Qu trnh hp hp

CHNG 5 S H HP T BO
T bo cn rt nhiu nng lng duy tr s sng: t vic sao chp, sa cha cc cu trc di truyn trong nhim sc th; to mi cc thnh phn cu to trong t bo, ly thc n vo, thi cht b ra, gi cho pH v nng ion c cn bng... Nu nng lng khng c cung cp cc phn ng khng th xy ra c v s sng ca t bo s ngng li. Tht ra, tt c nng lng cung cp cho s sng ngy nay l t mt tri, c cy hp thu v qua qu trnh quang hp to ra hp cht hu c giu nng lng. Hu ht cc sinh vt khng quang hp ly nng lng bng cch tiu ha cc sinh vt quang hp c v nhng sinh vt khc. Trong t bo, s h hp to ra nng lng cung cp cho tt c cc hot ng ca t bo.
I. I CNG 1. S tin dng v thoi dng

Trong t bo, nng lng c d tr trong cc hp cht c tng hp do qu trnh quang hp. Nng lng ny c gii phng qua qu trnh h hp hiu kh, trong glucoz c chuyn ha qua nhiu sn phm trung gian n sn phm cui cng l CO2 v H2O. Nng lng t do c to ra trong qu trnh ny c s dng nh l ngun nng lng cho nhng phn ng khc xy ra trong t bo. Tt c cc phn ng ny c gi l s bin dng ca t bo (cellular metabolism) v thng c chia thnh hai giai on: s tin dng (anabolism) l qu trnh nhng cht hu c phc tp c tng hp t nhng phn t n gin hn, v s thoi dng (catabolism) l qu trnh trong c th sng ly nng lng t thc n bng cch phn ct nhng phn t hu c phc tp thnh nhng phn t n gin hn. S h hp hiu kh (aerobic respiration) l mt kiu thoi dng chnh xy ra trong t bo.
2. Ty th

Ty th l bo quan tham gia vo s h hp hiu kh. Di knh hin vi in t, mi ty th c bao bc bi hai mng, mng ngoi v mng trong chia ty th ra lm hai ngn: ngn ngoi l khong gia ca hai mng, ngn trong t mng trong tr vo. Mng trong l nhng crista, nn b mt tip xc tng rt nhiu (Hnh 8, chng 1). Mng ngoi thm c nhng phn t nh, nhng mng trong th khng thm. Trn mng trong c cc protein knh, knh ng

m, bm iu tit chn lc nhng phn t i ra v i vo ngn trong. Mng trong cng cha nhng protein ca chui dntruyn in t tng t nh chui dn truyn in t trn mng thylakoid ca lc lp. V tht ra, s h hp ging nh s quang hp cng bao gm nhng phn ng oxy ha kh.
II. S H HP CARBOHYDRAT

Nhng carbohydrat phc tp nh tinh bt thc vt v glycogen ng vt u c thy phn thnh nhng phn t glucoz trc khi i vo qu trnh h hp. S h hp hon ton mt hp cht giu nng lng nh glucoz tri qua mt chui di nhng phn ng, trong c nhng phn ng xy ra khng cn s hin din ca oxy v c nhng chui phn ng l thuc vo oxy.
1. ng phn (glycolysis) (giai on I) (Hnh 1)

Hnh 1. Cc phn ng chnh ca ng phn

ng phn l giai on u tin ca qu trnh h hp glucoz xy ra khng cn s hin din ca O2. ng phn xy ra trong dch t bo cht ca tt c t bo sng, v l chui phn ng xy ra nhng sinh vt u tin khi m tri t cn cha c O2. Glucoz l mt hp cht bn vng, t c xu hng phn ct ra thnh nhng cht n gin hn, do t bo mun ly nng lng t glucoz trc tin phi u t cho n mt t nng lng hot ha phn t. Do , giai on u ca ng phn l cung cp ATP cho phn t glucoz (Hnh 1). Trong cc phn ng chun b hai phn t ATP gn gc phosphat cui cng ca n vo phn t glucoz.

