1

Chng 1
Trao i cht v nng lng sinh hc

1.1. Khi nim chung v trao i cht
Mi c th sng u tn ti trong mi trng v lin h mt thit vi
mi trng . Hin tng c th ly mt s cht t mi trng kin to
nn sinh cht ca mnh v thi ra ngoi nhng cht cn b c gi l s
trao i cht.
S trao i cht gii v sinh khc vi gii hu sinh. gii v
sinh, trao i cht lm cho cc cht hu c v v c b phn hu. V d,
vi (canxi carbonate) b xi mn v H
2
CO
3
c trong nc tc dng vi
vi thnh canxi bicarbonate, m b i ho thnh mt s cht khc l do tc
dng vi oxy.
th gii sinh vt, mi c th sng lun lun trao i cht vi mi
trng, ly thc n vo chuyn ho thnh cc cht s dng cho c th v
thi ra ngoi cc cht cn b. Qu trnh c thc hin l do cc bin
i ho hc lin tc xy ra trong c th. Ton b cc bin i ho hc
c gi l s trao i cht.
Qu trnh trao i cht gm nhiu khu chuyn ho trung gian. Mi
chuyn ho l mt mt xch ca mt trong hai qu trnh c bn: ng ho
v d ho.
ng ho v d ho l hai qu trnh i lp, nhng li thng nht vi
nhau trong mt c th: chng xy ra ng thi v lin quan mt thit vi
nhau. Cc cht c tng hp nn trong qu trnh ng ho l nguyn liu
cho qu trnh d ho (v d gluxit l sn phm ca qu trnh quang hp, l
nguyn liu cho qu trnh h hp). Nng lng gii phng ra trong qu
trnh d ho c s dng mt phn cho qu trnh tng hp.

1.2. Nng lng sinh hc

H thng sng cn nng lng chuyn ng, ln ln, tng hp
cc phn t sinh hc v vn chuyn ion, phn t qua mng.
Cc c th ly nng lng t mi trng sng v s dng nng
lng thc hin cc qu trnh sng c hiu qu.
nghin cu nng lng sinh hc i hi phi c hiu bit v
nhit ng hc, mt s nh lut, nguyn l m t ngun, trao i nhit,
nng lng v vt cht trong h thng nghin cu.
2
Nhit ng hc cho chng ta xc nh qu trnh ho hc v phn ng
c th t xy ra hay khng.
Mc d nhit ng hc l khi nim phc tp, nhng n da trn ba
nh lut tng i n gin v d hiu.
Mt vi nguyn l ca nhit ng hc c bn c a ra trong
chng ny bao gm phn tch ngun nhit, sn sinh entropy, hm nng
lng t do v mi lin quan gia entropy v thng tin.
Chng ny cng cp n ATP v nhng hp cht cao nng
khc.

Khi nim v nhit ng hc c bn
Bt k s quan tm no ca nhit ng hc cng phi phn bit gia
h thng v mi trng.
H thng l mt phn ca v tr m chng ta quan tm, trong khi
mi trng l gm tt c nhng g cn li. C ba trng thi c bn: h
thng c lp, h thng ng v h thng m.
H thng c lp: Khng c s trao i cht v nng lng vi mi
trng.
H thng ng: C trao i nng lng, nhng khng c trao i
cht vi mi trng.
H thng m: C trao i cht v nng lng vi mi trng.
C th sng l h thng m in hnh c trao i cht (dinh dng
v sn phm thi ra) v nng lng (nhit t trao i cht) vi mi trng.
nh lut 1: Nhit, cng v cc dng nng lng khc
Trc y trong s pht trin ca nhit ng hc ngi ta cho rng
nhit c th bin i thnh nhng dng nng lng khc v tt c cc
dng nng lng mt cch c bn c th bin i thnh mt s dng khc.
nh lut 1 ni rng: tng nng lng ca mt h thng c lp l
khng thay i.
Cc nh nhit ng hc m phng thnh mt hm ton hc
nghin cu s bin i nhit v s dng cng trong nhng h thng nhit
ng hc. Hm ny c gi l nng lng ni nng, thng k hiu l E
hoc U. Nng lng ny ch ph thuc vo trng thi hin ti ca mt h
thng v v vy c coi l hm trng thi. Nng lng ni nng khng
ph thuc vo h thng xy ra nh th no v v vy khng ph thuc vo
ng hng. Ni mt cch khc l chng ta c th thay i h thng
bng bt c con ng no v cho n khi no h thng tr v trng thi
ban u, nng lng ni nng s khng thay i.
Nng lng ni nng, E ca h thng c th thay i nu ngun
nng lng vo hoc ra khi h thng dng nhit hoc cng cho qu
3
trnh no bin i mt trng thi ny (1) sang mt trng thi khc (2) thay
i nng lng ni nng l:
E = E
2
- E
1
= q + w (1.1)
q l lng nhit c h thng hp th t mi trng
w l cng thc hin trn h thng do mi trng
Cng c hc c nh ngha l s chuyn ng t ch ny n ch
khc, gy ra do s dng lc. C hai phi xy ra cng mi c thc hin.
V d: Mt tu ch khch cha y khch nhng khng di
chuyn, theo nh ngha nhit ng hc cng khng c thc hin.
Trong h thng ho sinh hc v ho hc cng thng lin quan vi
p sut v th tch ca h thng. Cng c hc c xc nh w = -P V
Trong P l p sut, V l s thay i th tch, V = V
2
-V
1
Cng c th c thc hin nhiu dng: c hc, in, t v ho hc.
E, q, w phi c cng n v: calorie (cal) v kilocalorie (kcal) c s
dng theo truyn thng, nhng theo n v SI: Joule c ngh nn
dng.
Enthalpy: Hm c nhiu tin li cho h thng sinh hc
Nu nh ngha cng c gii hn bi cng c hc, trong trng
hp ny E ch l thay i nhit th tch khng i. V vy nu V khng
i, cng khng c thc hin. E = q. V vy E l mt nh lng rt
tin li trong qu trnh th tch khng thay i. E khng cn thit bng
bin i nhit. V l do ny cc nh ho sinh hc, ho hc xc nh mt
hm c bit ph hp cho qu trnh p sut khng i. N c gi l
enthalpy, H c nh ngha: H = E + PV (1.2)
Nu p sut khng thay i chng ta c:
H = E + P V = q + w + P V = q - P V + P V = q (1.3)
R rng H tng ng vi bin i nhit trong qu trnh p sut
khng i.
V cc phn ng ho sinh thng xy ra trong th lng hoc rn hn
l th kh nn thay i th tch l nh v enthalpy v nng lng ni nng
thng l nh nhau.
thun li khi so snh cc ch s nhit ng hc ca cc phn ng
khc nhau th ngi ta xc nh iu kin tiu chun. Mt dung dch
ho tan trng thi tiu chun, thng s dng n v n gin l nng
1M. Enthalpy, nng lng ni nng v nhng nh lng nhit ng
hc khc thng a ra hoc xc nh cho nhng iu kin tiu chun v
c k hiu l H
0
, E
0
...
Enthalpy thay i cc qu trnh ho sinh c th c xc nh
bng vic o nhit hp th (hoc to ra) bng mt calorimeter.
4
Mt khc cho bt k qu trnh no A B trng thi cn bng, s
thay i enthalpy trng thi tiu chun c xc nh t s ph thuc
vo nhit v h s cn bng:
d (ln K
eq
)
H
0
= ---------------- (1.4)
d (1/T)




















Hnh 1.1 S thay i enthapy, H
0
ca 1 phn ng c xc nh
dc ca s RlnKeq ngc vi 1/T. minh ha phng php ny nhng
gi tr hai bn ca 327K (54,5
0
C) c nu ra. S liu c s dng tnh
H
0
54,5
0
C.
y R l hng s kh = 8.314 J/mol K
V d: trong s bin tnh nhit ca protein chymotripsinogen (qu
trnh thun nghch).
Trng thi nguyn thu (N) Trng thi bin tnh (D)
K
eq
= D / N
John F. Brandts o hng s cn bng cho s bin tnh ca mt s
protein mt s gi tr pH v nhit khc nhau (bng 1.1).

Gi tr H
0
c ngha g i vi bin tnh ca protein? Gi tr
dng ca H
0
biu din s b gy lin kt hydro cng nh gii phng
-3,21-(-17,63)
=14,42
1000
T
(K
-1
)
2,98 3,00 3,02 3,04 3,06 3,08 3,10
30
20
10
0
-10
-20
-30
R
l
n
K
e
q

3,04-3,067
=-0,027
54,5
o
C
5
nhng nhm a nc t bn trong phn t protein ban u trong qa trnh
bin tnh, nh vy s nng nng lng ca dung dch protein.

Bng 1.1 Cc ch s nhit ng hc cho s bin tnh protein

Protein
(v iu kin)
H
0

kJ/mol
S
0

kJ/mol.K
G
0

kJ/mol
G
p

kJ/mol.K
Chymotrypsinogen
(pH 3; 25
0
C)
164 0,440 31 10,9
b- Lactoglobulin (5 M
urea; pH 3; 25
0
C)
-88 -0,300 2,5 9,0
Myoglobin (pH 9;
25
0
C)
180 0,400 57 5,9
Ribonuclease (pH 2,5;
30
0
C)
240 0,780 3,8 8,4


nh lut th hai v entropy:
nh lut th hai ca nhit ng hc c m t v th hin trong
nhiu cch bao gm nhng im sau:
H thng c xu hng tin t trng thi trt t sang trng thi khng
trt t (tng entropy).
Entropy ca h thng + mi trng l khng i bi qu trnh thun
nghch. Entropy ca h thng + mi trng tng do qu trnh khng thun
nghch.
Tt c cc qu trnh xy ra trong t nhin hng ti trng thi cn
bng, l trng thi nng lng nh nht.
Mt s im ca nh lut 2 dn n khi nim entropy, l thc
o s mt trt t ca h thng, trong trng thi mt trt t l trng thi
c entropy cao.
Entropy c th c xc nh theo mt vi cch. Nu W l s cch
sp xp thnh phn ca mt h thng m khng thay i nng lng
ni nng hoc enthalpy ( l s lng ca trng thi knh hin vi c
a ra nhit , nh sng v tng vt cht). Entropy c tnh:
S = klnW (1.5)
k l hng s Boltzmann = 1,38.10
-23
J/K
nh ngha ny tin li cho tnh ton thng k, nhng dng ph bin
hn lin quan entropy n s bin i nhit trong mt qu trnh l:
dS
thun nghch
=
T
dQ
(1.6)
6
dS
thun nghch
l thay i entropy ca h thng trong mt qu trnh
thun nghch,
q l nhit c bin i, T l nhit s bin i nhit xy
ra.

nh lut 3: Ti sao0 tuyt i quan trng nh vy?
nh lut 3 ca nhit ng hc ni rng: entropy ca bt k cht no
hon ton c trt t, tinh th phi tin n 0. nhit tin n 0 K v T=
0 K entropy chnh xc = 0. Da trn iu ny c kh nng thit lp mt h
thng t l entropy tuyt i, s lng
S = Cp
T
0
dlnT (1.7)
Cp: kh nng bin i nhit p sut khng i. Kh nng nhit ca
mt cht l tng s nhit ca 1M c th d tr khi nhit ca cht
c nng ln 1 . i vi qu trnh p sut khng i n c m t
bng ton hc
Cp =
dt
dH
(1.8)
Nu kh nng nhit c th c tnh tt c nhit gia 0 K v
nhit no , entropy tuyt i c tnh i vi qu trnh sinh hc
thay i entropy c nhiu tin li hn entropy tuyt i. Thay i entropy
cho mt qu trnh c th c tnh nu thay i enthalpy v nng lng t
do bit.

Nng lng t do: Mt gi thuyt nhng l cng c tin li
Mt cu hi quan trng i vi nh ho hc, c bit i vi nh
ho sinh hc l: Phn ng s xy ra theo hng t phi sang tri? Gibbs,
mt trong nhng ngi xy dng nn nhit ng hc nhn thy cu tr li
cho cu hi ny nm trong s so snh thay i enthalpy v thay i
entropy mt nhit no , nng lng t do Gibbs c nh ngha
nh sau:
G = H - TS (1.9)
Cho bt k qu trnh A B nhit v p sut khng i.
S thay i nng lng t do c tnh:
G = H - T S (1.10)
Nu G gn = 0 qu trnh cn bng, khng i theo hng thun
hoc ngc li khi G = 0
S = H/T v thay i enthalpy v entropy l cn bng chnh xc.
Bt k qu trnh vi G khc 0 thc hin t ng n trng thi cui cng
7
c nng lng t do thp. Nu G m th qu trnh xy ra theo hng t
tri sang phi.
Nu G 0 phn ng xy ra theo hng ngc li (k hiu v gi
tr ca G cho php xc nh qu trnh s xy ra nhanh nh th no). Nu
qu trnh c G m th qu trnh t xy ra, nu G dng th qu trnh
khng t xy ra (hay t xy ra theo chiu nghch).

Thay i nng lng t do tiu chun
Thay i nng lng t do G cho bt k phn ng no ph thuc
vo cht tham gia phn ng v sn phm phn ng v cng b nh hng
bi iu kin phn ng k c nht , p sut v pH v nng ca cht
phn ng v sn phm.
Nu thay i nng lng t do cho mt phn ng l nhy cm vi
iu kin ho tan, iu g c bit c ngha cho s thay i nng lng
t do trng thi tiu chun. tr li cho cu hi ny xem xt mt phn
ng gia hai cht A v B to nn sn phm C v D
A + B C + D (1.11)
Thay i nng lng t do cho nng khng trng thi tiu
chun l
G = G
0
+ RT ln
[B] [A]
[D] [C]
(1.12)

[B] [A]
[D] [C]
= K
eq

trng thi cn bng G = 0
Chng ta c
G
0
= - RT ln K
eq
(1.13)
hoc logarit c s 10

G
0
= - 2,3 RT/ log
10
K
eq
(1.14)

N c bin i K
eq
=10
- G/2,3 RT
(1.15)
Trong bt c dng no mi lin h cho php xc nh thay i nng
lng t do tiu chun cho bt k qu trnh no nu hng s cn bng
c bit.
Quan trng hn, iu ni rng cn bng thit lp cho mt phn
ng trong dung dch l mt hm ca s thay i nng lng t do tiu
chun cho qu trnh, ngha l G
0
l cch vit khc ca hng s cn bng.
8
V d hng s cn bng xc nh bi Brandts mt s nhit vi
s bin tnh ca chymotrypsinogen c th c dng tnh s thay i
nng lng t do cho qu trnh bin tnh. V d h s cn bng 54,5
0
C l
0,27, nh vy
G
0
= - (8,314 J/mol. K (327,5K) ln (0,27)
G
0
= - (2,72 kJ/mol . ln (0,27)
G
0
= 3,56 kJ/mol
K hiu dng ca G
0
ngha l qu trnh bin tnh khng u th.
Dng gp l dng bn ca protein 54,5
0
C. Mt khc ln tng i
nh ca G
0
ngha l dng gp chim u th nh.































50 52 54 56 58 60 62
10
8
6
4
2
0
-2
-4
-6
-8
-10
G
,

k
j
/
m
o
l

Nhit
o
C
9
Hnh 1.2 S ph thuc ca G
0
vo nhit nhit trong qu trnh
bin tnh ca chymotrypsinogen

Hnh 1.2 ch s ph thuc ca G
0
vo nhit bin tnh pH = 3.
Tnh c H
0
v G
0
ca s bin tnh chymotrypsin, c th tnh S
0

s dng phng trnh (3.10).
S
0
=
T
) H G (
0 0
(1.16)
54,5
0
C (327,5 K)
S
0
= - (3560 - 533,000 J/mol) / 327,5 K
S
0
= 1,620J/mol.K

Hnh 1.3 biu din s ph thuc ca S
0
vo nhit bin tnh ca
chymotrypsin pH = 3. Gi tr dng S
0
ch rng dung dch protein
tr nn khng trt t khi protein b bin tnh. So snh gi tr 1,62 kJ/mol.K
vi gi tr S
0
bng 1.1 ch ra rng gi tr hin ti cho chymotrypsin
54,5
0
C l hon ton ln. ngha vt l ca ch s nhit ng hc cho s
bin tnh ca chymotrypsin s r hn trong phn sau.

ngha vt l ca c tnh nhit ng hc
Nhng ch s nhit ng hc cho ta bit nhng hin tng sinh ho
g? Cch tt nht tr li cu hi ny l mt ch s ring r (v d H
hoc G

) khng c nhiu ngha. Mt gi tr dng H
0
cho s bin tnh
ca mt protein c th phn nh hoc l s gy cc lin kt hydro trong
protein hoc s xut hin cc nhm a nc ra bn ngoi. Tuy vy s so
snh mt s ch tiu nhit ng hc c th cung cp nhng hiu bit bn
trong c ngha v mt qu trnh.












52 54 56 58 60
2,4
2,3
2,2
2,1
2,0
1,9
1,8
1,7
1,6
1,5
1,4
G
,

k
j
/
m
o
l
-
K

Nhit
o
C
10






Hnh 1.3 S ph thuc ca S
0
vo nhit trong qu trnh bin tnh ca
chymotrypsinogen

nh hng ca nng n thay i nng lng t do thc t
Phng trnh (3.12) ch ra rng thay i nng lng t do i vi
mt phn ng rt khc nhau so vi gi tr trng thi tiu chun nu nng
ca cht phn ng v sn phm khc vi nng hot ng (1M cho
dung dch).
Xem xt s thu phn ca phosphocreatine:
Phosphocreatine + H
2
O Creatine + Pi
Phn ng ny to nhit rt mnh v G
0
37
0
C l -42,8 kJ/mol
Nng sinh l ca phosphocreatine, creatine v Pi thng l gia 1
mM v 10 mM.
Cho rng nng 1 mM v s dng phng trnh (3.12) G

cho
thu phn phosphocreatine l



G

= - 42,8 kJ/M + (8.314J/M) (310 K)
] 001 , 0 [
] 001 , 0 ][ 001 , 0 [
ln

G

= - 60,5 kJ/M
37
0
C s khc nhau gia trng thi tiu chun v nng 1 mM
cho mt phn ng nh vy l khong -17,7 kJ/mol

Tm quan trng ca cc qu trnh kt hp trong c th sng

Nhiu phn ng cn thit gi t bo v c th chng li th nhit
ng hc, l theo hng G

dng.
Trong c s tng hp ATP v nhng phn t cao nng khc v
to nn gradient ion trong tt c t bo ng vt c v. Nhng qu trnh
ny c thc hin theo hng bt buc nhit ng hc.
11
Bng s kt hp vi cc qu trnh c kh nng t xy ra nhiu qu
trnh gn lin vi nhau c cp phn sau ca chng ny. Chng rt
quan trng trong trao i cht trung gian, phosphoryl ho oxy ho v vn
chuyn qua mng.
Chng ta c th on nhng cp phn ng kt hp s xy ra t ng
bng s thay i tng nng lng t do cho mi phn ng. V d, xem xt
phn ng t qu trnh ng phn lin quan n s bin i
phosphoenolpyruvic acid n pyruvat. S thu phn PEP gii phng nng
lng v c s dng phosphoryl ho ADP thnh ATP, mt qu trnh
m v mt nng lng khng t xy ra.

1.2.1 c tnh nng lng ca s trao i cht
Nng lng ca cc qu trnh trao i cht (nng lng sinh hc)
khc vi nng lng c thc hin trong bn cht khng sng ba c
im sau y:
c tnh th nht l s chuyn ho nng lng thnh cng v thnh
nhng dng khc m khng km theo s chuyn ho s b nng lng ny
thnh nhit nng. Xut pht t nguyn tc ny chng ta cn xem h thng
sng nh l mt ng c ho ng hc ch khng phi l ng c nhit.
c tnh th hai l vic gii phng nng lng trong cc qu trnh
oxy ho sinh hc sinh ra t t, tng phn mt, trong mt chui di cc qu
trnh k tip nhau cho n khi no tt c cc nguyn t hydro v cacbon
u bin thnh cc sn phm cui cng l CO
2
v H
2
O. V d s oxy ho
mt phn t gam ng gii phng ra 686 kcal. Nu nng lng ny c
gii phng ra cng mt lc th s gy ting n v h thng sng khng th
s dng ton b nng lng trong mt khong thi gian ngn nh vy.














12
































Hnh 1.4 Tin trnh gii phng nng lng ho hc trong s trao i cht
c chia lm 3 giai on

c tnh th ba l nng lng ho hc c gii phng ra khi phn
gii glucid, lipid v nhng hp cht cao phn t khc u c th c tch
lu trong nhng hp cht tch tr nng lng c th, c gi l hp cht
cao nng.
Tin trnh ca vic gii phng nng lng ho hc c bn c chia
lm 3 giai on (hnh 1.4):
Giai on th nht cc hp cht cao phn t (tinh bt, glycogen,
proteine, lipid...) b thu phn thnh cc cht c phn t b
Protein Polysaccharide Lipid
Aminoacid ng n
Glycerin v
acid bo
Glucose
Pyruvate
Acetyl CoA
Giai on 1:
B gy cc i phn t
thnh cc n phn t
Giai on 2:
Phn gii cc n phn t thnh
Acetyl CoA, to ra ATP v NADH
mc nht nh
Giai on 3:
Oxy ha hon ton Acetyl CoA
thnh H
2
O v CO
2
, gii phng
mt lng ln NADH v ATP
ty th
Mng t
bo sinh
vt nhn
chun
Mng
ty th

Chu trnh
citric acid
13
(monosaccharide, amino acid, axit bo, glycerine...). Nng lng gii
phng ra trong giai on ny khng ng k, ch bng gn 1% d tr nng
lng t do ca cc cht ny c gii phng ra di dng nhit.
Giai on th hai l qu trnh ng phn, oxy ho cc axit bo v
cc amino acid. Nng lng gii phng ra trong giai on ny gn bng
1/3 nng lng t do d tr trong cc cht . Sn phm chnh ca giai
on ny l acetyl-CoA, -xetoglutaric acid v oxaloaxetic acid.
Giai on th ba l oxy ho tip tc cc cht trn trong chu trnh
Krebs. Khong 2/3 nng lng c gii phng ra giai on ny.

1.2.2 Cc hp cht cao nng
Nhng hp cht cao nng: Tt c s sng trn tri t ph thuc vo
nng lng mt tri, trong nhng dng sng c mt h thng th bc v
nng lng. Mt s tip nhn nng lng mt tri trc tip, mt s khc
nhn nng lng t nhm trn trong nhng qu trnh tip theo.
Nhng sinh vt hp thu nng lng nh sng trc tip c gi l c
th t dng. Nhng c th ny d tr nng lng mt tri trong cc phn
t hu c khc nhau. Nhng sinh vt s dng nhng phn t , gii
phng nng lng d tr trong mt lot cc phn ng oxy ho kh c
gi l sinh vt ho dng.
Mc d khc nhau c hai loi u c c ch chung v ti sinh mt
dng nng lng ho hc, nng lng c th c gii phng trong nhng
phn ng to nhit thc hin cc qu trnh sng a dng (cn nng
lng).
Mt nhm nh cc phn t l cht trung gian chuyn nng lng t
cc phn ng gii phng nng lng n cc phn ng cn nng lng ca
c th. Nhng phn t ny l coenzyme dng kh, nhng hp cht
phosphate c gi l cao nng nu chng gii phng ra nng lng t do
c gi tr m ln khi thu phn ( G
0
c gi tr m ln hn -25 kJ/M).
bng 1.2 sau y l danh sch nhng hp cht cao nng quan
trng nht, nhng phn t nh phosphoric anhydric (ATP, ADP), enol
phosphate (PEP), acyl phosphate (acetyl phosphate), guanidinophosphate
(creatine phosphate). C nhng hp cht thioeste, nh acetyl CoA khng
cha phospho nhng gii phng mt nng lng t do ln khi thu phn.






14
Bng 1.2 Nng lng t do gii phng khi thy phn mt s cht cao nng




Cc cht G
o
(kJ/mol) Cng tc cu to
Phosphoenolpyruvat (Pyruvate + P
i
) -62,2
3,5-Cyclic adenosin monophosphate -50,5
(5-AMP)
1,3-Bisphosphoglycerate -49,6
(3-phosphoglycerate + P
i
)
Creatine phosphate -43,3
(creatine + P
i
)
Acetyl phosphate (acetate + P
i
) -43,3

Adenosine-5 triphosphate -35,7
(ADP + P
i
)

Adenosine-5 triphosphate -30,5
(ADP + P
i
) (nng Mg
++
qu cao)

Adenosine-5 diphosphate -35,7
(AMP + P
i
)


P
15





















Tng s nng lng chnh xc gii phng ra khi thu phn ph thuc vo
nng , pH, nhit ... nhng gi tr G
0
khi thu phn nhng hp cht
ny c gi tr dng ln hn ng k so vi nhng cht trao i khc.
Chng c hai c im quan trng:
Nhng cht phosphate cao nng (high- energy phosphate
compounds) khng phi l cht d tr nng lng lu di, chng l nhng
cht chuyn tip nng lng d tr, l cht mang nng lng t im ny
sang im khc, t mt h thng ny n mt h thng khc.
Nng lng hot ho c cung cp ng k t ATP khi thu phn
nhm -phosphat.
Nng lng lm gy lin kt O-P thng l 200-400 kJ/M, ln
hn ng k so vi 30,5 kJ/M khi thu phn ATP.
Cc nh ho sinh hc quan tm nhiu n nng lng gii phng
thc t.

Vai tr trung tm ca ATP trong nng lng sinh hc
ATP cha hai pyrophosphoryl (hnh 1.4). Nhng phn t c lin kt
anhydric, ADP, GTP, GDP v cc nucleoside triphosphate khc,
nucleotide-ng nh UDP-glucose v pyrophosphate v c th hin nng
Cc cht phosphate G
o
(kJ/mol) Cng tc cu to
nng lng thp hn
Glucose-1-P (glucose+ P
i
) -21,0
Fructose-1-P (fructose+ P
i
) -16,0
Glucose-6-P (glucose+ P
i
) -13,9
Glycerol-3-P (glycerol+ P
i
) -9,2
Adenosine-5 monophosphate -9,2
(adenosin + P
i
)

16
lng t do G
0
ln khi thu phn. Nguyn nhn ho hc ca gi tr G
0

m ln l do s khng bn vng ca cht phn ng do s cng lin kt gy
ra bi s y tnh in. S bn vng ca sn phm phn ng do s ion
ho, s cng hng v nhng yu t entropy gy ra do thu phn v s ion
ho tip theo.
Mc d PEP, cyclic AMP, 13 DPG, phosphocreatine,
acetylphosphate v pyrophosphate u c gi tr G
0
ln hn, nhng ATP
l duy nht nh v gia cc cht phosphate cao nng, (ATP c tng hp
khi phn gii cc cht hu c) v cc cht nhn nng lng (khi cc cht
ny c phosphoryl ho tham gia cc phn ng tip theo trong trao
i cht). Ni mt cch khc ATP l mt xch ni lin hai qu trnh ngc
nhau, l ng ho v d ho.
Vic hnh thnh tt c cc hp cht cao nng khc cng xy ra do s
tiu ph nng lng vn tch lu trong ATP.
ADP c th nhn c phosphate v nng lng t cc phosphate cao
nng.
ATP cho c gc phosphate v nng lng i vi cc phn t c
nng lng thp. Nh vy ATP c vai tr d tr nng lng cng nh tiu
hao nng lng.
Xt v c ch bin i v chuyn ho nng lng trong s phn gii ATP
v cc hp cht cao nng tng t ATP ta thy nng lng cn thit
thc hin phn ng ho hc c gii phng ra mt im, c th c
chuyn n mt im khc, y nng lng c s dng mt cch trc
tip. iu ny c ngha l trong c th sng khng nht thit phi tip xc
vi nhau bng cch va chm (c trng cho ngoi c th sng) gia cc
phn t cho v nhn nng lng.














17































Hnh 1.5 Chui ba gc phosphat ca ATP cha 2 lin kt pyrophosphat, c
hai lin kt ny gii phng nhiu nng lng khi thu phn

Cu hi 1. Nng lng sinh hc?
2. Cc ng hng tng hp ATP trong thc vt ?
3. Cc dng hp cht cao nng, cho v d?

Lin kt
Phosphoric anhydride
ATP
(adenosin-5-triphosphate)
Adenosine
AMP
ADP
ATP
1
Chng 2
Vitamin
Vitamin l nhng cht hu c c trng lng phn t b, c cu to
ho hc rt khc nhau v c hot tnh sinh hc nhm m bo cho cc qu
trnh ho sinh v sinh l trong c th tin hnh c bnh thng, v do
, c nh hng rt ln i vi s trao i cht.
Vitamin khng c tng hp ng vt bc cao, v vy chng
phi c tip nhn cng vi thc n. Nhiu vitamin l tin cht ca
cofactor (vitamin nhm B) tham gia vo cc phn ng enzyme, trong khi
nhng vitamin khc tham gia vo qu trnh nhn v iu khin s sao
chp (vitamin A), cc phn ng kh (vitamin C v E), to xng (vitamin
D), ng mu (vitamin K) v.v
Cc vitamin c cu to ho hc rt khc nhau. Thng chng c
phn loi da vo ho tan:
Nhm vitamin ho tan trong nc: B
1
, B
2
, B
6
, B
12
, C, folate,
pantothenate, biotin. Chng c tch lu ch vi lng t. Lng d tha
c thi ra qua nc tiu.
Nhm vitamin ho tan trong cht bo: A, D, E, K. Lng d tha
c tch ly v dn n hin tng tha vitamin (c bit vitamin A v
E).

2.1 Cc vitamin ho tan trong nc
2.1.1 Thiamin (vitamin B
1
)
Thiamin c pyrophosphoryl ho thnh coenzyme
thiaminpyrophosphate (TPP), tham gia xc tc phn ng kh carboxyl ho
bng cch oxy ho v phn ng chuyn nhm aldehyd hot ho. Phn ng
ny ng mt vai tr quan trng trong trao i cht.
TPP l nhm prostetic ca pyruvat-dehydrogenase, pyruvat-
decarboxylase, 2-oxoglutarat-dehydrogenase v transcetolase v nh vy
n tham gia vo qu trnh ng phn, chu trnh citrate, pentose-
phosphate v Calvin.
a. Decarboxylase: xc tc cho phn ng loi nhm carboxyl ca
pyruvic acid, -cetoglutaric acid.
b. Transcetolase: xc tc cho phn ng vn chuyn glycoaldehyd
(CH
2
OH-CO-). V d phn ng chuyn on 2C (C
1
v C
2
) ca
xylulose 5-phosphate n ribose 5-phosphate to thnh
sedoheptulose 7-phosphate v glyceraldehyd-3-phosphate.
2

Sinh tng hp
Hai thnh phn ca thiamine l pyrimidine v thiazol c tng hp
ring v sau c kt hp li vi nhau. Cc ng hng tng hp l
khc nhau sinh vt nhn s v sinh vt nhn chun.
Trong E.coli v S.thyphimurium 5-phosphoribosyl-5-aminoimidazol
(AIR) l tin cht ca pyrimidine. E.coli pyruvat v D-glyceraldehyd l
cc tin cht ca thiazol. Chng lng kt thnh 1-desoxy-D-xylulose, cht
ny c gn cc nguyn t C 4, 4, 5, 6, 7. Ngun gc ca C
2
v N
3
l
tyrosine, nguyn t S bt ngun t cysteine. Pyrophosphatester ca thnh
phn pyrimidine tham gia vo phn ng gn. Bng mt phn ng
pyrophosphoryl ho vitamin c chuyn thnh coenzyme.









