
підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin
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~'\Q w B IP
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CH20H |
CH20H |
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HO a Op |
CH20H |
H~~ HO |
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OH H |
1O4 OH H OH |
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OH H |
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H HO |
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H1 0 4 |
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H HO |
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OH H |
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OH |
H OH |
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H HO |
H OH |
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Caxapo3a |
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naKT038 |
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ManbTOJa |
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PMc. 7-8. JlMcaxapMAbl nMIJ.IM.
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• crpyKTYPHhie rroJittcaxapH,L\hI, o6ecrreY:ttBaIOIIlHe KJierKaM 11 opraHaM MexaHHY:eCKYJO rrpoY:- HOCTh (CM. pa3l(eJI 15);
fOl(apH a-KOH¢HrypauHH rJIIOK03HOro OCTaTKa, rroJIHCaxapHl(HaH uerrh ttMeer KOH¢opMau1110 crr11paJIH. CttHffil OKpacKa rrp11 l(06aBJieHHH HOl(a K pacrBopy KpaxMaJia o6ycJIOBJieHa HaJIHY:HeM TaKOH crr11paJI11. AM1111orreKTHH HMeer pa3Bern11eHHYIO CTDYKTYPY· B Mecrax BerBJieHHH ocrarKH rnIOK03hI coe)J,HHeHhl a-l ,6-rJIHK0311l(HhlMH CB51351MH. JlHHeHHhie yy:acTKH COl(ep)!<aT rrpHMepHO 20-25 OCTaTKOB rJIIOK03hJ. Ilp11 3TOM ¢opM11pyercH n:peBOBH)J,HaH
•ITOJIHCaxapHl(hI, BXOL(HIIJHe B cocTaB Me)f(- CTpYKrypa, B KOTOpo:H HMeeTCH JIHlllh Ol(Ha aHOMep-
KJieTOY:HOro MaTpMKCa, rrpttHHMaIOT yy:aCTHe B o6pa3oBaHttH TKaHeiil, a TaK)f(e B rrpoJimpepaUHH H l(tt<P<PepeHIJ:HPOBKe KJieTOK. IlOJIHcaxap11)J.hl Me)f(KJieTOY:HOro MaTpMKCa BO)J,0- paCTBOpHMhl H CHJihHO rnn:parttpoBaHhl (CM.
pa3l(eJI 15).
B ITMIIJe Y:eJIOBeKa B OCHOBHOM COl(ep)f(aTCH rro-
JIMCaxap11l(hl paCTHTeJibHOro rrpOHCXO)f(l(eHHH - KpaxMan, ueJIJIIOJI03a. B MeHhIIIeM KOJIH'-lecrnerro-
cryrraer IIOJIHCaxapHl( )f(l1BOTHhIX - fJIHKoreH. KpaxMaJI - Hatt6oJiee Ba)f(HhIH yrneBOl(HhIH
KOMrrottettT rr11II1esoro pau11otta. 3ro pe3epBtthlH ITOJIHCaxapHl( pacTeHHH' con:ep)f(all!HHCH B ttaH60JihIIIeM KOJIMY:ecrne (l(O 45% OT MaCChI cyxoro semecrna) B 3epttax 3JiaKoB (rrIIIeHttua, K)'Kypy3a, pHC H l(p.), a TaIOICe JIYKOBHUaX, cre6m1x H KJiy6H51X pacreHHM (B Kapro<t>eJie rrp11Meptto 65%). KpaxMan - pa3sern11etttthIH rroJittcaxapttl(, cocroHIIJHH H3 ocrarKoB rnIOK03hI (roMornttKatt). Ott ttaXOl(MTCH B KJieTKax pacreHHH B BHl(e rpaHyJI, rrpaKTHY:eCKH ttepacTBOPHM B BO)J,e.
KpaXMaJI COCTOHT H3 aMHJI03hl H aMHJIOIIeKTHHa (pttc. 7-9). AMttJio3a - ttepa3Bern11ettHhIH rroJIHcaxapHl\, BKJIIO'-IaIOIIJHH200-300 OCTaTKOB rJIIOK03hl, CBH3aHHhIX a- l ,4-rnHK03Hl(HO:H CBH3hIO. E11a-
HaH OH-rpyrrrra. KpaxMaJI - BhICOKOMOJieKy-
JIHpHoe coel(HHemre, BKJIIOY:aIOIIIee COTHH ThICH'-l
ocrarKoB rnIOK03hI. Ero MoJieKyAApttaH Macca cocraBJIHer rropHl(Ka I05-] 0 8 LI,.
.QeJIJIIOJI03a (KJieT'-IaTKa) - OCHOBHOH crpyKryp-
HhlH noJittcaxapHl( pacreHHH. 3ro caMoe pacrrpocrpatteHHOe oprattMY:ecKoe coel(11HeHHe Ha 3eM11e. Ll,oM ueJIJIIOJI03hI B KJieTOY:Hhix creHKax pacrettHH cocraBJIHeT 40-50%. UeJIJIIOJio3a HMeer MOJieKyJIHPHYJO Macey rropHL(Ka 106 LI,, l(JIHHa MOJieKyJihl MO)[(eT l(OXOl(HTh .IlO 6-8 MKM.
