Добавил:
Upload Опубликованный материал нарушает ваши авторские права? Сообщите нам.
Вуз: Предмет: Файл:

підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

.pdf
Скачиваний:
32
Добавлен:
25.02.2016
Размер:
44.13 Mб
Скачать

~'\Q w B IP

 

©;~

CH20H

CH20H

 

©;x5-

 

 

 

 

 

 

 

 

HO a Op

CH20H

H~~ HO

a

 

1

2

 

 

 

 

 

OH H

1O4 OH H OH

 

OH H

 

H HO

OH

H1 0 4

OH

OH

 

 

 

 

 

H HO

 

OH H

H

OH

H OH

 

H HO

H OH

 

Caxapo3a

 

 

naKT038

 

 

ManbTOJa

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

PMc. 7-8. JlMcaxapMAbl nMIJ.IM.

xapH,L\hl Mettee paCTBOPHMhl, Y:eM MOHOCaxaPHl\hl, CJiel(oBaTeJibHO OHM He BJIIDUOT Ha ocMOTHY:eCKOe )J.aBJieHMe H I103TOM)'MOryr HaKarrJlHBaTbCH B KJieTKe, HarrpHMep, KpaxMaJI - B KJieTKaX pacreHHH, fJIHKOreH - B KJieTKax )f(H- BOTHhIX;

• 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

7-10).
B CJIJOHe npHCYTCTByeT mn.poJIHTITTeCKHH

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

YrnEBOJlOB

3rr11TeJIHaJihHhie KJieTKH KHIIIe'lHHKacnoco6HhI BCaCbIBaTh TOJibKO MOHOCaxaplf)lbl. TI03TOMY npouecc nepeBapHBaHH51 3aKJIJOtJ:aeTC51 B ¢epMeHTaTHBHOM I'H)lp0JIH3e fJIHK03H)lHhIX CB513eH B yrneBOII.ax, HMelOIUHX OJIHI'O- HJIH IIOJIHCaxapH)lHOe cTpoeH11e (p11c.

OJQ

r-H,O

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

305

TaK)!(e npyrnx yrneBOLl,OB IIHWH npOHCXOLl,HT B TOHKOM KHilleY:HHKe B pa3HbIX ero OTLl,eJiax non Ll,eHCTB.11eM rHLl,pOJIHTHY:eCKHX Q:>epMeHTOB - fJil1K03HLl,a3.

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:-

306

HO BhICOKOe, HO OHO CHM)K.aeTCR B npOKCHMaJihHOH H ,n:HCTaJihHOti: qacn1x KHWe'-IHHKa.

f JIHKOaMHJia3HhlH KOMDJieKc

3ToT <PepMeHTaTHBHbIH KOMnJTeKC KaTa.JIH3HpyeT rM,UpOJIH3 a- l ,4-CB513H Me)l(Uy fJIIOK03HhlMl1 ocTaTKaMH B OJIHrocaxapM,Uax, ,lleHCTBYR c BOCCTaHaBJIHBaJOw,ero KOHUa. no MeXaHM3MY JJ,ettCTBHR 3TOT

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

OnHrocaxapH.Qbl

KpaxMan

PMC. 7-12. Caxapa30-M30ManbT33HblM KOMnneKC. 1 - caxapasa; 2 - H30MaJlbTa3a; 3 - CB513b1Bal0Ll\"1iii AOMeH; 4 - rpaHCMeM6- paHHbl~ AOMeH; 5 - LlHTOnna3MarnLteCK"1iii AOMeH.

307

TperaJiaJa -

a-1,4-CBR3b

ManbT03a HoOoOoH

T

ManbTa3a

,}-Q, f,}-Q,\ ,?-Q,

ManbTOTp11103a HO-\__)-o-\__)-o-\__)-

PMc. 7-13. AeMCTBMe caxapa30-M30ManbTa3Horo KOMnneKca Ha ManbT03Y M ManbTOTPM03y.

lt130M0JlbT030 1-toro

~a-1,6-CBR3b

~ZJ-oH

1-/o~./lt130M0JlbT030

ro- H ITOJI11caxapH)lOB c o6pa30BaIDieM MOHOCaxapll)J,OB, octtoBttoii 113 KOTOPhIX - rn10Ko3a. KpoMe fJIIOK03hl, H3 yrneBO)lOB ITHlll,H TaIOKe o6pa3yIOTCH cPPYKT03a H raJiaKT03a, B MeHbIIIeM KOJIH'leCTBe - MaHtt03a, KCHJI03a, apa611tto3a. 06IUaH cxeMa ITepeBapHBattHH yrneBO)lOB ITpe)lCTaBJieHa Ha pttc. 7-17.

