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CHNG 6. H HP CA THC VT
3. C ch qu trnh h hp Khc vi qu trnh t chy cht hu c ngoi c th, qu trnh oxy ha trong c th phi tri qua nhiu chng, bao gm nhiu phn ng ha sinh cui cng gii phng CO2, H2O v nng lng vi dng ATP. C th chia qu trnh ny thnh 2 giai on: Giai on 1: tch hydro ra khi c cht h hp. Giai on ny c thc hin bng ba con ng khc nhau: - ng phn v ln men - ng phn v chu trnh krebs. - oxy ha trc tip ng qua chu trnh pentozophotphat. Giai on 2: oxy ha cc cofecment kh tng hp ATP. Giai on ny xy ra trn mng trong ca ty th, bao gm 2 qu trnh din ra ng thi v song song nhau: qu trnh chuyn vn electron v qu trnh photphoryl ha. 3.1. H hp hiu kh v h hp ym kh (ln men) Cy xanh phng thch CO2 khng ch to iu kin bnh thng ca kh quyn m c trong mi trng khng c oxi. Bng nhng dn liu chnh xc Pasteur ch ra rng s gii phng CO2 ca cy xanh trong iu kin khng c oxy khng ph thuc vo s c mt ca vi sinh vt m l kt qu ca cc qu trnh din ra trc tip ngay trong m thc vt. ng cn chng minh ln men l s sng khng cn oxy v cho rng qu trnh ym kh c vai tr ln ngay c trong cc hot ng sng ca sinh vt hiu kh. Pluger ln u tin a ra kin h hp tin hnh theo hai giai on: giai on u cc nguyn liu h hp phn gii thnh cc hp cht d b oxy ho nh ru etylic, ti giai on sau cc sn phm ca s ym kh khi b oxy ho bi oxy ca kh quyn n nhng cht cui cng l kh cacbonic v nc. Quan im c cc nh thc vt hc ni ting nht hi cui th k 19 tha nhn. Pleffer coi ru l mt sn phm trung gian ca qu trnh h hp hiu kh thng thng ca thc vt v qu trnh ny c lin quan vi s ln men. Nhng hc thuyt ca Cotxtuxev khng nh: h hp v ln men hon ton khc nhau nhng c quan h vi nhau. Giai on u bin i t ng n cc sn phm trung gian (axit pyruvic) l chung cho c h hp v ln men, sau phn ly thnh hai qu trnh khc nhau.

S sai khc gia h hp v ln men l trong qu trnh h hp hiu kh cc sn phm trung gian b oxy ho bng oxy t do dn n s hnh thnh CO2 v H2O. qu trnh ln men nu nh cc sn phm trung gian c to nn ca giai on u bin i tip tc trong iu kin khng c oxy th sn phm cui cng s l ru v CO2. 3.2. Qu trnh ng phn Con ng E.M.P. (Embden Meyerhof Parnas) Giai an chun b chung ca qu trnh h hp v ln men. Qu trnh ng phn bao gm trn mi phn ng ring bit din ra theo mt trnh t xc nh v c cc h enzym tng ng xc tc, chia thnh 4 chng: a) Chng hot ha phn t ng: nh ATP cc enzym tham gia, glucoza bin i ln lt thnh dng este mono v k thnh dng este diphosphat rt linh ng v d bin i. Vic chuyn ha dng gluco-piranoza thnh fructofuranoza cng lm tng thm tnh linh ng ca nguyn liu h hp. b) Chng phn ct ng 6C thnh ng 2C3: Cu trc i xng ca F1,6-P d b gy gia C3 v C4 thnh DOAP v Al-3-PG. DOAP d dng bin i thnh Al-3-PG tip tc bin i v trnh vic mt lt glucoza tham gia trong phn ng. c) Chng oxy ha Al-3-PG thnh Al-2-PG. chng ny sinh ra nng lng gii phng trong phn ng oxy ha. H c tch ra v chuyn n NAD v sau chuyn cho enzym khc v cui cng cho oxy kh tri trong trng hp h hp hiu kh hoc c s dng kh cc nguyn liu hu c khc nh axit piruvic, axetaldehyt trong hnh thc ln men. d) Chng chuyn ha Al-2-PG thnh axit piruvic. Chng cui din ra phn ng tch nng lng th hai v bin mt oxy axit thnh mt xeto axit c kh nng phn ng cao v c v tr quan trng trong qu trnh trao i cht. Phng trnh cn bng tng qut ca ng phn c th tm tt nh sau: C6H12O6 + 2NAD + 2ADP + 2H3PO4 2C3H4O3 + 2NADH + 2H+ + 2ATP + 2H+ Cng cn phi ch rng, nhng phn ng ca qu trnh ng phn cho ta nhng cht tin thn c th tham gia trong s hnh thnh mt s Axit amin. Bng cch kh hydro ha, APG c th to thnh axit phosphohydroxypiruvic v axit ny l tin cht ca serin. Serin c th to thnh mch bn ca Axit amin c nhn benzen (phenylalanin, tyrozin) v nhn indol (tryptophan); serin cng l cht tin thn ca Cystein hay glysin. APEP c th nh CO2 v cho axit oxaloaxetic. Axit