UeJIJIIOJI03a - JIHHettHhIH rroJI11caxapl1l( roMor-
JIHKaH, rrocrpoeHHhIH H3 OCTaTKOB rJIIOK03hl, coe)J,H- HeHHhIX Me)[(l(y co6o:H ~-1,4-rJIHK03Hl(HhIMH CB51351-
MH. IlHIIIeBap11reJihHaH CHCTeMa Y:eJioBeKa He HMeer ¢epMeHTOB, rnl(pOJIH3YJOIIIHX ~-CB5I3H B IIOJIHCaxaPH.IlaX. Ilo3TOMY ueJIJIIOJI03a - ttettcnoJih3yeMhIH yrneBO.Il, HO 3TOT IlHIIJeBOH KOMITOHeHT He06XOL(HM l(JIH HOpMaJihHOro npoTeKaHHH rrepesapHBaHHH.
rJIHKOfeH - TIOJIHCaxapHl( )[(HBOTHhIX H Y:eJIOBeKa. TaK )f(e, KaK KpaxMan B pacreHHHX, rJIHKOreH B KJieTKax )[(HBOTHhIX BhIITOJIHHeT pe3epBHYJO <PYHKUHIO, HO, TaK KaK B IlHIIJe conep)!(HTCH JIHII!h He60Jihll!Oe KOJIHY:eCTBO fJIHKOreHa, OH He HMeer rrttIIJeBoro 3HaY:eHHH.
304
1. AMlllI1038 20% |
2. AM111noneKT111H 80% |
P111c. 7-9. CTpoeH111e KpaxMana.
f m1KoreH rrpeacrnBJrneT co6otl: CTPYKTYPHhIH aHanor KpaxMana, HO HMeeT 6oJibIIIYJO cTeneHh BeTBJieHmi: np11MepHO Ha KmK)lbie 10 OCTaTKOB fJIIOK03bl npHXO)lHTC51 O)lHa cx- l ,6-rnHK03H)lHa51 CB513b.
11.nEPEBAP~BAH~E
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OJQ
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rru.1K03111,Q83blO HHo
OH HO
P111c. 7-10. f111Apon111s rn111K03MAHOiil ceS1s111.
A. IlEPEBAPMBAHME YfJIEBO.Ii:OB B POTOBOM
IIOJIOCTM
B poTOBOH noJIOCTH rrnwa H3MeJibtJ:aeTrn np11 ne-
pe)l(eBbIBaHHH, CMa'IHBa51Cb np11 3TOM CJJIOHOH. CJJJOHa Ha 99% COCTOHT H3 BOD.bl H o6bI'lHOHMeeT pH 6,8.
¢epMeHT cx-aMHJia3a (cx-1,4-rJIHK031f,lJ;a3a), paciuen-
JI51JOIUa51 B KpaxMaJie cx- l ,4-rJIHK03HD.Hhie CB513H.
B poTOBOM rroJIOCTH He MO)l(eT npo11cxo)lHTh noJIHoe paciuenJieHHe KpaxMarm, TaK KaK n.eti.crnHe ¢epMeHTa Ha KpaxMaJI KpaTKOBpeMeHHO. KpoMe Toro, aMHJJa3a CJIIOHbl He paCllleIIJI51eT cx-1,6-rnHK03H,U- Hble CB513H (CB513H B Mecrnx pa3BeTBJieHHM), I103TOMY KpaxMaJI nepeBapHBaeTC51 JJHWh tJ:aCTH'IHOc o6pa30BaHHeM KpynHhIX ¢parMeHTOB - neKCTpHHOB H He6oJihIIIoro KOJiw1ecrna MaJihT03hI. CJien.yeT OTMeTHTh, 'ITOaMHJia3a CJIIOHbl He nr,l.l,pomnyeT rnHK03H,UHbie CB513H B II.HCaxapl1JlaX.
Ll,etrcrn11e aMHJia3hI CJIJOHhI npeKpalllaeTC51 B pe3KO KHCJIOM cpe,ne con.ep)l(HMOfO )l(eJiy,UKa (pH
1,5-2,5). 0,nHaKo BHYTPH n11iueBoro KOMKa aKTHBHOCTh aMHJia3bl MO)!(eT HeKOTOpoe BpeMH coxpaH51.TbC51, noKa pH He H3MeHHTC51 B KHCJIYFO CTOPOHy. )KeJiy)lO'IHbIHCOK He co,nep)l(HT ¢epMeHTOB, pacwenm1.10iu11x yrneBOllht. B A<.eJJyII.O'lHOM con.ep)l(HMOM B03MO)l(eH JIHIIIb He3HalJMTeJihHbIH KHCJIOTHblH fH1lPOJIH3 fJIHK03H)lHhIX CB513eH.
Ii. IlEPEBAPMBAHME YfJIEB0.1J:OB
B KMIIIEqHMKE
TiocJJen.y10w11e 3TaI1hI nepeBapttBaHH51 HepaclllenJieHHoro HJIH qacTHlJ:Ho paclllenJieHHoro KpaxMaJJa, a
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IlauKpeaTll'lecKaHa-aMuna3a
B L1,BeHanuaT.11nepcTHOH KHWKe pH cpeLl,hI )!(eJiy-
Ll,OY:Horo conep)!(HMOro HeHTpaJIH3yeTC51, TaK KaK ceKpeT flOLl,)!(eJiyLl,OY:HOH )!(eJie3bl HMeeT pH 7,5-8,0 11 conep)!(HT 6.11Kap60HaTb1 (Heon. c ceKpeToM HOLl,)!(eJiyLl,OY:HOH )!(eJie3bl B Kl1WeY:Hl1K nocTynaeT
DattKpeaTll'leCKaH a-aMuna3a. 3TOT Q:>epMeHT rHLl,- po11113yeT a- I ,4-rnHK03HLl,Hble CB513l1 B KpaXMaJie H
neKCTPHHax.
npoLl,yKTbl nepeBapHBaHl151 KpaXMaJia Ha 3TOM
3Tane - n.11caxapHL1, MMbTOJa, conep)!(awa51 2 ocTaTKa fJIIOK03bl, CB513aHHble a- I,4-CB513b10. 113 Tex OCTaTKOB fJIIOK03bl, KOTOpb!e B MOJieKyJie KpaXMaJia HaXOLl,5JTC51 B Mecrax pa3BeTBJieHH51 11 COeLl,HHeHbl a- I ,6-rn.11K03l1Ll,HOH CB513bIO, 06pa3yeTC5J Ll,HCaxapHLI, H30MMbTOJa. KpoMe Toro, o6pa3yIOTC51 011.11rocaxaPHLLhI, conep)J(am.11e 3-8 ocraTKOB rnIOK03hI, CB513aHHbie a- l ,4- l1 a- l ,6-CB51351Ml1 (pHC. 7-11 ).
