Chng IV

NNG LNG V

s TRAO l CHT

KHI NIM CHNG

4.1. nh lut nhit ng hc cho bit nng lng bin i nh th no
Dng nng lng trong sinh vt: th nng c trong cc in t ca nguyn t

v nh vy c th c truyn t nguyn t ny n nguyn t khc.

Cc nh lut ca nhit ng hc. Nng lng khng bao gi b m t i nhng

khi n c dn truyn, ngy cng nhiu nng lng b tiu tn di dng

nhit.
Nng lng t do; trong mt phn ng ho hc, nng lng c gii phng ra

hoc c cung cp l hiu trong nng lng ca lin kt ho hc gia cht

phn ng v sn phm, c hiu chnh cho s hn n.
Nng lng hot ho: b t u mt phn ng ho hc, cn mt lng nh

nng lng ph v tnh n nh ca cc lin kt ho hc vn c.

4.2. Emym l cht xc tc sinh hc
Enzym. Protein hnh cu gi l enzyra c6 chc nng xc tc cc phn ng ho

hc trong t bo.
Hot ng ca enzym: enzym c to hnh khp hon ton vi c cht,

thc y cc nhm phn ng ho hc vo gn phn ng xy ra nhanh

chng.
Cc nhn t nh hng ln hot tnh enzym: enzym hot ng hiu qu nht

nhit v pH ti u.
Nhm ngoi ca enzym: ion kim loi v cc cht khc thng c tc dng h

tr enzym thc hin s xc tc.

Enzym cn nhiu dng. Mt s enzym kt hp theo cc nhm phc hp, cht

xc tc khc thm ch khng phi protein.

117

4.3. ATP l tin t nng lng ca s sng

ATP l g? T bo d tr v gii phng nng lng t

ca ATP - tin t nng lng ca t bo.

cc lin

kt photphat

trao i

cht (chuyn

4.4. Trao i cht l i sng ho hc ca t ho

Cc con ng sinh ho. n v t chc ca qu trnh

ho vt cht). Con ng sinh ho l n v t chc ca qu trnh trao i

cht.
S tin ho ca qu trn h trao i cht. Cc qu trnh chuyn ho ch yu
tin ho trn mt chu k di, da vo gi thuyt ci g xut hin trc.

C th coi s sng nh mt dng nng lng lin tc c sinh vt chuyn ti

sinh cng hu ch cho hot ng sng. Mi c im quan trng m nh
chng ta nh ngha s sng - l tnh tr t t, s sinh trng, sinh sn, trng
thi phn ng v s iu chnh ni sinh - u i hi s cung cp nng lng

thng xuyn. Nu ngun nng lng b mt i th s sng s dng li. Do , s

khng th c mt phn tch ton din v s sng nu khng bn lun v vn
nng lng sinh hc {bioenergetics) - s phn tch v nng lng c cung cp nh

th no cho hot ng ca cc h thng shg. Trong chng ny, chng ta s tp

trung v nng lng m sinh vt hp th, d tr v s dng nh th no trong i

sng ca mnh.

4.1. Cc nh ut nhit ng hc ni r nng lng bin i nh

th no
4.1.1. Dng nng lng trong sinh vt
Nng lng c nh ngha nh kh nng sinh cng. C th coi nng lng
tn ti di hai dng: ng nng v th nng. ng nng (kinetic energy) l nng

lng ca s vn ng. Vt ang chuyn ng thc hin cng nh lm cho vt khc

chuyn ng. Th nng (potential energy) l nng lng d tr. Cc vt khng vn

ng ch ng nhng c kh nng thc hin vn ng v chng c th nng. Mt

tng c t trn nh ni c th nng, khi n bt u ln xuhg ni th mt

phn th nng chuyn ho thnh ng nng. Nhiu hot ng m c th sng thc

hin l qu trnh bin i th nng thnh ng nng.
Nng lng c th c nhiu dng; c nng, nhit nng, m nng, in nng,
nh sng hoc tia phng x. Do nng lng c th tn ti trong nhiu dng nh vy
nn c nhiu phng php o nng lng. Thun tin nht l cch o theo nhit
lng bi v tt c cc dng nng lng khc c th bin i thnh nhit nng. Thc
th, nghin cu v nng lng c gi l nhit ng hc {thermo dynamics) l gii
thch v cc bin i ca nhit nng. n v ca nhit c s dng ph bin nht
trong sinh hc l kilocalo {kilocalorie = kcal). 1 kilocalo bng 1000 calo (cal) v 1
118

calo l lng nhit cn nng nhit ca mt gam nc ln mt bch phn

(C). (iu quan trng l khng nhm ln calo vi thut ng c lin quan vi khu
phn v s dinh dng cng l Calo vi ch vit hoa C). n v nng lng khc
thng c dng trong vt l l jun oule). Mt jun bng 0,239 calo.

S oxi ho kh: Dng nng lng trong c th sng

T m t tri, nng lng chy vo gii sinh vt. Mt tri chiu nh sng lin
tc vo tri t. Ngi ta tnh rng mt tri cung cp cho tri t vi hn
13.10^^ calo/nm hoc (40 triu t calo/giy). Thc vt, to v mt s loi vi khun
nht nh hp th mt phn nng lng ny thng qua qu trnh quang hp. Trong
quang hp, nng lng thu c t nh sng mt tri c s dng kt hp cc
phn t nh (nc v kh cacbonic) thnh phn t phc tp hn (ng). Nng
lng c d tr di dng th nng trong cc lin kt cng ho tr gia cc
nguyn t trong phn t ng. Ai cng bit mt nguyn t gm mt nhn trung
tm c bao quanh bng mt hoc nhiu in t qu o v mt lin kt cng ho
tr to nn khi hai nhn nguyn t gp chung in t. Vic ph v mt lin kt nh
vy cn nng lng ph v h t nhn. Thc vy, lc ca mt lin kt cng ho tr
o c l tng ng nng lng cn ph v n. Th d, cn 98,8 kca ph
vd mt phn t gam (6,023.10^) ca lin lt cacbon - hidro (C-H).
Trong mt phn ng ho hc, nng lng d tr trong cc lin kt ho hc c
th chuyn n cc lin kt mi. Trong mt s phn ng ny, in t thc s
chuyn t mt nguyn t hoc phn t n nguyn t, phn t khc. Khi mt
nguyn t hoc phn t m t in t th n b oxi ho v qu trnh ny c gi l
s oxi ho (oxidation). T hut ng ny phn nh s tht cho rng trong cc h sinh
hc th oxi c tc dng h t in t mnh l cht nhn in t ph bin nht. Ngc
li, khi mt nguyn t hoc phn t nhn in t th n b kh v qu trnh gi l
s kh {reduction). S oxi ho v kh lun lun xy ra vi nhau, bi v mi in t
b m t mt nguyn t thng qua s oxi ho th c mt s nguyn t khc thu
nhn thng qua s kh. Do , phn ng ho hc thuc loi ny c gi l phn
ng oxi ho - kh (oxidation reduction reactions: redox reactions) (Hnh 4.1). Nng
lng c chuyn t mt phn t n phn t khc thng qua phn ng oxi ho
kh. Do , dng kh ca mt phn t c mt mc nng lng cao hn 8 0 vi dng
oxi ho.
c
S m ftdint (s _o x i ho)--------

Hnh 4.1. Phn ng oxi ho

kh, 0x1 ho l w t t air l, w
kh l s nhn in t. Trong th
d ny, ln tch ca phn t A v
B c biu th bng vng trn
nh ph(a trn bn phi ca mi
phn t. Phn t mt nng

------ ^

S nhn din t (s kh)

ing khi n mt mi in t.

GDNng lng thp

Nng i ^ g cao

khi phn t B nhn nng

'^"9
119

Phn ng oxi ho kh ng vai tr ch yu trong dng nng lng thng qua

h thng sinh vt, do in t c chuyn t nguyn t ny n nguyn t kia c
mang theo nng lng. Lng nng lng m mt in t c6 c ph thuc vo
khong cch ca n ti h t nhn v h t nhn h t n vi lc m nh nh th no.
nh sng (v cc dng nng lng khc) c th nhng nng lng cho mt in t
v nng n n mt mc nng lng cao hn. Khi in t ny ri khi mt nguyn
t (s oxi ho) v chuyn n nguyn t khc (s kh), nng lng c gia tng
ca in t c chuyn theo n v in t chuyn ng trn qu o ca nhn
in t th 2 mc nng lng cao hdn. Nng lng gia tng c d tr nh l
th nng ho hc v v sau ngruyn t c th gii phng khi in t quay li mc
nng lng ban u ca n.

Nng lng l kh nng sinh cng hoc hot ng (ng nng)

hoc c d tr s dng v sau (th nng). Nng lng thng
dc truyn cng vi din t. Oxi ho s mt din t, s kh l s
nhn din t.
4.1.2. Cc nh lut ca nhit ng hc
Tt c cc hot ng trong v tr t s sng v ci cht ca t bo, s chy,
suy ngh, ca h t v mi hot ng ca con ngi... cho n cc v n bom nguyn
t, mt phn ng trong nh my in h t nhn, s sng v ci cht ca cc ngi
sao... t t c u tri qua cc bin i nng lng v chu s chi phi, iu khin ca
hai nh lut nng lng - cc nh lu t nhit ng hc.

4.1.2.1, nh lu t th n h t n h it ng hc
Theo nh lut th n h t ca nhit ng hc, cng c gi nh lut bo
ton nng ng {the law of cpnservation ofenergy), nng lng ca v tr l khng
i. Cng nh vy, tng nng lng ca bt k h thng no, ngha ca bt k
vt no v mi trng xung quanh n gi khng i. T mi trng xung quanh
mun ni ti phn cn li ca v tr. Khi mt vt tri qua mt bin i th n c
th nhn nng lng t mi trng xung quanh (vt xung quanh) hoc phn tn
nng ng vo mi trng xung quanh. S khc nhau v hm lng nng lng
ca vt trong giai on ban u v cu cng phi bng nhau do mt bin i tng
ng trong hm lng nng lng ca mi tnlng xung quanh.
nh lut th n h t v nhit ng hc cho rng trong cc qu trn h ho hc
hoc vt l bnh thng, nng lng c th c chuyn i v dng th d t th
nng thnh ng nng nhng khng bao gi c th b m t i, cng khng th to ra
nng lng mi. Tng nng lng trong v tr gi khng i bi v v tr l mt h
thng kn khi n trao i nng lng. Nh chng ta bit, nng lng c hnh
thnh khong 20 t nm trc, nhng n khng th c cng thm hoc b tr i.

120

Mc d sinh vt khng th to ra cng khng th ph hu nng lng nhng

n c th hp th u nng lng t mi trng v dng nng lng theo yu cu
ring ca n. Sinh vt cng c th bin nng lng t dng ny thnh dng khc.
Th d, trong qu trnh quang hp, t bo thc vt bin nng lng nh sng thnh
in nng v v sau thnh ho nng c d tr trong cc lin kt ho hc ca cc
cht hu c nh hidratcacbon, protein, lipit... trong cc c quan ca thc vt. Mt
s ng vt n thc vt li bin ho nng ny thnh c nng ca s co c hoc mt
s dng cn thit khc. Khi cc qu trnh bin i ang lng ny xy ra th mt s
nng lng b bin thnh nhit nng v b tiu tn vo mi trng v sinh vt
khng bao gi c th s dng tr li nng Ing . V nh vy, dng nng Ing
lin tc thng qua gii sinh vt theo mt chiu vi nng lng mi t m t tri
thng xuyn i vo h thng thay th cho nng lng b tiu tn di dng
nhit ieat). N hit l sT o v s chuyn ng hn n ca phn t (v do l s
o ca mt dng ng nng). R rng, nng lng thc s khng b "mt i", n vn
tn ti trong mi trng vt l bao quanh.
C th li dng nhit sinh cng ch khi c mt gradient nhit, ngha l c
mt hiu nhit gia hai vng (nh hot ng ca ng c hi nc). T bo l qu
b duy tr hiu nhit cao bn trong, do nhit nng khng c kh nng
sinh cng ca t bo. Mc d, tng nng lng trong v tr gi khng i, nhng
nng lng sn sng sinh cng gim xung, trong khi ngy cng nhiu nng
lng tiu tn di dng nhit. Qua nhng g trnh by trn, nh lut th
nht gii thch ti sao sinh vt khng th to nng lng m phi vay mn n lin
tc t mt ni no . Bi v mi t bo, mi c th sinh vt hay ton b th gii
sng u l h thng h nhng tii ti trong v tr - mt h thng kn vi tng
nng lng lun khng i.

4.I.2.2. nh lu t th hai nhit ng he

nh lut th hai nhit ng hc ni v s bin i ca th nng thnh nhit
hay s chuyn ng hn n ca phn t. N khng nh rng tnh hn n (c
o bng entropy) trong v tr lin tc tng ln. C th nh ngha entropy nh mt
trng thi hn n ca nng lng m khng dng c sinh cng. nh lut
th hai khng nh cc qu trnh vt l v ho hc tin hnh theo cch m entropy
ca h thng tng ln. Trong hu nh t t c cc qu trnh bin i nng lng th
c s m t mt s nng lng di dng nhit vo mi trng. Do , nng lng b
m t khng cn sinh cng c na.
Thc ra, nhit l nng lng ca s vn ng hn n ca cc phn t v l
dng nng lng v t chc nht. Nhit c th dng sinh cng ch khi xut hin
mt gradient nhit (mt hiu nhit: temperature difference) lm cho n chuyn
t vng m hn n vng lnh hn. Trong mt t bo sng th nhit ti rtii
im l nh nhau, do vy nhit khng th c dng sinh cng sinh hc.
121

iu quan trng ta hiu c rng nh lut th hai ca nhit ng hc ph

hp vi nh lut th nht. Tng nng lng trong v tr khng gim theo thi
gian, nhng nng lng sn c sinh cng b suy thoi th n h chuyn ng phn
t hn n.
nh lut th hai v nhit ng hc cn cho rng nng lng khng th bin
i t dng ny thnh dng khc m khng c s m t nng lng c ch v khng
c6 qu trnh no yu cu nng lng lun lun t hiu qu 100%. Thc th, s s
dng nng lng ca t bo t hiu qu khong 55% cn 45% nng lng b m t
i di dng nhit. Cc qu trn h sinh hc r t hiu qu nu 8 0 snh vi phn ln
my do con ngi s dng, th d nh ng c dng xng ch t hiu qu t 17%
n 25%, ho nng ca xng bin thnh s chuyn ng ca xe v phn ho nng
cn li b m t i di dng nhit. T t nhin, nhit l mt dng ca nng lng,
nhng nhit l dng nng lng hn n nht v nhanh chng tiu tn vo mi
trng. ng vt, s co c bin ho nng trong ATP th n h c nng, mt s nng
lng ny ngay lp tc bin thnh nhit. Vi s bin i nng lng lin tip nh
vy, cui cng t t c dng nng lng c ch u bin thnh nhit tiu tn vo mi
trng v khng th bin i tr li thnh mt dng ca th nng.
V c th sng c t chc cao, chng r t khng n nh. Thc th, s sng l
mt cuc u tran h khng ngng chng li nh lut th hai ca nhit ng hc. S
sng cn ca c th sinh vt cng nh ca cc h sinh thi ph thuc vo s b
sung nng lng lin tc. Do , cc sinh vt sn xut phi tin hnh qu trnh
quang hp, cn sinh vt tiu th v sinh vt phn hu phi n thc vt v n ln
nhau mi tn ti c.

4.I.2.3. Entropy
Entropy l thc o v tnh hn n ca h thng, nh vy nh lut th hai
ca nhit ng hc c th ph t biu n gin nh l "s tng entropy". Dng nng
lng hu ch, c t chc th c entropy thp, cn dng nng lng v t chc, t n
nh hn nh nhit th c entropy cao. Mt phng ngn np, gn gng c entropy
thp hn nhiu so vi phng hn n, ba bi. Chng ta bit rng phng ngn np
lun lun c khuynh hng hng n tnh hn n. Theo cch , cui cng cc
qu trnh bin nng ng u sinh ra nhit v do entropy ca v tr lun
lun tng ln. Ni chung, cc qu trn h bin i nng lng tin hnh mt cch t
pht chuyn ho vt cht t dng c tr t t nhng t n nh thnh dng t tr t
t hn nhng n nh hn.
Mt x hi vn m inh nh chng ta ang s dng nhiu nng lng c entropy
thp. Nng lng ca nhin liu ho thch nh than v du m c s dng
pht trin v x l lng thc m chng ta cn v c s dng duy tr mi
trng c chng ta c t chc. Vic xy dng cc to cao c, trng hc v ng
s u i hi vic chi tiu nng lng c ch'm cui cng bin th n h nhit. X hi
122

vn minh ca chng ta ngy nay ang ngy cng lm tng entropy ca v tr

mt tc cao hn nhiu so vi bt k x hi no qua.
nh lut th n h t ca nhit dng hc gii thich rng nng lng
khng th c to ra hoc khng th b ph hu. N ch c th tri
qua s bin i t dng ny thnh dng khc. nh lut th hai ca
nhit ng hc cho rng tnh hn n (entropy) trong v tr ang
tng ln. S sng bin nng lng t mi tri thnh cc dng nng
lng khc h tr cc qu trnh sng. Nng lng khng bao gi
b m t i nhng khi n c s dng th ngy cng nhiu nng
lng b bin th n h nhit - nng lng ca s chuyn ng phn t
hn n.
Nh vy, cc nh lu t nhit ng gii thch ti sao entropy ca v tr tng
ln mt cch t pht v ti sao cc sinh vt cn s cung cp thng xuyn nng
lng c ch duy tr t chc ca chng.

4.1.3. Nng lng t do

Lc pht ng t t c cc qu trnh trong h thng sng v khng sng l
khuynh hng ca h thng t iu kin ca entropy cc i. Nng lng
dng nhit c gii phng hoc c vt hp th lm cho h thng t trng thi
entropy cc i. Tng nhit hm {total heat content) hay entanpy H ca mt h
thng l tng th nng ca n. Trong mt phn ng ho hc, entanpy ca cc cht
phn ng hay cc sn phm bng vi tng nng lng ca lin kt ho hc. Khi lin
kt c hnh thnh hay b ph v, entanpy c hp th hoc c phng thch.
Entropy v entanpy c quan h theo phng din th ba ca nng lng gi l
nng lng t do (free energy). Chng ta c th coi nng lng t do nh l thnh
phn ca tng nng lng ca mt h thng sn c sinh cng di iu kin
nhit v p sut khng i. Do , nng lng t o l kha cnh nhit ng hc
ng ch nh t trong sinh hc.
Entropy v nng lng t do c quan h theo t l nghch. Hai i lng c
quan h theo phng trn h sau:
AG = AH - TAS
Trong :
S: entropy
G: nnglng t do
H: entanpy ca h thng
T: nhit tuyt theo Kelvin
A: ngha "bin i"
T t c cc qu trn h vt l v ho hc u tin hnh theo s gim nng lng
t do cho n khi chng t trng thi cn bng, nng lng t do ca h
123

thng l cc tiu v entropy l cc i. Nng lng t do l nng lng c ch trong

cc h sinh hc, entropy l trn g thi nng lng b phn r, v ch.
S bin i nng lng t do (hay G) l tnh cht c bn ca cc phn ng
ho hc. Trong mt s phn ng, AG l dng. iu c ngha l cc sn phm
ca phn ng cha nhiu nng lng t do hn so vi cht phn ng, nng lng
lin kt (H) cao hn hay tnh hn n (S) trong h thng thp hn. Cc phn ng
kiu n khng t ph t tin hnh. Chng i hi s cung cp nng ng v do
l phn ng thu nhit {endergonic reaction = "invuard energy"). Trong cc phn ng
khc, G l m. Cc sn phm ca phn ng cha t nng ng t do hdn cht
phn ng, hoc nng ng lin kt thp hn hoc tnh hn n cao hn hoc c
hai. Phn ng nh vy c khuynh hng tin hnh t pht. Bt k phn ng ho
hc no s c khuynh hng tin hnh t pht nu hiu trong tnh hn n (TAS)
ln hn hiu trong nng lng lin kt gia cht phn ng v sn phm (AH). Cc
phn ng ny gii phng nng lng t do di ra di dng nhit v do gi l
phn ng th u nhit {exergonic reaction = "cutivard energy"). H nh 4.2 tm t t cc
phn ng ny.

oc

| |

I
z e
Thu nhit
(b)

(a)

Hlnh 4.2. Nng lng trong cc phn ng ho hc

(a) Trong phn ng thu nhit (thu nng lng), sn phm ca phn ng cha nhiu nng
lng hn so VI cht phn ng v nang lng d6i ra phi c b sung cho phn ng
tin hnh
(b) Trong phn ng pht nhit, sn phm cha t nng lng hn so vi cht phn ng
v nng lng di ra c gii phng

Nng lng t do l n n g lng sn c sinh cng. T rong cc p h n

ng h o hc c ti n h n h bn tro n g t bo, s bin di n n g lng
t do (AG) l h iu tro n g n n g ing lin k t gia c h t p h n ng v

124

sn phm (H), tr i bt k s bin i no trong mc ca tnh

hn n ca h thng (TS). Bt k phn ng no m sn phm ca
n cha t nng lng t do hn so vi cht phn ng (AG m) s c
khuynh hng tin hnh t pht.
4.1.4. N n g l n g h o t h o : s c h u n b p h n t h o t n g

Nu t t c cc phn ng ho hc gii phng nng lng t do u c khuynh

hng tin hnh mt cch t pht, vy ti sao tt c cc phn ng u khng
xy ra? L do m chng khng xy ra l do phn ln phn ng cn cung cp
nng lng khi ng phn ng. Trc khi phn ng to cc lin kt ho hc
mi, ngay c cc lin kt cha t nng lng th trc tin n cn ph v cc lin
kt vn c trong phn t cht phn ng nn n cn chi dng nng lng. Nng
lng ph tri (nng lng ph thm = extra energy) cn ph v cc lin kt ho
hc vn c v khi ng mt phn ng ho hc gi l nng lng hot ho
(activation energy) (Hnh 4.3a).

oc ^

I s
^

Nng
lng
hot ho

II
0c

li
s I;
i

Cht
phn tiig

\
%

zI

Snph&n
(a)
Hnh 4.3. Nng lng hot ho v s xc tc
(a) Phn ng pht nhit khng cn tin hnh nhanh chng do nng lng phi c
cung cp ph v (lm mt n nh) cc lin kt ho hc vn c. Nang lng ph thm
ny l nng lng hot ho cho phn ng
(b) Cht xc tc lm tng tc cc phn ng ring nh lm gim lng nng lng hot
ho cn khi ng phn ng

Tc ca phn ng pht nhit ph thuc vo nng lng hot ho cn

phn ng bt u. Cc phn ng vi nng lng hot ho ln hn c khusmh hng
tin hnh chm hn do c t cc phn t thnh cng trong vic vt qua hng ro
nng lng ban u. Song, nng lng hot ho khng phi l hng s khng i.

125

Gy mt tc ng no ln cc lin kt ho hc ring c th lm cho chng d b

ph vd hn. Qu trin h tc ng ln cc lin kt ho hc theo cch lm gim nng
lng hot ho cn khi ng mt phn ng gi l s xc tc (catalysis) v cht
thc hin qu trn h ny c gi l cht xc tc (catalsts) (Hnh 4.3b).
C ht xc tc khng th vi phm nh lut c bn ca nhit ng hc. Th d,
chng khng th lm cho m t phn ng thu nhit tin h n h m t cch t pht. Nh
lm gim nng ng hot ho, cht xc tc lm tng tc c phn ng thun v
nghch chnh xc theo cng mc v t , n khng lm th ay i t l cui cng
ca cht phn ng bin th n h sn phm.
nm c iu ny, ta hnh dung c6 mt qu bng g nm trong mt h
nng trn mt pha dy ni. Ch mt vnh t hp di qu bng c tc dng
chn bng ln xung ni. By gi ta b vnh t hp y i th qu bng s bt u
ln xung; nhng vic oi b t pha di qu bng s khng bao gi lm cho
qu bng n ngc n ni. Vic loi b mp t n gin ch cho php qu bng
vn ng t do v trng lc xc nh chiu hng n ln xung chn ni nh th
no. Lm gim lc cn i vi 8 vn ng ca qu bng s c6 tc dng kch thch
s vn ng c hng nh v tr ca bng trn qu ni.
Nh vy, chiu hng mt phn ng ho hc tin h n h ch c xc nh
nh hiu trong nng lng t do. Ging nh vic b vnh t nm di qu bng g
trn ni, cht xc tc lm gim hng ro nng lng ngn cn p h n ng khng tin
hnh. C ht xc tc khng to b t k u tin thun i no cho phn ng thu nhit
hn l vic o b vnh t di qu bng g gi nh n ln ngc ln ni. Ch
phn ng pht nhit tin hn h t pht v cht xc tc khng th lm thay i quy
lut . iu m cht xc tc c th lm l khin cho phn ng tin hn h nhanh
hn nhiu m thi.

TC ca ph&n ng ph thuc vo n&ng lng hot ho cn khi

ng n. C h t xc t c lm gim n&ng lng h o t h o v n h vy lm
t n g tc p h n ng, m c d ch n g khng lm th a y di t l cu i
c n g c a c h t p h n ng v sn phm p h n ng.

4.2. Enzym cht xc tc sinh hc

4.2.1. Enzym: "ngi th cn mn" ca t bo
Cc phn ng ho hc bn trong c th sng c iu ho nh s iu tit cc
v tr xy ra 8 xc tc. Do , bn thn s sng c iu chnh bng cc cht xc
tc. Cc tc nhn thc hin phn ln s xc tc trong c th sng l cc protein
mang tn enzym mymes). (Nhng enzym protein khng phi l cht xc tc duy
nht trong c th sng. Nm 1989, Thomas Cech v Sidney A ltm an ng nhn gii
Nobel v ho hc cho cng trn h chng minh ARN ng vai tr nh mt enzym m
hai ng gi l ribozim. Do , khng phi mi enzym u l protein). Dng cu trc

126

khng gian ba chiu c bit ca enzym khin cho n c vai tr lm n nh kt hp

tm thi gia cc phn t c cht (phn t s tri qua phn ng). Nh m ang hai
phn t c cht cng vi nhau theo chiu hng chnh xc, hoc nh gy tc ng
ln cc lin kt ho hc ring ca c cht, nn enzym lm gim nng lng hot
ho cn to cc lin kt mi. Do , phn ng tin hnh nhanh chng hn nhiu
so vi trng hp khng c enzym xc tc. Do bn thn enzym khng b bin i
hoc b tiu th trong qu trn h phn ng nn cn cn mt lng nh enzym l
v n c th c s dng lp i lp li.
gii thch enzym hot ng nh th no, ta hy xem phn ng ca kh
cacbonic v nc to axit cacbonic. Phn ng c enzym xctc quan trng ny
xy ra trong t bo hng cu ca ng vt c xng sng:
CO2

kh cacbonic

H2O

nc

>

H2CO3

axit cacbonic

Phn ng ny c th tin hnh theo c hai chiu, nhng do n c nng lng

hot ho ln, phn ng tin hnh r t chm khi thiu enzym; trong mt gi ch c
khong 200 phn t axit cacbonic c hnh thnh trong t bo. Kiu phn ng
chm ny t c s dng trong t bo. T bo vt qua vn ny nh tuyn chn
mt enzym bn trong t bo cht gi l cacbonic anhidraza (carbonic anhydrase).
Di cng iu kin, nhng trong s c m t ca cacbonic anhidraza c n khong
100.000 phn t axit cacbonic hnh thnh mi giy. Do , enzym lm tng tc
phn ng hn 10 triu ln.
bit hng nghn loi enzym khc nhau v mi loi xc tc cho mt hoc mt
vi phn ng ho hc c hiu. Nh xc tc cc phn ng ho hc ring nn enzym
trong t bo quyt nh tin trn h trao i cht - tp hp ton b phn ng ho hc
trong t bo . Cc loi t bo khc nhau cha cc tp hp enzym khc nhau v s
sai khc ny dn n cc bin i v cu trc v chc nng gia cc loi t bo.
Phn ng ho hc xy ra trong t bo hng cu khc vi phn ng xy ra trong t
bo th n kinh, mt phn do t bo cht v mng ca t bo hng cu, t bo thn
kinh cha cc h enzym khc nhau.

T bo s dng cc protein gi l enzym nh l cht xc tc im

gim nng lng hot ho.

4.2.2. Enz3rm hot ng nh th no?

Phn ln enzym l protein hnh cu vi mt hoc nhiu ti hoc khe trn b
m t gi l v tr hot ng (active sites) (Hnh 4.4). C cht lin kt vi enzym v
tr hot ng ny, to phc h enzym - c cht {ernym - substrate complex). cho
s xc tc xy ra bn trong phc h, phn t c cht phi khp chnh xc vo v tr
hot ng.'* ng thi cc nhm bn axit amin ca enzym tip xc trc tip vi cc
in kt ca c cht. Cc nhm bn ny tng tc ho hc vi c cht, thng gy
127

tc ng hoc lm mo m lin k ring no v do lm gim nng lng hot

ho cn ph v lin kt. Lc ny c cht bin th n h sn phm v tip tch
khi enzym.
Hinh 4.4. Enzym phn gii (lysozyme) hot
ng nh th no?
(a) Rnh xuyn qua enzym phn gii khp
vi dng polisaccart (mt chui ng) to
vch t bo vi khun
(b) Khi chui ng di chuyn n rnh
v xm nhp vo rnh lm cho protein enzym
bin i hnh dng v m ly phn t c cht
mt cch cht ch hn. V tr khp cm ng
ny xy ra gc axit glutamic trong protein
bn cnh lin kt gia hai ng ln cn v
axit glutamic 'cp or mt in t khi lin
kt lm phn gii phn t c cht

( D C cht. saccaroz. gm glucoz

----- v fructoz lin kt V(M nhau

Fructoz

(2)ccht lin kt \ i enzym.

to pht h enzym - c cht

Lin kt

V tr
hot ng

Enzym

( 4) Sn phm c
gii phng. enzym
t o lin kt v
cc c cht khc

@ S lin kt
c cht v enzym ^
gy tc ng ln
lin kt gluc02-fruct
v lin kt b ph v

Hlnh 4.5. Chu trinh xc tc ca enzym. Enzym lm tng vn tc nh xy ra phn ng

ho hc. Trong phn ng c minh ho y, enzym sucraza tch ng saccaroz (c
trong phn ln bnh ngt) thnh hai ng an gin hn; glucoz v fructoz.
(1) u tin, c cht saccaroz Hn kt vo v tr hot ng ca enzym, khp vo khe im
trong b mt enzym
(2) S l^n kt a saccaroz vi v bf hot ng to nn phc h enzym-c cht, gy tc ng
cm ng ln phn t sucraza lm n ttiay i hnh dng, khp cht hn xung quanh saccaroz
(3) Cc gc axit amin trong v trf phn ng tip cn st vo lin kt gia thnh phn
glucoz v fructoz ca saccaroz, ph v lin kt
(4) Enzym gii phng cc mnh glucoz v fructoz (l sn phm ca phn ng), v sau
enzym sn sng lin kt vi phn t saccaroz khc v thng qua chu trnh xc tc mt ln
na. Chu trinh ny thng c tm tt bng phng trnh E + s - [ES] -* E + p, y E:
enzym, S: co cht, ES; phc h enzym - c cht v P: sn phm

128

Protein khng phi l dng phn t cng n h t m sng ng. S lin kt ca

c cht lm enzym cm ng iu chnh hnh dng no , dn n mt khp cm
ng (induced fit) tt hn gia enzym v c cht (Hnh 4.5). S tng tc ny cng
c th thc y s lin kt ca cc c cht khc v trong cc trng hp , bn
th n c cht "hot ho" enzym nhn cc c cht khc.