B
1
c nhiu trong cm go, gan, thn, Mm ng cc v nm men
l ngun rt giu vitamin ny.
C th ngi hng ngy cn 1-1,5 mg vitamin B
1
. Thiu vitamin
ny nh hng n qu trnh trao i carbohydrate dn n bnh ph
thng, hay cn gi l bnh beri-beri, ri lon thn kinh v nh hng n
chc nng ca tim. B
1
ch bn vi nhit trong mi trng acid, cn trong
mi trng kim n b phn hu nhanh chng khi un nng. Khi oxy ho
B
1
chuyn thnh mt hp cht gi l thiocrome pht hunh quang. Tnh
cht ny c s dng nh lng vitamin B
1
.
Hm lng B
1
trong nguyn liu c th thay i tu thuc iu kin
bo qun v ch bin. V d: go xt k hm lng B
1
b gim 4 ln so vi
ban u. m khi bo qun nguyn liu (thc, go) cng cao, hm lng
vitamin B
1
b gim cng mnh.

2.1.2 Riboflavin (vitamin B
2
)
NH
2
N
H
3
C
CH
2
*
N
S
CH
2
CH
3
H
CH
3
Thiamin (vitamin B
1
)
3
Trc kia B
2
c gi l lactoflavin v ln u tin c tch ra t
sa. Bn cnh nicotinamid-nucleotide NAD
+
v NADP
+
flavin-nucleotide
FAD v FMN l nhng nhm vn chuyn hydro quan trng. Chng tham
gia hn 100 phn ng oxy ho kh. V d, qu trnh kh carboxyl ho
bng cch oxy ho pyruvic acid, oxy ho acid bo, kh amin ho bng
cch oxy ho amino acid... Coenzyme ny c to nn t riboflavin bng
phosphoryl ho (FMN) v tip theo adenyl ho (FAD).





























Riboflavin (vitamin B
2
)
N H
3
C
N
NH
N
O
O
H
3
C

CH
2

HC-OH

HC-OH

HC-OH

CH
2
OH
Flavinmononucleotide (FMN)
N H
3
C
N
NH
N
O
O
H
3
C

CH
2

HC-OH

HC-OH

HC-OH

CH
2
OP
4

FAD

Sinh tng hp v bin i
Riboflavin c to nn t GTP v ribulose-5-P trong thc vt, nm
men v nhiu vi sinh vt.
Phn ng tng hp bt u bng vic m vng imidazol ca GTP v
tch ra mt gc pyrophosphate. Bng phn ng kh amin ho, phn ng
kh v tch ra gc phosphate cn li lm xut hin 5-amino-6-
ribitylamino-2,4-pyrimidindion. Phn ng ca cht ny vi 3,4-dihydroxy-
2-butanon-4-P (cht ny xut hin t ribulose-5-P) to nn phn t c hai
vng 6,7 dimethyl-8-ribityllumazin. Hp cht ny kt hp vi 5-amino-6-
ribitylamino-2,4-pyrimidindion thnh riboflavin. Phosphoryl ho
riboflavin to nn flavinmononucleotide (FMN). Bng phn ng adenyl
ho tip theo lm xut hin flavin-adenin-dinucleotide (FAD). S
phosphoryl ho riboflavin ng vt c iu khin chnh xc bng
hormone tuyn gip trng.



5

























Thiu vitamin B
2
nh hng n qu trnh oxy ho kh lm nh
hng n qu trnh to nng lng cn thit cho s sinh trng v pht
trin ca c th. Thiu vitamin gy ra nhng ri lon v sinh trng v cc
bnh v da. Tuy nhin nhng bnh ny tng i him.
Nhu cu vitamin B
2
hng ngy ca ngi ln khong 1,4 - 2 mg.
Sn phm phn gii v sn phm thi ra ngi l riboflavin. Cc bc
phn gii khc (c bit vi khun) l hydroxyl ho cc gc methyl v
lm ngn chui ribityl.
Chc nng ho sinh
Guanosin
triphosphate
GTP
Hydrolase II
2-diamino-6-
ribosylamino-3 (3H)-
pyrimidine 3-P
3H
2
O
HCOOH
PP
i
5-diamino-6-ribosylamino-2,4
(1H,3H)-pyrimidinedione 5-P
Pirimidine
Desaminase
H
2
O
NH
3

NADPH
NADPH + H
+
5-diamino-6-ribotylamino-2,4
(1H,3H)-pyrimidinedione 5-P
NADPH + H
+
NADPH
NH
3

H
2
O
Uracil
reductase
5-diamino-6-ribotylamino-2,4
(1H,3H)-pyrimidinedione
H
2
O
P
i
Phosphatase
Ribulose-5-
phosphate
3,4-Dihydroxy-2-
butanone-4-P
6,7-Dimethyl-8-
ribityllumazine
Riboflavin (vitamin B
2
)

P
i
HCOOH
Formate
Mg
++
6,7-Dimethyl-8-
ribityllumazine
synthase
Riboflavin
synthase
6
Chc nng oxy ho kh ca FMN v FAD l do vng isoalloxazin
chu trch nhim. mt phn ng kh hon ton hai in t v hai proton
c tip nhn.
Dng oxy ho ca riboflavin l nhng tinh th vng da cam, hp th
bc sng 370 v 450 nm. Dng kh ca n khng mu v mt tnh hp
th 450 nm. Cc coenzyme flavin nucleotide dng kh cng c th b
oxy ho tr li khi c cc cht nhn in t nh xanh methylen, 2,6-
diclorophenolindophenol
FADH
2
+ xanh methylen FAD + xanh methylen
(dng oxy ho mu xanh) (dng kh khng mu)
C th s dng cc tnh cht trn theo di cc phn ng do flavin
dehydrogenase xc tc.
Tinh th vitamin B
2
dng kh tng i bn vi nhit hn vitamin
B
1
, tuy nhin vitamin ny khng bn di tc dng ca nh sng. Ngc
vi vitamin B
1
, hm lng vitamin B
2
trong go, tht, trng, sa bin i
khng nhiu trong qu trnh bo qun v ch bin.

2.1.3 Pyridoxin (vitamin B
6
)

Pyridoxin (pyridoxol), pyridoxal v pyridoxamine to nn nhm
vitamin B
6
. C 3 dng ny d dng chuyn ho ln nhau. Dn xut ca
vitamin B
6
l pyridoxalphosphate (PLP) l coenzyme ca nhiu enzyme.
Pyridoxal- v pyridoxamin-phosphate l coenzyme quan trng cng tc
ng mt s ln cc phn ng bin i amino acid, v d phn ng
transaminase, decarboxylase v dehydratase. Vitamin B
6
cng l thnh
phn cu to ca phosphorylase xc tc cho phn ng phn gii glycogen.
Ngoi ra vitamin B
6
cn c vai tr quan trng trong qu trnh sinh tng
hp NAD
+
, coenzyme A... Do thiu vitamin ny nh hng n nhiu
qu trnh trao i proteine, saccharide, lipid, dn n ri lon v thn kinh
v trao i amino acid.
Sinh tng hp v bin i
Mt s vi khun, nm men v thc vt tng hp c pyridoxine.
ng vt nhai li cng khng cn c vitamin B
6
trong thc n v vi sinh
vt trong rut ca chng c th tng hp vitamin ny cung cp cho
ng vt ch. Cc ng vt khc cn phi c cung cp vitamin ny t
bn ngoi.
Sn phm ngng t ca pyruvate v D-glyceraldehyd l 1-desoxy-D-
xylulose cung cp nguyn t C 2, 2, 3, 4 v 4. Erythrose 4-P c bin
7
i thnh 4-hydroxy-L-threonine. Hp cht ny cung cp nguyn t C 5,
5, 6 v N-1. Pyridoxine iu khin qu trnh tng hp bng c ch ngc.
Loi hydro ca pyridoxine thnh pyridoxal cho n nay ch c quan st
thy trong vi khun.







ng vt pyridoxine, pyridoxal v pyridoxamine sau khi hp th
c phosphoryl ho nh enzym kinase. Kinase ny b c ch bng sn
phm ca n. Bng phn ng oxy ho pyridoxinephosphate v
pyridoxaminephosphate bin i thnh pyridoxalphosphate. Phn ng ny
cng b c ch bng sn phm ca n.
Lng b sung hng ngy cn thit ngi ln khong 2 mg. S
phn gii bt u bng phn ng do phosphatase xc tc. S phn gii tip
theo ca vitamin ny thc hin bng mt phn ng oxy ho to thnh
carbonic acid-4-pyridoxat, cht khng c hot tnh sinh hc.

2.1.4 Cobalamin (vitamin B
12
)

Cobalamin l mt trong nhng phn t sinh hc khng trng hp
ln nht. Nguyn t coban trung tm gn lin hp vi 6 thnh phn: Bn
thnh phn l pyrrol ca vng corrin, mt thnh phn l nucleotide bt
thng dimethylbenzimidazol v thnh phn th su c th l gc
hydroxyl (vitamin B
12
), gc methyl (methylcobalamin), hoc gc 5-
desoxyadenosyl (desoxyadenosylcobalamin, coenzyme B
12
). Dng vitamin
B
12
trn th trng: thnh phn th 6 l mt gc cyanid.
Sinh tng hp coenzyme v phn ng kh vitamin
Coenzyme B
12
c tng hp nhiu bc trong vi sinh vt. Bc
th nht ging nhau gia hem v chlorophyll. Nhng phn ng tip theo
c nhiu bin i.


H
2
C-OH
HO
H
3
C
CH
2
-O-P
H
H
N

H-C=O
HO
H
3
C
CH
2
-O-P
H
H
N

H
2
C-NH
2
HO
H
3
C
CH
2
-O-P
H
H
N

Pyridoxamine-P
Oxydase
Pyridoxamine-P
Oxydase
O
2
H
2
O O
2
NH
3
H
2
O
2
8


















Cobalamin

Sau nhiu bc methyl ho vng porphyrin bin thnh vng corrin.
Sau nhiu ln methyl ho tip theo v amid ho Co
++
c a vo v
c kh thnh Co
+
, ri kt hp vi adenosine. Tip theo -ribazol c
gn vo qua mt cu propionate-aminpropanol v to nn thnh phn th
6 ca Co. Co nm trong phc h lin hp l Co
+++
. ng vt c v nhu
cu B
12
( ngi 10 g/ngy) c p ng do vi khun rut.
hp thu vitamin B
12
t rut cn protein c hiu (50 kDa). Nu
thiu loi protein ny s dn n hin tng thiu vitamin B
12
trm trng
(thiu mu c tnh). vn chuyn trong mu B
12
lin kt vi nhiu
plasmglobulin khc nhau, c gi l transcobalamin. Phn ln B
12
c
tip nhn dng oxy ho (hydroxy-cobalamin, Co
++
). Sau hai bc phn
ng xut hin tr li dng desoxyadenosyl (coenzym B
12
).
Vitamin B
12
tham gia trong thnh phn cu to ca cc enzyme xc
tc cho cc phn ng:
- Phn ng chuyn ribonucleotide thnh desoxyribonucleotide
H
3
C
CH
3

CH
3

CH
3

CH
3

H
3
C
H
3
C
CH
3

H
3
C
NH
NH
2

CH
3

H
3
C
H
2
N
H
2
N
H
2
N
H
2
N
O
NH
2

O
O
O
O
O
O
O
-
O
O
P
N
N N
N
N
N
N
H
H
H
H
O
O HO
HO
Co
9
- Phn ng chuyn nhm methyl
- Sp xp li cc nhm th ca 2 carbon lin nhau trong phn t.

2.1.5 Vitamin H (Biotin)
Biotin l mt vitamin cn thit cho cc phn ng carboxyl ho, c
gn kt ng ho tr vi enzyme bng nhm -amin ca gc lysine.
Sinh tng hp v bin i
Qu trnh tng hp xy ra vi khun (c vi khun rut ca ng
vt), nm men v thc vt bc cao.
Phn t ban u l pimeloyl-CoA. Ngun gc ca phn t ny cha
c gii thch y , c th do s ngng t ca 3 phn t malonyl-CoA
v gii phng ra 2 phn t CO
2
. Tip theo n ngng t vi L-alanine tch
ra pyridoxal-pp, CO
2
v HS-CoA. 8-amino-7-oxopelargonat (7-KAP) xut
hin c bin i thnh 7,8-diaminopelargonate bng mt phn ng do
transaminase xc tc. phn ng ny, S-adenosylmethionine l cht cho
nhm amin (thay v chc nng thng xuyn l cht cho nhm methyl).
Bng s gn CO
2
vo (ph thuc vo ATP) vng imidazol c ng li.
Sau xy ra s gn S (t L-cysteine) v ng vng thiophan. Kt qu l
to nn (+)-Biotin.
S gn kt biotin vo enzyme tng ng cn hot ho bi ATP,
c xc tc bng enzyme c hiu biotin-ligase.
Ngi ln cn 100-200 g/ngy. Biotin c trong nhiu thc phm
v c tng hp bi vi khun ng rut, v vy t khi b bnh thiu
vitamin ny tr khi n nhiu trng sng c nhiu avidin kt hp rt cht
vi biotin, ngn cch vic hp thu n qua rut non.







Biotin


Phn gii bng vi khun
O
HN NH
S
OH
O
10
Pseudomonas c th lm ngn chui bng -oxy ho v nguyn t C
ca vng thiophan c tch ra. Gc ca vng imidazol c thi ra
nc tiu. ng vt c v ch c th tch chui bn v to nn hp cht
sulfoxid. Tuy nhin thnh phn chnh ca biotin c thi ra dng khng
bin i.

2.1.6 Ascorbate (vitamin C)
Vitamin C tham gia vo nhiu phn ng bin i cht v thng c
chc nng bo v.
Sinh tng hp v trao i cht







Vitamin C

Thc vt bc cao, to, nm men v phn ln ng vt c th tng
hp ascorbate. Mt s ng vt c v (ngi) cng nh cn trng, phn
ln cc loi c khng c kh nng ny. nhng ng vt c v cn cung
cp vitamin C, v thiu enzyme L-Gu-ionolacton-oxidase.
Sinh tng hp ng vt v phn ln thc vt t D-glucose v i
qua UDP-glucose, UDP-glucuronate, D-glucuronate, L-gulonate, L-
gulonolacton v 2-dehydro-L-gulonolacton. Vitamin C l sn phm oxy
ho ca cc ng. N c tng hp t glucose ( ng vt v thc vt),
t galactose ( mt s thc vt). Phn t vitamin C c cha en-iol, d
dng b loi hydro to thnh dehydroascorbic acid (cng c hot tnh
vitamin C). Vitamin C mt hot tnh khi vng lacton b thu phn. Trong
t nhin, vitamin C c th tn ti dng lin kt vi cc cht khc, gi l
ascorbigen l dng d tr vitamin C quan trng. Vitamin C ho tan tt
trong nc, dung dch c v acid, c tnh hot quang v c tnh kh mnh.
Ascorbic acid c kh nng kh dung dch Fehling, AgNO
3
. C th s dng
tnh cht ny nh lng ascorbic acid. Ngoi ra, cng c th s dng
vitamin ny lm cht chng oxy ho trong cng nghip thc phm.
Ascorbate i vo cng thc n c vn chuyn n cc c quan
khc nhau. mt s c quan s tip nhn l mt qu trnh ch ng, c
kch thch bi insulin. Nng ascorbate cao nht tm thy nang trn
O
O
OH
OH
HO
HO
11
thn, no thu, mt, gan, d dy v no. Bnh hoi huyt ngi c
ngn nga khi tip nhn 10mg ascorbate/ngy. Ngi ht thuc v ngi
c thai cn nhiu ascorbate hn. Ngi ta khng nh rng, mt liu lng
cao ascorbate tng cng h thng min dch, gim s mt mi v nguy c
bnh ung th. Tuy nhin s phn gii ascorbate xy ra mnh m, v vy
khi lng cn hng ngy l 50 mg th cn a vo c th l 1 500 mg.
Vitamin C c nhiu trong rau qu ti, c bit l cc loi: cam,
chanh, da chut... Vitamin C tham gia trong nhiu qu trnh quan trng
ca c th sng nh:
- Qu trnh hydroxyl ho do oxygenase. V d hydroxyl ho proline,
lysine trong phn t collagen. Thiu vitamin C, collagen mi c
to thnh khng c hydroxyl ho, khng to c dng xon ba
nn c bn km, do da b tn thng v thnh mch mu
mng manh d b v, gy ra bnh hoi huyt. Vitamin C tham gia
trong qu trnh chuyn ho protocollagen thnh collagen nn c tc
dng lm cho vt thng chng thnh so
- Vn chuyn hydro trong qu trnh oxy ho - kh thc vt
- Lm tng tnh khng ca c th i vi nhng iu kin khng
thun li ca mi trng ngoi, cc c t nhim trng. Ngoi ra
vitamin C cn tham gia trong nhiu qu trnh tng hp, chuyn ho
cc cht khc nhau.
- Duy tr cn bng gia cc ion Fe
2+
/Fe
3+
, Cu
+
/Cu
2+

Phn gii v s loi thi
loi linh trng (primat) mt phn ascorbate tn ti trong c th
c thi ra ngoi dng khng bin i qua nc tiu. Nhng sn phm
loi thi khc l oxalate v dioxogulonate (xut hin t dehydroascorbate
bng mt phn ng khng thun nghch v khng xc tc). Ascorbate c
th c oxy ho hon ton thnh CO
2
.
2.2 Cc vitamin ho tan trong cht bo
2.2.1 Retinol (vitamin A)
Vitamin A l tn thng gi cho ru retinol, aldehyd retinal v
retinic acid, chng l nhng isoprenoide c c tnh lipid. Chng tn ti
cc dng ng phn cis v trans khc nhau.
Sinh tng hp v bin i
Tin cht tng hp vitamin l provitamin -carotene, s tng hp
chng xy ra trong thc vt. ng vt c th tch carotene thnh retinal,
bng mt phn ng dioxygenase (xy ra trong rut) v s dng n nh
l vitamin.
12
Vitamin A c hp th vo cng vi cht bo v c tch lu
dng retinylpalmitat trong gan sau khi kh v ester ho. vn chuyn
n nhng c quan khc nhau ester c thu phn v retinol c a
vo mu, n c tip nhn bng mt protein kt hp c hiu vi
retinol. S bin i gia retinol, retinal v retinic acid c thc hin bng
nhng phn ng ph thuc NADP
+
.
Carotene c a vo cng vi thc n ca ngi c ngun gc
trc ht l t thc vt, trong khi retinol c ngun gc ng vt (du gan
c thu). Nhu cu hng ngy i vi ngi ln l khong 1 mg retinol hoc
6 mg -carotene. Thiu vitamin A rt ph bin nhm dn s suy dinh
dng, dn n chng qung g, c ch s pht trin ca xng v s
phn ho m biu b, hn ch s pht trin v chc nng sinh sn.

Chc nng ho sinh
Retinic acid l mt cht iu ho sao chp. Sau khi i qua mng t bo
phn t ny kt hp vi cht nhn c hiu: all-trans-retinic acid vi
RAR-receptor, 9-cis retinic acid vi RXR-receptor. Sau phc h
retinic acid -RAR-RXR kt hp vi hormone c to ra t DNA trong
nhn.
Thuc nhng gen c iu khin theo cch ny l gen m ho cho
protein kt hp vi retinol, enzyme PEP-carboxylase v apolipoprotein
A
1
. Retinic acid cng tham gia vo s iu khin s pht sinh phi v pht
sinh hnh thi ( liu lng nh vitamin A trong thi gian c thai l
nguyn nhn gy qui thai), pht trin, phn ho v kh nng sinh sn.
S bin i t 11-cis thnh all-cis-retinal nh nh sng l c s cho
qu trnh nhn ng vt. Trong vitamin nhm A c 2 dng quan trng l
vitamin A
1
v A
2
. Vitamin A
2
khc vi A
1
ch trong vng c thm mt
ni i gia C
3
v C
4
. Di tc dng ca enzyme retinoldehydrogenase
nhm alcol ca vitamin A d dng b oxy ho n aldehyd (all-cis-retinal,
vn c hot tnh vitamin A). Lin kt i gia C
11
v C
12
ca retinal c th
chuyn thnh dng cis (11-cis-retinal), 11-cis-retinal kt hp vi protein
opxin to nn sc t ca mt l rodopxin. y l protein nhn nh sng
(photoreceptor protein) c trong t bo hnh que ca mng li mt ngi
v ng vt c v. T bo ny hot ng trong nh sng yu, thch nghi
vi bng ti. Khi c nh sng, nhm thm ca rodopxin chuyn t dng
cis sang dng trans nn mt kh nng kt hp vi opxin. Ngc li trong
ti s ti to li dng 11-cis-retinal v rodopxin c tng hp tr li, lm
tng nhy cm ca mt trong nh sng yu.
13
Halobacterium s dng s bin i t all-trans thnh 13-cis-retinal
nh nh sng lm ng lc cho nhng bm proton ca chng.
Retinylphosphate tham gia vo qu trnh sinh tng hp cc
glycosaminoglycane trong cc t bo biu b, y n c chc nng
tng t nh dolicholphosphate. Bng cch ny n tham gia gin tip vo
s ton ven ca mng t bo v ty th, n nh biu b ca da v ca mng
nhy v gim nhng ri lon v pht trin.


















2.2.2 Calciferol (vitamin D)
ng vt bc cao nhng sn phm hydroxyl ca vitamin D ng
mt vai tr trung tm trong trao i calcium.






CHO
CH
2
O
H
-carotene
-carotene-15,
15-Dioxygenase


Retinol
dehydrogenase

NADH + H
+

NAD
+

All-trans-Retinal
All-trans-Retinol
(Vitamin A)
Cholecalciferol
(Vitamin D
3
)

CH
3
CH
3
CH
3
CH
2
HO

CH
3
14






Thc t khi nim vitamin D l gm mt nhm cc cht h hng,
khc nhau v ngun gc. Nu ni cht ch th chng khng phi l
vitamin, v da c th tng hp mt trong chng (D
3
) t mt steroid (7-
Dehydrocholesterin, provitamin D) khi c nh sng t ngoi.
Sinh tng hp v s bin i
Cc vitamin D khc nhau c to nn t
5,7
-steroid (sn phm
trung gian tng hp zoo-, phyto- v mycosteroid) bng vic m vng B
nh nh sng. Chng ch khc nhau cc mch bn. Hai vitamin D u
tin sau y l quan trng nht:
- Vitamin D
2
(ergocalciferol): to nn t ergosterin, cht ny tn ti
trong thc vt v nm.
- Vitamin D
3
(cholecalciferol): to nn t 7-dehydrocholesterin, cht
ny c trong ng vt bc cao.
- Vitamin D
4
: xut hin t 22-dihydroergosterin, c trong thc vt
v nm
- Vitamin D
5
: to nn t 7-dehydrositosterin, c trong thc vt.
Tng hp ni sinh
Trong phn ng u tin khi chiu nh sng t ngoi dn n s
ng phn ho ng thi vi vic m vng B ca cu trc sterin, lm xut
hin provitamin D. di bc sng thch hp nht cho quang phn ng
l 295 nm. Chiu sng vi bc sng ln hn s to lumisterin, vi bc
sng ngn hn to tachysterin. nh sng mnh hn xut hin suprasterin
I v II. Vitamin D c to nn do s ng phn ho ph thuc vo nhit
. Vitamin D i vo mu v gn vi protein c hiu (DBP: vitamin D-
bindende-protein).


a vo cng vi thc n:
Mc d s chiu sng c th cung cp y lng vitamin D, vic
b sung thm vn l iu cn thit ( ngi ln 5 g/ngy, tr em 10
15
g/ngy). Sau khi hp th (c h tr bi acid mt) vitamin D c gn
vi chylomikronen hoc DBP v c vn chuyn n gan.
trong gan xy ra s hydroxyl ho C25 bng mt enzyme ph
thuc vo cytochrome P-450. Cht ny c chuyn n thn, y xut
hin hp cht c hot tnh sinh hc 1 , 25-Dihydroxycalciferol (calcitriol,
DHCC) do s hydroxyl ho tip theo v tr C
1
ca cholecalciferol.












S phn gii thc hin bng s hydroxyl ho v/hoc oxy ho nhm
hydroxyl thnh nhm carboxyl. Nhng hp cht ny phn cc mnh hn,
th hin hot tnh sinh hc thp, c thi ra qua phn hoc nc tiu.
Chc nng sinh ho
ng vt c v vitamin D (thc t l 1 25-Dihydroxycalciferol)
iu khin s trao i calcium v phosphate.
N hot ho trc tip s hp thu (ti hp th) Ca
++
v phosphate
rut, thn. Mt khc n gii phng Ca
++
t xng cho s iu chnh tm
thi mc Ca
++
trong mu.
Hiu qu sao chp c thc hin bi vitamin D-receptor. y
vitamin c vai tr nh mt hormon. trong rut, d dy v xng n
kch thch s sao chp ca cc gen m ho cho cc protein gn vi calcium
(CaBP, calbindin, osteocalcin...). Nhng protein ny tham gia vo s tip
nhn calcium v khong ho xng.
Lnh vc y hc: Nhng bnh gn lin vi mc vitamin D khng bnh
thng:
Cholecalciferol
(Vitamin D
3
)

CH
3
CH
3
CH
3
CH
2
HO

CH
3
Ca
++
Phosphate
Hydroxyl ha
C25 (gan)
Hydroxyl ha
C1 (thn)
1 , 25-Dihydroxycalciferol
(Calcitriol, DHCC)
HO
CH
3
CH
3
CH
2
H
3
C

CH
3
HO
OH
H
H
16
Ci xng: (khong ho xng thp, phn ln tr em) sinh ra do
s hp th Ca
++
gim khi thiu vitamin D, thnh thong cng do
tng hp khng calcitriol t calciferol.
Bnh long xng: xut hin khi tng hp calcitriol b ri lon, do
thn khng hot ng hoc mc ostrogen thp v n kch thch
bc hydroxyl ho.

2.2.3 Tocopherol (vitamin E)
Tocopherol (tocos = con chu, pherol = sinh ra) kh ph bin cy
xanh, rau x lch, ht ng cc, du thc vt, gan b, lng trng.
Nhm tocopherol gm 8 hp cht, bao gm vng 8 - chromanol v
mt chui bn isoprenoide Sn phm cui cng ca qu trnh sinh tng
hp l -tocopherol (5,7,8-trimethyltocopherol), th hin hot tnh sinh
hc cao nht. thc vt v ng vt tocopherol bo v lipid trc s thit
hi do oxy ho.
Cc tocopherol phn bit vi nhau bng s lng v v tr ca nhm
methyl vng v cc lin kt i trong chui bn. Tt c chng u a
lipid.
Tocopherol c tng hp dng ng nht ch thc vt, trc
ht trong lc lp ca l. ng vt v thc vt n mng t bo (mng ty
th, hng huyt cu, v lp th).
Lng b sung hng ngy i vi ngi ln l 8-12 mg. Cn lng
ln hn khi thc phm c cha nhiu acid bo cha no. Him khi thy
thiu vitamin ny, ngoi tr trng hp non v ri lon hp thu
lipid. Tuy nhin trong chn nui ch dinh dng cung cp khng y
th gy ra s thoi ho cc c quan v v sinh.
Chc nng ho sinh ca tocopherol l bo v lipid mng trc
phn ng peroxid-ho (c bit acid bo c nhiu lin kt cha no).
Tocopherol l ni ether ca cc hydroquinon. Chng l nhng cht nhn
gc v chuyn sang trng thi tocopheroxy (semiquinon). y l mt gc
phn ng yu vi thi gian bn hu l nhiu gi v lm gy cc phn ng
chui. Phn ng ny l thun nghch, c th ascorbate thc hin s kh tr
li thnh tocopherol. y n b oxy ho thnh monodehydroascorbate.
Gia hai vitamin ny c s ng tc dng. -carotene cng c coi l
cht kh cho tocopherol. Ngc li semiquinon c oxy ho tip tc
thnh quinon, cht ny dn n s m vng thun nghch.


CH
3
CH
3
H
3
C

HO
O
CH
3
CH
3
CH
3
CH
3
-Tocopherol

17






2.2.4 Phylloquinon v menoquinon (vitamin K)
Phylloquinon l mt thnh phn ca h thng quang ho ca thc
vt quang hp. Menoquinon ng mt vai tr trong h hp ym kh ca vi
khun. ng vt hai hp cht (vitamin K
1
cng nh vitamin K
2
) l
cofactor s cacboxyl ho ca glutamat. l monomethyl-
naphtochinon vi mt chui bn isoprenoid.
Phyllochinon v menachinon phn bit nhau ch s lng lin kt
i v chiu di ca chui bn.
Lng tip nhn hng ngy cn thit ngi ln 70-140 g. Lng
ny c p ng bng thc n v sn phm ca vi khun ng rut.
Trc khi c thi phn ln ch cc chui bn c lm ngn bng oxy
ho.
Chc nng ho sinh
ng vt chng l cofactor to nn -cacboxylglutamate trong
protein xy ra mng li ni cht (endoplamatische reticulum) th nh
enzyme carboxylase. Protein to nn c phn gii. Qu trnh bin i
sau l cn thit cho kh nng kt hp Ca
++
v cho s c nh protein
bng cc cu Ca
++
mng phospholipid (v d qu trnh ng mu).
qu trnh bin i protein ny vitamin-K-hydroquinon tch ra mt
proton ca glutamate, tip theo n c carboxyl ho. ng thi xut hin
mt vitamin K-quinon-2,3-epoxid (nhng chi tit vn cn cha bit).
Epoxid c kh bng quinon-epoxid-reductase gn vi dithiol thnh
quinon v sau thnh hydroquinon. Dn xut ca cumarine c th km
hm reductase, v d dicumarol v warfarin c tc dng l cht khng
ng.





CH
3
CH
3


H
2
C CH = C CH
2
CH
2
CH
2
C CH
2
H

H 3

O O
CH
3
Phyloquinon
18














Cu hi
1. Chc nng ha sinh ca cc vitamin?
2. Tng hp cc vitamin ha tan trong nc v ha tan trong cht bo?

CH
3


H
2
C CH = C CH
2
H

n

O
O
CH
3
Menoquinon
37
CHNG III
ENZYME V S XC TC SINH HC

sng v pht trin, c th sinh vt phi thng xuyn trao i
cht v trao i nng lng vi mi trng bn ngoi. Cc qu trnh ny
tin hnh c l nh cc phn ng ha sinh xy ra trong c th theo quy
lut c t chc v c lin quan cht ch vi nhau.
c im chung ca cc phn ng ny l c th xy ra mt cch
c hiu, d dng vi vn tc ln ngay trong iu kin mi trng sinh
l v nhit ca c th. S d cc phn ng tin hnh trong c th c
nhng c im trn chnh l nh cc cht xc tc c bit c kh nng
lm tng vn tc phn ng ln gp bi, l cc cht xc tc sinh hc.
Enzyme l cht xc tc sinh hc c bn cht protein lm nhim v
xc tc cho cc phn ng ha sinh xy ra trong c th sinh vt.
Enzyme tn ti trong tt c cc t bo sng ca ng vt, thc vt,
vi sinh vt. Cc phn ng do enzyme xc tc c th xy ra ngay trong c
th sng hoc ngoi c th. Hin nay tch c rt nhiu enzyme
khc nhau t ngun ng vt, thc vt, c bit l t vi sinh vt, trong
c nhiu loi c thu nhn vi thun khit cao. Xc tc enzyme c
nhiu u im hn hn xc tc thng thng nh cng lc xc tc ln,
tnh c hiu cao, khng c, c th tc dng trong iu kin nh nhng,
n gin. Hn na c th sn xut enzyme t cc nguyn liu gi r nhng
em li hiu qu kinh t cao, nh vy vic ng dng enzyme rng ri
trong cc ngnh cng nghip nh, cng nghip thc phm, nng nghip, y
hc v nghin cu khoa hc nhanh chng c nhng bc tin di v
ngy cng c nhiu trin vng tt p.

