підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin
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1,3-6111ccpoccporrn111.iepar
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<t>occpor.n1111..jepar- |
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2,3-6111ccpoccpor.n111L1epar |
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P111c. 7-43. 06pa3oaaH111e 111 npeapall.leHMe 2,3-6111c<j>occj>orn1111.1epara.
qecKH TIOJlHOCTbIO 11C'leprrbIBaIOTCHnocne CYJO'lHOfO rono)J,aHllil.. B 3TOM cnyq:ae B ne'lett11 tta'l11ttaeTCH cHHTeJ rJIIOKOObI de novo - rJIIOKOHeoreaeJ. f nIOKO-
tteorette3 - npouecc c11ttTe3a fJ1JOK03hI 113 Bern,ecrn HeyrneBOllHOH np11pOllhI. Ero ocHOBHOH <l>YHKu11ei1: HBJIHeTCH II01l]lep)l{att11e ypOBHH fJIIOK03bl B Kp0Bl1 B rrep1101l 1lJ111TeJibHOf0 fOJIO)J,amrn 11 11HTeHC11BHbIX <t>11311qecKHX ttarpy30K. Ilpouecc rrpoTeKaeT B ocHOBHOM B rre'leHMM Mettee MHTeHCMBHO B KOPKOBOM Bemecrne ITO'leK, a TaK)l{e B CJI11311CTOH 060JlO'IKeKHllle'IH11Ka.3Tl1 TKaH11 MOryr 06ecne'l11BaTb CHHTe3 80-100 r fJ1IOK03hI B cyrK11. Ha llOJIIO M03ra rrp11 fOJlO]laH1111 npHXOllMTCH 60JlblllaH qacTh rroTpe6HOCT11 opraHM3Ma B fJ1JOK03e. 3TO 06bHCIDieTCH TeM, 'ITOKJieTI<tt: M03fa He cnoco6Hbl, B OTJIM'll1e OT 1lPYfIIX TKaHett, o6ecne'll1BaThTIOTPe6HOCT11 B 3Heprn11 3a C'leTOKHCJiemrn )1{11pHbIX KHCJIOT (CM. pa3]leJI 8).
KpoMe Mo3ra, B rnIOK03e H~aIOTCSJ TKatt11 11 KJieTI<M, B KOTOpbTX a3p06HhlH ITYJb pacna,IJ,a He-
B03MO)l(ett 11JIM orpaHM'leH,ttanp11Mep 3Pl1TPOUHTbI (om1 mnueHbI MHTOXOH]lpttif), KJieTKM ceT'laTKl1, M03fOBOfO CJIOH Ha,L(TIO'le'IH11KOB Jlp.
IlepBH'IHhiecy6cTpaThI rnJOKOHeorette3a - naKTaT, aMHHOKMcnoTbI 11 rnttuepon. BKJ1IO'leH11e3Tttx cy6cTpaTOB B rJIJOKOHeorette3 3aB11Cl1T OT <l>tt3HOJ10fM'leCKOfOCOCTOHH11H opraHM3Ma.
•JlaKTaT - npOllYKT atta3po6ttoro rn11KoJI113a. Ott o6pa3yeTCH nptt JII06bIX COCTOSIHMSIX opraHl13Ma B 3pMTPOUMTaX 11 pa6oTaIOll.l,HX MbIIIIuax. TaK11M o6pa30M, JiaKTaT 11cnOJih3yeTCH B fJllOKOtteorette3e TIOCTOHHHO.
•fn11uepon BhICB060)1{1laeTCH rrpw fHllPOJIH3e )l{HpOB B )1{11p0BOH TKaHM B rrep1101l fOJIO]laHHH HJlH npH MHTeJlbHOH <l>tt3H'leCKOH Harpy3Ke.
•AMHHOKMCJIOThI 06pa3y10TcH B pe3yJihTaTe pacna,n:a Mhlllle'IHbIX6eJIKOB H BKJIIO'laIOTCHB fJ1IOKOHeoreHe3 nptt MHTeJihHOM fOJIO)J,aHHH HJIH IlPOllOJI)l{MTeJibHOH MbIIIIe'!HOHpa6oTe.
