YK 578.1 BBK 28.072873 Hag Cepus ocnoeana 6 2006 2. Heascon JI. H49 ——Octosst GHoxumun Jlenutuxepa : a 37. T.2 : BuosnepretKa v meraGosusm / JI, Hescou, M. Koxe ; nep. ¢ ann. —M. : BHHOM. JiaGopatopns ananuii, 2014. —636 c. : uz. —(Jlysumit sapybexxetit yueGunk). ISBN 978-5-94774-366-1 (T. 2) ISBN 978-5-94774-364-7 B yweGHoM HaNaHHH, HaNHCAMHOM aMepHRAHCKHMIt YEHBIMH, KOTOPHE TO- “YM SceOBUlee MPHIHAHMe KAK TANAHTANBKIE MpenonasaTeNH yHuBEpCHTETCKO- TO YPOBHR, PACCMOTPCHBI COBPEMCHHME KOHILEMIAH GHOXHMMM B COOTRETETBHK ¢ waMexMoulelicn nneonorHell ato HayKH. B tom 2 souina wacrs I «Buosneprernka # meTaGonusv0». Jans oGune TepMORHHAMMYECKHE TOHATHA MpHMEHHTEMbHO K GHONOTHYECKHM cHCTEMAM, KTaCcHpHKALWNR XHMHYECKHX PeaKUMi, MPOHCXONALLX 8 >KHBBIX OPraHHsMAX, nonpo6Ho paccMOTpeHt OCHOBHBIE MeTAOOMHYECKHe MYTH —FAKKOMHS, TMOKO- HeoreHes, NeHTosopocbarHstii nyt, lIMKi AHMORHOH KHeAOTE, KaTaGOmHSM XoTo- ocoprmnposanne, mpouecce! GHocHHTesa H jerpanalHH CCHOBHAIX 6HOMO- JeKya, 8 TOM YMCNe XHPOR, & TAKKE NPHHIANA perynsuMH MeTaGoniaa, ‘Zl crynertos GHONOTHNeCKHX, XHMHYECKHX H MCJIMILHHCKHX BYI0B, @ TaIOKe naysnbix. paGOTHHKon, YAK 578.1 BBK 28.072873 Tlepsstii rpax ocyutecranen pH dunancosolt nomtepxKe Pocewiicxoro owe dysaaMentansbx uecnenoBannli no mpoexry Ne 06-04-62066 Yuebnoe wsdanue Cepus: «lysumnit sapySexnsiit yseGunn Heascon Js, Koxe Malin OCHOBbI BHOXHMHH JIEHHHJDKEPA B tpex Tomax Tom 2 BHOSHEPFETHKA H METABOIH3M Beayumit penaxrop kana. xum. nayk TH, Mowxacea. Penaxtop wana. 6uon. wayx TE, Toncmuxuna Xynoxnux HA. Hosax. Texunuecknii penaxtop E. B. Jlewoxoea. Koppextop Jl. H. Mypadan Komnpiotepiian weperka: 7. 9. Buyxooa Tloannearo 8 nevars 27.08.13. Dopmar 84x 108/16. Yen. new. 1. 67,20. Tupax 1700 9x3. 3aKa Hanatensctso «BMHOM. JlaGopatopus sani» 125167, Mocksa, mpoesa Aaponopra, 2. 3 Tenecpou: (499) 157-5272, e-mail: binom@Lbz.ru, http:/www.Lbz.ru First published in the United States by W.H. FREEMAN AND COMPANY, New York Copyright © 2008 by W.H. Freeman and Co. All rights reserved. ISBN 978-5-94774-366-1 (T. 2) © Mepeson wa pycennii sxsux, ISBN 978-5-94774-364-7 BHHOM. JlaGopatopua snannii, 2014YACTH II Buo3sHepretnka Mv MeTa6onn3M JesTeMbHOCTb KeTKH, MPH KOTOpoit Mponc- XOJMT KOOMIepaTHBHOe BIaMMOTeHICTBHE MHO- THX MyssrHcbepMeHTHBIX cHicTeM (MeTaGomHeckHe nyt) A18 Toro, ToGn: (1) HSBIEND XHMITYECKYIO aHepriio H3 oKpyxkaloulelt cpenbi (s1H60 nyTeM T10- HIOMLeHHS BHEprHH CoMHesHOTO CBeTa, M60 MpH Ae rpagaunn 6oraTbix aHeprieli mratesbHBIX BenLeCTB); (2) npesparirrs Mouexy.1b1 murrarembHBIX BeutecTB B coBcTBeHHBIe, XapakTepHBIe ALIA KIETKH MOmEKY.II, BIOMOYA MpeNeCTBEHHHKH MakpomozeKyzt; (3) H3 MpemuuecTBeHHHKOB-MOHOMepOB mMpoBecTH cGOpKy MakpoMorlekys1: GesIKOB, HYKJIeHHOBBIX KHCJIOT 1 110- amcaxapujios; n (4) ocymectBHTb CHHTes H erpana- M eraGou143M — 9TO BBICOKOKOOpAMHMpOBAHHas 13 OcHosei 6uoaHepreTHKn. Tunbi xuMMYeCKHX peakunih 11 14 [nwkonw3, rntokoHeoreHes W MeHTO30pocdatHbii nyTb 65 15 Mpwxyunet perynaynn meta6onu3ma 121 16 Ukr nMMOHHOW KMCHOThI 181 17 Kata6onn3m KMpHbIxX KMCNOT 225 18 OkvcneHMe KMpHBIX KMCHOT 4 O6paz0BaHne MoueBHHbI 261 19 OkucautenbHoe cboctbopunuposaHue wu cotococtopunnposanve 305 20 Buocuutes yrnesonos y pacrenui u 6aktepni 403 21 Buocuute3s nunugos 445 22 BuocuHte3 aMMHOKMCAOT, HYKM@OTHAOB M CBAAHHBIX (WX MeTa6onM3MoM MoneKyn 505 23 Mxterpayua m ropMonanbuaan perynayna Metabonu3ma ymnekonutaiuyx 565 UHO GHOMOTeKys1, HEOOXOAMMBIX KJIETKE JIA BBITION- HenHa ee byHkunii, — mimi08 MemOpan, BHYTpH- KJIETOUHBIX MECCEHIDKEPOB Ht IMT MeHTOB. MeraGosusm BkOYaeT COTHH pasMYHBIX Ka- Tanuanpyembix dbepMextamit peauuii. B aroii wac- TH KaMrH (4acrs IT) MBI paccMoTpHM LeHTpambHBIe MeraGomMYeCKHe MYTH, KOTOPHIE COBCEM He CTONb MHOTOUHCAEHHDI H YAMBHTEABHO CXO/HBI Y BCEX KH- BbIx (opm. IKuppie opranuamer MoxkHO pagnenitTb Ha ape GoXbIIMe Tpynnsi B 3aBHCHMOCTH OT Toro, B BUME KAKHX MONEKY. OHH NosyYAIOT yrnepon U3 oxpykatomteti cpenbi. ABrorpodpss (Takue Kak ¢poTo- CHHTesHpyloulme 6akTepHM M COCyAMCTbIe pacTeHHA) B KaecTBe €MHCTBEHHOTO HCTOUHHKa yrleposla MO-[6] Yacte IT. BmoaHeprerika u MeraGonnam Oo; e ~¥ pymaiesmorpob Terepoms bb co, Puc. 1. Kpyropopor yrnekucnoro raza M KACnOpona Me>KAy astorpoueim (‘porocnuTespyiouyny) 4 reteporpodHbin nomenann (akocucremann) 6Hocipepet. B stor xpyrosopor ‘BOBNeYEHE OrpoMHTe Macht BeMKECTB; 6HOCepHENK O6opor ouexnpaerca B ~4 - 10" r yrnepona 8 ron. TYT HCNOMb3OBATH AHOKCH/ yruiepota armoccepbi, 3 KOTOPOTO OHH CTPOAT Bee CBOH yrseposCOsep%Ka- ute 6uOMOneKy.1b1 (cM. puc. 1-5). Hexoropsie apro- Tpobabie opraniamel, HanpHMep mHanoGaxrepin, MOFYT TaK2KE HCTONBSOBATH aTMOCtpepHEIii as0T ATA TOFO, YTOGbI MPOH3BOANTb BCE CBO MOJEKYJBI, KO- Topple conep>kat asor. Tereporpodpst He MoryT Hc- ToNb3OBATS YrsleKHcmTBIi a3 aTMoctbepsl 1 [OZDKHDL nosy4atb yrnepon 3 OKpyxkalouleli cpest B ope OTHOCHTENBHO CIOXKHBIX OPraHHyecKHX MOJeKys1, TaKHX Kak TJ0KO3a. MHOrOK/eTOUHbIe %KMBOTHBIC H 6obUIMHCTBO MHKPOopraHHsMoB — reteporpodti. ABTOTpocHble K1eTKH H OpraHHsMbI OTHOCHTeTBHO CaMOAOCTATOYHEI, B TO BpeMA Kak reTepoTpocpHbie KJIETKH H OPFaHH3MBI JUIA YsOBsETBOPeHHs CBOHX norpe6xocteli B yrnepone HyksIa10TCa B Goulee C10%K- HBIX OPraHHYECKHX MOMEKYsIaX HM MOSTOMY JOJDKHBL TMTATBCA MPOAYKTAMM 2%KH3HEeATEABHOCTH ApyrHxX opraHusMos. Muorue aprorpodrule oprannamst Hcnomayior cboTocuntes, rile B KaveCTBe MCTOYHHKa SHeprHn pa- 6oraer connesHttit cpet. lereporpodpHbie opraHnsMbt Hosy'4aloT aHepriio MpH paculervienMn OpranHyecknx luraTenbHBIx BeMteCTB, Obpasyembrx aBtorpocbamn. B Guoccepe asrorpodpst H rereporporbbi *KUBYT BMeECTe 1 GombILIOM BaaHIMO3ABHCHMOM Lute. ABTOTPOSHDIE OpraHH3MBI JUIN NOCTPoeHHA CBOMX OpraHHeckHx Guomonexy. ucnombaylor yraeKHcmbi ras armoctbe- pbi. B aTom mpouecce HeKOTOpbie aBTOTPOrpbi TeHepH- pylor Kucaopon 13 Bost. Fereporporpet 6 KaecrBe TMTaTeNbHBIX BeMLECTB HCTOsMb3yWT OprannyeckHe MpoayKTH1, O6pasyemute aBrorpocbaM, t BELETAIOT B aTMmoc(pepy yraexncapiii ras. Ja HeKOTOpEIX OKHCTH- ‘TeNbHBIX peakUnil c O6pasoBaHHeM AMOKCHMIa yrsiepo- 4a HeobxO_MM KHC/OPO,,, KOTOpbIii B PeaKIHAX OKHC- sleHHA Mpespauiaerca B Bosly. TakHM O6pasoM, Mexuty TeteporpodHBIM MH ABTOTpodHBIM. ecTecTBeHHBIMH SKOCHCTEMaMM NOCTOAHHO MPOHCXOAHMT KpyroBopoT yraeposta, kucsopojta 1 Bost. [Broker 9THM r106asb- HbIM TIpoeccoM comHednan aHepris (puc. 1). Bee oKHBbIe OpraHH3MBI HY KMAIOTCA TaKKe B HCTOWHNKe as0Ta, KOTOpEIii HeOGXOAHM sUIK CHHITeSa AMHHOKHCHOT, HYKEOTHLOB H ApyrHX coeAMHENMi. PacteHist B KaYeCTBE HCTONHHKa a30Ta HCIOB3yIOT TaaBHBiM O6pasom aMMHaK HH HHTpaTEt. To3B0- HOWHbIE KUBOTHBIE