I KHI NIM V S XC TC NI CHUNG
Tc phn ng ha hc c xc nh bi gi tr nng lng hot
ha tc l nng lng cc cht tham gia phn ng phi t c trn mc
nng lng bnh thng ca chng ct t cc lin kt cn thit v hnh
thnh cc lin kt mi. Nng lng hot ha cc phn ng no cng ln
th tc phn ng cng chm v ngc li.
Vic a mt s cht no vo trong h thng vn c tc dng lm
tng tc phn ng ha hc c gi l s xc tc. Trong qu trnh xc
tc, cc cht xc tc lm gim nng lng hot ha ca phn ng ha
hc. Nng lng cn xy ra s va chm c hiu lc dn n phn ng
ha hc c gi l nng lng hot ha.
38
Trong qu trnh xc tc, cc cht xc tc ch tham gia vo cc phn
ng trung gian sau chng li c phc hi nhanh chng, cht xc tc
khng ng vai tr nh nhng cht tham gia phn ng.
Tc cc phn ng thun nghch cng c cc cht xc tc lm
tng theo c hai chiu.
iu ni ln rng: cht xc tc khng quyt nh chiu hng ca
phn ng, chng ch thc y cho phn ng t n cn bng mt cch
nhanh chng.
nh hng ca cht xc tc l lm gim nng lng hot ha. Trong
th sau: A l cht phn ng, B l sn phm ca phn ng.



















II ENZYME L CHT XC TC SINH HC
Nhng lun c sau y chng minh bn cht xc tc ca enzyme:
a) Cng nh cc cht xc tc ni chung, enzyme lm tng nhanh tc
phn ng. Enzyme khng quyt nh chiu hng ca phn ng.
b) Enzyme khng ng vai tr l cht tham gia phn ng trong phng
trnh phn ng. Trong qu trnh xc tc, lng enzyme khng thay i.
c) Cng nh mi cht xc tc khc, enzyme lm gim nng lng hot
ha cn thit ca cc phn ng ha hc. Ni cch khc nng lng hot
ha phn ng c gim i nhiu khi c s xc tc ca enzyme.




Phn ng khng c
xc tc Nng lng
Nng lng hot ha
Phn ng c hot ha c
enzyme xc tc enzyme xc tc
A

B

AG
1
AG
2

Chiu phn ng

n
g

l

n
g

t

d
o

39
III - BN CHT HA HC CA ENZYME
3.1. Enzyme l nhng cht xc tc c bn cht protein.
Trong s pht trin ca ha sinh hc, bc nhy vt t c khi
ngi ta thc hin thnh cng vic tch rt cc cht xc tc sinh hc ra
khi t bo v nghin cu tnh cht ca chng, lc ngi ta nhn bit
rng enzyme c bn cht protein. Nm 1926, Sumner l ngi u tin thu
c urease dng kt tinh. Cho n nay c khong hn 150 enzyme
c rt ra dng tinh khit. Trong s cc enzyme , mt s c
bit trn vn v cu trc bc I nh ribonuclease, trypsin, chymotrypsin,
Ngy nay ngi ta xc nhn rng, cc enzyme chnh l nhm
protein quan trng. Chng c hnh thnh trong t bo nh cc protein
n gin (enzyme mt thnh phn) hoc nh cc protein phc tp (enzyme
hai thnh phn). Trong s cc enzyme th a s l enzyme hai thnh phn.
Dng hot ng ca enzyme hai thnh phn bao gm phn protein v
phn khng c bn cht protein gi l nhm prostetic (nhm ngoi, nhm
ghp, )
Enzyme mt thnh phn l cc protein n gin thc hin chc nng
xc tc. V d: Ribonuclease A v mt s enzyme thy phn protein v
mt s enzyme khc.

3.2. Cc nhm ghp, cc coenzyme
Bn cnh phn protein th enzyme hai thnh phn cn cha phn
khng c bn cht protein. Ngi ta gi phn khng phi protein cn thit
bt buc i vi hot ng ca enzyme l nhm ghp (nhm ngoi, nhm
thm, yu t ph, ) v phn protein vn lin kt vi nhm l
apoenzyme, phc hp ca hai thnh phn trn l holoenzyme (enzyme hai
thnh phn).
Trong trng hp nhm ghp l nhng cht hu c c trng lng
phn t b c lin kt vi phn protein th nhm ghp c gi l
coenzyme. Theo cch th:




Nu ng ring r th c coenzyme cng nh apoenzyme u khng c
kh nng xc tc. Ch c lc no 2 phn ny kt hp vi nhau th hot tnh
xc tc ca enzyme mi th hin.
Bn cht ha hc ca coenzyme rt khc nhau:
- Mt s loi ny chnh l cc vitamin. S lin quan v chc nng gia
cc vitamin v cc coenzyme c gii thiu bng sau:
Apoenzyme + Coenzyme Holoenzyme
40

Coenzyme Chc nng Vitamin tng
ng
NAD, NADP
FAD. FMN
Coenzyme A


Thiaminpyro(P)

Pyridoxal(P)
- Chuyn H
+
v e
-
- Chuyn H
+
v e
-
- Vn chuyn gc acyl
- Phn gii ho kh v tng hp
acid bo
- Kh carboxyl ha
- Chuyn nhm aldehyd
- Chuyn amine ha
- Kh carboxyl ha

PP
B
2

Pantotenic acid
(B
3
)

Thiamine(B
1
)

Pyridoxine(B
6
)

- Cc ion kim loi c vai tr cn thit bt buc cho s hot ng ca
enzyme. Cc ion (cation) c chc nng ging vi cc coenzyme.
Ngi ta gi cc ion kim loi l cc coenzyme n gin. Cc
enzyme cn ion kim loi cho vic thc hin chc nng ca mnh c gi
l metalloenzyme. Chc nng ca cc ion kim loi ni chung phn nhiu
l chng to ra cc phc hp kiu chelate gia mt nhm nht nh no
ca c cht v enzyme. Trc tin cc cation to phc ion kim loi c
cht, sau phc hp ny mi phn ng vi enzyme. Cc ion Ca, Cu,
Mg, Mn, Mo, Zn, u l nhng ion tham gia trong s hot ng ca cc
enzyme.

3.3. Tnh c hiu ca enzyme
Kh nng xc tc vi tnh c hiu cao l mt trong nhng c tnh c
bn v quan trng nht ca enzyme. Tnh c hiu ca enzyme th hin
ch: enzyme ch xc tc cho mt trong v s nhng chuyn ha c th c
c i vi cc cht. C hai loi c hiu c bn l c hiu phn ng
v c hiu c cht.

* Tnh c hiu phn ng: c th hin ch enzyme ch c kh
nng la chn mt dng phn ng trong s cc phn ng v xc tc cho
phn ng . iu thy r trong v d sau y:





41




















Di tc dng ca oxidase, aminoacid b kh amine ha bng cch
oxy ha to ra cetoacid v NH
3
. Vi s c mt ca oxidase, mt phn
ng khc kh carboxyl ha li khng th xy ra. Vic xc tc ny i hi
phi c enzyme khc, l decarboxylase. Cng nh vy, phn ng
chuyn amine ha i hi phi c transaminase.

* Tnh c hiu c cht: Enzyme c th la chn i vi cc cht
tham gia phn ng. Khng phi mi c cht c kh nng phn ng u
c enzyme tip nhn nh nhau.
Mi enzyme ch chuyn xc tc cho mt hoc mt vi c cht nht
nh v mc c hiu ca n ty thuc vo tng loi enzyme. C 3
mc c hiu c cht ch yu:

a) c hiu tng i: Enzyme ch c th tc dng ln mt kiu
lin kt ha hc nht nh m khng ph thuc vo cc nhm ha hc nm
hai bn lin kt. V d cc esterase c th tc dng ln hng lot cc
ester ca phosphoric acid.

b) c hiu nhm: biu hin l enzyme ch c th tc dng ln mt
kiu lin kt ha hc nht nh v mt trong hai nhm nm hai bn lin
R
1
C COOH + NH
3

2H
oxidase O Cetoacid(1)
H
2
O

R
1
CH COOH decarboxylase R
1
CH
2
+ CO
2


NH
2
NH
2
Amine
Aminoacid (1) Transaminase

R
2
CH COOH

NH
2
Aminoacid (2)


R
2
C COOH R
1
C COOH

O O
Cetoacid (2) Cetoacid (1)





42
kt cng phi c cu to nht nh. V d carboxylpeptidase c kh nng
phn hy lin kt peptide gn nhm COOH t do, ngha l lin kt
peptide cui mch polypeptide,





c) c hiu tuyt i: Enzyme ch tc dng ln mt kiu lin kt
nht nh v cc nhm ha hc hai bn lin kt cng phi xc nh.
V d: Enzyme trypsine thy phn cc lin kt peptide gia lysine hoc
arginine vi bt c aminoacid no. Sn phm l nhng on peptide c
lysine hoc arginine cha nhm COOH t do pha tn cng ca peptide.








- Enzyme Trombine cn c tnh c hiu cao hn trypsine: n ch
thy phn lin kt peptide pha carboxyl ca gc arginine no c gc
glycine ng lin k sau n:








* Ngoi cc tnh c hiu trn nhiu enzyme cn biu hin rt cao
v tnh c hiu ha hc lp th (c hiu quang hc): Enzyme ch tc
dng ln nhng dng ng phn lp th no ca cc cht hu c.
V d: Enzyme L-lactatdehydrogenase ch tc dng ln L-lactic
acid m khng tc dng ln D-Lactic acid. Mun tc dng ln D-Lactic
acid phi c enzyme D-lactatdehydrogenase.
IV CC YU T NH HNG TI HOT TNH XC
TC CA ENZYME
O O

C NH CH C

R
V tr thy phn ca trypsine


Lysine hay
Arginine
O O

C NH CH C



V tr thy phn ca trombine

Arginine
Glycine
.. CH CONH CH COOH .. CH COOH + H
2
N CH COOH

R
1
R
2
R
1
R
2

43
4.1. Nhit
Trong phm vi l hc, tc ca phn ng tng ln cng vi s tng
ca nhit . Nhng khi vt qu phm vi no , cc phn ng c
enzyme xc tc b nh hng do s bin tnh ca phn t protein-enzyme.
Kt qu ny ph thuc vo nhit ti thch ca enzyme, l nhit m
ti tc phn ng enzyme t cc i.
Mi enzyme c nhit ti thch khc nhau. S khc nhau ny ty
thuc vo ngun gc ca cc enzyme, ty theo tng iu kin hoc tng
s khc nhau v tnh nhy cm vi nhit ca phn t protein-enzyme.
a s enzyme mt hot tnh xc tc nhit cao(>80
o
C), tr papain,
myokinase c th tn ti 100
o
C.











4.2. nh hng ca pH
Mi enzyme u c tr s pH ti thch no i vi hot tnh
ca chng. ngoi phm vi ca tr s ny hot tnh ca enzyme u b
gim thp.
Tr s pH ti thch ca mt s enzyme nh sau:

Enzyme pH ti thch
Pepsine
Amylase(mch nha)
Amylase(nc bt)
Trypsine
Arginase
Catalase
Peroxidase
1,5 2,5
4,6 5,0
6,8 7,2
7,8 9,5
9,8
6,8 7,0
6,0
Nhng nguyn nhn sau y c th dn ti s ph thuc vo pH
ca enzyme:
Hot
tnh
Nhit ti thch





20 30 40 50 60 70 80 t
o
C
44
a) Nu trong s cc nhm bn tham gia trc tip trong s hot
ng ca enzyme cha nhm c kh nng phn ly.
b) pH nh hng ti cc nhm phn ly khc ca protein-enzyme
vn c tc dng trong vic duy tr cu hnh c hot tnh ca enzyme.
c) S thay i pH ca mi trng c th nh hng ti cc nhm
phn ly ca c cht hay ca coenzyme vn c kt hp vi enzyme.

4.3. nh hng ca cht hot ha v cht km hm enzyme
Nhng cht no c kh nng lm tng hot tnh xc tc ca
enzyme th c gi l cht hot ha enzyme. Cc cht thng l cc
ion kim loi nh: K
+
, Na
+
, Mg
+2
, Ca
+2
, Co
+2
, Zn
+2
, Mn
+2
,
V d: Mg
+2
lm tng hot tnh phosphatase
Ca
+2
lm tng hot tnh lypase.
S hot ng ca cc enzyme u c th b km hm bi cc tc ng
gy bin tnh protein. Ngi ta phn bit cc hnh thc km hm enzyme
v phn bit cc cht km hm enzyme nh sau:
a) Cht km hm chung: cc cht ny km hm hot tnh xc tc
ca tt c cc enzyme. Cc cht ny l cc mui kim loi nng, cht
tannin.
b) Cht km hm ring: c tc dng km hm mt hay mt nhm
enzyme c cu to gn ging nhau. V d: cc cht cha nhm CN km
hm enzyme h hp.

4.4. Nng c cht v nng enzyme
Khi mi trng c y c cht th tc phn ng t l thun
vi lng enzyme. Khi nng c cht thp, khng li ko tt c
lng enzyme vo phn ng th tc phn ng tng t l thun vi nng
c cht. Tc phn ng t ti a khi tt c enzyme u kt hp vo
c cht.

V C CH XC TC CA ENZYME
Khi t vn nghin cu v c ch xc tc ca enzyme ngi ta
xut pht t gi thit cho rng trong cc phn ng c enzyme xc tc,
phc tm thi Enzyme C cht c to thnh. Qu trnh ny gm 3
giai on:


giai on 1 phn ng xy ra tng i nhanh, c cht (S) c lin
kt vi enzyme (E) nh cc lin kt yu. Lc ny s lin kt khng gian
E + S ES ES
*
E + P
1 2 3
45
gia cc phn t c cht v enzyme cha hiu qu i vi s xc tc
ca enzyme.
giai on tip theo xy ra s bin i ca c cht (S) c lin quan
ti vic ph v hay hnh thnh cc lin kt cng ha tr. giai on ny,
c cht c hot ha (mt hoc vi phc cht ES chuyn tip c hot
ha). y, cu trc bc 3 ca enzyme lun bin i to kh nng tip
xc gia cc nhm hot ng ca enzyme vi c cht ang bin i.
Enzyme lm bin i phn t c cht lm cho cc lin kt bn trong
phn t tr nn lng lo hn, do ch cn mt lng nng lng nh
cng lm cho c cht bin thnh cc sn phm (P) khc nhau.
Ngi ta chng minh rng trong khi hnh thnh phc cht ES c 2
qu trnh ng thi xy ra nhanh chng l:
a) S thay i mt in t gy nn s phn cc ha cc lin kt.
b) S bin dng v mt ha hc ca cc lin kt ko cng trong
phn t c cht.
C hai yu t ny (s bin hnh v s phn cc ha cc lin kt
ng ha tr) u lm tng th nng nhit ng hc ca cc lin kt ny,
ngha l xc tin vic vt qua hng ro nng lng hot ha ca qu
trnh chuyn tip phc Enzyme-C cht.

VI - NG HC CA PHN NG ENZYME
6.1. ng hc ca phn ng enzyme trong trng hp khng
c cht km hm.
6.1.1. giai on: E + S ES





K
+1
: hng s vn tc ca phn ng thun.
K
-1
: hng s vn tc ca phn ng nghch.
Gi V
1
l tc phn ng thun.
V
-1
l tc phn ng nghch.
[E]: nng enzyme.
[S]: nng c cht.
Ta c: V
1
= K
+1
([E]. [S])
V
-1
= K
-1
[ES] (1)
Khi phn ng t n cn bng (enzyme phn ng ht vi c cht)
th V
1
= V
-1
ngha l:
K
+1
([E]. [S]) = K
-1
[ES]
K
+1

E + S ES
K
-1

46
t ta c:
1
1
K
K
+

=
| | | |
| | S E
S E

Nu gi K
s
l hng s cn bng cc phn ng bc I, ta c:

s
K =
| || |
| | S E
S . E

s
K =
1
1
K
K
+

(2)
* Nu K
s
c gi tr ln th K
-1
s ln v K
+1
s nh. T ta thy
phc hp ES d phn gii thnh cc cht S v E. Phn ng enzyme tin
hnh chm.
* Nu K
s
c gi tr nh th tc to ES s nhanh ng thi phn
ng enzyme cng tin hnh nhanh. Vy nu K
s
cng nh th nng ES
cng cao.
giai on 1 ta tnh c:
s
K =
| | | |
| | S E
S E

Hay vit di dng khc l: [E] [S] = K
s
[ES] (3)
Nu gi [E
o
] l nng chung ca enzyme trc khi bt u tham
gia phn ng.
[E]: nng enzyme dng t do (khng to phc hp)
[ES]: nng ca phc h ES.
th nng enzyme khng to phc hp ( dng t do) l:
[E] = [E
o
] - [ES] (4)
Thay vo phng trnh (3), ta c:
([E
o
] - [ES]). [S] = K
s
. [ES]
Trin khai:
[E
o
] [S] - [ES] [S] = K
s
. [ES] hay l:
K
s
[ES] + [ES] [S] = [E
o
] [S] hay l:
[ES](K
s
+ [S]) = [E
o
] [S]
T
[ES] = [E
o
]
| |
| | S K
S
s
+

| |
| |
o
E
ES
=
| |
| | S K
S
s
+
(5)
Nh vy nng [ES] cng ln th tc phn ng enzyme cng
ln (v K
s
lc s nh). Tc s t ti ti a khi nng phc h
[ES] bng nng enzyme ban u [E
o
] ([ES] = [E
o
]) ngha l khi tt c
enzyme u c kt hp vi c cht th phn ng enzyme l ti a.
T c th lp thnh t l:
max
V
V
=
| |
| |
o
E
ES

V: tc phn ng enzyme ( giai on u)
V
max
: tc ti a.
47
T phng trnh (5) ta c:
| |
| |
o
E
ES
=
| |
| | S K
S
s
+

Vy
max
V
V
=
| |
| | S K
S
s
+

T : V = V
max
| |
| | S K
S
s
+

y l phng trnh Michaelis v Menten (1913) dng tnh tc
phn ng enzyme t E + S ES.
Nu K
s
= [S] th V =
2
1
V
max


6.1.2. giai on: ES
*
E + P
Khi ta c tc to ES l: V
+1
= K
+1
([E] [S])
Tc phn li ES theo phn ng nghch V
-1
= K
-1
[ES]
Tnh tc phn gii phc h ES to E + P, hng s tc ny
bng K
+2





Khi nng c cht [S] cao hn nhiu so vi nng [E] th s
t nhanh chng trng thi cn bng gia s hnh thnh ES v s phn gii
phc h ES .
Nh vy giai on ny ta c:






Hay l: (K
-1
+ K
+2
). [ES] = K
+1
[E] [S] (6)
T phng trnh (4): [E] = [E
o
] - [ES]
Thay vo phng trnh (6), ta c:
(K
-1
+ K
+2
). [ES] = K
+1
([E
o
] - [ES]). [S]


t :
K
+1
K
+2
E + S ES E + P
K
-1
K
-2

K
-1
[ES] + K
+2
[ES] = K
+1
[E] [S]


Tc phn gii Tc phn gii Tc to phc h ES
phc h ES E + S phc h ES E + P
48
1
2 1
K
K K
+
+
+
=
| | | | ( )| |
| | ES
S . S E E
o

=
| || | | || |
| | ES
S S E S E
o



t
1
2 1
K
K K
+
+
+
= K
m
K
m
=
| || | | || |
| | ES
S S E S E
o


K
m
c gi l hng s Michaelis (o bng mol/lit) (tnh theo khi
m s to [ES] v phn li [ES] bng nhau)
K
m
[ES] = [E
o
] [S] - [ES] [S]
K
m
[ES] + [ES] [S] = [E
o
] [S]
[ES](K
m
+ [S]) = [E
o
] [S]

[ES] =
| || |
| | S K
S . E
m
o
+
(7)
Mt khc tc phn ng tnh theo s hnh thnh sn phm P t
[ES] bng:
V = K
+2
[ES]
t : [ES] =
2
K
V
+

Thay vo (7) ta c:
2
K
V
+
=
| || |
| | S K
S . E
m
o
+

Vy V =
| || |
| | S K
S . E . K
m
o 2
+
+
(8)

Phng trnh ny cho php tnh c s ph thuc tc phn
ng vo nng ca enzyme v c cht.
V t cc i khi [ES] = [E
o
] ngha l khi E c kt hp hon
ton vi S.

max
V
V
=
| |
| |
0
E
ES
t phng trnh 7 ta c: ES =
| || |
| | S K
S E
m
0
+

Vy
max
V
V
=
| |
0
E
1
.
| || |
| | S K
S E
m
0
+
=
| |
| | S K
S
m
+



T : y l phng trnh
Michaelis v Menten dng
V = V
max
.
| |
| | S K
S
m
+

49
tnh tc phn ng t
ES
*
E + P
* Vy V =
2
1
.V
max
th

K
m
= [S]
Lc hng s Michaelis c o bng mol/lt
So snh gi tr K
m
v K
s
ta thy:
K
m
=
1
2 1
K
K K
+
+
+

m K
s
=
1
1
K
K
+

Vy K
m
= K
s
+
1
2
K
K
+
+

Khi K
+2
= K
+1
th K
m
= K
s
+ 1
T phng trnh (8) V = V
max
.
| |
| | S K
S
m
+
c th vit di dng
phng trnh:





Theo phng trnh ny th khi [S] tng th tc phn ng tng.
Nhng khi nng c cht [S] tng n mt mc no th t ti gi
tr tc cc i:V = V
max
. Sau th V khng tng theo nng c cht
[S] v V lun lun =V
max
.
l phng trnh biu din s ph thuc ca tc phn ng vi
nng c cht. C th biu din bng dng hiperbol:









C th gii thch ca ng biu din dng hiperbol nh sau:
Trong phn ng c enzyme xc tc s phn t c cht ln hn s phn t
| |
1
S
K
V
V
m
max
+
=
Tc
V
max C

2
V
max
B

0
K
m
[S]


50
enzyme rt nhiu (S>E). C cht ch c chuyn ho khi to thnh phc
h enzyme. Do tc phn ng ph thuc vo nng phc h [ES].
Nng [ES] 3 im A, B, C khng ging nhau. im A v B
cn vi phn t enzyme cha kt hp vi c cht, do nu tng hay
gim nng c cht [S] s lm tng hay gim s va chm gia E v S
ngha l lm tng hay gim nng [ES] v s nh hng ti tc phn
ng v tc l hm s ca nng c cht [S].
Ti im C tt c phn t enzyme kt hp vi S nn khi tng
nng c cht c th lm tng dn s va chm gia E v S nhng khng
lm tng tc phn ng.
V khng cn enzyme t do c th gn thm vo c cht v tc
t c l cc i.
im B ng
2
1
s phn t enzyme kt hp vi c cht,
tc =
2
1
V
max
v nng c cht tng ng vi im B bng
hng s K
m
.
T phng trnh Michaelis Menten




Nu V =
2
1
V
max
th K
m
= [S]. Nh vy ngha l hng s Michaelis
bng nng ca c cht khi m tc phn ng bng
2
1
V
max
.
* Phng trnh Michaelis cng c th vit di dng khc theo
ngh ca Lineweaver v Burk (1934) bng cch ly s nghch o:

V
1
=
| |
| | S . V
S K
max
m
+

V
1
=
max
m
V
K
.
| | S
1
+
max
V
1


Phng trnh c dng y = ax + b v ng biu din c dng
ng thng nh sau:




V = V
max
.
| |
| | S K
S
m
+


V
1




max
V
1



51











Qua th c th xc nh c V
max
v K
m
.

6.2. ng hc ca cc phn ng enzyme trong trng hp
c cht km hm
Trong trng hp c cht km hm cnh tranh hay khng cnh
tranh th gi tr V ca phn ng s bao gm c hng s K
i
(K
i
l hng s
phn li ca phc hp enzyme-cht km hm EI hay cn gi l hng s
km hm).
Hng s K
i
c tnh nh phng trnh sau:




K
i
=
1 i
1 i
K
K
+

=
| | | |
| | EI
I E

* Cc cht km hm cnh tranh: l cc cht thng c cu to gn
ging vi c cht. Cc cht km hm cnh tranh c tnh c hiu rt cao,
ngha l ch km hm mt enzyme ring bit.
V d: Malonic acid l cht km hm cnh tranh ca enzyme
succinat-dehydrogenase v n c cu to gn ging succinic acid.





Khi tng nng c cht th mt km hm cnh tranh.
* Cc cht km hm khng cnh tranh: l cc cht c tc dng km
hm hot ng ca enzyme bng cch gn vo cc v tr khc vi cc v tr
K
i+1
E + I EI
K
i-1
CH
2
COOH

COOH
Malonic acid
CH
2
COOH

CH
2
COOH
Succinic acid
52
gn c cht ca enzyme. Trong trng hp ny ch lm gim V cc i
ca phn ng ch khng nh hng g n hng s K
m
.
V d: Cc ion kim loi nng Ag
+
; Hg
+
; Trichloroacetic acid.

6.2.1. ng hc ca phn ng enzyme trong trng hp c cht km
hm cnh tranh.
Vic tnh ton V
i
s phc tp hn v ta c ng thi vi qu trnh gn
E vi c cht, cn c c qu trnh gn E vo cht km hm I.










Bng cch tnh ton nh trn ta c :

V
i
=
| |
| | | |
|
|
.
|

\
|
+ + I .
K
K
S K
S . V
i
m
m
max i

Phng trnh ny ging phng trnh Michaelis nhng c thm
| | I .
K
K
i
m
phn nh qu trnh km hm bi cc cht km hm cnh tranh.
* Cng c th tnh V
i
bng cch :

i
V
1
=
max i
V
1
.
| |
|
|
.
|

\
|
+
i
m
m
K
I . K
K .
| | S
1
+
max i
V
1


6.2.2. ng hc ca phn ng enzyme trong trng hp c cht
km hm khng cnh tranh:
Trong trng hp ny ngi ta thy rng cht km hm khng cnh
tranh c th gn vo enzyme t do v c vo phc h ES theo cc phn
ng sau:



K
+1
K
+2
E + S ES E + P
K
-1

K
i+1
E + I EI
K
i-1


K
+1
K
+2
E + S ES E + P
K
-1




53
















Cch tnh phc tp hn:
- Gi thit rng K
+2
rt nh so vi K
+1
v K
-2
, v phc hp IES khng
to thnh sn phm tng ng v cc hng s cn bng c gi tr nh nhau
(tng ng vi cc phn ng ghi trn), bng cch tnh nh trn s c:
V
i
=
| || |
| | ( )
| |
|
|
.
|

\
|
+ +
+
i
s
o 2
K
I
1 . S K
S E . K

hoc tnh theo dng khc:

i
V
1
=
| |
|
|
.
|

\
|
+
i
K
I
1
| |
|
|
.
|

\
|
+
S
1
.
V
K
V
1
max
m
max


Khi so snh tc phn ng khng b km hm vi tc phn ng c
cht km hm khng cnh tranh ngi ta thu c t l sau:

i
o
V
V
=
| |
i
K
I
1+
V
o
l tc ban u ca phn ng enzyme khng c cht km hm.
V
o
=
| || |
| | S K
S E . K
s
o 2
+
+
t (8)
y K
s
thay vo ch ca K
m
v K
+2
rt nh so vi K
-1
ngha l
K
m
= K
s
. Nh vy c th thy rng: S bin i tng i ca hot tnh
K
i+1
E + I EI
K
i-1

K
i+2
ES + I IES
K
i-2

K
i+3
EI + S IES
K
i-3

54
enzyme c cht km hm khng cnh tranh ch ph thuc vo nng ca
cht km hm m khng ph thuc vo nng ca c cht [S].
* i vi cht km hm cnh tranh th t l
i
o
V
V
c dng:
i
o
V
V
= 1 +
| |
| | I
S K
K K
m
i m
+
t phng trnh ny ta thy s bin i ca
hot tnh enzyme khi c cht km hm cnh tranh ph thuc c vo nng
c cht v nng cht km hm.
C th biu din tc phn ng enzyme trong trng hp c cht
km hm cnh tranh v khng cnh tranh bng th:



























Tm tt: (Tt c nhng phng trnh sau y ch dng cho trng hp
khi 1 c cht tham gia vo phn ng)
V V
V
max
= V
imax
V
max
V
imax
1 2 1 2




K
m
K
mi
[S] K
m
= K
i
[S]


V
1
2 1
V
1
2

max
V
1
=
max i
V
1

max i
V
1
1

max
V
1



m
K
1

mi
K
1

| | S
1

m
K
1
=
mi
K
1

| | S
1

1 c cht km hm.
2 khng c cht km hm.

C cht km hm cnh tranh C cht km hm khng cnh tranh

55



Khng c
cht km hm
C cht km hm
cnh tranh

C cht km hm
khng cnh tranh

Phng
trnh

V
1
=
max
m
V
K
.
| | S
1
+
max
V
1


i
V
1
=
max i
V
1
.
| |
|
|
.
|

\
|
+
i
m
m
K
I . K
K .
| | S
1
+
max i
V
1



i
V
1
=
| |
|
|
.
|

\
|
+
i
K
I
1 .
| |
|
|
.
|

\
|
+ +
S
1
V
K
V
1
max i
m
max i

on
thng trn
trc tung
ti im
max
V
1




max
V
1



max
V
1



max
V
1
.
| |
|
|
.
|

\
|
+
i
K
I
1
on
thng ct
trn trc
honh ti
im
| | S
1



m
K
1



| |
|
|
.
|

\
|
+

i
m
K
I
1 K
1



m
K
1



VII . CCH GI TN V PHN LOI ENZYME
Trong thi gian c ch tc dng ca enzyme cha c nu ra, ngi
ta t cho mt s enzyme nhng tn ring bit nh pepsine, trypsine,
chymotrypsine, papaine, bromeline,
Khi con s cc enzyme c bit ngy cng nhiu tn enzyme c
gi theo nguyn tc sau y: Tn enzyme = tn c cht m enzyme xc tc
+ loi phn ng m enzyme xc tc + tip v ng ase.

V d:


Ascorbat-
Ascorbic acid + O
2
Dehydroascorbic acid + H
2
O
oxidase
56







Vic phn loi enzyme l cng vic kh khn v s enzyme m c ch
xc tc ca n c hiu bit mt cch tng tn khng nhiu.
Vic phn loi enzyme c da theo nguyn tc l: ly c s kiu
phn ng do enzyme xc tc m Hip hi Ha sinh Quc t xut
nm 1964. Nm 1973 h thng phn loi ny li tip tc c hon thin
bi y ban danh php Ha sinh thuc Hip hi ha hc c bn v ng
dng Quc t (IUPAC). Trn c s tt c cc enzyme c phn chia 6
nhm ch yu sau:
1. Oxydoreductase 4. Lyase
2. Transferase 5. Isomerase
3. Hydrolase 6. Synthetase (Ligase)
Cc s thp phn trn bng phn loi c ngha nh sau:
- S th nht ch nhm chnh (V d nhm 2: transferase)
- S th hai qui nh mt s c tnh ca phn ng (V d: 2.1 cho bit
enzyme vn chuyn gc 1 carbon)
- S tip theo cho bit chi tit hn (V d 2.1.1: c ngha l enzyme
vn chuyn nhm methyl)
Tuy nhin cho n nay cc tn gi truyn thng ca enzyme vn cn
c s dng.