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Ha pucyHKe 7-44 noKa3aHbI nYHKTbI BKJI10qe- |
canoauernrn qepe3 M11ToxoH,l1,p11aJihlfYIO MeM6paHy, |
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tt1rn nepBHqHbIX cy6cTpaTOB B rJIIOKOHeoreHe3. |
KOTOPaH MH Hero HenpoH11uaeMa. 0Kca.rroauernT B |
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Ml1TOXOH,l1,pl1aJihHOM MaTP11KCe BOCCTaHaBJI11BaeTCH |
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A. PEAKu:1m: rJIIOKOHEOrEHE3A |
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c o6pa30BaH11eM MaJiarn (p11c. 7-47) np11 yqacT1111 |
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ooJiblllHHCTBO |
peaKUHtt rJIIOKOHeoreHe3a npo- |
NADH (o6parnaH peaKU11H u11Tparnoro u11KJia). |
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06pa30BaBllll1HCH MaJiaT 3aTeM npOXO)ll1T qepe3 Ml1- |
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TeKaeT |
3a cqeT o6paT11MbIX peaKU11H rJI11KOJIH3a |
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TOXOHJipl1aJihttyIO MeM6patty c IlOMOIUhlO cneu11anb- |
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(p11c. 7-45, peaKU11H 9, 8, 7, 6, 5, 4, 2) H KaTaJIH- |
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HhIX nepeHocq11KOB. KpoMe rnro, oKcanoauernT cno- |
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3HpyeTCH TeM11 )Ke cpepMeHTaM11. O;:J,HaKo 3 peaK- |
co6eH TpaHcnopTHpOBaTbCH 113 MllTOXOH)lpl1M B |
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Ul1H rJIHKOJIH3a TepMO)lHHaM11qecKH Heo6paT11Mbl. |
Ul1T030Jib B BH)le acnaprnrn B xo)le MaJiaT-acnap- |
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Ha 3THX CTM11Hx peaKU1111 rJIIOKOHeoreHe3a npo- |
TaTHoro qeJIHO'IHOfOMexaH113Ma, paCCMQTPeHHOro |
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TeKaIOT J:IPYrHMl1 nyrnMl1. |
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paHee (pnc. 7-39). |
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Heo6xo)l11Mo OTMeT11Tb, 'ITOrnHKOJIH3 npoTe- |
B Ul1T030Jie MaJiaT BHOBb npeBpamaeTCH B OKCa- |
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KaeT B UHT030Jie, a 'IaCTbpeaKU11H fJIIOKOHeoreHe- |
JioaueraT B XO)le peaKUl111 OKHCJieHllH c yqaCTHeM |
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3a npoHCXO)ll1T B MHTOXOH)lpHHX. |
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KocpepMeHTa NAD+. 06e peaKU11n: BoccrnHoBJie- |
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PaccM0Tp11M 6onee no)lp06Ho Te peaKU1111 rn10- |
H11e OKCaJioaueTaTa l1 OKHCJieHHe MaJiaTa KaTaJil1- |
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KOHeoreHe3a, KOTOpbre OTJIH'IaIOTCH OT peaKU11H |
3l1PyIOT MaJiaT)lef11)lpOreHa3a, HO B nepBOM cnyqae |
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fJil1KOJIH3a 11 npol1CXOJ]JIT B fJIIOKOHeoreHe3e c l1C- |
3TO Ml1TOXOH)lpl1aJibHbIH cpepMeHT, a BO BTOpOM - |
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nOJib30BaHHeM )lpyrnx cpepMeHTOB. PaccMOTpHM |
Ul1T030JlbHhIH. 06pa30BaHHblH B UllT030Jie 113 Ma- |
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npouecc Cl1HTe3a rnIOK03bI 113 nHpYBaTa. |
JiaTa oKcanoauernr 3aTeM npeBpamaeTcH B cpoccpo- |
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1. 06pa1oeanue <J>oc<J>oenOJ1nupyeama |
eHoJinnpyBaT B XO)le peaKU11ll, KaTaJI113ttpyeMOH |
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¢oc¢oeHoJin11pYBaTKap60Kc11K11Ha3ott - |
IT<ll-3a- |
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UJ nupyeama - |
nepeflR UJ neo6pamUMbtx |
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Bl1Cl1MbIM ¢epMeHTOM (puc. 7-48). Ha3BaHHe ¢ep- |
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cmaiJuii l.!l10Koneoeene1a |
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MeHTa )laHO no o6paTHOH peaKUI1l1. |
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06pa30BaH11e cpoccpoeHOJin11pyBarn |
113 n11pyBarn |
CxeMa Bcex peaKUHH, npoTeKa10iu11x Ha nepBott |
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npol1CXO)ll1T B XO)le )lByx peaKU11H (p11c. |
7-45, peaK- |
Heo6paTl1MOH CTa):ll111 fJIIOKOHeoreHe3a, npe)lCTaB- |
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U1111 11, |
12)' nepBaH H3 KOTOpbIX npoTeKaeT B MHTO- |
JieHa Ha pnc. 7-49. |
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XOH,Il,pHHX. IlttPYBaT'o6pa3yIOIUHHCH l13 JiaKTaTa 11Jll1 |
Cne)lyeT OTMeTHTh, 'ITO3TOT o6xO)lHOH yqacroK |
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H3 HeKOTOPbIX aM11HOKl1CJIOT, TpaHCilOPT11PYeTCH B |
rJIIOKOHeoreHe3a rpe6yeT pacxo)la J:IBYX MOJieKyJI c |
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MaTPHKC MHTOXOH)lpHtt H TaM Kap60KC11J1HpyeTCH c |
MaKp03prH'leCK11MllCBH3HMH (AT<fl I1 IT<fl) B pac- |
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o6pa30BaH11eM oKcanoaueTarn (p11c. 7-46). Il1tpymtT- |
qfae Ha O)lHY MOJieKYJIY l1CXO)lHOf0 Bemecrna - |
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Kap60KcuJJa3a, KaTaJI11311PYIOIUaH )laHHYIO peaK- |
n11pyBarn. B nepecqere Ha c11HTe3 O)lHOtt MOJieKY- |
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Ul1IO, - |
Ml1TOXOH,l1,pl1aJihHbIH cpepMeHT, KOcpepMeH- |
JibI fJIIOK03bl 113 )lByx MOJieKyJI nnpyBaTa paCXO)l |
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TOM Kornporo HBJIHeTCH 6110T11H. PeaKUHH nporeKaeT |
cocTaBJIHeT 2 MOJih A T<fl 11 |
2 MOJih |
fT<fl 11JIH |
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c 11cnoJib30BaH11eM AT<fl. |
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4 MOJib A T<fl (MH y)lo6crna |
pacc~eH11tt npe)l- |
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Li,a.rrhHetfurne npeBpameHHH oKcanoauernrn npo- |
naraeTcH cqnrnTh, qTo 3Hepr03aTpaThr Ha CHHTe3 |
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TeKaIOT B UHT030Jie. Cne)lOBaTeJibHO, Ha 3TOM 3Ta- |
AT<fl H IT<fl paBHhI). IloCJie o6pa3oBaHttH ¢oc- |
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ne )lOJI)l(Ha cyiuecrnoBaTb cucTeMa TPaHcnoprn OK- |
¢oeHoJinttpYBaTa Bee ocTaJihHhie peaKU.HH TaKXe |
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rmoK03A \ |
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~I OKcanoa1.1ernT I •I <l>occpoeHonn1-1pyeaT I |
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,Q1-1r1-1,QpoKc1-1a1.1ernHcpoccpaT |
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AMl-1HOKl-1CJ10Tbl |
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PMC. 7-44. 810110..eHMe cy6cTp8TOB B rmOKOHeoreHe3.