Enzym thng ch xc tc cho mt hoc mt vi phn ng ho hc

tng t do chng dc hiu trong cch chn la c cht. Tinh c
hiu ny l do v tri hot ng ca enzym m to hnh sa cho ch
mt phn t c cht nht nh s lp khp vo n.
4.2.3. Cc nhn t nh hng n hot ng ca enzym
Bt k nhn t ho hc hay vt l no lm bin i hnh dng ba chiu ca
en^ym nh nhit , pH, nng mui v s lin kt ca cc phn t iu chnh
c hiu u c th n h hdng ln kh nng xc tc phn ng ca enzyrn.

4.2.3.I. N hit
Khi khng c xc tc, nhit tng
s lm tc phn ng tng do s vn
ng hn n ca phn t tng ln. Tc
ca phn ng c enzym xc tc cng
tn g ln khi tng nhit , nhng ch
ln n mt im gi l nhit ti u
{temperature optimum) (Hnh 4.6a).
Di nhit ny, cc lin kt hidro v
cc tng tc k nc xc nh hnh
dng ca enzym l khng linh ng
cho php xy ra khp cm ng m
lm cho s xc tc t ti u. Trn
nhit t'i u th cc lc ny l qu
vii duy tr hnh dng ca enzym
chng li s vn ng hn n tng ln
c.a cc nguyn t trong enzym. cc
nhit cao hn ny, enzym bin tnh.
Phn ln enzym ca ngi c nhit
t'ii u t 35C n 40c - nhit bnh
thng ca c th cng nm trong
khong nhit ny. Vi khun sng
sui nc nng c enzym n nh hn
v nhit ti u cho cc enzym ny c
th ln n 70c hay cao hn.

Nhit tl u
ca enzym ngi

Nhit ti u ca
enzym vi khun
sud ac nog

Nhit ca phn hg (C)

(a)

O
i
a

pH tl u

pH t u

cho pepsin

cho ypsin

(b)

pH ca phn bg

Hlnh 4.6. Enzym mn cm vi mi trng

sng. Hot ng ca enzym chu tc ng bi
c (a) nhit v (b) pH. Phn ln enzym ca
ngi nh enzytn phn gii protein trpsin
hot ng tt nht nhit khong
trong phm vl pH t 6 - 8

40c

129

4.2.3.2, p H
Tng tc ion gia cc gc axit amin tch in tri du nh axit glutamic (-) v
lizin (+) cng c tc dng gi enzym li vi nhau. Cc tng tc ny nhy cm vi
nng ion hidro ca dch m enzym ho tan do n lm bin i nng lm
chuyn dch cn bng gia cc gc axit amin tch in dng v in m. V l do
, nn phn ln enzym c pH ti u (pH optimum) thng nm trong khong pH
6-8. Cc enzym no c th hot ng trong mi trng r t axit l protein c th duy
tr dng ba chiu thm ch khi c m t nng ion hidro cao. Th d, enzym pepsin
tiu ho protein trong d dy pH = 2, mi trng mang tnh axit (Hnh 4.6b).

4.2.3.3. Cht c c h v ch t hot ho

Hot ng ca enzym nhy cm vi s c m t ca cc cht c hiu c tc
dng lin kt vi enzym v lm bin i hnh dng ca n. Thng qua cc cht ny,
t bo c th iu chnh enzym no hot ng v enzym no khng hot ng mt
thi gian no . iu cho php t bo lm tng hiu qu hot ng v iu tit
cc bin i v c tnh trong qu trnh pht trin. Cht lin kt vi enzym v lm
gim (decrease) hot tnh enzym gi l cht c ch (inhibitor). Sn phm cui cng
ca con ng sinh ho hot ng nh mt cht c ch ca phn ng u tin trong
con ng - qu trnh ny c gi l s c ch lin h ngc ifeedback
inhibition).
S c ch enzym xy ra theo hai cch: cht c ch cnh tran h {competitive
inhihitor) th cnh tran h vi c cht cng v tr lin kt, lm i ch mt t l
phn t c cht khi enzym; trong khi , cht c ch khng cnh tran h
S tfc ch cnh anh

S tfc chkhdog cnh anh

Cht fc chcnh anh

gy nhiu lon v
hot ng cia enzym
khii cho c cht
khng th lin kt

(a)

Cht t ch df
lp th lm ia
hlnh dng cda
enzym khi n khng
(b) th lin kt vi c cht
Hlnh 4.7. Erym c6 th b c ch th no?

(a) Trong c ch cnh tranh, cht c ch gy nhiu lon v tri hot ng ca enzym
(b) Trong c ch khng cnh tranh, cht c ch lin kt vinzym mt v tr ngoi vj tr
hot ng, gy tc ng lm b^n i cu hnh trong enzym khin cho enzym khng th lin
kt vi c cht c na
130

(noncompetitive inhibitor) lin kt vi enzym mt v tr khc ngoi v tr hot

ng, c tc dng lm bin i hnh dng ca enzym v lm cho enzym khng th
lin kt vi c cht (Hnh 4.7). Phn ln cht c ch khng cnh tranh lin kt vi
mt phn c hiu ca enzym gi l v tr d lp th (allosteric site). Cc v tr ny
c chc nng nh ci chuyn mch on/off ho hc: s lin kt ca mt cht vo v
tr c th chuyn enzym gia cu dng hot ng v khng hot ng. Cht lin kt
vo v tr d lp th v lm gim hot tnh enzym gi l cht c ch d lp th
(allosteric inhibitor) (Hnh 4.7b). Cn cht hot ho lin kt vi v tr d lp th v
duy tr hot tnh enzym, nh lm tng hot tnh enzym.
Enzym c mt nhit d v pH ti u, d enzym hot ng hiu qu

nht. Cc cht c ch lm gim hot tnh enzym cn cht hot ho

m tng hot ng.
4.2.4. Cc nhm ngoi ca enzym
Hot ng ca enzym thng c h tr nh c thm cc thnh phn ho hc
gi l nhm ngoi (fiofactor). Th d, v tr hot ng ca nhiu enzym cha ion kim
loi c tc dng gip ht in t khi phn t c cht. Enzym cacboxipeptidaza
tiu ho protein nh s dng ion km (Zn*0 trong v tr hot ng loi b in t
khi cc lin kt ni cc axit amin. Cc nguyn t khc nh molipden v mangan
cng c s dng nh cc nhm ngoi. Ging nh km, cc cht ny cn vi mt
lng nh trong khu phn thc n. Khi nhm ngoi l mt phn t hu c khng
phi protein, n c gi l coenzym (coemyme). Nhiu vitamin l thnh phn ca
coenzym.
Trong nhiu phn ng oxi ho - kh c enzym xc tc, in t c chuyn
thnh i t v tr hot ng ca enzym n coenzym c chc nng nh cht nhn
in t. V sau, coenzym chuyn in t cho mt enzym khc gii phng chng (v
nng lng chng mang) cho c cht trong phn ng khc. Thng thng, cc in
t cp i vi proton (H*) di dng nguyn t hidro. Theo cch ny, coenzym lm
con thoi nng lng (shuttle energy) di dng nguyn t hidro t enzym ny n
enzym khc trong t bo.
Mt trong nhng coenzym quan trng n h t l ch't nhn hidro nicotinamit
adenin dinucleotit (NADO (Hnh 4.8). Phn t NAD* gm hai nucleotit lin kt vi
nhau. Nucleotit l mt ng 5 cacbon vi mt hoc nhiu nhm photphat gn vo
mt u v mt baz hu c gn vo u kia. Hai nucleotit c chc nng khc nhau
trong phn t NAD*: AMP hot ng nh ci li (nhn) to nn hnh dng nhiu
enzym nhn din, cn NMP l phn hot ng ca phn t, ng gp v tr d b
kh (d nhn in t).
Khi NAD* nhn in t v nguyn t hidro (gm hai in t v mt proton) t
v tr hot ng ca enzym, n b kh thnh NADH. Lc ny, phn t NADH mang
hai in t cao nng v proton. S oxi ho ca cc phn t cha nng lng c tc
131

dng cung cp nng lng cho t bo, km theo vic tc b in t khi cc phn
t v chuyn chng cho NAD* v trong hot ng trao i cht ca t bo th
nhiu nng lng ca NADH c chuyn cho phn t khc.

Hot ng ca enzym thng c cc nhm ngoi h tr m c th

i cc ion kim loi hoc cc cht khc. Nhm ngoi l cc phn t
hu c khng phi protein c gi l coenzym.

C -N H .

Nhm
phn ng
NMP

0
1
0 = P - 0

CHo

= P -0

Hlnh 4.8. Cu trc ho hc ca nlcotinamlt adenin dlnudeotil (NAD*). Nhm ngoi ch

yu ny gm hal nucleotit NMP v AMP gn u vo nhau

4.2.5. Enzym c nhiu dng

Trong khi nhiu enzym tri ni trong t bo cht ca t bo, t do vn ng v
khng gn kt vi bt k cu trc no, th cc enzym khc hot ng nh l b
phn kt hp ca cc cu trc v bo quan t bo.
132

4.2.5.1. Cc phc h a em ym
Trong t bo, mt s enzym xc tc cc bc khc nhau ca mt chui phn
ng kt hp lng lo vi nhau trong t hp lin kt khng cng ho tr gi l phc
h a enzym (multi emyme complexes). Th d trong hnh 4.9, phc h a enzym
mang tn piruvat dehidrogenaza vi khun cha cc enzym thc hin ba phn ng
k tip nhau trong qu trnh chuyn ho oxi ho. Mi phc h c nhiu bn sao ca
mt trong ba enzym gm tt c 60 tiu n v protein. Nhiu tiu n v kt hp
hot ng vi nhau ging nh mt nh my con.

(a)
Hlnh 4.9. Enzym piruvat dehidrogenaza
(a) Enzym (m hnh) tin hnh oxi ho piruvat l mt trong s cc enzym phc tp nht
bit, n c 60 tiu n v
(b) Nhiu trong c6 th thy trong nh hin vi in t (200.000X)

Cc phc h a enzym th hin nhng u im quan trng trong hiu qu xc tc:

1. Vn tc ca phn ng c enzym c gii hn nh tn s' m enzym bt gp
c cht ca n. Nu mt lot cc phn ng k tip nhau xy ra bn trong mt phc
h a enzym, sn phm ca mt phn ng c th c phn pht cho enzym tip
theo m khng gii phng n khuch tn i ra khi chui phn ng.
2. Do c cht phn ng khng bao gi ri khi phc h trong qu trnh thng
qua lot phn ng nn xc sut ca cc phn ng ph khng mong mun b loi b.
3. Tt c cc phn ng xy ra bn trong phc h a enzym c th c iu tit
nh mt n v.

133

Ngoi piruvat dehidrogenaza iu tit 8 thm nhp vo chu trn h Krebs, mt

8 cc qu trnh ch yu khc trong t bo c cc phc h a enzym xc tc. Mt
h thng c nghin cu tt l phc h synthetaza ixit bo c chc nng xc tc
qu trnh tng hp cc axit bo t cc cht tin th n hai cacbon. C by enzym
khc nhau trong phc h a enzym ny v cc cht trung gian ca phn ng vn
kt hp vi phc h cho ton b lot phn ng.

4.2.S.2. Khng p h i t t c cc ch t xc tc sinh hc l p ro tein

Cho n mt vi nm trc y, phn ln cc sch gio khoa sinh hc u cho
rng "Enzym l cht xc tc ca cc h sinh hc" v "protein l i phn t xc tc
duy nht trong h sinh hc". Song khi nim ny b ph nhn vi vic pht hin
cc phn t ARN xc tc u tin hay ribozim (Cech, T.R etal, 1981; G uerrier Takad, c , etal, 1983). Theo Cech v cng s th cc phn ng ko theo phn t
ARN c xc tc bi bn th n ARN, ch khng phi nh enzym. Quan im ban
u ny c khng nh nh b sung cc th d v s xc tc ca ARN trong
mt s nm qua. Thc nghim cho thy ribozim kch thch m nh vn tc cc phn
ng sinh ho ring bit v th hin tnh c hiu c bit vi cc c cht m chng
tc ng trn . Nh trong phn 4.2.1 trnh by v enzym, vi pht hin mi
ny cng nhiu thc nghim v sau, n nm 1989, Thomas Cech v Sidney
Altman dnh gii Nobel ho hc v ARN ng vai tr nh mt enzym thc th
m h gi l rbozim.
C t nht hai loi ribozim. Ribozim thc hin phn ng xc tc ni phn t
(ntramolecular catalysis) c cu trc gp np th xc tc phn ng trn bn th n
mnh. Cn ribozim no thc hin s xc tc gia cc phn t (intermolecular
catalyss) th hot ng trn cc phn t khc v khng t bin i trong qu trnh.
Nhiu phn ng t bo quan trng ko theo cc phn t nh ARN bao gm cc
phn ng ct b on khng cn thit khi cc bn sa ARN ca gen. l phn
ng t ct b intron nhm I (intron l on nucleotit khng c m ho trong
phn t ARN thng tin sd cp trong nhn to mARN chnh thc) chun b cho
ribosom tng hp protein. Ribozim cng thc y qu trn h sao chp ADN bn
trong ty th. Trong t t c cc trng hp ny, kh nng xc tc ca RN ang c
nghin cu m nh m. c bit trong qu trnh quang hp phc tp m c enzym v
ARN u ng vai tr xc tc quan trng.
Kh nng ca ARN, mt phn t thng tin hot ng nh mt cht xc tc,
kch thch nim hng phn ln gia cc nh sinh hc v c lin quan vi ngun gc
t pht (t sinh) ca gi thuyt v s sng. Ci ^ xut hin u tin, protein hay
axit nucleic? Gi y, t ra cng c th cho rng ARN c6 th tin ho u tin v
xc tc cho s hnh th n h cc protein u tin.

134

Khng phi mi cht xc tc sinh hc l lng t do trong t bo

cht. Mt s l b phn ca cc cu trc khc v mt s thm chi
khng phi protein.

4.3. ATP l tin t nng ng ca s sng

ATP l g?
Tin t nng lng ch yu m tt c t bo s dng l phn t adenozin
triphotphate (ATP). Khi nng lng m cy thu c trong qu trnh quang hp
c chuyn giao sn xut ATP nh phn ln nng lng d tr trong cht bo
v tinh bt. T bo dng ATP d tr cung cp hu ht mi qu trnh cn nng
lng chng thc hin t vic cung cp nng lng hot ho cho cc phn ng hc
v qu trnh dn truyn ch ng vt cht qua mng n vic vn ng qua mi
trng v sinh trng.

4.3.1. Cu trc ca phn t ATP

Mi phn t ATP gm ba thnh phn nh hn (Hnh 4.10). Thnh phn u
tin l ng 5 cacbon riboz, c chc nng nh b khung hai tiu n v khc
gn vo. Thnh phn th hai l adenin - mt phn t hu c gm hai vng cacbon nit. Mi nguyn t nit trong vng c mt i in t khng gp chung v ht yu
ion hidro. Do , adenin hot ng ho hc nh mt baz v thng c ni n
nh l mt baz nit (n l mt trong bn baz nit c m t trong ADN). Thnh
phn th ba ca ATP l nhm triphotphat (ba gc photphat lin kt thnh mt
chui).

4.3.2. ATP d tr nng ng nh th no?

iu mu cht gii thch ATP d tr nng lng nh th no nm nhm
triphotphat ca phn t. Cc nhm photphat nhm tch in m cao, nn chng
y nhau r t mnh. Do lc y tnh in gia cc nhm photphat tch in nn hai
lin kt cng ho tr nl cc gc photphat khng n nh, Phn t ATP thng c
v nh l mt "l xo xon c" icoiled spring)y cc nhm photphat b ko cng ra xa
nhau.
Cc lin kt khng n nh gi cc photphat vi nhau trong phn t ATP c
nng lng hot ho thp v d dng b ph v. Khi b ph v, chng c th chuyn
mt lng nng lng ln. Trong phn ln phn ng c ATP tham gia th ch lin
kt photphat cao nng ngoi cng b thu phn, tch ring nhm photphat u
cu. Khi iu ny xy ra, ATP bin thnh adenozin diphotphat (ADP) v nng
lng tng ng 7,3 kcal/mol c gii phng di iu kin tiu chun. Nhm
photphat c gii phng thng gn tm thi vi mt s' phn t trung gian. Khi
phn t ny loi nhm photphat (dephosphorylated), nhm photphat c gii
phng di dng photphat v c (Pi).
135

Nhm
triphotphat

o1
0

p o~

Cc lin kt
cao nng
0

= p o -

Adenne

L i A M P

= p o
0

(b)
Hlnh4.10. PhntATP
(a) M hnh
(b) S cu trc u chng t rng ging nh NAD*. ATP c li AMP. Song trong
trng hp ny, nhm phn ng c b sung vo u cui ca nhm photphat AMP khng
phi l nucleotit khc m ng hn l mt chui gm hai nhm thm photphat. Cc lin kt
ni hai nhm photphat vi nhau v vi AMP l cc Hn kt d tr nng lng

4.3.3. ATP cung cp nng lng cho cc phn ng cn nng lng nh

th no?
T bo dng ATP thc y cc phn ng thu nhit. Cc phn ng khng
tin hnh t pht do sn phm ca chng c nng lng t do nhiu hn cht phn
ng. Song, nu 8 tch lin kt cao nng tn cng ca ATP gii phng nhiu nng
lng hn phn ng tiu th nng lng th s bin i nng lng chung ca hai
phn ng lin kt s pht nhit (gii phng nng lng) v chng s cng tin
hnh. Do hu nh mi phn ng th u nhit cn t nng lng hn so vi nng lng
136

c gii phng nh tch ATP, ATP c th cung cp phn ln nng lng m t bo

cn dng.
Nt c trng ca ATP khng nhng l cht cho nng lng hiu qu m cn
c tnh khng n nh ca cc lin kt photphat, nhng ATP khng phi l phn t
d tr nng lng di hiu qu. Cc cht bo v hidratcacbon thc hin chc nng
ny t"t hdn. Phn ln t bo khng duy tr kho d tr ln ca ATP. Thay vo ,
chng thng ch c mt vi s cung cp ATP th yu bt k thi gian no v
chng lin tc to nhiu ATP t ADP v Pj.
Tinh khng n nh ca cc lin kt photphat khin cho ATP trd
thnh mt cht cho nng lng tuyt vi.

4,4. Trao i ch t l i sng ho hc ca t bo

4.4.1. Cc con n g sin h ho : n v t ch c c a tr a o i c h t
Ho hc s sng, l ton b
cc phn ng ho hc c sinh vt
thc hin gi l s trao i ch't
(metabolism) hay chuyn ho vt
cht. Phn ng no tiu th nng
lng to ra hay bin i cc
lin kt ho hc gi l phn ng ng
ho (anabolic reaction) hay s ng
ho {anabolism). Cc phn ng thu
nng lng khi cc lin kt ho hc b
ph v gi l phn ng d ho
{cataboic reaction) hay s d ho
(catabolsm).
Sinh vt cha r t nhiu loi
enzym khc nhau xc tc cho v s
phn ng. Cc phn ng trong t bo
to thnh chui gi l con ng sinh
ho {biochemical pathiuays). Trong
cc con ng , sn phm ca phn
ng ny tr th n h c ch^t cho phn
ng tip theo (Hnh 4.11). Con ng
sinh ho l n v t chc ca qu
trn h trao i cht - yu t m mt
sinh vt iu tit t c hot
ng cht ch ca qu trnh trao i

C cht

Sn phm

Hnh 4.11. Con ng sinh ho

Co cht u tin c hot ho nh enzym 1,
bin CX3 cht thnh dng mi v c nhn
bit nh enzym 2. Trong con ng ny, m
enzym li hot ho sn phm ca ga on rc

137

cht. Trong con ng sinh ho xy ra cc xoang (ngn) chuyn ho ca t bo,

phn ln cc bc m enzym xc tc u i theo trn h t, th d cc bc ca chu
trnh axit xitric (ch(ig^) xy ra bn trong ty th. Nh xc nh nhiu enzym xc
tc cc bc ny nh v u, chng ta c th "minh ho" (mp out) 8 cc qu
trnh chuyn ho vt cht trong t bo.

4A.1.1. Cc con ng sn h ho tin ho nh th no?

nhng t bo nguyn thu nht, cc qu trn h sirih ho ban u c l bao
gm nhng phn t giu nng lng c ly t mi trng. H u ht cc phn t
cn cho qu trnh ny tn ti trong "mn sp hu c" ca cc i dng nguyn
thu. Cc phn ng c xc tc u tin th n gin, cc phn ng mt bc th
mang nhng phn t cng nhau trong liu lin kt khc nhau. Cui cng, cc
phn t giu nng lng mi trng ngoi tr nn cn kit, ch nhng sinh vt
no tin ho theo cch to ra nhng phn t giu nng ng t cc c cht khc
trong mi trng mi c th tn ti. Do , c phn ng gi nh:
F

-> H

G
y hai phn t giu nng lng (F v G) phn ng to ra hpcht H v
gii phng nng ng, tr nn phc tp hn khi s cung cp ca F trong mi
trng cn kit. Mt phn ng mi c b sung trong phn t b cn kit (F)
c to ra t phn t khc (E) cng c trong mi trng:

E - F
+ - H
G
Khi 8 cung cp ca E li tr nn cn kit, cc sinh vt c th to ra n t mt
s tin cht sn c khc (D). Khi D b cn kit, cc sinh vt li c thay th
bng nhng sinh vt c kh nng tng hp D t phn t khc (C):

- D

F
+

G
Con ng sinh ho gi nh ny tin ho chm theo thi gian, vi phn ng
cui cng th xy ra trc tin, cn cc phn ng pha trc li tin ho sau. Nhn
vo con ng ny, ta s thy rng sinh vt, bt u vi hp cht c , c th tng hp
H qua nhiu bc. Con ng sinh ho trong c th sinh vt c cho rng tin
ho khng phi ngay lp tc, m tng bc mt, li li theo thi gian.
138

4.4.I.2. Cc con ng sinh ho c iu chnh nh th n o f

con ng sinh ho hot ng hiu qu, t bo phi iu phi v iu chnh
hot ng . T bo khng nht thit phi tng hp mt hp cht khi c mt lng
ln cht , v lm nh vy s to ra nhiu nng lng v nguyn liu th m t
bo c th a i s dng ni khc. Do , iu quan trng i vi mt t bo l c
ch tm thi con ng sinh ho khi sn phm ca chng cha cn thit cho t bo.

Hlnh 4.12. S c ch lin h ngc

(a) Con ng sinh ho khng c c ch lin h nguc
(b) Con ng sinh ho trong sn phm cui cng tr thnh cht tc ng d| lp th
i VI enzym u tin trong con ng. Ni cch khc, s hnh thnh sn phm cui cng
ca con ng lm ngng con ng

S iu chnh con ng sinh ho n gin thng ph thuc vo c ch lin h

ngc tinh t: sn phm cui cng ca con ng lin kt vi v tr d lp th trn
enzym xc tc phn ng u tin trong con dng. Trong con ng gi nh m
chng ta va trn h by, enzym xc tc phn ng c -> D phi c v tr d lp th i
vi H (sn phm cui cng ca con ng). Khi con ng sn xut hng lot sn
139

phm v lng H trong t bo tng ln th t t yu mt trong nhng phn t H s

bt gp v tr d lp th trn enzym c
D. Nu sn phm H hot ng nh mt c
ch d lp th ca enzym th s lin kt ca n vi enzym s c ch phn ng c ->
D. S c ch phn ng ny - phn ng u tin trong con ng s c ch mt cch
hiu qu ton b con ng. Do , khi t bo sn xut ra lng sn phm H ngy
cng tng th t bo s t ng c ch kh nng to ra nhiu sn phm hn. y l
m hnh iu chnh m ang tn s c ch lin h ngc (Hnh 4.12).
Con ng sinh ho l mt lot cc phn ng c t chc thng
c iu chnh nh m t n v.

4.4.2. S tin ho ca qu trnh trao i cht

Qu trnh trao i cht hay chuyn ho vt cht thay i nhiu khi s sng
trn tri t pht trin. l s th t hin nhin v cc phn ng m sinh vt
s dng hp th nng lng t m t tri tng hp cc phn t hu c (s ng
ho) v v sau phn gii cc phn t hu c thu nhn nng lng (s d ho).
Hai qu trnh ny tin ho trong m quan h vi nhau v y chnh l ch
ca hai chng tip theo.

4.4.2.1. S ph n hu
Dng nguyn thu n h t ca s shg nhn nng lng ho hc nh s phn hu
idegrading) hay phn gii (breaking doivn) cc phn t hu c do cc qu trnh
khng sng to ra.
S kin ch yu u tin trong qu trn h tin ho ca s trao i cht l
ngun gc ca kh nng khai thc sn xut nng lng lin kt ho hc. giai
on u tin, sinh vt bt u d tr nng lng ny trong cc lin kt ca ATP cht mang nng lng c mi sinh vt s dng.

4.4.2.2. Qu trnh ng p h n
S kin chnh th hai trong s tin ho ca qu trn h trao i cht l ng
phn (glycolysis) - s phn gii u tin ca glucoz. Khi protein pht trin cc
chc nng xc tc a dng, t bo c kh nng thu nhn phn ln nng lng ca
cc lin kt ho hc trong cc phn t hu c nh phn gii cc lin kt ho hc
trong lot cc bc. Th d, s phn gii t t ng 6-cacbon glucoz thnh phn t
3-cacbon c thc hin qua 10 bc, to hai phn t ATP. Nng lng tng hp
ATP l nh phn gii cc lin kt ho hc v hnh th n h cc lin kt mi vi nng
lng lin kt nh hn v hiu nng lng gia hai trng thi c chuyn giao
tng hp ATP. Con ng sinh ho ny c gi l ng phn.
Khng nghi ng g na, ng phn tin ho r t sm trong lch s ca s
sng trn hnh tinh, t khi con ng sinh ho ny c trong mi sinh vt sng.
l mt qu trnh ho hc m khng b bin i trong sut 3 t nm qua.
140

4.4.2.3. Q uang hp k k h i
S kin ch yu th ba trong qu trnh tin ho ca s trao i cht l quang
hp k kh {anaerobic photosynthesis). giai on u, mt s sinh vt tin ho
theo mt phng thc khc ca qu trnh sn sinh ATP gi l quang hp. l vic
nhn nng lng tng hp ATP nh thay cc lin kt ho hc nh trong ng
phn. Cc sinh vt ny pht trin kh nng dng nh sng bm cc proton
(HO ra ngoi t bo v dng gradient proton c c cung cp nng lng cho qu
trnh tng hp ATP - qu trnh ny c gi l s ho thm (chemiosmosis).
Quang hp tin ho khi khng c oxi v hot ng tt trong trng hdp .
H 2S ho tan c trong cc i dng, bn di tng kh quyn khng c kh oxi v
HgS c chc nng nh mt ngun nguyn t hidro sn c tng hp cc phn t
hu c. Lu hunh t do c to ra nh mt sn phm ph ca phn ng ny.

4.4.2.4. S c in
h n it
C nh nit l bc ch yu th t trong con ng tin ho ca qu trnh
trao i cht. Protein v axit nucleic khng th c tng hp t cc sn phm ca
quang hp, v c hai phn t sinh hc quan trng ny u cha nit. Vic thu nhn
nguyn t nit t kh N 2 , qu trn h ny gi l s c nh nit (nitrogen fixation), i
hi s phn gii lin kt ba NsN. Phn ng quan trng ny tin ho u tin
trong bu kh quyn giu hidro, khng c oxi ca qu t. Oxi hot ng nh mt
cht c i vi s c nh ni t nn qu trnh ny ch xy ra trong mi trng
khng c oxi hoc bn trong cc xoang khng c oxi ca mt s' vi khun no y.

4.4.2.5. Quang hp g i i p h n g oxi (quang hp hnh thnh oxi)

S thay th H 2O cho H 2S trong quang hp l s kin ch yu th nm trong
lch s ca qu trnh trao i cht. Quang hp gii phng oxi do dng H 2O ch
khng phi HS nh mt ngun nguyn t hidro v in t kt hp. V quang hp
thu nguyn t hidro t oxi kh hn l t lu hunh kh, cho nn n pht sinh kh
oxi ch khng to lu hunh t do.
Hn hai t nm trc, cc t bo b nh c th thc hin qu trnh quang hp
to oxi ny nh khun lam, bi v l dng sng u th trn hnh tinh. Kh oxi
bt u tch lu trong kh quyn. l s bt u ca mt chuyn bin ln lm
bin i vnh vin iu kin trn tri t. Gi y, kh quyn ca chng ta c 20,9%
oxi v mi phn t c bt ngun t mt phn ng quang hp to oxi.

4.4.2.6. H hp hiu khi

H hp hiu kh l s kin th su v cui cng trong lch s pht trin ca
con ng trao i cht. Qu trn h t bo ny hp th nng lng nh ly cc in
t cao nng t phn t hu c. H hp hiu kh s dng cng loi bm proton nh
quang hp v tin ho nh mt bin i ca b my quang hp c bn. Song,

141

hidro v cc in t kt hp ca chng khng phi nhn t H2 S hay H2 O nh trong

quang hp m ng hn l t s phn gii cc phn t hu c.
Cc nh sinh hc cho rng kh nng thc hin quang hp khng c H 2S
tin ho u tin trong cc vi khun ta khng lu hunh purple nonsulur hacteri)
do chng thu hidro t cc hp cht hu c. iu chc chn xy ra l trong s cc
th h con chu ca vi khun quang hp h hp ny, mt s' s khng hon ton
quang hp v tn ti nh nng lng v hidro bt ngun t s phn gii cc phn
t hu c. Ty th trong t t c t bo nhn thc c xem l con chu ca chng.
Su s i mi ch yu nu bt s tin ho ca qu trn h tra o i
cht nh chng ta bit ngy nay.