VIII. CC NHM ENZYME RING BIT
8.1. Oxydoreductase (cc enzyme oxy ha kh)
Cc enzyme nhm ny lm nhim v xc tc cho cc phn ng oxy
ha kh. Chng chuyn vn H
+
hay cc in t v xc tc cho s oxy ha
sinh hc. Cc enzyme ny c vai tr to ln trong qu trnh h hp v trao
i nng lng trong c th. S tng qut ca cc phn ng c xc
tc bi oxydoreductase nh sau:



Trong AH
2
l c cht; B l cht nhn H no , c th l O
2
.
Trong t bo oxydoreductase to nn cc h thng (c gi l
chui enzyme oxy ha kh), trong vic vn chuyn cc nguyn t
hydro (H
+
v e
-
) qua nhiu giai on, t c cht n cht nhn cui cng l
CH
2
COOH Succinat- CH COOH
+ FAD + FAD.H
2

CH
2
COOH dehydrogenase CH COOH
Succinic acid Fumaric acid
AH
2
+ B A + BH
2

oxydoreductase
57
O
2
. Kt qu l cc nguyn t hydro c vn chuyn n O
2
v s to
thnh H
2
O.
S h thng enzyme h hp v vic vn chuyn H
+
v e
-
c
trnh by tng qut sau y:
















Ty theo kh nng ca oxydoreductase chuyn H
2
va ly c t c
cht (AH
2
) n O
2
ca khng kh, ngi ta chia oxydoreductase thnh hai
phn nhm:
- Dehydrogenase ho kh (oxidase)
- Dehydrogenase ym kh.

a) Dehydrogenase ym kh: l nhng enzyme khng c kh nng
vn chuyn H
+
v e
-
t c cht oxy ha n trc tip cho O
2
ca khng
kh, m chng s chuyn H
+
v e
-
cho cc cht vn chuyn trung
gian khc.
Nhm ghp ca dehydrogenase ym kh l dn xut ca vitamin
PP (B
5
) l NAD v NADP.










NH
2

O N N

C NH
2
(NAD
+
)
N
N
O CH
2
O (P)
-
O (P)
-
O CH
2
O
N
(+)

H H H H

2H
+
, 2e
-
2H
+
, 2e
-

AH
2
NAD NADH
2
FAD FADH
2

(NADP) (NADPH
2
) 2H
+
, 2e
-

Q

2H
+
QH
2

2e
-

2cytochrome b
2e
-

2cytochrome c
2e
-

2cytochrome a
O
-2
O
2
2e
- H
2
O
58












b) Dehydrogenase ho kh: l nhng enzyme c kh nng vn
chuyn H
+
v e
-
n trc tip cho O
2
ca khng kh. Chng cng l nhng
enzyme hai thnh phn. Nhm ghp ca chng l dn xut ca vitamin B
2
(Riboflavin) l FMN v FAD.





















Cc enzyme cha FMN, FAD thng khng oxy ha trc tip c cht
m chng ly H
2
t NADH
2
(hoc t NADPH
2
) ri chuyn in t cho h
cytochrome.
O

N
H
3
C NH

NH
2
H
3
C
N N O
N N
CH
2


(CH OH)
3
N N FAD

CH
2
O (P) O (P) CH
2
O

H H

FMN H OH HO H
59
a s enzyme cha FMN khi tc dng trc tip vi O
2
khng kh s
to thnh H
2
O
2
. V d: enzyme glycolatoxidase xc tc cho phn ng oxy
ha glycolic acid theo s sau:












8.2. Nhm transferase (cc enzyme vn chuyn)
Cc enzyme nhm ny lm nhim v vn chuyn cc nhm ha hc t
hp cht ny sang hp cht khc m khng lm thay i ha tr ca cc
nguyn t.
S tng qut:


x: l nhm c vn chuyn.
Ty thuc vo x m ta c cc tn enzyme vn chuyn khc nhau.
V d: Acyltransferase: x l gc cc acid bo trong c gc acetic
acid. y l enzyme hai thnh phn trong nhm hot ng l HS-CoA
c cu to nh sau:










- Glycozyltransferase: Nu x l gc glycozyl.
- Aminotransferase: Nu x l nhm amin ( NH
2
).
- Methyltransferase: Nu x l nhm (CH
3
).
COOH Glycolatoxidase

CH
2
OH FMN H
2
O
2
Glycolic acid
O
C H FMNH
2
O
2

COOH
Glyoxilic acid
Ax + B A + Bx
transferase
NH
2
H
3
C CH
3
N N
O O SH

CH
2
O (P) O (P) O
O NH NH
N N H H OH
H H
OH O (P)


60
- Aldotransferase: Nu x l nhm aldehyd.
- Cetotransferase: Nu x l nhm cetone.
- Phosphotransferase: Nu x l gc H
3
PO
4
.
Enzyme ny cn c tn l:
a) Kinase: Nu xc tc cho phn ng chuyn gc H
3
PO
4
m ATP
l nhm hot ng ca enzyme ny. V d:




b) Mutase: l enzyme vn chuyn gc H
3
PO
4
trong ni ti phn t
ca 1 cht.
V d: Enzyme hexoso(P)-mutase xc tc cho phn ng sau:











8.3. Nhm hydrolase (cc enzyme thy phn):
Cc enzyme nhm ny xc tc cho cc phn ng thy phn. S
tng qut:




Da vo lin kt gia R
1
v R
2
m ta c tn cc enzyme thy phn
khc nhau:
- Peptidase: Thy phn lin kt peptide.
- Glycosidase: Thy phn lin kt glycoside.
- Esterase: Thy phn lin kt ester.
- Amidase: Thy phn amide.




Glycerine + ATP Glycerol(P) + ADP
Kinase
R
1
R
2
+ H
2
O R
1
H + R
2
OH
Hydrolase
R C NH
2
+ H
2
O R COOH + NH
3
Amidase
O
Amide
6 6
CH
2
OH CH
2
O (P)
5 O 5 O
H H H Hexoso(P) - H H H
4 1 4 1
HO OH H O (P) mutase HO OH H OH
3 2 3 2
H OH H OH
Glucoso-1(P) Glucoso-6(P)
61


V d:















8.4. Nhm Liase (cc enzyme phn ct)
Enzyme nhm ny phn ct mt nhm no ra khi mt hp cht m
khng c s tham gia ca H
2
O. C trng hp kt qu ca phn ng l to
lin kt i.
V d: * Decarboxylase: xc tc cho phn ng sau:








* Aldolase: xc tc cho phn ng thun nghch sau:








NH
2
C = O + H
2
O CO
2
+ 2NH
3

Urease
NH
2

Ure
H
2
N CH COOH H
2
N CH COOH
NH
3

CH
2
+ H
2
O CH
2

Asparaginase
CONH
2
COOH
Asparagin Aspactic acid
CO
2

R CH COOH R CH
2

Decarboxylase
NH
2
NH
2

Aminoacid Amine
Fructosodi(P) Aldehyd(P)glyceric + Phosphodioxiacetone
CH
2
O (P) O

C = O C H CH
2
OH
Aldolase
(CH OH)
3
CH OH + C = O

CH
2
O(P) CH
2
O(P) CH
2
O(P)
Fructoso 1,6 di(P) Aldehyd(P)glyceric Phosphodioxiacetone
62






* Hydratase (hydroliase): gm cc enzyme xc tc cho phn ng
loi tr v kt hp H
2
O.














8.5. Nhm Isomerase (cc enzyme ng phn ha):
Cc enzyme nhm ny xc tc cho phn ng chuyn ha cc cht hu
c thnh ng phn ca chng. V d chuyn ha gia dng cetone v
dng aldehyd. Gia dng L v dng D
V d: Enzyme trioso(P)isomerase xc tc cho phn ng sau:









Enzyme alaninracemase xc tc cho phn ng bin i L-alanine thnh
D-alanine v ngc li.


CH
2
COOH H
2
O CH
2
- COOH

C(OH).COOH C COOH
Aconitat-hydratase
CH
2
COOH CH - COOH
Citric acid Cis-aconitic acid
CH
2
COOH
H
2
O
CH COOH Aconitat-hydratase

CH.OH.COOH
Isocitric acid
COOH COOH
Alaninracemase
H
2
N C H H C NH
2


CH
3
CH
3

L-Alanine D-Alanine
O

CH
2
OH C H

C = O CH OH

CH
2
O(P) CH
2
O(P)
Phosphodioxiacetone Aldehyd(P)glyceric
63







8.6. Nhm synthetase (cc enzyme tng hp)
Cc enzyme nhm ny xc tc cho cc phn ng tng hp cc cht
hu c. Nhm ny ch xc tc khi c mt ca ATP v cc hp cht cao
nng khc. V cc phn ng tng hp i hi phi cung cp nhiu nng
lng. Mt s v d sau y:






















Cu hi
1. nh ngha enzyme? Phn nhm enzyme v cho v d?
2. Cch gi tn enzyme v cho v d?
3. Mi lin quan gia enzyme v vitamin?
4. Ti sao ni: Enzyme l nhng cht xc tc c bn cht
protein?.
ATP ADP + H
3
PO
4

* Aspactic acid + NH
3
Asparagine
H
2
O
Asparaginsynthetase
ATP ADP + H
3
PO
4

* Glutamic acid + NH
3
Glutamine
H
2
O
Glutaminsynthetase
ATP AMP + H
4
P
2
O
7

* CH
3
COOH + HS-CoA CH
3
C ~ S.CoA
Acetic acid Acetyl-CoA-Synthetase
O
Acetyl-CoA
64
5. Cho bit s khc nhau (v mt chc nng) gia Trombin v Tripxin?
6. C ch tc ng ca enzyme?
7. Cc iu kin nh hng n tc phn ng ca enzyme?
56
CHNG 4
CARBOHYDRATE V S TRAO I
CARBOHYDRATE TRONG C TH THC
VT

4.1 Cu to, tnh cht v vai tr ca carbohydrate
Carbohydrate l nhm cht hu c ph bin kh rng ri trong c
th sinh vt. Nhn chung hm lng carbohydrate thc vt cao hn
ng vt. thc vt carbohydrate tp trung ch yu thnh t bo, m
nng v m d tr. Tuy nhin hm lng carbohydrate thay i tu
theo loi, giai on sinh trng, pht trin. Trong c th ngi v ng vt
carbohydrate tp trung ch yu trong gan.
Thc vt xanh c kh nng s dng nng lng nh sng tng
hp carbohydrate t CO
2
v H
2
O. Carbohydrate thc vt l ngun dinh
dng quan trng ca ngi v ng vt.
Trong c th sng carbohydrate gi nhiu vai tr quan trng nh:
- Cung cp nng lng cho c th, carbohydrate m bo khong
60% nng lng cho cc qu trnh sng.
- C vai tr cu trc, to hnh (v d: cellulose, peptidglican...)
- C vai tr bo v (mucopolysaccharide)
- Gp phn bo m tng tc c hiu ca t bo (polysaccharide
trn mng t bo hng cu, thnh t bo mt s vi sinh vt).
Da vo cu to, tnh cht carbohydrate c chia lm hai nhm
ln: monosaccharide v polysaccharide.

4.2 Cc monosaccharide
Monosaccharide l cc aldehyd hoc cetone c cha mt hoc nhiu
nhm hydroxyl. S lng carbon trong phn t monosaccharide t nht l
3, l glyceraldehyd v dihydroxyacetone. Chng c cng thc nh sau:

O H

C

HOCH

CH
2
OH
L-glyceraldehyd
O H

C

HCOH

CH
2
OH
D-glyceraldehyd
CH
2
OH

C = O

CH
2
OH


Dihydroxyacetone
57
Glyceraldehyd c cha 1 cacbon bt i (C*), c hai ng phn D
v L, cn dihydroxyaketone khng cha carbon bt i. S ng phn lp
th ca monosaccharide tnh theo cng thc X = 2
n
(n l s C* trong phn
t).
Nu s cacbon trong phn t monosaccharide l 4, 5, 6 hoc 7 chng c
tn gi tng ng l tetrose, pentose, hexose v heptose. Cng thc cu
to ca mt s monosaccharide thng gp c trnh by di y:



CHO

HCOH

HCOH

CH
2
OH



D-Erythrose

CHO

HCOH

HCOH

HCOH

CH
2
OH

D-Ribose

CH
2
OH

C = O

HC-OH

HCOH

CH
2
OH

D-Ribulose

CH
2
OH

C = O

HO-CH

HCOH

CH
2
OH

D-Xilulose



CHO

HCOH

HOCH

HCOH

HCOH

CH
2
OH


D-Glucose

CH
2
OH

C = O

HO-CH

HCOH

HCOH

CH
2
OH


D-Fructose

CH
2
OH

C = O

HO-CH

HCOH

HCOH

HCOH

CH
2
OH

D-Sedoheptulose

Khi nh s th t cc nguyn t carbon trong phn t
monosaccharide, bt u t carbon ca nhm carbonyl ca aldose, hoc
carbon u gn nhm carbonyl ca cc ketose.
Cn c vo v tr ca H v OH carbon bt i mang s th t ln
nht (carbon bt i xa nhm carbonyl) ging vi D hoc L-
58
glyceraldehyd xp monosaccharide vo dng D hoc L. Cc
monosaccharide c kh nng lm quay mt phng nh sng phn cc v
bn phi (k hiu du +) hoc bn tri (k hiu du -).
Trong ho sinh hc ng 6C v 5C c ngha ln. Mt s ng
v d 1,6 fructosediphosphate v ng ribose c nu trn. Hexose
v pentose c th tn ti dng thng hoc vng. Theo ngh ca
Harworth phn t ng c vit dng vng. y ngi ta tng
tng mt mt phng nm ngang c 6 hoc 5 gc, nhng gc ca n sp
xp pha trn hoc pha di nhng chui bn vung gc vi mt phng.
dng vng xut hin mt nhm hydroxyl glycoside, nhm ny c kh
nng phn ng cao, d dng to lin kt glycoside v c tnh kh.




















Nhm hydroxyl glycoside l c trng ca ng kh, nhm ny c
th pha di hoc pha trn mt bng phn t. Nu n nm pha di th
ngi ta gi l dng , v d -glucose, nu pha trn th c gi l
dng . Glucose tn ti ch yu dng 6 cnh (pyranose), fructose v
pentose dng vng 5 cnh (furanose).
Mt lin kt glycoside xut hin khi mt phn t nguyn t H
c thay th bng mt gc ng (glycosyl), v d adenine c thay
th bi 1 ribosyl (gc ng ca ribose).
CH
2
OH
OH
HO
OH
OH
O
HO-CH
2

OH OH
OH
O
HO-CH
2

OH
HO
OH
O
CH
2
-OH
OH glycoside
- glucose
OH glycoside
- ribose
OH glycoside
- fructose
59
































Hnh 4.1 Tng hp hexosephosphate t triosephosphate

Trong trng hp ny nguyn t N ni vi 2 thnh phn, v vy ngi
ta gi N-glycoside. Tng t nh vy S-glycoside v O-glycoside cng
c to nn. Trong trng hp th nht nguyn t H ca mt nhm SH,
trong trng hp th hai nguyn t H ca mt nhm OH c thay th
bi mt gc glycosyl.
N N
N N
H
NH
2

N N
N N
NH
2

HO-CH
2

OH OH
Ribosyl
O
R
HO-CH
2

OH OH
O
R-H
CH
2
OH
OH
HO
OH
O
O P
CH
2
-O P
OH
HO
OH
O
OH
P O-CH
2

OH
HO
OH
O
CH
2
-OH Isomerase
Isomerase
Glucose 1- phosphate Glucose 6- phosphate Fructose 6- phosphate
P O-CH
2

OH
HO
OH
O
CH
2
-O P
HOH
P OH
Phosphatase
PGA
DHAP
CH
2
- O P
C = O
H
2
C-OH

OCH
HCOH
CH
2
-O P

CH
2
- O P
C = O
H
2
C-OH

OCH
HCOH
CH
2
-O P
Aldolase
Bin i thnh
dng vng
Fructose 1,6-diphosphate
60
Tnh cht chung ca cc monosaccharide
Cc monosaccharide l nhng cht khng mu, phn ln c v ngt.
Chng ho tan tt trong nc, khng tan trong dung mi hu c khng
phn cc, tan trong dung dch ethanol 80%.
Tnh cht l hc c trng ca monosaccharide l tnh hot quang
ca chng, ngha l c kh nng lm quay mt phng ca nh sng phn
cc sang phi hoc sang tri. C th o c gc quay ny bng my phn
cc k.
Ho tnh quan trng ca monosaccharide l nhng tnh cht ca
nhm chc aldehyd hoc ketone, in hnh l tnh kh.
a) Tu iu kin oxy ho monosaccharide b oxy ho thnh cc acid
khc nhau
- Khi oxy ho nh bng dung dch clo, brom, iod trong mi trng
kim, hoc dung dch kim ca cc ion kim loi, nhm chc
aldehyd ca monosaccharide b oxy ho. Ion kim loi b kh thnh
dng c ho tr thp hn hoc n kim loi t do (vi d Cu
2+

Cu
+1
, Ag
+
Ag
0
).

CHO COOH

HCOH HCOH

HOCH Oxy ho nh HOCH

HCOH HCOH

HCOH HCOH

CH
2
OH CH
2
OH

D-glucose D-gluconic acid

- Khi cht oxy ho l HNO
3
c, c hai nhm aldehyd v alcohol
bc 1 ca monosaccharide u b oxy ho to thnh acid cha 2
nhm carboxyl gi l saccharic acid:







61
CHO COOH

HCOH HCOH

HOCH HNO
3
HOCH

HCOH HCOH

HCOH HCOH

CH
2
OH COOH

D-glucose Saccharic acid

b) Di tc dng ca cc cht kh nhm carbonyl ca
monosaccharide b kh to thnh cc polyol tng ng
tin hnh phn ng kh c th dng dng kh hydro c cht xc
tc l kim loi (hn hp Hg v Na). V d khi b kh fructose to thnh 2
polyol ng phn l sorbitol v mannitol. Sorbitol cng l sn phm ca
phn ng kh glucose.

CHO CH
2
OH

HCOH HCOH

HOCH HOCH

HCOH HCOH

HCOH HCOH

CH
2
OH COOH

D-glucose D-sorbitol

Trong c th sinh vt, glucose c th b kh to thnh polyol vng
gi l inosite, v d mezoinosite l yu t sinh trng ca cc m thc vt
nui cy. Estephosphoric ca n (fitin) l ngun d tr phospho trong ht.







CH
2
OH
OH
HO
OH
OH
O
62
c) Phn ng to este thng xy ra vi nhm alcohol bc 1, OH
glycosite ca monosaccharide. Cc este phosphat ca
monosaccharide l nhng sn phm trung gian quan trng nht ca
nhiu qu trnh trao i cht trong c th sinh vt.
d) Phn ng ca nhm OH glycosite to thnh cc hp cht glycoside
Nhm OH glycoside c th phn ng vi ru to thnh este tng
ng gi l glycosite. Tu theo v tr ca nhm OH ny m - hoc -
glycoside c to thnh. Trong phn t glycoside, phn khng phi
carbonhydrate c gi l aglicon.
C th phn bit cc kiu lin kt glycosite khc nhau:
- Lin kt O-glycoside: (G-C-O-A) gc aglicon (A) kt hp vi
carbonhydrate (G) qua O. Lin kt O-glycoside l dng lin kt ca i-,
tri-, oligo- v polysaccharide.
- Lin kt S-glycoside: (G-C-S-A) gc aglicon (A) kt hp vi
carbonhydrate (G) qua S
- Lin kt N-glycoside: (G-C-N-A) gc aglicon (A) kt hp vi
gluxit (G) qua N
- Lin kt C-glycoside: (G-C-C-A) gc aglicon (A) kt hp vi
carbonhydrate (G) qua C
Lin kt glycoside khng bn vi acid, tng i bn vi kim.
Di tc dng ca acid hoc cc enzyme tng ng, glycoside b thu
phn to thnh monosaccharide v aglicon.

4.3 Cc polysaccharide
Polysaccharide do nhiu gc monosaccharide kt hp vi nhau, c
khi lng phn t ln, do polysaccharide khng c tnh kh. Cc
monosaccharide c th thuc mt loi hay nhiu loi khc nhau. Cc lin
kt glycoside trong phn t polysaccharide c th l - hoc -glycoside.
Tn polysaccharide da theo tn ca monosaccharide cu to nn n
nhng i ui ose thnh ui an. V d D-glucane, D-fructane, D-
galactane. Sau y ch cp chi tit hn mt s polysaccharide ph bin
v quan trng.
Tinh bt l polysaccharide d tr thc vt ph bin nht, l mt hn
hp ca amylose v amylopectin, trong amylopectin chim khong
80%. Amylose l mt chui khng phn nhnh gm nhiu gc glucose. C
khong 200-300 gc glucose kt hp vi nhau theo kiu -glycoside. Lin
kt lun lun nm gia C
1
v C
4
. Ngc li amylopectin l chui phn
nhnh. S phn nhnh l do lin kt glycoside gia C
1
v C
6
. Amylopectin
63
c to thnh t amylose. Gc amylose (khong 40 gc glucose) c
gn kt vi mt chui amylose khc bng lin kt -1-6-glycoside. Ngi
ta cho rng chui amylose di v c nhng chui amylopectin ngn hn c
cu to xon.
C quan d tr tinh bt l v sc lp (amyloplast), tng t nh lc
lp v pht trin t tin lc lp. C quan cha t bo d tr (ht ng cc,
c khoai ty) gm cc v sc lp xp st nhau.


Amylose


Amylopectin vi s phn nhnh 1,6



64
Tinh bt c tch lu trong v sc lp v lc lp dng ht tinh bt
v to nn v tr tinh bt tp trung. Cu trc ny i vi tng loi thc vt
l khc nhau nh hnh 4.2. Ngi ta c th nhn dng ht tinh bt c
ngun gc t cc loi thc vt khc nhau. nhng v tr tinh bt tp trung
th hin s kt hp ca cc ht tinh bt dy hn hoc tha hn. S kt hp
dy hn th hin vo ban ngy, tha hn vo ban m. Tinh bt c nhiu
trong cc loi qu v c khc nhau, nh trong ht ng cc (60-70 %) trong
c khoai ty (15-20%).





















Hnh 4.2 Cu trc ca cc ht tinh bt khc nhau

Trong lc lp tinh bt ch c tch lu tm thi v thc ra ch khi
hot ng quang hp mnh, vo ban ngy. Ban m nhng ht tinh bt
c phn gii thnh cc monosacaride, c bit l cc triose v c i ra
khi lc lp.
Glycogen l mt chui mch nhnh, i phn t c to nn t
nhiu gc glucose. Glycogen (tinh bt ca gan) l carbohydrate d tr ca
ng vt bc cao. N tn ti ch yu trong gan v trong m c. Nng
glycogen trong gan cao hn trong m c, tuy nhin v trng lng ca
m c ln hn trng lng ca gan nn m c c nng glycogen tng
s cao hn gan. Glycogen nm trong t bo cht dng ht nh, c
Khoai ty
La m
La
Ng
65
ng knh 10-40 nm. S tng hp glycogen thc hin t UDP-glucose.
N l cht cho gc glycosyl i vi enzyme tng hp glycogensynthetase.
Glycogen trong gan chu trch nhim iu khin nng ng trong
mu. Nng ng trong mu nm gia 4,5-7mM. Khi cung cp ng
qua thc n th glucose trong mu c tng hp thnh glycogen, khi cn
glucose, v d hot ng ca c th glucose c gii phng t glycogen.
Cellulose l mt thnh phn quan trng ca thnh t bo thc vt.
thnh t bo s cp n chim khong 30%, t bo gi thnh phn ny
cn cao hn. Cellulose l mt glucan, mt polysaccharide, c to nn t
cc gc glycosyl. Lin kt nm gia C
1
v C
4
v tng t lin kt trong
cellobiose. lin kt -glycoside nhm OH C
1
nm pha trn, trong
khi nhm OH C
4
nm pha di (cng thc Harworth). Nh vy
nhng nhm OH ca hai gc nh 2 pha i din nhau, ngi ta cho
rng mt bng ca phn t t quay 180
0
. Sau phn t cellulose c mt
hnh dng, m nguyn t C
6
lc nm pha trn v lc th pha di
ca phn t.



















Cellulose (mt on ct)

CH
2
OH
OH
HO
OH
OH
O
CH
2
OH
OH
HO
OH
OH
O
CH
2
OH
OH
HO
OH
OH
O
CH
2
OH
OH
HO
OH
OH
O
180
o
CH
2
OH
O
CH
2
OH
O
O
CH
2
OH
O
CH
2
OH
O
O
O
66
C khong 36 chui cellulose ring bit kt hp vi nhau thnh 1 vi
si, y cc chui lin kt vi nhau bng lin kt hydro. Cc vi si ny
c nhng vng kt tinh v dng ny enzyme kh c th tc ng. Nhng
vi si kt hp li thnh si, v b si, nh bng vi l dng si tinh
khit. Cht cho glucosyl l UDP-glucose. Enzyme, cellulose synthetase,
c gian bo.
4.4. Ho sinh quang hp
4.4.1 Pha sng trong quang hp
Trong th gii sinh vt th quang hp l mt qu trnh c bn. Thng
qua quang hp m nng lng ca nh sng mt tri chiu xung c
bin thnh nng lng ho hc. Hu ht sinh vt ca hnh tinh chng ta
sng trc tip hoc gin tip nh nng lng nh sng mt tri. Nh vy
nng lng c tch lu l dng nng lng ho hc.
Phn ng tng qut ca qu trnh quang hp thng c biu din
bng phng trnh sau, mc d n th hin khng hon ton y .
6CO
2
+ 6H
2
O + Nng lng C
6
H
12
O
6
+ 6O
2

qu trnh quang hp nh nng lng nh sng mt tri m ng
c to thnh v quang hp thc vt gn lin vi vic gii phng
oxy.
Qu trnh quang hp c chia lm hai giai on:
- Phn ng sng
- S ng ho CO
2

Giai on th nht l qu trnh chuyn nng lng nh sng thnh
nng lng ho hc.
Giai on th hai l to ra dng nng lng ho hc bn vng v
d d tr hn.












67


Hnh 4.3 Cu trc ca dip lc a v b

nh sng cn cho qu trnh quang hp c tip nhn bi hai nhm
sc t: chlorophyll v carotenoid. mt s sinh vt bc thp hn cn c
phycobillin. Cu trc c bn ca dip lc l vng porphyrin, c to nn
t 4 vng pyrrol ring l (hnh 4.3). trung tm ca vng c mt nguyn
t Mg lin kt vi cc nguyn t nit bi hai lin kt ng ho tr v hai
lin kt phi tr. Nguyn t Mg c th c tch ra khi c tc ng ca
axit long. Phn t dip lc khng cha Mg c gi l pheophytin cng
c mt vai tr quan trng trong phn ng sng quang hp. Chui bn ca
phn t dip lc c tnh k nc v nh vy m ton b phn t c c
tnh k nc. Ngi ta bit nhng loi chlorophyll khc nhau: chlorophyll
a, chlorophyll b v chlorophyll vi khun. S khc nhau ca nhng loi ny
l do s khc nhau ca cc chui bn. chlorophyll b c nhm aldehyde,
nh hnh 4.3, c thay th bng nhm methyl chlorophyll a. iu th
v hn l chui bn di k nc c to nn t cc dn xut isopren.
68
chlorophyll a v b chui bn l mt phytol (ru) lin kt ester vi mt
chui bn ca vng porphyrine. mt s vi khun chlorophyll cha mt
farnesol (alkohol) c lin kt ester vi chui bn ca vng.
iu c ngha c bit hn l nhng lin kt i lin hp trong
vng porphyrine. Chng c kh nng hp thu nh sng v v vy nhng
lin i ny l yu t quan trng ca chlorophyll. Nhng lin kt lin hp
ca carotenoide cng hp thu nh sng. Cc cht thuc nhm carotenoide
c mu vng n mu da cam, thy r l cy vo ma thu, lc ny
dip lc b phn hu nn khng cn che ph sc t carotenoid.
hiu hn v qu trnh quang hp chng ta phi cp n c
quan m quang hp xy ra, l lc lp. Lc lp c hnh trn n
hnh bu dc, c ng knh khong 10 m. N c bao quanh bi
mng trong v mng ngoi, nhng mng ny c ngha i vi vic i
vo v i ra ca cc cht. Mng trong ca lc lp c to nn t h thng
mng ko dn, mt s v tr c dng xp chng ln nhau.














Hnh 4.4 S biu din mt lc lp

Nhng mng ny xut hin di knh hin vi in t nh nhng ti
dt. V vy ngi ta gi chng l thylacoid. Mng thylacoid bao quanh
mt khng gian bn trong gi l c cht ca lp th v to nn s ngn
cch gia bn trong thylacoid vi mi trng ngoi. S phn cch gia
bn trong thylacoid vi stroma l cn thit cho phn ng sng ca quang
Stroma
Bn trong
Thylakoid
Grana Thylakoid

Mng Thylakoid
Mng ngoi
Mng trong
Stroma - Thylakoid
69
hp. mt s v tr c nhiu thylacoid xp ln nhau thnh chng rt dy.
Chng xut hin di knh hin vi nh nhng ht (grana), v vy ngi ta
gi chng l grana-thylacoid. Chng c ni kt vi nhng thylacoid
ring l nh mng kp xuyn qua c cht lc lp. Chng c gi l
stroma-thylacoid (hnh 4.4).
Mng thylacoid l ni m nh sng c tip nhn bi cc sc t
m t trn v nh cc h thng oxy ho kh khc nhau m nng lng
nh sng mt tri c bin i thnh nng lng ho hc v thm thu.
Qu trnh ny phc tp v cha c gii thch chi tit.
Mng thylacoid cha ba cu thnh siu phn t, chng chim ton
b b rng ca mng v l thnh phn quan trng ca chui vn chuyn
in t trong quang hp. Ba phc h l h thng quang ho I, h thng
quang ho II v phc h cytochrome b/f (hnh 4.5).