a-AMHJia3a IIOLl,)J(eJiyLLOY:HOH )J(eJie3bI, TaK )Ke, KaK a-aM1111a3a CJIIOHhI, netfcrnyer KaK 3HL1,orn11K0311Ll,a- 3a. naHKpeaTHY:eCKa51 a-aMHJia3a He paCIUenJI5IeT a- I ,6-rnHK03HLl,Hble CB5l3l1 B KpaxMMe. 3roT Q:>epMeHT TaK)J(e He rnnpo11113yeT P- l ,4-rn11K03HLl,Hbie CB513l1, KOTOPbIMH COeLl,HHeHhl OCTaTKH fJIIOK03bl B MOJieKYJie ueJIJIIOJI03hI. Ue11mo11ma, TaKHM o6pa3oM, npoxomrr Y:epe3 KHWeLJHHK HeH3MeHeHHOH. TeM He MeHee HerrepeBapeHHM ue11mo1103a BhinOJIH5Jer B~
HYIO Q:>yHKUHIO 6aJIJiaCTHOro Bewecrna, rrpHLl,aBa51 HHIUe Ll,OHOJIHHTeJibHbIH 06beM l1 HOJIO)J(l1TeJibHO BJI.115151 Ha npouecc rrepesap.11BaH.115I. KpoMe rnro, B TOJICTOM KHWelJHHKe ueJIJIIOJI03a MO)J(eT no,LI,BepraTbC}I Ll,eHCTBl1IO 6aKrepl1aJihHbIX Q:>epMeHTOB 11 Y:aCTHY:HO pacweruIBTbC51 c o6pa30BaHHeM CHHPTOB, opramtLJeCKHX _Kl1CJIOT l1 C02'flpOLl,YKTbl 6aKTepl1MbHOf0 pacwenJieH.1151 ueJIJIIOJI03hl B~Hbl KaK CTHMYJI51TOpbl rrep.11CTaJibTl1Kl1 KHWelJHHKa.
ManbT03a, 1130MaJibT03a .11 Tp1103ocaxapHL1,bI, o6pa3yIOw.11ec51 B BepxHHX OTLl,eJiax Kl1WelJHl1Ka .113 KpaxMaJia, - npoMe)!(yTOLJHbie rrponyKTbI. )..l,aJihHetfwee 11X nepeBapHBaHHe npOl1CXOLl,11T IIOLI, Ll,eHCTBHeM crreu.11Q:>11LJeCKl1X Q:>epMeHTOB B TOHKOM KHmelJHHKec )..l,.11caxap.11L1,bI n.11m.11 caxapma .11 11aKTOJa TaK)J(e rHL1,pOJil13YIOTCH crreu11Q:>11Y:eCKHMl1 Ll,l1CaxapHL1,a3aMH B TOHKOM KHWellHHKe.
Oco6eHHOCTb nepeBap.11BaH11H yrneBOLLOB B TOHKOM KHWel!HH.Ke 3aKJIIOl!aeTC51 B TOM, Y:TO aKTHBHOCTb cneu.11Q:>.11Y:ecK.11x 01111rol1 n:11caxap.11n:a3 B
npocseTe KHWeY:HHKa HH3Ka51. Ho Q:>epMeHTbI aKTHBHO Ll,eHCTBYIOT Ha noBepXHOCTH 3HHTeJIHaJibHblX KJieTOK Kl1WeY:HHKa.
ToHKHH KHWel!HHK H3Hyrp11 11MeeT Q:>opMy nMh-
ueo6pa3HbIX BblpOCTOB - BOpCHHOK, noKpbITblX
3IIHTeJil1aJlbHb!Ml1 KJieTKaMH. 3n11TeJIHaJibHbie KJieTKH, B CBOIO OY:epeLl,b, noKpblTbl MHKpOBOpCHHKaM.11, o6paweHHb!Ml1 B npOCBeT KHWeY:HHKa. 3r.11 KJieTKH BMeCTe C BOpC11HKaMH o6pa3ylOT rueTOY:HYIO KaeMKY, 611aroL1,ap51 KOTOpOH yBeJIHY:l1BaeTCH HOBepxHOCTb KOHTaKTa rHLl,pOJIHTHl!eCKHX Q:>epMeHTOB l1 HX cy6cTpaTOB B conep)J(HMOM K11weY:H11Ka. Ha 1 MM2 rIOBepxHOCTH TOHKOH KHWKH y Y:eJIOBeKa np11xoLl,l1TCH 80-140 MJIH BOpCHHOK.
<PepMeHTbl, pacwenJI51IOW11e fJil1K03HLl,Hbie CB513l1 B n11caxapwrnx (n11caxap11na3bI), o6pa3YJOT Q:>ep-
MeHraT11BHb1e KOMHJieKCbl, JIOKaJil130BaHHbie Ha
Hapy)!(HOH noBepXHOCTH UHTOHJia3MaTHlJeCKOH MeM6paHhl 3HTepOUHTOB.
