |
" ro1100Ka,. |
O=P-0-
C=OC=O
l |
I |
|
|
|
|
|
|
CH2 |
CH2 |
|
|
|
|
|
|
l |
I |
|
|
|
|
|
CH3 |
CH2 |
CH2 |
|
|
|
+ |
||
I |
I |
|
|
|
0 - CH2- CH2- N ~ CH3 |
||
CH2 |
CH2 |
|
|
|
O=P-0- |
CH3 |
|
I |
l |
|
|
|
|||
CH2 |
CH2 |
|
|
|
|
|
|
I |
l |
|
|
|
|
|
|
CH2 |
CH2 |
|
|
|
|
|
|
I |
I |
|
|
|
|
|
|
CH2 |
CH2 |
|
|
|
|
|
|
I |
I |
|
|
|
|
|
|
CH2 |
CH2 |
f111ApOcj:>06Hbllil |
|
C=OC=O |
|
||
I |
l |
«XBOCT» |
|
|
|||
CH2 |
CH |
|
I |
I |
|
||
|
|
CH2 |
CH2 |
|
|||
I |
U |
|
|
|
|||
CH2 |
CH |
|
|
I |
I |
|
|
I |
I |
|
|
R1 |
R2 |
|
|
CH2 |
CH2 |
|
|
|
|
|
|
|
|
|
|
|
|
||
I |
I |
|
|
AllfllHHble |
|
||
CH2 |
CH2 |
|
|
|
|||
|
'am1¢ar1114ecK111e |
|
|||||
I |
I |
|
|
||||
CH2 |
CH2 |
|
|
1..1en111 |
|
||
I |
I |
|
|
|
|||
|
|
|
|
|
|
||
CH2 |
CH2 |
|
|
|
|
|
|
! |
! |
|
|
|
|
|
|
CH2 |
CH2 |
|
<t>occj:>aTlllAlllilXOJlf/IH |
|
|||
!!
CH2 CH2
I I
CH2 CH2
!I
CH3 CH3
<t>occj:>aTl!IAHas:i Kf/ICJlOTa
OCHzCHOH -CHzO
I I
-0-P=O O=P-0-
'
O=C |
C=O |
I |
I |
CH2 |
CH2 |
I |
I |
R1 |
R2 |
|
|
|
' ---- y ----- " |
|
|
||
AllfllHHble |
||||
AllfllHHble |
||||
am..icpar1114ecK111e |
am'1cpan14ecK111e |
|||
|
4en111 |
4em1 |
||
Kap.Q111on111n111H
+
0 - CH2- CHzNH3
I
O=P-0-
C=OC=O
I I
|
CH2 |
CH2 |
|
! |
I |
|
R1 |
R2 |
' |
---- v |
---- ' |
AllfllHHble an111¢ar1114ecK111e 1..1en111
<t>occpaTl!IAlllJl3TaHOJlaMlllH
,J'.'.l:.
f'HbH
0 OH
!