TM TT KHI NIM
4.1. nh lut nhit ng hc cho bit nng lng bin i nh th no
Nng lng l kh nng gy ra bin i, to ra s vn ng chng li mt
lc hoc sinh cng.
ng nng tham gia ch ng trong vic sinh cng, cn th nng c kh nng
thc hin iu . Nhiu s bin i nng lng ca sinh vt ko theo s
bin i ca th nng thnh ng nng.
Phn ng oxi ho kh (redox reaction) l phn ng trong in t c ly
t mt nguyn t hoc phn t (oxi ho) v chuyn cho cht khc (kh).
nh lut th nh t nhit ng hc khng nh lng nng lng trong v tr
khng i. Nng lng khng b m t i v cng khng c to ra.
nh lut th hai nhit ng hc khng nh tnh hn n trong v tr
(entropy) c khuynh hng tng ln. Kt qu l, nng lng bin i t pht
thnh cc dng t tr t t hn.
Bt k phn ng ho hc no m sn phm ca chng cha t nng lng t
do hn cht phn ng ban u s c khuynh hng t pht. Song, hiu (s sai
khc) trong nng lng t do khng xc nh tc ca phn ng.
Tc ca phn ng ph thuc vo lng nng lng hot ho cn ph v
cc lin kt vn c.
Xc tc l qu trnh lm gim nng lng hot ho nh tc ng ln cc lin
kt ho hc.

4.2. Enzim cht xc tc sinh hc

Enzym l cht xc tc ch yu ca t bo. Chng gy tc ng ln tc phn
ng nhng khng lm nh hng ln s cn bng ch yu gia cht phn ng
v sn phm.
142

 T bo cha nhiu enzym khc nhau v mi enzym xc tc mt phn ng c

hiu.
Tnh chuyn ho ca mt enzym l do v tr hot ng ca enzym ch khp vi
mt hoc mt vi loi phn t c cht.

4J3. ATP tin t nng lng ca s sng

T bo nhn nng lng t quang hp v oxi ho cc phn t hu c, dng
nng lng gii phng ra to ATP t ADP v photphat.
Nng lng d tr trong ATP c s dng thc y cc phn ng thu
nhit.

4.4.

S trao i cht hay chuyn ho vt cht l i sng ho hc ca t bo

Ni chung, cc phn ng cui cng ca mt con ng sinh ho tin ho

u tin, cc phn ng xy ra trc trong con ng c b sung v sau mi
ln mt bc.

GII THCH THUT NG

1. A c tiv a tio n e n e rg y (nng lng hot ho): Nng lng cn c cung cp
ph v tnh n nh ca cc lin kt ho hc vh c v khi ng mt phn
ng ho hc. Ngay c phn ng t pht cng yu cu nng lng hot ho ban
u ny.
2. E n e rg y (nng lng): Mi hot ng ca t bo c thc y bng nng lng
c m t tri cung cp hoc cha trong cc lin kt ho hc.
3. C a ta y sis (s xc tc); S gia tng tc ca phn ng ho hc nh lm gim
nng lng hot ho.
4. C oenzym e (coenzym): Phn t hu c khng phi protein, ng vai tr ph tr
trong phn ng c enzym xc tc v thng hot ng nh mt cht cho hay
cht nhn in t. NAD^ l mt coenzym.
5. E n d e rg o n ic r e a c tio n (phn ng thu nhit): Phn ng ho hc phi c ngun
nng lng bn ngoi cung cp trc khi phn ng tin hnh - ngc vi phn
ng pht nhit.
6. E n tro p y (entropy): s o v tnh hn n hoc v tr t t ca h thng. Trong t
bo, entropy l mt s' o gii thch c bao nhiu nng lng b pht tn (thng
l nhit phn b ng u) m n khng cn sinh cng c na.
Entropy cn c hiu nh l nng lng bin i t pht thnh dng t
tr t t hn. Do , m quan h gia cc phn t c khuynh hng tr nn v
tr t t nhiu hn (entropy tng ln) khi nng lng hng n dng c t chc
nh nh t - l s vn ng phn t hn n nh nhit chng hn.
143

7. E x erg o n ic re a c tio n (phn ng pht nhit): Mt phn ng ho hc to nng

lng. Phn ng pht nhit c khuynh hng tin hnh t pht mc d nng
lng hot ho l cn khi ng chng.
8. F re e e n e rg y (nng lng t do): Nng lng c sn sinh cng. Nng lng t
do c m t trong cc lin kt ho hc ca phn t l nng lng t do ca phn
t tr i nng lng v ch do s v trt t. Mt cch chung hn, nng lng t
do l nng lng sn c sinh cng.
9. K ilo calo rie (kilocalo = 1000 calo): Mt calo l nhit cn nng nhit ca 1
gam nc ln 1 (1C).
10. E nzym e sp e c ific ity (tnh chuyn ho ca enzym): Mt enzym ch xc tc cho
mt hoc mt vi phn ng ring do ch cc phn t c cht nht nh c th
khp cht vo cc rnh hay hc (v tr hot ng ca enzym) trn b m t ca
enzym.
11. M etab o lism (s chuyn ho vt cht hay s trao i cht): Bao gm t t c cc
phn ng ho hc trong t bo sng, cho php sinh vt nm bt v hp th nng
lng.
12. O x id a tio n (s oxi ho) l s mt in t nguyn t hay phn t. N xy ra
ng thi vi s kh mt s nguyn t hoc phn t khc do in t b mt d
mt nguyn t th c nguyn t khc nhn.
13. R e d u c tio n (s kh) l s nhn in t mt nguyn t hay phn t. Phn ng
oxi ho - kh l mt phng thc quan trng ca qu trn h truyn nng lng
bn trong h thng shg.
14. S u b s tra te (c cht) l phn t m enzym tc ng, cht phn ng ban u
trong phn ng c enzym xc tc.
15. Lysozym e (enzym gy tan): Enzym phn hu polysaccarit trong vch t bo.
Enzym phn b' rng trong th thc khun, lng trng trng v nc m t ngi.
N xc tc phn ng thu phn cc lin kt c hiu gia cc gc trong
polysaccarit. Do , chng c vai tr phn gii vch t bo, dn n tiu hu v
lm vi khun cht.

CU HI N TP
1. Trnh by s khc nhau gia qu trnh ng ho v d ho.
2. Hai trng thi c bn trong nng lng c th tn ti l g? Chng khc nhau
nh th no?
3. nh ngha s oxi ho v s kh. Ti sao hai phn ng ny phi lun lun xy ra
vi nhau? Trong phn ng 0 X1 ho kh, cht phn ng v sn phm trao i ci g?

144

4. Trnh by nh lu t th n h t v th hai ca nhit ng hc

5. Trnh by nh ngha nhit, entropy v nng lng t do.
6. S khc nhau gia phn ng pht nhit v phn ng thu nhit l g? Loi phn
ng no c khuynh hng tin hnh t pht?
7. Nng lng hot ho l g? N c lin quan vi tc phn ng nh th no?
Cht xc tc nh hng ln nng lng hot ho, tic phn ng v nh hng
ln t l cui cng ca cht phn ng bin thnh sn phm nh th no?
8. Tic ca phn ng c enzym xc tc chu nh hng ca nhit , pH nh th
no? C s phn t v cc tc ng ca cc iu kin ny ln tc phn ng l
g?
9. S khc nhau gia v tr hot ng v v tr d lp th trn enzym l g? Cht c
ch cnh tranh hot ng ca enzym khc vi cht c ch khng cnh tranh
nh th no?
10. Nhm ngoi v coenzym tham gia trong phn ng ho hc xy ra bn trong t
bo nh th no?
11. Phn no ca phn t ATP cha lin kt c dng cung cp nng lng cho
phn ln cc phn ng thu nhit trong t bo?
12. Con ng sinh ho l g? c ch lin h ngc iu chnh hot ng ca con
ng sinh ho nh th no?

CU HI SUY LUN
1. Hu nh khng c nh sng m t tri thm nhp vo i dng su thm. Song
nhiu loi c sng hp dn con mi v gi phi m nh nh to nh sng
ring ca chng. Vy nh sng ny bt ngun t u? S pht sinh nh sng
c cn nng lng khng?
2. Cc phn ng oxi ho kh c th ko theo nhiu loi phn t. Ti sao hidro v
oxi tham gia trong phn ng ny li c tm quan trng sng cn trong cc h
sinh hc?
3. Mt s ngi cho rng: s tin ho thng kt hp vi s tng dn v phc tp
(trt t) ca sinh vt khng th xy ra do entropy (tnh hn n) ang tng ln
trong v tr. Quan nim ny c ng khng?

145

Chng V
T BO TH U HI NNG LNG NH T H NO

KHI NIM CHNG

5.1. T bo thu hi nng lng trong cc lin kt ho hc
Dng nng lng ho hc thc y qu trnh trao i cht. Nng lng
trong cc lin kt c - H, c - 0 v cc lin kt ho hc khc c th c thu
ly v s dng lm nhin liu cho tng hp ATP.

5.2. H hp t bo oxi ho cc phn t thc n

Khi qut v qu trn h d ho glucoz; nng lng ho hc trong ng c
thu hi nh photphoril ho mc c cht v h hp hiu kh
Giai on 1: ng phn. 10 phn ng ca ng phn thu nng lng t
glucoz nh thay i cc lin kt.
Giai on 2: s oxi ho piruvat. Piruvat - sn phm ca ng phn b oxi

ho thnh axetyl-CoA.
Giai on 3; chu trn h Krebs. Trong mt lot cc phn ng, in t b tc

ot khi axetyl-CoA.
Thu hi nng lng nh chit r t in t. H hp vi c cht glucoz l mt
lot cc phn ng oxi ho - kh gii phng nng lng nh chuyn in t

n gn hn vi nguyn t oxi.
Giai on 4: chui truyn in t. in t c thu hi t glucoz thng qua
mt chui cc protein m ng dng nng lng bm proton, thc y s tng
hp ATP.
Tm t t h hp hiu kh. H hp hiu kh thu nhn hn mt na nng lng
trong cc lin kt ho hc ca glucoz.
iu chnh h hp hiu kh. Mc ATP cao c khuynh hng lm ngng h
hp t bo do c ch cc phn ng ch yu.
147

5.3. Qu trinh d ho priotein v cht bo c th to nhiu nng lng

* Glucoz khnig phi l nguyn liu duy nht. Protein v cht bo b phn gii
to sn phm lim nguvn liu cho h hp t bo.
5.4. Tbo c thchuym ho nguyn liu thc n khng c oxi
S ln men. Ln. men cho php chuyn ho tip khi khng c oxi nh chuyn
in t thu nhn t ng phn cho cc phn t hu c.
S sng tin trin c l nh nng lng. Mi hot ng m sinh vt tin
hnh nh s bi ca vi khun, ting ku ca con mo, bn c cc dng ch ny
u dng nng lng. Trong chng ny, chng ta s trnh by cc qu trnh m
mi t bo s dag tlhu nhn nng lng ho hc t cc phn t hu c v bin
nng lng thnh ATP. Chng ta s xem xt quang hp - qu trnh dng nng
lng nh sng ch ktomg phi nng lng ho hc mt cch chi tit trong chng
6 - chng cui ca gio trinh ny. Chng ta xem xt qu trn h bin i nng lng
ho hc thnh ATP truc quang hp bi v mi sinh vt k c sinh vt quang hp v
sinh vt dng c th qniaang hp lm thc n u c kh nng thu hi nng lng t
cc lin kt ho hc mc d qu trnh ny v quang hp c nhiu im chung.

5.1. T bo thu hi nng Ing trong cc Hn kt ho hc

* D n g n n g l ng ho hc th c y q u tr in h tr a o i c h t
Thc vt, to v Knt fi' vi khun thu nhn nng lng ca nh sng m t tri
thng qua quang top, bin nng lng bc x thnh nng lng ho hc. Cc sinh
vt ny cng mt vi c th khc dng nng lng ho hc theo cch ging nhau
gi l sinh vt t dig (autotrophs). Tt c cc sinh vt khc sng nh vo sinh
vt t dng to nni| ng c goi l sinh vt d dng hterotrophs). C t
n ht 95% loi sinh vtt tn hnh tinh gm t t c ng vt, nm, phn ln sinh vt
nguyn sinh v vi khuini u l sinh vt d dng.
Nng lng ho hc trong thc n nm u v sinh vt d dng thu hi
nng lng nh th no thc hin nhiu chc nng ca s sng? Phn ln
thc n cha nhiu hidratcacbon, protein v cht bo, t t c cha y nng lng
trong cc lin kt ho hc. Th d, hidratcacbon v cht bo cha nhiu lin kt
ccbon-hidro (C'H cjng nh lin kt cacbon-oxi (C-0). Qu trnh chit r t nng
lng t hn hp hu C(d phc tp ny c x l trong cc giai on khc nhau.
u tin, enzym phn gi cc phn t ln thnh phn t nh hn, l qu trnh
tiu ho (digestiom). v sau, cc enzym phn gii cc m nh phn t nh mi ln
mt t, thu nng lulng t cc lin kt C-H v cc lin kt ho hc khc mi giai
on. Qu trnh ny e gi l s d ho (catabolism).
Trong khi ta ang thu nhn nng lng t nhiu hp phn ca thc n th
trc ht ta hy tp tr-ung vo qu trnh d ho ca hidratcacbon. Chng ta s theo
148

di ng su cacbon - glucoz (CgHiaOe) khi cc lin kt ho hc c chit rt nng

lng mt cch t t. v sau, s xem xt qu trnh d ho protein v cht bo.

5.1.1. H hp t bo
Nng lng trong cc lin kt ho hc c th c hnh dung nh l th nng
do in t mang m to nn lin kt cng ho tr. T bo thu hi nng lng ny
nh a in t vo hot ng thng l tng hp ATP - tin t nng lng ca t
bo. Ngay sau , in t ngho nng lng (energy - depleted electron) (kt hp vi
proton di dng nguyn t hidro) c chuyn cho mt s phn t khc. Khi kh
oxi (O2) nhn nguyn t hidro th nc hnh thnh v qu trnh ny c gi l h
hp hiu kh {aerobic respiration). Khi phn t v c (khng phi oxi) nhn hidro,
qu trnh m ang tn h hp k kh (anaerohic respiration). Khi phn t hu c nhn
nguyn t hidro, gi l s ln men ermentation).
V m t ho hc, c t khc nhau gia qu trnh d ho hidratcacbon trong t
bo v s t chy g trong l si. Trong c hai trng hp, cht phn ng l
hidratcacbon v oxi, sn phm l kh cacbonic, nc v nng lng:
CeHiaOe + 6O2 -> 6CO2 + 6H2O + nng lng (nhit hoc A T P )

S bin i v nng lng t do trong phn ng ny l -720 kcal (-3012 kilo

jun)/mol glucoz di iu kin xy ra trong t bo (gi tr truyn thng l 686 kcal
hay -2870 kJ/mol di iu kin tiu chun: nhit phng, p sut lat...). S bin
i nng lng t do ny ch yu bt ngun t vic ph v 6 lin kt c - H trong
phn t glucoz (-) chng t rng sn phm c nng lng t do nh hdn cht phn
ng. Cng lng nng lng c gii phng hoc do glucoz thng qua qu trnh d
ho hoc b t chy, nhng khi glucoz b t chy trong l si th phn ln nng
lng c gii phng ra di dng nhit. Nhit ny khng th c dng sinh
cng trong t bo. iu mu cht vi kh nng ca t' bo thu nng lng
hu ch t qu trnh d ho cc phn t thc n nh glucoz l vic bin i mt
phn nng lng th n h dng hu ch hn. T bo tin hnh vic ny nh dng mt
s nng lng thc y qu trnh tng hp ATP - phn t c th cung cp nng
lng cho cc hot ng ca t bo.
5.1.2. P h n t A TP
Adenozin triphotphat (ATP) l tin t nng lng ca t bo - phn t c chc
nng truyn nng lng hp th c trong qu trnh h hp cho nhiu v tr dng
nng lng trong t bo. Nh vy, ATP c th sn sng truyn nng lng nh th
no? T chng 4, ta thy rng ATP gm ng (riboz) lin kt vi mt baz hu c
(adenin) v mt chui gm ba nhm photphat. Nh trong hnh 5.1, mi nhm
photphat tch in m. Ging nh cc in tch y nhau, nhm photphat lin kt
y ngc li lin kt m gi chng vi nhau. Nh mt by chut gi sn, cc
nhm photphat lin kt d tr nng lng ca lc y tnh in. Vic truyn mt
nhm photphat n phn t khc c tc dng ni lng l xo tnh in ca ATP,
149

ng thi kho (hm: cocking) l xo phn t c photphoril ho. v sau, phn

t ny c th dng nng lng tri qua mt s bin i c6 i hi cng hu ch.
Adenin
NHa

Nhm
triphotphat

II

0 - p - o
o

Jp

1
0'

"

~ 0 - P - 0 -C H 2
1
O'

Loi tch
in m

Hnh 5.1. Cu trc ca phn t ATP

ATP gm mt baz hu c v mt chui photphat gn vo u i din cOa ng 5 cacbon. Ch rng cc vng tch in ca chui photphat gn k nhau. Cc in tch kiu
ny c khuynh hng y nha to cho cc lin kt gi chng vi nhau mt th truyn nng
lng rt cao

5.1.3. T b o s d n g A TP n h t h no?
T bo dng ATP thc hin phn ln hot ng no i hi cng. Mt trong
nhng hot ng r rng nht l s vn ng. Mt s vi khun bi, t y ti pha
trc nh lm quay nhanh lng roi (tin mao) kiu ui di, ging nh tu thu di
ng nh lm quay chn vt. Trong qu trnh pht trin ca con ngi khi trng
thi phi, nhiu t bo vn ng qua li, trn ln nhau tm v tr mi, s vn
ng cng xy ra bn trong t bo. Cc si b t bn trong t bo c y nhanh co c.
Ty th di chuyn mt mt hoc hn dc theo cc t bo thn kinh hp nl chn vi
xng sng. Cc nhim sc th b cc vi qun ko i trong qu trnh phn chia t bo.
Tt c cc vn ng ny t bo u cn nng lng ATP.
Phng thc ch yu th hai t bo dng ATP l thc y phn ng thu nhit
{endergonic reaction). Nhiu hot ng tng hp ca t bo l thu nhit do cc phn
t tng hp (building molecules) ly nng lng. Cc lin kt ho hc ca cc sn
phm ca phn ng ny cha nhiu nng lng hn hoc c t chc hn so vi
cht phn ng. Phn ng khng th tin hnh cho n khi nng lng thm c
cung cp cho phn ng. l ATP c vai tr cung ng cho nng lng cn thit ny.
150

5.1.4. ATP thc y phn ng thu nhit nh th no?

ATP thc y phn ng thu
nhit nh th no? Enzym xc
tc phn ng thu nhit c hai v
tr lin kt trn b mt, mt v tr

^-A D P

Hot ng
Hnh 5.2. ATP thc y phn ng thu nhit nh

lng ny y cht phn ng v

tr th hai "ln dc" (uphill) thc
y phn ng thu nhit.

th no? Trong nhiu trng hp. mt nhm photphat

tch khi ATP c tc dng hot ho protein, xc tc
qu trnh iu nhit

Khi qu trnh tch phn t ATP thc y phn ng yu cu nng lng trong
t bo, hai phn ca phn ng - tch ATP v thu nhit xy ra vi nhau. Trong mt
s trng hp, hai phn u xy ra trn b m t ca cng enzym, chng lin kt
hoc "cp i" ging nh hai chn an< i. Trong cc trng hp khc, nhm
photphat cao nng t ATP gn vo protein xc tc qu trnh thu nhit, hot ho
protein (Hnh 5.2). Phn ng cp i cn nng lng tch ATP theo cch ny l
mt trong cc cng c ch yu c t bo s dng qun l nng lng.
Qu trnh d ho glucoz thnh khi cacbonic v nc trong c th'
sng gii phng khong 720 kcal (khong 3012 kJ) nng ng trn
mol glucoz. Nng lng ny c th u gi trong ATP phn t c
chc nng d tr nng lng nh lin kt cc nhm photphat tch
in nm gn nhau. Khi cc lin kt photphat trong ATP b thu
phn, nng lng c gii phng v sn sng d sinh cng.

5.2. H hp t bo oxi ho cc phn t thc n

5.2.1. Khi qut v qu trinh d ho glucoz
T bo c th tng hp ATP t qu trnh d ho cc phn t hu c theo hai
cch khc nhau:
1. Photphoril ho mc c cht {substrate - level phosphoryraton). Trong qu
trnh ny, ATP c tng hp nh chuyn mt nhm photphat trc tip cho ADP t
cht trung gian m ang photphat (Hnh 5.3). qu trnh ng phn (xem phn tip
theo), cc lin kt ho hc ca glucoz c dch chuyn trong cc phn ng, cung
cp nng lng cn to ATP.
2. H hp hiu kh {aerobc respiration). Theo cch th hai ny, ATP c
hnh th n h khi in t c thu hi, chuyn dc theo chui truyn in t v cui
cng c chuyn cho oxi. Cc sinh vt nhn thc to phn ln ATP t glucoz
theo cch ny.
151

Trong phn l0n sinh vt, hai qu trnh ny kt hp vi nhau. thu nng
lng tng hp ATP t ng glucoz khi c m t oxi, t bo thc hin mt lot cc
phn ng c enzym xc tc phc tp v xy ra trong bn giai on: giai on u
thu hi nng lng nh photphoril ho mc c cht thng qua ng phn, ba giai
on tip theo tin hnh h hp hiu kh nh oxi ho sn phm cui cng ca
ng phn.

-A D P

Hinh 5.3. Photphorll ho mc c cht. Mt s phn t nh photphoenol pimvat (PEP) c

mt lin kt photphat cao nang ging nh cc lin kt trong ATP. Khi nhm photphat ca
PEP c chuyn nh enzym xc tc n ADP, nng lung trong lin kt c bo ton v
ATP c to ra

5.2.1.1. Qu trinh ng p h n
Giai on 1: ng phn. Giai on u tin ca qu trn h chit r t nng
lng t glucoz l con ng sinh ho 10 phn ng gi l qu trn h ng phn
(glycolyss) c chc nng tng hp ATP nh photphoril ho mc cd cht. Cc enzym
xc tc phn ng ng phn tn ti trong t bo cht ca t bo, khng in kt
vi bt k mng hay bo quan no. u tin t bo tiu th hai phn t ATP trong
con ng tng hp bn phn t ATP nh photphoril ho mc c cht. T to di
ra hai phn t ATP cho mi phn t gucoz thng qua qu trn h d ho. Ngoi ra, t
bo thu c bn in t di dng NADH. Tuy nhin, tng hiu qu ATP l thp.
Khi qu trnh ng phn hon thnh, hai phn t piruvat va c hnh thnh
vn cha phn n nng ng m phn t gucoz ban u d tr.
ng phn xy ra trong mi sinh vt v cc phn ng ng phn to ATP
theo photphoril ho mc c cht c th xy ra vi oxi hoc khng c oxi. Song, trong
ng vt, vic thu hi cc in t cao nng nh ng phn khng th xy ra v
thi hn khi khng c oxi. Kt qu l, nu khng c oxi th t bo ng vt s cht.

152

5.2.I.2. H hp hiu khi

Giai on 2: oxi ho piruvat. Trong giai on ny, piruvat - sn phm cui
cng ca ng phn - b bin i thnh kh cacbonic v phn t hai cacbon gi l
axetyl-CoA. Khi mi phn t piruvat b bin i, mt phn t NADH c tng hp.
Giai on 3: chu trnh Krebs. Giai on ba nhp axetyl-CoA vo chu trnh gm
9 phn ng gi l chu trnh Krebs mang tn nh sinh ho ngi Anh Hans Krebs,
ngi p h t hin ra chu trnh ny (chu trnh KreLs cng c gi l chu trnh axit
xitric v axit xitric hay xitrat c hnh thnh trong bc u tin ca chu trnh v
t ph bin, chu trn h Krebs cng c gi l chu trnh tricacboxilic axit bi v citrat
c ba nhm cacboxin). Trong chu trnh Krebs, hai phn t ATP c chit r t theo
photphoril ho mc c cht v mt lng ln cc in t b loi ra nh NADH.

Hinh 5.4. Bc tranh khi qut v h hp hiu kh

Giai on 4: chui truyn in t. Trong giai on th t, cc in t cao nng

do NADM mang c s dng thc y qu trnh tng hp mt lng ln ATP
nh chui truyn in t.
S oxi ho piruvat, cc phn ng ca chu trnh Krebs v qu trnh tng hp
ATP nh chui truyn in t xy ra bn trong ty th ca mi sinh vt nhn thc
cng nh trong nhiu dng vi khun (ty th tin ho t vi khun - theo chng
1). Mc d, thc vt v to c th to ATP bng quang hp, nhng chng cng to
ATP bng h hp hiu kh ging nh ng vt v cc sinh vt khng quang hp
khc. Hnh 5.4 gii thiu bc tranh khi qut v h hp hiu kh.

153

5.2.I.3. H hp k khi
Vi s c m t ca oxi, t bo c th h hp hiu kh. Khi thiu oxi, mt s' sinh
vt h hp k kh, dng cht nhn in t v c khc, khng phi oxi. Th d,
nhiu vi khun dng lu hunh, n itrat hoc cc hp cht v c khc nh l cht
nhn in t thay cho oxi.
Vi khun sinh kh m etan {methanogen). Trong s cc sinh vt d dng tin
hnh h hp k kh l c khun nguyn thu (primtive archaebacteria) nh vi
khun a nhit (thermophiles). Mt s vi khun ny gi l vi khun sinh kh
metan, dng CO2 nh cht nhn in t, kh CO2 thnh CH 4 (metan) vi hidro bt
ngun t cc phn t hu c do sinh vt khc to ra.
Vi khun lu hunh: bng chng v qu trnh h hp k kh th hai trong s
cc vi khun nguyn thu xut hin trong nhm khong 2,7 t nm trc.
Nguyn liu hu c trong ny l kh giu v ng v nh lu hunh
so vi
ng v nng s*. Cha bit qu trnh a ho to ra s giu c ny nh th no,
nhng qu trnh kh lu hunh sinh hc vn tin hnh ngy nay cc vi khun
nguyn thu nh t nh. Trong h hp sunphat, vi khun nhn nng lng t qu
trn h kh cc sunphat v c (SO4* ) thnh H 2S. Nguyn t hidro nhn t cc phn
t hu c khc do sinh vt to ra. Do , cc vi khun ny cng tin hnh qu trnh
nh vi khun sinh kh m etan, nhng chng dng S 4^ nh tc nhn oxi ho (ngha
l nhn in t) thay cho CO2 .
Cc cht kh sunphat xc lp trng thi cho s tin ho ca quang hp, to ra
mi trng phong ph v H 2 S. Nh trnh by trong chng 4, dng quang hp
u tin nhn hidro t H 2S dng nng lng nh sng m t tri.

Trong h hp hiu kh, t bo thu nng lng t cc phn t glucoz

theo mt chui gm bn con ng chinh: ng phn, oxi ho
piruvat, chu trnh Krebs v chui truyn in t. Oxi l cht nhn
in t cui cng. H hp k khi chuyn giao cc din t thu c
cho cc hp cht v c khc.
5.2.2. Giai on 1: ng phn
Qu trnh chuyn ho vt cht ca cc sinh vt nguyn thu tp trung vo
glucoz. Phn t glucoz c th b phn gii theo nhiu cch, nhng sinh vt nguyn
thu pht trin nn qu trnh d ho glucoz, gii phng nng lng thc
y s tng hp ATP trong cc phn ng lin kt. l qu trnh ng phn xy
ra trong t bo cht v ko theo mt chui 10 phn ng (Hnh 5.5) bin glucoz
thnh hai phn t piruvat 3 cacbon. mi phn t glucoz thng qua s bin i
ny, t bo to ring ra hai phn t ATP nh photphoril ho mc bn th

154

CH OH

Glucoz
1. S photphoryl ho
glucz bi ATP

CH -O ig)

Hexokinaza

-o

Gluco 6 -photphat
2-3. S sp xp li, k
tip bng photphryl ho
ATP th hai
_____

|^otphoglucoizomeraza

Photphofructoknaza

ADP

7. Loi photphat cao

nng bng hai phn t
ADP to hai phn tATP
v lu li hai phn t 3PG

-0 -C H 2

Pructo 1 , 6 biphotphat
^ zo m erazVAldolaza

- 0 -CH2
. S oxi ho tip theo
bng photphoryl ho to
hai phn t NADH v hai
phn t BPG v mi
phn t ny c mt lin
kt photpht cao nng

CH2 OH

Pructo 6 -photphat

4-5. Phn t 6 -cacbon

tch nh 2 phn t 3cacbon: mt G3P, phn
t th hai bin i thnh
G3P trong phn ng khc

>

c =0

9 " Dihydroxyaceton Glyceraldehit

CH2 OH photphat
3-DhotDhat (G3P)

CHOH
C H 2 -0 -

____ ^

Triozo photphat

^'>-^1 NAD|

dfhvHrivfna7a
dehydrogenaza

1,3-Biphotphoglycerat
(BPG)

,3-Biphotphoglycerat
(BPG)

-o -c= o
CHOH
C H - 0 -<B)

Photphoglycerokinaza

O'
3-Photphoglycerat
(3PG)

3-Photphoglycert
z PG)

c =o
CHOH
CH2 - 0 -

lotphoglyceromuta;

c =0

2-PhotphoglycetBt
(2PG)

2-Photphoglycerat
(2PG)

J
:

Enolaza

10. Loi photphat cao

nng bng 2 phn tADP
to 2 phn tATP v 2
phn t pyruvat

CH2- 0 -

(D
8-9. Loi nc to hai
phn t P EP v mi phn
t ny c mt lin kt
photphat cao nng

,0 -

Photphoenopyruvat
tDhoenoi
(PEP)

HjO

Photphoernolpyruvat
(PEP)

CH2 OH
'

c =0
cII - 0 -

OH

Pyruvat kinaza
Pyruvat

H -C -0 -< g |

c =0
Pyaivat

Hlnh 5.5. Con ng itng phn. Nm phn ng u bin mt phn t glucoz thnh hai
phn t G3P. Nm phn ng tti hai bin G3P thnh plmvat

155

5.2.2.1. Schun b (priming)

Na u ca ng phn gm 5 phn ng k tip nhau, bin mt phn t
glucoz thnh hai phn t c ba cacbon - glixerandehit 3 photphat (G3P). Phn ng
ny i hi chi dng ATP. Nh vy, chng l qu trnh cn nng lng.
Bc A: s chun b ca gucoz. Ba phn ng "chun b" glucoz nh bin i n
thnh mt hp cht m c th b tch d dng thnh hai phn t b photphoril ho
ba cacbon. Hai phn ng ny cn tch ATP, nh vy bc ny i hi t bo dng
hai phn t ATP.
Bc B: tch v sp xp li. Trong phn ng u tin ca i phn ng cn li,
sn phm 6 - cacbon ca bc A tch thnh hai phn t 3 cacbon. Mt phn t l
G3P, phn t khc v sau bin thnh G3P nh phn ng th hai.