Trong phn ng ny hexokinaz xc tc chuyn mt gc phosphat vo glucoz. Phn ng k tip l phn ng chuyn i glucoz-6-phosphat thnh fructoz-6-phosphat.

Sau khi to ra sn phm trong bc (2), mt phn t ATP na c tiu th thm mt gc phosphat na vo phn t.

K tip fructoz-1,6-bisphosphat b ct i gia C th ba v C th t to ra hai cht 3C tng t nhau trong bc (4). Mt cht l PGAL (phosphoglyceraldehyd) v mt cht trung gian thng chuyn i ngay thnh PGAL trong bc (5). PGAL l mt ng trung gian 3C, l cha kha trung gian trong c qu trnh ng phn v quang hp.

n giai on ny qu trnh ng phn s dng 2 phn t ATP. Phn ng k tip, hi phc tp hn, bt u dn n s thnh lp ATP mi, tht s l hai phn ng. Phn ng u l mt phn ng oxy ha kh: hai in t v mt ion H+ c ly t mi phn t PGAL (nh vy phn t ny b oxy ha) bi phn t nhn in t nicotinamid adenin dinucleotid, hay NAD+, cht ny b kh. NAD+ rt gn vi NADP+ tm thy trong lc lp. Trong trng hp ny sn phm trung gian l NADH thay v l NADPH. Phn ng th hai l s phosphoryl ha PGAL. Nng lng c gii phng t s oxy ha PGAL c dng gn mt gc phosphat v c P vo PGAL, gc phosphat c gn vo bng mt cu ni giu nng lng.

Trong phn ng k tip, gc phosphat mi c chuyn vo ADP to ra ATP. Trong qu trnh ny, mt gc phosphat giu nng lng c chuyn vo mt c cht ADP to thnh ATP, phn ng ny c gi l phosphoryl ha mc c cht (substrate-level phosphorylation). Sn phm 3C l PGA, mt cht trung gian trong chu trnh Calvin-Benson, mt ln na cho thy s tng quan gia hai qu trnh:

giai on ny, t bo thu li c 2 phn t ATP dng cho s phosphoryl ha glucoz trong lc bt u ng phn. Nng lng u t ban u c tr li. Qua phn ng k tip, cui cng l nc c tch ra t PGA, v sau gc phosphat c chuyn i v c gn li bi cu ni giu nng lng:

Sau phn ng sp xp li trong bc (9) gc phosphat c chuyn vo ADP theo s phosphoryl ha mc c cht thnh lp ATP, kt qu to ra hai phn t ATP v hai phn t acidpyruvic:

V hai phn t ATP s dng trong bc (1 v 3) c ly li trong bc (7), nn hai phn t ATP ny l c tng hp thm cho t bo.

Cc im quan trng cn ch trong s ng phn l: Mi phn t glucoz (C6 H12O6) b phn tch thnh hai phn t acid pyruvic (C3H4O3). Hai phn t ATP s dng trong lc u ca qu trnh, sau c bn phn t c to ra, nh vy t bo cn c hai phn t ny. Hai phn t NADH c thnh lp. V khng s dng oxy, qu trnh c th xy ra d c s hin din ca O2 hay khng. Cc phn ng ca ng phn xy ra trong dch t bo cht ca t bo, bn ngoi ty th.
2. S ln men (fermentation) (Hnh 2)

Trong ng phn, hai phn t NAD+ c kh thnh NADH. Chc nng ca phn t NAD+ trong t bo l vn chuyn in t, trao i ion H+ v in t gia cht ny v cht khc. Do , NAD+ ch l mt cht tm thi ti ion H+ v in t, sau khi chuyn ion H+ v in t cho cht khc th n tr li lm cht ti tip tc. Nu NADH c thnh lp khng nhanh chng loi b ion H+ v in t th NAD+ trong t bo s thiu; khi bc (6) trong ng phn khng th xy ra v qu trnh ng phn tm dng li. Nh vy, s oxy ha NADH thnh NAD+ l cn thit cho qu trnh ng phn tip tc. Trong hu ht t bo, nu c O2, n s l cht nhn in t cui cng t NADH. Nhng di iu kin ym kh, khng c O2 nhn hydro v in t th acid pyruvic c to ra trong qu trnh ng phn s nhn hydro v in t t NADH, qu trnh ny c gi l s ln men.