O= P-0-
1
AllfllHHble a11111¢ar1114ecK111e 1..1en111
<t>occpaTM.QlllJlf/IH03111TOJl
230
Ta6n111i.ia 5-1. <t>occj>on1tnMAHbliii COCTaB KlleTO'IHblXopraHenn It nna3M8TM'leCKOiiiMeM6paHbl renaTOLIMTOB
<l>octlJoJJunu,!l,bI |
|
|
|
,[(oJJH OT cyMMapuoro KOJIH'leCTBatlJoctlJoJJHDH,!l,OB, % |
|||||
|
|
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|
|
|
MeM6paHbl |
|
||
c pa3HbIM CTpoeuueM |
Ml!TO- |
|
R,D;epHaR |
IIJia3MaT1111ecKaR |
|||||
Jil130C0Mbl |
arrrrapaTa |
||||||||
UOJJHpHblX <<rOJJOBOK» |
XOH,D;p1111 |
MeM6paHa |
MeM6paHa |
||||||
|
fOJib,ll;)Kl! |
||||||||
|
|
|
|
|
|
|
|
||
|
|
|
|
|
|
|
|
|
|
Kap,D;11orr11rr11H |
18 |
|
|
1 |
4 |
1 |
1 |
||
<l>occpaTH):(HJI3TaHOJiaMl1H |
35 |
|
|
14 |
13 |
20 |
23 |
||
<l>occpaT!1,!l,11JIXOJil1H |
40 |
|
|
40 |
55 |
50 |
39 |
||
<l>occpaTH,!l,11Jil1H0311TOJI |
5 |
|
|
5 |
10 |
12 |
8 |
||
<l>occpaT11,D;11rrcep11H |
1 |
|
|
2 |
3 |
6 |
9 |
||
<l>occpaTl!,D;HaR Kl!CJIOTa |
- |
|
|
1 |
2 |
1 |
1 |
||
|
|
||||||||
Ccp11HroM11err11H |
1 |
|
|
20 |
3 |
8 |
16 |
||
|
|
|
|
|
|
|
|
|
|
Ta6nMLl8 5-2. JK1tpHO-KMCnOTHbliii COCTaB HeKOTOpblX MeM6paH ne'leHM
|
|
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|
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|
|
JKupub1e |
|
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|
MeM6pauuaH tlJpaKJ)HH |
|
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||
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||
|
KHCJIOTbl, |
|
MeM6paHbI MMTOXOH,!l,p11il |
|
ArrrrapaT r OJib,D;)Kl! |
IIrraJMaT1111ecKaR |
||||
|
% (no Macce) |
|
|
|
3P |
|||||
|
Hapy)KHaR |
BHYTPeHHRR |
MeM6paHa |
|||||||
|
|
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|||||
|
|
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|
|
|
|
|
|
||
|
M11p11CTHHOBaR |
0,4 |
0,3 |
0,4 |
0,9 |
|
0,9 |
|||
14:0 |
|
|
|
|
|
|
|
|
|
|
|
IJaJibMl!Tl!HOBaR |
4,0 |
3,6 |
3,1 |
- |
|
- |
|||
16:0 |
|
|
|
|
|
|
|
|
|
|
|
IJaJibMl!TO- |
21,0 |
18,0 |
26,5 |
22,5 |
|
31,2 |
|||
|
orre11HoBaR |
|
|
|
|
|
|
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|
|
16:1 |
|
|
|
|
|
|
|
|
|
|
|
CTeap11Hosa11 |
13,5 |
15,8 |
14,9 |
18,5 |
|
12,9 |
|||
18:0 |
|
|
|
|
|
|
|
|
|
|
|
Apax11,!l,0HoBaR |
15,7 |
18,5 |
14,0 |
14,5 |
|
|
|
||
|
|
11, 1 |
||||||||
20:4 |
|
3,5 |
3,8 |
0,7 |
- |
|
|
|
||
|
IJ;epBOHOBaR |
|
- |
|||||||
22:6 |
|
|
|
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|
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' |
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||
B cocrnB <Poc<Parn,L(HJ111ttoJ11ToJI-4,5-6ttc<Poc<PaToB BXOJUIT B OCHOBHOM auHJihHbie OCTaTl(l1 CTeap11HOBOW HJil1 naJihM11Tl1HOBOW (no nepBOMY TIOJIO)l(e- Hl1l0 nrnuepona) 11 apaxtt,L(ottoBo:ti (no BTOPOMY noJIO)l(ett1110) )1(11pHbIX Kl1CJ10T.
Cneu11<Pw1ecK11e <Poc<Pon11nH,n:h1 BHYTpettHew MeM6paHhl MHTOXOH.n:Pl1M - Kapn1101111nl1Hbl (.n:11- <Poc<PaTl11lHJlrJil1UepoJibI), TIOCTpoeHHhie Ha ocHOBe rnttuepona 11 ,n:Byx ocTaTKOB <Poc<Pan1.n:- ttow Kl1CJ10Tbl. 0HH CHHTe3HpyIOTCH <PepMeHTa-
MH BHYTPeHHeM MeM6paHbl MHTOXOHllPHM H coCTaBJlHIOT OKOJlO 22% OT Bcex <Poc<Pon11m1)J,OB MeM6paHbl.