5.2.2.2. Photphoril ho mc ccht

Trong na th hai ca qu trnh ng phn, c nm phn ng bin G3P
thnh piruvat trong qu trnh to nng lng pht sinh ATP. Ni chung, ng
phn l mt lot gm 10 phn ng c enzym xc tc, trong dng mt s ATP
to ra nhiu hn.
Bc C: 8 oxi ho. T bo chuyn hai in t v mt proton t G3P cho NAD*,
to NADH. Lu rng, NAD* l mt ion v c hai in t trong lin kt cng ho
tr mi bt ngun t G3P.
Bc D: s pht sinh ATP. Bn phn ng bin G3P thnh phn t 3 cacbon
khc piruvat. Qu trnh ny lm pht sinh hai phn t ATP (Hnh 5.3). Do mi
phn t glucoz tch thnh hai phn t G3P nn chui phn ng chung to hai
phn t ATP cng nh hai phn t NADH v hai piruvat.

4 ATP (2 ATP cho mt trong hai phn t G3P trong bc D)

-

ATP (c chi dng trong hai phn ng trong bc A)

2ATP
Di iu kin khng chun trong t bo, mi phn t ATP c to ra biu
hin s thu gi khong 12 kcal (50 kJ) nng lng/mol glucoz ch khng phi 7,3
theo tnh ton truyn thng cho iu kin tiu chun. iu c ngha rng ng
phn thu gi khong 24 kcal/mol (100 kJ/mol). y khng phi lng nng lng
ln. Tng hm nng lng trong cc lin kt ho hc ca glucoz l 6 86 kcal (2870
kJ)/mol, nh vy, ng phn ch thu c 3,5% nng lng ho hc ca gIucoz.

Qu trnh ng phn gii phng ra bao nhiu nng ng th cha r nhng

bit c rng ng phn to ra ATP. Hn mt t nm trong cc giai on k kh
u tin ca s sng trn qu t, ng phn l phng thc ch yu m sinh vt
d dng tng hp ATP t phn t hu c. Ging nh nhiu con ng sinh ho,
ng phn c coi l tin ho tht li vi cc bc cui cng trong qu trnh
156

l nguyn thu nht. Do trong na th hai ca ng phn, s phn gii G3P

to ATP c th l qu trnh ban u m cc sinh vt nguyn thu s dng tng
hp ATP. S tng hp ca G3P t glucoz s phi xut hin mun hn, c l khi cc
ngun G3P khc cn kit

5.2.2.3. Mi t bo usdng ngphn

Chui phn ng ng phn l nguyn thu nht trong tt c cc qu trnh
sinh ho. ng phn khng dng oxi phn t v xy ra d dng trong mi trng
k kh. Tt c phn ng ca ng phn xy ra t do trong t bo cht, khng mt
phn ng no kt hp vi bt k bo quan hay cu trc mng no. Mi sinh vt
sng c th thc hin ng phn. Song phn ln sinh vt ang tn ti ngy nay c
th chit rt phn ln nng lng t glucoz thng qua h hp hiu kh.
Hiu qu nng lng khi khng c oxi tng i l khng ng k, vy ti sao
ngy nay ng phn vn tn ti? C quan nim cho rng tin ho l mt qu trnh
gia tng { iic r e m e n ta l p ro cess): s bin i xy ra nh s ci tin trn cc thnh
cng qua. Trong qu trnh chuyn ho d ho (c a ta b o lic m e ta b o lis m ) ng phn
lm tho mn mt tiu chun tin ho c bn: n l mt s ci tin (im p r o v e m e n t).

T bo khng th tin hnh ng phn s th bt li v cnh tranh v ch t bo

no c th ng phn th mi sng st v vt flua s cnh tranh ban u ca s
sng. Cc ci tin v sau trong qu trnh chuyn ho d ho thit lp nn s thnh
cng ny. ng phn khng b vt b trong lch trnh tin ho. Hn th, ng
phn c chc nng nh im khi u cho qu trnh chit rt thm nng lng ho
hc. Qu trnh chuyn ho tin ho nh mt tng gm cc phn ng b sung cho
nhau ging nh cc tng sn k tip ph ln bc vch ca mt to nh c. Hu nh
mi sinh vt tn ti ngy nay tin hnh ng phn nh s hi tng chuyn ho
ca qu kh tin ho sinh vt.

s.2.2.4. Kt thc chu trnh chuyn ho: 8ti snh NA*

Phn ng ca chui ng phn:
Glucoz +2ADP +2Pi + 2 NAD" -> 2 piruvat + 2 ATP + 2 NADH + 2 H" + 2 H2O
'Ta c th thy ba bin i xy ra trong ng phn: (1) glucoz b bin thnh
hai phn t piruvat, (2) 2 phn t ADP bin thnh ATP v (3) hai phn t NAD*
bin thnh NADH.

5.2.2.S. Scn thit quay vngNADH

Chng no phn t thc n c th b bin i thnh glucoz, t bo c th sn
xut nhiu ATP thc y hot ng sng bnh thng. Song lm nh vy, t
bo tch lu NADH v dng ht NAD*. T bo khng cha lng ln NAD*
ng phn tip tc, NADH phi c quay vng thnh NAD* Mt s phn t
khc ch khng phi NAD* cui cng phi nhn nguyn t hidro ly t G3P v b
kh. Hai phn t c th thc hin nhim v ch yu ny (Hnh 5.6):
157

1.

H hp hiu kh. Oxi l cht nhn in t tuyt vi. Thng qua mt lot
cht truyn in t, nguyn t hidro ly t G3P c th c chuyn cho oxi, to ra
nc. iu xy ra trong t bo ca sinh vt nhn thc khi c mt oxi. Do
khng kh c nhiu oxi, nn qu trnh ny cng c xem nh s chuyn ho hiu
kh {aerobicmetabolsm).
2. S ln men. Khi oxi khng c sn, mt phn t hu c c th nhn nguyn
t hidro thay th. S ln men ny ng vai tr quan trng trong qu trnh chuyn
ho ca phn ln sinh vt, thm ch c cc sinh vt c6 th h hp hiu kh.
C6 oxi

Khng c oxi

Hlnh 5.6. iu g xy ra vi pinivat sn phm ca ng phn?

Khi c6 oxi, plruvat b| oxi ho thnh axetyl CoA v thftm nhp vo chu trnh Krebs. Khi
thiu oxi, piruvat bj kh li nhn in t c chit rt trong ung phn v c NADH
mang theo. Khl pinivat b kh fre tip nh trong t bo cd thi sn phm l lactat. Kht

co, b

loi u tln khi pimvat v sn phm l axetaldehit v tip tc bj kh nh trong t bo nm

men, v sn phm l aiu etylic

S phn ca piruvat c to ra nh ng phn ph thuc vo qu trnh no

trong hai qu trnh ny xy ra. Con ng h hp hiu kh bt u vi s oxi ho
piruvat thnh axetyl-CoA, v sau tip tc b oxi ho theo mt lot cc phn ng gi
l chu trnh Krebs. Ngc li, con ng ln men ko theo 8 kh tt c hoc mt
phn piruvat. Chng ta s bt u bng vic l gii h hp hiu kh ri gii thch

ngn gn s ln men.
ng p h n lm p h t sin h m t lng nh ATP n h lm th a y i hay
ci to cc in k t c a p h n t glucoz. T rong ng p hn, h a i p h n
t NAD*^ b k h th n h NADH. ND* p h i c t i sin h ng
p h n ti p t c khng yu i.

158

5.2.3. Giai on 2: s oxi ho piruvat

Khi oxi c mt, s oxi ho glucoz bt u trong ng phn tip tc d u m

ng phn li piruvat. Trong cc sinh vt nhn thc, s chit rt thm nng
lng t piruvat ch xy ra bn trong ty th. T bo thu nng lng ca piruvat
theo hai cch: u tin nh oxi ho piruvat to axetyl-CoA v tip theo nh oxi
ho axetyl-CoA theo chu trnh Krebs.

5.2.3.I. Sto axetyl-CoA

Piruvat b oxi ho trong

ng phn

phn ng n "tch nhm

cacboxyl" (d e c a r b o x y la to n ) v
li mt trong ba cacbon ca

piruvat. Cacbon ny ri khi

chui phn ng di dng CO2
(Hnh 5.7a). Phn ng ny to
mt mnh hai cacbon gi l
nhm axetyl, cng nh pht
sinh mt i in t v hidro
kt hp v kh NAD* thnh

NADH. y l phn ng phc

hp ko theo ba giai on trung
gian v c xc tc bn trong
ty th nh phc h a enzym
(m ultiem ym e complex). Nh
chng 4 trnh by, phc h
t chc mt lot cc bc c
enzym xc tc sao cho sn phm
ho hc trung gian khng
khuch tn i hoc khng tri

qua cc phn ng khc. Bn

trong phc h, thnh phn
polypeptit chuyn cd cht t
enzym n enzym tip theo v
khng phng thch chng. Phc
h enzym piruvat dehidrogenaza
c chc nng loi CO2 khi
piruvat, l mt trong cc enzym
ln nht bit. N6 c 60 tiu
n v. Trong qu trnh phn
ng, nhm axetyl b loi khi

Nhm
axetyl

Hlnh 5.7. Soxi ho plruvat

(a) Phn ng phc tp ny ko ttieo s kh NAD*
thnh NADH v do n l ngun nng lng trao l
cht quan bng ca t bo
(b) Sn phm ca phn ng axetyl CoA l nguyn
liu khi u cho chu binh Krebs. Hu nh tt c phn
t tri qua qu trnh dj ho nng lng u bin
thnh axetyl CoA, v sau c chuyn cho qu trinh
tng hp cht bo hoc tng hp ATP

159

piruvat kt hp vi nhm ngoi coenzym A (CoA), hnh thnh hp cht mang tn

axetyl-CoA.
Piruvat + NAD^ + CoA -> Axetyl- CoA + NADH + CO2
Phn ng ny to mt phn t NADH v sau c dng tng hp ATP.
Song, ngha ln lao hdn so vi s kh ca NAD* thnh NADH l s to axetylCoA (Hnh 5.7b). Axetyl-CoA quan trng nh vy bi v nhiu qu trnh chuyn ho
khc nhau u to ra n. IChng ch s oxi ho piruvat - cht trung gian trong qu
trnh d ho hidratcacbon to ra n, m cn s phn gii chuyn ho imetabolic
hreakdovn) ca cc protein, cht bo v cc lipit khc cng pht sinh axetyl-CoA.
Thc th, hu nh mi phn t tri qua qu trnh d ho thu nng lng u
bin thnh axetyl-CoA. V sau, axetyl-CoA c chuyn cho qu trnh tng hp
cht bo hoc tng hp ATP, ph thuc vo yu cu nng lng ca c th.
Axetyl-CoA l tiu im chnh cho nhiu ATP d ho ca t bo nhn thc.

5.2.3.3. Vic sdng axetyl-CoA

Mc d t bo to axetyl-CoA theo nhiu cch, nhng ch mt s" gii hn qu
trnh dng axetyl-CoA. Phn ln trong l hoc nh hng hng n s d
tr nng lng (th d tng hp lipit) hoc b oxi ho trong chu trnh Krebs to
ATP. Phn no trong hai phn ny chim u th th ph thuc vo mc ATP trong
t bo. Khi ATP trong t bo cao, con ng oxi ho b c ch v axetyl-CoA c
chuyn vo qu trinh tng hp axit bo. iu gii thch ti sao nhiu ng vt
(k c con ngi) pht trin m d tr cht bo khi chng tiu th nhiu thc n
hn so vi yu cu ca c th. Ni khc i, khi mc ATP thp, con ng oxi ho
c kch thch v axetyl-CoA i vo qu trnh chuyn ho oxi ho to nng lng.
T rong giai on th u n n g lng th hai ca q u trn h d h o glucoz,
p iru v at b t c h nhm cacboxy (deeacboxy ho) to axetyl-Co,
NADH v CO,. Q u tr n h n y xy ra bn tro n g ty th,
5.2.4. G ia i o n 3: c h u t r n h K re b s

Sau khi ng phn d ho glucoz to piruvat v piruvat b oxi ho to

axetyl-CoA, giai on th ba ca qu trnh chit rt nng lng t glucoz bt u.
Trong giai on th ba ny, axetyl-CoA b oxi ho trong mt lot chn phn ng gi
l chu trnh Krebs. Cc phn ng ny xy ra trong cht nn (matrix) ca ty th
(Hnh 5.8). Trong chu trnh ny, nhm axetyl - hai cacbon ca axetyl-CoA kt hp
vi phn t bn - cacbon gi l oxaloaxetat to nn phn t su - cacbon v v
sau i qua mt lot cc phn ng oxi ho to in t trong hai phn t CO2 tch
ra, li oxaloaxetat. Tip , oxaloaxetat quay vng kt hp vi nhm axetyl
khc. Trong mi vng ca chu trnh, mt nhm axetyl mi thay th hai phn t
CO2 b mt v nhiu in t c chit rt thc y bm proton pht sinh ATP.
160

5.2.4.I. Bc tranh khi qut v chu trnh Krebs

Chn phn ng ca chu trnh Krebs xy ra trong hai bc:
Bc A: chun b. Ba phn ng chun b a phn t 6 - cacbon vo qu trnh
chit rt nng lng. u tin, axetyl-CoA kt hp vo chu trnh v v sau cc
nhm ho hc c sp xp li.
Bc B: chit rt nng lng. Bn trong su phn ng trong bc ny l phn
ng oxi ho trong in t b loi ra v mt phn ng pht sinh mt ng lng
ATP trc tip theo photphoril ho mc c cht.

5.2A.2. Ccphn ng ca chu trnh Krebs

Chu trnh Krebs gm chn phn ng k tip nhau m t bo dng chit rt
cc in t cao nng v thc y 8 tng hp ATP. Nhm hai cacbon t axetyl-CoA
thm nhp vo chu trnh im khi u, hai phn t CO2 v mt s in t b loi
ra.
Phn ng 1 ; s ngng t. Nhm hai cacbon t axetyl-CoA kt hp vi mt
phn t 4-cacbon oxaloaxetat to phn t 6 -cacbon-xitrat. Phn ng ngng t
ny l khng thun nghch chuyn giao nhm axetyl 2-cacbon cho chu trnh Krebs.
Phn ng b c ch khi nng ATP ca t bo cao v b kch thch khi ATP thp.
T , khi t bo c lng ln ATP, chu trnh Krebs b c ch v axetyl-CoA c
chuyn vo qu trnh tng hp cht bo.
Phn ng 2 v 3: s ng phn ho. Trc khi phn ng oxi ho c th bt u,
nhm hidroxin (-0H) ca xitrat phi c thay i v tr. Phn ng tin hnh theo
hai bc: u tin, phn t nc b loi khi mt cacbon, ri nc c b sung cho
cacbon khc. Kt qu l nhm -H v nhm -OH thay i v tr. Sn phm l mt
cht ng phn ca xitrat gi l izoxitrat.

Phn ng 4: s oxi ho u tin. Trong bc to nng lng u tin ca chu

trnh, izoxitrat tri qua phn ng decacboxyl ho. u tin, izoxitrat b oxi ho, to
mt i in t v c vai tr kh phn t NAD^ thnh NADH. v sau, cht trung
gian oxi ho b loi nhm cacboxyl, nguyn t cacbon trung tm b tch ra to
CO2, v pht sinh phn t 5- cacbon gi l a-ketoglutarat.
Phn ng 5; s oxi ho th hai. Tip theo, a-ketoglutarat b loi nhm
cacbonxin nh phc h a enzym ging nh phc h piruvat dehidrogenaza. Sau
khi loi CO2, nhm sucxinin cn li kt hp vi coenzym A to sucxinin-CoA. Trong
qu trnh, hai in t b chit rt v chng kh phn t NAD* khc to NADH.
Phn ng 6 : photphoril ho mc c cht. S lin kt gia nhm sucxinin 4cacbon v CoA mt lin kt cao nng. Trong mt phn ng kt hp (phn ng
cp i) ging nh phn ng xy ra trong ng phn, lin kt ny b tch ra v
nng lng c gii phng, thc y qu trnh photphoril ho guanozin
diphotphat (GDP) to guanozin triphotphat (GTP). GTP d dng bin i thnh
ATP, v mnh 4-cacbon lu li c gi l sucxinat.
161

S oxi ho pyruvat
NAD*

SHH]
Chu trnh bt u
khi n v 2C ca
acetyl-CA phn
ng vi phn t
4C (oxaoacetat)
to citrat (6 C)

M
I
CHa

Acetyl-CoA(2C)

Chu trnh Krebs

loi hidro ca
malat to phn
NADHthbav
chu nh quay ll
m.kh y___

COO"

CoA-SH

CHa

M ng ty th

H O -C -C O O '
0

Oxaloacetat (4C)

Citrat
synthetaza Ciat (6 C)

H2
I
COO'

coo
i

CH

Aconitazav

HC - C 0 0 "

()

H O -C H

Malat
dehydrogenaza
Malat (4C)

Isociat (6 C)
Isocitrat
dehydrogenaza

' n-F^^W nh5m

ccbxinto
NADM cng
vi vic gii
phng t

(g)Fumaraza
COO'

ICH
H

Puramat (4C)
ruiaiiHSi

a-Ketoglutarat(5C)

Succinat

HC
I

NAD*

a-Ketogutarat

coo
F A D -' \
Sucdnal(4C) Sucdnyl<A

Soxi ho
succinat
to FADH,

co

ucdnyl-CoA (4C)

H2

coo"

1
Hg

CH 2
I

CH2
I

coo

c=0

S loi nhm
cacboxin oxi ho
thhaitoNADH
th hai vi vic
gii phng c o ,
th hai

S -C o A

Hnh S.8 . Chu trinh Krebs

Lot phn ng ny xy ra bn trong cht nn ca ty th. phn gii hon ton mt
phn t glucoz, haf phn t axetyl CoA uc to ra do ng phn v s oxi ho piruvat,
mi phn t s phi lm mt cuc hnh trnh xung quanh chu nh Krebs. Ch cc bin
i xy ra trong khung cacbon ca phn t khl chng tin hnh thng qua chu trnh

162

Phn ng 7: s oxi ho th ba. Tip theo, sucxinat b oxi ho thnh uramat.

S bin i nng lng t do trong phn ng ny khng ln kh NAD*. Thay
vo , Aavin adenin dinucleotit (FAD) l cht nhn in t. Khng ging nh
NAD^, FAD* khng c t do khuch tn bn trong ty th. N l b phn kt hp
ca mng trong ty th. Dng kh PADHg gp in t cho chui truyn in t
trong mng.
Phn ng 8 v 9: s ti sinh oxaloaxetat. Trong hai phn ng CUI cng ca
chu trnh, mt phn t nc c b sung cho umarat, to maat. v sau malat b
oxi ho, to phn H- cacbon oxaloaxetat v hai in t kh phn t NAD* thnh
NADH. Oxaloaxetat - phn t khi u chu trnh t do kt hp vi nhm axetyl
2 - cacbon khc t axetyl-CoA v khi ng li chu trnh.

s.2.4.3. Cc snphm ca chu trinh Krebs

Trong qu trnh h hp hiu kh, glucoz hon ton b phn gii. Phn t glucoz
6 -cacbon u tin tch thnh mt i phn t piruvat 3-cacbon trong qu trnh
ng phn. V sau, mt trong cc cacbon ca mi piruvat b mt i di dng CO2
trong qu trnh bin i ca piruvat thrnh axetyl-CoA v hai cacbon khc b mt i
di dng CO2 trong qu trnh oxi ho ca chu trnh Krebs. Sn phm quan trng
m t bo li kh chuyn ho phn t gucoz thnh su phh t COz l nng
lng ca n c bo ton trong bn phn t ATP v c bit trong trng thi kh
ca 12 cht mang in t. 10 cht mang ny l phn t NADH v hai cht mang
khclPADH
C hu tr in h K re b s p h t sin h h a i p h n t ATP tr n ph&n t glucoz,
c n g s ATP n h d n g ph&n to ra . Q uan tr n g h n , c h u tr in h
K rebs v s oxi h o c a p iru v a t th u n h i u i n t c ao n&ng c
th c h u y n c h o c h u i tru y n in t d th c y t n g hp n h u
ATP hn.

5.2.5. Thu nng lng nh chit rt in t

hiu t bo dng mt s nng lng c gii phng trong qu trnh d ho
glucoz cho tng hp ATP nh th no, chng ta cn xem xt k cc in t trong
cc lin kt C-H ca phn t gucoz. Nh nu trong chng 4, khi mt in t b
tch khi mt nguyn t v chuyn cho cht khc, th nng v tr ca in t cng
c dn truyn. Trong qu trnh ny, nguyn t nhn in t b kh. S kh theo
kiu hon ton hoc khng hon ton trong trng hp n ko theo s truyn hon

ton mt in t t nguyn t n nguyn t. Song i khi s kh ch lm bin i

mc gp chung in t (degree of s h a r n g ) bn trong mt lin kt cng ho tr.
Vy iu p xy ra khi s truyn in t khng hon ton.

163

s.2.5.1. Tip cn s oxi ho kh

Qu trnh d ho glucoz l mt phn ng oxi ho kh. Cc in t cng ho tr

trong lin kt C-H ca glucoz gp chung gn bng nhau gia nguyn t c v H
do nhn ca cacbon v hidro c i lc tng ng i vi cc in t ho tr (ngha
l chng biu th tnh in m nh nhau). Song khi nguyn t cacbon ca glucoz
phn ng vi oxi to kh cacbonic th in t trong cc lin kt cng ho tr mi
chim mt v tr khc. Thay v gp chung nh nhau, in t kt hp vi nguyn t
cacbon trong glucoz dch chuyn hng n nguyn t oxi trong CO2 do oxi rt m
tnh (rt in m). Do cc in t ny b ko cch xa nguyn t cacbon, nguyn t
cacbon ca glucoz b oxi ho
(mt in t) v nguyn t oxi
b kh (nhn in t). Nh vy,
khi nguyn t hidro ca glucoz
kt hp vi nguyn t oxi to
Nng lng
cho tng
nc, nguyn t oxi ht cc in
o
t gp chung mt cch mnh
m hng n chng, oxi li b
kh v glucoz b oxi ho. Trong
phn ng ny, oxi l mt tc
nhn oxi ho (ht in t bi v
n oxi ho cc nguyn t
glucoz).

5^.5.2. Giiphngnnglng
Vn mu cht hiu s
oxi ho glucoz l nng lng ca
Nng lng thp
cc in t gp chung. Trong
mt lin kt cng ho tr, nng
lng phi c b sung loi
mt in t khi nguyn t
ging nh nng lng phi c
Hinh 5.9. S truyn in t hot ng nh th no?
s dng ln hn ln ni.
S chng minh ATP c pht sinh khi in t
Nguyn t cng c in m hn,
chuyn t mt mc nng lng n mc tip theo.
th hng ro nng lng phi
Khng ging vi s gi phng nng lng trong v n
vt qua ko in t ra cch
ca thuc n bnh thng, in t "ror trong cc ItHic
n
mc nng lng ngy cng thp, gii phng nng
xa nguyn t cng dc hn.
lng d tr vi mi bc ri khi chng ri xung cht
Song, nng lng c gii
nhn in t thp nht (ln m nht)
phng khi mt in t b dch
chuyn khi nguyn t c6 in m nh hn v gn hn vdi nguyn t c in m
ln ging y nh nng lng c gii phng khi hn ln xung ni. Trong s d
ho ca glucoz, nng lng c gii phng khi glucoa b oxi ho, in t chuyn
n gn hn vi oxi (Hnh 5.9).
164

Glucoz l thc n giu nng lng v n c nhiu lin kt C-H. Theo quan im
oxi ho kh, glucoz c nhiu in t duy tr cch xa nguyn t ca chng v tt c
u c th nng vn ng n gn hn hng n oxi. Trong h hp oxi ho,
nng lng c gii phng khng ch do nguyn t hidro ca cc lin kt C-H c
chuyn t glucoz n oxi m cn do v tr ca cc in t ho tr dch chuyn. S
dch chuyn ny c tc dng gii phng nng lng, c th c s dng tng
hp ATP.

s.2.5.3. Thu nng lngtrong ccgiai on

gii thch t bo thu nng lng trong cc bc chit rt in t nh th
no, trc ht ta hy hnh dung hnh nh c th v s t chy xng trong t. Khi
xng b t chy, lng nng lng c gii phng ra nhiu hay t ph thuc vo
lng xng cung cp cho b ch ho kh l bao nhiu. Nu tt c xng trong bnh
kh ca xe b t chy cng mt lc th nng lng c gii phng ra bng vi
nng lng c gii phng khi xng c t chy trong mt lot cc bc rt nh
bn trong xilanh. Thc th, s gii phng nng lng trong xng mi ln mt t v
xe c th dng nhiu nng lng y pittng nn xe vn ng c.

NAD"*": dng oxi ho ca nicotinamit

NADH: dng kh ca nicotinamit

Hlnh 5.10. NAD* v NADH. Dinucleotit ny c chc nng nh l "con thoi in t*

(e/ectmn shuttle) trong h hp t bo. NAO* nhn in t t cht cao phn t b d ho v b
kh thnh NADH

ng dng nguyn l tng t l gii s oxi ho glucoz trong t bo. Nu tt

c hidro c chuyn n oxi trong mt bc oxi ho, gii phng tt c nng lng
165

t do cung mt lc th t bo s thu hi rt t nng lng trong dng hu ch.

Thay vo , t b(0 coxii hio thc n nhiu bc ging nh t t chy xng nhiu
ln, mi ln niit t,. Sm Ihidro trong cc lin kt C-H ca glucoz c tch ra trong
cc giai on theo mit, lot cc phn ng c enzym xc tc trong qu trnh ng
phn v chu trnh Kebs m c trnh by trong cc phn trc. Lu rng
hidro c tch ra nihal chuyn chng n cht mang coenzym NAD* (Hnh 5.10).
Nh trnh bty tiomg cihng 4, NAD* l mt cht nhn in t rt hu hiu, lm
con thoi chuyn (tim t mang nng lng trong su^t t bo (energy - bering
electrons). Trong qni fcrmh thu nng Ing ca glucoz, NAD* hot ng nh cht
nhn in t ch yui.

s.2.5.4. Di theo in t
Khi theo di cc J)himng chit rt in t cao nng, ta c khng nhm ln
do cc bin i tomg.din tch. Lun lun di theo in t ollow the electrons).
Enzym chit rt hai Biitdno ngha l ly i hai in t,v hai proton khi gucoz v
chuyn c hai in
v nt proton cho NAD^. Proton th hai c gii phng di
dng ion hidro H* voi dbng dch xung quanh. S truyn ny bin NAD* thnh
NADH, ngha l haii in t m v mt prton dng c giao cho mt NAD* tch
in dng t NM)II - dng trung ho in.
Nng lng <(C INADH thu gi khng phi c t bo thu ly (di dng
ATP) tt c mt lm. Thaiy v c truyn trc tip n oxi, hai in t do NADH
mang theo c chiuyra dlc theo chui truyn in t nu oxi c mt. Chui phn
ng ny gm mt lott phn t ch yu l cc enzym c gn bn trong mng
trong ca ty th. NABH pin phi cc in t n nh ca chui truyn in t v
oxi thu gi chng y. V sau, oxi kt hp vi ion hidro to ra nc. mi
bc trong chui, uitt t di chuyn n mt cht mang c in m hn mt t v v
tr ca chng dch (Chtuym mt t. Do , in t vn ng xui theo mt gradient
nng ng. Tom ) (qu trnh gii phng mt tng khong 53 kcal/mol (2 2 2
kJ/mol) di iiii kdm Itiu chun. S truyn in t dc theo chui ny cho php
nng lng c (chift rTlt t t. Trong phn tip theo, chng ta s gii thch nng
ng c a vo sinh cng, thc y qu trnh tng hp ATP nh th no.
Qu trinh d ho [lucoz ko theo mt lot cc phn ng oxi ho k h gs p h n g n&ng lng nh chuyn v tr (thay i v tr) cc
in t gm VH Itguyn
hn. Do , nng lng c t bo
thu ly t plim tr glucz trong cc bc tun t, dng NAD* nh
m t c h t m aag; m t.
5.2.6. G ia i o n 4 : c b u i tru y n i n t

Cc phn t NADH v PADHa c hnh thnh trong ba giai on u ca h

hp hiu kh v rmi phn t ny mang mt i in t thu c khi NAD* v FAD"^
b kh. Cc phni tt NADH mang in t n mng tirong ty th, chng
166

chuyn in t n mt lot cc protein kt hp mng mang tn chung chui

truyn in t.

5.2.6.I. in tvn ng qua chui truyn in t

Protein u tin nhn in t
l enzym phc gn mng gi l
NADH dehidrogenaza. Mt cht
mang gi l ubiquinon tiuyn cc
in t n phc h protein xitocrom (gi l phc h bci). Phc
h ny cng vi phc h khc trong
chui hot ng nh mt bm
proton y proton qua mng ra
ngoi. Xitocrom l protein h hp
c cha nhm hem, cc vng cacbon
phc tp vi nhiu lin kt n v
i xen k v mt nguyn t st
trung tm.
Tip theo, in t c cht
mang khc thu nhn l xitocrom c
thnh phc h xitocrom oxidaza
(Hnh 5.11). Phc h ny s dng
bn in t kh phn t oxi, v

Hlnh 5.11. Phc h xitocrom oxidaza. Protein

mng ln nht vi cu trc bit hon ton.
Xltocrom oxidaza gm 11 tiu on v protein to
knh xuyn qua mng cho proton v in t

mi oxi kt hp vi hai ion hidro to nc:

O2 + 4H" + 4e -> 2H2O

Lot cc cht mang in t kt hp mng ny mang tn chung l chui truyn

in t (Hnh 5.12).
NADH gp in t ca n cho protein u tin ca chui truyn in t NADH dehidrogenaza. FADH2 lun lun gn vo mng trong ty th, cung cp in
t cho ubiquinon i vo chui truyn in t.
Vn l s c6 sn ca cht nhn in t mnh (thng l oxi) khin cho h
hp oxi ho c th tin hnh. Nh chng ta s thy trong chng 6 , chui truyn
in t c s dng trong h hp hiu kh l ging v rt c th tin ho t
chui c tn dng trong quang hp hiu kh.
C hui tru y n in t m t o t gm 5 protein k t hp m ng. in
t c p h n p h t n h NADH v FAD2 dc chuyn t p ro te in ny
n p ro te in ti p th eo dc th eo chui, ging n h m t que g tru y n
tay tro n g cuc c hy ti p sc.