Caxapa30-H30Ma.rtbTa3tthIH KOMnneKc
3ror Q:>epMeHTaTHBHblH KOMnJieKC COCTOHT 113
LLBYX no11.11nenTHL1,HbIX uenei1 11 11MeeT noMeHHoe cTpoeH11e. Caxapa30-H30MaJibTa3HbIH KOMnJieKc np.11Kpen11HeTCH K MeM6paHe MHKPOBOPCl1HOK Kl1Wel!Hl1Ka c fIOMOIUbIO mnpoQ:>o6Horo (TpaHCMeM6paHHOro) Ll,OMeHa, o6pa3oBaHHOro N-KOHUeBoH Y:aCTbIO no1111nerrT11n:a. KaTan.11r11qecK11tl. ueHTP BbIcrynaeT B npocseT K11weY:HHKa (p11c. 7-12). CB513h 3TOfO n11weBap11TeJibHOfO Q:>epMeHra c MeM6paHOH cnoco6crnyeT 3Q:>Q:>eKTHBHOMY rrornomeHHIO rrpo-
D:YKTOB I'l1).l,p0JIH3aKJieTKOH.
Caxapa30-H30MMbTa3HbIH KOMnJieKc rHL1,po11.113y-
eT caxapmy 11 1130MMbTOJy, pacmen115151 |
a-1,2- .11 |
a- l ,6-rn11K03HJlHbie CB513H. KpoMe rnro, |
o6a Q:>ep- |
MeHTHbIX ).l,OMeHa HMelOT MaJibTa3HYJO 11. MaJibTOTpl1a3HYIO aKTl1BHOCHI, rnn:pOJil13YH a- I ,4-rJil1K0311Ll,Hbie CB51311 B MaJibT03e 11 MaJibTOTp1103e (TP11Caxapl1LI,, o6pa3ylOW11HC51 113 Kpaxiv1aJia). Ha ).l,OJIIO caxapa30-113o- MMbTa3Horo KOMrIJieKca np11XO).l,11TCH 80% OT Beet!. MaJihTa3HOH aKTl1BHocr11 K11wetrn11Ka. Ho HecMOTP51 Ha npHCYWYIO eMy BbICOKYIO MaJibTa3HYJO aKT11B-
HOCTb, '.:.lTOT Q:>epMeHTaTl1BHblH KOMnJieKC Ha3BaH B
COOTBeTCTBl111 c OCHOBHOH cneu11Q:>11Y:HOCTblO. K TOMY )Ke caxapa3Ha51 cy6be).l,11H11ua - eL1,11HcTBeHHbIH Q:>epMem B KHWeY:Hl1Ke, rHLl,pOJil13YIOllLH.H caxapmy. l1JoMaJibTa3Ha51 cy6beLl,l1H11Ua c 60JihWetl. CKOpOCTblO r11L1,- pOJil13YeT fJI11K03HLl,Hble CB5.1311. B l130MaJihT03e, l!eM B MaJibT03e 11 MaJibTOTpl103e (p.11c. 7-13, 7-14).
B Toruetl. KHWKe coL1,ep)!(aH11e caxapa30-1130MMb- ra3HOro Q:>epMeHTaTl1BHOro KOMnJieKca ,llOCTaTOY:-
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HO BhICOKOe, HO OHO CHM)K.aeTCR B npOKCHMaJihHOH H ,n:HCTaJihHOti: qacn1x KHWe'-IHHKa.
f JIHKOaMHJia3HhlH KOMDJieKc
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cj:lepMeHT ornocRT K 3K30fJIHK03H.n;a3aM. KoMnJieKc pacw,enmreT TaIOKe CB513H B MaJihT03e, ,lleHCTBYH KaK MaJibTa3a. B rnMKOaMHJia3ttbrti KOMnJieKc BXOLlRT LlBe pa3Hble KaTaJIHTM'-IeCKMe cy6beJJ,HHHUhI, MMeJOWMe He60Jibllll1e pa3JIH'-IHR B cy6cTpaTHOH cneuHcj:IH'-1- HOCTH. fJIMKOaMMJia3HaR aKTMBHOCTh KOMilJleKca HaH60JihlliaR B HH)K.HHX omenax TOHKOfO KMWe<t- HMKa.
cco
cco
u-aMHJla3a ManbTOTpH03a
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KpaxMan
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T
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ManbTOTp11103a HO-\__)-o-\__)-o-\__)-
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lt130M0JlbT030 1-toro
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B KllETKVI
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PMC. 7-14. AeMCTBMe caxapa30-M30ManbTa3HOro KOMnneKca Ha M30ManbT03Y M onMrocaxapMA.
~-fJIHK0311,LJ,a3HbIH KOMDJieKC (JiaKTa3a)
JlaKTa3a paclll,ernrneT ~-1,4-rn11K03ll)J,Hbie CBH311 Me)l():(y rarraKT030H H rJIIOK030H B JiaKT03e (pl1C. 7-15).
mOT c):>epMeHTaTl1BHhIH KOMIUieKC ITO XHMWieCKOH ITp11po.ue HBJUieTCSI rn11KoITpoTeHHOM. JlaKTa3a, KaK 11 .upyrne rJIHK03ll)J,a3Hhie KOMIUieKCbl, CBH3atta c lll,eTO'lHOH KaeMKOH 11 pacITpe)leJietta ttepaBHOMeptto ITO BCeMY TOHKOMY KHIIIe'lHHKY.AKTHBHOCTb JlaI<Ta3bl KOJie6JieTCH B 3aBHCl1MOCTH OT B03pacrn. TaK, aKTHBHOCTh JiaKTa3hI y ITJIO)la oco6etttto ITOBhIIIIetta B IT03)lHHe cpoKH 6epeMeHHOCTl1 H coxpaHHeTCH Ha BhICOKOM ypOBHe )JO 5-7-JieTttero B03pacrn. 3aTeM aKTHBHOCTb <f>epMeHTa CHmI<:aeTCH, COCTaBJI5HJ y B3pOCJiblX 10% OT ypOBHH: aKTl1BHOCTH, xapaKTepttoro )lJIH )leTeii.