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r----, |
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rmoK03a |
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1_, ___ _ |
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._/H P0 |
I C02 : |
AT<I> |
A,Q<I> + Pi |
C=O |
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CH3 |
CH2 |
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rnlOK030-6-¢oc¢aT |
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n 11pyBaTKap60KCL1fla3a |
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n11pyBaT |
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<l>pyKToso-6-¢oc¢aT |
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PMc. 7-46. 06paaosaHMe 0Kcanoal41!TaTa M3 nMpysaTa. |
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<l>pyKToso-1 ,6-611cqmcQ:JaT
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rn114epan1>.QerM,QQ:JocQ:JaT
611--:~;·
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1-9_':.9_! |
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0Kcanoa4erar |
Manar |
PMC. 7-47. npespa11.1eHMe OKCanoa4eTST8 B ManaT.
PMc. 7-45. rnMKOnMa MrntoKoHeoreHea. <t>epMeHffii-06parnMb1x
peaK4111~ rn111Kon1113a 111 rnlOKOHeoreHeaa: 2 - |
cpoccpormoK0111soMe- |
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pasa; 4 - |
anbAOnaaa; 5 - |
rp111oaocpoccpar1130Mepa3a; |
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6 - rnM4epanbAernA¢occparAerMAporeHaaa; 7 - |
cpoccporn1114epar- |
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KMHaaa; 8 - |
cpoccporn1o14eparMyra3a; 9 - |
eHonaaa. <t>epMeHTbr |
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Heo6paTl1MblX peaK4111~ rnlOKOH00reHe3a: 11 - |
n1o1pyBSTKSp6oKCM- |
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naaa; 12 - |
¢oc¢oeHonn111pyBaTKap60KCMKMH83a; 13 - <PPYKT030- |
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1,6-6111ccpoccpara3a; 14-rnlOK03o-6-cpoccpara3a. l-lll-cy6crpar- Hble l..llllKnbl.
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:coo-: |
rTCIJ |
r---- |
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'co2 : r,o,<i> |
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CH2 |
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CH2 |
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C=O |
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4>occpoeHonn111pysar- |
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coo- |
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Kap60Kc"1KJ11Ha3a |
coo- |
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ClloccjJoeHonn1o1pysar |
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PMC. 7·48. npespa11.1eHMe OKCanoa4eTST8 B cj>OCcj>OeHOnnM· pyBST.
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Acn---> OA---> ¢oc¢oeHonnHpyeaT
ManaT~
P11c. 7-49. 06pa3oeaH11e 0Kcanoa1.1eTaTa, TpaHcnopT e l.IMT030nb 11 npeepall.16HMe e cpoccpoeHonn11pyeaT. 1 - TpaHcnopT n11py0aTa 113 u1-noson5i B Ml1TOXOHJ:1p111t0; 2 - npe0pa11jeH111e n111py0aTa B oKcanoaueTaT (OA); 3 - npe0palJ.leH11e OA B Mana1 11n11 acnap1a1; 4 - TpaHcnop1 acnap1a1a 111 ManaTa 1113 Ml1TOXOHJ:1p111111 B u111rnsonb; 5 - npe0palJ.leH111e acnapTaTa 111 ManaTa B OA; 6 - npe0pa11jeH11e OA s cpoccpoeHonn11py0a1.
rrpoTeKalOT B UHT030.JJ.e BTIJIOTh ).1,0 o6pa30BaHHH <l>PYKT030-l ,6-6Hc<t>oc<t>arn H KaTa.JIH3HPYJOTCH fJUiKO.JIHTHqeCKl1MH <PepMeHTaMH.
2. I'uiJpoAU'J </Jpy1emo30-l,6-6uc<ftoc<ftama u i?.llJOK03o-6-<ftoc<ftama
OnueTIJieHwe <t>oc<t>arnoH. rpyrrnhI H3 <l>PYKT030- I ,6-6Hc<t>oc<t>arn 11 rnIOK03o-6-<t>oc<t>arn - TaK- )Ke Heo6paTHMhie peam1111 rnIOKOHeoreHe3a. B xone f.JIHKO.JIH3a 3TH peaKUHH KaTaJIH3HPYJOT crreu11<t>11qecKl1e Kl1Ha3bI c 11crro.JJ.h30BaH11eM 3Hepr1111 AT<l>. B f.JIIOKOHeoreHe3e OHH rrpoTeKaIOT 6e3 yqacTHH AT<l> 11 AIJ:<l> 11 ycKopHIOTCH He KHHa3aMH, a <t>oc- <t>arn3aMH - <t>epMeHTaMH, rrp11Hau.rre)KaIUHMH K K.JJ.accy rH).l,pO.JJ.a3. <l>epMeHTbI <l>PYKT030-1,6-611c- <t>oc<t>arn3a H rnIOK03o-6-<t>oc<t>arn3a KaTaJIH3HpyIOT OTIUeTIJieHHe <t>oc<t>aTHOH rpynIThl OT <l>PYKT030- l ,6-6Hc<t>oc<t>arn 11 rnIOK03o-6-<t>oc<t>arn. IIoc.JJ.e qero CB060).l,HaH f.JIIOK03a BhIXO).l,HT H3 K.JJ.eTKH B KPOBh. CxeMa Bcex peamwH. f.JIIOKOHeoreHe3a rrpenCTaB.JJ.eHa Ha pwc. 7-45.