Hnh 4.5 Cc phc h vn chuyn quan trng nht ca mng
thylacoid: H thng quang ho II (HTQHII), phc h cytochrome b/f, h
thng quang ho I (HTQHI)
H thng quang ho l v tr cha cc sc t hp th nh sng
(nhng anten). H thng quang ho I cha t nht l 13 loi protein khc
nhau, khong 200 phn t dip lc, mt s lng carotenoid cha bit v
ba phc h Fe-S. Nhng polypeptide kt hp v nh hng cc phn t
chlorophyll v carotenoid.
H thng quang ho II cha t nht 11 phn t polypeptide khc
nhau, khong 200 phn t dip lc a v 100 phn t dip lc b, hai phn t
pheophytin, quinon, 4 Mn cng nh Ca v chlorid. C hai h thng tip
nhn nh sng v chuyn ho chng trong s vn chuyn in t.
Chlorophyll
Carotenoide
Protein
Phc h Fe
2
-S
2
Chlorophyll
Carotenoide
Protein
Quinon
Phc h
Cyto-
chrome b,f
T bo cht
2NADP
+
+ 2H
+
2NADPH
4e
-
4e
-
Ferredoxin
Mng
Thylakoid
2H
2
O
4e
-
4H
+
+ O
2
HTQH II HTQH I
Plasto
cyanin
P682

P700

Pl.chinon
Pl.chinol
70
H thng quang ho II cn c nhim v l tch in t ra t H
2
O,
nh vy l oxy ho H
2
O. S oxy ho ny l qu trnh phc tp, gm 4
bc (mi bc 1e
-
c vn chuyn v y c s tham gia ca Mn vi
s thay i ho tr).
2H
2
O 4e
-
+ 4H
+
+ O
2
.
Qu trnh tch nc trong quang hp c gi l quang phn ly
nc, n gii phng ra O
2
v c trng cho quang hp sinh vt nhn
chun.
H thng quang ho I v II khng ch cha sc t tip nhn nh sng
m mi h thng cn cha mt phn t dip lc hon ton c bit th
hin v tr ca n, phn t dip lc ny khi tip nhn nh sng c kh
nng bn ra 1 in t. Qu trnh ny nh l s khi u cho s vn chuyn
in t trong quang hp. Phn t dip lc c bit ca h thng quang ho
I hp thu cc i nh sng c bc sng 700 nm. V vy gi l P700
(Pigment 700) v phn t dip lc c bit ca h thng quang ho II hp
thu cc i 682 nm.
bn ra 1 in t th phn t dip lc cn nng lng. Nng lng
c hp th dng nng lng nh sng t nhng sc t khc
(chlorophyll v carotenoid) ca mi h thng quang ho v c chuyn
n P682 cng nh P700 qua s cng hng in t. Bng cch ny hai
h thng quang ho to ra mt s vn chuyn in t khi c nh sng, s
vn chuyn in t ny i t H
2
O nh l ngun in t, qua h thng
quang ho II, plastochinone, phc h cytochrom b/f, plastocianin n h
thng quang ho I v cui cng n NADP
+
v n c kh thnh
NADPH.
NADP
+
+ H
+
+ 2e
-
NADPH
Trong chui kh ny h thng quang ho II v h thng quang ho I
khi ng k tip nhau. Plastochinone cng nh phc h cytochrome b/f
v plastocyanin c chc nng vn chuyn e
-
gia hai h thng quang ho
ny v chng nh v nhng v tr khc nhau. S vn chuyn e
-
lm
cho nng lng nh sng c bin i thnh nng lng ho hc, dng
kh NADPH (1 mol NADPH 220 kJ).
Thnh vin vn chuyn in t c bn trong chui vn chuyn l
phc cytochrome b/f, cng l protein c heme l nhm prostetic, v vn
chuyn in t l do s thay i ho tr ca Fe. Bn cnh phc
cytochrome b/f l plastocyanin, mt protein cha Cu (protein mu xanh)
vn chuyn in t gia phc cytochrome b/f v h thng quang ho I. S
vn chuyn ny c th l do s thay i ho tr ca Cu. H thng
71
plastochinon-plastochinol vn chuyn H t h thng quang ho II n
phc cytochrome b/f. Khong cch vn chuyn y ln hn. N khuch
tn trong vng ui ca lp mng kp lipid. Thnh phn mng cha
nhiu acid bo cha no th s khuch tn ca plastochinone d dng hn.
Sau y l phn t plastochinone v s kh n thnh plastochinol
(plastohydrochinone). c tnh k nc ca phn t l do cc nhm CH
3

v chui bn di vi 9 gc isopren. Phn ng kh vng chinone thc hin
qua s tip nhn ca 2e
-
v 2H
+
v vng quinone chuyn sang mt h
thng lin kt i ca bensoide.









H thng plastochinone/plastochinol vn chuyn H gia h thng
quang ho II v phc h cytochrome b/f. Phc h dng oxy ho-
plastochinon gn vo h thng quang ho II trc ht nhn 2e
-
v to
thnh semichinone (dng anion) v sau nhn 2H
+
ca h thng quang
ho II to thnh dng kh plastochinol. Sau n tch ra khi h thng
quang ho II v khuch tn n phc h cytochrome b/f. n y n b
oxy ho khi nhng 2e
-
v 2H
+
, to nn plastochinone v li khuch tn v
h thng quang II (hnh 4.5).
Chc nng c bn ca hai h thng quang ho l hp thu nh sng
v tp trung nng lng ca chng n P682 v P700, ri dn n qu
trnh quang ho, thc cht l bn ra 1e
-
. Chc nng ca 2 h thng tng
t nhau, tuy nhin nhng qu trnh chi tit hai h thng khc nhau, v v
chi tit cng cha c gii thch ht. S nhng 1 in t ca P682 lm
xut hin mt cht oxy ho rt mnh (phn t c kh nng oxy ho cht
khc) c th nng khong 1200 mV. P682 dng oxy ho (P
+
682) c kh
nng oxy ho H
2
O, ngha l tch e- ra khi H
2
O. Giai on quan trng
qu trnh ny l s tip nhn e
-
t P682 bn ra.


OH
OH
O
O
H
3
C
H
3
C
CH
2

CH
CH
3

C
CH
2
H
9
2e
-
2H
+
Plastochinone

Plastochinol

72










Ngy nay ngi ta cho rng, cht nhn in t bn ra l nhng
hp cht chinone. y chinone c kh thnh bn chinone
(semichinone mang in tch m). Semichinone l 1 radical, tip tc phn
ng nhanh, c th chuyn e
-
ca n n plastochinone. Dng semichinone
c kh to thnh plastochinone.
Pheophytin cng tham gia vo s vn chuyn e
-
. in t c bn ra
t P682 c th khng c tip nhn ngay, di tc dng ca nh sng
to nn rt nhanh l hng in t ca P682. Dng oxy ho ca P682
(P
+
682) tn ti rt ngn vi thi gian khong 10
-12
giy.
Nh c h thng quang ho II, di tc dng ca nh sng 1e
-
t
H
2
O c th nng oxy ho kh cao (th nng oxy ho kh tiu chun l
+800 mV) c chuyn n plastochinone c th oxy ho kh thp (th
kh tiu chun 100 mV). Nh vy kh nng kh ca e
-
c tng ln rt
nhiu.
Trong h thng quang ho I e
-
c bn ra t P700 c tip nhn
bi 1 phc h Fe-S. N chuyn e
-
ny tip tc n ferredoxin dng ho
tan nm trong stroma ca lp th. y cng thc hin mt phn ng
ngc chiu ca e
-
ngha l in t c chuyn n ni c in th m
ln hn, mt qu trnh cn nng lng, c thc hin nh nh sng mt
tri tip nhn c. Th kh tiu chun ca P700 nm khong +400 mV,
ca ferredoxin khong - 400 mV. Nng lng c nng ln r rt nh h
thng quang ho I. Ferredoxin kh c to nn l mt cht kh mnh, c
kh nng kh NADP
+
.



O
O
O
-
O
*

Semichinone
mang in m
(radical)
e
-
2x Ferredoxin Fe
II
2x Ferredoxin Fe
III
NADP
+
+ H
+
NADPH

73



hnh 4.6 a ra s c gi l s Z, ch ra hai s nng
nng lng c thc hin t h thng quang ho I v II trong chui vn
chuyn e ca quang hp. y e- ca H
2
O c chuyn n NADP
+

to nn NADPH l mt cht kh mnh cn cho nhiu qu trnh tng hp.












Hnh 4.6 S Z biu din s vn chuyn in t t in th dng n
in th m, s vn chuyn nng lng thc hin qua P682 v P700


Ferredoxin l mt cht kh mnh, c kh nng kh nhiu cht, c
ch ra hnh 4.7. Ngoi ra NADP
+
cn c ngha c bit trong vic kh
nitrite, sulfate v glutarate. Phn ng kh nitrite v sulfate l mt bc
quan trng trong s ng ho cc ion v c. V vy phn ng sng ca
quang hp khng ch l s ng ho CO
2
m cn ng ho c N v S.







Ferredoxin

NADP
+
-400

e
-
P700

Quang phosphoryl
ha vng
0

+400

+800

P682

H
2
O
e
-
Plasto-
chinone
e
-
Cyto-
chrome b,f
PC

e
-
e
-
Eo mV
74









Hnh 4.7 Nhng kh nng phn ng ca ferredoxin

Trong hnh 4.6 ch ra t ferredoxin e
-
khng nhng c th c
chuyn n NADP-reductase m cn tr v plastochinone. Nh vy to
nn mt dng in t khp kn. y in t li i qua cytochrome f v
plastocyanin tr v li P700 v t y li nh nng lng nh sng
c a n ferredoxin. Dng in t khp kn ny cng to nn s phn
tch proton v nh vy m tng hp ATP, dng in t khp kn ny c
gi l quang phosphoryl ho vng.
Quang phosphoryl ho vng ch sn xut ATP, quang phosphoryl
ho khng vng to nn ATP v NADPH. Bng cch ny qu trnh quang
hp c th t ph hp vi nhu cu ATP v NADPH. Qu trnh tng hp
tinh bt ch yu cn ATP, trong khi tng hp acid bo cn tng i
nhiu NADPH. ng ho CO
2
, cn ATP v NADPH theo t l 3:2.
Ferredoxin cng c th chuyn e
-
ca n n O
2
(hnh 4.7). Sau
H
2
O xut hin, v O
2
b kh lp tc phn ng vi H
+
trong mi trng
nc.
1/2O
2
+ 2e
-
O
2-

O
2-
+ 2H
+
H
2
O
Phn ng ny c ngha l tiu hao e
-
v trit tiu gradient H
+
. V vy
n lm quang phosphoryl ho tch ri s vn chuyn e
-
.
Tng kt chuyn ho nng lng quang hp ngi ta cho rng 4 mol
proton nh sng cn cho vic vn chuyn 2 mol e
-
v qu trnh quang
hp chui vn chuyn e
-
to nn 1 mol ATP v 1 mol NADPH.

Bng 4.1 Tng qut nng lng ca qu trnh quang hp
u vo:
4 Einstein (= mol proton) 710 x 10
3
J
Plastochinol

Nitrite

Sulfate

Glutarate

Cytochrome

O
2
e
-
Ferredoxin

HTQHI

NADP
+
e
-
e
-
e
-
e
-
e
-
e
-
e
-
75

u ra:
1 mol NADPH 220 x 10
3
J
1 mol ATP 32 x 10
3
J
Hiu sut: 35,5 % 252 x 10
3
J
Nh vy khong 35% nng lng nh sng hp thu c chuyn sang
dng nng lng ho hc.

4.4.2 Pha ti trong quang hp (chu trnh Calvin)
S kt hp CO
2
vo mt cht nhn hu c (s ng ho CO
2
) l mt
trong nhng qu trnh sinh hc quan trng nht. Nh qu trnh ny m
cacbon v c c a vo th gii sng. Phn ng c xc tc bi
enzyme ribulosediphosphate-carboxylase (RuDP-carboxylase), enzyme
ny c nhiu trong l xanh v n chim khong 30-50% protein ho tan
trong l. Enzyme gm 8 tiu n v ln v 8 tiu n v nh, trong ch
c cc tiu n v ln c hot tnh xc tc, cc tiu n v nh c chc
nng iu khin. Th v hn l thng tin di truyn ca cc tiu n v ln
nh v trn DNA ca lp th (DNA l cht mang gen).
xc tc enzyme cn nhng iu kin hon ton xc nh, l
gi tr pH (pH 8,3-8,6) v nng Mg
2+
tng i cao. C hai u b tc
ng bi s vn chuyn e
-
qua mng thylacoid. V vy enzyme ch hot
ng trong iu kin chiu sng. Phn ng ny gii phng nhit ( G
01
= -
40 x 10
3
J) v tng ng vi phng trnh sau:
RuDP + CO
2
+ H
2
O 2 phosphoglyceric acid.













H
2
C

O P

C=O

H C - OH

HC - OH

H
2
C - O P
H
2
C

O P

HC - OH

COO H

C O OH

HC - OH

H
2
C - O P
+ CO
2
+ H
2
O
RuDP Phosphoglycerate
76
Ngi ta cho rng carboxyl ho ribulosediphosphate (RuDP) dn n
to thnh mt hp cht trung gian c 6 nguyn t C vi enzyme, sau
c tch ra thnh hai phn t phosphoglyceric acid. Nh s trn CO
2

c tip nhn c mt trong phosphoglyceric acid pha trn v O t
phn t H
2
O c tip nhn th trong phosphoglyceric acid pha di.
Mt thi gian di ngi ta nh gi thp hot ng ca enzyme ny
v khi phn lp enzyme thng mt hot tnh. Kh khn na l CO
2
khng
nhng l c cht m cn nh hng trc tip n xc tc. Ngy nay ngi
ta bit hot tnh ca enzyme c c trng bi h s K
m
(CO
2
) t 10-
20 m l s ng ho CO
2
xy ra trong t nhin.
Sn phm carboxyl ho, phosphoglyceric acid c kh to thnh
aldehydphosphoglyceric. Qu trnh ny thu nng lng, cn ATP v
NADPH. C hai coenzyme tn ti stroma ca lc lp nh phn ng
sng. Khi khng c nh sng, nng ATP v NADPH thp th phn ng
xy ra ngc li, ngha l aldehydphosphoglyceric b oxy ho thnh
phosphoglyceric acid. Phn ng kh phosphoglyceric acid to ra sn phm
trung gian l 1,3 iphosphoglyceric acid. qu trnh ny bn cnh ATP
v NADPH cn c mt SH-enzyme-photphoglycerate-dehydrogenase
tham gia, c biu din s sau:



















COOH

HCOH

H
2
C-O - P
Aldehydphospho-
glyceric
O

CO~ P H S Enzyme

HCOH

H
2
C-O - P
ATP

ADP
O

CH

HCOH

H
2
C-O - P
O

C~ S Enzyme

HCOH

H
2
C-O - P
NADP
+
NADPH + H
+
HS-Enzyme
P OH (=P
i
)
77
Aldehydphosphoglyceric l ng u tin xut hin qu trnh
ng ho CO
2
. N d dng chuyn sang dng ng phn
phosphodioxyaketone (PDOA). Cn bng phn ng nghing mnh v pha
phosphodioxyaketone (PDOA) v khong 90% triosophosphate nm
dng PDOA.
c s ng ho CO
2
lin tc th cht nhn CO
2
, RuDP phi c
to nn mt nng nht nh. ng 3C c to ra l
aldehydphosphoglyceric v phosphodioxyaketone c bin i mt phn
thnh RuDP. Trnh t phn ng l mt qu trnh khp kn. S ng ho
CO
2
v bin i ca triosephosphate c gii thch bi Calvin v cng s
ca ng. V vy chu trnh ny c gi l chu trnh Calvin hoc l chu
trnh kh pentosophosphate v s bin i t phosphoglyceric acid thnh
aldehydphosphoglyceric l phn ng kh.







C ba loi phn ng c trng cho chu trnh Calvin:
- Phn ng phosphatase
- Phn ng aldolase
- Phn ng transcetolase
Phn ng phosphatase c cp chng enzyme. Nhng
enzyme aldolase c th gn kt mt aldose vi mt cetose, v d
aldehydphosphoglyceric v photphodioxyacetone. Phn ng ny ch c
mt s bin i nh v nng lng, y l phn ng thun nghch, n
c biu din nh sau:







CHO

HCOH

H
2
C-O - P
Aldehydphospho-
glyceric (aldose)
H
2
C-OH

C=O

H
2
C-O - P
Phosphodioxy-
aceton (cetose)
78












phn ng transcetolase 2C ca cetose c chuyn ln mt aldose
(v d aldehydphosphoglyceric).
Bng cch tng t 2C ca fructosemonophosphate c chuyn ln
aldehydphosphoglyceric, to thnh mt pentosephosphate v mt
tetrosephosphate.
Chu trnh Calvin bt u vi mt phn ng aldolase (hnh 4.8). Khi
tn ti nhiu triosephosphate th aldehydphosphoglyceric phn ng vi
phosphodioxyacetone to nn fructose1,6 diphosphate.












Bng cch ny ng 6C u tin c to nn. T fructose1,6
diphosphate mt phosphate c tch ra nh enzyme phosphatase, lm
xut hin fructosemonophotphate. Fructosemonophosphate cng vi mt
CHO

HCOH

H
2
C-O - P
Phosphodioxy aceton (cetose) + Aldehydphosphoglyceric (aldose)
H
2
C-O P

C=O

HOCH
2
Fructose 1,6 diphosphate
HCOH

HCOH

H
2
C-O - P
H
2
C-O P

C=O

HOCH
CH
2
- OH

C=O

HOCH

HCOH

HCOH

CH
2
O P
Fructosemono-
phosphate
CHO

HCOH

H
2
C-O - P
Aldehydphospho-
glyceric
H
2
C-OH

C=O

HOCH

HCOH

H
2
C-O P
Pentose-
phosphate
CHO

HCOH

HCOH

H
2
C O P
Tetrose-
phosphate
Transcetolase
79
aldehydphosphoglyceric khc c bin i thnh mt ng c 4C
(erythrosephosphate) v mt ng c 5C (xylulosephosphate) bng phn
ng transcetolase. Erythrosephosphate phn ng vi mt triosephosphate
bng phn ng aldolase to thnh mt ng c 7C
(sedoheptulosediphosphate). T y mt phosphate c tch ra nh
enzyme phosphatase to nn sedoheptulosephosphate.
Sedoheptulosephosphate phn ng vi mt triosephosphate khc nh
enzyme transcetolase to thnh hai ng c 5C l xylulosephosphate
v ribosephosphate.
Trong phn ng tng qut th cn 5 ng triosephosphate to
nn 3 ng pentosephosphate, trong 1 ribosephosphate v 2
xylulosephosphate. Ba ng pentosephosphate c bin i sang dng
ng phn ca n l ribulosephosphate, ng ny c photphoryl ho
nh ATP to thnh ribulosediphosphate. Nh vy m cht nhn CO
2

c ti to.
Ba phn t ribulosodiphosphate c th kt hp vi 3 phn t CO
2
v
nh vy s to nn 6 phn t triosephosphate. y 5 phn t
triosephosphate cn ti to nn 3 phn t ribulosediphosphate. Mt
phn t triosephosphate cn li cho mc ch tng hp. Tng ng c
phng trnh nh sau:

3CO
2
+ 9 ATP + 6NADPH 1 triosephosphate + 9 ADP + 6NADP
+


T phng trnh trn ta thy kh mt phn t phosphoglyceric
acid cn 1 ATP v 1 NADPH. to nn 1 phn t ribulosediphosphate
t ribulosephosphate th cn thm mt phn t ATP. Chu trnh tng qut
cn 3 ATP v 2 phn t NADPH cho c nh 1 phn t CO
2
. Vi qu trnh
quang phosphoryl ho vng th h thng s to c nhiu phn t ATP
hn NADPH, ngha l t l yu cu theo l thuyt ATP/ NADPH ph hp
l 3/2.







80











Hnh 4.8 Chu trnh Calvin, PGA: phosphoglyceric acid,
PDOA: phosphodioxyacetone, FMP: fructosemonophosphate,
FDP: fructosediphosphate

4.4.3 H hp sng- Chu trnh Glycolate

Cy xanh khi c nh sng cn tip nhn O
2
v gii phng CO
2
. H
hp ph ny c gi l h hp sng. S tip nhn O
2
c th do nhiu qu
trnh v d ferredoxin tham gia vo quang hp chuyn e
-
ca n trc tip
n O
2
, nh vy H
2
O xut hin (hnh 4.7).
Warburg ch ra rng O
2
c ch s ng ho CO
2
. Enzyme xc tc
cho ng ho CO
2
l ribulosediphosphate-carboxylase th hin khng
nhng hot tnh carboxylase m cn hot tnh oxygenase. S th hin hot
tnh no l ph thuc vo nng CO
2
v nng O
2
. CO
2
c ch
oxygenase, O
2
c ch carboxylase. Carboxyl ho v oxy ho c cng trung
tm hot ho, l gc lysine. Nhng iu kin nh l Mg
2+
kch thch
carboxyl ho v gi tr pH cao kch thch s oxy ho.








NADP
+

+ P
i
NADPH + H
+
CO
2
+ H
2
O
Diphospho
glyceric acid
Triose-
phosphate
APG
AlPG
AlPG
AlPG
PDOA
ATP
PDOA
ADP
H
2
O
H
2
O
FMN
P
i
P
i
RuDP

1,6 FDP

Ribulose-
monophosphate

Sedoheptulose-
diphosphate

Erythroso-
phosphate

Xilulosophosphate

Sedoheptulose-
monophosphate

Xilulosephosphate

Ribosephosphate

1
2
3
4
5
81






Hnh 4.9 S cnh tranh gia CO
2
v O
2
vo cng mt v tr ca enzyme
ribulosephosphate-carboxylase-oxygenase

Cui cng th c hai phn ng c cng c cht, l
ribulosediphosphate. S cnh tranh gia O
2
v CO
2
vi phn t enzyme
c minh ha hnh 4.9.
S cnh tranh ca CO
2
v O
2
cng mt v tr ca enzyme to ra s
lin quan vi qu trnh quang hp. Khi cng quang hp cao th nng
O
2
trong t bo mesophyll tng ln do qu trnh quang phn ly nc,
trong khi s ng ho CO
2
th nng CO
2
gim. Nhng iu kin ny
thun li cho phn ng oxy ho. Nh vy nhng iu kin thun li cho
quang hp nh nhit thch hp, cng nh sng mnh kch thch s
oxy ho ribulosediphosphate. Phn ng c biu din nh sau:













y phn t O
2
c tch ra lm 2 nguyn t: mt nguyn t gn
vi gc 2C (phosphoglycolate), nguyn t oxy khc gn vi gc c 3C.
Phosphoglycolic acid l dn xut ca acetic acid. Bng phn ng
phosphatase t phosphoglycolic acid, glycolic acid s c to thnh.
CO
2
RuDP

O
2
RuDP
Carboxylase
Oxygenase

H
2
C

O P

C=O

H C - OH

HC - OH

H
2
C - O P
RuDP
COOH

HC - OH

H
2
C - O P


H
2
C - O P

COOH



H
2
C - OH

COOH


O
2
Oxygenase
Phosphoglycolate
H
2
O
P OH
Glycolate
Phosphoglycerate
82
S oxy ho ribulosediphosphate to nn chu trnh glyoxylate, y l
nhng phn ng quan trng ca quang h hp. Phn ng to thnh
phosphoglycolate v phn ng kh phosphoryl ho ca n c xc tc
bi enzyme phosphatase xy ra trong c cht ca lc lp. Hot tnh ca
glycolate-phosphatase c kch thch bi nng Mg
2+
v pH cao v
y l nhng iu kin thun li cho quang hp.
Glycolate t lc lp i n peroxisome l vi c quan t nm st cnh
lc lp, c bao bc bi mng v cha cc enzyme khc nhau nh
transaminase, oxidase v catalase.







Trong peroxisome glycolate b oxy ho to thnh glyoxilic acid
nh mt oxidase. H
2
O
2
xut hin c phn gii thnh H
2
O + 1/2 O
2
v
glyoxilic acid b bin i thnh mt amino acid, l glycine nh mt
transaminase. Transaminase s c cp k hn chng proteine,
y ch nu ln nguyn l ca n l nhm amin ca mt amino acid c
chuyn ln mt nhm ceto hoc nhm aldehyd. Amino acid tr thnh
cetoacid.












H
2
C - OH

COOH



CHO

COOH


Glycolic acid
O
2

H
2
O
2
Glyoxylic acid

CHO

COOH


Aminoacid
Glyoxylic acid

COOH

H
2
C NH
2





COOH

C=O

R



COOH

H
2
N -C H

R


Cetoacid
Glycine
83
Glycine i t peroxisome vo ty th, l c quan t m trong h
hp xy ra. Glycine b oxy ho di tc dng ca NAD
+
v sau b kh
amin ho (gii phng NH
3
), xut hin li glyoxylic acid.







NADH to ra phn ng kh v l c cht cho chui enzyme h
hp. Glyoxilic acid b kh cacboxyl ho v phn ng vi gc CH-OH t
do.










Tetrahydrofolic acid (THFA) kt hp vi mt gc hydroxylmethyl,
gc ny c chuyn n mt glycine, to nn aminoacid l serin v
THFA li tr v trng thi ban u ca n.









CHO

COOH

Glyoxylic acid

COOH

HC = NH

Iminoacid

COOH

H
2
C - NH
2



Glycine
NAD
+
NADH + H
+
H
2
O
NH
3
Hydroxymethyl

COO H

CHO = CH - OH
CO
2
THFA

H

THFA

H
2
C - OH

COOH

HC NH
2

H


Glycine

THFA

H
2
C - OH

THFA

H

COOH

HC NH
2

H
2
C - OH


Serine
+ +
84
Serine b kh amin ho to nn hydroxylpyruvate v cht ny c
th c bin i thnh phosphoglyceric acid bng phn ng kh v phn
ng phosphoryl ha.
















Phn ng cn 1 NADH. Phosphoglyceric acid l mt cht trao i
ca chu trnh Calvin. Cht ny li i vo lc lp v n c s dng
to nn triosephosphat. Nh hnh 4.10 ch ra, chu trnh glyoxylate xy
ra trong cc c quan t khc nhau. Tng qut l c hai phn t glycolate
c s dng tng hp nn mt phn t phosphoglyceric acid.












COOH

HC NH
2

H
2
C - OH


Serine
COOH

C = O

H
2
C - OH
Pyruvic acid
COOH

HC - OH

H
2
C - O P

Phosphoglyceric acid
COOH

HC - OH

H
2
C - OH
Cetoacid
Aminoacid
ADP
ATP
Kinase
NADH + H
+
NAD
+
Hydroxypyruvic acid
Lc lp
Ty th
2x Glycolate

P-Glycerate
2x Glyoxylate

2x Glycine
Peroxisome
O
2
O
2
CO
2
NH
3
85
Hnh 4.10 Nhng cht trao i quan trng ca chu trnh glycolate cc c
quan t khc nhau
Chu trnh glycolate c mi lin h cht ch vi chu trnh Calvin qua
ribulosediphosphate-carboxylase-oxygenase. Tng qut ca chu trnh
glyolate l 2 v tr O
2
c tip nhn (phn ng oxygenase v phn ng
oxidase) v CO
2
v NH
3
c gii phng ra. Nh vy chu trnh glycolate
phn gii C v N hu c. ngha sinh l ca s phn gii ny vn cha
c gii thch r rng.
Quang h hp b nh hng bi nhiu yu t ngoi cnh. Nhng yu
t kch thch quang hp nh cng chiu sng cao, nhit thch hp
lm tiu hao nhiu CO
2
trong m l v sn sinh nhiu O
2
do quang phn ly
nc lm cho h hp sng xy ra mnh m. CO
2
v NH
3
gii phng ra
trong h hp sng li c ng ho. c bit l NH
3
.
Cht km hm t nhin ca chu trnh glyolate l glutamate, aspactate
v glycolate. nhng thc vt C4 h hp sng khng c ngha. y
c th do c ch ng ho CO
2
c bit m nng CO
2
tng i cao
trong lc lp, v vy m hot tnh oxygenase ca ribulosediphosphat-
carboxylase-oxygenase khng th hin.

4.4.4. Chu trnh Hatch v Slack

Ribulosediphosphate khng phi l cht nhn CO
2
duy nht cho
phn ng u tin ca s ng ho CO
2
. mt s loi thc vt, c bit
loi cy h c nhit i (ng, cao lng, ma) cht nhn CO
2
l
phosphoenolpyruvate (PEP). Qu trnh bt u bng phn ng kt hp
HCO
3
-
vo PEP. Bng cch ny nh enzyme PEP-carboxylase, mt hp
cht 4C- oxaloaxetic acid, c to thnh.










PEP
COOH

C O P

CH
2
COO
-


CH
2


C = O + P OH

COOH
Oxaloacetate
O

+ HO C O
-

86

Nh ch ra s , HCO
3
-
c tch ra thnh 1 gc hydroxyl v
1 gc carboxyl. Gc hydroxyl phn ng vi gc phosphoryl ca PEP
to nn phosphate v c (H
3
PO
4
). Gc carboxyl kt hp vi cht c 3C
(PEP) to nn cht c 4C (oxaloaxetic acid).





















Hnh 4.11 S ng ho CO
2
theo chu trnh C
4
nh enzyme PEP-carboxylase

Phn ng ny gii phng nng lng, v lin kt cao nng trong PEP
c phn gii. Oxaloaxetic acid c to nn (anion oxalacetate) sau
tip tc b kh to thnh malate hoc b amin ho to thnh
aspartate. V nhng phn ng ny cha nhng hp cht c 4C, nn s
ng ho CO
2
nh PEP-carboxylase c gi l chu trnh C
4
. Nhng loi
thc vt m ng ho CO
2
u tin bng con ng ny, c gi l thc
vt C
4
ngc vi thc vt C
3
, l nhng cy ng ho trc tip CO
2
qua
PEP
COOH

C O P

CH
2

*
COO
-


CH
2


C = O

COOH
Oxaloacetate
COOH

H
2
N-CH

CH
2


COO
-

Aspartate
Aminoacid

COOH

H
2
N -C H

R



COOH

C=O

R


Cetoacid
H
+
CO
3
-
Pi

1

3

2

NADPH
+ H
+
NADP
+
COOH

C = O

CH
3

Pyruvate

*
COO H

CH
2


HCOH

COO
-

Malate
NADP
+
NADPH + H
+
+

CO
2
ATP + Pi

P ~ P
+ AMP

4

5

Chu trnh Calvin
87
chu trnh Calvin. Trnh t phn ng ca chu trnh C
4
c ch ra hnh
4.11.
Phn ng 1: HCO
3
-
c kt hp vi PEP to thnh oxalacetate.
Phn ng ny c xc tc bi enzyme PEP-carboxylase.
Phn ng 2: Oxalacetate b kh to thnh malate nh NADPH
Phn ng 3: Oxalacetate c bin i thnh aspartate bng phn
ng chuyn amin ho.
Phn ng 4: Nh cc malatenzyme m malate b kh carboxyl ho
v oxy ho to nn pyruvate. CO
2
c gii phng ra do kh cacboxyl
ho c a n chu trnh Calvin.
Phn ng 5: Pyruvate c bin i tr li thnh PEP bng phn
ng kh carboxyl ho. Phn ng cn ATP v photphate v c.
ATP c tch ra thnh AMP v pyrophosphate. Phn ng ny c
mt gi tr m G
01
cao, nn phn ng xy ra nhanh nn cht nhn CO
2
l
PEP c to ra nhiu. Enzyme xc tc gi l pyruvate-phosphat-
dikinase, gn vo enolpyruvate theo phn ng di y:








Trao i CO
2
ca thc vt C
4
c hiu khi ngi ta chia ra nhng
qu trnh ring l nhng m l khc nhau v nhn bit nhng v tr ca
t bo. Phn ln nhng thc vt C
4
c kiu gii phu Kranz c trng, mt
khi nim c Haberlandt (1904) ngh. y l nhng t bo xanh
bao quanh b mch dng xon kp xp st xung quanh nhng t bo b
mch, trong khi nhng t bo mesophyll nh hn sp xp lng lo hn
(hnh 4.12).
COOH

C - OH

CH
2
Enolpyruvate PEP
COOH

C O P

CH
2
ATP + P
i
AMP + P ~ P

88

Hnh 4.12 S sp xp ca cc t bo tht l v cc t bo b mch cu trc
Kranz ca thc vt C
4
1. Biu b trn, 2. T bo m giu cha lc lp, 3. Mch dn,
4. Khoang trng gian bo, 5. Biu b di vi kh khng.

Trong nhng t bo mesophyll CO
2
c ng ho nh PEP-
carboxylase v oxaloacetate c to ra b bin i thnh malate hoc
aspartate. Nhng t bo mesophyll ca thc vt C
4
khng cha RuDP-
carboxylase. Sn phm ng ho l malate v aspartate, loi ny l
malate v loi kia l aspartate, c vn chuyn qua gian bo n cc t
bo bao quanh b mch. y chng c kh carboxyl ho. Pyruvate
xut hin li tr v t bo mesophyll. Qu trnh ny xy ra hai loi t bo
v c ch ra hnh 4.13.