LOIKHBI TOy4aTb azoT B dpopMe aMHHOKHCNOT HAM ApyrHX OpranMyeckHx coeqnHe- uni. TouibKo HeKoTOpbie OpraHH3Mbi — UHaHoGakTe- PHH H MHOrHe BHAbI NOYBeHHBIX GaKTepHit, O6HTaIO- LHX B KaYeCTBe CHMGUOHTOB Ha KOpHAX HeKOTOpLIX pacrenmii, — cnioco61i mpespauiarb (pHxcHpoBarb) armoccepubiii asoT No B ammiak. JIpyrue GaxrepHnt (auTpriuumpyioume Gaxrepi) oKHcAAtOT aMMHaK 2O HATPHTOB H HHTPaTOB, HO eCrb H TakHe, KoTOpbIe npespamator nmrpatbi B cBoGoswBIii asor Np. Takum o6pason, B LonOnHEHKE K r106aTBHOMY Kpyronopo- Ty yrlepona H KUCHOposa B 6uoccepe mpoucxosuT KpyrOBopor a3ora, B KOTOPOM yyYacrByeT OrpoMHOe KONMUECTBO aTOTO atementa (pute. 2). KpyroBopors, KHCAOPOHa, YTseposta H az0Ta, B KOTOPHIE B KOHCH- Asorduxen- . Aemrrpudaumpyrounte pytouine Armoctbep- “Garcrepu, apxent Gaxtepiit unaii ago HTpHOb Awmonnit | Hirrparsa Tiaone- NOs | Garcrepan Hurrpudmuur yyroutite jaxrepit Hurrpapaumpyioune Gaictepait it apxen Pacrenma 2KupoTune Pacrenwa Puc. 2. Kpyropopor asora B Guoctpepe. Tasoobpasni asor N; cocrapnaer 80% 3emoi armoctbeput.HOM CYETE BOBJIEKAIOTCA BCe BHULbI, 3aBHCHT OT ecTe- CTBeHHOTO 6aslaHCa MEXK/y AKTHBHOCTBIO NpOsyeH- ‘ToB (aBrorpocos) H KOHCyMentToR (rereporporpos) B Hamieti 6noccepe. Kpyrosoporbt 9tHx auementos npHBosatca 8 ABIDKeHHe TpOMasHBIM MOTOKOM 9HEprin, HOCTYTa~ toutuM B GHoctbepy H3aBHe, a 3arem mpeobpasyembim B Guocthepe; Be HaumHaerca c nornomleHHs comed- Holi aHeprHn dorocHHTesHpy!oNLUMH OpranH3mamn € HOCEAYIONIMM HCMONb3OBaHHeEM gTOI aHeprun AAA cosnanua Gorarbix aHeprHeii yrieBonoB H Apy- THX OpraHWyeckHX coeqMHeHHil. ITH MHTaTeAbHBIE BEULECTBA BBICTYMAIOT B KAYECTBE HCTOUHHKa 9HEp- HH ja rereporpotpHEIx opranHsmoB. B meta6o- JINYECKHX MIponeccax H pH mO6KX npespamtenuax aieprint Yactb cao6osHOii aHeprim Tepaetcs nyTeM BBUIeIEHMA Tera B OKPy2Kaloulylo Cpenly M yBemM- NeHus atporim cucrembs. TakHM o6paso, B 610- chepe Marepua yuactsyer B HenpeKpamtaiomenca Kpyronopore nenlects, a 9Hepria yTHIHsHpyeTca — opranisuis He croco6unt perenupupoBars aHeprito, KoTOpad paccenBaeTcA B BULLE TEIIOTBI M SHTPOLHH. Kpyronopor yrnepona, kucropozia 1 as0Ta ocymtect- BIAETCA HeMpepbIBHO, B TO BpeMa KaK aHeprHa 0- croaHHo mpespamaerca 8 dopmy, B KoTOpoii ona yxke He Moxker GxITb Neniomss0BaHa, — B TenTOByIO aHeprHio. MeraGommam — copoxytinocth Bcex xHMive- CKHX NpeBpailleHHii, KOTOpbIe MPOMCXOLAT B KIeTKE WIM OpraHH3Me HM OcyULecTBsAIOTCA MOCcpeCTBOM cepu nocnenopaTenbHBIX KaTaTH3HpyembIx ¢ep- MeHTaMH peakiii, HaabiBAeMbIX MeraGonH¥ecKHMH, nytamn. Peaxunn (craaun) meraGommueckoro nyTH CAAYIOT /Ipyr 3a LPYTOM B OMIpesLesIEHHOM NOpALKE H Ha Kaki CTaHM B CHCTeMy MIPHBHOCHTCA He6Ob- moe cneumHyeckoe H3MeHeHHe ee xXHMHYECKOrO cocrasa, O6biuHO TipH 3TOM MpovcxoAMT yaaeHne, Tepemenenne ium j06annenme ofnoro aroma Wit (bynkunonatbxoii rpynnet. [pespamenne mpestie- CTBEHHHKa B KOHE4HBII IpoAyKT HAeT Yepes cepHO NPOMEKYTOWBIX NpomyKTOB MeTa6ONM3Ma, HadbI- BaeMbIx Meta6orHTamH. TepMui mpomMexyTOuHbHi MetaGoM3M YacTO MIpHMeHAIOT K CoBOKyNHOcTH (pepMentarHpHbix peakunii Bcex meTaGouMuecKHx yteii, 8 KOTOpBIX NPOHCXOAHT BsaNMHOe MpeBpalle- He npemuecrBentikon, Mera6omtToR 1 HU3KOMOMTe- KYMAPHBIX BeuLecTB (O6bI4HO cM,< 1000). KaraGoauam o6beannset npoueccsi aerpana- UM, Np KOTOPbIX OpraHHdeckHe MONEKYBI NHL (yraesonti, HEI H Genk) npespamtaiorca B HH3- 7] KOMOJIEKYAAPHBIC H Goulee NPOCTbIe KOHEYHBIE T1po- aykrbt (TaKHe kak MoOUHas KHCROTAa, CO, NH3). KataGonnsm conponoxaaetca Bbicsoboxtentien aueprin, KoTOpas sanacaerca B cbopme ATP 1 Boc- ctaopnennex nepenocunkos Boxopona (NADH, NADPH, FADH,). Ocratox aeprim paccenpaerca B BHyle Teruia. AnaGommaM, HasbiBaeMbiii TaKoKe 6140- CHHTe3OM, BKMOYaeT NPOLECCHI, MPH KOTOpEIX 13 Me/IKHX MPOCTbIX MpesecTBeHHHKOB cHHTTesHPy- ores Goutee KPYIINBIE Ht CHODKHBIE MOTEKY.tB1, B TOM ‘“MCJIe XKUpbI, NOsMCAXapHAbI, GeIKH HM HYKsIeHHOBBIe kucaorta, Peaxituit anabosmama nporeKaror ¢ norpe- Grenvem axeprHn, o6sruHO noayyaeMoli pH pas- ppipe cboctbarupix cpaseii ATP u poccraossennn NADH, NADPH, FADH, (pute. 3). Maxpomonexy.nst Conepaxamne ‘RAETKH aneprino mitra~ Beaku ‘eabHute BeulecTBa Hoancaxapases ‘Yrsenonts Janna Monexy:i- mpenmectennmtat Anmoxnexorot Caxapa CO, 2Kupupie xucnore: HO Asoruerue NH ochowattist Puc. 3. 3Hepreruveckoe conpamenne nytel KaTa6onnaMa WanaGonnama. KaraGonnveckwe MyTH MOcraBnavoT XMMUYE- cry sHeprwio 8 dopme ATP, NADH, NADPH » FADH,. 37m Tepenocunkn sHeprun «paGoraiot» B axaGonuyecknx ny- ‘TAX, Te MPOWCKORAT MpespaljexWA HM3KOMONEKYNAPHEX peujecre B MaKpoMOneKynbs.[8] Yacte II. BmoaHeprerinka H MeTaGonn3m Osa MeraGonmueckne nytH mpencrasnennt Anueiinoii yenoukoii mocnenoBaTeAbHBIX mpe- Bpamenuii. [pyrue MeraGoamueckne nyTH HMe1OT PasBeTBAEHHA, T. €. M3 OMHOFO NpeuEcTBEHHUKA COsMaeTCA MHOXKECTBO [OJeSHBIX KOHEYHBIX TIpO- AYKTOB, WAH M3 HECKONBKMX MCXOAHBIX BeMeCTB oGpaayercs oun nporykt. Boobue, KataGomue- cxie nYTH Xoneepeeumnes, Torna KaK anaGomHye- ckne — dusepzenmnni (prc. 4). Hexotoppie nyTH npecrapasior coboii peakUMOHHbie WAKbI, Kora. MouleKy.1a HCXOMHOTO BeULeCTBa pereHepupyeTcA B cepitu peakutnii, np aToM HeckonDKo MozeKy.1, 06- PasoBaHHBIX H3 MCXO;HOTO PeareHTa, BHOBD AaioT HCXOAHOe BeULeCTBO Kak NpoayKT. B cneayiounx Pochoanmnnta ‘Tpraruurzmepunaepekupunie kuesroTet Kpaxman = Anan Denna auansnt Tastxoremi> Crioxosaseqi> Enpysar Caxay Cepun Jetix Meoneiinntt Konneprenrunuii xaraGonmam —, a Ustrpaer Oxcanoanerat ruaBax paccMOTpeHbt HpHMeph! paaiM4HBIX MeTa- Goamyeckux nyteii, Y GonbuHcrBa KIeTOK ecrb dbepmentel AA nposeyeHusa kak peakunii pacnaya, Tak M peakunii cunresa saxHaix rpynn GviomoneKya, HanpHMep 2KUpHBIX KHCAOT. OsHaKo OAHOBpeMeHHOe MpoTe- kanHe CHHTesa M pacriazla KMPHBIX KHCAOT Gui 6bI pacTounTenbHO, 3TO MpexoTBpallaeTca B3a- uMuoii perysauneli anabonmueckux HM KaTa6onH- yeCKHX MOCAeLOBaTeNbHOCTeli peaKUii — KorAa ouHa peakUMOHHas LenouKa akTHBH3Mpyerca, Apy- raa nonapasetca. Takaa perynauna He Moria 6b ocymectBaATECA, cmH 661 anaGomMecKHe Ht KaTa- GommueckHe NYTH KaTaTHsHpoBamHch OAHMM H TEM Kaysyk t sonerrin emanie mmpotbocear > Kerecrepaareene an ~ ‘Menaonar Depuput xoueerepiita Kaporinionanme Crepowmuinie TnuEMenT ropMoubi Burana K Aueroauetit-CoA Siikosaxonm J 2Kupmute KucnoTss a TpHansrrmuepmrEL ‘X CDP-zvaunarannep Hi a> Dochommrs Apineprenrasrit anaGomam 6 ‘ tS CO, co, -_ Lbncneeckuit nyt Puc. 4. Tp THNa HeNMHeNHEIX MeTAGOANYECKHX nyTei. a — KoHBeprenTHutit Kara6onwyeckiit Tyre; 6 — puseprentiuri anaGonwieckuit myrb; 6 — IMKRMYeCKMit MYT, B KOTOPOM OAHO ¥3 MC- XONHEX Rejects (B RaHHOM cnywae OXcanoayleTaT) pereHepMpyerca, M UMKN BHOBb HaMMHaeTCA. Auerar — xmoyenoi mponexyrousiti mponykt MeraGon¥3Ma — BosMKaer MpH pacriane yenoro pana Goransx sHeprelt onexyn (a); OH CHYMHT B Ka¥ecrBe MIpenMIecrReMHuKa MOTH MPORyX- ‘708 (6) W Nornonjaerca B KaTaGONMNECKOM MYTH, WIBECTHOM KaK UMKN MMMOHHO! KUCHOTH (8).oe Ha6opom chepMenTop, AeiicTByIOUUIX B OZHOM HanpaBjleHHn [W181 anaGosIH3Ma HB pOTHBOTOIOX- Hom — s19 KaraGonnsma. MaruGuposanne cbep- MeHTa, ROBNeYeHHOTO B KaTaGONHSM, NpHBeO Oot K MHrHOMpoBaHMIO TOCTeAOBaTeABHOCTH peakuii aHa6onnueckoro HanpasieHna. Kataboanyeckne H ana6onuueckHe NYT, KOTOpbIe HMeIoT OnMHa- KoBbIe KOHLeBbIe TOUKH (HampHMep, TJOKOa —>—> UMpysat K nMpyBat > rH0Kosa) MoryT HcnOAb- 30BaTh MHOXKECTBO OAUHAKOBLIX cbepmeHtos. Ho BOT 4T0 O6s3aTeNbHO — No Kpaiiteii Mepe omHa 13 ctaanit kara6ommeckoro H anabonnuecKoro nyTeit karauanpyerca pasnuimn cbepMenramn HM HMeeT PasMYHBIE MexaHH3Mbl peryaaunH; aTH cepMeH- Th M ABAAOTCA MecTaMM OTAeNbHOIL peryaAUMH. Bonee toro, yro6u axaGoamuecknii u KataGonn- yeckuii nyTH Opin Heo6paTHMBIMH, yHHKambHBIE ALIA KaKAOFO HallpaBAeHHA MOcAeLOBaTeLbHOCTH Peakuntit omkHBI BKMOYaT xOTA GuI OAHY peaK- WHO, KOTOpan TepMOAMHAaMHYeCKH RecbMa 6.1aro- TpHaTHa, ApyrHMH cHOBAMH, OGpaTHas eit peaKuua TepMOAMHAaMMYeCKH HeBbiroqHa. HesapncumocTb perynannm Kata6onmyecknx m anabomyecknx Npoueccos ycHanBaeTcs H TeM, ¥TO MapHBIe KaTa~ Gonnuecknii 1 ana6onnuecknit nytH O6buHO Npo- HCXOJAT B pasHBIX yYacrKax KeTKH. HanpuMep, KaTa6043M 2%KMPHBIX KHCOT NPOMCXOAMT B MHTO- XOHAPHAX, a cHHTes — B uNTormasMe. Konnentpa- WMH MpoMexyTOUHDIX MeTaGomnTOB, depMeHToB H peryaTopos MoryT nosepxHBaTECA B pasmny- HBIX YWACTKAX KuIeTKH Ha pasHbIx yposHax. Buraro- japa TOMY, WTO MeTaGonMYeCKHe NYTH KHHeTHUe- Ck KOHTpoaMpylorca KOHUeHTpanHeii cy6ctpata, OTAENHLIE MIPOMEXYTOYHBIE NPOAYKTHI aHaGoH3- Ma Mt kaTaGOsIM3Ma TOKE KOHTPOAUpYIOT CKOPOCT meta6onmueckHx mpoueccos. MexaHu3MaM TakHXx anabosmuecknx H KaTaGommuecKHXx Mpoueccos MBI yaenum oco6oe simmanne. Mera6onmyeckne nyTH peryaupyiotca Ha HeCKOABKUX YPOBHAX, KAK BHYTPH KNETKH, TAK HL BiieKnerouno. HanGonee 6sicrpo MeraGonnueckue Mpouteccst pearupyior Ha nanuune cy6crpata. B obtem caysae BHYTPHKAeTOUNAA KOHLEHTpALITA cyGcrpara Menbiue Ky; pH 9ToM cKopoctb pe- akUMM ONnpeserseTcA KOHWeHTpauneii cy6crpata (cm. puc. 6-11). Bropoit cnoco6 Buytpukzerouro yupaBaAtb CKOpoctbio MeTaGorM¥ecKHx Mpouec- CoB cBsigzaH ¢ alocrepuueckoii perynauneli (r. 1, c. 220) npoMexyrounnim npoxyKTOM MeTa6ouH3- Ma HJIM KObepMeHTOM, HanpHMep aMHHOKHCAOTOI [9] um ATP, xotopue curnatusnpytor o cocroaunn Mera6oausMa BHYTpH KuIerkH. Korma KnerKa co- ACPKUT LOCTATOYHOE JA CBOMX HACyULHBIX T10- tpe6nocteit KomM4eCTBO, CKaxKeM, acnapTaTa WAM korja yposenb ATP B kleTKe Takoii, 4TO AanbHeli- wee norpe6eHHe sHeprun B AaHHbIit MoMeHT He HYKHO, 9TH CHTHaNB anOCTepHYeckH HHTHOUpY- oT aKTHBHOCTS OnNOTO HAH Gonee (bepmeHToR B coornercrsyiomeli nocnenosatenbuoctu peaxunii. Y MuoroxserTounbix opranu3Mos Metabomnueckaa AKTHBHOCTD pasaMYHBIX TkaHeii perympyerca H MHTerpupyeTca pocToBLimn (bakTopamMH H TOpMo- Hamu, KoTopwle selicrayioT cHapy2xu KneTKH. B HEKOTOPBIX CAYYAAX 3Ta peryANNA MpoMcXoANT chakrnyeckn MrHopento (iHorsa 6sicrpee, YeM 3a MIUUINCeKyHAy) Yepes W3MeHeHUA copeprKanust BHYTPHKNETOUHBIX MeCCeH/DKePOB, KOTOPBIe 13- MeHsIOT aKTHBHOCT cbepMeHTOB myTem asocre- pieckoli pery.ain WH Hx KOBameHTHO! Mom (buxannn, Hanpumep npx docopnanponannn. B APYPUX CHYYAAX BHEKIETOUNBI CHEHAN NpHBOAUT kK HSMeHEHMIO KOHUeHTPALMH (pepMeHTa B KieTKe, BUIMAA Ha CKOpOCTE ero cHiTTesa WT pacnana. Ta- Koli adtbext npostBanerca TOAbKO Yepes MUKYTEI MIM Yacet. Yacrb II MbI HauHHaeM C ONMCAHHs OCHOBHBIX aHepreTHYeckKHX 3aKoHOMepHocreii MetaGanHaMa (ra. 13). 3atem o6cyaum raaBnbie nyTH KaTaGomHs- Ma, 110 KOTOPBIM KIETKa NOY 4aeT IHEPIHIO, OKHCIAA pasnoo6pasiible Beutectsa (ra. 14-19). Toxpo610 aHepreTHyeckHe acnekTHl MeTaGos1NaMa paccmorpe- HI B Tui, 19. Ona 11ocBsimteHa XeMHOCMOTHMeCKOMY conpsKeHHI0 aHEpruH — YHUBepcalbHOMy Mexa- HHaMy, 10 KoTOpomy cHHTes ATP onpenensetca TpaHcMeMO6paHHBIM 31€KTPOXMMHYECKHM TIOTeHUM- @QJIOM, BOSHHKAMOUIHM J1HGO B Mpouecce OKHCHeHHA cy6crpara, 60 B mpoulecce normomeHHa comHed- voit aueprun. B ra, 20-22 paccmarpupatotca raprbie ana- Gonmueckue nyrH, rae sa cuer oueprun ATP us Gostee MpocTbIx Mo/eKY.1-TIpenMecTBEHHUKOB CHH- TesMpyIOTCA YFNEBOABI, AMMMAbI, AMMHOKHCHOTBI KH nykaeorin. B ra. 23 mpi nepeiiem k neTambHOMy oGcyxxneHiIo MeTaGorNeCKHX HyTeii y pasansHEIX opranH3Mos, Haumtas c Echerichia coli u saxanunean YeOREKOM, 4 TOPMOHAMBHBIX MeXAHHSMOB MX pery- AAU H MHTerpalin y MJleKOmMTAIOMHX. Vi nocae toro mei, HakoHeu, nepeiitem K usy- NeHul0 npoMexyTo¥Horo MeraGonuama. Mayyaa KuleTOWHEIT MeTa602M3M, HajlO TOMHUTb, YTO MHO-[10] Yacts II. BuosHeprernka # mera6onnam FOUMCAeHHbIe PeakUMH, ONMCAHHbIe B ITO! KHHTe, aelicTBHTeAbHO NpoHcXOAAT H UTpaioT peula‘outy1o ports p xuBEIx opranuamax. Berna crapaiitece 10- HsITb, KAKOBa POb TOFO WH HHOTO XHMMYECKOTO npoutecca (peakusm WAH Beero MeTaGomHHeCKOTO nyta) 6 opranusme. Kaxum o6pasom nsyyaembie PeakuMM cBAgaHBI C APyTAMH peaKMAMH, HeNpe- PHBHO NPOHCXOAAUIMH B TOI %e CaMOii KneTKE C lenbio NOAyYeHNA aHeprHn H BemecTB, HeObxomH- MEX UIA TOMIEpRaKHA KHSHECTIOCOOHOCTH KuteT- kn? Kak MHOroypoBHesble MexaHM3MbI peryAuHH MPHBOAAT B PABHOBECHE HOTOMeHHeE H BbYIeNEHHE Beullecrs H 9HeprHn, 7. e. OGecneyHBaIOT NOCTIDKE- HMe CTAal|MOHapHO-/MHaMMYeCKOrO COCTOAHHA BCe- To opranu3ma? Vsayyaa meta6o.nH3M c TakHXx TosH- UMii, BbI HCTIbITaeTe 3aXBaTbIBAIOUIee M BIOIHE T10- YUHTeNbHOE NOrpyxeHHe B OCHOBHI CaMoii 7USHH, H NonyuenHbte BaMH rayGoKHe aHaHHs, Geaycnon- HO, HaiiyT MHOXeCTBO NpHMeHeHHii B MeqMUHHe, CeNbCKOM XO3aiicTBe H GHOTeEXHONOTHAX.O6mar sHeprua BceneHHol octaerca NocToRHHON; o6mlaA HTpOTTAA HeTtpe- pHBHO yeennunBaerca. Pydonsq Knaysuyc, The Mechanical Theory of Heat with Its Applications to the Steam-Engine and to the Physical Properties of Bodies (Mexanuyeckas reopua Tena 8 npwnoKeHnn Mapowomy ABuratenw vu du3sMYecKMM CBOuCTBaM Ten), 1865 Bnaronapa nsoMoppusmy SHTpOTMN H MH(POPMALMH MOKAO ycTaHOBMT® BIa- MMOCBRSK MeXRY NBYMA (OPMAMK 3HErHU: 3HeprMek coBepulare nevicrene v sHeprves Hampasnars Nelicreue, KoTOpoe coBepuiaeTcA. (lornxa xan: pancya Kax06, La logique du vivant: une hitoire de I'heredite CTOPMA HacnencrBeHHOCTH), 1970 OcHoBb! 6v03HepreTHKH. Tunbl XAMNYeCKMX peakynn 13.1, Bao3Heprerka M TepMOAMHaMHKa 12 13.2. XAMMYECKHe OCHOBbI GHOXHMMYeCKHX peakywi 20 13.3. Mepenoc doctattbix rpynn w ATP 28 13.4. OKHCAMTeENbHO-BOCCTAHOBHTeMbHbIC Peakywn B Guonormyecknx cucTemax 43 Mpble KAETKH H OpraHH3MBl jOmKHBI OCy- IeCTBAATS MpOUECcHI TOsULp»kAHA %HS- HH, pocta H paamnioxkenna, Crioco6Hoct» uapseKaT iM mIpeo6pazonbiwarb aHepriio, KoTOpas pacxosyerca wa OHONOTHMECKHE mpOUeCcss, ABI erca ¢yraMenTabHbIM CBOICTBOM BCEX. 2KMBEIX opraivamop. 