167

Khong gian mng

NADH
dehidrogenaza

Phc h bc,

Phc h
ixitocrom oxidaza

H"

H"

Cht nn ty th
Hlnh 5.12. Chui truyn in t
Cc in t cao nng c thu ly t phn t b d ho c dn truyn (theo mi tn)
nh cc cht mang in t linh ng (ubiquinon - Q v xitocrom
dc theo chui gm
cc protein mng. Ba protein dng phn nng lng ca in t bm proton (mi tn) ra
ngoi cht nn v vo xoang (khong) gian mng. Cui cng, in t c chuyn cho oxi
hinh thnh nc

c -c

5.2..2. Xc lp mt gradient in ho
Trong t bo nhn thc, qu trnh chuyn ho hiu kh xy ra bn trong ty th
thc s c mt trong mi t bo. Xoang trong hay cht nn {matrix) ca ty th cha
enzym thc hin cc phn ng ca chu trnh Krebs. Khi cc in t thu c nh
h hp oxi ho chuyn dc theo chui truyn in t th nng lng chng gii
phng truyn in t ra khi vng cht nn i vo xoang ngoi m i khi c gi
l khong gian mng. Ba protein xuyn qua mng d mng trong ty th (Hnh 5.12)
thc hin qu trnh dn truyn. Dng in t trng thi kch thch (in t cao
nng) gy ra s bin i v hnh dng ca cc protein bm ny v do lm cho
chng truyn proton qua mng. Cc in t do NADH ng gp c tc dng hot
ho c ba bm proton ny, trong khi in t do FAD2 ng gp ch hot ho hai
bm proton.

s.2.6.3. Stng hp ATP: ho thm

Khi nng proton xoang ngoi tng ln trn nng vng cht nn, cht
nn tr nn m hn. Tnh m bn trong ny c tc dng ht cc proton tch in
dng v lm cho chng quay tr li vng cht nn. Nng bn ngoi cao hn c
khu)Tih hng y proton quay li nh khuch tn. Do mng tng l khng thm
cc ion, nn phn ln proton quay tr li vng cht nn u phi i qua cc knh
168

proton chuyn ho nh v Khong gian mng ^

mng trong ty th. Khi cc
proton i qua, cc knh ny
tng hp ATP t ADP + Pj bn
trong vng cht nn. v sau,
ATP c truyn bng qu trnh
khuch tn nhanh ra ngoi ty
th v i vo t bo cht ca t
bo. Do s hnh thnh ho hc
ATP {chemical formation of
ATP) c thc y bng lc
khuch tn ging vi s thm
Mng
thu, nn qu trnh ny c v

nh

s
ho
thm
(chemiosmosis) (Hnh 5.13).

trong
ty th

f41 ch nn

t M
Hnh 5.13. Ho thm

Nh vy, chui truyn in

NADH truyn in t cao nng thu c t qu trnh
t dng in t thu c trong
d ho cc cht cao phn t cho "bam proton m dng
h hp hiu kh bm mt
nng lng bm proton ra khi cht nn ty th. Kt
lng ln proton qua mng trong
qu l nng proton bn ngoi mng trong ty th
ty th. S tr li ca proton vo
tng ln lm cho proton khuch tn tr li vo cht nn
ty th. Nhiu proton i qua knh chuyn ho lln kt s
vng cht nn ty th thc y
trd li ca proton vi s tng hp ATP
qu trnh tng hp ATP nh qu
trnh ho thm. Hnh 5.14 tm tt mt cch tng qut qu trnh tng hp ATP.
Pyruvat t
t bo cht

Khong gian mng

Mng trong
ty th

[ trnh \^ e b y j

H uyn
gNin t

3. Oxy kt hp vi
proton tao nc

4. Protein khuch
tn tr li, thc dy
ng hp TP

Cht nn ty th

Protein
knh

Hinh 5.14. M hnh khi qut v s pht sinh ATP. Qu trnh ny bt u vi

piruvat - sn phm ca ng phn v kt thc VI s tng hp ATP
169

in t thu c t gIucoz c bm ra ngoi vng cht nn ty th

nh chui truyn din t. S quay li ca proton vo vng cht nn
nh khuch tn thng qua cc knh chuyn ho c lin kt vi s
pht sinh ATP.
5.2.7. T m t t h h p h i u k h

Vi cc in t thu t phn t glucoz dng cho chui truyn in t to ATP

ho thm, t bo thc s thu c bao nhiu nng lng chuyn ho?

5.2.7.1. Hiu qu l thuyt

M hnh ho thm cho rng mi bm proton c chui truyn in t hot
ho th pht sinh mt phn t ATP. Do in t t NADH hot ho ba bm v in
t t PA D H hot ho hai bm nn chng ta s cho rng mi phn t NADH pht
sinh 3ATP v FADH2 to 2ATP. Song do't bo nhn thc tin hnh ng phn
trong t bo cht v chu trnh Krebs trong ty th nn chng phi chuyn 2 phn t
NADH do ng phn to ra qua mng ty th m cn lATP cho phn t NADH. Do
, s sn xut thun ATP gim xung khong hai. T , s sn xut ATP chung
bt ngun t h hp hiu kh v mt l thuyt phi l 4 (t photphoril ho mc c
cht trong ng phn) + 30 (3 ATP t mi phn t NADH X 10 phn t NADH) +
4 (2 ATP t mi phn t FAD2 X 2 phn t PA D H - 2 vn chuyn NADH

ng phn qua mng ty th) = 36 phn t ATP (Hnh 5.15).

s.2.7.2. Hiu qu thc t

Lng ATP thc s c to ra trong t bo nhn thc trong h hp hiu kh
thp hn 36 cht t v hai l do. Th nht, mng trong ty th b "r thm" (leaky) t
nhiu vi proton, cho php mt s proton thm nhp tr li vng cht nn, khng
i qua knh pht sinh ATP. Th hai, ty th thng s dng gradient proton do qu
trnh ho thm pht sinh cho cc mc ch khc hn l tng hp ATP (nh dn
truyn piruvat vo cht nn). Do , gi tr o c thc s ca ATP do NADH v
FADH2 pht sinh th gn hn vi 2,5 cho mi NADH v 1,5 cho mi PADH. Vi cc
hiu chnh ny, thu hoch chung ATP t mt phn t glucoz trong t bo nhn
thc l: 4 (t photphoril ho mc c cht) + 25 (2,5 t mi NADH ca 10 phn t) +
3 (1,5 t mi PA D H ca 2 phn t) - 2 (vn chuyn NADH ng phn) = 30 phn
t ATP.
Khc vi ng phn, qu trnh d ho glucoz nh h hp hiu kh rt hiu
qu. H hp hiu kh trong t bo nhn thc thu c khong (12x30):686 = 52%
nng lng c trong glucoz (Theo so snh, mt t in hnh ch bin i khong
25% nng lng trong xng thnh nng lng hu ch). Tnh hiu qu ca h hp
oxi ho nng lng thu c xc lp mt gii hn t nhin v di cc i ca
cc chui thc n.
170

Hiu qu cao ca h hp hiu kh l mt trong nhng nhn t ch yu c tc

dng kch thch s tin ho ca cc sinh vt d dng (heterotrophs). Vi c ch ny
tng hp ATP, cc sinh vt khng quang hp c th c quyn thu nhn nng
lng chuyn ho t qu trnh phn gii oxi ho cc sinh vt khc. Chng no mt
s sinh vt vn thu gi nng lng nh quang hp th cc sinh vt khc ch c th
tn ti nh n sinh vt quang hp.
t bo nhn thc, h hp oxi ho to ra khong 30 phn t ATP t
mi phn t glucoz. Lng ATP ny chim hn mt na nng lng
trong cc lin kt ho hc ca glucoz.

Glucoz
E)ng
phn
2

PdH

m si

m si
2 |f a d h , |

Tng hiu qu iun ATP = 36

Hnh 5.15. Hiu qu l thuyt ATP thu c t glucoz nh h hp hiu kh tng cng 36
phntATP

171

5.2.8. S iu chnh h hp hiu khi

Khi t bo c lng ATP
phong ph, cc phn ng ch yu
ca ng phn, chu trnh Krebs
v phn gii axit bo b c ch,
lm chm s hnh thnh ATP. S
iu chnh cc con ng sinh ho
ny do mc ATP l mt th d v
s c ch lin h ngc. Ngc
li, khi mc ATP trong t bo
thp, mc ADP cao v ADP hot
ho cc enzym trong con ng d
ho hidratcacbon kch thch s
hnh thnh ATP nhiu hn,

cch

Glcoz

fA D P

Fructoz 6-photphat

Hot ho

-^ h o s p h o fru c to k in a z ^
Fructoz 1,6-bisphotphat

Hot ho

Piruvat

Piruvat decacboxylaza

S iu tit qu trnh d ho
Acetyl-CoA
glucoz xy ra hai im chnh
ca con ng d ho (Hnh 5.16).
im iu tit trong ng phn
l enzym photphofructokinaza m
xc tc phn ng 3, bin fructoz
photphat
thnh
fructoz
bisphotphat ructoz diphotphat).
y l phn ng u tin ca
Chui uyn
ng phn m d thun nghch,
din t
chuyn giao c cht cho chui
v ho thm
ng phn. Mc ADP cao so vi
ATP (biu hin 8 cn thit
bin nhiu ADP thnh ATP) c
Hlnh 5.16. S iu tit ca qu trnh d ho glucoz.
tc
dng
kch
thch
Mc tng l ca ADP v ATP c 6 tc dng iu tit
photphofructokinaza, chuyn giao
con ng d ho d hai im ch yu: phn ng chuyn
nhiu ng hn cho con ng d
giao ca ung phn v chu trinh Krebs
ho, nh vy lm cho mc xitrat
thp hn (biu hin chu trnh khng hot ng ht cng sut v cn lng u vo
nhiu hn). im iu tit ch yu trong oxi ho piruvat xy ra bc chuyn giao
{committings te p ) trong chu trnh Krebs vi enzym piruvat decacboxilaza. N b c
ch do mc NADH cao (mun ni khng cn nhiu hn).
im iu tit khc trong chu trnh Krebs l enzym xitrat sinthaza, xc tc
phn ng u tin bin oxaloaxetat v axetyl-CoA thnh xitrat. Mc ATP cao c
ch xitrat sinthaza (cng nh piruvat decacboxilaza v hai enzym khc ca chu
trnh Krebs) c ch (lm ngng) con ng d ho.
Mc tdng i ca ADP v ATP iu chnh qu trnh d ho glucoz
cc phn ng chuyn giao ch yu.
172

5.3. Qu trnh d ho ca protein v cht bo c th to nhiu nng

lng

Glucoz khng phi l thc n duy nht

Chng ta d tho lun nhiu v h hp oxi ho glucoz m sinh vt nhn c
c cht t qu trnh tiu ho (phn gii) hidratcacbon hoc t quang hp. Cc phn
t hu c khc c bit protein v cht bo cng l ngun nng lng quan trng
(Hnh 5.17),
I

________ __________

s p h n g i i
cao plian t

!Axit nuclcic

Protein I

'

---------------,

Polsaccarit

._____ _J . ____

Lipit v
cit bo I

1 .................. l
Khc k i i

tr cteb o

I
\

.............
_ I.
' Axi bo ;
Nucleotil .z 1 Axil auiin I I
S loai
loi nhm
iihu amiK
aiiK ng phn
p- oxi ho
1
s
pl

n g

I Piruval

! lp

AcetUCoA

oxi ho

"TiTsarpIiu

cu cng
cia quil triih
chuyn
h o

Hnh 5.17. T bo chit rt nng lng ho hc nh th no?

Mi t bo nhn thc, nhiu t bo tin nhn chit rt nng lng t phn t hu c nh
ho chng. Giai on u ca qu trinh ny l s phn gii cc cao phn t thnh cc
hp phn ca chng, to t nng lng. Giai on th hai l h hp oxi ho hay h hp hiu
kh chit rt nng lng ch yu di dng cc in l cao nng, to nc v kh
cacbonic
0X1

5.3.1. H hp t bo vi protein
Lc u. protein b phn gii thnh cc axit amin thnh phn. Nhm bn cha
nit (nhr amin) b loi khi mi axit amin trong qu trinh loi nhm amin
(deamin ho: d ea m in a tio n ). Mt lot cc phn ng bin chui cacbon thnh phn t
tham gia trong ng phn v chu trnh Krebs. Th d, alanin bin thnh piruvat,
173

glutamin thnh a-ketoglutarat v atpactat thnh oxaloaxetat. v sau, cc phn

ng ca ng phn v chu trnh Krebs chit rt cc in t cao nng t cc phn
t ny v a chng vo hot ng to ATP:

5.3.2. H hp t bo vi c h t bo
Cht bo b phn gii thnh axit bo v glixeron. Phn ui ca axit bo
thng c 16 lin kt -CH2 hoc nhiu hn v nhiu nguyn t hidro phn ui
di cung cp nhiu in t cao nng thu nhiu nng ing ny. Cc axit bo b
oxi ho trong phn cht nn ca ty th. Cc enzym trong cht nn loi nhm axetyl
2 cacbon khi u cu ca mi ui axit bo cho n khi ton b axit bo bin
thnh cc nhm axetyl. v sau mi nhm axetyl kt hp vi coenzym A to
axetyl-CoA. y l qu trnh P-oxi ho (p-oxidation).
Qu trnh d ho cc axit bo to ra bao nhiu ATP? Chng ta hy so snh mt
axit bo c 6-cacbon gi nh vi phn t glucoz 6-cacbon. Trong phn trc, chng
ta trnh by rng khi phn gii oxi ho glucoz 6-cacbon to ra khong 30 phn
t ATP trong t bo nhn thc, y, hai vng ca P-oxi ho s bin axit bo
thnh 3 phn t axetyl-CoA. Mi vng cn mt phn t ATP chun b qu trnh,
nhng n cng to ra mt phn t NADH v mt FAD2. Cc phn t ny cng to
ra bn phn t ATP (to ra 2,5 ATP/NADH v 1,5 ATP/ PADHa). S oxi ho mi
axetyl-CoA trong chu trnh Krebs cui cng to ra thm 10 phn t ATP. Mt cch
tng qut, hiu qu ATP ca mt axit bo 6 -cacbon s xp x 8 ATP (t hai vng Poxi ho) -2ATP ( chun b cho hai vng ) +30ATP (t oxi ho ba axetyl-CoA) =
36 phn t ATP. Do , qu trnh h hp vi axit bo 6-cacbon to hn 20% ATP 80
vi h hp bng glucoz. Hn na, mt axit bo c ny s c trng lng nh hn hai
phn ba glucoz, nh th mt gam axit bo cha hn hai ln kilocalo 80 vi mt
glucoz. iu gii thch t sao cht bo l phn t d tr nng lng tha trong
nhiu loi ng vt. Nu nng lng tha c d tr thay hidratcacbon th c th
ng vt s s hn.
P ro te in , c h t bo v phn t hu c k h c c n g c ch u y n ho
th u n n g Ing. Cc axit am in c a p ro te in b oi nhm am n u
ti n , tro n g k h i cc cht bo phi tr i q u a q u tr in h gi l P-oxi ho.

5.4. T bo c th chuyn ho thc n khng c oxi

* S ln men
Khi thiu oxi, qu trnh chuyn ho hiu kh khng th xy ra v t bo phi
da vo ng phn to TP. Di cc iu kin , nguyn t hidro pht sinh
nh ng phn c chuyn cho cc phn t hu c trong qu trnh gi l s ln
men ermentation).
174

Vi khun tin hnh nhiu loi ln men, tt c u s dng mt s dng phn

t hu c nhn nguyn t hidro t NADH v do quay vng NAD*:
Phn t hu c + NADH

Phn t hu c b kh + NAD*

Thng th phn t hu c b kh l mt axit hu c nh axit axetic, axit

butyric, axit propionic hoc
axit lactic hoc ru etylic.
Olucoz
H
I

6.4.1. S ln men ru etyc

T bo nhn thc ch c
mt vi loi ln men. Mt loi
xy ra trong nm n bo gi
l nm men (yeast), phn t
nhn hidro t NADH l
piruvat - sn phm cu cng
ca ng phn. Cc enzym
ca nm men loi nhm CO2
cui cng khi piruvat thng
qua phn ng loi nhm
cacboxyl (decacboxyl ho) to
phn t 2 -cacbon gi l
axetaldehit. CO2 c gii
phng ra lm cho bnh m
phng n, trong khi bnh m
sn xut khng c nm men
(bnh m khng c men) thi
khng phng ln, axetaldehit
nhn nguyn t hidro t
NADH, to NAD* v ru
etylic (ethanol = ethyl alcohol).
x)i ln men c bit ny c
con ngi quan tm nhiu v
n l ngun ru etylic trong
ru nho v bia (Hnh 5.18).
Ru etyic l sn phm ph
ca qu trnh ln men v thc
s c i vi nm men khi n
t nng khong 1 2 % v bt
u lm cht nm men. iu
ny gii thch ti sao nlu nho

ln men t nhin ch cha

khong

1 2 % ru

etylic.

2ADP-

-2NAI>-

H --O H
I
H3
2Btano

^T
H
1

Or

=o

=o
I
c=0

CH,
2 Axetaldehit

_
Pyruvat

CH,
Ln men riu etylc ong nm men
Glucoz

=o
1

2ADP

H - -O H

CH,

2NAET-

2Lactat

2 Pinivat

Ln men axit lacc ong t bo c

Hlnh 5.18. Ru nho c sn xut nh th no?
Nm men tin hnh bin l plruvat thnh n/u etylic.
Qu trnh ny xy ra t nhin bong chm nho li men
trn cy nho, cng nh trong chum (thng) ln men cha
chm nho bj nghln nt. Khl nng ruu etylic t
1 2 %, tc dng c ca niu git nm men, phn cn li
l ruu nho. T bo c bin pimvat ttinh lactat, c
hn aru etylic. Song lactat vn c hi to cm
gic au n trong c khi luyn tp nng do 0X1 toong c
b tiu th ht

175

5.4.2. S ln m en axit lactic

Phn ln t bo ng vt ti sinh NAD* m khng tri qua phn ng loi nhm

cacboxyl. Th d, cc t bo c dng enzym gi l lactat dehidrogenaza truyn
nguyn t hidro t NADH quay li piruvat do ng phn tp ra. Phn ng ny
bin piruvat thnh axit lactic v ti sinh NAD* t NADH. Do , phn ng tip cn
chu trnh chuyn ho, cho php ng phn tip tc chng no sn c glucoz. Mu
tun hon loi b lactat tha (dng ion ca axit lactic) khi c, nhng khi s loi b
khng th theo kp vi s tng hp, th axit lactic tch lu gy nhiu vi chc nng

ca c v khin cho c mt mi.

S ln m en xy ra khi thiu oxi, in t b t ngun t qu trn h phn

gii dng phn ca glucoz c chuyn cho mt phn t hu c, ti

sinh NAD* t NADH.

TM TT KHI NIM
5.1. T b o t h u n n g l n g tr o n g c c i n k t h o h c

Sinh vt thu nng lng A T P t quang hp hoc h hp t bo - l qu

trnh trong t bo thu nng lng ho hc trong cc lin kt ca cc
phn t hu c.
Cc phn ng ca h hp t bo l phn ng oxi ho kh (redoxreaction). Cc
phn ng i hi s cung cp nng lng th lin kt vi s phn gii ATP v
gii phng nng lng t do.
C ch ca h hp t bo thng c iu chnh bi tp tnh in t (electron
behavor) m li b nh hng do s c mt ca cc cht nhn in t. Mt s
nguyn t nh oxi rt m tnh v do c tnh cht nh l tc nhn oxi ho
tt.
5.2. H h p t b o o x i h o cc p h n t th c n .

Trong t bo nhn thc, h hp oxi ho ca piruvat xy ra bn trong cht nn

ty th.
Cc in t pht sinh trong qu trnh c chuyn dc theo chui truyn in
t - mt lot gm cc cht mang in t trong mng ty th.
Mt s nng lng c gii phng nh chuyn in t dc theo chui truyn
in t c t bo s dng bm proton ra ngoi xoang cht nn ty th. S
thm nhp tr li ca proton vo cht nn c lin kt vi s tng hp ATP.
Qu trnh ny c gi l s ho thm, v sau ATP ri khi ty th bng c
ch khuch tn nhanh acilitateddiffusion).
176

5.3. Qu trn h d h o p ro tein v cht bo c th to nhiu n n g lng

S d ho ca cc axit bo bt u vdi P-oxi ho v cung cp nhiu nng lng

hn qu trnh d ho hidratcacbon.
.4. T bo c th chuyn ho thc n khi khng c oxi
Ln men l qu trnh h hp k kh, dng mt phn t hu c thay cho oxi
nh cht nhn in t cui cng

Ln men xy ra trong vi khun cng nh trong t bo nhn thc, gm nm

men v t bo c ca ng vt.
GII THCH THUT NG
1.

Aerobic respiration (h hp hiu kh); l h hp t bo cn oxi nh cht nhn

in t. N bao gm s oxi ho piruvat v chu trnh Krebs.

2.

Anaerobic respiration (h hp k khO: l h hp t bo trong t bo dng

cht nhn in t v c hn l oxi. N bao gm ng phn.

3. Cellular respiration (h hp t bo): S oxi ho cc phn t hu cd bn trong

t bo v c tin hnh nh thu in t cao nng, c s dng tng hdp

ATP v v sau chuyn cho phn t v c. Trong h hp hiu kh, cht nhn in
t cui cng l oxi. H hp t bo bao gm c h hp hiu kh v h hp k kh.
4. Chemiosmosis (s ho thm): S chuyn cc in t cao nng dc theo chui
truyn in t lin kt vi s bm proton qua mng v s quay tr li ca
proton vo pha ban u ca mng thng qua knh pht sinh ATP. C th hn,
khi proton c bm ra ngoi cht nn ty th, cht nn tr nn m tnh hn v
do to ra mt lc pht ng cc proton tch in dng quay tr li cht
nn. Phn ln proton quay tr li u thng qua knh to ATP. Ho thm l s
tng hp ATP c thc y nh s vn ng ca proton hng n cht nn
tch in m.
5. Electronegativity (tnh m in - m in); l s ht cc in t ho tr m
nguyn t c. Mt s nguyn t c tnh m in mnh hn nhiu so vi nguyn
t khc. Cc nguyn t c tnh m in cao nh oxi th ht in t mnh v l
tc nhn oxi ho tt.
6.

Electron transport Chain (chui truyn in t): Mt lot cc cht mang in

t bn trong mng trong ty th m thng qua in t thu c t cc phn t
hu c c chuyn qua. Mi cht mang th m tnh hn so vi cht ng trc.
Ba cht mang thc hin s truyn in t nh bm proton qua mng t cht
nn ca ty th n t bo cht.
177

7. Permentation (s ln men): l qu trnh m nh t bo (khi thiu oxi)

chuyn hidro do ng phn pht sinh cho mt phn t hu c. C th nh
ngha ln men nh l con ng to ATP theo kiu khc c tin hnh mt s'
t bo khi thiu oxi, trong cht nhn in t cui cng l mt phn t hu c.
8.

Maximvim effciency (hiu qu cc i); l s cc i phn t ATP c pht

sinh nh qu trnh oxi ho mt cht so vi nng lng t do ca cht . Trong
sinh vt th hiu qu thc thng nh hn hiu qu cc i.

9. Oxidation (s oxi ho): l s mt in t. Trong h hp t bo, cc in t cao

nng b loi khi phn t thc n v oxi ho chng.
10. p-oxidation (P-oxi ho): axit bo, n v c bn ca cht bo, b oxi ho nh loi
mnh 2 -cacbon v hng chng vo con ng t bo cho h hp hiu kh.
11. Photosynthesis (quang hp): S pht sinh ATP ho thm v cc phn t hu
c phc tp c cung cp nng lng nh nng Ing t nh sng mt tri.
12. Redox reaction (phn ng oxi ho kh): S cho v nhn in t t nguyn t
ny n nguyn t khc. Nng lng ca in t b gim nu n vn ng
n mc nng lng thp hn trong nguyn t nhn.
13. Substrate - level phosphorylation (photphori ho mc c cht): S pht
sinh ATP nh truyn trc tip mt nhm photphat n ADP t phn t b
photphoril ho khc.

CU HI N TP
1.

S khc nhau gia 8nh vt t ddng v d dng? Mi nhm nhn nng lng
nh th no?

2. S khc nhau gia s tiu ho {digestion) v s d ho? Dng no cung cp nhiu

nng lng hn?
3. Photphoril ho mc c cht l g? Trong qu trnh oxi ho glucoz trong t bo
nhn thc, qu trnh ny xy ra lc no?
4. ng phn xy ra u trong t bo nhn thc? Phn t no c to ra t
nguyn t cacbon trong glucoz nh l kt qu ca ng phn? S to thun
ATP trong ng phn l g v ti sao n khc vi s lng phn t ATP c
tng hp trong ng phn?
5. ng phn b nh hng nh th no do thiu oxi? Nu glucoz c cung cp
khng hn ch thi cu cng nhn t no hn ch ng phn?
6.

Nh hai c ch no c th NADH bt ngun t ng phn b bin i tr li

thnh NAD*? Cc cd ch ny khc nhau nh th no theo cht nhn in t v
s sn xut ATP? C ch no l hiu kh v c ch no l k kh?

178

7. Trong t bo nhn thc, qu trnh oxi ho piruvat, chu trnh Krebs v s truyn
in t dc theo chui truyn in t xy ra u? S phn t ATP cc i theo
l thuyt c to ra trong qu trnh oxi ho phn t glucoz nh cc qu trnh
ny l bao nhiu? Ti sao s phn t ATP thc s c to ra thng thp hn
s cc i l thuyt?
8.

S chuyn cc in t dc theo chui truyn in t lin kt nh th no vi s

to ATP? S lin kt ny b nh hng nh th no do s thiu oxi?

9. Axetyl-CoA c to ra nh th no trong qu trnh oxi ho hiu kh ca

hidratcacbon v iu g xy ra vi n? N c to ra nh th no trong qu
trnh oxi ho hiu kh cc axit bo v v sau iu g xy ra vi n?
10. So snh nh th no lng ATP c to ra nh oxi ho hiu kh glucoz v oxi
ho hiu kh cc axit bo? Loi cht no cha nhiu nng lng hn trn c s
so snh trng lng {onaper~weight basis)?

CU HI SUY LUN
1 . Nh

gii thch trong chng 1 , ty th c xem l tin ho t vi khun m

b ng ho (engue) v sng cng sinh bn t bo nhn thc nguyn thu.
Ti sao t bo nhn chun tn ti ngy nay khng b qua ty th, t tt c gen
ty th trong nhn v thc hin tt c chc nng chuyn ho ca ty th bn
trong t bo chV

2.

Ti sao cy thng d tr nng lng tha di dng hidratcacbon ch khng

phi cht bo?

179

Chng VI

QUANG HP

KHI NIM CHUNG

6.1. Quanghpxy ra tronglclp

Lc lp nh mt b my quang hp
Quang hp trong thc vt xy ra bn trong mt h thng mng c t chc cao
6.2. N g h i n c u v q u a n g hp: m t c h n g n g th c n g h i m

Vai tr ca t v nc: sinh kh tng ch yu bt ngun t quang hp, ch

mt lng nh ly t t. Trong cy, nc cung cp in t dng kh kh
cacbonic.
Vai tr ca nh sng: quang hp l mt qu trnh gm hai giai on, ch giai
on u cn nh sng.
6.3. S c t h p th n n g l n g t n h s n g m t tri

L sinh ca nh sng. Nng lng nh sng mt tri tn ti trong cc "b" gi

l photon c sc t' hp th.
Dip lc v sc t vng. Cc sc t quang hp hp th nh sng v thu nng
lng ca nh sng.
T chc sc t thnh cc quang h: Quang h l mt nhm c t chc, c chc
nng hi t nng lng nh sng vo mt phn t sc t y bt ra mt
in t cao nng.
Quang h bin nng lng nh sng thnh nng lng ho hc nh th no?
Mt s' vi khun da vo quang h n to ATP. Thc vt dng hai quang
h theo mt dy, pht sinh nng lng kh NADP* v pht sinh ATP
Hai quang h ca thc vt hot ng' vi nhau nh th no? Quang h II v I
thc y tng hp ATP v NADPH cn to cc phn t hu c.

181

6.4. Tbodngnnglngvlckhdophnngsngthuc toccphn

thuc
Cc phn ng khng ph thuc nh sng. Thc vt dng ATP v NADPH
cung cp nng lng tng hp ribuloz diphotphat - cht lin kt vi CO2
trong qu trnh c nh cacbon.
H hp sng: Enzym xc tc c nh cacbon cng c vai tr xc tc gii
phng CO2 c bit nhit cao.
Chu trnh nng lng; Cc con ng x l nng lng ca lc lp v ty th
c lin quan mt thit vi nhau.
Quang hp l qu trnh dng nng lng nh sng lin kt to oxi phn t
vi s cc nh CO2 v nh duy tr hot ng hu nh ton b s shg trn hnh
tinh ny. Trong chng 5, chng ta gii thch t bo chit rt nng lng ho
hc t phn t thc n v dng nng lng nh th no cung cp nng lng
cho cc hot ng ca chng. Trong chng ny, chng ta s xem xt quang hp qu trnh m nh thc vt, to lc v khun lam... hp thu nng lng t nh
sng mt tri v dng n tng hp cc phn t thc n giu nng lng ho hc
nh th no.

6.1. Quang hp xy ra trong c lp

* Lc p nh mt b my quang hp
nh sng mt tri cung cp nng lng cho s sng. Nng lng m phn ln
t bo sng s dng ch yu bt ngun t mt tri do thc vt, to v vi khun hp

th thng qua qu trnh quang hp. Tnh a dng ca s sng ch c th do hnh

tinh ca chng ta tm mnh trong dng nng lng t mt tri hng v qu t.
Mi ngy, nng lng bc x n qu t bng khong mt triu qu bom nguyn
t c Hiroshima. Quang hp hp thu khong 1% ngun nng lng khng l ny,
dng n cung cp nng ng, thc y ton b s sng trn hnh tinh.

6.1.1. Qu trnh quang hp

Quang hp xy ra trong nhiu loi vi khun, to, l cy v i khi trong thn
ca cy xanh. Hnh 6.1 m t mc t chc trong l cy. T chng 1, ta bit rng
t bo ca l cy cha cc bo quan gi l lc lp thc hin qu trnh quang hp.
Khng c cu trc no khc trong cy c th tin hnh quang hp. Quang hp xy
ra trong ba giai on: ( 1 ) hp thu nng lng t nh sng mt tri; (2 ) dng nng
lng to ATP v lc kh dng mt hp cht gi l NADPH; (3) dng ATP v
NADPH cung cp nng lng tng hp cc phn t hu c t CO2 trong khng
kh (s Cnh cacbon).
182

.Tng cutin
-Biu b
Tng tht l

Mng ngoi

Lc lp

Mng ong

Khng
bo

Granum
Nhn

Cht nn

Vch
t bo

Thilacoit

Hlnh 6.1. Con ng vo l

L cy c mt tng dy cc t bo (tng tht l) cha nhiu lc lp. Cc thilacoit dt trong
lc lp chng thnh cc ct gi l grana (t s It: granum). Phn ng ph thuc nh sng
xy ra trn mng thilacoit

183

Thilacoit

Cht nn lc lp

Hnh 6.1 (tip), v pht sinh ATP v NADPH cung cp nng lng cho cc phn ng
khng ph thuc nh sng. Cht nn bn trong lc lp l stroma (cht nn ca lc lp) cha
cc enzym thc hin cc phn ng khng ph thuc nh sng

Hai qu trnh u xy ra khi c nh sng v thng c gi l cc phn ng

ph thuc nh sng (light - dependent reactions). Qu trnh th ba (to phn t
hu cd t CO2 kh quyn) c th xy ra c hoc khng c nh sng v bao gm cc
phn ng khng ph thuc nh sng ca quang hp (light - independent reactions).
Chng no ATP v NADPH sn c trong t bo th cc phn ng khng ph thuc
nh sng xy ra mt cch d dng khi khng c nh sng cng nh khi c nh
sng.