lfraK, B rreqeHH cymeCTBYJOT 4 <t>epMeHTa, KOTOpbie rrpl1Hl1MaIOT yqaCTHe TO.JJ.hKO B f.JJ.lOKOHeoreHe3e H KaTaJIH3HPYIOT o6XO).l,Hhie peaKUHH Heo6panl..MhIX CTa).l,HH f.JIHKO.JIH3a. 3To - TTHpyBaTKap60KCH.JJ.a3a, <t>oc<t>oeHO.JIITHPYBaTKap6oKCHKHHa3a, <PPYKT030- l ,6-611c<t>oc<t>arn3a H rn10Ko3o-6- <t>oc<t>arn3a.
3. 3uepzemu11ec1euii 6ll.!lauc 2.ll101Coueozeue3a U'3 nupy6ama
B xo.LJ.e 3TOro rrpouecca pacxo).l,YJOTCH 6 MO.JJ.b AT<l> Ha CHHTe3 1 MO.JJ.h rnIOK03hI H3 2 MO.JJ.h rr11-
pyBarn. qeThipe MO.JJ.b AT<l> pacxonyIOTCH Ha crn- ).1,HH CHHTe3a <t>oc<t>oeHO.JIITHpyBaTa 113 OKCaJIOauernrn 11 eme 2 MO.JJ.h AT<l> Ha CTa).l,HHX o6pa30BaHHH 1,3-611c<t>oc<t>orn11ueparn 113 3-<t>oc<t>o- rnwuepaTa.
CYMMapHbIH pe3y.JJ.hTaT rmoKoHeoreHe3a H3 rr11PYBaTa Bhipa)KaeTrn c.JJ.eJJ.YIOIUHM ypaBttett11eM:
2 IlHpysaT + 4 AT<l> + 2 IT<l> + 2 (NADH+ H+) + 4 Hp -t fmoK03a + 4 AIJ:<l> + 2 f)l<l> +
6 H3P(\ + 2 NAO+.
Ii. CMHTE3 rJIIOK03hl 113 JIAKTATA
JlaKTaT, o6pa3oBaHHhlH B aHa3p06HOM fJIHKOmne, He HBIDl.eTCH KOHeqHhIM rrpO).l,j'KTOMMeTa60.JIH3Ma. llcrrO.JJ.h30BaH11e .JJ.aKTaTa CBH3aHO c ero rrpeBpameHHeM B rreqem1 B ITHPYBaT. JlaKTaT KaK HCToqHHK rrHpYBaTa Ba)KeH He CTO.JJ.hKO rrp11 ro.JJ.o- naHHH, CKO.JJ.hKO rrp11 ttopMa.JJ.bHOH )Kl13tte;neHTe.JJ.h- HocTH opraHH3Ma. Ero rrpeBpameH11e B TTHPYBaT H JJ.aJihHefnuee HCI10.JJ.h30BaHHe ITOC.JJ.e).l,Hero HBllil.IOTCH crroco6oM YTH.JIH3aUHH naKrnrn.
JlaKTaT, o6pa30BaBiliHHCH B HHTeHCHBHO pa6oTalOIUHX MhIIliUax HJIH B K.JJ.eTKax c rrpeo6Jia).l,aIOIUHM aHa3p06HhIM crroco6oM KaTa60.JIH3Ma f.JIIOK03bl, rrocrynaeT B KPOBb, a 3aTeM B rreqeHh. 8 rreqettH OTHorneH11e NADH/NAD+ HIDKe, qeM B coKpammomeifcH Mhirnue, no3TOMY naKTaT).l,erl1).l,poreHa3HaH peaKUHH npoTeKaeT B o6paTHOM HanpaBJieHHH, T.e. B CTOPOHY o6pa3oBaHHH rr11pyBaTa H3
.JJ.aKrnrn. )lanee rrHpYBaT BK.JJ.10qaeTCH B fJIIOKoHeoreHe3, a o6pa30BaBiliaHCH fJIIOK03a IlOCTYTTaeT B KpOBb H nornomaeTCSI CKeJieTHbIMH MhIIliUaMH. 3ry
347
IIOCJie,UOBaTeJibHOCTb C06bITHH Ha3bIBaJOT «rJIIOK0- 30-JIRKTaTHblM U,HKJIOM», HJIH «U,HKJIOM Kopu» (pwc. UwKJI Kopw Bbmonmi:eT 2 BIDKHe:H- urne cPYHKUHH: 1 - 06ecrre1mBaeT ynmmaumo naKTaTa; 2 - rrpe,aornpamaeT ttaKoIIJiemrn naKTma H, KaK CJie,UCTBHe 3TOfO, orracttoe CHmKeHHe pH (JiaKTOaUH,!J.03). lfacTb IIHpyBaTa, o6pa30BaHHOfO H3 JiaKTaTa, OKHCmieTCH rre11eHblO ,UO co2 H H20. 3tteprHH OKMCJieHHH MmKeT HCIIOJib30BaTbCH ,UJIH CHHTe3a AT<l>, Heo6xo,awMoro ,UJIH peaKUMH rnioKotteorette3a.