Hnh 2. Cc phn ng chnh ca s ln men S bin dng acid pyruvic trong s ln men thay i theo c th sinh vt (Hnh 2). t bo ng vt v nhiu vi sinh vt s kh acid pyruvic to ra acid lactic:

hu ht t bo thc vt v men (yeast), sn phm ln men l ru ethyl v CO2, qu trnh ny c p dng trong sn xut:

V vy, trong iu kin ym kh, NAD+ nh con thoi i qua i li gia bc (6) v bc (11), ly hydro v in t to ra NADH trong bc (6) v tr li hydro v in t trong bc (11). S ln men l s ni tip ca qu trnh ng phn, bng cch ny glucoz c bin i thnh ru hay thnh acid lactic di iu kin ym kh, v tn gi ty thuc vo tn ca loi sn phm cui cng. D sn phm cui cng l cht no th s ln men ch ly c mt phn nng lng rt nh t glucoz, cc sn phm ca qu trnh ln men l cc cht cn cha rt nhiu nng lng t do. S ln men ca nhng t bo men v nhng vi sinh vt c ng dng trong nhiu k ngh quan trng nh lm bnh m, pho ma, da ua, sn xut ru v cc loi thc ung c ru; mt s sn phm khc c to ra t cc vi sinh vt khc nh mi c trng ca pho ma Thy s l do sn phm ln men l acid proprionic.
3. S oxy ha pyruvic acid (giai on II)

Nu c s hin din ca O2, th O2 l cht nhn in t cui cng t NADH, do acid pyruvic s c a vo ty th v y s c tip tc bin dng v ng thi to ra nhiu ATP mi. Acid pyruvic trong dch t bo cht c chuyn vo ngn trong ca ty th. Qua mt chui phn ng phc tp, acid pyruvic b oxy ha thnh CO2 v mt gc acetyl 2C, cht ny gn vi mt coenzim c gi l coenzim A (CoA) to ra cht acetyl-CoA. Khi acid pyruvic c oxy ha, in t v ion H+ b ly i, v mt ln na NAD+ l cht nhn in t v ion H+ to ra NADH. Chui phn ng phc tp ny c th c tm tt trong phng trnh sau:

Cui giai on II, 2 trong 6C trong glucoz ban u c gii phng ra di dng CO2.
4. Chu trnh Krebs (giai on III)

K tip acetyl-CoA i vo mt chui phn ng ca mt chu trnh gi l chu trnh Krebs (do nh khoa hc ngi Anh, Hans Krebs, c gii thng Nobel nh lm sng t c chu trnh ny) hay chu trnh acid citric. Nhng im chnh trong chu trnh c trnh by trong hnh 3. Mi phn t acetyl-CoA c to ra t phn t glucoz ban u kt hp vi mt hp cht 4C, acid oxaloacetic, hin din trong t bo to ra mt hp cht 6C mi l acid citric. Trong cc phn ng tip theo, 2C b mt i di dng CO2, nh vy hp cht ch cn 4C v c bin i tr li cht 4C ban u v chu trnh li

tip tc. V mi phn t glucoz to ra hai phn t acetyl CoA, nn c hai vng acid citric xy ra v tng cng l c 4C c gii phng di dng CO2; cng thm 2C c gii phng di dng CO2 trong giai on II, nh vy tt c l 6C ca phn t glucoz ban u.