TaIOKe rnHK03H)la3Hhii1: KOMrmeKc, fH)lPOJIH3yIOlll,HH CB51:3H Me)l():(y MOHOMepaMH B TPefaJI03e - .u11caxapH)le, co.uep:>1<:alll,eMcH B rpn6ax. TperaJI03a COCTOHT H3 ,I:UJyx fJIIOK03HhIX OCTaTKOB, CBH3aHHhIX fJIHK03ll)J,HOH CB51:3hl0 Me)l():(y ITepBhlMl1 aHOMepHbIMl1 aTOMaM11 yrnepo.ua (pHC. 7-16).
CoBMecrnoe .uei1:crn11e Bcex ITepeq11cJieHHhIX <f>epMeHToB 3aBepIIIaeT ITepeBapHBaHHe I1Hlll,eBbIX OJIH-
A. BCAChIBAHHE MOHOCAXAPH)J,OB
B KHIIIEqHHKE
TpattcITopT MOHocaxapH)lOB B KJieTKH CJIH3HCToi1: o60JIO'-lKHKHIIIe'lHHKaMO)f(eT OCylll,eCTBJI51:TbCH pa3HhIMl1 cIToco6aMH: ITYTeM o6Jier'leHHoi1:.utt<P- <f>y31111 11 aKTl1Bttoro TpattcnopTa. B cJiyqae aKTl1BHoro TpaHCITOpTa rn10K03a H Na+ ITPOXO)lH:T qepe3 MeM6paHbl c JIIOMl1HaJibHOH CTOPOHbl, CBH3bIBaHCb c pa3HhIMH y'-laCTKaMtt 6eJIKa-nepettoc- 'lHKa.Tip11 3TOM Na+ ITOcTynaeT B KJieTKY ITO rpa- )lHeHTY KOHI1eHTPa111111, l1 O)lHOBpeMeHHO fJIIOK03a TPaHCITOpTttpyeTCH ITPOTHB rpa.unettrn Kott11ettTpaUHl1 (BTOpM'lHO-aKTl1BHbIH TpaHCITOpT, CM. pa3- )leJI 5). CJie)loBaTeJihHO, 'leM6oJihIIIe rpa.r:i:tteHT Na+, TeM 6oJihIIIe ITOCTYITJieHtte rnIDK03hI B 3HTePOIIHThI. EcJIH KOHI1eHTpa11HH: Na+ BO BHeKJieTo'l- HOH )K.H)JKOCTl1 YMeHhIIIaeTcH, TPaHCITOPT rn10Ko3hI cttmKaeTcSI. fpa.r:i:tteHT KOHUettTpautt11 Na+, H:BAAIOIUHHCH )lBmKylll,eii CHJIOH aKTHBHOfO CHMITOpTa, co3.uaeTcH pa6ornl1 Na+ ,K+ -AT<l>-a3hI. Tiepettoc B KJieTKH CJII13l1CTOH o60JIO'IKHKHIIIe'IHHKa ITO MexaHH3MY BTOpH'-lHO-aKTHBHOro TPaHCITOpTa xapaKTepeH TaIOKe )lAA raJiaKT03hI.
TipH pa3HOH KOHI1eHTPa1IHH fJIIOK03bl B ITPOCBeTe KHIIIe'IHHKa«pa60TaIOT» pa31Il1'1HhleMexaHH3Mbl TpaHCITOpTa. DJiaro.uapH aKTHBHOMY TpaHCITOpTy 3ITHTeJIHaJibHble KJieTKl1 KHIIIe'IHHKaMOryT ITOfJIOlll,aTb fJIIOK03Y ITpH ee O'leHbHH3KOH KOHUeHTPaIIHH B ITpocneTe Killlle'IHHKa.Ecmr )f(e KOHU:eHTIJa-
308

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"~="'" |
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fiaKTa3a |
'4' |
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+ |
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OH |
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fiaKTo3a |
D-ranaKro3a |
D-rmoKo3a |
PMC. 7-15. .QeMCTBMe naKT83bl.
H OH |
H |
rnlOK038 |
rnlOK038 |
PMc. 7-16. CTpoeHMe Tperanoab1.
UIDI rJIIOK03hl B rrpocBeTe KHllle'rnHKaBeJIHKa, TO Otta MO)J(eT TpattcrropTHPOBaTbC51 B KJieTKY rryreM o6nerqettttoti: llHcPcPY3HH. TaKHM )J(e cnoco6oM MO- )l(eT BCacbrnaThC5l H <l>PYKT03a. Cne.IJ,yeT OTMeTHTh, qTo CKOpOCTh BCaCbIBattH51 rJIIOK03bl H raJiaKT03hl ropa3,a:o BhrIIIe, qeM ,llpyrnx MOttocaxaptt,Il:OB. Cnoco6b1 TPattcrropTa MottocaxapM,llOB qepe3 MeM6patty 3I1HTeJ111aJihHbIX KJieTOK KHweqm1Ka npe.ll,CTaMeHbl tta p11c. 7-18.
IlocJie BcachIBamrn MottocaxapM,llhI (rnaBHhIM o6pa3oM, rJIIOK03a) ITOKH.IJ,aIOT KJieTKM CJIM3MCTOH 060JIO'!KMKHIIIeqm1Ka qepe3 MeM6patty, o6pamettttyl0 K KpOBeHOCHOMY KaITHJIJI51py, c ITOMO!llhlO o6nerqeHHOll ,a:11¢cpy3l111. qaCTb rJIIOK03bl (6oJiee I10J10Bl1Hbl) qepe3 Kamuurnpbl KHllleqHhIX BOpCHHOK rrona,llaeT B KPOBettOCHYIO CHCTeMY 11 no BOpOTttOH Bette
.ll,OCTaBmieTCH B neqeHh. 0CTaJJbHOe KOJil1'-leCTBO fJIIOK03hl nOCTYJ1aeT B KJieTKM .ll,pyrnx TKatteil.