Phng trnh n gin sau y tm tt qu trnh chung ca quang hp:

6

CO2 + 6 H2 O + nh sng -> C6I2 0 6 + 6 H2 O + 6 O2

Kh cacbonic nc

glucoz

nc

oxi

6.1.2. Phn bn trong c lp

Mng bn trong (mng ni b) ca lc lp c t chc thnh ti gi l
thilacoit (thylakoids) v thng c v s thilacoit chng ln nhau to thnh ct gi
l grana. Bao quanh h mng thilacoit l cht na lng gi l cht nn (stroma).
Trong cc mng ca thilacoit, sc t' quang hp nhm vi nhau thnh mng gi l
quang h (photosystem). Nh vy, mng thilacoit l h mng th ba ca lc lp m
ty th khng c.

Mi phn t sc t'bn trong mng ca quang h c kh nng hp th photon.

Mng li gm cc protein gi sc t tip xc nhau. Khi nh sng c bc sng
thch hpva p vo mt phn t sc t' trong quang h dn n s kch thch v
184

tc ng kch thch ny c dn truyn t mt phn t dip lc ny n phn t

tip theo. in t b kch thch {excitedelectron) khng c dn truyn theo c ch
vt l. l nng lng (energy) c chuyn t mt phn t n phn t khc. C
th so snh dng truyn nng lng ny ging nh c ph bng u tin chy bi
trong tr chi bi a. Nu gy chi bi a nh (p) vo im vung gc qu bi ca dy
tam gic gm 15 bi, hai bi gc xa ca hnh tarn gic bay i nhng khng c qu bi
no gia vn ng. Nng lng truyn qua cc V'n bi gia n vin bi xa nht.
Cui cng nng lng n mt phn t dip lc ch yu bn trong trung tm
phn ng - dip lc nm st vo mt protein lin kt mng. Ti y nng lng
c truyn di dng mt in t b kch thch cho protein ny v t , in t b
kch thch c chuyn n mt lot cc protein lin kt mng khc v a nng
lng vo hot ng to ATP v NADPH m tng hp cc phn t hu c. Do ,
quang h hot ng nh mt mng anten ln c tc dng khuch i cng sut ca
cc phn t sc t c th thu thp nh sng.
Cc phn ng ca quang hp xy ra bn trong mng thilacoit ca
lc p trong t bo l.

6.2. Nghin cu v quang hp: mt chng ng thc nghim

6.2.1. Vai tr ca t v nc
Nhng nghin cu nh th no v quang hp l cu chuyn th v trong khoa
hc v n c vai tr nh s nhp mn tt cho qu trnh phc tp ny. Cu chuyn
bt u hn 300 nm trc, tin s ngi B, Jan Baptista Van Helmont (1577 1644), thit k mt th nghim n gin nhng cn thn. T thi , ngi Hy
Lp bit rng cy nhn thc n t t, theo ng nguyn vn cy ht thc n
qua r ca chng. Van Helmont ngh ra mt cch n gin th nghim
tng. ng trng mt cy liu nh trong chu t sau khi cn cy v t. Cy
sinh trng trong chu qua mt s nm, trong thi gian Van Helmont ch thm
nc cho cy. Sau nm nm, cy ln ln nhiu v trng ng cy tng n 74,4
kg. Song, tt c khi tng thm ny khng th bt ngun t t, bi v t trong
chu cn c ch nh hn 57 gam so vi t c 5 nm trc. Vi th nghim ny,
Van Helmont chng minh rng vt cht ca cy khng ch c to ra t t.
ng kt lun khng ng rng khi lng tng ca cy ch yu do nc b sung.
Hn mt trm nm qua k t th nghim ca Van Helmont, cu chuyn tr
nn r rng hn. Bng chng ch yu c nh khoa hc ngi Anh Joseph Priestly
b sung trong cc nghin cu tin phong v tnh cht ca khng kh. Ngy 17-81771, Priestly bt ng tm ra phng php lm phc hi khng kh b tn hi do
s chy ca ngn nn. ng cho mt cnh nh cy bc h cn sng vo khng kh
trong c mt ngn sp nn chy ht v thy rng vo ngy th 27 cng
185

thng, ngn nn khc c th chy trong cng khng kh . Bng cch ny, hnh
nh cy xanh c kh nng phc hi khng kh. Priestly tm thy rng trong khi
chut khng th th trong khng kh m ngn nn tt, thi khng kh "c phc
hi" nh cy xanh khng phi hon ton khng thun li cho chut. Theo , bng
chng ch yu l ch cy xanh b sung mt ci gi cho khng kh.
Cy xanh "phc hi" khng kh nh th no? Hai mi lm nm sau, thy
thuc ngi H Lan Jan Ingenhousz gii quyt vn rc ri ny. Sau mt s
nm lm vic, Ingenh0 U8 Z ti hin v m rng ng k nhng kt qu th

nghim ca Priestly, chng minh rng khng kh ch c phc hi khi c nh sng

mt tri v ch nh l xanh ca thc vt m khng phi bng r ca n. Theo
Ingenhousz, cc phn xanh ca cy thc hin mt qu trnh (m ngy nay chng ta
gi l quang hp) l dng nh sng mt tri tch kh cacbonic (CO2) thnh
cacbon v oxi. ng cho rng oxi c gii phng di dng kh O2 vo khng kh,
cn nguyn t cacbon kt hp vi nc to ra hidratcacbon. iu ngh ca ng

l s phng on tt: v sau cc nh ho hc tm thy rng t l ca nguyn t

cacbon, oxi v hidro trong cc hidratcacbon thc s vo khong mt nguyn t
cacbon/phn t nc (nh thut ng hidratcacbon nu ln). Phn ng tng qut
ca Ingenhousz cho quang hp l:
CO2 + HO + nng lng nh sng -> (CH2O) + O2
Gi thuyt rt c l v n gin ny c tha nhn nh l s thc trong
hn mt trm nm, nhng nh thng xy ra trong khoa hc, cui cng quang hp

c minh chng khng phi n gin nh vy.

ngha thc s ca phng trnh dn gin nhng d lm ln ca Ingenhou8z
c gii quyt vo u nhng nm 1930 nh C.B.Van Niel trng i hc
Stanford - ngi nghin cu quang hp trong vi khun. Mt trong cc loi vi
khun m Van Niel nghin cu l vi khun liu hunh ta th khng gii phng oxi
trong quang hp, thay vo chng bin hidro sunphit (H2S) thnh ht nguyn t
lu hunh tinh khit tch lu trong c th chng. Qu trnh m Van Niel nghin
cu l:
CO2 + 2 H2S + nng lng nh sng -> (CHjO) + H2O + 2 S
S tng t r rng gia phng trnh ny v phng trnh ca Ingenhousz
dn Van Niel nu gi nh cho rng qu trnh khi qut ca quang h^p l:

CO2 + 2H2A + nng lng nh sng

(CH2 O) + H2 O + 2A

Trong phng trnh ny, cht HgA c chc nng nh mt cht cho in t.
Trong quang hp c cy xanh tin hnh, H2A l nc, trong khi vi khun lu

hunh ta, HA l hidro sunphit. Sn phm A bt ngun t s tch (phn ly) ca

186

HA. Do , O2 c to ra trong quang hp ca cy xanh bt ngun t s tch nc

khng phi tch kh cacbonic.
Khi cht ng v phng x c s dng ph bin trong sinh hc vo u
nhng nm 1950, ngi ta tin hnh kim tra gi nh c tnh cch mng ca
Van Niel. Cc nh ghin cu nghin cu quang hp trong cy xanh c cung
cp vi nc c 0 *v h tm thy rng 0 *nh du xut hin trong kh oxi ch
khng phi trong hidratcacbon ng nh Van Niel d on:
CO2 + 2 H2 0 + nng lng nh sng -> (CH2O) + H2O + 2
Trong to v cy xanh, hidratcacbon c quang hp thng l ng glucoz c
su cacbon. Do , phng trnh cn bng hon ton cho quang hp trong cc c th
ny tr thnh:
6 CO2 + I 2 H2O + nng lng nh

sng

CeHOe + 6 O2 + H2O

Ban u, cc nh nghin cu cho rng t b sung khi lng cho

cy ang sinh trng. Van Helmont ch ra iu l khng dng.
Van Niel pht hin ra rng quang hp phn ly phn t nc, kt
hp nguyn t cacbon ca kh cacbonic v nguyn t hidro ca nc
thnh phn t hu c v gii phng khi oxi.
6.2.2. Vai tr c a nh sn g

C phng trnh ban u v quang hp ca Ingenhousz v phng trnh c

tha nhn gn y ca Van Niel u bao gm mt nhn t m chng ta cha bn
n, l nng lng nh sng. nh sng ng vai tr nh th no trong quang
hp? Vo u th k 20, nh sinh l thc vt ngi Anh F.F.Blackman bt u nu
ln vn ny. Nm 1905, Blackman nu kt lun mi rng quang hp l mt qu
trnh hai giai on v ch mt trong cn nh sng.
Blackman xc nh c tc ng ca cng nh sng v nhit khc
nhau ln quang hp. Khi cng nh sng tdng thp, ng tm thy rng
quang hp c th c tng tc nh lm tng lng nh sng, nhng khng tng

quang hp do lm tng nhit (Hnh 6.2). Song, cng nh sng Co, khi tng
nhit th lm tng ng k quang hp. Da trn cc s liu ny, Blackman
kt lun rng quang hp bao gm mt nhm phn ng u tin m ng gi l phn

ng "sng" thng khng ph thuc nhit v nhm phn ng th hai l phn

ng "t" khng ph thuc nh sng. Khng nhm ln vi biu tng ca Blackman
cng gi l phn ng "t" xy ra ngoi sng (thc ra, chng yu cu cc sn phm
ca phn ng sng). Tn gi ca phn ng ny n gin cho rng nh sng khng
trc tip tham gia trong cc phn ng . Phn ln cc nh sinh l thc vt hin
nay gi chng l "phn ng khng ph thuc nh sng" nhn mnh im ny.
187

Tc Cfc i
COj cao qu
mc; 3 5 c

Nhit b gi(M hn
CO ca o qu mfc;

20c

COj b g i hn
COj khng (0,01% )- 20c
2000

2500

Hlnh 6.2. S pht hin v phn ng ti

(a) Blackman o cng quang hp di cng nh sng, nng CO2 v nhit

khc nhau
(b) Nh so ny cho thy, nh sng l nhn t hn ch cng nh sng Uip
trong khi nhit v nng COj l nhn t hn ch cng nh sng cao. )dn v chn
nn (ft)0 candles) l s o tiu chun v chi sng (n v khc l lux, 1 n v chn nn
= 10 lux). Nang lung thc ca nh sng c o theo jun/gidy (jun/sec) hoc Watt. nh
sng mt trdl sng chi c khong i. n v chn nn, 108.00 lux hoc 500W/m

Blackman tm thy rng nhit tng ln th lm tng tc ca phn ng

t nhng ch ln n khong 35C. Nhit cao hdn lm cho tc gim nhanh
chng, v 35C nhiu enzym thc vt tr nn b bin tnh (cc lin kt hidro c
chc nng gi enzym dng xc tc ring bt u b ph vd). Blackman kt lun
rng enzym phi thc hin cc phn ng t. By gi chng ta bit rng, giai on
u tin ca quang hp l phn ng ph thuc nh sng hay phn ng "sng",
dng nng lng ca nh sng kh NADH (mt phn t cht mang in t)
thnh NADPH v tng hp ATP. NADPH v ATP t giai on u ca quang hp
tip tc c s dng trong giai on th hai - cc phn ng khng ph thuc nh
sng hay phn ng "ti" kh cacbon trong kh cacbonic v to ra ng n m
khung cacbon ca n c th c s dng tng hp cc phn t hu ckhc. S
kt hp ca cc nguyn t cacbon ca CO2 thnh phn t hu ctrong phn ng ti
c gi l s c nh cacbon (carbonfixation).
188

T cng trnh ca Blackman v cng trnh ca nhiu nh nghin cu khc, vn

tr nn r rng: nng lng ca nh sng mt tri c s dng trong cc phn
ng sng ca quang hp thc y qu trnh kh cc phn t cht mang, hay ni

khc i, quang hp l mt qu trnh oxi ho kh. Hy nh li t chdng 5 rng tt

c cc qu trnh oxi ho kh u ko theo s truyn nng lng. Trong h hp oxi
ho, nng lng c gii phng t phn t tng khi in t kt hp vi cc

nguyn t hidro trong ng c dn truyn nh cht mang nh NADH n oxi,

hnh thnh nc nh mt sn phm ph cui cng. in t c chit rt t ng
mt th nng khi chng b ht vo chui truyn in t do cc nguyn t oxi c
in m mnh v ty th dng nng lng c gii phng ra tng hp ATP.
Trong quang hp, con ng dn truyn in t b o ngc. Nc b phn ly v
in t (cng vi nguyn t hidro kt hp) c chit rt v c truyn n

nguyn t cacbon trong kh cacbonic, kh chng v to ra ng. i theo chiu

hng ny, in t thu c th nng khi chng chuyn t nc n ng. Vai tr
ca nh sng trong quang hp l cung cp nng lng ny.
Blackman d ch ra rng thu nng ing quang hp thi cn nh

sng mt tri, cn vic tng hp phn t hu c th khng.

6.3. Sc t thu nng lng t nh sng mt tri

6.3.1. L sinh ca nh sng
Nng lng trong nh sng nm u? Trong nh sng mt tri, cy c th
dng ci g kh kh cacbonic? l b mt ca quang hp - mt nhn t khc c
bn vi cc qu trnh nh h hp. tr li nhng vn ny, chng ta cn xem
xt v tnh cht vt l ca bn thn nh sng. C l ni tt nht bt u l th
nghim ni ting c thc hin trong phng th nghim c nm 1887. Nh vt

l tr Heinrich Hertz c gng gii thch l thuyt ton hc cao cp d on v s

tn ti ca cc sng in t. thy liu cc sng nh vy c tn ti hay khng,
Hertz thit k mt th nghim ti tnh. Trn mt pha ca phng ng dng
mt my pht tia in mnh gm hai qu cu kim loi ln sng bng ng cnh
nhau trn thanh thon nh, cao vt ln. Khi in tch tnh rt cao c tch t trn
mt qu cu, tia in s nhy qua qu cu kia.
Sau khi p t tht b ny, Hertz d nh nghin cu liu bt tia in s to
ra cc sng in t khng nhn thy gi l sng v tuyn nh d on theo
thuyt ton hc hay khng. Trn pha kia ca phng ng t my thu v tuyn u
tin trn th gii - mt ci vnh kim loi mng trn gi cch in. C mt khe nh
y ca vnh sao cho vnh khng to nn vng trn y . Khi Hertz m
my pht tia in xuyn qua phng, ng nhn thy cc tia in b t xuyn qua
khe trong vnh. l minh chng u tin v sng v tuyn. Nhng Hertz nu
hin tng l lng khc. Khi nh sng chiu qua khe trn vnh th cc tia in
c to ra mt cch d dng hn. S t ph bt ng ny c gi l hiu ng
189

quang in
nm.

(photoelectriceffect) gy bi r cho cc nh nghin cu trong nhiu

Hiu ng quang in cui cng Albert Einstein gii thch nm 1905. nh sng
gm n v nng lng gi l photon v mt s cc photon ny c cc nguyn t
kim loi vnh hp th. Trong qu trnh ny, mt s' in t trong nguyn t kim
loi c nng ln mc nng lng cao hn v lm cho chng b bt ra khi nguyn
t. Thc t l nh sng lm bt in t t b mt kim loi cc u cui ca vnh
kim loi, to ra cc ion v do khai thng ng dn tia in t do sng v tuyn

gy ra.

6.3.I.I. Nng lng trongphoton

Cc photon u khng c cng mc nng lng. Mc nng lng ca photon t
l nghch vi bc sng nh sng: nh sng c bc sng ngn cha photon c nng
ng cao hn nh gng c bc sng di. Tia X cha nhiu nng lng c bc sng
rt ngn - ngn hn nhiu so vi nh sng thy, khin cho chng l l tng cho
cc knh hin vi c6 phn gii cao.
[

---------- Nng l(;tag tng ln-------------------------------------------- 1

i n

i / v w

\ A

/ v v \ y \

----------------vBc sng tng ln-----

0.001 nm
1

1 nm lOnm lOOOnm

Tia
Tia gamma

400nm

0.01 cm

lo n

.......... - ]

430nm

T ia X

uv

500 nm

1m
1

100 m
1

Tia hQg
Sng v tuyn

560 nm

600 nm

650 nm

740ran

Hlnh 6.3. Ph ln t. nh sng l dng nng lng in t coi nh sng. Bc sng

nh sng cng ngn, nng lng ca n cng ln. Anh sng thy ch th hin mt phn nh
ca ph in t gia 400 v 740 nm (nm = namomet = mt phn t mt = 1/10*m)

Hertz lu rng cng ca hiu ng quang in ph thuc vo bc

sng nh sng. Cc bc sng ngn hiu qu hn nhiu so vi bc sng di trong
190

vic to ra hiu ng quang in. L thuyt ca Einstein v hiu ng quang in

a ra li gii thch: nh sng mt tri cha cc photon gm nhiu mc nng lng
khc nhau v ch mt s trong mt ta thu nhn nh l nh sng nhn thy.
Photon c nng ng cao nht cui bc sng ngn ca ph in t (Hnh 6.3) l
tia gamma c bc sng nh hn Inm. Cc photon c nng lng thp nht vi bc
sng ln n hng ngn mt l sng v tuyn. Bn trong phn ph nhn thy, nh
sng tm c6 bc sng ngn nht v mang cc pl'oton c nng lng cao nht. nh

sng trong vng ph nhn thy c bc sng di nht v do c photon sinh

nng lng thp nht.

6.3.1.2. nh sng cc tm
nh sng mt tri cha mt lng ln nh sng cc tm (UV). Do nh sng u v
c bc sng ngn hn nn c nng lng cao hn nhiu so vi nh sng nhn thy.
nh sng v l ngun nng lng quan trng trn tri t nguyn thu khi s
sng c khi sinh. Kh quyn ngy nay cha ozon (bt ngun t kh oxi) m hp
th phn ln cc photon ca u v trong nh sng mt tri, nhng bt k ai b sm
nng (rm nng) u bit rng mt lng ng k ca nh sng u v vn thot ra v
thm nhp vo kh quyn. Gn y, ngi ta pht hin nhng "l" y b n
trong tng ozon. L ny xut hin l do hot ng ca con ngi v bo trc s
tng ng k t l tc ng nh hdng v cn bnh ung th da ca ngi trn ton
th gii.

6.3.1.3, Phhp th v cc loi sc t

Mt phn t bt gi (capture) nng lng nh sng nh th no? C th hnh
dung photon nh mt ti nng lng vn ng rt nhanh (a very fast - moving
packet ofenerg). Khi n df> vo mt phn t th nng lng ca n hoc mt i
di dng nhit hoc c in t ca phn t hp th nng cc in t ny n
mc nng lng cao hn. Nng ng ca photon c hp th ph thuc vo n
mang bao nhiu nng lng (c xc nh theo bc sng ca n) v vo tnh cht
ho hc ca phn t n va chm. Nh chng ta bit, in t chim mc nng
lng ling trong cc qu o xung quanh nhn nguyn t. nng mt in t
ln mc nng lng khc t i hi mc nng lng thch hp tng t nh vic
bn tro thang, bn mun vn ln mt nc thang tip theo th bn cn nng chn
ln khong cch thch hp. Do , mt nguyn t ring ch c th hp th cc photon
nht nh ca nh sng, c th l cc photon no tng ng vi mc nng lng kh
dng ca in t nguyn t. Kt qu l mi phn t c mt ph hp th c trng
(absorptionspectrum) - khong photon (phm vi photon) n c th hp th.
Phn t c kh nng hp th tt nh sng gi l sc t (pigment). Sinh vt
pht trin nhiu loi sc t khc nhau nhng ch c6 hai loi ph bin c s dng
trong quang hp ca cy xanh: cc sc t vng v dip lc. Dip lc hp th photon
bn trong khong nng lng thp. Thc vt c hai loi dip lc: dip lc a v b
tng ng hp th nh sng xanh - tm v nh sng (Hnh 6.4). Khng c sc t
191

no trong cc sc t ny hp th photon vi bc sng khong gia 500 v 600 nm

v do nh sng ca cc bc sng ny b cy xanh phn x. V sau khi cc
photon ny c sc t hp th nh sng trong mt ta hp th (retinal) th chng
ta thy c chng di dng mu lc.
Dip lc b

550

Dip lc a

600

Bc sng (nm)
Hlnh 6.4. Ph hp th ca dip lc. inh th ch bc sng ca nh sng mt tri m
hai dng sc t quang hp ph bin - dip lc a (ng lin nhau) v dip lc b (ng
chm chm) hp th mnh. Cc sc t ny hp tti ch yu nh sng xanh tm v nh sng
trong hai bng hp ca ph v phn x nh sng lc vng gia ca ph. Cc sc t
vng (khng trnh by y) hp th ch yu nh sng xanh v nh sng lc, phn x nh
sng da cam v nh sng vng

Dip lc a l sc t quang hp ch yu v l sc t duy nht m c th hot

ng trc tip bin nng lng nh sng thnh nng lng ho hc. Song, dip
lc b hot ng nh mt sc t ph hay sc t th cp hp th nh sng, c chc
nng h tr v b sung cho s hp th nh sng ca sc t' chnh - dip lc a. Dip
lc b c mt ph hp th dch chuyn hng n bc sng lc. Do , dip lc b c
th hp th cc photon m dip lc a khng th hp th. V vy dip lc b lm tng
ng k t l photon trong nh sng mt tri m cy c th hp th. Nhm sc t
ph quan trng gi l sc t vng

(carotenoit) cng h tr quang hp nh hp th

nng lng t nh sng gm cc bc sng m khng c dip lc hp th mt

cch hiu qu.
192

T rong q u an g hp, cc p hoton c a n h sn g c cc sc t h'p th .

Bc sng c a n h sn g dc h p th th p h th u c vo sc t
chuyn ho.

6.3.2. Dip lc v sc t vng

6.3.2.1. Dip lc
Dip lc hp th photon nh qu trnh kch thch tng t vi 'hiu ng quang
in. Cc sc t ny cha mt cu trc vng phc hp gi l vng pocphirin vi cc
lin kt n v i xen k. trung tm ca vng l mt nguyn t magie. Photon
c phn t sc t"hp th kch thch in t trong vng v tip tc chuyn qua h
lin kt cacbon. Mt s nhm bn nh b c gn vo pha ngoi vng c tc dng lm
thay i tnh cht hp th ca phn t trong cc loi dip lc khc nhau (Hnh 6.5).
Chloro Jhyll
Chlorohll

Phn t dip
lc c ga
ong mng

a:.-CHj
b:CHO

H jC = C H

Mng

thilacoit
H
CH H

0=0

'

COCH,

V-

Granum
Hlnh 6.5. Dip lc
Phn t dip lc gm phn u poqhirn v phn ui hidrocacbon c chc nng neo
gi phn t sc t vo vng ky nuc ca proteln c gn bn trong mng thilacolt. S sai
khc duy nht gia hai phn t sc t l s thay th ca nhm aldehlt (-CHO) ong dip lc b
cho nhm metyl (-CH3) trong dip lc a

Trong phn trc, Ingenhouz chng xiinh rng ch cc b phn xanh ca

cy c th "phc hi" khng kh. Cc nh nghin cu nghi ng quan im coi
dip lc l sc t ch yu m cy dng hp th nh sng trong quang hp. Cc
193

th nghim c tin hnh trong nhng nm 1800 chng minh mt cch r rng
s nghi ng ny. Mt th nghim nh vy do T.W.Englemaun tin hnh nm 1882
(Hnh 6 .6 ) c tc dng nh mt th d rt mi m: n gin v thit k v r rng
v kt qu. Englemann trnh by ph hot ng ca quang hp (action spectrum)
ngha l hiu qu tng ca cc bc sng khc nhau ca nh sng trong vic
kch thch quang hp. ng tin hnh trn vn th nghim dng tiu bn n gn

trn knh hin vi. thu nhn cc bc sng nh sng khc nhau, ng t mt
lng knh di knh hin vi tch nh sng chiu vo tiu bn thnh ph mu

(spectrumofcolors). ng cho si t bo to ngang qua ph sao cho cc phn khc

nhau ca si to c chiu sng vi bc sng khc nhau to tin hnh quang

hp. nh gi quang hp tin hnh nhanh chng nh th no, Englemann theo

di tc gii phng oxi. Mc d lc by gi cn thiu khi ph k v cc thit b
hin i khc, nhng ng b sung cho tiu bn vi khun theo kh (aerotactic
bacteri) hay cn gi l vi khun tm oxi {oxygen- seeking bacteria) v ng bit
vi khun ny s tp trung dc theo sdi to cc v tr oxi c gii phng, ng

tm thy rng vi khun t hp li cc vng c chiu sng bng nh sng v

nh sng tm - hai mu nh sng c dip lc hp th mnh m nht.

Hlnh 6 .6 . y l th nghim ca T.VV.Englemann chng minh ph hot ng ca quang

hp trong to SI Spirogyra nm 1882. Englemann dng tc to 0X1 o tc ca
quang hp. n g chn vi khun b oxi dn d nh l vt ch th. v tr ca guang v mn
chn sng ttiitng c dng chiu sng mu vt di knh hin vi, ng thay th "b vi
ph" {mlcmspectral apparatus) c6 tc dng to ra mt ph mu b nh chiu ln vt knh
dui kfnh hin vi. Sau , ng t mt si t bo to song song vi s lan truyn ca ph.
Cc vi khun tim kim oxi tp hp thnh m dy c ch yu trong vng bc sng tm v
gp si to. Dng th v ph hot ng cho quang hp. Ph hot ng cho quang hp
m Englemann trnh by trong th nghim ny song song vi ph hp th ca dip lc.

Mi thc vt v to, ch tr mt nhm nguyn thu ca vi khun quang hp

u s dng dip lc a nh l sc t' ch yu trong quang hp. Vn c l hi
ti sao cc c th quang hp ny khng dng sc t nh retinal m c mt bng ph
hp th rng ph ln khong bc sng nh sng t 500 - 600nm? Gi thuyt thch
hp nht c lin quan n tnh hiu qu ca nh sng hay gi l quang hiu qu

194

(photo efficiency). Mc d sc t retinal hp th mt khong rng ca bc sng

nhng vi hiu qu tng i thp. Ngc li, dip lc ch hp th nh sng trong
hai bng hp, nhng tin hnh vi hiu qu cao. Do , thc vt v phn ln cc c
th quang hp khc vi dip lc th t c tc hp th photon chung kh cao
so vi cc sc t khc.

s.3.2.2. Sc tvng
Bao gm cc vng cacbon lin kt thnh chui vi cc lin kt n v i xen
k. Chng c th hp th photon vi mt khong rng ca nng lng, mc d
chng khng phi lun lun c hiu qu cao trong vic truyn nng lng ny. sc
t vng h tr quang hp nh hp th nng lng t nh sng gm cc bc sng

m khng c dip lc hp th hiu qu (Hnh 6.7).

L si
ong ma h

v^H ^yLsi
{M k
trong ma thu

Hnh 6.7. Mu sc ca ma thu c to ra nh cc sc t vng v cc sc t ph khc.

Trong ma xun v ma h, dip lc trong l che lp s hin din ca cc sc t vng v
cc sc t ph khc. Khi nhit ma thu lnh lm cho l ngng tng hp dip lc, dip lc
khng cn phn x nh sng lc, l phn x nh sng da cam v nh sng m cc sc
t vng v cc sc t khc khng hp Ui

Mt Sc t vng in hnh l P-caroten c hai vng cacbon ni nhau bng chui

gm 18 nguyn t cacbon vi cc lin kt n v i xen k. Tch mt phn t Pcaroten thnh hai na bng nhau th to nn hai phn t vitamin A. S oxi ho
vitamin A to nn sc t hp th nh sng retinal - sc t c s dng trong s
nhn ca ng vt c xng sng. iu gii thch ti sao c rt giu P-caroten c
tc dng tng cng s nhn ca ngi v cc ng vt c xng sng khc.
Sc t l phn t hp th nh sng. Bc sng dc mt sc t' ring
hp th th ph thuc vo mc nng lng m in t c nh sng
kich thich c th c nng ln trong sc t.
6.3.3. S t ch c sc t th n h quang h

Phn ng ph thuc nh sng ca quang hp xy ra trn mng (Hnh 6 .8 ). vi

khun nh vi khun do Van Niel nghin cu, bn thn mng sinh cht l mng
quang hp. Ngc li, thc vt v to, quang hp xy ra bo quan m l dng di
195

(con chu) tin ho ca vi khun quang hp - lc lp gm cc mng quang hp tn ti

bn trong lc lp. Cc phn ng ph thuc nh sng xy ra trong ba giai on;

Hlnh 6 .8 . S ly mu cc mng quang tip trong cc c th t dng

(a) Cy xanh Coleus blumei (500x)
(b) To lc on bo Chlorella

(c) Khun lam c nh nit Nostoc museorum (6000x)

1. Quang tc ng s cp (primary photoevent): Photon nh sng c sc t

hp th. Kt qu ca quang tc ng s cp ny l 8 kch thch mt in t bn
trong sc t.
2. Truyn in t {electron transport): in t kch thch di chuyn dc theo
mt lot cc phn t cht mang in t c gn bn trong mng quang hp cho
n khi n n knh bm proton xuyn qua mng (i khi n c gi l "bm oxi
ho kh"). Vic in t n bm dn n s truyn proton qua mng. K , in t
c chuyn n mt cht nhn.
3. S ho thm {chemosmosis)-. S truyn proton quay tr li qua mng c tc
dng thc y tng hp ho thm ATP, ging nh qu irkih xy ra trong h hp
hiu kh.