ny trong s chuyn v amin s thnh axit aspactic v aspactic l tin thn ca methionin, threonin v -alanin. Axit piruvic c th bin thnh alanin bng chuyn v amin. 3.3. Qu trnh ln men Trong iu kin khng c oxy, axit pyruvic bin i theo con ng ym kh theo cc hnh thc ln men to ra cc hp cht n gin hn cn cha nhiu nng lng. Ty thuc vo sn phm hnh thnh trong qu trnh ln men ngi ta phn bit cc dng ln men: ln men ru, ln men lactic, butyric 3.3.1. Ln men ru etylic Di tc dng ca enzym decacboxylaza, axit pyruvic s b kh cacboxin ha gii phng kh cacbonic v aldehytacetic. Phn ng xy ra nh sau:

Nh c NADH2 aldehyt acetic b kh thnh ru:

Sn phm cui cng ca ln men ru l ru ethylic v kh CO2. 3.3.2. Ln men axit lactic S ln men lactic tin hnh bi cc vi khun lactic (lc mui da, lm sa chua, lm tng) Axit pyruvic b kh trc tip bi NADH2 di tc dng ca enzym dehydrogenaza to axit lactic.

3.4. Qu trnh phn gii hiu kh axit pyruvic 3.4.1. Chu trnh Krebs Khi c mt ca oxy, axit pyruvic s chu nhng bin i phc tp. Nhng bin i ny l mt chu trnh ca cc phn ng c tn l chu trnh Krebs. Chu trnh

Krebs cn c tn l chu trnh axit citric hay chu trnh axit di- v tricacboxylic pht hin nm 1937. Chu trnh ny l s k tc trc tip ca cc qu trnh ng phn trong t bo sng, n rt ph bin trong m thc vt bc cao v m ng vt. Chu trnh Krebs din ra ba nhm qu trnh quan trng: loi tr hoc kt hp nc, kh cacboxyl v oxy ha. Nh kin ca Palain 1914 xut. nc l mt nhn t tham gia tch cc trong qu trnh phn gii cht hu c trong c th sng. Nc ng vai tr l cht oxy ha, cung cp oxy oxy ha cacbon ca nguyn liu to nn sn phm cui cng l CO2. Cc phn ng kh cacboxyl trong chu trnh c tnh cht l phn ng ym kh. Ngun gc CO2 l cc cetoaxit, nhm cacboxyl cc cht ny d dng tch ra di tc ng ca nhng cacboxylaza tng ng. Ngun gc CO2 l mt trong cc im sai bit gia h hp v t chy v trong qu trnh t chy, CO2 that ra l kt qu trc tip ca s oxy ha cacbon nguyn liu bng oxy ca kh tri ht vo, cn trong qu trnh h hp CO2 c gii phng khng ph thuc vo s hp th oxy. Cc phn ng oxy ha kh tch H+ v in t vn chuyn qua dy chuyn in t l nhng phn ng quan trng din ra trong chu trnh Kreb v chnh chng cui cng ny nng lung v H 2O c hnh thnh, chu trnh c tm tt bng phng trnh:

Rt nhiu thc nghim xc nhn tnh ph bin ca chu trnh krebs trong cc m thc vt. Ngi ta pht hin c tt c cc axit hu c v cc enzym tham gia trong chu trnh krebs. Nu thm axit hu c ca chu trnh s kch thch s h hp ca m thc vt. Malonat l cht km hm cc phn ng chu trnh Krebs c ch qu trnh h hp ca m thc vt. Nh nu trong qu trnh ng phn cng din ra mt phn ng oxy ha bin AlPG thnh APG v ng vi 1 phn t glucoza, 2 cp H2 c dehydraza ym kh tch ra s tc dng vi 1 phn t oxy ca kh tri to thnh 2H2O. Tng hp li ta c phng trnh tng qut ca h hp hiu kh:

C6H12O6 + 6 O2 + 6 H2O

6 CO2 + 12 H2O

Rt gn li:

C6H12O6 + 6 O2

6 CO2 + 6 H2O

3.4.2. Chu trnh axit Glyoxylic Nm 1957 Kornberg v Krebs pht hin c chu trnh axit glyoxylic, mt dng ci bin ca chu trnh crep. S sai bit c bt u axit isocitric tc ng ca enzym isoitraza, axit isocitric bin thnh axit succinic v axit glyoxylic. Axit glyoxylic s ngng t vi axit acetic nh enzym malatsynthetaza xc tc v to thnh axit malic. Chu trnh axit glyoxylic pht hin thy nm mc, vi khun v mt s thc vt, cc phn ng ca n l c s ca vic bin i lipit thnh gluxit, thng xy ra trong s ny mm ca ht c du.

Hnh 6.1. Chu trnh glyoxylat bin i lipit thnh ng cn cho s ny mm ht c du 3.5. Con ng pentozo photphat Con ng oxy ha cc cht hu c cn c tn l s oxy ha hexozomonophosphat hoc oxy ha trc tip hoc con ng chuyn ha ca pentoza. l con ng bin i hiu kh gluxit thng gp thc vt bc cao v

vi sinh vt. phn ng u ca n trng vi phn ng ng phn. S khc bit c bt u t G-6-P. Khc vi ng phn, trong qu trnh ny G-6-P khng qua chng ng b i phn t thnh 2 trioza m n b oxy ha thnh cc hp cht cacbon. Cc hp cht ny qua nhiu phn ng trung gian v cui cng li to thnh G-6-P. Hp cht ny lm ng kn qu trnh trn. V th qu trnh ny gi l chu trnh pentozo photphat hoc hexozomonophosphat. Tt c cc giai an ring bit ca chu trnh ny uc chng minh bng thc nghim trn cc i tng nm men, ng vt cng nh thc vt. Mc tham gia ca n cc loi cy, trong cc giai an pht trin, trong cc iu kin ngoi cnh khc nhau thng khng ging nhau. Pentoza c bit c ngha i vi qu trnh sinh trng v qu trnh sinh tng hp nhiu hp cht hu c quan trng nh axit nucleic, Coenzym NADP, ATP Pentoza tham gia vo qu trnh quang hp l hp cht u tin nhn CO2 v ta c th xem v c bn qu trnh ny ging chu trnh calvin trong quang hp nhng din ra theo chiu ngc li. Hai giai an u l nhng phn ng oxy ha i hi phi c NADP tham gia. phn ng th nht G-6-P oxy ha to thnh 6-phosphogluconat, cht ny s b kh cacboxyl v oxy ha. Kt qu l to thnh pentoza v mt phn t CO2 c gii phng. Pentoza c to thnh dng Ru-5-P, cht ny b ng phn ha bng cch nh sau: c 3 phn t ribuloza to thnh th 2 phn t bin thnh xilulozo-5-P(X-5P) v mt phn t bin thnh R-5-P. Trong cc phn ng tip theo ca chu trnh c s tham gia ca cc enzym transxetolaza v transaldolaza chuyn tan b mch cc nguyn t cacbon. Nh enzym transxetolaza cc pentoza va to thnh s phn ng vi nhau, transxetolaza s chuyn nhm Glycolaldehyt hot ng t X-5-P n R-5-P. kt qu l to thnh sedoheptulozo-7-phosphat (Se-7-P) v gc trioza-3-P. Cc sn phm ca phn ng ny mt ln na li tc dng vi nhau nh transaldolaza. Kt qu l to F-6-P v E-4-I. E-4-P phn ng vi X-5-P to thnh F-6-P v triozo-P. Hai trioza-P ngng t vi nhau nh aldolaza cho ta phn ng Fr-6-P th ba. Chu trnh pentozophosphat c biu din nh sau: 6 G-6-P + 6 H2O + 12 NADP 6 R-5-P + 6 CO2 + 12 NADPH2 6 R-5-P 5 G-6-P + H2PO3 G-6-P + 12 NADP 6 CO2 + H2PO3 + 12 NADPH2