B. TPAHCTIOPT fJIIOK03hl H3 KPOBH
B K.JIETKH
11oTPe6JJeHHe fJIIOK03bl KJleTKaMl1 113 KPOBOTOKa
rrpottCXO,llMT TaK)J(e rryreM o6JieI"-lettHOH ,llH<pcpy3HH.
CJie.IJ,oBaTeJihHO, cKopocTh TPaHCMeM6paHHoro no-
TOKa rJIIOK03bl 3aBl1Cl1T TOJibKO OT rpa,U11ettTa ee
KOHUeHTpa11,1111. J1CKJIIO!.[eH11e COCTaBJ151IOT KJieTKH
MhIIIIU M )l(HPOBOH TKattH, me o6Jierlfetttta51 ,lll1cpcpy31151 peryJI11pyeTc5l HHCYJJHHOM (ropMott no.ll,)J(e- JIYllO':!HOH)J(eJie3hI). B 0Tcyrcrn11e ttHcyn11tta nJia3- MaTw-recKaH MeM6patta 3TMX KJieTOK ttenpomrnaeMa llJ151 fJIIOK03bl, TaK KaK Otta He conep)J(HT 6eJIKH- nepeHOCl-fl1KH (TpattcnopTeph1) rnroK03hr. Tpattcrrop1ephr rnIOK03hr tta3hisaroT TaJOKe peu,enTopaM11 rnIOK03hI. Hanp11Mep, on11caH TpaHcnopTep rnroK03hI, BhWeJieHHhIH 113 3pttTpo11,ttTOB. 3To TPattcMeM6patttthIM 6eJIOK, I10JI11nenTH).l,Ha5l 11,enb KOTOporo ITOCTPOetta H3 492 aMHHOKMCJ10THbIX OCTaTKOB H HMeeT ,llOMettttylO CTPYKTYPY· nom1pHbie !lOMeHhl 6eJIKa pacnoJIO)J(etthl no pa3ttble CTOPOHbl MeM6paHhI, rnnpo¢06tth1e pacnonaraIOTCH B MeM6patte, nepeceKa5l ee HeCKOJihKO pa3. TpattcnopTep 11MeeT yqacTOK CB5!3bIBaHH5l fJ1IOK03bl Ha Bttewttetl cTopotte MeM6patthI. Iloc11e npttcoe,llttHettHH rmoK03hI KOH¢opMaUH5I 6eJIKa H3MettHeTC51, B pe3yJ1hTaTe qero [JIIOK03a OKa3bIBaeTC51 CB513aHttOH c 6eJJKOM B yl-faCTKe, o6pameHHOM BHyrpb KJleTKH. 3aTeM fJJIOK03a OT.IJ,eJIHeTCH OT TpaHcnopTepa, nepeXO,llH BHYTPh KJieTK11 (CM. pa3,lleJJ 5).
Cl-fttTaroT, 1.-fTO crroco6 o6Jierl-feHHOH Lll1cPcPY3Hl1 no cpaBttettHIO c aKTHBHblM TpattcrropTOM rrpe- !lOTBpamaeT TpattcnopT HOHOB BMeCTe c fJIIOK030H, ecJIH Otta TpattcnopnrpyeTCH no rpa,U11ettry KOHIJ,ettTpa11,1111.
309

a-aM1-1na3a cnlOHbt
KpaxMan,
caxapo3a,
naKT03a
KpaxMan,
A6KCTp1-1Hbl
l-130ManbT03a,
Tp1111 ormro- caxap1-1Abt
l-130ManbTa3a
caxapa3a
@ - rn10K03a, D - cppyKT03a,
P11c. 7-17. nepeeap111eaH111e yrneBOl'.IOB.
noA)f(enyA04Hafl
)f(ene3a
/K1-1W64Hl-1K
8HT6p0[.11-1T
I
@
D
Q - ranaKT03a.
310

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Na+ |
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<t>pyKT038 |
rmoK038 |
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ranaKT03a |
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® |
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6 |
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/0 |
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l~ |
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i |
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u |
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!VU |
P; |
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3Na+ |
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A,Q<I> |
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® |
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D |
Na+ |
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l |
l/ |
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2K+ |
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2K+ |
D ®O KpOBb
-6enK111-nepeHOCY111K111 (rpaHcnoprepb1) cppyKT03bl;
-6enK111-nepeHoc11111K111 (rpaHcnoprepb1) rnt0K03b1;
-Na+-3as111c111Mbliii 6enoK-nepeHoc11111K;
&_ _ Na+,K+ - AT<t>-a3a.
w=
PMC. 7-18. BcaCblBaHMe yrneBOAOB B KMWe'-IHMKe. BcaCblBaHllle MOHOcaxaplllAOB 1113 KlllW9'iHlllKanpOlllCXOAlllT nyTeM o6nerYeHHOH Alllcpcpy3111111 c noMOU1blO cneu111anbHblX 6enKos-nepeHOC4111KOB (TpaHcnopTepos). KpoMe Toro, rn10K03a 111 ranaKTo3a TpaHcnoprnpy10Tc5l B 3HTepou111T nyTeM BTOp1114HO-aKTlllBHOro TpaHcnopTa, 3aBlllClllMOro OT rpaA1119HTa KOHU9HTpau111111 lllOHOB HaTp11151. 6enKlll-TpaHcnopTepbl, 3aBlllClllMbl9 OT rpaA1119HTa Na+, o6ecne4111BalOT BCaCblBaHlll9 rnlOK03bl 1113 npocBeTa KlllW94H111Ka B 3HT9p0UlllT npOTlllB rpas:i111eHTa KOHU9HTpau111111. KOHU9HTpaU1115l Na+, Heo6XOA111Mal'lAnl'l3TOro TpaHcnoprn, 06ecneY111BaeTC5l Na•,K•-ATct>-a30H, KOTOpal'lpa6oTaeT KaK Hacoc, OTKa4111Ba5l 1113 Kn9TKlll Na• B o6MeH Ha K·. B OTnlil'illleOT rnlOK03bl, ¢PYKT03a TpaHcnoprnpyeTCl'lClllCTeMOH, He 3aBlllC5llJ19H OT rpaAl19HTa HaTp11151.