JlaKTOalJ,ll,!J;o3. TepMHH «aUH,U03» 0603Ha11aeT yseJIHIJ:ettwe KHCJIOTHOCTH cpe,Ubl opraHH3Ma (CHmKeHMe pH) ,UO 3Ha11eHHH, BbIXO,UHill,HX 3a rrpe,aeJibl HOpMbl. flpw a11,H,U03e JIH60 yseJIHlJ:HBaeTCH rrpo- ,UYKUHH rrpoTOHOB, JIH6o rrpOHCXO,UHT CHH)l(eHHe HX 3KCKpell,HH (B HeKOTOpbIX cnyqaHX M TO M ,apyroe). Mern6onw11ecKH:i1 auw,ao3 B03HHKaeT rrpw yseJIHIJ:eHMH KOHUeHTpaUHH rrpoMe~OlJ:HbIX rrpo- ,UYKTOB o6MeHa (KMCJIOTHOfO xapaKTepa) BCJie,U- CTBHe yseJIHIJ:eHHH HX CHHTe3a HJIH yMeHhIIIeHHH CKOpOCTM pacrra,aa HJIH BbIBe,UeHHH. flpH HapyIIIeHHH KMCJIOTHO-OCHOBHOfO COCTOHHHH opraHH3Ma 6bICTpO BKJIIOIJ:alOTCH 6y<t>epHbie CHCTeMbl KOMIIeH-
cauww ('-1epe3 MHH). Jiero11HaH KOMnettca-
UHH o6ecrre11HBaeT CTa6HJIH3aUHIO COOTHOWeHHH HC03-/H2C03, KOTopaH B HOpMe coorneTcrnyeT
1:20, a rrpH auH,U03e yMeHbIIIaeTCH. Jiero11HaH KOMneHCaUHH ,UOCTHraeTCH yseJIMIJ:eHHeM 06'beMaBeH-
THJIHUHH H, CJie,UOBaTeJibHO, ycKopem1eM BbIBe,UeHHH C02 H3 opraHH3Ma. 0,attaKO OCHOBHYJO pOJib B KOMIIeHCaUHM all,H,U03a HrpalOT II011e11Hbie MexaHH3Mbl c yqaCTHeM aMMHaIJ:HOro 6y¢epa (CM. pa3- ,UeJI 9). 0,UHOH H3 rrpH11HH MeTa60JIH11eCKOfO arrn- ,U03a MO)l(eT 6bITb HaKOIIJieHHe MOJIOIJ:HOH KHCJIOTbl.
B ttopMe naKTaT B rre11eHH rrpeBpamaeTcH o6paTHO B fJIIOK03y rryTeM rJIIOKOHeoreHe3a JIH60 OKHCAAeTCH. KpoMe rre11eHH, ,UPYTHM rrOTpe6wTeJieM naKTaTa CJIY)i(aT IIOIJ:KM H cep,aeIJ:HaH MbIIIIUa, r,ae JiaKTaT
MO)l(eT OKMCAATbCH ,UO C02 M H 20 H HCIIOJib30BaTbCH KaK HCTOIJ:HHK 3HeprMH, oco6eHHO npH cPH3HIJ:eCKOH pa6oTe.
YpoBeHb naKTaTa B KPOBM - pe3yJihTaT pasHOBeCHH Me)l(,!ly rrpoueccaMH era o6pa3oBaHHH H yTHJIH3aUMH. KpaTKOBpeMeHHbIH KOMrrettcwpoBaHHbIH JiaKTOaUH,!J.03 BCTpe11aeTCH ,UOBOJibHO '-la- CTO ,Ua)l(e y 3,UOPOBbIX JIIO,Ue:i1 rrpw MHTeHCHBHOH MbIIIIeIJ:HOH pa60Te. y HeTpeHHPOBaHHbIX JIIO,Ue:H JiaKTOall,H,!J.03 rrpM cPM3H11eCKOH pa60Te B03HHKaeT KaK CJie,UCTBHe OTHOCHTeJibHOfO He,UOCTaTKa KHCJiopo,aa B MbIIIIUaX H pa3BHBaeTCH ,UOCTaTOIJHO 6bICTpo. KoMrrettcauwH ocymecTBJIHeTcH rrYTeM rnrrepBeHTHJIHUHH.
IlpH HeKOMIIeHCHpOBaHHOM JiaKTOaUH,U03e co- ,aep)l(aHHe naKTaTa B KPOBH yBeJIH'-IHBaeTCH ,ao
5 MMOJib/JI (B HOpMe ,UO 2 MMOJib/JI). IlpH 3TOM
pH KPOBH MO)l(eT COCTaBJIHTb 7,25 M Mettee (B HOpMe
7,36-7,44).
PMc. 7-50. UMK11 KopM (rnt0Koao-naKTaTHblM l.IMK11). 1 - nocl)'nneH111enaKTaTa 1113 c0Kpall.lato111ei1csi MblWL.lbl c TOKOM Kpos111 B ne<ieHb; 2 - ClllHTe3 rntoK03bl 1113 naKTara B ne<ieH111; 3 - nOCT)'nneHl!lerntoK03bl 1113 ne<ieHlll c TOKOM KpOBlll B pa60TBIOlllYIO MblWL.ly; 4 - 111cnonb30BaH111e rntoK03bl KaK aHeprern<iecKoro cy6crpara c0Kpall.lato111eiksi MblWL.lei1 111 06pasosaH111e naKTaTa.
348
floBhilllem1e co,nep:)l<amrn naKrnrn B KPOBH MO-
)l(eT 6hITh cne,ncTBHeM ttapyrnemur Mern6omnMa nttpyBaTa (pHC. 7-51).
naKrnT
111
n11pysaT~ --- t illlOK038
n,QK f2 1
A1..1ern11-KoA
I
I
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PMc. 7-51. HapyweHMR MeTa6onM3Ma nMpyBaTa npM naKToa- l.IMA03e. 1 - HapyweH11e 11cnonb30BaH1M1 n11pysarn B rmoKoHeoreHese; 2 - HapyweH11e oK11cneH11R n11pysarn.