B nnaJMaTWiecKHx MeM6pattax KJieTOK B JHaqttTeJibHhIX KOJ111'1eCTBaXCOJiep)!(aTCH CcPl1HfOMl1eJll1Hbl (p11c. 5-4). C<P11ttroM11eJIHHhI nOCTPOeHhI Ha OCHOBe uepaMHJia - au11n11poBaHHOro aMl1HOcn11pTa CcPHHf03HHa. 00JIHpttaH rpynna COCTOl1T 113 OCTaTKa cPOCcPOPHOM KHCJlOTbl l1 XOJIHHa, 3TaHOJlaM11Ha HJ111 cep1rna. C<P11ttroM11eJittHbI - rnaBHhie JIHnH,!l,bl MHeJIHHOBOW 06onoqK11 HepBHbIX BOJlOKOH.
fJIHKOJIHDH,!l,bl. B rJIHKOJIHnH,L(ax rHJipo<Po6ttaH qaCTb npe,n:CTaBJieHa uepaMHJIOM. fHJIPOcPHJibHWI rpyrma - yrneBO,L(HblW OCTaTOK, np11coe,!l,11HeHHbIW rJIHK03HJIHOM CBH3bl0 K rHJipOKCHJlbHOM rpynne y
231
nomlpHafl «fOnOBKa»
HC |
|
|
II |
|
|
CH |
|
|
I |
|
|
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
|
rHApOcp06Hbli:I |
CH2 |
|
|
I |
|
«XBOCT» |
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
|
|
CH2 |
|
|
I |
J |
|
CH3 |
||
|
||
CQJVIHf03VIH
H
H |
C=O |
|
|
I |
|
HC |
CH2 |
|
I |
||
II |
||
CH2 |
||
CH |
||
I |
||
I |
CH2 |
|
CH2 |
I |
|
I |
CH2 |
|
CH2 |
I |
|
I |
CH2 |
|
CH2 |
||
I |
||
I |
CH2 |
|
CH2 |
||
I |
||
I |
CH2 |
|
CH2 |
||
I |
||
I |
||
CH2 |
||
CH2 |
||
I |
||
I |
||
CH2 |
||
CH2 |
||
I |
||
I |
CH2 |
|
CH2 |
||
I |
||
I |
CH2 |
|
CH2 |
||
I |
||
I |
CH2 |
|
CH2 |
||
I |
||
I |
CH2 |
|
CH2 |
||
I |
||
I |
||
CH2 |
||
CH2 |
||
I |
||
I |
CH2 |
|
CH3 |
||
I |
||
|
CH2 |
|
|
I |
|
|
CHa |
LlepaMVIA
PMc. 5-4. Ccj>MHrocpoccponMnMAbl - npoM3BOAHble 1.1epaMMJ18.
CH3
I
CH3- N+-CH3
I
CH2
I
CH2
I
0
I
O=P-0-
nomlpHaR «fOnOBKa»
AnviHHbte anvicpaTw-iecKvie 1..1en111
r1-1ApOcpo6Hb1e «XBOCTbl»
Cqn11--iroM111en111H
nepBoro yrnepo,LJ.Horo aroMa uepaMH)J,a (pMc.
B 3aBMCMMOCTM or MMHhI w crpoeHM>I yn1eBo,LJ.- Hoil qacrw pa3JIH'!aIOTn;epe6po3H,!J,bt, co)J,ep)l(arn,Me MOHOMJIM OJrnrocaxapM,LJ.HhIH OCTaTOK, If raurnu-
03H,!J,bl, K OH-rpynne KOTOph1x npwcoe)J,wtteH cJIO)l(- HhIH, pa3BeTBJieHHbIM OJIMrocaxapH)J,, CO)J,ep)l(aill,MH N-auerMJitteilpaMMHOBy10 KMCJIOTY (NANA) (cM. pa3).l,eJI 7).