311
(fJUOT) o6ttapy;KeHbl BO Bcex TKallilX. Cyrn:ecrnyer HeCKOJihKO pa:moBHllHOCTe:H fJHOT (Ta6JI. 7-1), OHH rrpottyMepoBaHhI B COOTBeTCTBllH c ITOPMKOM HX o6Hapy;Kemrn.
CTpyKrypa 6eJIKOB ceMeH:crna fJJIOT ornwmeTCH OT 6eJIKOB, TpaHCITOPTllPYJOIUllX fJIIOK03Y qepe3 MeM6patty B KHUie'IHHKeH IIO'IKaxrrpOTHB rpa-
ntteHTa KOHUeHTpaUHH.
01rncaHHhie 5 TlllIOB f JJIOT llMeIOT CXO)lHhie
rrepBH'IHYJOCTpYKTYPY ll )lOMeHHYJO opraHJBaUHIO.
• f JJIOT-1 o6ecne'IHBaeTCTa6HJihHbIM IIOTOK
|
fJIJOK03bJ B M03r; |
• |
fJJIOT-2 06Hapy)((eH B KJieTKax opraHOB, BbI- |
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neJIHJOIUHX rnIOKo3y B KPOBh. l1MeHHo rrptt yqa- |
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cmH f JJIOT-2 rJIIOK03a nepexonm B KPOBh H3 |
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3HTepOUHTOB ll ne'leHH.f JJIOT-2 yqacrnyeT B |
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TpaHCIIOPTe fJIIOK03hI B P-KJieTKH no,LOKenyaoq- |
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ttoil: )((eJie3h1; |
• |
f JJIOT-3 o6JiaLJ;aeT 6oJiblllllM, 'leMfJJIOT-1, |
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cpoacrnoM K fJIJOK03e. Ott TaK)((e o6ecrreqn- |
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BaeT IIOCTOHHHblH rrpMTOK fJIIOK03bl K KJieT- |
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KaM HepBHOll ll ,i:i:pyrnx TKaHeil:; |
• |
f JJIOT-4 - fJiaBHblll nepeHOC'lllKfJIIOK03hI |
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B KJieTKH MhilllU ll )((HpOBOH TKaHH; |
• fJJIOT-5 BCTpeqaeTCH, fJiaBHhIM o6pa30M, B |
|
|
KJieTKax TOHKoro Kttllle'lHHKa. Ero <l>YHKUHH |
|
ll3BeCTHbl He)lOCTaTO'IHO. |
Bee rnrrhI fJJIOT MOryT HaxoattThCH KaK B IIJia3MaTtt'leCKoil:MeM6paHe, TaK ll B UHT030JihHhIX Be3MKYJiax. fJJIOT-4 (H B MeHhllle:H Mepe fJJIOT-1)
ITO'ITllIIOJIHOCTbIO HaXOLJ;HTCH B UHTOIIJia3Me KJieTOK. BJIHHHtte MHCYJIHHa Ha rnKHe KJieTKH rrpHBOLJ:HT K rrepeMemeHHIO Be3MKYJI, coaep)KamHX fJJIOT,
K IIJia3MaTH'leCKOHMeM6paHe, CJIIDIHHIO c He:H ll BCTpaMBaHHIO TpaHCIIOPTepOB B MeM6paHy. IlOCJie qero B03MO)((eH o6Jief'leHHhIBTpaHCIIOPT fJIIOK03bI B 3TH KJieTKH. IlOCJie CHH)KeHIDI KOHUeHTpaUIDI MHCYJIHHa B KPOBH TpattcrropTeph1 fJIJOK03hI cttoBa rre-
peMemaIOTCH B UHTOIIJia3MY, H IIOCTYJIJiem1e fJIIOK03bI B KJieTKy rrpeKpamaeTcH (pttc. 7-19).
IlepeMemeHHe fJJIOK03bl H3 nepBH'lHOHMO'lH B KJieTKH IIO'le'IHblXKaHaJibUeB rrpollCXOLJ:HT BIOPM'l- HO-aKTHBHblM TpaHCIIOPTOM, IIOLJ;OOHO TOMY, KaK 3TO ocyrn:eCTBJIHeTCH rrptt BCaCbJBaHHM fJIJOK03bI H3 rrpoCBeTa KMllle'lHHKaB 3HTepouttThI. EnaronapH 3TOMY fJIIOK03a MO)KeT ITOCTYJiaTh B K.JleTKM aaxe B TOM cny- qae, ecmr ee KOHUeHTpaUHH B rrepBH'lHOllMO'leMeHhUie, qeM B K.JleTKax. Ilptt 3TOM fJIJOK03a pea6cop6ttPYeTCH H3 nepBWIHOM MO'lHITO'ITHTIOJIHOCThIO (99%).
Yl3BeCTHhl pa31IM'IHbieHapyrneHHH B pa6oTe TpaHCIIOPTePOB rnIOK03hI. HacnencrneHHhrH: neQ:>eKT 3THX 6eJIKOB MO)KeT Jie)KaTb B OCHOBe HHCYJIMHOHe3aBHCllMOf0 caxapttoro LJ;Ha6ern (cM. pa3,Uen 11). Bro )((e BpeMH npH'IMHOMHapyrneHIDI pa60Tbl TpaHcrroprepa fJIIOK03hl MO)((eT 6hITh He TOJihKO ne<t:>eKT caMoro 6eJIKa. HapyrneHHH ¢>YHKUMM fJJIOT-4 B03MO)((HhI Ha CJieeyIOIUHX 3Tarrax:
•rrepeaaqa cttrttana MHCYJittHa o rrepeMemeHHM 3TOfO TpaHCIIOpTepa K MeM6patte;
•nepeMemetttte TpaHcrropTepa B UHTOTIJia3Me;
•BKJIIO'leHtteB COCTaB MeM6paHhI;
•OTUJHYPOBbIBaHtte OT MeM6paHbl ll T.LJ;.