6.3.3.I.Spht minh cc quang h

Phng php nghin cu sc t hp th nh sng l nh gi s ph thuc v
hiu sut ca quang hp ln cng chiu sng, ngha l xem xt c bao nhiu

hiu sut quang hp c to ra vi bao nhiu nh sng c s dng. Khi th

nghim ny c tin hnh trn thc vt thi ngi ta bit rng hiu sut ca
quang hp tng ln tuyn tnh cng cao hn v cu^ cng bo ho cng

nh sng cao. S bo ho xy ra do tt c kh nng hp th nh sng ca thc

vt hin c dng nn vic tng thm nhiu nh sng khng lm tng hiu sut
do khng cn g hp th thm photon.

196

Ngi ta quan nim rng lc bo ho th tt c cc phn t sc t ca cy c

s dng ht. Nm 1932, cc nh sinh l thc vt Robert Emerson v William
Arnold d nh kim tra gi thuyt ny trong c th thc vt m h c th o c v
s phn t dip lc v hiu sut quang hp. Trong th nghim ny, h o hiu qu
gii phng oxi ca quang hp khi chlorella (to lc n bo) c chiu vi cc chp
sng rt ngn ch tri qua mt vi microgiy. Gi d cho gi thuyt v s bo ho
ca sc t l ng, h hi vng tm ra rng khi tng cng ca chp sng th hiu
sut trn chp sng s tng ln cho n khi mi phn t dip lc hp th mt
photon m v sau c s dng trong cc phn ng ph thuc nh sng to ra
mt phn t O2.
Sboho ichi

tt c phn t '
Khng ng, iu khng xy
dip lc ang
ra (Hnh 6.9). Thay vo , s bo
s dng
ho t c sm hn nhiu vi ch
mt phn t O2 trn 2500 phn t
S bo ho khi tt c
dip lc. Kt qu ny dn IMI<v
^ ^ u a n g h ang s dng
Emerson v Arnold kt lun rng
nh sng c hp th khng phi

D on

do cc phn t sc t ring (phn

---------- Quan st
t sc t' c lp) m ng hn l
Cng ca chp sng
do mt nhm gm cc phn t dip
lc v phn t sc t' ph gi l
Hlnh 6.9. Th nghim ca Emerson v Amold.
quang h (photosystems). Trong
Khi t c s bo ho quang hp thl s tng tip
mt quang h nh quang h ca
tc v citng nh sng khng lm tng hiu
Chlorella, nh sng c hp th sut quang hp
nh bt k mt trong hng trm
phn t sc t trong quang h v nng lng kch thch c truyn n mt phn
t sc t c mc nng lng thp hn phn t khc - l trung tm phn ng
(reaction center) ca quang h hot ng nh mt im tho nng lng (energy
s in k ) - c tc dng by nng lng kch thch. l s bo ho ca cc trung tm
phn ng ny, khng phi bo ho cc phn t sc t c th - kt qu th nghim do
Emerson v Arnold quan st c.

6.3.3.2. Kin trc ca mt quang h

Trong lc lp v gn nh phn ln vi
khunnguyn thu, nh sng c
th nh cc quang h . Mi quang h l mt mng gm ccphnt dip lc av
cc sc t ph c duy tr bn trong cht nn protein trn b mt ca mng quang

hp. Ging nh mt knh lp lm hi t nh sng trn mt im chnh xc, quang

h chuyn nng lng kch thch do bt k mt phn t sc t no tp hp cho mt

phn t sc t chuyn ho - dip lc trung tm phn ng. v sau, phn t ny

truyn nng lng ra ngoi quang h sao cho n a nng lng vo hot ng
thc y qu trnh tng hp ATP v cc phn t hu c.
197

hp

Do , quang h gm hai thnh phn lin kt cht ch vi nhau: ( 1 ) phc h

anten gm hng trm phn t dip lc c chc nng thu thp (tp hp) photon v
cung cp nng lng nh sng thu c cho trung tm phn ng v (2) trung tm
phn ng (re a c tio n c e n te r ) gm mt hoc nhiu phn t dip lc a trong cht nn

gm protein c chc nng truyn nng lng ra ngoi quang h.

* Phchanten
Phc h anten hp th
photon t nh sng mt tri
(Hnh 6.10). Trong lc lp, phc
h anten l mt mng gm cc
phn t dip lc lin kt vi
Cht cho
nhau v c gi cht trn
in t
mng thilacoit nh mt cht
nn gm cc protein. S lng
cc sc t ph carotenoit cng
c th bin i. Cht nn
protein c chc nng nh mt Phn t
loi khung hay gin gio dip lc
Quang h
(scaffold) gi cc phn t sc
t c th theo hng ti thch
Hinh 6.10. Phc h anten hot ng nh th no?
cho s dn truyn nng lng.
Khi nh sng ca bc sng thlch hp p vo bt
k phn t sc t no bn trong quang h, nh sng
Nng lng kch thch xut
uc
phn t sc t hp th. v sau, nng lng
pht t s hp th ca photon
kch thlch c uyn t phn t n phn t k tip
c truyn t mt phn t sc
bn trong nhm (b) phn t sc t cho n khl n bt
t n phn t tip theo trn
gp dip lc a trung tm phn ng. Dip lc trung tm
con ng n trung tm phn
phn ng truyn nng lng dng in t cao nng
ra khi quang h cho b my hp thu nng lng
ng. Sau khi dn truyn, in
(energy harvestng machinery)
t kch thch trong mi phn t
sc t quay li mc nng lng
thp m n c trc khi photon c hp th. Do , nng lng ch khng phi
bn thn in t kch thch c truyn t mt phn t sc t ny n phn t tip
theo. Phc h anten c chc nng thu ht v dn t nng lng t nhiu in t
cho trung tm phn ng.

s.3.3.3. Trung tmphn ng

Trung tm phn ng l phc h protein - sc t xuyn qua mng thilacoit.
trung tm phn ng ca vi khun ta quang hp (n gin hn so vi lc lp
nhng c nghin cu y hn) c mt i phn t dip lc a hot ng nh
ci by vi nng lng photon, ri chuyn in t kch thch cho mt cht
198

nhn thch hp nh l phn t ln cn ca n. y l bc chuyn s kch thch

ca photon i khi dip lc v l s bin i ch yu ca nng lng nh sng
thnh nng lng ho hc.
Hnh 6.11 ch r s truyn
Phn t dip lc
Dip lc b
b idc thch
nng lng t trung tm phn
ox ho
C h t nhn C h t cho Cht nhn
ng n cht nhn in t s C h t cho Ch't nhn
in t
>i kh
b oj ho b kh
in t
cp. Nh kch hot mt in t
ca dip lc trung tm phn
ng, nh sng to mt cht cho
in t mnh m khng tn ti
trc . Dip lc truyn in
t cao nng (in t c kch
Hlnh 6.11. S bin i nng litng nh sng thnh
hot; energed electron) n
nng lng ho hc. Dip lc trung tm phn ng
cht nhn s cp - mt phn t
chuyn giao cht nhn in t cao nng cho cht nhn
in
t s cp v kh cht ny. Dip lc b oxi ho lp
quinon ri kh quinon v bin
y "l trng" in t (l trng electron) nh ox ho phn
n thnh cht cho in t
t cht cho
mnh, v sau, cht cho in t
yu {weak electrondonor) truyn in t c nng lng thp (oiv - energy electron)
cho dip lc ri phc hi n tr li trng thi ban u. vi khun ta, cht cho ny
l mt xitocrom v chng ng ca in t i theo mt con ng vng nh chng
ta s trnh by trong phn tip theo. Trong lc lp thc vt, ndc ng vai tr nh
cht cho in t.
Cc sc t trong quang h hp thu nng lng photon t nh sng,
ri hi t nng lng cho trung tm phn ng v trung tm ny
dng nng lng kch thch in t thnh in t cao nng, c
chuyn vo h truyn in t quang hp d sinh cng ho hc.
6.3.4. Q u a n g h b i n n n g l ng n h s n g th n h n n g l n g h o h c n h
th no?

6.3.4.I. Vi khun dng quang h n

Quan nim chung coi n v quang hp tin ho hn 3 t nm trc trong vi
khun ging vi khun lu hunh do Van Niel nghin cu. Trong vi khun ny, s
hp th mt photon nh sng nh hp th 870 nanomet nh quang h dn n
vic truyn mt in t cao nng dc theo chui truyn in t, cui cng kt hp

vi proton to ra nguyn t hidro. Trong vi khun lu hunh (Hnh 6.12), proton

c chit rt t hidro sunfua (HS), li nguyn t' lu hunh nh l sn phm
ph. vi khun tin ho v sau, cng nh trong to v thc vt, proton bt ngun
t nc, to oxi nh l sn phm ph.
199

Hinh 6.12. Vi khun lu hunh ta. Do h quang ca chng tng i n gin nn vi

khun lu hunh l i tng nghin cu chuyn su

S y bt ra mt in t khi trung tm phn ng vi khun khin trung tm

thiu mt in t. TrC khi quang h ca vi khun lu hunh c th hot ng tr

li th mt in t phi quay tr li. Vi khun ny chuyn in t quay li sc t'

thng qua h truyn in t ging vi h thng trnh by trong chng 5 v s
truyn ca in t c tc dng kch hot bm proton pht ng qu trnh tng
hp ho thm ATP. Mt phn t ATP c to ra mi khi ba in t i theo con
ng ny. NMn khi qMt (Hnh 6.13) th con ng ca in t c chit rt ban
u t quang h II m vng: o ; nh ho hc gi qu trnh ny l photphoril
ho quang hp vng hay quang photphoril ho vng {cclic p h o to p h o s p h o r y la tio n ).

Song, lu rng in t ri khi trung tm phn ng Ps70 l in t cao nng, cn

in t quay li ch c nng lng nh n c trc khi photon c hp th.
Hiu nng trong nng llng ca in t l kt qu quang hp (hiu qu quang
hp = photosynthetiicpayoff) -nng lng c tc dng kch hot bm proton.
Trong hdn mt t nm, quang photphoril ho vng l dng phn ng sng
quang hp duy nht, m sinh vt s dng. Song, hn ch ch yu ca n l ch
n ch ch hng phc v cho s sn xut nng lng, khng hng n qu trnh
sinh tng hp. Phn ln c th quang hp thng kt hp kh cacbonic kh quyn

thnh hidratcacboi. Do phn t hidratcacbon b kh hn (c nhiu nguyn t hidro

hn) so vi kh cacbonic nn mt ngun lc kh (ngha l hidro) phi c b sung.
Quang photphoril ho vng khng lm iu ny v vi khun b hn ch vi qu
trnh ny phi tm kim (thu gom: scavenge) hidro t cc ngun khc - mt cng
vic khng hiu qu.
200

Cht ahn
in t

Peedoxin

<S-

00
I
o

Hnh 6.13. Con ng ca in t trong vi khun lu hunh ta. Khi mt in t c nh

sng hot ho bt ra khi trung tm phn ng ca quang h (Pg^o), n chuyn theo mt
vng, cui cng quay li quang h m t n b bt ra

6.3.4.2. Ti 800 thc vt dng hai quang h?

Sau khi vi khun lu hunh xut hin, cc loi vi khun khc pht trin
nn kiu quang h ci tin, vt qua s hn ch ca quang photphoril ho vng
theo cch kho lo v n gin: mt quang h th hai, mnh hn dng cch sp
xp dip lc a theo kiu khc c lin quan vi bn gc. S tn b tin ho ln ny
xy ra khi khun lam xut hin, khng t hn 2,8 t nm trc.
Quang h th hai ny c gi l quang h II

(photosystemII), trong phn

t dip lc a c sp xp theo hnh dng khc sao cho cng nhiu bc sng ngn
hn mang photon nng lng cao hn c hp th so vi quang h nguyn thu

gi l quang h I

(photosstemI). Cng nh trong quang h nguyn thu, nng

lng c truyn t phn t sc t' ny n phn t khc bn trong phc h anten

ca quang h ny cho n khi n n trung tm phn ng - mt phn t sc t c

bit nm gn mt cht nhn in t mnh lin kt mng. Trong quang h II, nh
hp th (cc i hp th - bc sng nh sng c hp th mnh nht) ca sc t
l khong 680 nanomet, do sc t trung tm phn ng mang tn Peso- nh hp
th ca sc t quang h I trong thc vt l 700 nanomet, nh vy sc t trung tm
phn ng ca n c gi l P 7 0 0 . Cng phi hp vi nhau, hai quang h thc hin
phn ng ph thuc nh sng khng vng.
201

Gradient proton
c hnh inh
tng hp ATP
^lastoquinonc

/
NADP + H

Photon

Photon

Plastocynin
Enzym
phn ly nc

Quang h I

NADP reductaza

Hlnh 6.14. S z gm quang h I v II. Hai quang h hot ng theo dy ni tip. u

tin photon nh sng lm bt mt in t cao nng t quang h II, in t ny c dng
bm proton qua mng, ng gp c ch ho thm cho qu trnh tng hp phn t ATP. Tip
,
t bj bt ra diuyn dc theo chui xitocrom n quang h I. Khi quang h I hp th
photon nh sng, n lm bt mt in t cao nng c dng thc y to ra NADPH

Vic s dng hai quang h gii quyt vn v vic thu nhn lc kh theo
cch n gin v trc tip nh khai thc s dng nng lng ca hai quang h.
S trong hnh .l4 gi s d z (z diagram) mnh ho hai bc kch hot in
t, mi mt bc c xc tc nh mt quang h. in t xut pht t nc (gi
in t rt cht) vi th oxi ho kh bng +820 mV v kt thc trong NADPH gi
in t lng lo hn nhiu (th oxi ho kh = -320 mV). Vic s dng hai quang h
ring trong mt dy cung cp nng lng hn nhiu kch hot in t sut
con ng t y ca quang h II n nh ca quang h I vi nng lng li
cho php chui truyn in t lin kt hai quang h bm proton qua mng
thilacoit v nh cung cp nng lng tng hp ATP ho thm.
T rong vi k h u n lu hunh, in t b kch thich b t ra khi tru n g
tm p h n ng chuyn theo con ng vng, kch h ot botai proton v
v sa u q u ay li quang h ban du ca chng. Thc vt dng hai
q u an g h tro n g mt dy v pht sinh nng lng kh NADP*
th n h NADPH vi nng lng li tng hp ATP.

202

6.3.5. Hai quang h ca thc vt hot dng vi nhau nh th no?

Thc vt dng hai quang h theo th t ci u ri tip theo, ng thi to c

ATP v NADPH. Qu trnh hai giai on ny c gi l quang photphoril ho
khng vng (n o n c y c lic p h o to p h o s p h o r y la tio n ) do con ng ca in t khng trong

mt vng - in t bt ra t quang h khng quay li quang h na m kt thc

trong NADPH. Thay vo quang h c lin y tr li (lm y cc l trng
in t trong quang h) bng in t do phn ly n c to ra. Quang h II hot ng
u tin. in t cao nng pht sinh nh quang h II c s dng tng hp
ATP v tip c chuyn n quang h I kch hot qu trnh tng hp
MADPH. C mi i in t t nc th pha sng to ra mt phn t NADPH v
hn mt phn t ATP.

6.3.5.1. Quang h II
Trung tm phn ng ca quang h II gi l Peso kh ging vi trung tm phn
ng ca vi khun ta. N gm hn 10 tiu n v protein xuyn qua mng. Phc h
anten hp thu nh sng gm khong 250 phn t dip lc a v cc sc t ph lin
kt vi mt s chui protein. Trong quang h II, cc nguyn t oxi ca hai phn t
nc lin kt vi mt b (nhm) cc nguyn t magi gn bn trong enzym trong
trung tm phn ng, k hiu z. Trong con ng cha hiu cn k, enzym ny
(enzym phn ly nc) loi mi ln mt in t lp y cc l trng in t li
trong trung tm phn ng do s xut pht ca cc in t cao nng. Ngay khi bn
in t b loi khi hai phn t nc, O2 c gii phng.
Tuy nhin chi tit phn t chnh xc ca qu trnh oxi ho nc do trung tm
gii phng oxi (oxygen-evolvingcenter: OEC) xc tc vn cha c cu tr li. Cng
c nhiu thng tin v c ch xc tc ca PSII t mt phm vi. lng cc nghin
cu tip cn khc nhau (B.A.Diner, G.T.Bbcoek, 1996-2002; J.Barber, 2003).
Nghin cu ca nhm tc gi c vin Max-Volmer do Athina Zouni ng u
(2001) lm sng t hn mt bc v thnh phn cu trc ca PSII. Dng khun
lam a nhit Synechococus elongatus vi in di gen SDS-poliacrilamit v khi ph
th PSII gm t ra 17 tiu n v polipeptit trong 14 nh v trong mng quang
hp: cc protein ca trung tm phn ng Dj v Dj, cc tiu n v anten bn trong
mng dip lc CP43 v CP47, tiu n v a- v p ca xitocrom b-559 v nhiu tiu
n v khc nh hn. y l bc tranh v cu trc tia X ca quang h II trn cs
ca cc tinh th hon ton hot ng trong qu trnh oxi ho nc. Cu trc chng
minh cc tiu n v protein v cc nhm ngoi (cofactor) c t chc theo khng
gian nh th no. xc nh cc tiu n v ln hn cng nh v tr v s nh
hng ca cc nhm ngoi. Thng tin mi cng b sung v tr, kch c v hnh dng
ca b mangan xc tc oxi ho nc. y l bc nhy vt trong tri thc ca chng
ta v s gii phng O2, l mt tin b quan trng v s kch thch nghin cu v
chc nng ring trong c ch oxi ho nc ca PSII.
203

Photon

Photon
H* + NADP*

aent
proton

2HjO
&izyin
phn ly nc

Fastocyatdn

Peredoxiii

4H*

Quang

II

phc h

Quang h I

NADP
reoctaza

Xoang thiacdt
Hnh 6.15. H truyn in t quang hp. Khi photon ca nh sng p vo phn t sc
t trong quang h II, n kch ich mt in t. Mn t ny lin kt vi mt proton b tuc
ot t nc do proteln z v c dn truyn dc theo ctiui gm cc cht mang n t
xltocrom lin kt mng (mi tn m). Khi nc b| phn ly, oxi c gii phng khi t bo
v cc ion hidro lu li trong xoang thilacoit. d bdm proton (phc h b,-f), nng lng do
proton cung cp c s dng truyn proton qua mng vo thilacolt. Nng cc ion
hldro bn trong thilacoK tip tc tng ln. Khi quang h I hp th photon khc ca nh sng,
sc t ca n chuyn in t cao nng th hai cho phc h kh, t pht sinh NADPH

Cng dng khun lam Thermosynechococus elongatus, nhm nghin cu Anh Nht do Kristina N.Perreira ng u (2004) thng bo xc nh cu trc ca
PSII phn gii 3,5 angstron ( = 1 0 cm) vi phn ln cc gc axit amin ca
phc h a tiu n v lng phn 650 kilodanton (50 kilodanton dimeric
multisubunit complex), cc v tr lin kt cho cc nhm ngoi v trnh by chi tit
v cu trc ca trung tm gii phng oxy. M hnh ca phc h PSII l n v tinh
th hc bt i xng {crystallograph asymmetrc unt) cha mt ime (cht nh
trng) ca PSII gm hai monome (n hp) trong ime gn ging ht nhau. Trong
m hnh, monome cha 19 tiu n v protein. Mi monome cha 36 dip lc a, 7
phn t P-caroten. Mi m hnh monome cng gm mt OEC, mt hem b, mt hem
c, hai plattoquinon, hai pheophitin, mt Fe khng hem v hai bicacbonat. Cu trc
ca OEC cha mt b Mn3Ca4 c dng hnh lp phng vi Mn th t bng lin
kt cu. Nh vy trong OEC c mt 4 ion Mn** v mt Ca** lin kt cht nm cnh
ion Mn** l iu kin tin quyt cho c ch oxi ho nc gii phng oxi phn t

(Siegbahn etal, 1999-2002). Ba ion Mn** v ion Ca** nm bn gc ca hnh lp

phng mo m vi cc nguyn t oxi cc gc khc ca hnh lp phng, y
khng tin trnh by mi chi tit cu trc ca OEC cng nh c ch oxi ho nc,
204

nhng mun nhn mnh rng kin trc ca trung tm gii phng oxi quang hp
ngy cng sng t hn nhiu 80 vi nhng g chng ta bit t trc n nay.

.3.5.2. Conng n quang h I

Cht nhn in t s cp cho in t cao nng ri khi quang h II l phn t
quinon ging nh trong quang h ca vi khun trnh by phn trc. Quinon
kh (plattoquinon, k hiu l Q) l mt cht cho in t mnh c vai tr chuyn
in t kch thch n bm proton gi l phc h be-r be-fcomplex) gn bn trong
mng thilacoit (Hnh 6.15). Phc h ny rt giHT phc h bCi trong chui truyn
in t h hp ca ty th trnh by chng 5. S ti ca in t cao nng c
tc dng kch hot phc h bg- bm proton vo xoang thilacoit. Protein b cha
ng gi l plattoxianin (k hiu PC) c chc nng mang in t n quang h I.

Photon

T bo thc vt
Hnh 6.16. Qu trnh ho thm trong lc lp. Phc h be-f gn trong mng illacot bm
proton vo phn bn trong ca thilgcoit. ATP c to ra trn b mt ngoi ca mng (pha
cht nn) khi proton khuch tn tr li ra ngoi thilacoit thng qua knh ATP synttiaza

205

6.3.S.3. Tng h*p ATP: qu trinh ho thm

Mi thilacoit l mt xoang kn cha proton do phc h bg-f bm t phn cht
nn vo n. Mng thilacoit khng thm vi proton, nh vy khi proton quay li
cht nn th hu nh ch thng qua knh gi l ATP synthaza (TPsynthases).
Knh ny nh li ra ging nh qu m trn b mt ngoi ca mng thilacoit. Khi
proton chuyn ra ngoi thilacoit thng qua knh ATP synthaza, ADP c
photphoril ho thnh ATP v c phng vo phn cht nn lng bn trong lc lp.
Cht nn cha enzvm c chc nng xc tc cc phn ng khng ph thuc nh
sng ca qu trnh c nh cacbon (Hnh 6.16).

.3.5,4. Quang hI
Trung tam phn ng ca quang h I mang tn P700 l mt phc h protein
xuyn qua mng, gm t nht 13 tiu n v protein. Nng lng c truyn cho
P700 nh phc h anten gm 130 dip lc a v cc phn t sc t' ph. Quang h I
nhn in t t plattoxianin lp l trng in t do s tch ri ca in t cao
nng to ra. in t n ny chc khng mt i tt c nng lng kch thch, vn
cn li gn mt na. Do , s hp th mt photon nh sng quang h I c tc
dng nng in t ri khi trung tm phn ng n mt mc nng lng rt cao.
Khng ging quang h II v quang h ca vi khun, quang h I khng da vo
quinon nh l cht nhn in t. Thay vo , n chuyn in t cho protein st lu hunh gi l feredoxin (Fd) (Hnh 6.14).
Trong nhng nm gn y, cu trc nguyn t ca PSI cng c ch . Ni
bt nht l nghin cu ca nhm ngi c do Patrick Jordan ng u (2001)
thng bo v cu trc ba chiu ca quang h I khun lam a nhit Synechococus
elongatus vi phn gii 2,5. y l bc tranh chi tit nguyn t vi 12 tiu n
v protein v 127 nhm ngo bao gm 96 dip lc, 2 phinoquinon, 3 b Fe4S<, 2 2
sc t vng, 4 lipit, mt ion
v 201 phn t nc. Thng tin cu trc v
protein, cc nhm ngai v s tng tc ca chng to c 8 tm hiu v hiu

qu cao ca quang h I tsrong vic hp thu nh sng v truyn in t t c nh

th no.

e.3.5.5. Tnghp NAPH

Quang h I truyn in t cho feredoxin trn pha cht nn mng ( ngoi
thilacoit). Peredoxin kh mang in t c th nng rt cao. Hai in t cao nng
ny t hai phn t PerecBoxin kh c chuyn cho phn t NADP* to NADPH.
Phn ng c enzym lin kt mng xc tc - NADP reductaza. Do phn ng xy
ra trn pha cht nn ca mng v ko theo s hp th proton trong khi hnh
thnh NADPH nn n tng gp xc lp gradient proton trong qu trnh truyn
in t quang hp (Hnh 6.14 v hnh 59, hnh 60 trang 152, 155 trong cun Sinh
hc i cng - Sinh hc phn t t bo, ti bn ln th 6 , Hong c C).
206

.3.5.6. To ATP nhiu hn

S truyn in t t nc n NADPH trong quang photphoril ho khng vng
c trnh by phn trc lm pht sinh mt phn t NADPH v hn mt phn
t ATP. Tuy nhin, vic tng hp cc phn t hu c cn nhiu nng lng hn. N
cn 1,5 phn t ATP trn phn t NADPH c nh cacbon. to thm ATP,
nhiu loi thc vt c th rt ngn quang h I, chuyn i quang hp thnh
phng thc quang photphoril ho vng {cyclicphotophosphorylationmode) sao cho
in t kch thch ri khi quang h I c s dn to ATP thay v to NADPH.
n gin l in t cao nng c chuyn tr li cho phc h bg- hn l chuyn cho
NADP^. Phc h bs-f bm proton ra ngoi xoang thilacoit, to nn gradient proton
v kch hot tng hp ho thm ATP. T l tng l ca quang photphoril ho
vng v khng vng trong thc vt ny xc nh lng ATP v NADPH tng i c
sn tng hp cc phn t hu c.
in t m quang hp s dng tng hp cc phn t hu c b
kh giu nng lng bt ngun t nc. Nguyn t oxi cn li ca
phn t nc kt hp to khi oxi.

6.4. T bo dng nng lng v lc kh do phn ng sng thu c

tng hp cc phn t hu c
Da vo s khc bit trong c ch ng ho CO2, c th phn chia thc vt
sng trn cn thnh ba kiu quang hp ch yu, c th l thc vt

C 3,

thc vt

C4

v thc vt chuyn ho axit cy mng nc (CAM). Mi kiu quang hp c mt

lot tnh cht c trng v gii phu, sinh l v sinh ho, lm cho chng thch nghi

S c nh CX)2
trong thc vt C3

S c nh CO2
ttong thc vt C4

Hnh 6.17. S c nh CO2. C th chia thc vt theo kiu

S c nh CO2
ong thc vt CAM

cnh CO;
207

vi t sinh thi iecological niche) ring. Thc vt C3 thc hin tt chc nng trong
vng kh hu n i, trong khi thc vt C4 pht trin mnh m trong mi trng
c nh sng mnh, kh hn v nng. Thc vt CAM c c trng bng vic hp
th CO2 ban m, thch nghi vi iu kin rt kh hn. Cc nghin cu v phn
loi v pht sinh chng loi nu ln rng thc vt CAM v thc vt C4 bt ngun
t thc vt C3 v s chuyn tip xy ra nhiu ln trong cc cp phn loi ring bit

trong lch trnh tin ho. Hnh 6.17 phc ho ba kiu quang hp in hnh trong
thc vt C3, C4 v CAM. Trong thc vt C3, chu trnh Calvin c nh CO2 mt cch

trc tip v phn t c th pht hin u tin i theo s c nh l PGA - mt phn

t C3. Thc vt C c nh CO2 nh to ra phn t C4 trc khi tham gia vo chu
trnh Calvin. Thc vt CAM c nh CO2 nh hnh thnh mt phn t C4 ban m

khi l kh c th m m khng mt nhiu nc.

6.4.1. Cc phn ng khng ph thuc nh sng

Quang hp l phng cch to cc phn t hu c t kh cacbonic (CO2).

Phn t hu cd l cht c trng thi kh cao so vi CO2 v chng cha nhiu lin
kt C-H. tng hp phn t hu c, t bo dng cc nguyn liu th {raw
materials) do phn ng ph thuc nh sng cung cp:
. Nng lng. ATP do quang h II cung cp c tc dng kch hot (thc y)
phn ng thu nhit.
1

2. Lc kh. NADPH do quang h I cung cp c tc dng b sung ngun hidro

v in t cao nng cn lin kt chng vi nguyn t cacbon. Phn ln nng
lng nh sng do quang hp hp th cu cng c u t vo cc lin kt C-H
giu nng lng ca ng.

6.4.1.1. Scinh cacbon

Giai on ch yu trong cc phn ng khng phu thuc nh sng l phn ng

kh CO2 - s gn hay lin kt CO2 vo phn t hu c. Mun vy, t bo cn mt
phn t rt chuyn ho. T bo quang hp to phn t ny bng cch ti hp
{ r e a s se m b lin g ) cc lin kt ca hai sn phm trung gian trong chui ng phn l

fructozo 6 -photphat v glixeron dehit 3-photphat to ra ng 5-cacbon giu

nng lng ribulozo 1,5 diphotphat (RuDP) hay cn gi l ribulozo 1,5 biphotphat
(RuBP).
CO2 lin kt thnh RuBP trong qu trnh ch yu gi l s c nh cacbon
(carbonixation) to ra hai phn t 3-cacbon l photphoglixerat (PGA) (Hnh 6.18).
Enzym thc hin vai tr xc tc cho phn ng ny l ribulozo biphotphat
cacboxilaza (thng gi mt cch ngn gn rubisco). y l mt enzym rt ln c
mt trong phn cht nn ca lc lp. N hot ng vi tc rt chm, ch x l
khong 3 phn t RuBP/giy, trong khi mt enzym in hnh thng x l khong
1000 phn t c chgiy. Do n hot ng chm nh vy nn t bo quang hp
cn c nhiu bn sao ca rubisco. Trong l cy in hnh, rubisco chim hdn 50%
208

tng protein ca t bo. Do , ngi ta coi rubisco l protein phong ph nht trn
hnh tinh.
H 2C -O -0

HC -O H
0

H 2C-O -0

c = o

phn t
photphoglycerat
(PG)
2

ICIO2 + H2O

HC-OH
I
HC - O H

Rubisco

H2C - o -

H 2C - 0 -
i

HC-OH

Ribuloz
1,5 biphotphat
(RuBP)

Hlnh 6.18. Bc ch yu trong qu trinh c nh cacbon. Melvin Calvin v ng s

tn/ng i hc Callomia pht hin ra bc u tin trong con ng c <^nh cacbon m
sau ny gi t chu trnh Calvin. H phdi to quang hp vi kh cacbonic c hot tnh
phng x ( COi). Nh theo di ng hung chuyn ho hay s phn (fate) ca nguyn t
cacbon c hot tnh phng x, h Uiy rng u tin, nguyn t cacbon ny lidn kt vi
phn t ribulozo 1,5 biphotphat (RuBP) v ngay sau thl tch ra to nn 2 phn t
photphogllxerat (PGA). Mt trong hai PGA cha nguyn t cact>on c6 hot tnh phng x.
Nm 1948, cc tc gi tch rlng erym xc tc phn ng c inh cacbon c bit ny;
mblsco

6.4.I.2. Chu trnh Calvin

Cc phn ng khng ph thuc nh sng ca quang hp c th xy ra d dng
trong ti. Phn ng c' nh cacbon tin hnh d dng do RuBP l phn t giu
nng lng. Song, vic to kh nng cung cp RuBP cn mt lot cc phn ng m
tiu th lng ln ATP v NADPH. Chng to nn chu trnh gm cc bc phn
ng c6 enzym xc tc ging nh chu trnh Krebs. Do chu trnh bt u khi CO2 lin
kt vi RuBP to ra PGA v PGA cha 3 nguyn t cacbon, nn qu trnh ny
c gi l quang hp C 3 ( C 3 p h o to s y n th e s is ).
Trong mt lot cc phn ng (Hnh 6.19), 3 phn t CO2 c rubisco c nh
to ra 6 phn t PGA (cha tt c 6x3=18 nguyn t cacbon, trong 3 cacbon
t CO2 v 15 t RuBP). V sau, 18 nguyn t cacbon tri qua mt chu trnh phn
ng ti sinh 3 phn t RuBP c s dng trong bc khi u (cha 3x5=15
nguyn t cacbon). Chu trnh li 1 phn t glixeraldehit 3-photphat (3 nguyn
t cacbon) nh l s thc thu (netgain) (Hnh 6.19).
209

3 phn t
lacbon dioxii
(CO.