31a cpoiicrso 6x10 npnobpereo Ha ‘Axryant Jlapyasee, 1743-1794 panieii craaun KneTOuHO! aBomMOuNH. B coBpemeH- HBIX OPraHH3Max COBepUlaeTCA MHOXKECTBO pastO- o6pasnbix npeo6pasozannii aHeprun, mpespamenuit omHoii bopMbI aHepruu B Apyryi0. 7Kupbie opranns- MBI HCTIONBSYIOT XUMHYECKYIO SHEPrHHO KETOUHEIX MONEKY JUIA CHHTESA CHOXKHBIX: BBI- COKOYNOpsOYEHHLIX MaKPOMOJeKy.1 43 MpocTEIx Moulekys-nipemmectBeHHHKoB. OHH Take mpespa- UlalOT XMMHYECKYIO 3HEPrHiO KETOUHOTO B rpadenTbt KoHUeHTpaUMit H rpanMeHTeI aueKTpH- 4eCKHX NOTeHUMAIOB, B ABMKeHHe Hi Ter10. HemHo- THe OpraHH3Mbl, TakHe Kak CBETIAKH M HeKOTOpbIe ray6oKoBosibie MopcKHe pHiGb!, clocoubl HaAy4aTb cpet. DorocnHtesupyioutne OpraHnsmbl mpeo6pasy- JT CBeToBy10 SHepritio B pyrHe (oper aHepriH. XHMMYECKHe MeXaHH3MbI, KOTOPDIE /exKaT B ochose 6uomornieckix npeo6pasopannii aHeprin, B Tevenne croneThii savaponbinan 6uos0ron 1 6po- cain uM BEIs0B. DpanuyscKuit xummK AnTyan Jla- BYa3be 3AMETHII, 4TO KMBOTHBIE KAKHM-TO O6pasoM Npewpallaior XHMHYECKOE , T. €. BeIeCTBA 13 IMUM, B TETLIO, H CENA BEIBOA, YT ALIXAHHe — 2KUSHEHHO BaXkHBIi Mpouecc. --B OOUIEM, JbIXaHHe He 4TO HHOe, KaK MeqleH- Hoe ropeHHe yruepoia 1 BonOpona, KOTOpoe OYeHD TOXOxKe Ha Nporecc, Mpoucxonsunit B kepocuiosoii zamne HH cree. C atoii TouKH SPeHHs! KHBOTHBIe, KOTOPHIE JLbILAT, ABIISIOTCA[12] Yacte II. 13. OcHossi 6oaHepreTMKn. Tanti xHMAYECKHX peakynit HACTOALLHMI TONMBHBIMH KOTaMH, KoTOpHIE @xuraior H nO-kUpaior cau ce6s. MoxHo cka- 3aTb, “TO 31a AHALOrHA Mek TOpeHHeM H ABI XaliHem He H36exkala BHUMAHHs To9TOB — m1pa- BuuibHee, (put0compos ApeBHoctH. Dror oron, yxpanennbiii c ne6ec, sor dbaxen Tpometes He TONDKO MeN, POKAEHHAA B FOsOBAX HHDKEHEpOB Hm109T0B, OH ABAAeTCA BepHBIM OTIcaHHeM m1po- UeCCOB, IPOHCXOAAWHX B IpUpose, 10 Kpaiitieli Mepe, B CyYae XKHBOTHBIX, KOTOPBIE LbIAT. Tlostomy, ceays airTiunbim TBOpeNHAM, MOXk- Ho H Ham noitocodpcrBoBars, cKasaB, 470 ba- Ken ICUSHU 3@KUTACTCA B TOT MOMEHT, KAK TOS KO MJlajleHell HaNHHaeT /UbllaTb, H OH He racHeT 410 Tex Mop, oka He HacTynaeT CMepTb.” B XX B. YeoBeyecTBO AOCTHTIO HayabHO- ro TOHUMAHHA GombINMHCTRa XHMHYeCKHX npo- yeccos <«akesa 2%xH3HH». Ipespallenna aHeprun B 6MONOrHYeCKHX CHCTeMaX MOAYHHAIOTCA TeM 2Ke cbuanveckum sakonam, KOTOpbIe ynpaBasor M BCe- MM ApyTMMH MpuposHEIMH mponeccamn. TloaToMy CTyeHTaM-GHOXHMMKaM O4eHb BaKHO TOHATS 3TH 3aKOHDI, a TAKE TO, KaK 9TH 3aKOHbI NIpHMEHHMBI K noToKy aHeprHH B GHOcKepe. B oroii riane Mp1 cHayasta o6cyAHM 3aKOHD TePpMOAMHAMMKH HM KOJIMYECTBEHHIe COOTHOMIEHHA Mexny cBo6osHoit sHeprueli, aHTanbNHelt H aHTpO- mueii. Jlanee paccMoTpuM THIBL MponcxonAMINX B OKMBBIX KIETKAX OCHOBHBIX OHOXHMMYECKHX peak- unii, KOTOpble HeOGXOAHMBI JUIN HCHOMB3OBAHHS, Xpanenita, tlepenaun HM BbicBoGoxneHHA aHeprii, BOCMPHHHMaeMOl OpraHigMoM 13 OKpyxkatoueli cpenbi. Sate Mp1 rloapo6Hee ocraoBuMcn Ha Tex PeakuMAX, KOTOpbIe UrpaloT OcoGyi0 poh B 6uo- NOFHYECKHX mpoweccax sHeprerHueckoro o6Me- Ha, B YACTHOCTH Ha peaxumMax ¢ yyacruem ATP. V1 naxoneu, nepeiitem x paccMorpenimo BakHetx OKHC/IHTEABHO-BOCCTANOBHTEABHBIX peakUHii B>*KU- BLIX KNeTKaX, aHeprerHuecKHX 3aKOHOMepHocTeit nepenoca azexrporios B GuosorHuecKHX CHCTeMaX, @ TawKE TEX MEPEHOCHHKOB 9IEKTPOHOB, KOTOpBIE Hanon yacto paGoraior B aTHX NpoUeccax B Kaye- cree Kompakropos. * Vis memyapon Apwaria Cerena Antyatia Jlapyaste, 1a- Tupopantiex 1789 1; weTHponano no Kune [Lavoiser A. (1862) Oeucres de Lavoisier, Imprimerie Imperiale, Paris]. 13.1. BaoaHepretuka M TepMoAMHaMMKa Buosnepreruxa usyyaer Kommuecrennbie cropo- Hot Mpeo6pasonannit aHepram 113 omHOli ¢PopabI B APYTYI0, KOTOPBIe NPOHCXOADT BKUBBIX KeTKAX, a TAKOKE XHMHMECKHE MPOUECCHI, JNexKAUINE B OCHOBE aTHx npeo6pasopanutii. Xora 3aKonel TepMoguHaMnt- KH yoke O6cY:ANATHCh B MPeALITYMLNX rz1aBax, a BOS- MOXKHO, XOpOUO BLAH HByYEHBI BaMN pane, 3necb 6yaer BecbMa nonesHo B OSuMHX YepTax BCHOMMITE HekoTopble KomMecTRETTHBIe aceKTHI. Npeo6pazoBanna 3Heprun B 6vonormueckux cHcTeMax NOAYMHAIOTCA 3aKOHaM TepMOAMHAMHKK MHorouHcuIeHHble KOMYeCTBEHHbIe HCCeOBAaHMA NO B3aMMONpenpamlenmio pasnMyHBIX (opm aHep- TMH, BEINOAHEHHbIC (bu3HKaMH H XHMHKaMH, 1103B0- aun 8 XIX B. cbopmyaupowars sea OCHOBHBIX 3a- Kora Tepmoamnanuin, Fepsstii 3akon — 310 3aKoH COXpaHeHua aHeprii: npu s0G0M dusuueckom une XUMUNECKOM UaMeHeHUU obujee KoRUYECMeD aHep- zuu 60 ecenennoii ocmaemca nocmosnuem; mnepeun 2uoocem nepexodume us oduoil popMet 6 Opyzyio uru ‘Momem nepepacnpedenumcs, Ho ne Momcem ucued- nym. Bropoii 3aKoH TepMOAMHaMHKH 0 TOM, YTO BCE MpoueccbI BO BCeNeHHOli CrpeMaTCA kK yBETHEHIIO Gecniopanka (nesoprannsaunn): 6 pesyaemame .0- Obix ecmecmeenneix npoueccos sumponua ecenenHot eo3zpacmaem. Ka- rouleii cpetb cocrapnseT Bcenennyto. B aabopa- TOpHM HeKOTOpBIe XUMHYECKHE MIM (UsHYECKHE MpOueccbI MOrYT NPOTeKaTb B MSOAMPOBAHHBIX HH B SaKPHITEIX CHCTeMax, He CHOCOOHEIX K O6MeHY BellecTB H SHEprHH c OKpy2xkatouleli cpeaoii. On- HaKO 2KHBbIE KETKM H OPraHHSMbI ABIIOTCH OT- KPBITLIMH CHCTeMaMH, KOTOpbie OOMeHHBAIOTCS Ht BeillecTBaMH, H aHeprneli c OKpyxkaroulelt cpenoii. 2Kupple CHcTeMbI HUKOrAa He MPHXOMAT K PaBHO- BecHiO C OKpy2kaloleli cpenoii. TloctoaHHbie BsaH- MogelicrBua Mexkly cHCTeMOii H oKpy>kalomteli cpe- oii OSbacHAOT, KAKUM O6pas0M OpraHHaMEl MoryT NOJWLEPKHBATh BHYTPEHHIOI YOPALOYEHHOCTh 1 Tip atom jelicrBOBaTb B paMkax BrOporo saKoHa TepMOAMHaMHKH. Bra. 1 (1. 