89












Hnh 4.13 S tng hp PEP, phn ng carboxyl ho, phn ng kh
oxaloacetate t bo tht l, s kh carboxyl ho v chu trnh Calvin xy ra
t bo b mch
loi cy malate pyruvate c to nn trc tip t malate bng
cch kh carboxyl ho oxy ho.









phn ng do malat-enzyme xc tc bn cnh CO
2
cn to nn
NADPH, l cht cn trong chu trnh Calvin. Vi malate khng nhng cc
nguyn t C m cn c cht kh c vn chuyn n chu trnh Calvin.
loi cy aspartate aspartate c kh carboxyl ho to thnh alanine.
Alanine c bin i thnh pyruvate nh phn ng chuyn amin ho.
Phn ng chuyn amin ho xy ra trong t bo cht ca t bo tht l.
Alanine c vn chuyn t t bo bao quanh b mch tr v t bo tht
l.


COOH

C = O

CH
3

Pyruvate
+ CO
2
COO H

CH
2


H C O H

COO H
Malate
NADP
+
NADPH + H
+
T bo bao quanh
b mch
T bo tht l
Malate
NADP
+
NADPH + H
+
ATP + Pi
AMP + Pyrophosphate
Oxaloacetate
NADPH
+ H
+
NADP
+
Pyruvate
Chu trnh
Calvin
CO
2

PEP
HCO
3
-
90












S phn chia cc phn ng ring l t bo cht v lc lp ca t
bo tht l tng ng vi t l nng lng. S phn chia ny c gii
thch trong hnh 4.14.
Pyruvate c a vo t bo tht l phi chui vo lc lp ca t bo
ny, v y c mt ca enzyme pyruvat-phosphate-dikinase v ATP,
ATP ny c to ra t quang hp. PEP to nn trong lc lp c vn
chuyn ra t bo cht bng s trao i vi phosphat v c. y PEP
cng vi CO
2
l c cht ca enzyme PEP-carboxylase. Oxalacetat c
to ra i vo lc lp ri b kh thnh malate nh NADPH. ATP v
NADPH c to ra trong lc lp l nh qu trnh quang hp.













Hnh 4.14 S vn chuyn nng lng v carbon gia lc lp v t bo cht
ca t bo tht l thc vt C
4

COOH

H
2
N-CH

CH
2


COO H
COOH

H
2
N-CH

CH
3

COOH

C = O

CH
3

Aspartate Alanine
Cetoacid
Aminoacid
Pyruvate
CO
2
Pyruvate
T bo
bao
quanh b
mch
T bo tht l
Malate
NADP
+
NADPH + H
+
Lc lp
ATP + Pi
Oxaloacetate
Pi
PEP
HCO
3
-
AMP ~ P ~ P
91

Malate sau c vn chuyn n t bo bao quanh b mch. T
bo tht l v t bo bao quanh b mch cha nhng lc lp c hnh thi
khc nhau: Lc lp ca t bo bao quanh b mch ln hn v khng c
grana v tch lu tinh bt. Lc lp ca t bo tht l nh hn v lun lun
cha grana.
u im ca s ng ho CO
2
thc vt C
4
l enzyme PEP-
carboxylase c trong t bo cht tip nhn CO
2
c trong m l v CO
2
t
qu trnh h hp tt hn s tip nhn CO
2
thc vt C
3
, v thc vt C
3

CO
2
trc ht phi chui vo lc lp. Tng ng im b CO
2
ca thc vt
C
4
c bn l thp hn thc vt C
3
, nm trong khong 0,5-1 l/LCO
2
,
im b CO
2
ca thc vt C
3
nm trong khong 3-5 l/LCO
2
. im b
quang hp l nng CO
2
m khng c s tch lu cht hu c.
Thc vt C
4
c kh nng gim nng CO
2
trong m l mnh hn thc
vt C
3
. V vy chng c kh nng s dng CO
2
trong khng kh c hiu
qu hn.
Hn na nng CO
2
cao c to ra trong t bo bao quanh b
mch nh qu trnh kh cacboxyl ho. Nh vy m im bo ho CO
2
ca
RuDP-carboxylase gn t c v hot tnh oxygenase ca enzyme b c
ch. V vy thc vt C
4
c kh nng s dng tt nh sng c cng
cao. V l do ny m chng thc t khng c im bo ho nh sng.
c im ny c th hin trn hnh 4.15. y ng cong ng ho
CO
2
ca hai cy trng C
4
l ma v ng tng cao hn ng cong ca thc
vt C
3
.















100 200 300 400
Cng nh sng Jm
-2
s
-1



50-
40-
30-
20-
10-
0
Ma
Ng
La m
C ci ng
Thuc l
C ci nc
M
g

C
O
2
-
1
0
0
c
m
-
2
.
h
-
1

92
Hnh 4.15 nh hng ca cng nh sng n s ng ho CO
2
mt s
cy trng
Tuy nhin thc vt C
4
cn nhiu nng lng cho s ng ho CO
2
,
v chu trnh C
4
khi ng trc chu trnh Calvin cn cho s kt hp 1 phn
t CO
2
hai phn ng phosphoryl ho (to nn hai lin kt cao nng)
tng hp nn ATP t AMP theo phng trnh sau:

AMP + ATP 2ADP
2 ADP + 2 phosphate 2ATP (quang phosphoryl ho)
-------------------------------------------------------------------
Tng qut: AMP + 2 phosphate 2ATP

Phn ln nhng thc vt C
4
c ngun gc t nhng vng c cng
nh sng mnh, v vy ATP c to nn nhiu nh quang phosphoryl
ho. i vi s ng ho CO
2
th cng nh sng l yu t t hn ch
hn so vi nng CO
2
. iu cng d hiu l trong qu trnh tin ho
mt s loi thc vt pht trin c ch ng ho CO
2
trong nhng iu kin
thi tit c bit. Do s thch nghi ny m enzyme pyruvate-phosphate-
dikinase mn cm vi nhit thp. V d iu kin kh hu Trung chu
u ma xun th hin s ngng pht trin v vng l ng, trong khi cc
loi ng cc khc (la m, la mch, yn mch, kiu mch) pht trin tt,
l do s mn cm ca enzyme pyruvate-phosphate-dikinase vi nhit
thp. Khi thi tit m ln th ng bt u pht trin vi tc nhanh, vt
nhiu so vi thc vt C
3
. Tc pht trin nhanh l do s ng ho CO
2

c hiu qu, v do thc vt C
4
s mt mt sn phm ng ho do h hp
sng thp hn thc vt C
3
.
Cu to gii phu kiu Kranz ca t bo l cng c mt ngha i
vi trao i nc. Nhng t bo xp st b mch c s trao i kh thp
hn v nh vy s mt nc do thot hi nc cng t hn. iu ny gii
thch ti sao thc vt C
4
c nhu cu nc tng i thp hn.

4.4.5 Quang hp theo chu trnh CAM (Crassulaceae Acid Metabolism)

Cc loi thc vt khc nhau ca h Crassulaceae v nhng i din
ca Agavaceae, Bromeliaceae, Cactaceae, Euphobiaceae, Liliaceae v
Orchideaceae c kh nng tip nhn CO
2
vo ban m nh kh khng m
v ban ngy th kh khng ng li trnh s mt nc. Tri vi a s
cc loi thc vt khc, kh khng ng li ban m v m ban ngy, kh
nng ny gn lin vi s trao i acid v CO
2
c bit, trc ht c
93
nghin cu loi Crassulaceae. Nhng thc vt c hnh thc trao i ny
c gi thc vt CAM.















Hnh 4.16 S ng ho CO
2
v trao i acid thc vt CAM

Nguyn l ca s trao i cht c bit ny c minh ha trong
hnh 4.16. Ban m CO
2
c tip nhn nh kh khng m, enzyme PEP-
carboxylase carboxyl ho PEP to thnh oxaloaxetate. Cht ny c
kh to thnh malate. Sau malate c d tr trong khng bo.
y nng malate c th ln n 100-200 mM, c bit nng ny cao
trong nhng gi u bui sng. Ban ngy khi kh khng ng, malate
trong khng bo c huy ng vo lc lp v y n c kh
carboxyl ho. CO
2
t do v NADPH to ra i vo chu trnh Calvin. Bn
cnh s ng ho CO
2
c cung cp bi ATP v NADPH c to ra
t quang hp. Sn phm ng ho ny mt phn c bin i thnh tinh
bt, tch lu trong lc lp. Ban m mt phn tinh bt ny c phn gii
to thnh PEP, l cht tip nhn CO
2
trong t bo cht. s kh
carboxyl ho oxy ho pyruvate c to thnh t malate, sau pyruvate
c bin i thnh PEP nh enzyme pyruvate-phosphate-kinase. Bn
cnh malate v oxaloaxetate th aspartate, glutamate, alanine, glycine bng
phn ng chuyn amin ho to nn nhng sn phm trung gian gn lin
vi s trao i cht ca thc vt CAM.
CO
2
(m)
Oxaloacetate
(m)
Malate
(m)
Malate
(ngy)
Khng bo
(m)
(m)
PEP
Tinh bt
Chu trnh Calvin
Lc lp
CO
2
+ NADPH
Pyruvate
(ngy)
94
S tch lu malate trong khng bo l mt qu trnh ch ng, c
iu khin bi sc trng ca khng bo. Th nc ca cy iu khin s
ng m kh khng. Ngi ta phn bit nhng thc vt CAM bt buc
v khng bt buc. Thc vt CAM khng bt buc ch thc hin theo
chu trnh CAM khi thiu nc, c bit l trong iu kin ngy di.
Kh khng ng ban ngy th s mt nc gim rt ng k, nh vy
m thc vt c kh nng vt qua c thi k kh hn. mt th nc
trong t l 2200 kPa, vt xa im ho ca cc cy trng khc, nhng
trong cc m ca loi cy ny vn duy tr c th nc t 500-1000 kPa.

4.4.6 Tng hp carbohydrate

Vi mt s phn ng tng i n gin thc vt c kh nng tng
hp cc hexosephosphate khc nhau, l nhng nguyn liu cn thit cho
tng hp oligo- v polysaccharide.
Nguyn l sinh tng hp l to nn lin kt glycoside trc ht c
gii thch bi Leloir. Nguyn l ny c ch ra v d tng hp glucan
(polysaccharide ca glucose) (hnh 4.17). phn ng ny c hai enzyme
tham gia: ADP-glucopyrophosphorylase v glucan- hoc ADP-gluco- -
glucan-4 -glucosyltransferase. Enzyme ADP- glucopyrophosphorylase
hot ho glucose1-phosphate nh ATP, vi s tch ra pyrophosphate xut
hin dng glucose hot ho, l ADP-glucose. phn ng ny nguyn
t H ca nhm phosphoryl c thay th bng nhm acyl ca
adenosinemonophosphate. Acyl tng ng gi l adenyl. Sau dng
glucosyl hot ho c chuyn ln chui glucan v thay th nguyn t H
ca nhm OH v tr C
4
(xem cng thc). Nh vy xut hin mt lin kt
glycoside gia C
1
ca glucosyl mi i vo v C
4
ca glucosyl chui
glucan. Lin kt nm pha di ca mt phng glucose v l lin kt
glycosidic. Theo nguyn tc ny mt chui di ca cc gc glucosyl c
to thnh.
Chui ko di theo hng C
1
C
4
. S tng hp polysaccharide
phn ln bt u bng mt phn t mi (primer), y glycosyl u tin
c nh vo. s tng hp tinh bt v glycogen th primer l 1
glycoprotein. Glucosyl l gc ng ca glucose, fructosyl ca fructose,
ribosyl ca ribose.



95
















Hnh 4.17 Tng hp ADP-glucose nh enzyme ADP-glucose-
pyrophosphorylase v s chuyn glucosyl ln mt chui glucan

Bn cnh tinh bt, saccharose l sn phm quan trng khc ca qu
trnh quang hp. Triosephosphate c to nn t chu trnh Calvin i t
lc lp ra t bo cht, bin i thnh UDP-Glucose, Fructose 6-P v sau
thnh saccharose 6-P nh enzyme saccharose-P- synthase. Cui cng
phosphate c tch ra bng enzyme saccharose-P-phosphatase.
iu khin enzyme saccharose-synthase thc hin bc fructose-
1,6-disphosphate bi nng ca fructose 2,6-P
2
(xem gluconeogenese).
Mt v tr iu khin khc l saccharose-P-synthase c hot ho bi c
cht glucose 1-P v b c ch bng phosphate v c. Ngoi ra hot tnh
ca enzyme gim xung do qu trnh phosphoryl ho v tng ln do s
kh phosphoryl ho (tng t nh glycogen-synthase). Nng
saccharose li iu khin s tng hp tinh bt.




O
CH
2
OH
O
O

PO H

OH
O O O

HOP~OP~O~PO Aden

OH OH OH
O O

HOPO~POH

OH OH
Pyrophosphorylase
+
CH
2
OH
O
O
O O

PO~PO Aden

OH OH
CH
2
OH
O
O
CH
2
OH
O
O
CH
2
OH
O
O
CH
2
OH
O
H O
S tng hp glucan
96





















Hnh 4.18 S tng hp saccharosephosphate v saccharose

Saccharose va l mt dng carbonhydrate vn chuyn va l tin
cht ca tinh bt (trong t bo, nm cch ra khi v tr quang hp) v ca
cellulose. Enzyme saccharose-synthase c tc dng thun nghch to nn
nucleosid - ng cn thit.
Saccharose + UDP = UDP-D-Glucose + Fructose







CH
2
OH
OH
HO
HO
O
HO-CH
2

OH
HO
O
O
CH
2
OH
CH
2
OH
P O-CH
2

OH
HO
O
O H
Saccharose
Saccharosephosphate
O
H
2
O
P
i
CH
2
OH
OH
HO
OH
UDP
O
CH
2
OH
OH
HO
OH
O
P O-CH
2

OH
HO
O
UDP
97























Tng t ATP l coenzyme ca ADP-pyrophotphorylase qu trnh
tng hp tinh bt, UTP cung cp nng lng cho s tng hp saccharose,
glycogen v cellulose. y UDP-pyrophosphorylase xc tc tng hp
UDP-glucose t glucose1-phosphate v UTP. s tng hp saccharose
mt glycosyl c vn chuyn t UDP-glucose n 1 fructosephosphate
to nn saccharosephosphate, t cht ny saccharose v phosphate v
c c tch ra. S tng hp saccharose sau khi ng ho CO
2
cng nh
s to nn saccharose t tinh bt trong ht ny mm c thc hin theo
con ng ny (hnh 4.18). Enzyme xc tc cho phn ng ny l
saccharosephosphat synthetase. Hng phn ng dch v pha
saccharosephosphate cng nh saccharose, v s to ra phosphate nh
phosphatase, l mt qu trnh thun li v nhit ng hc. Khng nn
nhm ln gia enzym saccharosephosphatsynthetase vi enzyme
saccharosesynthetase. Enzyme saccharosesynthetase ch tng hp
saccharose trong nhng trng hp ngoi l. Chc nng quan trng ca n
O O

H OP~OP~O

OH OH
CH
2

OH OH
O
N
N
O
OH
UDP Saccharose
HOCH
2

OH
HO
OH
O
CH
2
OH
CH
2
OH
OH
HO
OH
O
O
O O

P~OP~O

OH OH
CH
2

OH OH
O
N
N
O
OH
Fructose UDP-glucose
+
CH
2
OH
OH
HO
OH
O
CH
2

OH
HO
CH
2
OH
O
O
98
l to nn UDP-glucose t saccharose v UDP theo phng trnh phn
ng sau:
Saccharose + UDP UDP-glucose + fructose
Cn bng phng trnh ny dch v pha to UDP-glucose. Enzyme
ny l saccharose:UDP-glycosyltransferase, ng vai tr quan trng
trong m sinh trng v trong nhng c quan d tr ang ln (c, ht) m
saccharose c chuyn n tng hp UDP-glucose. Sau UDP-
glucose c s dng ch yu cho vic tng hp cc cht ca thnh t
bo, nh cellulose.
C ch ca phn ng ca saccharose:UDP-glycosyltransferase c
th hin s trn.
Nhng polysaccharide m t trn c cha nhiu nhm OH, nn
phn t c c tnh a nc v c kh nng to nn cc cu hydro. Cc
nhm acid, carboxyl, nhm sulfate lm cho phn t c tnh acid. Trong
iu kin pH sinh l (pH 5,5-7,5) cc proton c tch ra, v vy cc i
phn t mang nhiu nhm anion, nn tch in m v c kh nng kt hp
vi cc cht tch in dng, v d nhm amin b proton ho to lin
kt ion.

4.5 Ho sinh h hp
4.5.1 nh ngha h hp
H hp l qu trnh phn gii, oxy ho cc hp cht hu c phc tp
thnh cc sn phm cui cng l CO
2
v H
2
O, ng thi gii phng nng
lng cn thit cho c th sng.
Phng trnh tng qut ca h hp:
C
6
H
12
O
6
+ 6O
2
6CO
2
+ 6H
2
O + Nng lng
V bn cht h hp l qu trnh oxy ho sinh hc c thc hin
bng s tch cc nguyn t hydro ra khi c cht v vn chuyn n (H
+
v
e
-
) n nhng cht nhn khc nhau, cht nhn cui cng l O
2
. H hp xy
ra hai v tr khc nhau trong t bo. Giai on th nht xy ra t bo
cht (qu trnh ng phn), hoc bn trong ty th (chu trnh Krebs), l s
tch hydro ra khi c cht v chuyn n n NAD v/hoc FAD to
nn NADH
2
v FADH
2
. giai on th hai NADH
2
v FADH
2
b oxy
ho, l qu trnh chuyn H
+
v e
-
cho cc cht nhn trung gian v cui
cng n O
2
. Qu trnh ny xy ra mng trong ca ty th. Kt qu ca
qu trnh ny to ra cc phn t ATP v H
2
O.
H hp ho kh:

99
c tnh ca h hp ho kh l gii phng CO
2
v tip nhn O
2
. H
hp ym kh theo nh ngha trc y l gii phng CO
2
trong iu kin
khng c O
2
. Ho sinh hc nh ngha h hp ym kh l s phn gii
carbohydrate qua qu trnh ng phn v tip theo l s ln men ru.
Ton b qu trnh ny khng cn O
2
, CO
2
c gii phng ra trong qu
trnh ln men bng s kh carboxyl ho pyruvate.
H hp ho kh xy ra trong ty th. Khi qu trnh quang hp v h
hp ho kh c gii thch cng chi tit th s ging nhau gia chng
nh l my bin i nng lng cng nhiu. qu trnh quang hp
ngi ta bit matrix cha nhng enzym ca chu trnh Calvin, h hp
matrix ca ty th cha nhng enzyme ca chu trnh Krebs. H thng oxy
ho kh ca quang hp (chui vn chuyn e
-
trong quang hp) nh v
mng thylacoid ca lc lp, h thng oxy ho kh ca h hp nh v trn
mng trong ca ty th. S ging nhau gia hai my bin i nng lng
n l ngc nhin.












Hnh 4.19 Chui vn chuyn e
-
ca h hp vi nhng h thng oxy ho kh
quan trng v cc phc h vn chuyn mng

Chui vn chuyn e
-
trong h hp, v s nh v ca chng trong mng
trong ca ty th, c gii thch hnh 4.19. Chui vn chuyn e
-
gm c 4
phc h proteine vn chuyn ca mng v hai phc h protein nh hn,
cuccinate dehydrogenase (FAD) v cytochrome. Ngoi ra cn c
ubichinol/ubichinone trong chui vn chuyn. Cht kh tng ng l NADH
v FADH
2
, c hai u c to ra trong chu trnh Krebs, c i vo chui
h hp qua phc h FMN-protein cng nh qua phc h FAD-protein. Hai
Pha ngoi ty th (pH 5)
2H
+
H
+
O
2
+ 2H
+
H
2
O
ADP + P
i
ATP

FMN
FAD
NAD
+
NADH + H
+
Succinate

Fumarate

Matrix (pH 8)

ATP Syn-
thetase

Cyto-
chrome
b,c
1
Cyto-
chrome
a,a
3
Cyt c

Ubichinol

Ubi-
chinol

Ubichinone

Ubi-
chinone

P
r
o
t
o
n
e
n
-
k
a
n
a
l

100
phc h vn chuyn H ca n n ubichinone v kh n thnh ubichinol. H
thng ubichinone/ ubchinol v h thng plastochinone/plastochinol c cu to
v chc nng rt ging nhau (xem s ).











Ubichinone/ ubichinol c kh nng khuch tn rt nhanh vo mng
nh c tnh a lipid ca n. H s khuch tn nm 10
-8
cm
2
/sec. Ubichinol
kh phc h cytochrome b, c
1
, y dn n s tch H thnh H
+
v e
-
.

















in t c tip nhn bi phc h cytochrome b, c
1
, trong khi
H
+
c thi ra pha ngoi mng. Phc h cytochrome b, c
1
, cn c
OH
OH
O
O
2e
_
2H
+
2x Cytochrome.Fe
3+
2x Cytochrome.Fe
2+
CH
CH
3

C
CH
2
H
n
O
O
H
3
CO
H
3
CO
CH
2

CH
3

2H

Ubichinol

Ubichinone

CH
CH
3

C
CH
2
H
n
OH
OH
H
3
CO
H
3
CO
CH
2

CH
3

101
gi cytochrome c-reductase, v n kh cytochrome c, in t ca n c
chuyn cho cytochrome c. y cng l 1 phc proteine nh hn, c th
khuch tn d dng ra pha mng ngoi. N chuyn e
-
n phc h
cytochrome a, cn c gi l cytochrome c-oxidase, v n oxy ho
cytochrome c. Phc h cytochrome a l thnh vin cui cng ca chui
vn chuyn e
-
. N chuyn e
-
n O
2
, ri phn ng vi H
+
ca h thng
to H
2
O. O
2
trong chui h hp c nhim v tip nhn e
-
cui cng.
Chui h hp trong hnh 4.19 lm r s vn chuyn H v e
-
qua
cc thnh vin ca chui dn n s chnh lch proton ca mng. S khc
bit ny xut hin s kh ca phc h cytochrom b, c
1
bi ubichinol. V
c s oxy ho NADH phc h FMN v s kh O
2
phc cytochrome a
lm cho matrix kim, v n lm gim H
+
, ngi ta c th nhn bit trn
hnh 4.19.
Bng cch ny m 1 th in ho c to nn. Matrix tch in m
v c mt nng H
+
thp (pH 8), ngoi mng tch in dng v c
nng H
+
tng i cao. S chnh lch th nng gia 2 mng c ln
l 200mV. Nng lng ca in th ho hc ny c s dng tng
hp ATP. Bn phc h vn chuyn mng c cha ATP-synthetase
(ATPase). Proton i qua cc knh vn chuyn ca phc h vn chuyn
mng t ngoi mng vo matrix, thc hin tng hp ATP. y c s
tng t nh sinh tng hp ATP trong quang hp.
Trc khi gii thch nng lng ca chui h hp, cn ni r hn h
thng oxy ho kh.
- Phc h FMN, NADH-dehydrogenase gm t nht 26
polypeptide khc nhau. N cha FMN trung tm phn ng v c
khong 20 nguyn t Fe v S linh ng/ FMN. Trng lng phn
t ca n l 850 kDa.
- Phc h FAD l cuccinate dehydrogenase, cha FAD v
phc Fe-S ho tr 2 v ho tr 4 trung tm phn ng.
- Phc h cytochrome b, c
1
cha cytochrome c
1
v 2 loi
cytochrome b, t nht 8 chui polypeptide ln hn v 1 phc Fe-S
c ho tr 2. Antimycin l 1 cht km hm ca phc h cytochrome
b, c
1
, n nh ch s vn chuyn e
-
gia cytochrome b v
cytochrome c
1
.
Cytochrome a, a
3
, cn c gi cytochrome c-oxidase gm 7 chui
polypeptide khc nhau. ng thi n tip nhn t cytochrome c 4 e
-
v
chuyn chng n 1 O
2
. s vn chuyn ny c s tham gia ca heme
trong cytochrome a
3
, cytochrome a v nguyn t Cu. Oxy dng kh (O
2
4-
)
phn ng vi 4H
+
c ngun gc t nc to H
2
O.
102
O
2
4-
+ 4H
+
H
2
O
Phn ng oxy ho kh cui cng ca chui h hp ny c gi l
s oxy ho cui cng. N s dng khong 90% nhu cu O
2
ca t bo.
S oxy ho ny b c ch bi CO
2
, cyanid v acid. Nhng cht ny kt
hp vi heme v phong to s kt hp ca hem vi O
2
, v vy chng l
cht c i vi h hp. Phc h vn chuyn mng, cha ATP-synthetase
gm nhiu chui polypeptide khc nhau, trong cha 1 knh proton,
thc hin s vn chuyn H
+
t pha mng ny sang pha mng khc. ATP-
synthetase ny rt ging ATP-synthetase trong mng thylacoid ca lc
lp v ATP- synthetase ca vi khun.
S vn chuyn e
-
trong chui h hp l s vn chuyn thun chiu,
theo th oxy ho kh tiu chun, nh trong hnh 4.20. Nng lng gii
phng ra y c bin i thnh nng lng thm thu, th hin s
chnh lch nng H (gradient H
+
). S chnh lch nng H
+
hai pha
ca mng to nn mt th in ho, m ln ca n c tnh theo
phng trnh sau y:
P -59 pH +
P lc
pH Grandient H
+

s khc nhau v in tch gia 2 mng


















FMN FAD
Cytochrome
b, c
1
Cytochrome
c

Cytochrome
a, a
3

-400
-200
0
+200
+400
+600
+800
Ubichinone
e
-
e
-
1/2O
2
+ 2H
+
H
2
O
Th oxy ha kh tiu chun, mV
103





Hnh 4.20 S vn chuyn e
-
trong chui h hp ph thuc vo th oxy ho
kh tiu chun
Phng trnh ny ng vi mt nhit nht nh, m t lc di
chuyn proton ngha l lc y 1 H
+
t 1 pha mng nng H
+
cao
(pha ngoi mng) n 1 pha khc ca mng vi nng H
+
thp
(matrix). Lc ny c biu din mV l s chnh lch in th, y 1
n v pH tng ng vi 59 mV, v vy l h s ca phng trnh. Lc di
chuyn proton phi ln, mi tng hp c ATP. Vy bao nhiu H
+

phi i vo matrix, mi tng hp c 1 ATP? Ngi ta cho rng s
chnh lch th (lc t pH v , xem phng trnh) l 220 mV s vn
chuyn 1 mol H
+
t mng ngoi n matrix th nng lng t do gii
phng ra l 21.3kJ. Khi 2H t NADH trong chui h hp c a vo,
n tch ra 5H
+
m ngi ta c th rt ra t hnh 4.19. Ba H
+
cn trong
matrix, hai H
+
c i ra ngoi mng. Khi 5 mol H
+
tr v matrix, to ra 5
x 21,3 106.5 kJ nng lng t do vi iu kin chnh lch th l 220
mV. Gi s nng lng ny tng hp 3 mol ATP, v G
0
ca 1 lin
kt cao nng trong ATP l 30kJ. Hai nguyn t H, i vo chui enzyme h
hp qua phc h FMN, cn s oxy ho cui cng 1/2O
2
. th nng 220
mV c th tng hp t 2H (t NADH) 3 ATP vi s tiu hao 1/2 O
2
. Nh
vy t l P/O l 3, l s phn t ATP c tng hp/nguyn t O. Tuy
nhin gi tr thu c t thc nghim chnh lch nhiu so vi t l P/O
ny. iu ny d hiu v t l ch ng khi chnh lch th l 220 mV. Khi
tnh ton nng lng bin i ngi ta cho rng c cht h hp NADH +
H
+
to ra 3ATP, c cht l FADH
2
l 2ATP. Tuy nhin t l P/O vi
NADH v FADH
2
theo gii thch trn c th lch nhiu so vi gi tr thc
t. S chnh lch ny cn do nng lng t do tng hp ATP c tnh
trong iu kin tiu chun, m trong thc t phn ln cc trng hp
khng phi trong iu kin ny. Nng lng cn cho tng hp ATP hng
v trng thi cn bng phn ng:
ADP + Pi ATP
T l ATP/ADP iu khin h hp rt hiu qu v c ngha.
nng ADP cao, tng ng vi s thiu ATP th ATP c to nn
nhiu v nhanh vi tiu hao nng lng t, ngc li vi nng ATP cao
104
th s tng hp ATP cn nhiu nng lng. y l s thch nghi tt vi
nhu cu nng lng. Khi ATP c s dng nhiu cho cc qu trnh sinh
trng (tng hp protein, nucleic acid) hoc cho lao ng c bp, th
cng h hp tng ln, lc ny nng ATP thp v ADP cao. Trong
giai on c bp t hot ng nng ATP cao, tng ng vi cng
h hp thp.
nng ATP cao phn ng tng hp ATP c th theo chiu
ngc li. y ATP c tch thnh ADP v Pi v nh nng lng gii
phng ra m H
+
c bm ra pha ngoi mng. ATP-synthetase tr thnh
mt bm proton. V l do ny ngi ta gi ATP-synthetase ngc cng l
ATPase. Thc t l cc bm proton rt ph bin trong t nhin. Ngc vi
ATP-synthetase trong chui h hp, ATP-synthetase c trong mng
thylacoid khng th tch ATP.
H hp c chia lm hai loi: H hp ym kh v h hp ho kh.
C hai kiu h hp ny u c mt qu trnh chung l qu trnh ng
phn.

4.5.2 ng phn, s ln men ru v ln men lactic

Nh nhng enzyme thu phn k trn m chui carbohydrate
d tr c bin i thnh hexose, nu khng phi l glucose hoc
fructose, th chng cng d dng bin i thnh hai dng monosaccharide
ny. Glucose v fructose l nhng c cht trc tip ca qu trnh ng
phn (glycolyse).
Glycolyse c ngha l phn gii ng. Cc phn ng c gii thch
bi Gustav Embden v Otto Mayerhof. V vy qu trnh ny c gi l
s phn ng Embden- Mayerhof. S ny gm s phn gii ng v
polysaccharide ym kh cho n pyruvate. Nhng enzyme ca qu trnh
ny c mt t bo cht. Nh tc dng ca phosphorylase glucose1-
phosphate c tch ra l c cht trc tip ca qu trnh ng phn.
Chui phn ng gm nhng phn ng ring l nh sau:

1. T chui amylose nh phosphorylase m glucose1-phosphate
c tch ra.
2. Glucose1-phosphate c bin i thnh glucose-6-phosphate
nh isomerase.
3. Glucose-6-phosphate c bin i thnh fructose-6-phosphate
nh enzym isomerase.
105
4. Fructose-6-phosphate c phosphoryl ho nh ATP v
enzyme phosphofructokinase to thnh fructose-1,6-
diphosphate
5. Fructose-1,6-diphosphate nh enzyme aldolase c tch ra
thnh aldehydphosphoglyceric v phosphodioxyacetone.
Khi aldehydphosphoglyceric (AlPG) c s dng trong cc
phn ng tip theo th cht ny li c to thnh t
phosphodioxyacetone (PDOA). trng thi cn bng c hn 90% l
dng PDOA.
6. Aldehydphosphoglyceric (triosephotphate) c kt hp vi
mt HS-enzyme. Phc h HS-Enzyme ny cn kt hp vi
NAD
+

7. Bng vic nhng 2H cho NAD
+
, AlPG c oxy ho thnh
phosphoglyceric acid. S thay i trong phn t lm cho nhm
acyl giu nng lng c gn vo S ca phc h enzyme.
8. NADH gn vo phc SH-Enzyme c oxy ho, trong H v
e
-
ca n c chuyn ln mt NAD
+
t do.
9. Nh gc phosphate v c m phosphoglycerinacyl c tch ra
khi phc h. y nhm SH li xut hin khi to thnh 1,3
phosphoglyceric acid.
qu trnh ny phosphoric acid (P-OH) c tch ra 1 nguyn t H v
1 gc P-O- (xem s ). Gc P-O- c gn kt vo acyl ca
phosphoglyceric acid, trong khi H c kt hp vi nguyn t S ca
enzyme. Enzyme li tr li trng thi ban u.