Hnh 3. S ca chu trnh Krebs Trong mt vng ca chu trnh (Hnh 3), mt phn t ATP c tng hp (bi s phosphoryl ha mc c cht) v 8 in t v 8 ion H+ c ly i bi cht nhn in t. 6 in t v 6 hydro c dng kh 3 phn t NAD+ (to ra 3 phn t NADH v 3 ion H+) v 2 in t v 2 hydro c nhn bi hp cht FAD (to ra FADH2). V s oxy ha mt phn t glucoz tri qua hai vng chu trnh Krebs nn tng cng c 2 ATP v 8 phn t cht kh (6 phn t NADH v 2 FADH2) (Hnh 5.4).
5. S trao i nng lng v iu ha trong qu trnh h hp (giai on IV)

Ch c tng cng l 4 phn t ATP mi c li ra (2 trong ng phn v 2 trong chu trnh Krebs). Nng lng cn li c d tr trong hai cht kh giu nng lng l NADH v FADH2, tng cng c tt c 12 phn t c tng hp trong sut qu trnh oxy ha glucoz: 2 phn t NADH trong ng phn, 2 phn t NADH trong qu trnh oxy ha acid pyruvic thnh acetyl-CoA, 6 phn t NADH v 2 phn t FADH2 trong chu trnh Krebs.

ATP c tng hp bng cch no? Di iu kin hiu kh, NADH chuyn in t n O2 v sinh ra NAD+. Ni mt cch khc, O2 l cht nhn in t v hydro cui cng to thnh nc:

Hnh 4. Tm tt cc sn phm trong ba giai on I, II v III t t NADH c chuyn qua mt chui dn truyn in t, cui chui l phn t O2 (Hnh 5). Cc phn t ti in t thng c gi l chui h hp (respiration chain) gn trn mng trong ca ty th (tng t s dn truyn in t trong chui quang hp trong lc lp) to ra mt gradient ha in, c dng tng hp ATP t ADP v phosphat v c. Qu trnh ny c gi l s phosphoryl ha chuyn in t (electron-transport phosphorylation) hay s phosphoryl oxy ha
Tuy nhin, khng phi NADH chuyn in t n O2 mt cch trc tip. in

(oxydative phosphorylation), bao gm c s ha thm thu (chemiosmotic) ca s dn truyn in t v s tng hp ATP. Protein dn truyn in t trong h hp l thnh phn chnh trong cho qu trnh phosphoryl oxy ha, to ra hu ht ATP sinh ra do s oxy ha glucoz. S phosphoryl oxy ha tng t s quang phosphoryl ha, s dng dng in t c chuyn trong chui dn truyn in t bm ion H+ qua mng, kt qu l sinh ra mt gradient ha in c dng tng hp ATP. Trong s h hp c hai cch to ra ATP: th nht l s phosphoryl ha mc c cht, trong mt gc phosphat c chuyn t mt hp cht cao nng lng vo ADP v th hai l s phosphoryl oxy ha, trong nng lng trong gradient ha in H+ c khai thc tng hp ATP.

Hnh 5. Chui dn truyn in t h hp S tng hp ATP trong ty th tng t s tng hp ATP trong lc lp. Nhng phn t dn truyn in t sp xp thnh 4 phc hp protein, gn