TaK, np11 mn0Kc1111, B03H11Ka10met1 BCJieLJ:CTB11e Hapyrnemrn CHa6)KeHJ.rn TKaHetl KMCJIOPOLJ:OM 11Jil1 KPOBhIO, YMeHhrnaeTCH aKTl1BHOCTh n11pYBaT.uerHLJ:- poreHa3Horo KOMnJieKca H CHlf)l(aeTCH OKMCmtTeJihHOe ,neKap60Kc11nttpoBaH11e n11pYBaTa. B 3THX ycJIOBl1HX paBHOBec11e peaKU1111 n11pyBaT~JJaKTaT
CLJ:Bl1HYTO B crnpoey o6pa3oBaHHH naKTaTa. KpoMe Toro, np11 r11n0Kc1111 YMeHhrnaeTcH c11ttTe3 ATCI>,
'lTOCJie,UOBaTeJJhHO, Be,UeT K CHJ1)KeHl1lO CKOpOCTl1 rn10KoHeoreHe3a - .upyroro nym yr11n11Jau1111 naKrnrn. 00Bh1rnett11e Kottuettrpau1111 naKTaTa 11 ctt11- )Kett11e BttyTpl1KJJeTO'lHOfOpH OTp11UaTeJJbHO BJil1HIOT Ha aKTHBHOCTh Bcex cpepMeHTOB, B TOM '111CJie 11 n11pyBaTKap60KCHna3hl, KaTaJJl1311PyIOIUetl Ha'laJih-
HYIO peaKUHIO rn10KoHeoreHe3a.
Bo3Hl1KHOBett1110 naKToauHLJ:o3a TaK)l(e cnoco6crny10T ttapyrneHHH m10Kotteorette3a rrp11 ne'leH01:1- Hotl He,UOCTaTO'lHOCTl1pa3JJHlJHOfO np011CXO)K,UeHHH. KpoMe Toro, naKToauHLJ:030M MO)l(eT corrpooo)K,UaThCH r11rroBHTaM11H03 B 1, rnK KaK rrpo113BOLJ:HOe 3Toro Bl1TaMl1Ha (T11aMl1H,Ul1cpoccpaT) BhIIIOJIWleT KOcpepMeHTttylO cpyHKUHIO B COCTaBe Il,l],K rrp11 OKHCJil1TeJihHOM ,UeKap60KCl1Jil1pOBamrn rr11pyBaTa (CM. pa3,Uen 6). D,ecpHUMT THaM11Ha MO)KeT B03Hl1KaTh, ttarrp11Mep, y aJIKOfOJil1KOB c HapyrneHHhIM pe)!(H- MOM Ill1TaHJ·IB.
l1TaK, rrp111:111HaM11 HaKOT1JleHl15I MOJIO'lHOHKHCJJOThI 11 pa3Bl1THH naKTOaUH,U03a MOryr 6hITh:
• aKT11Baum1 atta3po6ttoro rn11KoJI113a Bcne.u- CTB11e TKaHeBOH fl1T10KCl111 pa3JIH'IHOf0rrpoHCXO)l(LJ:ettm1;
•nop~eHHH ne'lemt(TOKCHlJeCKHe ,Ul1CTpocpH11, UHppo3 11 ,Up.);
•HapyweHHe HCT10Jlh30BaHHH JiaKTaTa BCJie,U- crn11e ttacne.ucrneHHhIX ,necpeKToB cpepMeHTOB fJJIOKOHeoreHe3a, He,UOCTaTO'lHOCTH fJIIOK030- 6-¢oc¢arn3hI;
•ttapyrnem1e pa60Thl CTD,K Bcne.ucTBHe ,ne¢eKTOB cpepMeHTOB HJIH fHT10BHTaMHH030B;
•npHMettett11e pH,Ua neKapcrneHHhIX npenapaTOB, HanpHMep 6HryaHH,UOB (6JIOKaTOphI fJIIOKOHeoreHe3a, ttcrroJih3yeMhre npH ne'leHHH caxapttoro .u11a6ern).
B.CHHTE3 fJIIOK03hl H3 AMHHOKHCJIOT
B ycnoBHHX rono.uaH11H 1:1acTh 6eJIKOB Mh1llle1:1- Hoti TKaHH pacrra.uaeTCH ,no aMHHOKHCJIOT, KOTOpble .uanee BKJJIO'laIOTCSI B rrpouecc Karn6on113Ma. AMMHOKHCJIOThI, KOTOpbie npH Karn60JIH3Me rrpeBpamaIOTCH B nttpYBaT HJIH MeTa60JJHThl UHrpaTHOro UMKJJa, Moryr paccMarpHBaThCH KaK noTeHUMaJihHbie rrpe.urnecrneHH11Kl1 rJIIOK03bI H fJIHKOfeHa H HOCHT Ha3BaHMe fJIHKOreHHhlX. HanpHMep, OKCaJioauernT, o6pa3yIOIUHHCH H3 acrraparHHOBOM KMCJIOThl, HBJIHeTCH npoMe)l(yTO'lHhlM rrpo.uyKTOM KaK u11rpaTHoro u11KJJa, TaK 11 rn10KotteoreHe3a.