P - O H

CO ~ S -

HCOH

CH
2
O P
O
C - O ~ P

C = O

CH
2
O P



E
n
z
y
m
e

E
n
z
y
m
e

HS -
106
10. Nhm phosphoryl mang nhiu nng lng ca 1,3
diphosphoglyceric acid c chuyn n ADP, tng hp nn
ATP.
11. 3-phosphoglyceric acid c bin i thnh 2-phosphoglyceric
acid nh enzyme isomerase
12. Nh enzyme hydratase H
2
O c tch ra t 2-phosphoglyceric
acid. Phosphoenolpyruvate c to thnh. Cu trc phn t
ca lin kt ny to nn nhm phosphoryl giu nng lng.
13. Nhm phosphoryl ca phosphoenolpyruvate c chuyn n
ADP. ATP c tng hp.
14. Enolpyruvate trng thi cn bng vi dng ceto ca pyruvic
acid (s ).












Hexose (glucose, fructose) cng nh triose (aldehydglyceric) c
phosphoryl ho nh kinase, trc khi chng tham gia vo phn ng ca
qu trnh ng phn.
Ton b qu trnh ng phn c chia lm 3 giai on:
1. Cc carbohydrate c bin i thnh triosephosphate do s
oxy ho ng.
2. Triosephosphate c bin i do s oxy ho thnh cc acid
hu c (glycerate).
3. Phosphoglycerate c bin i thnh pyruvate. Vi s gii
phng ra nng lng t do, ATP c tng hp.


COO H

C - OH

CH
2
Enolpyruvate
COO H

C = O

CH
3
Pyruvate
107

Hnh 4.21. Qu trnh ng phn

108
Tng qut: khi a vo 1 glucosephosphate v kt qu to ra 2 phn
t triosephosphate:
u vo u ra
1 glucosephosphate +1ATP + 2 NAD
+
ADP + 2NADH + 2H
+
+ 2 HS-Enzyme + 2HS- Enzyme +4ATP +2H
2
O
2 P-OH + 4 ADP + 2 pyruvic acid
Tng qut: glucosephosphate + 2 NAD
+
+ 3 ADP + 2 P-OH
2 pyruvic acid + 2 NADH + 2H
+
+ 3ATP +2 H
2
O
S tng qut ny lm r vai tr sinh l ca qu trnh ng
phn: hexosephosphate c phn gii thnh apyruvic acid. Nng lng
gii phng ra c dng tng hp ATP v kh NAD
+
. V vy qu
trnh ng phn v ngha sinh l tng ng vi phn ng sng ca qu
trnh quang hp. Qu trnh quang hp cn nng lng nh sng mt tri
tng hp ATP v NADPH, trong khi qu trnh ng phn th nng
lng ho hc c s dng tng hp ATP v NADH. S tng hp
ATP qu trnh ng phn xy ra trong iu kin ym kh. V vy ngi
ta gi phosphoryl ho ym kh, ngha l phosphoryl ho ADP tng hp
ATP khng cn O
2
.

4.5.3. Cc loi h hp
Pyruvate c coi l sn phm cui cng ca qu trnh ng phn.
N c th ln men ru to ethanol hoc ln men lactic to lactic
acid hoc c phn gii qua chu trnh Krebs to CO
2
v H
2
O.
s ln men ru trc ht pyruvate c kh cacboxyl ho thnh
acetaldehyde, sau di tc dng ca NADH n c kh thnh
ethanol.







NADH c ra trong qu trnh ng phn c s dng oxy
ho acetaldehyde, nh vy s ti to NADH l do oxy ho
aldehydphosphoglyceric (phn ng 8 ng phn) cung cp cho phn ng
ny.
Ethanol
COO H

C = O

CH
3
Pyruvic acid
NAD
+
NADH + H
+


CO
2
CHO

CH
3
CH
2
- OH

CH
3
Acetaldehyd
109
Qu trnh ln men ru gm qu trnh ng phn t carbohydrate
cho n pyruvic acid, t cht ny n acetaldehyde v ethanol l sn phm
cui cng. ng hexose (glucose, fructose) l nguyn liu ban u ca
qu trnh ln men, tng qut c biu din nh sau:

1 Hexose + 2ATP fructose-1,6-diphosphate
Fructose-1,6-diphosphate 2 aldehydphosphoglyceric
2 aldehydphosphoglyceric + 2 NAD
+
2 pyruvic acid + 4ATP +2NADH
2 pyruvic acid + 2 NADH 2 ethanol + 2 NAD
+
+ 2 CO
2

Cui cng: 1 hexose 2 ethanol + 2 CO
2
+ 2 ATP

Nhiu c quan nh r ca thc vt bc cao, c kh nng thc hin s
ln men ru trong iu kin ym kh. i vi cng ngh quan trng l
Saccharomyces cerevisiae, l mt loi men k kh khng bt buc, ngha
l n c th pht trin trong iu kin c hoc khng c O
2
. Khi khng c
O
2
s phn gii glucose xy ra nhanh hn. Ngi ta gi hiu ng ny l
hiu ng Pasteur. Pasteur pht hin ra rng, s phn gii ng trong
iu kin ym kh xy ra 7 ln nhanh hn so vi iu kin ho kh. Ngy
nay chng ta bit rng, hiu ng ny l do s iu khin bin i cu trc
(bin cu) ca enzyme fructosephosphatkinase. s phn gii ho kh th
ATP c to ra tng i nhiu, v khi nng ATP cao, n c ch hot
tnh ca fructosephosphatkinase lm cho s i vo ca glucosephosphat b
gim xung. Ngc li khi phn gii ng trong iu kin ym kh, nng
AMP cao, n kch thch hot tnh ca fructosephosphatkinase. ADP
c to ra nhiu phn ng do kinase xc tc, c bin i thnh ATP
v AMP nh sau:
2 ADP ATP + AMP
Ln men lactic cng gm qu trnh bin i t carbohydrate n
pyruvate, cht ny c kh to thnh lactic acid nh NADH c to
ra trong qu trnh ng phn.







Lactic acid
COO H

C = O

CH
3
Pyruvic acid
NAD
+
NADH + H
+


COO H

HCOH

CH
3
110
Qu trnh ny c tng qut nh sau:
1 glucose 2 lactic acid + 2 ATP

Nhng vi khun lactic quan trng thuc nhng chng loi sau:

Streptococcus khng mn cm vi O
2

Lactobacillus khng mn cm vi O
2

Clostridium ym kh nghim ngt
Vi khun streptococus v lactobacillus khng mn cm vi O
2
,
ngha l chng c enzyme superoxiddismutase, c kh nng lm mt c
tnh ca gc O
2
. Streptococcus v lactobacillus c kh nng s dng O
2

nhng khng phi cho s phn gii ho kh (h hp), v chng khng c
kh nng tng hp heme. Chng khng c cytochrome, l thnh phn
quan trng ca h hp. Clostridium sng nghim ngt trong iu kin ym
kh, nn O
2
l cht c i vi chng. Clostridium botulinum tng hp
c cht c t nhin mnh nht l botulinus-toxin. Mt lng rt nh
0.03 ng/kg trng lng c th gy cht.
Hu ht vi khun lactic khng mn cm vi nhit v pH thp.
qu trnh ln men lactic nhanh th pH mi trng gim nhanh v v nhiu
vi sinh vt mn cm vi pH thp, v d vi khun gy thi, nn chng
khng c c hi cnh tranh vi vi khun lactic, nh vy m nng sn c
bo v.
Khi sn xut thc n chua th s gy chua nhanh nh vi khun lactic
l mt iu kin c bn cho thnh cng ca ln men lactic v bo qun.
Ln men ch yu l nh lactobacillus plantarum, loi vi khun sng trn
l cy v bng cch no m c a vo cng vi nng sn. ln
men lactic nhanh iu quan trng l thc vt c carbohydrate, c bit
l saccharose. Nhng bp ci trng gi tho mn iu kin ny. Thc
phm xanh, non th hm lng carbohydrate t, v cy thc n giu m
th s ln men lactic xy ra chm, pH ch c gim rt t v nhng vi
khun khc, nh vi khun butyric c th phn gii thc phm xanh. Vic
cho thm mt vo c th hn ch nhc im trn, v mt cha nhiu
saccharose v raffinose, nhng cht ny sau khi thu phn c phn gii
nhanh bi vi khun lactic. Ng chn sa l nguyn liu ln men l tng,
v trong qu ca n cha nhiu carbohydrate d phn gii.
Khi sn xut phomat s ln men lactic cng c ngha quan trng.
Sa chua c gy ra phn ln l do streptococcus lactis v streptococcus
cremonis. pH thp dn n s kt ta protein (sa c li), c gi l
phomat trng. Phomat ny khng nhng cha protein m cn c cht bo
111
sa. Phomat ny l nguyn liu sn xut cc loi phomat khc nhau.
sn xut phomat cn c s tham gia ca cc vi khun lactic, streptococus,
lactobacillus, chng khng nhng to nn lactic acid m cn thu phn
protein, ngha l phn gii thnh peptide v aminoacid. Trong phomat
lactic acid cn c chc nng bo qun.
Nguyn l ca s phophoryl ho ym kh c nghin cu k. Nh
hnh 4.21 phn t c cht c bin i cho n nhm phosphoryl ca
n dng lin kt giu nng lng. Nhm phosphoryl ny c chuyn
n ADP, tng hp ATP. V vy ngi ta gi s phosphoryl ho ny l
phosphoryl ho c cht.
So snh hai giai on ca phn gii carbohydrate (h hp ho kh)
th nng lng thu c (ATP) t qu trnh ng phn ch chim khong
20% tng nng lng d tr trong phn t glucose. Phn nng lng ln
nht c to ra s giai on phn gii tip theo.
Qu trnh ng phn c iu khin bi enzyme
fructosephosphatkinase (phn ng 4 trong s ). cng l enzyme
quan trng ca qu trnh ng phn. Enzyme c 4 tiu n v b c ch
theo c ch tc ng bin cu bi ATP v citrate, ngc li c hot
ho bi AMP. V vy t l AMP/ATP cao kch thch v t l ATP/AMP
cao th c ch hot tnh ca enzyme. C ch ny rt c ngha, v khi
trong h thng c nhiu ATP th khng cn tng hp qua qu trnh ng
phn. Ngc li khi t l AMP/ATP cao biu hin s thiu ATP, xy ra
s tng hp ATP. Mt nng citrate cao l cht trao i cung cp cho
chu trnh Krebs. V qu trnh ng phn cung cp cc cht trung gian cho
chu trnh Krebs, nn s iu khin qu trnh ng phn qua hiu qu bin
cu ca citrate n fructosephosphatkinase l mt c ch rt c ngha.

4.5.4 H hp ho kh (chu trinh Krebs)

Chu trnh tricarboxylic, hay l chu trnh Krebs c gii thch bi
Hans Krebs, v Schueler Otto Warburg. Nh pht minh ny m Krebs
nhn c gii thng Nobel vo nm 1953.
Chu trnh ny xy ra trong ty th. C quan t ny c th so snh
c vi lc lp, thc ra n nh hn lc lp (ng knh 0.5-1 m), tuy
nhin v cu to v chc nng chng tng t nhau (hnh 4.22)



112
.















Hnh 4.22 S cu to ca ty th: hnh pha trn l s ct ngang qua
ty th, hnh pha di l ct mt phn ty th

Ngi ta cho rng cng nh lc lp, cc ty th xut hin trc
y khong 10
9
nm trong nhng c th c lp, trong qu trnh tin ho
chng c sinh vt nhn chun bt v ho nhp. Nhng t bo c s
bin i nng lng ln cha nhiu ty th, nh t bo m c hoc t bo
gan, chng c hn 1000 ty th trong mt t bo. Ty th xut hin di
knh hin vi l nhng ht nh hoc si.
Tuy nhin, di knh hin vi in t, ty th c bao bc bi mng
ngoi v bn trong l mt h thng mng ko dn vi nhiu s un cong
vo. Nh cu trc ny m mng trong c din tch b mt ln hn mng
ngoi khong 5 ln. Mng ngoi cho nhng phn t c trng lng ln
(n 10.000 Da) thm qua, v mng ny cha nhng protein nh l
nhng knh vn chuyn. Mng trong kh thm qua hn v vn chuyn cc
cht c tnh chn lc. Mng trong bao quanh stroma, trong stroma cha
cc enzyme ca chu trnh Krebs v enzyme phn gii cht bo (hnh 4.22).
Ngc li trn mng trong ty th c cc enzyme v h thng oxy ho.
y tng t nh lc lp, chui vn chuyn e
-
nh v trn mng thylacoid
v cc enzyme ca chu trnh Calvin trong matrix.
Mng ngoi
Mng trong
Matrix
Khong gia
hai mng
Khong gia
hai mng Matrix
Mng ngoi
Mng trong
113
C cht ca chu trnh Krebs, pyruvate l sn phm ca qu trnh
ng phn. N c vn chuyn t t bo cht qua mng trong ty th n
stroma v y n c kh carboxyl ho oxy ho nh enzyme pyruvate
dehydrogenase. Pyruvate dehydrogenase l mt v d v a enzyme, gm
mt phc h cha carboxylase, transacetylase v dehydrogenase. Bn cnh
trong phn ng cn c cc nhm prostetic l thyaminpyrophosphate,
liponic acid, coenzyme NAD
+
v Coenzyme A. Sau y l cc bc c
th ca chu trnh Krebs (hnh 4.23). Pyruvate b kh carboxyl ho oxy ho
trc khi i vo chu trnh.
Tng t qu trnh ng phn chu trnh Krebs l mt v d ni ln rng
trong c th sng cc phn ng phn gii cc cht cha nhiu nng lng
thc hin theo tng bc. mi chu trnh Krebs ly vo 1 phn t acetyl
CoA.
1. Pyruvic acid c chuyn thnh acetyl CoA bng phn ng kh
carboxyl ho oxy ho. phn ng oxy ho nng lng gii phng ra
c lin kt trong NADH v trong lin kt cao nng ca acetyl-CoA.








2. Nng lng cha trong lin kt cao nng ca acetyl-CoA gn
acetate vo oxaloacetate to thnh citrate.









3. Qua s ngng t ny 1 phn t c 6 nguyn t C c to nn. Phn
ng c xc tc bi enzyme citrate synthetase. Bng phn ng 3 v 4
citric acid c bin i thnh isocitric acid qua aconitic acid.
OH

CH
2
C CH
2
+ CoASH

COOH COOH COOH
Citric acid

+ H
2
O
Oxaloacetic acid

O
C
SCoA
O

CH
2
C + HCH
2
-

COOH COOH
CO ~ SCoA

CH
3

Acetyl - CoA
COO H

C = O

CH
3
Pyruvic acid
CoASH + NAD
+
NADH + H
+
+ CO
2
114








4. Di tc dng ca aconitase (hydratase) xy s kt hp vi nc.
trng thi cn bng citric acid chim 91%, aconitic acid 3% v
isocitric acid 6%.









5. hnh 4.23, phn ng 5 v 6 c thc hin bi enzyme isocitrate
dehydrogenase. Trc ht isocitric acid c oxy ho thnh
oxalosuccinic acid v sau c kh carboxyl ho thnh axit -
cetoglutaric acid.
6. C hai dng isocitrate dehydrogenase, 1 dng cng vi NAD
+
v 1
dng cng vi NADP
+
. ty th isocitrate dehydrogenase dng NAD
+

quan trng hn.
7. Bng phn ng kh carboxyl ho oxy ho - cetoglutaric acid c
bin i thnh succinyl-CoA. Enzyme xc tc l 1 phc h -
cetoglutarate dehydrogenase. Phn ng v enzyme tng t s kh
carboxyl ho oxy ho pyruvic acid.





COOH

CH
2


CH
2

O
C
SCoA
Succinyl-CoA

COOH

CH
2


CH
2


CO- COO H
NAD
+
NADH + H
+
+ CO
2 + CoA-SH

CH
2
- COOH

CH - COOH

H OC H - COOH
Isocitric acid
CH
2
- COOH

CH - COO H

O=C - COOH
Oxalosuccinic acid
COOH

CH
2


CH
2


C=O

COOH
-cetoglutaric acid
NAD
+
NADH + H
+
CO
2
CH
2
- COOH

CH - COOH

HOCH - COOH
Isocitric acid
CH
2
- COOH

HO C - COOH

H CH - COOH
Citric acid
CH
2
- COOH

CH - COOH

CH - COOH
Aconitic acid
- H
2
O + H
2
O
115






8. T succinyl-CoA, succinic acid c to thnh. Lin kt giu nng
lng ca succinyl-CoA c b gy, nng lng gii phng ra dng
tng hp guanosintriphosphate (GTP) t guanosindiphosphate
(GDP) v phosphate v c. Trong c th thc vt GTP c to thnh
tng t nh ATP.










9. Di tc dng ca FAD succcinate b oxy ho to thnh
fumarate.










Enzyme ny l mt flavoprotein, c nhm prostetic l FAD c gi
l succinate dehydrogenase. s oxy ho succinate thnh fumarate nng
lng gii phng khng kh NAD
+
. Tuy nhin n kh FAD
(xem th kh tiu chun).
10. Di tc dng ca H
2
O, fumarate bin thnh malate
COOH

CH

CH

COOH
Fumaric acid

COOH

HC H

HC H

COOH
Succinic acid

FAD

NADH
2
COOH

CH
2


CH
2


COOH
Succinic acid

GDP + P

GTP

COOH

CH
2


CH
2

O
C
SCoA
Succinyl-CoA

116











11. Malate dehydrogenase kh malate thnh oxaloacetate, l cht
khi u ca 1 chu trnh mi.









Nhng phn ng c bn trong chu trnh Krebs l s kh carboxyl
ho v phn ng oxy ho. Nu tnh phn ng t pyruvate n acetyl-CoA
vo chu trnh Krebs th c 3 v tr khc nhau CO
2
c gii phng (phn
ng 1,6,7) v 5 v tr khc nhau c cht b oxy ho (phn ng 1,5,7,9 v
11). nhng phn ng oxy ho lun lun c 1 coenzyme (NAD
+
) hoc 1
nhm prostetic (FAD) tham gia. Chng chuyn thnh dng kh bng phn
ng oxy ho. Nng lng c gii phng ra do s oxy ho c cht c
s dng cho vic kh coenzyme cng nh prostetic. Quan trng l nng
lng phn ng 8 c dng tng hp 1 phn t GTP, tng ng
vi 1 ATP, v GTP c th phn ng vi ADP nh sau:
GTP + ADP ATP + GDP
T hnh 4.23 rt ra rng, trong chu trnh, CO
2
c tch ra nhng O
2

khng c tip nhn vo. Nguyn t O c trong CO
2
khng phi l oxy
cho h hp. ngha ca chu trnh Krebs l cung cp nhng coenzyme kh
(NADH, FADH
2
), nhng cht kh ny l cn thit cho h hp. Ngoi ra
COOH

CH
2


HCOH

COOH
COOH

CH
2


C=O

COOH
NAD
+
NADH + H
+
Malic acid

Oxaloacetic acid

COOH

CH
2


HCOH

COOH
Malic acid

COOH

CH

CH

COOH
Fumaric acid

+ H
2
O
117
cc acid khc nhau ca chu trnh cn tham gia vo nhng qu trnh trao
i cht khc.



















Hnh 4.23 Chu trnh Krebs

Qua chu trnh Krebs chui carbon t carbohydrate n trao i lipid
v trao i ca aminoacid, protein.


4.5.5 Chu trnh pentosephosphate- oxy ho trc tip glucose

Bn cnh qu trnh ng phn trong t bo cht cn xy ra mt
con ng khc, gi l chu trnh pentosephosphate v qu trnh phn ng
c phn tng t chu trnh Calvin. Phn ng bt u vi vic oxy ho trc
tip glucose-6-phosphate, v vy gi l oxy ho trc tip glucose. Enzyme
xc tc cho phn ng u tin l glucose-6-phosphate-dehydrogenase,
Oxaloacetate
Acetyl-CoA
Pyruvate
1
HSCoA + NAD
+
NADH+ CO
2
HSCoA

Citrate

H
2
O
H
2
O
Aconitate
H
2
O
Isocitrate
Oxalosuccinate
HSCoA + NAD
+
NADH+ CO
2
-cetoglutarate
CO
2
Succinyl-CoA
Succinate
Fumarate
Malate
H
2
O
NADH

NADH

NAD
+
NAD
+
FAD

FADH
2
GTP + HSCoA

GPP + phosphate

2
3
4
5
6
7
8
9
10
11
118
c Otto Warburg pht hin nm 1931. Cc bc oxy ho c ch ra
trong hnh 4.24.
- Phn ng 1: T glucose-6-phosphate 2 nguyn t H nh C
1

c tch ra nh enzyme glucose-6-phosphate dehydrogenase,
xut hin dng lactone. Glucose-6-phosphate dehydrogenase c
hiu i vi NADP
+
. i lc ca n i vi NADP
+
cao hn 1000
ln so vi NAD
+
.





















Hnh 4.24 S oxy ho trc tip glucose trong chu trnh pentosephosphate

- Phn ng 2: Dng lactone c thu phn, to nn 6
phosphogluconic acid.
- Phn ng 3: 6-phosphogluconate c kh carboxyl ho oxy ho.
Enzyme xc tc l 6-phosphogluconate dehydrogenase.



CH
2
O P
OH
HO
OH
H
O
OH
CH
2
O P
OH
HO
OH
O
O
COOH

HC-OH

HOCH

HCOH

HCOH

CH
2
O P
6-phosphogluconic
acid
Glucose 6-
phosphate
Lactonphosphogluconic acid
NADP
+
NADPH + H
+
+ HOH

1

2

CHO

HC-OH

HC-OH

HC-OH

CH
2
O P Ribose 5-phosphate
CO
2
NADP
+
NADPH + H
+
3

119


























Trong chu trnh ny 2 v tr (phn ng 1 v 3) NADP
+
c kh.
NADPH c to ra c th s dng cho phn ng tng hp, v d tng
hp cc acid bo. Ribulose-1-phosphate khng phi l sn phm cui cng
ca chui phn ng ny. N c bin i d dng thnh cc dng ng
phn nh ribose-5-phosphate hoc xylulose-5-phosphate, theo nh sau.
Chu trnh pentosephosphate cung cp nguyn liu cho cc qu trnh
tng hp khc. Ribose-5-phosphate tham gia tng hp nucleotide, nh vy
l tng hp cc nucleic acid v nhiu enzyme. Xylulose-5-phosphate cng
vi ribose-5-phosphate nh phn ng transcetolase to nn
sedoheptulosephosphate v aldehydphosphoglycic.


CHO

HC-OH

HC-OH

HC-OH

CH
2
O P
Ribose
5-phosphate
Xylulose
5-phosphate
Ribulose 5-
phosphate
Epimerase
Isomerase
CH
2
- OH

C=O

HCOH

HCOH

CH
2
O P
CH
2
- OH

C=O

HOCH

HCOH

CH
2
O P
120
















Transcetolase vn chuyn 2 nguyn t C t ng ceto n aldose,
nhm prostetic ca chng l thiaminpyrophosphate, vng thiazol ca n
ng ceto c kt hp vo khi bt u phn ng.
Sedoheptulosephosphate phn ng vi aldehydphosphoglyceric trong mt
phn ng transcetolase to thnh erythrose-4-phosphate. phn ng
ny 3 nguyn t C c vn chuyn n aldehydphosphoglyceric.












CH
2
- OH

C=O

HOCH

HCOH

HCOH

CH
2
O P
Fructose 6-
phosphate
CHO

HOCH

HCOH

CH
2
O P
Erythrose 4-
phosphate
CHO

HCOH

CH
2
O P
Aldehydphospho-
glyceric
Sedoheptulose
7-phosphate
CH
2
- OH

C=O

HOCH

HCOH

HCOH

HCOH

CH
2
O P
+ +
CH
2
- OH

C=O

HOCH

HCOH

CH
2
O P
Xylulose
5-phosphate
CHO

HC-OH

HC-OH

HC-OH

CH
2
O P
Ribose
5-phosphate
CHO

HCOH

CH
2
O P
Aldehydphospho-
glyceric
Sedoheptulose
7-phosphate
CH
2
- OH

C=O

HOCH

HCOH

HCOH

HCOH

CH
2
O P
+ +
121

Fructose-6-photphate l cht trao i ca qu trnh ng phn, c
th i vo cc qu trnh khc. Erythrose-4-phosphate phn ng vi
xylulose-5-photphate to thnh fructose-6-photphate v
aldehydphosphoglyceric.












Phn ng ny cng vn chuyn 2C (transcetolase). Hai sn phm
ca phn ng l cht trao i ca qu trnh ng phn, c th c phn
gii tip tc trong qua trnh ny.
Ton b cc phn ng ca chu trnh pentosephosphate c biu
din trong hnh 4.25.












CH
2
- OH

C=O

HOCH

HCOH

CH
2
O P
CHO

HOCH

HCOH

CH
2
O P
CH
2
- OH

C=O

HOCH

HCOH

HCOH

CH
2
O P
CHO

HCOH

CH
2
O P
+
+
Xylulose
5-phosphate
Erythrose
4-phosphate
Fructose
6-phosphate
Aldehydphospho-
glyceric
122

















Hnh 4.25. Chu trnh pentosephosphate

Cc bc oxy ho kh u tin dn n 2 v tr kh NADP
+
thnh
NADPH. Dng kh ca coenzyme cn cho phn ng tng hp v 1 H
c i vo cu to mt cht hu c. NADPH ng vai tr quan trng
trong tng hp cht bo. Nhng t bo trong acid bo c tng hp
nhiu th trao i cht theo chu trnh pentosephosphate xy ra mnh m, v
d trong cc m tuyn ca ng vt b sa. Ribulose-5-phosphate l tin
cht ca ribose-5-phosphate, l thnh phn ca nhiu nucleotide v nh
vy l ca nucleic acid. c bit nhng m sinh trng, tng hp nucleic
acid thng xuyn cn ribosephosphate, y trnh t phn ng ca chu
trnh cho n ribulose-5-phosphate thc hin vi tc ln. Erythrose-4-
phosphate l tin cht ca cc aminoacid thm (phenylalanine, tyrosine)
c s dng tng hp protein c bit trong cc m sinh trng. Khi
c nhu cu ln v NADPH th phn ng xy ra vi tc ln. T 6C ca
1 phn t glucose th 5C i vo qu trnh ng phn. Chu trnh
pentosephosphate c kh nng thch nghi vi nhu cu sinh l, tng t chu
trnh Krebs cung cp cho cc con ng trao i cht khc nhau vi chui
carbon hu c v cht kh tng ng.
Cu hi
6-phosphogluco-
lacton
6-phosphoglucuronate
Ribulose 5-phosphate
Nucleic acid
Ribose 5-phosphate
Pentose
-5 (P)
Glucose 6-phosphate
Fructose 6-phosphate
Glycolyse
Fructose 6-(P)
(P) Glyceraldehyde
Amino acid thm
Erythrose
4-phosphate
Xylulose 5-phosphate
Phospho-
Glyceraldehyde
Xylulose
5-phosphate
Ribulose
5-phosphate
Sedo 7-phosphate
H
2
O
CO
2
NADP
+
NADPH + H
+
NADP
+
NADPH + H
+
123
1. Quang hp; h hp l g ? Cc loi h hp?
2. Ni dung v c ch pha sng trong quang hp ?
3. C ch v ngha ca chu trnh C
3
v chu trnh C
4
?
4. Cho bit nhng im khc nhau gia chu trnh C
3
v chu trnh C
4
?
5. C ch v ngha ca qu trnh ng phn ?
6. C ch ca s bin i pyruvic acid trong iu kin ho kh.
7. C ch v ngha ca chu trnh KREBS ?
8. Ti sao ni: nh sng l iu kin cn thit cho qu trnh quang hp?
9. C ch v ngha ca h hp sng (Quang h hp) ?





131
CHNG V
LIPID V S TRAO I LIPID
TRONG C TH THC VT

Lipid l nhm cht hu c c cc c tnh ha l ging nhau, chng
khng tan trong nc, ch tan trong cc dung mi hu c nh ether,
cloroform, benzene, acetone, Khng phi tt c lipid u ha tan nh
nhau trong cc dung mi hu c ni trn m mi lipid ha tan trong dung
mi hu c tng ng ca mnh, nh c tnh ny ngi ta c th phn
tch ring tng loi lipid. V mt ha hc lipid l nhng ester gia ru v
acid bo, in hnh l triglycerid.


R
1
, R
2
, R
3
c th ging nhau,
c th khc nhau, c th bo
ha hoc cha bo ha.



Ngoi ru v cc acid bo, cc lipid phc tp (lipoid), trong phn
t ca chng cn cha cc dn xut c phospho, nit,
Vai tr ca lipid:
- L cht d tr nng lng, khi oxy ha mt gam lipid c th thu
c 9,3 Kcal.
- Lipid cu t ca t bo cht l thnh phn cu to ca t bo v cha
trong t bo vi s lng n nh. Lipid l thnh phn cu trc ca mng
t bo, mng ty lp th, Trong mng sinh hc lipid trng thi kt hp
vi protein to thnh hp cht lipoproteid. Chnh nh hp cht ny to
cho mng sinh hc c c tnh thm thu chn lc.
- Lipid di da ng vt c tc dng gi m v gi m cho c th.
- Lipid l dung mi cho nhiu vitamin quan trng (nh A, D, E, K).
- i vi loi ng vt ng ng, ng vt di c, cc loi su kn,
lipid cn l ngun cung cp nc, v khi oxy ha lipid cho mt lng
nc sinh ra.
- Cc ht cy trng khc nhau c hm lng lipid khc nhau.
V d: u tng (20 30%); go (2,2%); ng (4,9%); la m (1,9%);
cao lng (3,9%); lc (44 56%); thu du (50 60%).