CToAApHble «fOJIOBKH» fJIHKOCcpMHfOJIHIIM)J,OB HaXO)J,>ITC>I Ha Hap~o:H IIOBepXHOCTM IIJia3MaTwqecKMX MeM6paH. B 3HaqwreJibHblX KOJIMqecTBax rnMKOJIMilH,1J,hI CO)J,ep)((aTC>I B MeM6paHax KJieTOK M03ra, 3pHTpOUMTOB, 3IlMTeJIHaJibHb!X KJieTOK. faHfJIH03H- )J,bl 3p.MTpOU.MTOB pa3Hh1X HH,LJ.MBH,LJ.YYMOB pa3Jlwqa10rc>I CTpOeHHeM OJUITOCaxapM,LJ.HblX ueneti:, npO>IBJIHJOill,HX aHTHfeHHhie CBOHCTBa.
232
UH CHpH
HO
0
|
HC |
HC |
||
|
II |
II |
||
|
CH |
CH |
||
|
I |
I |
||
|
R1 |
R1 |
||
|
|
|
|
|
|
A.nYIHHbl6 |
Am1HHb1e |
||
|
anY1cparn1..1ecKY1e |
am1cpan1YeCK1.1e |
||
|
1.1em1 |
1..1en11 |
||
L1epe6po31'1.Q(ra11a1<To1.1epe6po31.1.Q) |
faHr!ll.1031'1,Q |
|||
PMc. 5-5. rn111Kon111nMAbl. Gal - ranaKT03a; Glc - mt0K0Ja; NANA (NeuAc) - |
N-auernnHei;ipaMl'1HOBa51Hnf.1 Cf.1anosa51 Kf.1cnorn. |
|||
XoJiecTepoJI. XoJiecTepon rrpHcyrcrnyeT BO Bcex MeM6pattax )KllBOTHhIX KJieroK. Ero MOJieKyJia cocroVIT H3 )J(eCTKOfO fHD,po¢o6Horo 51)],pa H rn6KOM ymeBOJ.lOPOJ.lHOM uenH, e)lHHCTBeHHaJI rHD,pOKCHJibHaH
rpyrrrra 5IBJrn:ercH «IJOllilpHOM ffiJlOBKOM» (pMC.
,U;rn )l(IIBOTHOH KJleTKH cpe).l,Hee MOAApHOe OTHO-
rneHMe xo11ecTepo11/cpoccpo11HrrH).l,hI paBHO
HO B IJJla3MaTH'!eCKOM MeM6paHe 3TO COOTHOllle-
HHe ropa3no Bhrwe Hanwme xoJiecTepo-
11a B MeM6paHax yMeHbWaeT IJO)lBH)l(HOCTb )l(HpHblX KHCJlOT, CHlDKaer JlaTepaJibHYIO )lMcpcpy3HIO JIHTIH- )lOB M 6eJIKOB, H IJ03TOMy MO)f(eT BJIH51.Tb Ha cpyHKUMH MeM6paHHbIX 6eJIKOB.
B cocraBe MeM6paH pacTeHHti. xo11ecTepoJia HeT, a rrpHcyrcTByroT paCTMTeJlbHbie CTepOH).l,bl - CHTOCTepoJI H CTHrMacrepoJI.
2. TpaHCMeM6paHHUR acUMMempu.a AunuiJo6
K~a.sI MeM6paHa KJieTKH JaMKHyra, r.e. HMeer BHyrpeHHIOIO H BHeWHIOIO IJOBepXHOCTH, pa3-
JIM'IaIOIIl,HeC51no JlMIJH)lHOMY H 6eJIKOBOMY cocra-
BaM - 3TY oco6eHHOCTb MeM6paH Ha3bIBaror
TpancMeM6pauuoii (norrepetrnotl.) acHMMeTpueii.
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