IV. HAPYWEHl'ISl nEPEBAPl'IBAHl'ISl l'IBCACblBAHl'ISlvrnEBO.QOB
B OCHOBe naTOJIOfMll rrepeBapMBaHHH M BCaChIBaHMH yrneBO,ll,OB MOryT 6bITh rrpll'IMHhI LJ;Byx THIIOB:
•ne<t:>eKTbl <t:>epMeHTOB, yqaCTBYJOIUHX B fHJlPOJIH3e yrneBOLJ;OB B Kmlle'IHMKe;
•HapyllleHHe BCaChlBaHIDI rrpo,UyKTOB rrepeBapllBaHHH yrneBO,ll,OB B KJieTKM CJIH3llCTOH 060JIO'IKHKllllie'IHHKa.
Ta6mu.1a 7-1. PacnpeAeneHMe 6enKoe-TpaHcnopTepoe rnioKo31>1 (rntOT)
Tunbl r JllOT |
JloKaJinJaum1 o oprauax |
||
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rnIOT-1 |
TipettMyrn;ecTBeHHO B M03re, rrnau;eHTe, TIO'IKax,TOJICTOM KllIUe'IHHKe |
||
rnIOT-2 |
TipettMyrn;ecTBeHHO B ne'leHH,no'IKax,P-KJieTKax ocTpOBKOB Jlattrepxattca, |
||
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3HTepOQHTaX |
|
rnIOT-3 |
Bo MHOrHX TKaIDIX, BKJIIO'laslM03f, nnau;ettzy, TIO'IKH |
||
rnIOT-4 |
B Mhiuru;ax (cKeneTttoii, cep)l;e'lttoH:),)lrnposoH: TKaHH |
||
( ttttcynttH3aBHCHMhIH) |
Co)l;ep)l<RTCSI B OTCYTCTBHe ttttcyntttta no'ITHnonHOCThIO B u;ttTonna3Me |
||
fJIIOT-5 |
|
B TOHKOM KHUie'IHHKe.Bo3MOJKHO, SIBmieTCSI nepeHOC'IHKOM¢pyKT03bI. |
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312

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rmoKo3a |
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/1~ |
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• |
• |
• |
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Pe1.1emoP.-----.... |
1 |
1 |
1 |
MeM6paHa |
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__y |
~· |
l |
1 |
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'\ |
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• |
• |
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' |
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'\
' |
,, |
' |
' |
\ |
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' |
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'',, |
/ / ' |
TpaHcnoprepb1
rmOK03bl (rnlOT-4)
PMC. 7-19. BnMRHMe MHCynMHa Ha nepeMe1J4eHMe TpaHcnopTepoe rnlOK03bl M3 4MTOnna3Mbl B nna3M8TM'leCKYIOMeM6paHy. 1 -
CBR3b1eaH1o1e 1o1Hcy1rnHa c pe4emopoM; 2 - y4acTOK HHcyn1o1Hoeoro pe4emopa, o6pal.LlEiHHblM eHyrpb KnSTKH, crnMyn1o1pyeT nepeMel.LleHHe TpaHcnopTepoe rntOK03bl. 3, 4 - TpaHcnopTSpbl B COCTaB9 COA9P>K8l.LlHX HX B93HKyn nepeM9l.Ll8IOTCR K nna3Marn49CKOM M9M6paH9 Kn9TKH, BKnl048IOTCR Bee COCTaB H nepeHOCsn mtOK03y B Kn9TKy.
B 06011x c11yqMx B03HHKaeT OCMOTH'lecKaH.u11apeH, KOTOpy10 BbI3hIBalOT HepacmenneHHhie ,[(lfCaxapHL(bI MJIH HeBCOCaBWHeC5l MOHOcaxapH)J,bl. 3-111
HeBOCTpe6oBaHHhle yrneBO,[(bl rroczynaIOT B ,[(HCTaJibHhie OT,[(eJlhl KHllie'IHHKa, H3Mellirn OCMOTH'leCKoe
.uaBJieHMe co.uep)l(HMOro Kmne'IHHKa. KpoMe Toro, OCTaBUIHeCH B rrpocBeTe KHllie'IHHKayrneBO,[(bI 'laCTH'IHOIIO,[(BepraIOTC5l $epMeHTaTHBHOMY pacmerrJieHHlO MHKPOopraHH3MaMM c o6prooBaHHeM opraHWiecKHX KHCJIOT H ra30B. Bee BMecTe npttBO.UHT K rrpHTOKY BO.Uhl B KHWe'IHHK,yneJIH'lemuoo6oeMa KHWe'IHOroco.uep)J<JfMOro, ycl1JleHHIO nepHCTaJibTHKH, Cila3MaM 11 60AAM, a TalOKe MeTeop113MY.
TepMHHOM «MaJiha6cop6UH5I» Ha3hIBaIOT He.uoCTaTO'IHOeBCaCbIBamte rrepeBapeHHblX IIPO.UYKTOB yrneBOL\OB. Ho nocKOJihKY KmrnwrecKtte rrpo5IBJieHH5l np11 tte.uoCTaTO'IHOMnepenap11BaH11H 11 ncaChrnamrn CXO,[(Hbl'TO TepMHHOM «MaIIha6cop6uH5l» Ha3bIBaIOT o6a BHL(a HapYllieHHll.
A. llAPYllIEHIDI IJEPEBAPHBAHHR YfJIEB0,11,0B
B KHillEqHHKE
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313