1, c. 22) mot aan onpenenenne tpem KOJIMMECTBEHHBIM TepMO/MMHaMH4eCKUM by HKLM, © NOMOULIO KOTOPHIX MOXXHO BbIPAsHTS aHepreTH- YeCKHe H3MeHeHHA, NPOHCXOAMUMe B XHMMYECKOH peaxunn: CsoGomnan sneprun Tr66ca G — 370 Ta vact SHeprimH, KOTOpaa cnocobria MpoHsBOAHT® pa6bo- TY B peakMH, MporeKaiomieli pH rlocrosHHO‘ ‘enmepatype MlocroaHHom japennn. B tex cayyasax, Kora peakiuis mpoteKaer ¢ BHICBOGOXK- denen cpoGoaHoit aneprin (7. e. usMeHeHH B ‘icreme nponicxonsT c yMeHbuteHHeM cuoGoaHO! aHeprHit), UsMeneHHe CBoGomHoii aHeprun AG — OTpHuarerbHaa BeNMMHA, M Takay peakytA Ha- SbIBaeTCA aK3eproHHueckoii. B aHAeproHHuecKHX 13.1 Bwo3Hepreruxa m Tepmonnnamnxa [13] peakyHax cHcreMa mpHoOperaer (noppiuaer) cBoGoaHyi0 aHeprinio HAG > 0. Outampnusa H — 310 Tak Ha3sbIBaemoe kaHHe> peakiHOHHOii cucremb. Ona or- paxkaer KONMYECTEO M LIpHpoxy XHMHYECKHX cpaseii B BellleCTBAX, BCTYNAIOWHX B peakUHio, HB BemectBax — nponykrax peaxunu. Xunm- yeCKad peaklMA, [pH KOTOPO!i POHCXOsMT BbI- neneHue Tera, HasbIBaeTca aK3orepMMUECKOii. Ecan atransnna o6pasonanns Hog, tIpolyktoB PpeakUHH MenbIlIe, YM (U1 BeIeCTB, BCTyTIaKO- MUX B peaKino, AH < 0. Peakinonnbte cucre- Mbl, KOTOPbi€ MOJ1YYal0T TeIIO OT OKpy2alouleii Cpembi, HasbIBaIoTCA SHAOTepMHYECKHMH; 71a nx AH — norokuTenbHaA BeNMUHHA. Surponua S— mepa xaormunoctn (HH Heyno- pasovennocrn) cncrempi (cm. on. 1-3). Korna MpomykTbl peakWHH MeHee croxkubIE 1 Gonee HeYNOPAMOYEHHBIE, HEM HCXO/HBIE BeULECTBA, cunTaerca, 4TO peakUMA NpoTeKaer Cc yBenHM4e- Hive SHTponHH. Equnnust nsmepenns: AG 1 AH — [bk/Mons (s0Ko- yib Ha Monb) 1 Kat/Mom» (KaTOpHH Ha Monb). Ha- noMHM, 1 Kan = 4,184 Jbx. Eannmust antpormn $= Tox /(mons- K) (koynb ta Moab Ha KenbouH) (ra6a. 13-1). B ycnopuax, xapakrepHbix juin 6HonorMyecKHX cucTeM (nocrosHHas Temmepatypa Ht lapsienMe), Ha LOTTE Koncratira Bosrbunatia, k = 1,381 - 10° Jbx/K Yucso Avoranpo, N= 6,022 - 10 moa * Koncranra Dapanes,J = 96 480 Jb /(B - wonp) Tasosas nocrosimas, R= 8,315 Jo«/(moan- K) (© 1,987 Ka/(mom - K)) Pasmepriocrs AG 4 AH — Jbk/Moap (vam Kan/ Mom) Pasepwoct AS Jl /(wo.nb K) (isi xat/(maxin « K)) f xaa = 4184/06 PasmepHoct a6comoruoit renepatypst T— K (keri) 25°C = 298K Tipu 25 °C RT = 2.479 lbox/ moat 592 xxast/ moms)[14] Hace IL, 13. Ochostr 6HoaHepretHkn. Tunbi xuMMdecKHX peak MeHeHHs cBo6oAHOit SHeprHH, 9HTATBNMK M SHTpO- MMM CBABAHBI KOMYECTBEHHO JIpyt € APYTOM caleay- 1OWLMM ypaBHennem: AG = AH—TAS (13-1) rye AG — naMenenne cBoGosHoii aHeprin peakiHoH- oii cucremti, AH — maMeHeHHe aHTaIbNHM peaKin- onnoit cucremm, T— Temnepatypa (B Kenbennax, K) 4 AS — uameHene sHTponnn cuctembl. YouOBUMCs, uro AS > 0, Kora SHTponHiA yBeAMUUBaeTCs, H, KaK ormeyaiocs Baie, AH < 0, Korma peakuHoHHas CH- crema oTmaet rerio oKpyzkaiouteli cpexe. JIA camo- MPpoHsBosbHO (crOHTaHHO) MpoTeKaroulNX peaxuuii H MpHpoaHBx mponeccos Bcerna AG <0. To Bropomy sakony TepMoanHaMHkn mpit xH- MHMECKHX peakUMsx HIM CpHSHYeCKHX nponeccax aHTponust ecerennoit yeamunpaerca. 3 Toro 3a- KOHa, OHaKO, He CulejlyeT, YTO BOspacraHMe 3HTpO- KH AOXKHO MpOHCXomHT OGasaTebHO B caNoii peaxyuonnow cucmeme. Buyrpukrerounaa ynopa- MOYEHHOCT, KOTOPad COsaeTCA B Mpouecce pocta H JeneHHA K1eTOK, Gonee YeM KOMIIeHCHpyeTCA BO3- HukKaloulel Np ITOM HeylOpsOYeHHOCTIO B OKpy- skarouleii cpeste. Koporko rogops, xkHBbIe OpraHH3Mbt COXpaHsIOT BHYTPEHHIOW yOpALONeHHOCTS, Nosy- wad 13 OKPY2KalOUleli Cpesb! CBOGOLHYIO 9HEPrHio B BHMe NMTATEsHBIX BeLLECTB HIM COMHEYHOTO CBeTA 1 BOSBpalllad B Hee TAKOE %Ke KOAMMECTRO 9HEPrHHt B sue Terula 4 9HTponHM. Knetkam Heo6xoqMmbt MCTOYHMKH CBO6OAHOK 3HEprHn Kuerku — nsorepmmueckwe cucrempl, oun ¢bynk- IMOHMPy!oT NpH MlocToaHHOit Temneparype (a TaKwKe pH nlocrosiHom japstenun). Teno (Harpepanne) He MOXET CIY2KHTb HCTOUHHKOM SHEPHH JIA KIETOK, NOcKONbKY TerLI0 cnocoGHO MpoHsBoANTS paGoTy JMU B TOM cuy¥ae, Kora OHO MepexomnT oT Gonee varpetoro Tea k Gotee XonOJNOMY WM H13 O6s1aCTH ¢ Gonee BbICoKolt Temnepatypoii B oGzacTb c Gonee Huskoii Temnepatypoii. KaerkH MOryT 1 NOsRHEI HIC- Nomb30BaTb CBOGoMHYIO aHeprHio [H66ca G, Koropaa CBA3AHa C MPOYHOCTBIO XHMMYECKHX cBABei MEKIY aTOMaMH, OnpeylenaeT COCTOAHHE PaBHOBECHA Hl Ha- Iipawienne XHMMUYECKHX peakuutit; 9r0 Ta TeoperH- MeCKM BOIMOKHAA XHMHYECKAA SHEPrHa CHCTEMBI, KoTopad MoxKeT 6bITb NpespauieHa B pabory, 7. €. BbI- JesleHa BO BCeneHHyI0 B BAe TeILIa H 9ATPOTMH MPH NpOTeKaHHtHt XHMHYECKO!i PeaKLMH MPH NOCTOSHHOI Temmepatype 4 MOcTOAHHOM jtaBiennn. Tereporpod- HBIE KIETKH H3BIEKAIOT HEOOXOAHMyI0 cBOGOAHYIO SHeprinio H3 MouIeKy.1 IHHTaTeMIbHDIX BEULECTB, a poTO- CHHTeSHpyIONIMe KIETKH NOrsOULaIoT aHEPTHIO CoN- Heunoro cBera. te, 1 pyre nepewostr cBo60HyIO aneprino B ATP 1 pyre coesut- HeHHia, KOTOpEIe CnocobHE! sanacarb aHepriwo 218 BEINOAHEHHA paGorb! (6HOOTHYECKHX MpoLeccoB) B YCNOBHAX NOCTOSIHHOII TeMreparypsl. V3menexne craHpapTHon cBobo_HOn 3Heprh HenOcpeACTBeHHO CBAIAHO (C KOHCTaHTOM paBHoBecna Cocras peakunonnolt cucrembl (cMecb Beutecte, BCTYNAIOMINX B PeaKUMH, H NPOAYKTOB peaKuntHt) Me- HAETCA {lO TEX MOP, NOKa He ycTaHaB/MBaeTCA paBHo- zecue. [lo ,ocrinkenM pabhoBecHst KOHUeHTpaLUH PearMpyioulMX BeLECTB HM MpOLYKTOB peak He MEHSOTCA, CKOPOCTH TIpAMOil H OGparHoii peaxunit CTaHOBATCA PaBHbIMN Ht ATbHEMMIME HaMEHEHHA B cucTeme mpekpatiaiorca. Konuentpaunn pearnpylo- LUX BeLIECTB HM NPOAYKTOB peakuMN @ cocmoanuu PAsHOBECUA OTIPELEIOT KOHCTAHTY PaBHOBECHA Keg (v1, c. 24). 3anmuem ana o606meHHO! peaKunHt aA + bB = cC+dD rae a, b, cu d — 4ncno monexya A, B, C uD, Kon- craHTy papHosecna _{crDy Keg IAVBE (13-2) rae [A]. [B], [C] 1 [D] — Monapaite konuerrpannn BeULECTB B COCTORHHM paBHonecHa. Hepapnonectiaa’ peaxunonnas cictema scerja caMONPOMSBONBHO CIpeMHTCA K yCTaHOBEHHIO paB- HOBECHOTO COCTOAHHA NIpH 3aaHHBIX YCAOBHAX (KOH- YeHTpauis, Temnepatypa, AaBneHHe); TIpH aTOM Mpo- HcxOMWT HBMeHEHHe cROboAHO! aHeprHm peaKL AG. IIpw cranmapTHbix yCnOBHAX, T. e. MpH Temnepa- type T = 298 K = 25 °C, nexoanttx KoHueHTpaunax BCeX KOMIOHEHTOB peakuuit 1 Movtb/s1 (HIM M), Wt Juis ragoo6pasHbix BeLICCTS NaplMaIbHOM AaBIeHUIL 101,3 xMla (xw1omackanb) = 1 arM, ABWKYUIAK CHA CHCTeMbl K COCTOAHMIO PaBHOBEcHA Onpenerserca H3MeHeHHeM CranfaprHoli crobosnoli axepran AG’. Cranaapribimn ycrosnsimn 21% peaxunit ¢ yuacra- em HoHOB BOOpona aBanerca [H*] = 1 M, 1H pH 0.Boabumicreo 6uoxumuvecknx peaxunii npoucxoaut pasGapnennux Gycepubix pacrsopax npu pH = 7. Tpit atom nt pH, 1 KoruerTpaninio Bonb1 (sU18 suctoit poabi [H,O] = 55,5 M) Moxktto cunrarp Tloctoarbr- ou. Jia yao6crba pacieTos B Ka4ecTBe CTaHjlapTHEIX yeuosuit ua GMoxMMuecKHX Mpoueccos nputis- THI CHeAYIOUE: KOULLEHTPALLHA BONOPOAHDIX HOHOB 107 M (pH 7), konnentpanns sone 55,5 Mu xou- uesrpauna Mg?* 1 mM (ui peaxuuii, npoexaroumx ¢ yyactnem Honos Mg”*, skmto4as GosbUIHHCTBO pe- axunii, rae Bsanmoneiicrayer ATP). KAHOYEBBIE AOFOBOPEHHOCTH. JLna yno6cra B Ka4e- cree cranjaprnoro coctosnna (cranaprHEx ycno- Bui) OHOXUMHYECKHX cHCTeM BHIGpaHBI yenoBHA, OTNYHBIE OT TX, YTO NPHHATE B XHMHH H CbHBHKe. A MMeHHO, B 6HOXHIMMM CTaHapTHOe COCTOAHHe CO- omererayet [H*] = 10-7 M (pH 7) #[H,0] = 55,5 M. Zina peakunit ¢ yyactnem nonos Mg* (k KoTopbiM OTHOCATCH MHOFHe peakuH c yuactHem ATP) KoH- ueHtpauna Mg?’ B pactBope o6ni4Ho cuntaeTcA T0- crostioii u pasnioit 1 WM. usnueckve napamerpbl, OTHOCAUIMECA K 6HOXH- MHMECKOMY CTaHJlapTHOMY COCTOAHIIO, Ha3blBaloTcA craijapribiMn NpHBejeHHbIMH NapaMeTpaMH 11 T1H- lyTca co UITpHxOM (HanpuMep, AG” Ht K,) L181 TOTO, 706 OTMYATh HX OT HenpHBEAeHHEX Mapamerpos, HIcTIO.TSYeMbIX XHMHKaMH H (pHSHKaMH. BaMerHM, To B Apyrnx yueGumkax 6HOXHMUH Yall HcTomsyeTcA oGostauerne AG” (a He AG”, Kak MbI BBE BBE). Tlapaerp AG” pexomersozan’ MexyHaposlHbim cO- 1030M XHMHKOB H 6HOXIMMKOB ¢ He:TbIO MORYEPKHYTD, “To HerIpHBezeHHbHi napamerp G' — ato KpHTeprtii pasnosecus. JLut npocrorsi guech 1 jase Gynem Ha- SbIBATb STH MIPHBCACHHBIC HlapaMeTpbl CTaHapTHbIMH M3MeHeHHAMH CBOGOTHOH sHEprHH. KAIOYEBBIE OTOBOPEHHOCTH. Comiacuo apyroii 20- robopernoctu, ecm HO, H* u/iin Mg?* sbicryna- JOT B KaYECTBE HCXOAHBIX BELECTB HIM MpoAyKTOB eaKUMH, HX KOHLeHTPAUItH He BKLIOUAIOTCH B ypaB- Hens, Takne Kak (13-2), a BMecTO aTorO BXOBAT B koncrantet Kg AG". i Vitak, 1A peakiun xapaktepa KoHcTaHTa pasroseciia K',, orcioaa AG’ — 370 xapakrepuctii- eckutit napamerp peakiuin, Kax sor ormeyau B 12. 6, Ki, AG” cassatint npocrbit cooTHomieniteM: AG” = -RT In Ky, (13-3) 13.1 BuoaHepreruka u tepmoquuammKa [15] mo npocmo odun us 6oamomne1x cnoco6os pacue- ma Koncmanme pasnosecua peaxyuu vepes uaMe- nenue cmandapmnoti ceoGodnoit anepeuu xunuue- cxoit peaxyuu. B ta6.a. 13-2 npwpenersr snayenna AG" it Ky. Ecattt Keg = 1,0, to AG" = 0,0 (raK KaKk In 1,0 = 0). Ecm Ki, > 1,0, 10 AG” < 0. Ecan xe Kg < 1,0, AG” > 0. Tockonbxy sasncumocts AG” ot Kiy sorapnepmirieckas, oTHOCHTeNbHO HeGob- le H3MeHeHus AG” npuBogsT K CHIbHBIM H3Me- nenua K, Tlonesnio take onpenenirs uaMenenne cBo- Gonnoit aHeprim m Apyrum nytem. AG” — sto pa3- HOCT Mexsly CBOSOAHO! aHeprHeli MpomyKToB pe- akUHH H cBoGOAHOI aHeprueii HcxoAHDIX BeULeCTB. Korna AG" < 0, cpo6oguas steprus o6pasonanna Gey (poaykto peaxtuttt) < Gasp (pearentos); Ta- Kasi Peaks IPH CTAHsaPTHBIX YCOBHAX TIPOHCXO- JT CaMONpoHSBoNbHO (CHOHTAHHO), HOCKOTBKY BCE XUMHYECKHE peakIHH CTPeMsITCA MATH B Hampapite- HHH, COOTBETCTBYIOUIEM YMeHBILeHHIO cBOGoAHOIi atieprun cuctembi, Ec AG™ > 0, 10 Gey (mpoayk- ‘ToB peak) > Gog, (pearetttos). [px Kontertpa- LusX HCXOAHBIX KoMONeHTOB 1,0 M (cranaaprHDre YeuoBia) Takast peakuuns Wet B OOparHom Harpas- aenun. B ta6.a. 13-3 o606menst 911 Moto%KeHHs. 10% ATA 41 10? 114 ~27 10! -57 14 1 00 00 10°' 57 14 10? 114 27 108 17,1 44 104 228 55. 105 285 68 10-6 34,2 82 * BCH pasmepnocrs aneprun — mxoysts (41 KiuIOmKOY:16); coxpamerno JL (Mm K/bx). OnwaKo OHoxmNnKH ame Bce- To TIpHBOAAT AG" B KKa/Mom (KHIOKaTOpHA Ha MomB). 1 [bx = 4,184 kKa.[16] Yacte IT. 13. OcHoes: 6uoaKepreHKn. Tanbi xaMMYeCKHx peaKynit eee Tipu navanbuoit xonerrpatun Keg AG” acex wemects { M >10 <0 Tipamas peaxnuist 1.0 0 Papitonecitoe cocrosime <1,0 >0 O6paruas peaxuns lm Mpumep 13-1 PACHET AG’? Haiiurre 1amenenne cpoGonnoit aHeprum peaxa, kaTasanpyeMoii bepmerttom (pociporoKomyrasoit: TatoKxo30-1-cpoccpat = ritoKo30-6-cpoccpat B nayare peau KoHUeHTpaunst rm10Ko30-1- cpoccpata cocrasrsita 20 MM, a rmoKo30-6-pocdar B peakuMOHHoii cpene oTcyTCTBOBAN; B paBloBec- noii cmecn mpu 25 °C u pH 7,0 coneprkurca 1,0 MM rm0Ko30-1-pocdara H 19 MM ra10K030-6-;poctpata. Tlorepeit nam yoemuueriem cno6omtoli areprin co- Mpopoxuaetca peakunsl, Wlyulat B Hanpapnenin o6pasoparia raoKo30-6-cboctpara? Pemenne. Creayet paccuntatb KoHcTaHTy paBHo- BecHs: [rmoKo30-6-«pocdar _leanca Sorta 19 mM/1,0 MM = 19 [rm0K030-1 0. act Peaxwii xJbx/Mom KKa/MOb Tuaposms. Kucaornetit miapoms Yecycussii anrnpu + HO — 2 anerara 91,1 -21,8 ATP + H,0— ADP + P, 305 -73 ‘ATP + H,0 — AMP + PP, 45,6 - 109 PP, + H,0 > 2P, 19,2 46 UDP-roxosa + HO > UMP + raoxoso-L-pocipar 43,0 103 Tuaponsts acbipon Stiuranerar + HO — sranon + anerar ~19,6 47 LitoKo30-6-thoctpar + H,O — rnioKosa + P, 13.8 -33 Hapoans asoon nw nenriyton Cayranam + HO — rayrawar + NHj 14.2 3A Cumuermaun + HO 2 rmnuna 9.2 22 Tuapoms raxosti0n Maasrosa + H,O > 2 rmoxoans e153 -37 Jaktosa + H,O > rmoxosa + raakrosa 159 38 Buytpumoxekyaspuie neperpynnuposKet Emoxox0--«poctbar > rmoKoa0-6-¢hocepar 13 Aa Dpyxros0-6-pocipar —> rmoKoso-6- ¢bywrapat + HO 34 08 Oxnexenne Mosiexyasptibim KHes10p010M Tmoxosa + 60, > 6CO, + 6H,O 2,840 686 Tanbantrar + 230, — 16CO, + 16H,0 -9,770 -2,338B ra. 13-4 npusexcnst yncaenupie suavenus HigMeHeHHii crangaptioi cBoGoaHoii aHeprum He- KOTOPEIX BAXHDIX XHMHYECKHX peaktutii. OrweriM, TO THAPOAMS MpocTLIx ahupoB, aMHAOB NenTHA0B 1 PAMKO3HOB, Kak H_BHYTpHMoNeKYaApHbIe Mepe- TPYMIMpOBKH HasUIMMLMpOBAHe, COMpOBOANAOTCA HeOONbIIMM H3MeHEHHeM CTaHsaprHoli cBo6ogHoit SHEPriH, B TO BPeMa KaK MPH rHApOIHge atIrHApH- JOB KUCIOT NPOHCXOsUT SHAuHTebHOE yMeHDUIeHHE ctaHaptHoii cBo6onHoii aneprun. Tomioe oxucne- He OpraHHueckHX BeuLeCTB, TAKUX Kak rOKOIa HTH NawbMHTHHOBAM KHCIOTa, 0 CO, 4 HO, nporeKato- lee B KAETKAX MHOTOCTYTIEHIATO, IPHBONT K O4CHD GoubWOMy yMeHBUIeHHIO craHaapTHOii cBobonHOit aueprun. Jannwle, npiwenenupie B tabs. 13-4, oT- HocaTca k cmandapmnen ycaoewam. Mapamerpsi pe- aKuHii, pOHcXOASLUNX B KNeTKe, MOXHO TOyUHTD, MpHBeAA BEAHAHHDI B CTAHLApTHDIX YCNOBHAX K pe- auIbHDIM youoBHAM (B K7eTKe). V3meHeHna cBo6oqHon sHeprun B peasibHbIxX CHCTeMaX 3aBMCAT OT KOHWeHTpaynii CXOpHbIx BeLYeCTB M NPoAYKTOB peakyHn Mot nosxtibt yeTKo yacHHTD, YO HaMeHEHHe CBO Goanoii aHeprin AG 1 HaMeHeHue ctarizapTHoii cBo- Goanoit axeprim AG” sncrenno pasmuaiotcs. Kark- Jad XUMMMECKaA peakIMIA XapakTepH3yeTcA ompe- JEICHHDIM H3MeHeHeM cratsaprHoit cBobonHoll SHEPriH, KOTOPOe MOXxET GEITH MOIOKUTERBHBIM, OTPHUATENDHbIM HIM PaBHBIM HYJHO B 3aBHCHMOCTHL OT KoHCraHT papHoBecua sanHoll peaxuun. Ho H3- MeHeHMio craHapTHoit cBoGoAHOii SHEprHKt MOKHO CYAMT 0 TOM, B KaKOM HanpaBnenin nporeKaer taH- Hast peak H Kak aleKa Oa OT paBHOREcHA mpu ucxodubix KonYenmpayusx ecex KoMnonenmos 1,0 M, pH 7,0, remneparype 25 °C u paprennn 101,3 KIa. Taku o6pasom, AG” — xapakrepucruyecksii napa- MeTp (KOHCTaHTa) jaHHOii peakuHM. OsHaKo B pe- AJbHBIX YOIOBHAX H3MeHEHHe CBOGOAHO! sHEprHN AG aapncut of KonuerTpautii HcxonMbIx BeMeCTB MH NposyKTOB peakitn, a TakKe OT H3MeHEHHA TeMIe- paypbi mo xoy peakunn. Pearbibte YCnOBHA MOrYT He CoBilajaTb co craHslaptHbimu. Bonee toro, AG s106oii peaxuiin, cnowTaHHo crpemamtedicn K paBHo- BECHIO, BCera OTPHUaTesIbHas BEMYHHA, HO Mepe mporekanna peakunn AG yseamunpaercs, crpemach Bnperese k Hym0. B MoMeHT yeratoB7enna paBHio- peciit AG = 0 — B zannoli peaxurnt 6osbine He MO- xkeT 6bITb coBeplteHO HUKaKOii paGorEl. 