113 Bcex aMMHOKHCJJOT, noczyrra10m11x s ne'leHh, npHMepHO 30% rrpMXO,UMTCH Ha ,UOJIIO anaHHHa. 3TO o6bHCHHeTCH TeM, ':!TOnpH pacmeT1JleHHH MhIWe':l- HhIX 6eJIKOB o6pa3ylOTCH aMl1HOKMCJIOTbl, MHOr11e 113 KOTOphIX npeBpama10TCH cpa3y B rr11pyBaT HJIH CHa'laJiaB OKCaJIOaUeTaT, a 3aTeM B T111pyBaT. CTocJie,LI,Hl1H rrpeBpamaeTCH B anatt11H, rrp1106peTaH aMHHOrpynny OT .upyr11x aMHHOKHCJJOT. AnaHHH 113 MhlWU nepeHOCl1TCH KPOBhlO B ne'leHb,r.ue CHOBa npeo6pa3yeTCH B rrnpyBaT' KOTOpbIH 1:1aCTH':IHO OKHCJIHeTCH 11 1:1aCTl1'lHOBKJIIO'laeTCHB fJIIOKOHeorette3. Cne.uoBaTeJihHO, cymecTByeT cne.uyIOIUaH nocJJe,UOBaTeJJhHOCTh C06hITl1M (rJllOK030-a.JiaHHHO- BhIH U,UKJI): fJIIOK03a B MhIWUaX ~ I111PYBaT B Mhllliuax ~ anaHHH B MhlWUaX ~ aJiaHl1H B rre'leHH~ rnlOK03a B ne'leHH ~ l'JJIOK03a B MbIIlIUaX (pnc.
7-52). Bech UHKJJ tte np11Bo,n11T K YBeJI11'1eH1110KOn11qecTBa fJIIOK03hI B MhIIlIUax, HO OH pelllaeT npo6JieMhl TpaHcnopTa aMl1HHOfO a3oTa 113 MhlilIU B rre'leHh11 npe.uornpamaeT naKToauHLJ:o3.
r. C11HTE3 rn10Ko3b1 113 rn11u:EP0JIA
fn11uepon 06pa3yercH rrp11 fl1Ll:pon113e rp11au11nrnm1eponoB, rnaBHblM o6pa30M B )!(HPOBOH TKaH11. l1cnOJih30BaTb ero MOryr TOJihKO Te TKaH11, B KOTO-
349
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------- rnlOK03a |
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n111pyeaT |
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P11c. 7-52. rmoKoao-anaH11HOBb1iil 1.111Kn. |
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phIX ttMeeTcsi ¢>epMeHT nrnuepon KHHaJa, HarrpttMep rreqeHh, rroqKJI, 3ToT AT<l>-3aBHCHMhIH ¢>epMeHT KaTaJIR3ttpyeT rrpeBpalll,em1e rnwuepona B a-rnwuepo¢>oc¢>aT (rnwuepon-3-¢>oc¢>aT). Ilptt BKmoqeHttH rnrruepon-3-¢>oc¢>arn B rmoKoHeoreHe3 rrpowcxomrr ero .uenmpwpoBaHtte NAD-3aBH- cwMoi1 .uem.uporetta3oi1 c o6pa3oBaHtteM .urrnmpoKcrraueTott¢>oc¢>arn (pttc. 7-53), KOTOphIH .uanee rrpeBpalll,aeTCSI B rJIIOK03Y.
x. PErYn~u~~ rn~KOfl~3A
~ rntOKOHEOrEHE3A
B nE'-IEH~
Ilo cpaBHeHmo c .upyrttMR oprattaMH rreqeHh OTnwmeTCSI Hatt6onee CJIO)J(HbIM o6MeHOM rJIIOK03hI. KpoMe rrapw rrpOTttBorrono)l(HhIX rrpoueccoB (CHHTe3a H pacrra.ua rnttKoretta), B rreqeHtt MOryT rrpottcxO.IJ:HTh elll,e .IJ:Ba rrpOTRBOIIOJIO)!(HO HarrpaBJieHHhIX rrpouecca - rJIHKOJIH3 H rJIIOKOHeorette3.
B 6oJihlllHHCTBe .upyrRX oprattoB rrporrcxo.uRT TOJihKO rJIHKOJIR3. IlepeKJI10qeHtte rreqeHR c rJIHKOJIH3a Ha rJIIOKOHeorette3 H o6parno rrpottcxomrr c yqacrneM HHCYJIRHa H rJIIOKaroHa H ocymecTBJISieTCSI c IIOMOlll,hIO:
•aJIJIOCTeprrqecKOH peryJISIURH aKTHBHOCTH ¢>epMeHTOB;
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PerymnopHhie B03.Uei1cTBHSI HarrpaBJieHhI Ha ¢>epMeHThl, KaTaJIH3RPYJOIUHe Heo6paTHMbie CTa-
.URH rJIHKOJIH3a H rJIIOKOHeoreHe3a, coqeTaHHe
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A. PErYJUIUHH CKOPOCTH PEAKU,HH
fJUIKOJIH3A H rJIIOKOHEOfEHE3A,
COCTABJIHIOllJ,HX CYBCTPATHhIE UHKJihl
«Cy6C'rpaTHbie» IJ,llKJlbl - rrapHhie KOM6HHaURH rrpoueccoB crrme3a H pacrra.ua MeTa60JIHTOB. KaK ~e yrroMHHaJIOCh, coqernttHe rrpoueccoB c11me3a ll pacrra.ua rJIRKOreHa HJIH Heo6paTHMhIX peaKURH rJIHKOJIH3a H COOTBeTCTBYJOIURX HM Heo6paTRMhIX peaKUHH rJIIOKOHeoreHe3a MO)!(eT COCTaBRTh rro.uo6HhIH UHKJI. Ha3BaHHe «cy6cTpaTHhii1 UHKJI» 03HaqaeT o6be.IJ:ttHettwe peaKUHH CHHTe3a 11 pacrra.ua cy6cTparn. Ha3Battwe «XOJIOCTOH» oTpIDKaeT pe3yJihTaT pa6oTbI rro.uo6ttoro UttKJia, JaKJI10qa10IUHttCS1 B 6ecrrone3HOM pacxo.uoBaHHH AT<l>. Xorn cyw,ecrnoBaHtte «XOJIOCThIX» UlfKJIOB HeJIOrRqHO, TeM He Mettee OHH MOrYT <l:>YHKUHOHttpoBaTh. Eonee Toro, 3Tlf UlfKJibI MOryT 6bITh MHllleHhIO peryJISITOpHbIX B03-
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l13MeHeHRe B rreqeHR rJIHKOJilfTiflleCKOro HarrpaBJieHRSI Ha rJIIOKOHeOreHe3 H o6paTHO rrpw CMeHe a6cop6TlfBHOro COCTOSIHRSI Ha IIOCTa6cop6THBHOe HJIH rrpw rono.uaHlflf rrpOlfCXO.IJ:HT rnaBHhIM o6pa30M B pe3yJihTaTe peryJifilIHH aKTRBHOCTH ¢>epMeHTOB, KaTaJIR3HPYJOIUHX peaKURH cy6CTpaTHhIX UHKJIOB. 3TH UHKJihI o603ttaqeHhI u11¢>paM11 I, II, III Ha pwc. 7-54, rrpe.ucTaBJISIIOIUeM o6IUYID KapTHHY peryAAURH rJIHKOJIH3a H rJIIOKOHeoreHe3a B rreqeHH.