Rubsco

3 phn t

6 phn t
-photphglycerat (3C ) (PGA]|

S c nh cacbon

3 ADP

6AD P

6 phn t
S Ui oh
RuBP

3 ATI

1,3-biphotphatglycerat (3C)

6NADPH

G 3 P d ebyd rogenazv
S o ngc
cia ng phn

5 phn

6 NADP*
6 phn t

Glyceraldehit 3-photphat (3C)

Glyceraldehit 3-photphat (3C ) (G3P)

1 phn t
Glyceraldehit 3-photphat (3C) (G3P)

T
G1 uco 2 v
cc teg khc

Hlnh 6.19. Chu trnh Calvin. m5f ln 3 phn t CO2 t vo chu tilnh, mt phn t ca
hp cht 3-cacbon l glbceraldehit 3-photphat c to ra. Lu rng qu trnh ydu cu
nang lung c d tr trong ATP v NADPH do phn mg ph Uiuc nh sng to ra. Qu
trnh ny xy ra trong phn cht nn ca lc lp

Phng trnh thc ca cc phn ng khng ph thuc nh sng l :

3 CO2 + 9 ATP + 6 NADPH + nc -> glixeraldehit 3-photphat + 8 P + 9 ADP + 6 NADP*
Chu trnh phn ng ny c gi l chu trnh Calvin {Calvin cycle) theo tn
nh pht minh chu trnh - Melvin Calvin thuc trng i hc California,
Berkeley. Vi ba vng tun hon kn ca chu trnh, ba phn t kh cacbonic i vo,
to ra mt phn t glixerandehit 3-photphat (G3P) v ti sinh 3 phn t RuBP.

210

6.4.I.3. Hiu aut ca chu trinh Calvin

Glixerandehit 3-photphat l sn phm ca chu trnh Calvin li l sn phm
trung gian ch yu trong ng phn (ng 3-cacbon). Phn ln G3P c bi
xut khi lc lp vo t bo cht ca t bo v ti xy ra s o ngc mt s
phn ng trong ng phn, cho php n bin i thnh fructozo 6 - photphat v
glucoz 1 - photphat v t thnh saccaroza (sucros) - mt ng dn truyn ch
yu trong thc vt (saccaroz - ng n ph bin l mt disaccarit c to ra t
fructoz v glucoz).
Trong thi gian quang hp ang din ra mnh m, mc glixerandehit 3photphat trong cht nn ca lc lp tng ln. Kt qu l, mt s G3P trong lc lp
bin i thnh glucoz 1 -photphat trong mt lot cc phn ng tng t vi phn
ng xy ra trong t bo cht nh mt s phn ng o ngc ca ng phn, v
sau, glucoz 1-photphat t hp thnh mt cao phn t khng ho tan, to nn chui
tinh bt di c d tr di dng ht tinh bt to ln trong lc lp.
P h n ln th c v t k t hp khi cacbonic th n h tag nh ch u trn h
p h n ng gi l chu trn h Calvin. Phn ng ph thuc n h sng to ra
ATP v NADPH c chc n n g kch hot cc phn ng ca ch u trn h
Calvin nh ng bn th n chu trin h Calvin c th xy ra tro n g ti.
6.4.2. H h p s n g

S tin ho khng nht

thit dn n cc gii php t
u. ng hn, n u tin cc
gii php d thc hin m c
th pht sinh t gii php khc
tn ti. Quang hp khng
phi l ngoi l. Rubisco enzym
xc tc phn ng c' nh cacbon
ch yu ca quang hp (HinK
6 .20 ) r rng b sung cho t bo
quang hp mt gii php ring.

Enzym ny c mt hot tnh

enzym th hai m ngn cn chu
trnh Calvin l oxi ho ribulozo
1,5 biphotphat. l qu trnh
mang tn h hp sng
iphotorespiration) trong CO
c gii phng m khng to
ra ATP hoc NADPH. V l do

Hlnh 6.20. cu trc ca domatn xc tc ca

rubisco. Kiu cu trc l mt thng ap. 8 si p song
song th hin bng mu vng, 8 si a c6 mu hng v
phn cn li ca phn t l mu xanh

, h hp sng lm tn hi hot ng ca quang hp.

211

Qu trnh cacboxyl ho v oxi ho ca ribulozo 1,5 biphotphat c xc tc

cng v tr trn rubisco v cnh tranh nhau. Di iu kin bnh thng 25c, tc
ca phn ng cacboxyl ho gp 4 ln tc phn ng oxi ho, chng t rng
20% cacbon do quang hp c"nh b mt i do h hp sng gy ra. S mt mt ny
tng ln ng k khi nhit tng ln, do tc ca phn ng oxi ho tng ln vi
nhit nhanh hn nhiu so vi phn ng cacboxyl ho.
S mt cacbon c nh do h hp sng gy ra l khng bnh thng. Thc
vt s dng chu trnh Calvin c' nh cacbon kh quyn gi l thc vt Cg hay cy
C3, b mt khong gia mt phn t cho n mt na cacbon do quang hp c nh

theo cch ny, ph thuc ch yu vo nhit . Trong vng kh hu nhit i, c

bit cc vng trong nhit thng trn 28c th y l mt vn nghim
trng v c mt tc ng khng nh ln nn nng nghip nhit i.
V mt c ch, ta c th hnh dung
h hp sng theo phng trnh n

gin sau;
RuBP+Oa - PGA+photphoglicolat

CO2

Thc ra, con ng h hp sng

phc tp hn nhiu (Hnh 6.21). Theo
C.R.Somerville (2001) h hp sng xy
ra trong ba bo quan: lc lp,
peroxisom v ty th thng nm cnh
nhau trong t bo cht ca t bo. Khi
c nh sng, photphoglicolat do hot
ng ca RuBP oxigenaza to ra b bin
i thnh glcoat nh xc tc ca
enzym

photphoglicolat

photphataza

trong lc lp. Glicolat thm nhp vo

peroxisom v bin i thnh glioxilat

Glycerate Glcerate

NDts

- 1 /2 0 ,

NADHOH-Pyruvate Glyoxylate

nh xc tc ca enzym glicolat oxidaza.

Gioxilat chuyn amin ho thnh gixin

(Gly) cng bng phn ng enzym.
Trong ty th glixin bin thnh CO2,

^ 20 G-Ser
Peroxisome

NH3 v nhm metylen ca metylen

tetrahidroMat (Ci-THF). Glixin v CiTHF ngng t to Serin (Ser). Serin i

vo peroxisom v b loi nhm amin
(deamin ho) bin thnh hidroxylpiruvat
(OH-piruvat) v b kh thnh glixerat
nh hidroxipiruvat reductaza. Glixerat

Ser

THF

Ty th

G l

Gly

C,-THF

NADH NAD

thm nhp vo lc lp v b photphoril

ho thnh 3-photphoglixerat (PGA) Hnh 6.21. S nit gn ca cn ng h hp sng

212

cht trung gian ca chu trnh Calvin. NH3 c gii phng trong qu trnh

decacbonxin ho glixin v c glutamin synthetaza s dng to glutamin (Gln).

Glutamic axit (Glu) synthaza ngng t 2 -oxoglutarat (2 -OG) v Gln to hai
phn t Glu. Phn t dn truyn dicacboxilat {dicarbolylate transporter) (DT)
mng bao lc lp c chc nng chuyn oxoglutarat, Glu v Gln qua mng bao lc
lp. Mt cch tng qut, hai phn t photphoglicolat (2x2C) bin i thnh mt
phn t photphogixerat (3C) v mt phn t CO2 (IC) hon thnh con ng h
hp sng. Chui phn ng ny c gi l h hp sng v vi s c mt ca nh
sng photo), oxi c hp th v CO2 c to ra.
R rng, h hp sng c nn nhn nh l mt qu trnh lm tht thot hiu
qu quang hp ca thc vt C3, c th l:
1 . Ribulozo
2.

1,5-bisphotphat b mt i khi chu trnh Calvin

S c nh CO2 b o ngc: O2 c tiu th v CO2 c gii phng

3. Ch mt phn ca cacbon quay tr li lc lp

4. ATP b tiu th mt cch khng cn thit
Vic nh gi bt k chc nng c li no ca h hp sng l rt kh khn.
Thc cht l h hp sng hot ng chng li ton b qu trnh quang hp, xo b
nhng g do quang hp to ra. V l do nn trong thi gian qua c nhiu n lc
nghin cu su rng nhm bin i di truyn thc vt C3 c bit cy lng thc

th gii nh la mi, la mch, la nc... sao cho chng mt i tp tnh h hp

sng. Vi cng ngh di truyn s cho php chng ta bin np gen loi C4 vo mt s
cy Csnh thuc l, c chua, Arabidopsis v la. Gn y nhm nghin cu Nht
v c (S.Suzuki, J.N.Burnell etal, 2000) v W.C.Taylor vin nghin cu la quc
t - Philippin (2001) c gng lm tng nng CO2 v tr ca Rubisco nh bin
np enzym decacboxin ho (enzym xc tc tch nhm cacboxin COO ) quang hp C4

- photphoenonpiruvat cacboxikinaza (PCK) vo lc lp ca t bo tht l la. Kt

qu th nghim cho thy rng s biu hin lc v tr (ectopic expression) ca PCK
trong lc lp la c th lm bin i mt phn dng cacbon trong t bo tht l la
thnh con ng quang hp kiu
C3 v hng n la C4 (C< ric).

lm tng kh nng quang hp ca thc vt

6.4.2.1. Conng axt cy mng nc

Mt s thc vt mng nc nh da, xdng rng thch nghi sng trong mi
trng kh hn nh sa mc v cc mi trng nng tin ho nn hai phng
thc c bn i ph vi vn ny. Mt cch tip cn l thc y quang hp
trong vng nng nhiu cy mng nc (d tr nc) thuc h Crassulaceae v
mt s' thnh vin khong hai t cc nhm cy khc (Hnh 6.22). Phng thc c
nh cacbon ny c gi l s chuyn ho axit cy mng nc (crasssulaceaeacid
metabolism - CAM) theo tn h thc vt Crassulaceae v cy mang tn thc vt
CAM (Hnh 6.17). Trong thc vt ny, l kh (l m) chuyn ho m trong m v
213

ng trong ngy. Kiu ng m l kh ny ngc vi kiu ng m trong phn ln

thc vt. L kh ng trong ngy c6 tc dng ngn chn s mt nc v lm gim
h hp sng nh cn tr CO2 ri khi l nh duy tr t s cao ca CO2 vi O2 bn
trong l. CO2 cn tng hp ng c b sung t cc phn t hu c do chu
trnh Calvin to ra trong m.
Vy nt c trng ca quang hp theo con ng CAM l g?
1.

Thc vt CAM s dng

pepco (PEP cacboxilaza =
photphoenonpiruvat
cacboxilaza) c dnh kh
cacbonic ban m khi l kh
m, to malat (axit malic) v
d tr malat trong khng bo
ln trong t bo tht l cho n
ngy tip theo.
2. Trong khi quang hp C4
(xem phn sau) th hin s
phn chia v khng gian - s c

nh cacbon xy ra trong t bo
tht l, cn chu trnh Calvin
xy ra trong t bo bao b
mch. CAM l mt th d v s
phn chia quang hp theo thi
gian. Malat c to ra lc
m th c phng thch khi
khng bo t bo tht l trong
ngy v n c chu trnh
Calvin thu nhn trong cng t
Hnh 6.22. Thc vt CAM quen thuc. Da v nhiu
bo m lc ny c mt NADH
cy mng nc nhlt l khc l thc vt CAM
v ATP t phn ng ph thuc
nh sng. L do c bn cho s phn chia theo thi gian l t bo phi bo ton
nc. bo ton nc, thc vt CAM m l kh lc m v do chl thi gian
ny th CO2 kh quyn mi sn c cho enzym pepco. Khi nh sng mt tri n cy,
l kh ng i v CO2 khng th thm nhp vo cy.
3. CAM thng khng hiu qu hoc nh quang hp

C3

hoc nh quang hp

C4, nhng n cho php cy sinh sng di cc iu kin bt li. Mi trong ba dng

quang hp c th li v bt li ring: Thc vt CAM khng phi l c th quang hp

hiu qu, nhng chng c th tn ti di cc iu kin kh hn, thc vt C4 l c
th quang hp hiu qu, nhng chng khng cnh tranh tt vi thc vt C3 di
nhit
Ni khc i, thc vt C4 mn cm vi nhit lnh.

25c.

214

Thc vt CAM s dng pepco tin hnh c nh kh cacbonic lc

m khi l kh m. Trong ngy, CO2 d tr c th thm nhp vo chu
trinh Calvin.
Bng 1. So snh tnh cht khc bit ca ba loi thc vt ny

Ba con ng c nh CO2
Cs
C4
T bo bao b T bo bao b
mch thiu lc lp mch c lc lp
Pepco
Rubisco
30 - 40C
15 - 25c
22 0,3
39 17(*)

CAM
Cutrcl
C khng bo ln
trong t bo tht l
Emymcsdng
Pepco
35C
Nhit ti u
Thp - c th bin
T s nng sut
i
(tn/ hecta/nm)
(*) Theo D.K.Northington & J.R.Goodin. The Botanical world. St.Louis, 1984)
Tnh cht

6.4.2.2. Conng C4
Cch tip cn th hai c mt s cy tho nh ng, ma, la min s dng
vt qua chu k kh hn, nng trong nm l quang hp C4 {C4 photosynthesis).
Quang hp C 4 xy ra trong hai loi t bo: t bo tht l nm ngoi bao ly t bo
bao b mch trong m bao quanh b mch dn hay gn l (Hnh 6.23). u th ca
thc vt C4 l c kh nng to ra nng CO2 cao bn trong t bo bao b mch,
to thun li enzym rubisco thc hin phn ng cacbonxin ho (Hnh 6.24). u
tin chng xc tc phn ng cacboxyl ho sn phm chuyn ho 3-cacbon gi l
photphoenonpiruvat, to ra oxaloaxetat (4 cacbon). Oxaloaxetat li bin i thnh
sn phm trung gian malat v malat c chuyn vo t bo bao b k bn. Khi
bn trong t bo bao b m ch, m alat b decacbXyl ho (li nhm

-COO ) to

ra

.T b o
m& k b a y t
ao b m cb

Lkhl

nr
kh

b o

Hnh 6.23. So snh gii phu l ca thc vt C3 v c,. l C3 , CO2 c t bo tht l

c khong gian bo rng hp th (t bo m khuyt), l C4. CO2 c gl phng t hp
cht C4 bn trong t bo bao b mch v t bo ny tng i khng thm VI COj nn to
ra nng cao ca COj

215

piruvat v gii phng CO2. Do t bo bao b mch khng thm i vi CO2, nn CO2
c lu gi bn trong t bo vi nng cao. M.S.B.Ku, G.E.Edwards v nhiu
ngi khc (2000 ) gi y l c ch tp trung CO2 trong thc vt C4 (CO2 concentrating mechanism) v rng chc nng ch yu ca con ng C4 l tp
trung CO2 hay "c c" CO2 trong cc t bo bao b mch bn trong ni Rubisco
nh v. Tip theo, puvat quay li t bo tht l, hai lin kt cao nng trong
phn t ATP tch ra bin piruvat tr li thnh photphoenopiruvat (PEP), do
hon thnh chu trnh.

T b o tht

Hlnh 6.24. S c nh CO2 trong thc vt C4 . Qu trinh ny gi l con ng C4 do hp

cht xut hin u tin l oxaloaxetat l phn t cha 4 cacbon

Cn liu rng phn ng cacbonxin ho C2 kh quyn xy ra u tin trong

phn bo tng (cytosol) ca t bo tht l do enzym PEP cacboxilaza xc tc
(PEPC), to ra oxaloaxetat (OAA) c 4- cacbon. OAA i vo t bo tht l v b kh
thnh malat. Malat i vo t bo bao b mch v din bin phn ng tip theo s
c trinh by theo hnh 6.25 phn s pha phi.

216

T bo bao b mch

T b o tht l
NADPH
OAA

Mal

M al
O A A
NADP-M DH

1
2

&
P
p E P PEP

PPDK

.
a
<o
s

pr

Pyr

ATP

HCO
CO

RuBP

s ^ADPH
d
^CO

t
CO.

Lc lp

Lc lp
Bo tng

Botg

Hlnh 6.25. Con ng quang hp C4 trong cy ng. Chc nng ch yu ca con ng

l tp trung COj teong t bo bao b mch, ni rubisco nh v v do c ch hot tnh
oxigenaza v h hp sng kt hp. OAA: oxaloaxetat, NADP-MDH; NADP malat
dehidrogenaza, Mal: malat, PEP: photphoenolpiruvat, PEPC: PEP cacboxitaza, Pyr: piruvat,
RuBP: ribulozo 1,5-blsphotphat, Rublsco: ribulozo blsphotphat cacboxilaza oxigenaza,
PPDK: piruvat octophotphat diklnaza, NADP ME enzym malic ph thuc NADP

T bo bao b mch

Tb o tht l
6 N A D P H ,6 A T P
6 tr io s e -p i 6 P G A 3 triose-P
glycerate-

3 glycerate

3
^ o
3 ATP. 3 NADPH^
33NH
NHj-------- ^ 3 glutamate

3 N A D P H .6 A T P

6C3

60,.3 CO,^
V

J *9
- 9 triose-P

9A T P
18 A T P
Tng -3 0 A T P .9 N A D P H
RuBP - 3.3 ATP, 1 NADPH

9 CO.

18 ATP.9NAPH
- 2 ATP, 1 NADPH (ATP/NADPH tioDg BS - 2)

Tng S trn RuBP

_________________ 5 .3 T P .2 N A D P H ________

___________

Hinh 6.26. H qu gi nh ca h hp sng v yu cu nng lng cho vic pht sinh

CO trong t bo bao b mch trong thc vt C4. S gii thiu s tiu th RuBP nh mt
h qu ca h hp sng Uong ttic vt C4. 6 0 j v 3C0j phn ng vi 9RuBP, 3C 0j c
gii phng trong t bo bao b mch trong con ng glicolat (con ng h hp sng). VI
mt vng ca chu lnh C4, RuBP ng 5.3ATP v 2NADPH s phi tiu th trong t bo
tht l v t bo bao b mch. BS: t bo bao b mch

217

T bo bao b mch l ni nh cc enzym thc hin chu trnh Calvin trong

thc vt C4. Nh c ch tp trung CO2 nn t bo bao b mch c nng CO2 kh
cao v do c ch qu trnh h hp sng. Nhng do mi phn t CO2 khi c
chuyn qua mng bao lc lp t bo bao b mch phi chi dng hai lin kt ATP
cao nng v do su cacbon phi c c' nh to mt phn t glucoz nn t bo
bao b mch cn dng thm 12 phn t ATP to ra mt phn t glucoz c 6cacbon. Do vy, tron g quang hp C 4 , ph tn n n g lng tn g hp glucoz l gn
gp hai ln ph tn ca quang hp C 3 : 30 phn t ATP tng phn vi 18 (Hnh
6.26). Tuy nhin, quang hp C 4 c li th trong vng kh hu nng.
H h p sng r t ngn chu trin h quang hp. Thc v t CAM v th c
vt C4 vt qua c iu kin bt li v kh hu thng qua cc bin
i v kin tr c l v chui phn ng ho sin h quan g hp m c tc
dng lm t n g t i ch nng COj. Thc v t CAM t c h rin g p h n
ng to CO] theo thi gian, cn thc vt C4 theo khng gian.

\nt B'

Hlnh 6.27. Lc lp v ty U: hon thnh chu trnh nng lng. Chu trnh nc v kh oxi
gia lc lp v ty th bn trong t bo thc vt to ra glucoz v CO2. T bo c lc lp i
hi ngun COj v nc bn ngoi pht sinh glucoz v 0X1. T bo khng c lc lp nh
t bo ng vt, cn ngun gtucoz v oxi bo ngoi v pht sinh COj v nuc

218

* Chu trnh nng lng

Qu trnh chuyn ho nng lng ca lc lp (chdng 6) v ca ty th (chng
5) c lin quan mt thit vi nhau. Quang hp s dng cc sn phm ca h hp
nh l c cht khi u v h hp dng sn phm ca quang hp di dng c cht
khi u (Hnh 6.27). Thm ch chu trnh Calvin cn dng mt phn ca con ng
ng phn nguyn thu tin hnh theo chiu ngc li to glucoz. Dng nh
cc protein c bn tham gia trong truyn in t thc vt c quan h vi protein
trong ty th v trong nhiu trng hp chng thc s ging nhau. Do quang hp
xut hin t lu trc khi thc vt ra i nn mi sinh vt u ph thuc trc tip
hoc gin tip vo quang hp ly nng lng cung cp cho 8 sng.
TM TT KHI NIM
6.1. Q uang hp xy ra trong lc lp
nh sng c thc vt, to v mt s vi khun s dng trong qu trnh
quang hp bin cacbon kh quyn (CO2) thnh hidratcacbon.
6.2. N ghin cu v quang hp: Mt ch n g ng thc nghim
Mt lot cc th nghim n gin chng minh rng thc vt hp th nng
lng t nh sng v dng n bin nguyn t cacbon ca CO2 v nguyn t
hidro ca nc thnh phn t hu c.
6.3. Sc t hp th nng lng t nh sn g m t tri
nh sng gm cc b nng lng gi l photon. Bc sng nh sng cng
ngn, nng lng ca photon cng nhiu.
Khi photon c sc t hp th, in t trong sc t c nng ln mt mc
nng lng cao hn.
Quang hp tin ho trong c th ng dng vi vi khun lu hunh ta m
s dng mt h gm phn t dip lc v phn t sc t ph (quang h)
truyn nng lng kch thch ca photon cho mt phn t n sc t.
Phn t chuyn mt in t cho chui truyn in t, nh kch hot
bm proton v cui cng mang in t tr li cho sc t trong qu trnh quang
photphoril ho vng.
Cc hu du (con chu = d e s c e n d a n t) ca vi khun quang hp ban u s
dng hai quang h m c th pht sinh nng lng dng H2O hn l
H2S nh mt ngun hidro.
Trong cc sinh vt c hai quang h, ban u nh sng c quang h II hp
th, tch mt in t khi mt phn t dip lc a l Peso- Qu trnh ny c hai
tc ng: (1) s thiu vng ca in t cao nng lm cho quang h II phi tm
219

kim in t khc lp l trng in t ca Pes. l in t b t ngun t

qu trn h p h n ly nc, gii phng O 2 nh mt s n phm p h v (2 ) in t
cao nng c chuyn cho quang h I c tc dng kch hot bm proton trong
qu trnh v n n tng hp ho thm ATP.

Khi in t t Peso n quang h I, P 700 hp th mt photon ca nh sng, tip

nng lng (nng ng ho) cho m t trong cc in t. in t t quang h I
c chuyn cho m t cht n h n in t s cp k h NADP* th n h NADPH.
Vic s dng NADPH ch khng dng NADH cho php thc v t v to duy
tr qu trnh quang hp v h hp oxi ho tch khi nhau.
6.4. T bo dng n n g lng v lc kh o phn ng s n g th u c
t n g hp c c p h n t h u c
ATP v lc kh do phn ng ph thuc n h sng to ra c t bo s dng
C n h cacbon trong m t lo t cc phn ng k h n g ph thuc n h s n g gi

l chu trn h Calvin.

Trong qu trnh ny, ribulozo 1,5- biphotphat (RuBP) b cacboxyl ho v sn

phm tch th n h hai phn t photphoglixerat (PGA). Cc phn t PGA tr i
qua mt ot phn ng to cc phn t glxerandehit photphat v mt s
cc phn t ny c t bo s dng t i sinh (ti to) RuBP. P h n cn li
ca cc phn t glixerandehit photphat thm nhp vo qu trnh chuyn ho
ca t bo di dng cacbon va c c nh (cacbon mi c nh).
RuBP cacboxilaza - enzym c chc nng c nh cacbon trong chu trn h Calvin
cng thc hin phn ng oxi ho vi cc sn phm ca quang hp - qu trnh
c gi h h p sng.

Nhiu thc vt nhit i c ch h hp sng nh tiu th ATP lm tng

nng CO ni bo. Qu trnh ny gi l con ng C4 lm tng gn gp i
ph tn nng lng cao ca s tng hp gucoz. kh hu nng, ph tn nng
lng nh hn ph tn to phn t gucoz m theo cch khc s b bin i
tr li thnh kh cacbonic do h hp sng gy ra v b mt i trong thc vt
C3 v c c nh li trong thc vt C4.
GII THCH CC THUT NG CHNH
1

. P h o to s y s te m (quang h). Cc sc t dip c c t chc th n h dn v t hp

(in te g r a te d u n it) gi l quang h c chc nng thu gp nng lng kch thch ch
yu khuch i kh nng thu nh sng ca cc phn t sc t c th.

2.

R e d u c in g p o w e r Oc kh), Ngun hidro cn k h COa tng hp phn t

hu c. sinh vt vi hai quang h, mt quang h-c6 vai tr kch hot qu trnh

tng hp NADPH, do b sung lc kh. Sinh v t.j^ m t quang h th chng
phi th u gom ngun hidro v c hiu qu nh hn.
220

3. Carbon x a tio n (s c nh cacbon). Quang hp c nh cacbon t mi trng

(thng dng CO2 kh quyn) v bin i thnh hidratcacbon thng qua cc
phn ng khng ph thuc nh sng ca quang hp.
4. P h otoresp iration (h hp sng) l qu trnh m nh thc vt v to thnh
thong bin cacbon c quang hp c' nh tr li th n h CO 2 v nhit lm tn
hi kt qu t c ca quang hp. S d xy ra qu trnh ny l do enzym
cacboxilaza lin kt vi oxi cng nh vi (^2. Do c ngi gi RuBP
cacboxilaza l enzym c chc nng kp trong thc vt.
5. C4 p h o to sy n th esis (quang hp C4). Mt qu trnh quang hp xen k (a lte r n a tiv e
p h o to s y n th e tic p r o c e s s ) thc vt trong vng kh hu nng s dng chng li
s mt CO2 thng qua h hp sng. Thc vt C4 c c ch tp trung CO2 bn
trong cc t bo chuyn ho (t bo bao b mch) cnh tranh vi O2 v tr
xc tc ca enzym cacboxilaza.

CU HI N TP
1

. Van Helmont tin hnh th nghim nh th no xc nh rng thc vt

khng nhn thc n t t?

2. Hy trnh by phng trnh tng qut ca quang hp. Cc nguyn t oxi trong
O2 c to ra trong qu trnh quang hp bt ngun t u? Cu tr li cho vn
ny c xc nh nh th no?
3. n v c bn ca nng lng nh sng l g? Bc sng no ca nh sng thy
c nng lng hot ho cao nht v bc sng no c nng lng hot ho b
nht?
4. Nng lng nh sng c phn t sc t hp th nh th no? Ti sao nh sng
do sc t dip lc phn x c mu lc?
5. Englemann tin hnh th nghim th no cho rng dip lc c th tham gia
trong quang hp?
*

6.

Hy trnh by ba giai on ch yu trong cc phn ng ph thuc nh sng ca

quang hp? Hy nu chc nng ca dip lc trung tm phn ng? Chc nng
ca cht nhn in t s cp l g?

7. Gii thch v quang hp trong vi khun lu hunh l mt qu trnh vng? Trnh

by hiu qu nng lng ca quang hp theo phn t ATP c tng hp trn
in t.
8.

Hai quang h trong thc vt v to hot ng nh th no? Giai on no p h t

sinh ATP v giai don no pht sinh NADPH?

221

9. iu g xy ra trong cc phn ng khng ph thuc nh sng ca quang hp?

Cc p h n ng khng p h thuc n h sng s dng ATP v NADPH th no
chng c to ra trong p h n ng ph thuc n h sng?
10. Trong chu trn h Calvin, iu g xy ra vi ribulozo 1,5-biphotphat? iu g xy
ra vdi gxerandehit 3-photphat (G3P)?

11. Phn ng ph thuc nh sng xy ra u trong thc vt

ph thuc n h sng xy ra u?
12

C 3?

Phn ng khng

. H hp sng l g? Li th no lm cho thc v t C4 vt tri thc v t C3 c h

hp sng? iu bt li no lm cho thc vt C 4 phi hn ch s phn b"ch yu
vo vng nng ca qu t?

C HI SUY LUN
1. Gii thch i th ca mi quang h c nhiu phn t sc t nhng ch mt dip
lc tru n g tm p h n ng. Ni cch khc, ti sao khng lin k t trc tip mi
phn t sc t vi m t ch t n h n in t?
2.

H ai quang h Peso v p 70QCU1 k h u n ix to v thc vt to IIGII m t cit 0 X1

ho c th p h n ly nc. Vy s tin ho tip sau ca h hp t bo c th khc
bit nh th no nu hai quang h khng tin ho v t t c cc sinh v t quan g
hp c h n c h vi q u an g photphoril ho vng do vi k h u n l u h u n h s

dng?

3. Theo l thuyt, mt thc vt c gi hon ton trong t vn c th to ra

glucoz nu n c b sung cc phn t no?

222

TI LIU THAM KHO

[1]

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[4]
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223

NHA XUT BN I HC QUC GIA HA n i

16 Hng Chui - Hai B Trng - H Ni
i n thoai: (0 4 ) 9 7 2 4 8 5 2 : (04) 9 7 2 4 7 7 0 . F a x : (04) 9 7 1 4 8 9 9

Chu trch nhim x u t bn:

Gim c:

PHNG QUC BO

Tng bin tp:

NGUYN b t h n h

Bin tp:

SAO MAI

Chbn:

SAO MAI

Trinh by ba:

HI ANH

SINH HC T BAO
Ms; 1K-39H2007
In 1.000 cun, kh 19 X 27 cm ti Nh in i hc Quc gia H Ni
S xut bn: 249 - 2007/CXB/08 - 46/HQHN, ngy 04/04/2007
Quyt nh xut bn s: 394 KH/XB
In xong v np iu chiu qu III nm 2007.