trn mng trong ca ty th (Hnh 5). Cht ti in t c ng nh phn t protein Q v cytochrom c (Q l mt coenzim ha tan trong lipid v c th di chuyn xuyn qua mng, cytochrome c (cyt.c) l protein tan trong nc nm mt ngoi ca mng trong ca ty th) chuyn in t t phc hp ny sang phc hp khc. NADH c to ra trong giai on II v III chuyn 2 in t v 1 ion H+ ca chng vo phc hp I v cng lc b oxy ha thnh NAD+. Cc in t c ti t cht ti ny sang mt cht ti khc theo mt trnh t c bit cui cng n O2 (Hnh 5). iu ny cho thy v sao phi cn c O2 trong s h hp hiu kh. Khng c O2 nhn in t, chui in t khng hot ng v NAD+ cn thit cho giai on II v III khng c to ra t NADH. Khi O2 nhn in t, n cng nhn lun ion H+ t cht c bn to ra nc, mt trong nhng sn phm cui cng ca s thoi dng glucoz. Cc nghin cu cho thy ch c in t c ti qua chui dn truyn in t, ion H+ khng i cng. Tuy vy, 3 trong 4 phc hp bm ion H+ t ngn trong xuyn qua mng trong ra ngoi. Kt qu l nng ion H+ ngn ngoi gia tng, ngn trong gim xung, v bn ngoi ca mng trong tr nn c in tch dng hn so vi bn trong v sinh ra gradient ha in xuyn mng. Nng lng ca gradient ha in ny c dng tng hp ATP t ADP v phosphat v c. Nhng phc hp protein c bit, c gi l ATP synthetaz, c gn vo bn trong ca mng trong ca ty th, tng t nh trn mng thylakoid. Nhng phc hp ny va l knh ion H+ cho ion H+ tr vo ngn trong va l mt enzim xc tc s tng hp ATP. V ion H+i t ni c nng cao n ni c nng thp qua ATP synthetaz nn nng lng c gii phng v ATP c tng hp. ATP c tng hp trong cht c bn s c protein ti qua mng ty th vo trong dch t bo cht.

Hnh 6. C ch ca qu trnh h hp Cc nghin cu cho thy t nht phi c 2 ion H+ i qua phc hp ATP synthetaz mi to ra c mt phn t ATP t ADP. V mi cp in t t NADH c dn truyn qua chui in t bm c khong 6 ion H+ i ra ngn ngoi (Hnh6). S oxy ha mt phn t NADH cho ra khong 3 phn t ATP. Tt c 8 NADH c sinh ra trong ty th (2 t s oxy ha acid pyruvic v 6 t chu trnh Krebs), cho ra tng cng 24 phn t ATP mi. Tuy nhin, 2 in t t FADH2 c t nng lng hn NADH; nhng in t ny i vo chui in t phc hp II (Hnh6), nn s ion H+ c bm ra t hn; do s oxy ha hp cht ny ch cho 2 phn t ATP (tng cng c 4 phn t ATP cho 2 phn t FADH2). 2 phn t NADH c to ra trong qu trnh ng phn trong dch t bo cht v c chuyn vo bn trong ty th nh mt phn t protein vn chuyn, nhng nng lng b mt i trong qu trnh ny. Kt qu l in t c i vo chui h hp cng v tr vi ni m in t ca FADH2 i vo (Hnh 6); do cng ch c 2 phn t ATP c to ra cho mi phn t NADH trong t bo cht (tng cng c 4 phn t ATP cho 2 phn t NADH). Tm li, s oxy ha hon ton mt phn t glucoz sinh ra 12 phn t cht kh. Khi cc i in t ca chng i vo chui h hp, 8 phn t sinh ra ATP nhiu hn, 3ATP cho mi phn t NADH; 4 phn t cn li ch to c 2 phn t ATP cho mi phn t FADH2 (Hnh 5 v 6). Tng cng c 32 phn t

ATP c to ra t s phosphoryl oxy ha ca chui dn truyn in t. Nh vy, s oxy ha hon ton mt phn t glucoz cho ra CO2, nc v 36 phn t ATP mi.

Hnh 7. Tng ATP c to ra do s h hp mt phn t glucoz Nu tnh theo t l %, th c khong 39% nng lng t do ban u trong phn t glucoz c gi li, 61% c gii phng ch yu di dng nhit. Ch c 2 trong s 36 phn t ATP c to ra trong s bin dng ym kh (t hn 6%), 34 phn t cn li c to ra do s h hp hiu kh (hn 94%). iu ny cho thy ti sao oxy thit yu cho con ngi v hu ht cc sinh vt khc. D vy, trong c th con ngi vn c cc qu trnh h hp ym kh xy ra nht l trong lc hot ng qu nhiu bng th lc; lc ny c cn rt nhiu nng lng trong khi O2 cung cp khng . Kt qu l s ln men to ra acid lactic lm mi c, nhng sau oxy c cung cp bnh thng tr

li th acid lactic s c chuyn v gan v s c bin dng tr li thnh glucoz ti y. Trong 34 phn t ATP c tng hp trong iu kin hiu kh c 32 phn t l c tng hp t gradient ion H+ sinh ra bi s dn truyn in t. Cyanid v mt s cc hp cht c khc c th lm nh hng trn cc protein ca chui, do c th lm ngng s dn truyn in t, hu qu l cc phn ng ca s h hp hiu kh khng th xy ra c, t bo s cht do khng c cung cp nng lng.