350
HanpaBJieuue peaKIUIH nepsoro cy6crpa111oro
mt perymrpyeTCH rnaBtthIM o6pa30M KOHUeHTPau11etl: fJllOK03bl. flpH ilHlll,eBapeHHH KOHUettTpaUI151 nnoK03bl B KPOBH noBhilllaeTrn Cao 8-10 MMOJih/II). AlcrttBHOCTb rJIIOKOKl1Ha3bl B 3Tl1X ycJIOBl15IX MaKCHMaJihHa. BcJieLJ:CTBHe 3Toro ycKopHeTCH rnHKOJIHTWiecKM peaKUHH o6pa3oBattttH rn10K03o-6-cpoc- cparn. KpoMe Toro, Httcymm HHAyu11pyeT CttHTe3 fJIIOKOKHHa3bl l1 ycKop5IeT TeM caMbIM cpoc¢opnJIHPOBaHHe rn10K03hl. ITocKOJihKY rnIOKOKllHaJa neqeHH He llttrtt6npyeTC51 rJIIOK030-6-cpoccpaTOM (BOTJI11qne OT reKCOKlltta3bl Mhilllll,), TO OCHOBHaH qacTh rnJOK03o-6-cpoccparn B a6cop6nrnHOM rrep110LJ:e HanpaBIDieTCH Ha CHHTe3 fJlllKOreHa l1 ITO fJIHKOJIHTllqecKOMY nyTH.
HanpaBJieHUe peaKU,Uii BTOporo cy6crpa111oro IJ)IK- mt 3aBHCHT OT aKTHBHOCTH cpoccpocppyKTOKl1Ha3bl II ¢oc¢aTa3hl cppyKT030- l ,6-6ttccpoccpaTa. AlcrHBHOCTh 3TIIX cpepMeHTOB 3aBHCHT OT KOHll,eHTpaUlllI ¢pyK- T030-2,6-6HccpoccpaTa. <l>pYKf03o-2,6-6uc<J>oc<J>aT - MeTa60JIHT, o6pa3yIOlll,HHC51 B He3HaqHTeJibHbIX KOJIWiecrnax 113 ¢PYKT03o-6-¢occparn 11 BhIIIOJIIDIIOmtttl: TOJibKO peryAATOpHhie cpyttKUIIH. 06pa30Bam1e cppyKTo3o-2,6-611ccpoccparn nYTeM ¢occpopl:IJIHPOBaHH51 cppyKT03o-6-cpoccparn KaTaJIH3ttpyeT 6n<J>yuKQHoua.rn.Hbill: <J>epMeHT (BH<I>), KOTOpblH KaTaJIH3ttpyeT TaIOKe 11 o6paTHyio pemru,mo (pnc. 7-55, A). OLJ:HaKO npeBpalll,eHHe ¢PYKT03o-2,6-611c¢oc¢arn B <PPYK- T03o-6-cpoccpaT tte HBmreTcH o6paTttMhIM rrpo11,eccoM. 06pa30BaHHe cppyKT03o-2,6-6nccpoccparn Tpe6yeT 3aTpaT AT<I>, a rrpn 06pa30Batt1111 ¢pyKTo3o-6-cpoc- cparn 113 cppyKT03o-2,6-6Hccpoccparn rttLJ:POJIHTttqecKH OTIUenJIHeTCH tteoprattmecKIIii ¢occpaT.
B peaK1lttl1 cpoccpop11JI11poBaHHH <PPYKTmo-6- cpoccpaTa cpepMeHT rrp05IBJI5leT Kl1Ha3HYJO aKTHBHOCTh, a rrp11 J.J,ecpoccpop11Jitt:poBaHHH o6pa30BaHHOfO ¢PYKT03o-2,6-6nccpoccparn - cpoc¢aTa3HyIO. 3rn o6cT051TeJibCTBO l1 onpeneJIHJIO Ha3BaHHe ¢epMeHTa «6HcpyttKl1110HaJihHhlH».
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353
12-2128