13.1 Bwoavepretuka u tepmonuHaMMKa \17] 71a mo6oiit peakunn aA + bB = cC + dD AG WAG” cpasaniel ypasnennem AG = AG" + RT| key De. 13-4 "Tarp &CD BKOTOPOM NlapaMeTPbI, BbI/eJIEHHBIE KpacHUM, mpe- obradaouue & 1aHHO! cHcteme. KonueHTpatnn 8 ypabnenun 13-4 npunsiro HagiiBarb elicrByio- IUIMH Maccamu, a orHomenne [C]*{D]*/[AP[B]® BbIpaxkaeT 3aKOH AelicTByiomux Macc Q. B kaye- cree mpumepa zanaitte npeanozoxum, ¥To peaK- uns A+B = C+D nporexaer npn crannapr- HbIx yculoBHAX Temnepatypbt (25 °C) H paBsEHHs (101,3 kIla), Ho 40 KonueHrpausn [A], [B], [C] [D] ne pasnet Mexay co6oii u HH onNa 13 Hux He papna crariaaprHoii KonueHTpaunn 1,0 M. Urobe1 naiitu usMeHeHue cBoboAHoOii aHeprun AG peak- LMM, MpoTeKaiomelt cena Hanpabo, MpH ITHX He- cTanmapTHDlx YcTOBMAX, MOCTABHM YHCMeHHBIE 3uavenHa KoUNeHTpanit KoMMOHeHTOB A, B, Cu D Bypabnenne 13-3. Mapamerpnt R, THAG" oTno- CATCA K CTaHsapTHbIM ycroBHaM. AG <0 m0 Mepe HpoTekaHHa peakUMH ITOT Mapamerp CTpeMHTCA K HYJO M3-3a Toro, ¥TO BeulectBa A u B pacxony- orca ([A] 1 [B] yenburaiorea), a nemecrsa Cu D napa6atpatorea ([C] u [D] ysennunpawrea). OreriM, 970 B cocroaumn papiopecna AG = 0 1 ypapnenne 13-4 ynpomaerca: Clea De o=aG=ac" + Rrin Cel TAla(Bla AG = — RT In Ke, 1. e. MBL CHOBA Momy4uiutt ypabHenne 13-3, caasEiBa- jomlee H3MeHeHHe cranapTHoii cBoGopHOM aHeprHn HW KOHCTaNTY paBHoBecns. Kpurepuii camonpoussosbHocrn peakunH cBs- san c AG, ane ¢ AG”. pu AG” > 0 peaxunsa naer TONbKO NpH ycLOBHH, YO AG <0. Takoe BosMOxKHO, Kora B ypasnennn 13-4 0-< RT In ({npoayxrss pe- axw|/[ucxoanbe BewectBal) > AG’. Hanpunep, Tp HeMeAeHHOM ynaneHHn MpoxyKTOB peakunn orHomenne [1pomyKTbl peaKun)/[Hcxo_Hble Be- uectBa] K 1, tak uto RT In [npoayKrei peaxunn]/ [ucxonule BemectBa] <0 (Gonbuloe orpnuatenb- Hoe uneno). Bentununt AG u AG” supaxator MaKcUManoHOe KONMYECTBO cBOGOAHO! aHeprnn, KOTOPoe meopemuvecku natiHan peakunA MOxKeT MpegoctaBHts, T. e. KoMHYECTBO aHEpruH, KoTOpoe[18] Yacte IT. 13. OcHost! 6vosHepreThKn. THnbi xHMMYeCKHX peakunit Moro Gurr pearis0Banto, ecu OBI HMECH BbICOKO- adtekTHBHBIi MexaHu3M ee Mcnonb30BaHHA. Tockonsky Taxoro Mexanvama Het (Bo Bpemst mo6oro mpouecca, Kakas-TO YacTb cBoboAHOl SHeprit BCerfa TepseTca B Be BHTpONH), KO- AMUECTBO paGoTHI, BLITIOAHEHHOE 3a CUeT AHHOit peakiMH pH MlocToaHHoii TeMmepatype H MocTO- ANHOM aBIeHHM, BCerAa MeHbUIe TeopeTHIEcKH okUaeMOrO. Apyroii saxnetii MoMenT 3akmoyaetcs B TOM, To HeKoTopple 6xaronpHATHBIE B TepMomHHaMmtue- CKOM OTHOLIEHHH peaklMH (a HMEHHO, peaKUHH, JVI Koropsix AG” < 0) npovcxoasT ¢ HenocrarouHo Gombmoii ckopoctnio. Hanpumep, ropene (peakua oKHicneHHa) ApoB 0 CO, w H,O tepMoannamnue- ckH O4eHD BBITOmHO. OsHako ApoBa xpansttes rona- MH — TOpeHHe HaunHaeTcH He CaMONpOUsBOIbHO, hlockosbKy aeprua akrupanun (cm. prc. 6-2 1 6-3) PeaKUWH TOpeHHA BbULIe, YEM AOCTYNHaA MPH KOM- Harxoit Temmepatype aueprust. Een npeaocrasurs neo6xoanMyto ateprino akTHBanun (tanpHMep, OF 3a0KCHHOH CHMYKH), HadHeTcA Mpouecc ropeHus, Mpespautarounit zepeso B Gonee crabuabHble mpo- ayxrot CO, u HzO u BpicsoboxKnaroutnit aHeprii0 B Hine Tenia cpeTa. Terstora, BoIneneHHaa MpH Toit ak3orepaiveckoit peaKnun, oecriesuBaeT aHepriio AKTHBALMM JIA TOpeHHA CELYIOWNX NOpunii ApoB, M TaKHM O6pa30M Mpouecc ropeHiaA MoxeT Mpoqos- 2KaTbCA HETIpepbIBHO. BOKMBBIX KIETKAX XUMHMECKHE peakIMH MpO- ‘Tekan 6bt upesBbtuaiiHo MeUIEHHO, ecaH Obt TAM He 6o.10 achpexmusnoix kamanusamopos — (pepMeHtos, KOTOpHIe BOBCE He OGecneyHBaIOT peakIMOHHy1O cucTemy AonoHHTeAHBIM TenIOM (aHeprueit), a TOWKaIOT sHepeuio axmueayuu. Depment «npen- Jlarae> abrepHaTusHniii nyt peaKinn c Gonee HNSKoit aHeprieli akTHBaLUH, YeM B CTyYae HeKa- TaIMsHpyeMoit peakuMH, {UIA Toro YTOGbI aHEprHst Gonbumuctea Mosley. cy6cTpata Np KOMHaTHOIt Temrlepatype OKasaacb AOCTaTOUKOi AIA Npeono- aeHnst aHepretuueckoro Gaphepa — aHepruM aKTH- Bann. Baarogapa depMetarnpiomy KaTausy CKOpOCTS peakUHM YBEAMYMBAeTCA BO MHOTO pas. Hamenenue ceoboduoit anepeuu peaxyun ne sasu- cum om nymu, no Komopomy peaxyus npomexaem, €3aBHCHT TOMIbKO OT MPHPOABI H KOHWEHTPALNH HC~ XO/HDIX BEMLECTE H KOHEYHBIX MPOAYKTOB peakUHH. TIpu dbepmenmamusnom Kamasuse He usMensemca eenununa KOHCmaHMe pagHosecuR Peaxkuu; HO NO; aelicrguem (bepMertoB MoxkeT yBemMuMBaTbCA IL aeiicrBHTeIbHO YBeMAMBAeTCA CKOPOCTH IIpAMOi u o6paruoii peakunii n mpouece Haer B Tepmonn- HaMmueckH pagpeulentiom Hanpapenun (31aKOM ugMenens cBoGoaHoli sHeprunt). WaMeHeHua CraHqapTHoii cBo6oAHOH 3HeprMn aAAMTHBHDI Paccmorpu nocaenosarenbubie peakunn A = B «BC. Kaxkaaa peakuna umeet co6cTBennyio KOHCTaHTy papHoBecns HM xapakrepHayerca onpe- Ae@NCHHBIM H3MeHeHHeM CTaHsapTHoii cBoGosHoOii aneprun AG? 1 AGS. Tockonbxy aH aBe peakumn Mporekaior MoctenopatestbHo, KoMTOHeHT B Mok Ho HeKMOUNTD 13 oBcyaxTeNMA. Torna paccMoTpHM cymMapxyto peaxnino A = C, xoropaa xapakrepu- BYeTCH KOHCTaHTO! paBHOBecHA H, culeNOBaTeNb- Ho, H3MeHeHHeM craHpaptHoit cpoboaHolt aHeprim AG Sue Aut nocnedosamenvubix xuMUYecKUx peaxyuit AG” addumuenw. Jia cyMMapHoii peakuun A = C o6mee HaMeneHHe cralaptHoii cBoGoAHOi anep- THH AG¢u PabHO CyMMe M3MeHeHHii CTaHapTHX cao6oarbix aHeprnit AGS n AG? apyx peakunii: ‘in Gy + AGS a) A+B aGr Q) B>C AGF Hmow: A>C AGr+AGe Dror npmnwuin GHosneprernKn oOacuAeT, KaK peaK- unis (aHaepronmieckas peakuHs), He BBIOMHas Tep- MomitHamnveckit, Moxker 6bITb Mponenenia B MpsMOM Hanpapaenun Mpit ConpwKeHHH ¢ BEICOKOaKeprO- Hmueckoii peakuneli sepes oOumnii npomexyTourtit npoaykt. Hanpumep, y Mitorux opranuaMos cuiTes rai0K030-6-cpoctpata sapasetca neproii cranneli rpm YCBOeHIH raOKOSBI: Triokosa + P, + rmioKo30-6-«bociar + HO AG" = 13,8 KJbx/onp Ana stoi peakuwin AG” > 0, 1 npn cranmaprHerx YCNOBHAX PeakI[HA CaMOMPOH3BONBHO He NpoHc- XO/MT B YKa3aHHOM HampaBzennn. J[pyroit mporecc B kjlerke — rnapoaus ATP go ADP uP; — Hao6opor BBICOKOaK3eproHH4eckuii: ATP +H,0 — ADP + P, AG” = -30,5 KJbok/MomsSTH ape peakwun cBssarbl OGM MpoMeEryTON- noiwit mpoaykrasn P; 1 H,O 1 x MoxkHO mpencra~ BIITD Kak rlocnelonatembutble peaKUNH: (1) Pmoxosa + Pj rntoKo30-6-cpoctbar + HO (2) ATP +H,O > ADP +P, Himoeo: ATP + rmoKosa > ADP + raioxoa0-6-poctpar O6umee nsmenenne craxjaprHoit cBoGonHOit aHep- THH MOxKHO TOyYHTb Kak cyMMy AG” otesbHbix peaxinit: AG" = 13,8 KJbx/monp + (~30,5 KJbx/som) = = 16,7 KJbk/Mom CymmapHas peakua ak3epronmyeckas, H aHep- ria, sanacentas B ATP, ucrospayeres 118 cHHTesa rmioKo30-6-tpociata, XoTA O6pasoBaHine aToro coe- AUIHEHHSL M3 FIIOKOSH H HeopranttyecKoro ¢octara P, — stimepronnueckuit npouecc. Pearoreul nymo o6pasoBanua rmoKos0-6-poccbara B KueTKe TyTeM nepentoca boctbarnoii rpyni or ATP oranuaeres oT mpupenentisix Borme peaxnuii (1) 1 (2), Ho KO- Heunbili pesyubtar noay4aercs TOT %KE caMBIii, YO H mp cyMMupowatit 3THx aByx peakunii, B tep- MOJIMHAMHYECKHX PacueTax COCTOAHMA CHCTEMBI B Hayate H KOHUe Mpoyecca MpeaCTaB.eHbI ITHMH Be mectRaMn. A NYT’ MeKAY HavacIbHBIM M KOHEYHbIN COCTOSIIMAMH He BaxKen. Mbt yxe ropopran 0 Tom, ¥to AG” — cnoco6 BbIP@KEHHA KOHCTAHTH paBHoBecns peakuwn. JL1a peaxustnt (1) _ [r110K030-6-ocibar] - ian “1.” —“Trmoxosa] [Py] 29" 10°M 3amerun, uo H,O ne BxomuT p 970 BLIparxKerne, 10- ckonbKY KoHUerTpanua Bomb (55,5 M) He H3MeHs- eTca B xone peakunn. KorictaHta paBlosecna peak- un riapossa ATP pasiia (ADP][ Pi] 7 = 2,0 -10°M " [ATP] @ KOHCTaHTa paBHOBeCHs! JLBYX COMps>KeHHBIX peakiuii [rm0Ko30-6-choctpar][ ADP] P,] Keay= [ruoxosa] [P,][ATP] = (Ky) (Keg,) = (3,9 10° M-) (2,0 10° M) =78- 10 13.1. BwoakeprermKa wtepmopwamnxa [19] Suece caenyer O6patHT BHHMaHHe Ha BaxKHbIii MO- Ment, Kacaiouumiica KoHCTaHTHL paBHosecua. Xora AG” apyx peaxunii, 13 KOTOpbIx CHaraeTcA TpeTbA, addumusnes, Ki, cyMMapnoli peakun paptia npo- useedenuo K', 9THx otTAeNbHBIX peakunii. Tozopst, 4TO KOHCTAHTH! paBHoBecHa peakunii Myzomuntu- Kamuenoi. Bnaronapa conpsxenmio rHmpomusa ATP c cunresom rtoKos0-6-cpocdpara K’, ypeuu- unpaetcs B 2,0 - 105 pasa. Dra crparerus, rae mpit cuntese BroprunLx MeTaGO.MTOB M KeTOUNLX KOMMOHEHTOR MpHcyT- creyer o6umii mpomexyTounbiii mpoayKr, Bcrpe- 4YaeTCA BO BCEX XKUBBIX KNeTKaX. OyeBHAHO, YTO ona paGotaer TOAbKO MIpH yesoBHH NocTomHoro HanH4na TakHX coenuHeHMii, Kak ATP. B cnenyio- MNX ru1aBax MBI OGcysIM HeKOTOpBIE H3 HanGo.ee BaxkHbIx nyteli cunresa ATP. Kpatkoe conepkanne pasgena 13.1 BMO3HEPTETMKA M TEPMOAMHAMUKA @ = Kuspie krerku nocrosHHo BbinoaKmor pabory. ZLas nonaepxxatins cBoux BbicoKooprannsoBalt- HBIX CTPYKTYP, CHHTe3a KIeTOUHBIX KOMITOHEH- TOB M MHOTHX APYTHX MpoueccoB KeTKaM He- o6xoquMa 9HEprHs. @ Buosnepreruxa sannMaercs KOuIMYECTBeHEBIM way4eHHeM fpespailieHHii 9Heprun B OvonorH- vecknx cucremax. Buosornyeckue npespauie- HMA SHEprHM TOAYMHAOTCA 3aKOHAM TepMomH- HaMHKH. @ = Ha xoa mo6oii xumuyeckoit peaKuim BrHsOT ipa cbaktopa: crpeMuienie K COcTOsMITIO ¢ Han Gonbuteli ycroiiusBoctb10 (powHocTBI0) cBxseli (xapakrepuayercs arranpnneit H) ucrpemmenne Kk Hanpbicuiell HeynopaoyennoctH cuctemEt (xapaxrepuayeres autpormeit 5). Cymmapuoit np KYWWel cHaolt peakUMM sBiseTcA UaMeHe- Hue cBo6onHoii aHeprun AG; sToT TepmMoanni Muueckuii MapaMerp yunTEIBaeT OOuHii acpcpeKT atx AByx (baxropos: AG = AH — TAS. @ Hamenenue cranaaptioii cnoboanoii anep- Tim — xapaxrepnicTieckuit pianko-xiMn- 4eckHit Hapamerp AaHHoii peakUNN; ero MOK HO paccuiirats 13 KOHCTaHTH paBHoBecns: AG" = — RT In Ki[20] Yacte II. 13. Ochoss 6uosHepreTHkM. Tunis xuMMYECKHx peaKunit = B peanpnoii cucteme uamencnue cBoGoanoii anepruu AG He KoucTatTa, a 3apncuT or AG™ 1 OT KOHUeHTpauntii HexXoMHBIX BeULeCTB H TIpO- AYKTOB peak: 6 mpoaykrst peakwin =Acr + poatykrel peak] ac ~ac" + Rtn resoame nemtecrpa @ = Koraa AG < 0, peaxuna npotekaer cnepa Ha- npavo (Tr. €. B MIpsMOM HampapaeHitn); Korma AG < 0, peakuna crpemutca natH B O6patHoM Hanpaprennn; 1 Koraa AG = 0, cucrema Haxo- ATCA B COCTOAHHM pannonecna. Hsmerienne cnoGoanoit aneprin peanut ne sa- Bueutt or NyTH MporeKanuia peaxuuut. ismene- Hite cBoGos Hoi SHEprHIH HeCKONBKHX MOcTenO- parembHBIX peaxtuii (cranuit) — ananTHBHBtii Hapamerp; o6utee uamenenne cBo6onHoit auep- rut cymMapnoii xumiueckoil pean, KoTOpas ABNACTCH PesysILTATOM MocMEOBATENBUDIX Pe- akunii c O6MIMM NpoMexKYTOUHBIM MposyKTOM, pasio cymMe AG ornenbubix peakunit. 13.2. XAMMY4eCKME OCHOBbI 6moxumuyeckux peakynin Bromormueckne mpouecchi c mepenayei sneprun, © KOTOPbIX MBI TOBOPHM 3/eCb, NpeNCTABIsIOT CO- Goi onpenenenpie xuMMYECKHe peakusH. B KneT- Ke MPOHCXOAAT He BCE THI peakunti, c KOTOPBIMIT OGBI4HO sHaKoMaTCA B O6uIeM Kypce opranHuecKoii xumun, To, Kakue peakuHH mpoTeKaioT B 61010- THYECKHX CHCTeMax, a KaKMe Het, ompeensetcs 1) ux porbio B KoNKpeTHOi 6uonorHyecKoii cHCTe- Me n 2) ux ckopoctamn. O6a aTHx ¢bakTopa UMeIoT BaxHoe sHaveHe 8 (popMHpoBaHH MeTaGosMue- CKHX nyteii, © KOTOpEIX Mbl GyjeM roBOpHtTD slaTee. ShainMas peakunA — 3TO Ta peakUMA, KOTOpan HC- nosbayer HMeoUUtiicA B HAHN cyOcrpar H Mpe- Bpauiaer ero B HY>kHbIit MpoayKt. OattaKo axe aH 3HaYMMbIe (MOTEHLMaHO) peak He Beer/a MO- TyT mpotekath B Knerke. Hexotoppre xuMuyecKHe TIpeBpallleHHisl HACTOsTBKO 3AME/IEHHBI (T. €. HMEIOT HACTONBKO BBICOKHE SHEPrHM AKTHBaMWH), YTO HE MOFYT IIPOHCXO/MTD B XKUBBIX CHCTEMAX 1a2Ke B IPH CYTCTBHH MOMHDIX KaTasHZaTopoR — depMentTos. Te peakunn, Kotopsie jelicranTenbHo mporeKalor B IRMBEIX KuTeTKAX, MpencTasamoT coGolt BosHHKUITi Blpollecce aBoMONN HaGop XUMHYECKHX «nHeTpy- MEHTOB>, MOSBOTAIOUINI OOXOAHTH ¢HeBOSMOXKHBIC> peakuun. Jia xopomero nonmManmia 61oxnMTH KJETKH HEOOXOAMMO YMeTb pacllo3HaBaTb Te peak- IIH, IpOTeKAHHe KOTOPHIX BOSMOXKHO. Hecmorps Ha Bce ykasanHible Orpansuerita, KO- uimuectBo MeTaGouIM¥ecKHX Mpenpailenuii B O6LY- Hoii Knerke orpomHo. Boabuniicrso KuleToK o6s1a- RaWT CHOCOSHOCTHIO OCyMlecTBAATD THICAYH crrett- uuyeckux cbepMentarneHbix peaxunit, Hanpumep MpeBpalllaTb MIpocToe BeULeCTBO, Takoe KaK TOKO3a, B aMMHOKHCJOTHI, HYKEOTHADL WJM MMHIbL, H3BIe- KaTb dHeprivio 13 BellecTB, MocTynaloumx ¢ mHueli, TYTEM HX OKHCEHHA HIM MPOBOAHTS MomHMepHsa- TUM MoHOMepHBIx MoxteKyz1 ¢ O6pasonarttem Makpo- MosieKys1. YroG1 uaysuTh Bee 9TH peaKUMH, HeobxosUM cucremaruyeckuii noon. XHMMYECKHE peaKin, MPOHCXOAAMIME B %*KUBBIX OPraH3MAX, MOXKHO pas- AEAUTD Ha TUMI, H Torna He HoHaLOGuTCH Hay4aTb K@KAYIO OTACAbHY!O PeakUNO, WOOBI MOHATH «10- TUKY®, 3A1OKEHHYIO B OMOXHMHYECKHX Mpolleccax. Muoxectso peakunii B XXMBbIX KeTKaX MOXKHO kalaccuHNMpoBarb, BUIeNHB NaTb THTOB: 1) peak- UNH, HAYLINe c OOpasoBaHHeM WIM paspbiBOM CRASH yraepoa—yrepox; 2) BHYTpUMONeKY.IApHBIe nepe- TPYMMMpOBKH, H3OMEpHsaUHA 1 |MUMHMpOBAKHE; 3) peaxum ¢ yuactnem cnobonnBIx pamitKaron: 4) peaxiun ¢ meperocom ¢byHKunoranietx rpynn; 5). oKNcANTeNbHO-BOCCTaHOBITENbHBIe peaKtunt. Alanee met noapoGHee oGcyanM pasni4unbie THIB peaxunii Ha KOHKpeTHDIX MpuMepax OMoxHNHE- cKHx npespatennii. 3amersre, yro peakuuu, OTHO- CAUIMe C pasHOMy THITY, He HCKONAIOT Apyr Apyra, HanpuMep peaxiuia H3OMEepH3alN MoxET HATH 10 cno6oqno-panitkabHOMy MexanuaMy. Onuako npexxne sem nponomKue obcyxnenHe THNOB peakiuii, HaM CleAyeT BCNOMHHTb Ba T1aB- HbIX XHMHMECKHX MpHiuitia. Tepepiii npr 3a- KMOYeTCA B TOM, 4TO KOBAIeHTHAA XHMHYECKaA cBa3b O6pasyerca B pesysbrate oG6obiwecTBTeHHA Tapbl as1eKTpoHOB, a paspbiB STO! CBASH MOXET TIPO- HCXOAMT ABYMA TlyTAMH (puc. 13-1). [pi romomH- THYECKOM paspbibe Kaxk [bili ATOM, YYaCTBOBABIINI B o6pa3z0BaHMn CBa3H, NpeBpalllaeTca B CBOGO/HBII pa- auKa, Hecyuutii ommH HecnapenHblii anexTpon. Tpit TeTeposMTHYeCKOM paspbiBe, KOTOPEIii NPOHCXOAHT ropa3Z0 Yale, O6a auleKTpOHa OCTAIOTCA CBAZAHHLIMH ¢ OHM 13 AByx atomos. Ha pu. 13-1 ns06paxentt actin, KoTOpHIe HanGostee YacTo ObpaayioTes pit pacurerienut cpsiseii C—C mw C—H. KapGannouns,