Hnh 8. So snh s tng hp ATP trong ty th v lc lp Qu trnh ha thm thu trong ty th v lc lp tng t nhau. Tuy nhin, c mt vi im khc bit: Ngun nng lng cho dng ti in t: lc lp nh sng mt tri cung cp nng lng, ty th nng lng ly t thc n (Hnh 8). Hng bm ion H+: trong thylakoid, ion H+ c bm vo bn trong thylakoid, ty th ion H+c bm ra khi mng trong, do nng ion H+ bn trong thylakoid cao hn bn ngoi, trong khi ty th nng ion H+ ngn ngoi cao hn ngn trong (Hnh 9).

lc lp in t t nc qua chui dn truyn in t n cht ti l NADP v O2 c to ra; trong khi ty th in t t cht ti NADH v FADH2 qua chui dn truyn n O2 v to thnh nc.

Hnh 9. So snh s ha thm thu ty th v lc lp

III. S H HP LIPID V PROTEIN 1. S h hp lipid TOP

S bin dng ca lipid bt u bng s thy gii chng thnh glycerol v nhng acid bo. Sau , glycerol (mt hp cht 3C) c bin i thnh PGAL v c a vo l trnh ng phn. Acid bo c chuyn vo ngn trong ca ty th, y chng c ct ra thnh acetyl-CoA v c a vo chu trnh Krebs. V lipid c thnh phn ca hydro cao hn carbohydrat, nn s oxy ha

hon ton cho nng lng cao hn tnh trn n v trng lng, mt gram cht bo thng cho nng lng cao hn hai ln so vi mt gram carbohydrat.

2. S h hp protein
Trc tin, protein c thy gii thnh acid amin v sau c bin dng theo nhiu cch khc nhau. Sau khi gc amino c tch ra, mt s acid amin c bin i thnh acid pyruvic, mt s khc thnh acetyl-CoA v c th thnh dng ny hay dng khc ca cc hp cht trong chu trnh Krebs. S oxy ha hon ton mt gram protein cho nng lng tng ng mt gram carbohydrat. Nhng hp cht nh acid pyruvic, acetyl-CoA v cc hp cht trong chu trnh Krebs c th c thoi dng theo nhiu cch khc nhau. Khng nhng chng l nhng cht giu nng lng c oxy ha thnh CO2 v H2O m chng cn c vai tr trong tin dng tng hp thnh acid amin, ng v cht bo. ly nng lng cn thit, cc cht c th thay i hng bin dng trong cc l trnh. Th d, PGAL v acetyl-CoA trong s oxy ha carbohydrat c th c dng tng hp cht bo; tng t nhiu acid amin c th bin i thnh carbohydrat qua mt s phn ng trung gian trong cc qu trnh bin dng ca chng (Hnh 10). Tuy nhin, khng phi tt c u c th i theo hai chiu. t bo ng vt ng c th bin i thnh cht bo, nhng acid bo khng th bin i thnh ng, bi v phn ng t acid pyruvic thnh acetyl-CoA khng th o ngc c. iu ny c lin quan n ch n ung ca con ngi. V glucoz l ngun nng lng duy nht cho t bo no, do thc n phi cung cp glucoz. Nu mc glucoz trong mu xung qu thp, th protein nh protein ca c phi c s dng oxy ha v sau mt s acid amin c th c bin i thnh glucoz cung cp cho t bo no. Do , ch n nhiu cht bo, t carbohydrat c th gy nguy him cho c th con ngi.

Hnh 10. Mi lin quan gia s bin dng cht ng, cht bo v protein