Ill. MEXAHVl3M

TPAHCMEM6PAHHOro

nEPEHOCA r11t0K03bl

VI llPYrVIX MOHOCAXAPVlllOB

B KllETKVI

MottocaxapH)lhI, o6pa3oBaBIIIHeCH B pe3yJihTaTe ITepeBap11BaHHH, BCaChIBalOTCSI 3ITHTeJIHaJihHbIMH KJieTKaMH TOlll,eH H ITO)lB3)lOIIIHOH KHIIIOK c ITOMOlll,hlO cITeI1HaJihHhIX Mexatttt3MOB TpaHcIToprn qepe3 MeM6paHhl 3THX KJieTOK.

HoOoOoH

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

 

 

"~="'"

fiaKTa3a

'4'

 

 

 

+

 

 

OH

 

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

 

 

Na+

 

 

 

<t>pyKT038

rmoK038

 

 

ranaKT03a

 

 

 

 

 

®

 

 

 

 

 

6

 

 

/0

 

 

l~

 

 

 

 

i

 

 

 

 

 

u

 

 

 

 

!VU

P;

 

3Na+

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

A,Q<I>

 

 

 

 

®

 

 

 

 

 

D

Na+

 

 

 

 

 

 

l

l/

 

 

 

2K+

 

 

 

 

 

 

 

 

 

 

 

 

 

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

fJIIOK03HLle TpaHCDOPTfpbl

(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-

 

neJIHJOIUHX rnIOKo3y B KPOBh. l1MeHHo rrptt yqa-

 

cmH f JJIOT-2 rJIIOK03a nepexonm B KPOBh H3

 

3HTepOUHTOB ll ne'leHH.f JJIOT-2 yqacrnyeT B

 

TpaHCIIOPTe fJIIOK03hI B P-KJieTKH no,LOKenyaoq-

 

ttoil: )((eJie3h1;

f JJIOT-3 o6JiaLJ;aeT 6oJiblllllM, 'leMfJJIOT-1,

 

cpoacrnoM K fJIJOK03e. Ott TaK)((e o6ecrreqn-

 

BaeT IIOCTOHHHblH rrpMTOK fJIIOK03bl K KJieT-

 

KaM HepBHOll ll ,i:i:pyrnx TKaHeil:;

f JJIOT-4 - fJiaBHblll nepeHOC'lllKfJIIOK03hI

 

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

 

 

 

 

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,

 

 

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.

 

312

 

 

 

 

 

rmoKo3a

 

 

 

 

 

 

 

/1~

 

 

 

 

 

 

 

 

Pe1.1emoP.-----....

1

1

1

MeM6paHa

 

 

 

 

 

__y

l

1

 

 

'\

 

 

 

'

 

 

 

 

 

'\

'

,,

'

'

\

'

'',,

/ / '

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

HapyweHHH rrepeBapHBaHHH MOfYT 6bITh CBH3aHhI KaK c He,[(OCTaTO'-IHOHaKTHBHOCTblO OT)l,eJlbHbIX ,[(HcaxapHL(a3, TaK H c He)l,OCTaT01:1HOCTblO BCero ¢epMeHTaTHBHoro KOMnJieKca, ttarrp11Mep caxapa- 30-H30MaIIhTa3HOro.

1'faBeCTHhIttacne.ucrneHHhie H rrpwo6peTeHHhre ¢opMbl He,[(OCTaTOlJHOCTH aKTHBHOCTH ¢epMeHTOB. CHMilTOMbl BpO)!()],eHHblX ¢opM npOHBJIHIOTC5I ,[(OCTaT01:1HO paHO, HanpttMep ITOCJie rrepBbIX KOpMJieHHll rpy.UHblM MOJIOKOM (rrpH .ue<t>ttUHTe JiaKTa3hI), nocne nepexo.ua Ha HCKJ'CCTBeHHoe BCKapMmrnaHHe MJIH rrpH .uo6aBJieHMH B pau110H caxapa H KpaxMana (rrp11 .ueqmu1ne a-aM11Jia3hI HJIH crreuH¢11qecK11x .u11caxaptt.Ua3). B CJIY1:1ae He-

.uocTaT01:1Horo Jie'leHHHBpo)!()],eHHbie ¢opMhI naTOJIOrHH conpoBO)!()],aIOTCH xpoHHlJeCKHM .u11c6aK-

313

Соседние файлы в папке підручники. біохімія