V c th s ha s bin i 6 R-5-P thnh 5 G-6-P nh sau:

Hnh 6.2. Chu trnh Pentozo-P

3.6. S oxy ha lipit v protein 3.6.1. Oxy ha lipit Hm lng lipit trong thc vt ni chung khng cao. Trong ht cc cy c du v mt s cy h u tch ly mt s lng lipit kh ln. S thy phn u tin thc hin bi lipaza xc tc s phn gii lipit thnh glycerin v axit bo.

Khi oxy ha glycerin c th bin thnh Al-3-PG v sau thnh axit pyruvic theo ng phn. Cn cc axit bo th chu s - oxy ha, s oxy ha xy ra nguyn t cacbon v tr so vi nhm cacboxyl.

giai on th nht, Coenzym A c kt hp vo phn t axit bo. Phn ng ny i hi phi c s tham gia ca ATP. Sau c s kh hydro ha gia nguyn t cacbon v ca axit bo. Phn ng tip theo l s kt hp nc vo v tr ni i (do enzym crotonaza), loi tr H nguyn t cacbon ca axit bo (do dehydraza c hiu vi NAD). Giai an cui cng l s phn gii cetoaxit v kt qu l to thnh acethyl CoA v gc axit bo c hai nguyn t cacbon t hn so vi axit bo ban u. Axit bo mi to thnh li c th b oxy ha nh trn v phn ng tip tc nh th mi cho n lc axit bo b phn gii han ton thnh acethyl CoA. Acetyl CoA sau tip tc bin i theo chu trnh krebs. 3.6.2. Oxy ha protein thc vt protit phn gii thnh axit amin. Trong thin nhin cc protit thc vt u c hnh thnh cc L.axitamin. Mt s lng ng k cc D-axitamin ch thy c trong nm mc v trong vi khun. Vic kh amin ha bng cch oxy ha l ng hng c bn ca s oxy ha axit amin trong c th sng. Lc u axitamin b oxy ha to thnh axit amin, sau nc s kt hp vi axit amin v loi tr NH3. Trong s kh axit amin ha ny c s tham gia ca enzym glutamicodehydrogenaza c hiu vi NAD. V d :

Ngoi s oxy ha nh glutamicodehydrogenaza cc axit amin cng c th b bin i nh cc enzym c bit c bn cht flavoprotein gi l cc oxydaza ca axit amin. Cc enzym ny cng thc hin c qu trnh kh amin ha bng cch oxy ha nh trn. So vi qu trnh trn, qu trnh kh amin ha axit amin nh oxydaza ch khc c mt im l H do oxydaza ly khi c cht trong trng hp ny s b oxy ha thnh peroxyt hydro:

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