CH
2
O CO R
1

CH O CO R
2

CH
2
O CO R
3




132

I - CU TO V TNH CHT CA LIPID
1.1. Cu to: Du, m c tng hp cc c th sng v ty theo
ngun gc m chng c phn ra du thc vt v m ng vt.
Glycerine l mt ru c 3 chc, do c th hnh thnh mono,
di hay triester. Cc ester ny c bit t lu vi cc tn mono, di v
triacylglycerid. Du, m c ngun gc t nhin lun l hn hp cc
triacylglycerid.
Cc acid bo ca du, m c ngun gc t nhin u c s nguyn
t carbon chn. Bi v cc acid bo u c tng hp t cc n v 2C
(gc acetyl).
Bn cnh cc acid bo bo ha, mt s acid bo khng bo ha
c tm thy trong du, m. Sau y l mt s acid bo bo ha
thng gp:
- Caproic acid (6C): CH
3
(CH
2
)
4
COOH
- Caprilic acid (8C): CH
3
(CH
2
)
6
COOH
- Caprinic acid (10C); lauric acid (12C); miristic acid (14C);
panmitic acid (16C); stearic acid (18C); arachidic acid (20C).
* Cc acid bo cha bo ha thng gp l:
- Oleic acid: CH
3
(CH
2
)
7
CH = CH (CH
2
)
7
COOH
- Linoleic acid:
CH
3
(CH
2
)
3
- [CH
2
CH = CH]
2
(CH
2
)
7
COOH
- Linolenic acid: CH
3
[CH
2
CH = CH]
3
(CH
2
)
7
COOH
- Eruxic acid: CH
3
(CH
2
)
7
CH = CH (CH
2
)
11
COOH
1.2. Tnh cht
a. Nhit nng chy: Ty thuc vo lng acid bo bo ha hay
cha bo ha chim u th trong thnh phn ca du m m nhit nng
chy khc nhau.
Nu trong thnh phn ca du, m c nhiu acid bo bo ha th
nhit nng chy cao, nu nhiu acid bo cha bo ha th nhit
nng chy thp v trng thi lng. a s du thc vt dng lng
nhit thng.
b. Ch s acid: l lng mg KOH cn thit trung ha cc acid bo
t do c trong 1 gam du, m. Ch s acid cng cao th lng acid bo t
do cng nhiu. Cht bo lu ngy, khng bo qun cn thn s c nhiu
acid bo t do. Ch s ny cho ta bit c cht lng ca cht bo.
c. Ch s x phng ha: l lng mg KOH cn thit trung ha cc
acid bo t do v cc acid bo kt hp vi glycerine khi x phng ha 1



133
gam cht bo. Ch s ny c trng cho phn t lng trung bnh ca
glyceride c trong du, m.
d. Ch s iod: l s gam iod c th kt hp vi 100 gam du, m.
Ch s ny dng m t mc khng bo ha ca cc acid bo c
trong thnh phn du, m. Ch s iod cng cao th du m cng long,
chng cng b oxy ha nhanh hn v cc acid bo b oxy ha d nht v
tr cc lin kt i.
S gn iod vo acid bo cha bo ha xy ra theo s sau:





Ch s iod ca m ng vt dao ng trong khong 30 70; cn ca
du thc vt trong khong 120 160.
e. S i ha du, m: Du, m lu ngy s c v hi, ng. Nguyn
nhn l do tc dng ca O
2
. Trng hp ny thng xy ra khi du m
cha nhiu acid bo cha bo ha. Oxy kt hp vo cc ni i ca acid
bo cha bo ha to thnh peroxid:






hoc O
2
kt hp vi nguyn t C bn cnh lin kt i to thnh
hydroperoxid:






Peroxid v hydroperoxid c to thnh lp tc b phn gii to thnh
aldehyd v cetone l nhng cht c mi v kh chu.

II. S PHN GII TRIGLYCERID
Du m l nhng cht dinh dng c gi tr nng lng cao, trong
ht cc cy ly du vn c t carbohydrate th du l cht d tr chnh v

C = C + I
2
C C

I I

C = C C + O
2
C = C C

O OH
Hydroperoxid

C = C + O
2
C C

O O
Peroxid



134
chng l ngun nng lng v ngun vt liu xy dng cho mm ang
pht trin.
Khi phn gii du ngi ta thy hm lng carbohydrate tng ln.
2.1. Phn ng thy phn triglycerid







2.2. Phn gii glycerine







Phn ng 1: do enzyme glycerolkinase xc tc.
Phn ng 2: do enzyme dehydrogenase xc tc.
- Phosphodioxiacetone di tc dng ca enzyme trioso(P)-isomerase
s chuyn thnh aldehydphosphoglyceric.








* Glycerine c quan h gn vi carbohydrate hoc l c s dng
tng hp fructose v carbohydrate khc hoc b phn gii nh
carbohydrate. Sau y l s cc ng hng trao i glycerine:






CH
2
O CO R
1
CH
2
OH R
1
COOH
lipase
CH O CO R
2
+ 3H
2
O CH OH + R
2
COOH
CH
2
O CO R
3
CH
2
OH R
3
COOH
Triglycerid Glycerine Cc acid bo



CH
2
OH CH
2
OH CH2 OH
ATP ADP NAD NADH
2

CH OH CH OH C = O
(1) (2)
CH
2
OH CH
2
O(P) CH
2
O(P)
Glycerine Glycerol(P) (P)dioxiacetone
CH
2
OH O
Trioso-(P) - C H
C = O
isomerase CH OH
CH
2
O(P)
(P)dioxiacetone CH
2
O(P)
Aldehyd(P)glyceric
Glycerine glycerol(P) (P)dioxiacetone Aldehyd(P)glyceric


Fructoso 1,6di(P) Pyruvic acid


Acetyl-CoA


Chu trnh Krebs
Carbohydrate khc



135



2.3. Phn gii acid bo
Mt acid bo mun c oxy ha phi tri qua mt s phn ng sau:

2.3.1. Hot ha acid bo:Nh h thng enzyme Acyl-CoA-Synthetase,
gm 2 bc sau:

Bc 1:



Bc 2:



Qu trnh ny c thc hin ngoi ty, lp th (t bo cht)

2.3.2. Gn Acyl-CoA vo carnitine to thnh acylcarnitine: Cht
ny i qua mng ty th. Trong ty th cc gc acyl ca acid bo c vn
chuyn li cho HS-CoA.














2.3.3. To Acyl-CoA tr li: qu trnh ny ngc li bc gn acyl
vo carnitine. Carnitine c gii phng v tr li mt ngoi ca ty th.
2.3.4. Qu trnh -oxy ha acid bo:
R CH
2
CH
2
COOH + ATP R CH
2
CH
2
CO AMP
-H
4
P
2
O
7
Acyl-AMP

R (CH
2
)
2
CO AMP + HS-CoA R CH
2
CH
2
CO ~ S.CoA
Acyl-AMP -AMP Acyl-CoA




R (CH
3
)
3
(CH
3
)
3


CH
2
+ N
+
Transferase N
+


CH
2
CH
2
-HS.CoA CH
2


CO ~ S.CoA CH OH CH O CO CH
2

Acyl-CoA
CH
2
- COOH CH
2
- COOH CH
2

Carnitine
R
Acylcarnitine




136
Trong c th sinh vt, s oxy ha acid bo xy ra bng cch oxy ha
nguyn t carbon v tr so vi nhm carboxyl, do qu trnh ny cn
c gi l qu trnh -oxy ha. Kt qu ca s -oxy ha l tng i
nguyn t carbon c tch ra di dng acetyl-CoA v acid bo mi to
thnh c mch carbon ngn hn trc 2 nguyn t carbon. S oxy ha
khng th t xy ra, c th tham gia phn ng, acid bo phi c hot
ha nh nng lng ca ATP, nhng y nng lng chuyn t ATP ti
cht bo khng thng qua con ng phosphoryl ha nh trong trng
hp oxy ha glucose m thng qua s to thnh hp cht acyl-CoA. Qu
trnh -oxi ha c Knoop (ngi c) a ra 1904, qu trnh ny xy ra
trong gian bo ty th.
Phn ng tng qut ca s to thnh acyl-CoA nh sau:







Sau acyl-CoA b oxy ha bi enzyme acyl-CoA-dehydrogenase
c nhm hot ng l FAD:








Di tc dng ca enzyme Enoyl-CoA-hydratase, phn t H
2
O kt
hp vo ni i v to thnh -oxyacyl-CoA:








Acyl-CoA-synthetase
R CH
2
CH
2
COOH + HS-CoA R CH
2
CH
2
C ~ S.CoA

ATP AMP + H
4
P
2
O
7
O
(1) Acyl-CoA
Acyl-CoA-dehydrogenase
R CH
2
CH
2
C ~ S.CoA R CH = CH C ~ S.CoA

O FAD FADH
2
O
Acyl-CoA (2) Enoyl-CoA
Enoyl-CoA-hydratase
R CH = CH C ~ S.CoA + H
2
O R CH CH
2
C ~ S.CoA
(3)
O OH O
Enoyl-CoA -oxyacyl-CoA




137
* -oxyacyl-CoA li b oxy ha ln th 2 di tc dng ca enzyme
-oxyacyl-CoA-dehydrogenase c coenzyme l NAD to thnh
-cetoacyl-CoA:










* -cetoacyl-CoA li phn ng vi HS-CoA to thnh acyl-CoA
mi v acetyl-CoA di tc dng ca enzyme -cetoacyl-CoA-thiolase:








Acyl-CoA mi to thnh ny cha gc acid bo c t hn acid bo ban
u 2 nguyn t carbon, n li c th tip tc tham gia cc phn ng 2, 3,
4, 5 to thnh acyl-CoA mi c mch carbon ngn hn 2 nguyn t
carbon v tch ra mt phn t acetyl-CoA na v qu trnh -oxy ha c
tip tc lp li nhiu ln cho n khi ton b phn t acid bo c
chuyn thnh cc acetyl-CoA.
S -oxy ha c Lynen m t di m hnh xon c.
Acetyl-CoA to thnh do kt qu ca s -oxy ha acid bo c th i
vo chu trnh Krebs, chu trnh glyoxilate hoc tham gia vo nhiu phn
ng khc.








NAD (4) NADH
2

R CH CH
2
C ~ S.CoA R C CH
2
C ~ S.CoA
-oxyacyl-CoA-
OH O dehydrogenase O O
-cetoacyl-CoA
(5)
R C CH
2
C ~ S.CoA + HS-CoA R C ~ SCoA + CH
3
C ~ S.CoA
-cetoacyl-
O O CoA-thiolase O O
Acyl-CoA(mi) Acetyl-CoA




138


























Hnh 5.1 - S ca s -oxy ha acid bo (theo Lynen)

Hiu qu nng lng trong qu trnh -oxy ha acid bo:
V d -oxy ha hon ton palmitic acid (16C) t bo thu c ngun
nng lng nh sau: 7 vng quay to ra 7FADH
2
v 7NADH
2
v c 8
phn t acetyl-CoA.





Nhng v phi tiu tn 1 ATP hot ha acid bo nn s ATP to ra
khi -oxy ha palmitic acid l: 35ATP 1ATP = 34ATP.
Nu 8 phn t acetyl-CoA i vo chu trnh Krebs s to ra:
7FADH
2
7 x 2ATP = 14ATP
7NADH
2
7 x 3ATP = 21ATP
Tng cng : 35ATP
R CH
2
CH
2
COOH
ATP
HS-CoA
AMP
1) H
4
P
2
O
7


R CH
2
CH
2
C ~ S.CoA

CH
3
C ~ S.CoA Acyl-CoA O FAD
2)
O Acetyl-CoA FADH
2

O R CH = CH C ~ S.CoA
Acyl-CoA
R C ~ S.CoA 2C 2C 2C 2C

O Enoyl-CoA

HS-CoA 2CH
3
C ~ S.CoA 3) H
2
O
5) Acetyl-CoA
O
R C CH
2
C S.CoA R CH CH
2
C ~ S.CoA
4)
O O NADH
2
NAD OH O
-cetoacyl-CoA -oxyacyl-CoA


Chu trnh
Krebs
Chu trnh
glyoxilate



139
8 x 12ATP = 96ATP
Tng cng: 34ATP + 96ATP = 130ATP.



Ta c th p dng cng thc tnh sau:
A = 12 x
2
n
1 1
2
n
5
*---------------* * -------*
(-oxy ha) (Krebs)

A l s phn t ATP.
n l s nguyn t C (n: chn).

III. S PHN GII CC ACID BO CHA BO HA
i vi acid bo cha bo ha nh oleic acid, linolenic acid qu
trnh -oxy ha din ra bnh thng, cc phn t acetyl-CoA c tch
dn ra, cho ti gn lin kt i. Ti y, ty theo v tr ca lin kt i m
s phn gii c th c cc ng hng sau:
3.1. nhng acid bo cha bo ha m c lin kt i khng
gia C

v C

th nh tc dng ca enzyme isomerase y lin kt i v

ng gia v tr C

v C

sau chu s -oxy ha.

Cng c nhng th nghim chng minh rng: nhng acid bo
cha bo ha m c lin kt i ng v tr gia C

v C

th n chu s
-oxy ha bnh thng.
3.2. Cc acid bo cha bo ha trc khi b oxy ha th chng
chuyn thnh acid bo ha bng cch gn thm H
2
v sau li chu s
-oxy ha nh bnh thng.

IV. S PHN GII CC ACID BO C S NGUYN T
CARBON L
S phn gii cc acid bo ny vn tin hnh bnh thng bng s -oxy
ha cho n khi hnh thnh (propionyl-CoA),



sau propionyl-CoA ( ty th thc vt v vi sinh vt) giai on u
ging nh s -oxy ha cho n khi hnh thnh -hydroxipropionyl-CoA.
CH
3
CH
2
C ~ S.CoA

O



140
Sau nguyn t C

b oxy ha 2 ln (cht nhn H

2
l NAD v NADP)
to malonyl-CoA. Sau malonyl-CoA b kh carboxyl ha to
acetyl-CoA. Cht ny c th i vo chu trnh Krebs hoc i vo chu trnh
glyoxilate.




































-oxy ha
Acid bo c C l CH
3
CH
2
C ~ S.CoA

O
Propionyl-CoA
(1) FAD

FADH
2
CH
2
= CH C ~ S.CoA

H
2
O O
(2) Propionolyl-CoA

CH
2
CH
2
C ~ S.CoA

OH O
-hydroxipropionyl-CoA
(3) NAD

NADH
2


CHO CH
2
C ~ S.CoA

O Aldehyd malonyl-CoA
H
2
O
(4) NADP

NADPH
2

HOOC CH
2
C ~ S.CoA

O Malonyl-CoA

(5) CO
2

CO
2

CH
3
C ~ S.CoA

O Acetyl-CoA
H
2
O
KREBS



141



Hnh 5.2 - S ca s phn gii acid bo c s nguyn t carbon l:

V. CHU TRNH GLYOXILATE
So vi chu trnh Krebs th chu trnh Glyoxilate khng c ngha
my v mt nng lng v phi tiu tn mi vng mt 2 phn t
Acetyl-CoA.
ngha ca chu trnh ny l s to thnh succinic acid, ngha l c
s chuyn ha cht bo thnh carbohydrate. V t succinic acid s hnh
thnh nn oxaloacetic acid v theo qu trnh ngc ng phn s to nn
carbohydrate. Ngoi ra t chu trnh ny s hnh thnh glyoxilic acid. Bng
cch kh carboxyl ha acid ny s chuyn thnh aldehyd formic l cht
tham gia vo hng lot qu trnh trao i cht khc. T glyoxilic acid s
tng hp c glycine tham gia to nn protein.



























H
2
O HS.C
H
2
O HS-CoA
Oxaloacetic acid
Citric acid
(1)
NADH
2
(2) H
2
O

NAD (6)
Malic acid Cis-Aconitic acid

(3) H
2
O

HS-CoA (5) CH
2
COOH

CH COOH

CH.OH.COOH
H
2
O O Isocitric acid
C
H (4)
COOH
Glyoxilic acid
CH
2
COOH

CH
2
COOH Carbohydrate
Succinic acid
Hnh 5.3 - S chu trnh Glyoxilate



Acetyl-CoA
Acetyl-CoA



142




Phn ng 1: Enzyme citratesynthetase xc tc.
Phn ng 2: Enzyme aconitate-hydratase xc tc.
Phn ng 3: Enzyme aconitat-hydratase xc tc.
Phn ng 4: Enzyme isocitrat-liase xc tc.
Phn ng 5: Enzyme synthetase xc tc.
Phn ng 6: Enzyme malatdehydrogenase xc tc.

VI. SINH TNG HP TRIGLYCERID
Do khng tan trong nc nn du khng th vn chuyn c trong
cy. V vy thc vt vic tng hp cht bo (du) phi tin hnh ti ch
trong tt c cc c quan ca thc vt. Nguyn liu tng hp cht bo l
t nhng cht ha tan c a vo cc c quan ny. D ng vt hay
vi sinh vt , thc vt, cht bo u c to thnh t carbohydrate. Trong
c th cht bo c to thnh t glycerol(P) v cc acid bo, bi vy
mun tng hp c cht bo trc ht phi tng hp c 2 cu t ny.

6.1. Tng hp glycerolphosphate
Nhng cht ban u to thnh glycerolphosphate l
aldehyd(P)glyceric hoc (P)dioxiacetone. Nhng cht ny c hnh thnh
trong qu trnh quang hp (sn phm ca chu trnh Calvin) hoc t s
phn gii carbohydrate m c (v d t qu trnh ng phn hoc t chu
trnh Pentoso(P)).
C ch hnh thnh glycerol(P) nh sau:










6.2. Tng hp acid bo theo vng xon Lynen-Wakil
- Acid bo c tng hp t acetyl-CoA trong t bo cht.
O

CH
2
OH C H CH
2
- OH
Trioso(P)- NADH
2
NAD
C = O CH OH CH - OH
isomerase Dehydrogenase
CH
2
O(P) CH
2
O(P) CH
2
O (P)
Phosphodioxiacetone Aldehyd(P)glyceric Glycerol(P)



143
- Qu trnh ny c s tham gia ca phc hp multienzyme, cu trc
ca phc hp ny nh sau:
+ gia l phn t protein-ACP (acyl-carrier-protein) khng c
hot tnh enzyme, c nhim v mang nhm acyl trong qu trnh tng hp
acid bo. ACP u tin c phn lp t E.coli do 77 gc aminoacid to
nn. ACP c coi nh mt ci nhn nm chnh gia 6 phn t enzyme
ca phc hp multienzyme. ACP gn vi mt phn t pantoten (vitamin
G) to thnh mt b phn hot ng tng t CoA c nhim v mang
nhm Acyl trong qu trnh sinh tng hp acid bo. Gc acyl trung gian
c mc vo ACP qua lin kt thioester vi nhm SH ca protein. Gc
phosphopantoten hot ng c v nh cnh tay di ng vn chuyn
gc acyl t enzyme ny n enzyme khc.
- Phc hp multienzyme c 2 nhm SH
+ Nhm SH ca ACP (SH-gia)
+ Nhm SH ca phn t enzyme (SH-bin)
sinh tng hp acid bo cn nhiu phn t acetyl-CoA t cc ngun
khc nhau v tr mt acetyl-CoA ban u i vo dng khng i, cn tt
c nhng acetyl-CoA sau u phi di dng carboxyl ha l
malonyl-CoA.




Theo Lynen-Wakil, qu trnh tng hp acid bo xy ra qua ba giai
on nh sau:
* Giai on 1: Hot ha acetyl-CoA.
T acetyl-CoA to thnh malonyl-CoA vi s tham gia xc tc ca
carboxylase cng vi s c mt ca ATP v Mn
+2
.








* Giai on 2: Ni di chui carbon trong acid bo:
Bc 1: Gn acetyl-CoA u tin vi ACP hnh thnh acyl-ACP


HOOC CH
2
C ~ S.CoA

O
SH-bin SH-bin
ACP-acyl transferase
CH
3
C ~ S.CoA + E.ACP E.ACP
HS.CoA
O SH-gia S C CH
3


O
ATP ADP+H
3
PO
4

carboxylase
CH
3
C ~ S.CoA + CO
2
HOOC CH
2
C ~ S.CoA
Biotin, Mn
+2

O O
Acetyl-CoA Malonyl-CoA



144

















Bc 2: Acyl-ACP kt hp vi malonyl-CoA hnh thnh diacyl-ACP.










Bc 3: Ngng t diacyl-ACP thnh -cetoacyl-ACP.









Bc 4: -cetoacyl-ACP b kh nh dehydrogenase c coenzyme l
NADPH
2
hnh thnh -oxybutiryl-ACP.
SH-bin S C CH
3

ACP-acyl transferase
E.ACP E.ACP O

S C CH
3
SH-gia

O Acyl-ACP
S C CH
3
S C CH
3


E.ACP O + HOOC CH
2
C ~ S.CoA E.ACP O + CO
2
+ HS-CoA

SH-gia O S C CH
3

Malonyl-CoA
O
Diacyl-ACP
S C CH
3
SH-bin
-cetoacyl-ACP-synthetase
E.ACP O E.ACP

S C CH
3
S C CH
2
C CH
3


O O O



145






Bc 5: Loi nc nh enzyme hydratase hnh thnh Crotonyl-ACP.












Bc 6: Crotonyl-ACP b kh nh enzyime dehydrogenase c
coenzyme l FMNH
2
hnh thnh Butiryl-ACP.









* Phn ng hon nguyn ca FMN:
FMN + NADPH
2
FMNH
2
+ NADP

+ Khi hnh thnh butiryl-ACP, qu trnh li tip tc nhn mt phn
t malonyl-CoA v tip tc nh trn. Mi vng quay li ni di thm mt
cp carbon cho n khi phn t acid bo c di p ng vi phn t
acid bo m t bo cn tng hp.

* Giai on 3: Kt thc tng hp acid bo: Giai on ny nh s
xc tc ca enzyme acyltransferase.
SH NADPH
2
NADP SH

E.ACP E.ACP
dehydrogenase
S C CH
2
C CH
3
S C CH
2
CH CH
3


O O O OH
-oxybutiryl-ACP
SH-bin SH-bin
H
2
O
E.ACP E.ACP
Hydratase
S C CH
2
CH CH
3
S C CH = CH CH
3


O OH O
Crotonyl-ACP
SH-bin FMNH
2
FMN SH-bin

E.ACP E.ACP
Dehydrogenase
S C CH = CH CH
3
S C CH
2
CH
2
CH
3


O O Butiryl-ACP



146








Hnh 5.4 - S SINH TNG HP ACID BO THEO LYNEN-WAKIL































SH-bin SH-bin

E.ACP E.ACP

S C (CH
2
)
n
CH
3
SH-gia
acyltransferase
Acyl-ACP O CH
3
(CH
2
)
n
C ~ S.CoA

HS-CoA O
Acyl-CoA


SH-bin Acetyl-CoA

E.ACP CO
2

ATP
S CO (CH
2
)
2
CH
3
ADP + H
3
PO
4
Butiryl-ACP
COOH
Acetyl-CoA
SH-bin SCOCH
3
CH
2


E.ACP E.ACP CO ~ S.CoA
Malonyl-CoA
SH-bin S COCH
3
SH-gia

E.ACP
HS-CoA
SH-gia CO
2


SH-bin SCOCH
3


E.ACP E.ACP

S CO CH = CH CH
3
SCOCH
3

Crotonyl-ACP diacyl-ACP
H
2
O


SH-bin NADP NADPH
2
SH-bin

E.ACP E.ACP

S CO CH
2
CH CH
3
S CO CH
2
C CH
3


OH O
-oxybutiryl-ACP -cetobutiryl-ACP
CH
3
CO ~ S.CoA
CH
3
CO ~ S.CoA



N
A
D
P













N
A
D
P
H
2






F
M
N
H
2















F
M
N

HS.CoA



147

Hnh 5.4 S sinh tng hp acid bo theo Lynen-Wakil


6.3. Tng hp triglycerid
Qu trnh tng hp acid bo ni trn to ra cc acid bo di dng
cc dn xut acyl-CoA. Trong t bo c sn glycerol(P). Qu trnh gn
acid bo c thc hin tng phn t ln lt v to thnh cc sn phm
mono, diglycerid(P) v cui cng mi to triglycerid.





















VII. SINH TNG HP V PHN GII PHOSPHOLIPID
Phospholipid l cc diglycerid lin kt vi phosphoric acid bng lin
kt ester v tr nhm hydroxyl th 3. Sau gc phosphoric acid b ester
ha bi ru cha nhm amin (choline, ethanolamine,). Cng thc tng
qut ca phospholipid nh sau:


(X) hp cht ph
R
1
; R
2
gc acid bo.

CH
2
O CO R
1


CH O CO R
2

OH
CH
2
O P = O

O (X)



CH
2
OH CH
2
O CO R
2HS.CoA
CH OH + 2R C ~ S.CoA CH O CO R
(1)
CH
2
O (P) O CH
2
O (P)
Glycerol(P) Acyl-CoA Diglycerid(P)
H
2
O
(2)

H
3
PO
4

CH
2
O CO R (3) CH
2
O CO - R

CH O CO R R
3
C ~ S.CoA CH O CO - R
HS-CoA
CH
2
O CO R
3
O CH
2
OH
Triglycerid Acyl-CoA Diglycerid




148




7.1. Sinh tng hp phospholipid
Nguyn liu cn c l: glycerine, cc acid bo v cc hp cht ph
(x). Qu trnh tng hp i hi chi ph mt lng nng lng do ATP v
CTP cung cp.
V d: Sinh tng hp phosphatidyl-choline (tn c l lecithin).
S sinh tng hp nh sau:



















7.2. Phn gii phospholipid (V d phn gii lecithin)
Qu trnh phn gii phospholipid xy ra mnh m khi ht ny mm, s
phn gii chng bng cch thy phn vi s tham gia ca H
2
O di tc
dng ca nhm enzyme c gi l phospholipase.
u tin phospholipase A xc tc cho s loi b bng cch thy phn
acid bo khi nguyn t C
2
ca glycerine; sau phospholipase B loi
acid bo khi nguyn t C
1
ca glycerine. Phospholipase C tch
phosphoryl-Choline. Cn phospholipase D thy phn Cholinephosphate.
Kt qu ca s tc dng hp nht ca cc phospholipase l to thnh cc
acid bo; glycerine t do; H
3
PO
4
v Choline.

Glycerol(P) Choline: (CH
3
)
3
N CH
2
CH
2
OH
R
1
CO ~ S.CoA ATP
2HS-CoA OH
R
2
CO ~ S.CoA ADP
Diglycerid(P) Choline(P)
H
2
O CTP
H
3
PO
4
H
4
P
2
O
7

Diglycerid Cytidyldi(P)choline

CMP
CH
2
O CO R
1


CH O CO R
2


CH
2
O (P) O
Phosphatidyl-Choline(lecithin)

Choline



149




S c trnh by nh sau:


















Cu hi
1.Vai tr v tnh cht ca cht bo?
2. Phn gii cht bo?
3. C ch ca s -oxi ha acid bo?
4. C ch v ngha ca chu trnh Glioxilate? Cho bit nhng
im khc nhau gia chu trnh Glioxilate v chu trnh KREBS ?
5. Tng hp cht bo (Triglycerid)?
6.Tnh nng lng (di dng ATP) khi phn gii, oxy ha hon
ton mt cht bo no .







Phospholipase B
CH
2
O C R
1


O
Phospholipase A
CH O C R
2


O
OH
CH
2
O P = O
O CH
2
CH
2
N(CH
3
)
3


OH
Phospholipase C Phospholipase D



150




222
Chng 6 NUCLEIC ACID V S CHUYN HO CA NUCLEIC ACID

6.1 Cu trc ca nucleic acid

Nucleic acid l nhng cht mang thng tin di truyn v chuyn nhng thng tin di
truyn ny n trao i cht. Cu to ca chng gm nhiu n phn c ba thnh phn:
base nit, phosphate v ribose hoc desoxyribose.
Base nit l dn xut ca pyrimidine hoc purine. Pyrimidine c tng hp t
carbamylphosphate v asparagine. Purine c tng hp t asparagine, glycine,
glutamine, CO
2
v formiate c hot ho bi tetrahydrofolic acid.











Adenine v guanine l nhng dn xut ca purine. Cytosine, thymine v uracil l dn
xut ca pyrimidine.



















trng thi cn bng nguyn t H ca nhm hydroxyl dch chuyn n nguyn t N.







Purine

Adenine

Guanine

C
H
C
HC
N
C
N
N
CH
N
H
C

C
HC
N
C
N
N
CH
N
H
NH
2
C

C
HC
N
C
N
N
CH
N
H
OH
Pyrimidine

Cytosine

Thymine

Uracil

CH
H
C
HC
N
CH
N
CH

C
HOC
N
CH
N
NH
2

C
HOC
N
CH
N
OH
CH

C
HOC
N
CH
N
OH

C-CH
3
Asp

Formiat

Glu-NH2
Glu-NH2
Formiat

CO2
N
N
N
N
H
Gly
CH

C
HOC
N
CH
N
NH
2
CH

C
H=C
N
CH
N
H
NH
2
223






dng ny nguyn t H ng cnh nguyn t N c th to lin kt N-glycoside vi
mt ribose.









Ribose l thnh phn ng ca ribonucleic acid (RNA). Desoxyribose l thnh
phn ng ca desoxyribonucleic acid (DNA). Desoxyribose khc vi ribose ch v
tr carbon th 2: desoxyribose c 1 nguyn t H thay cho nhm OH.
Desoxyribose c to nn t ribose sinh vt nhn chun bng phn ng kh nh
enzyme nucleosidiphosphate-reductase, phn ng kh c thc hin nh 2 nhm SH
ca enzyme. Phn ng tng qut c biu din nh sau:











ng (ribose, desoxyribose) lin kt vi base nit bng lin kt glycoside. Ngi
ta gi chng l nucleoside. S biu din nhng base nit quan trng nht v nucleoside
nh sau:
Base nit Nucleoside
Adenine Adenosine
Guanine Guanosine
Cytosine Cytidine
Thymine Thymidine
Uracil Uridine
S biu din cc nucleotide v nucleotidphosphate cng theo quy lut ny, v d:
thymidinmonophosphate hoc adenosintriphosphate. nucleoside nguyn t H ca
nhm OH nh v tr carbon th 5 c thay th bng nhm phosphate, nh vy ta
c mt nucleotide.
Nucleoside: Ribose- Base nit
Nucleotide: Phosphate - Ribose - Base nit
HO-CH
2

OH OH
OH
O HO-CH
2

OH H
OH
O
Ribose Desoxyribose
+ H
2
O
P ~O P OCH
2

OH OH
N-base
O
P ~O P OCH
2

OH H
N-base
O
SH
R
SH
S
R
S
224
ng lin kt vi base nit bng lin kt glycoside, vi phosphate bng lin kt
ester. nucleotide ny, mt nucleotide th 2 kt hp vo, y H ca nhm OH nh
v tr carbon th 3 c thay th bng mt nucleotidyl, c biu din nh sau:
















Nguyn t H ca nhm hydroxyl v tr C
3
ca nucleotide th hai c thay th
bng mt nucleotidyl tip theo, ngha l chui ko di theo hng ca ribose v tr
C
3
. Nucleotidyl l gc ca mt nucleotide, l nucleosidphosphoryl.



























Hnh 6.1 Cu to chung ca mt ribonucleic acid
O

HO-P-O-CH
2


OH
O OH
N-base
O
Nhm phosphate


O=P-O-CH
2


OH
O OH
N-base
O


O=P-O-CH
2


OH
OH OH
N-base
O
Nhm hydroxyl C
3
n

P OCH
2

O H OH
N-base
O
P OCH
2

O OH
N-base
O

HO-P=O

CH
2

O H OH
N-base
O
Nucleotide
Nucleotidyl
Dinucleotide
OH

HO-
*
P=O

CH
2

O H OH
N-base
O
O

HO
- *
P-O-CH
2


OH
O H OH
N-base
O
Nucleosidphosphoryl
225
Theo nguyn tc ny c th to nn mt chui rt di. mt u n mang mt
nhm phosphate v u kia mt nhm OH t do ca ng ribose. hnh 6.1 biu din
cu trc ca 1 si n ca RNA, y n c ln t 10
2
n 10
4
.
Nucleic acid l mt polymer, c to nn rt h thng, vi mt nhm phosphate
mt u v mt nhm OH v tr carbon th 3 ca ribose hoc desoxyribose u kia.
Gc phosphate ca nucleic acid cn cha mt nhm OH t do, nhm ny c kh nng
phn ly H
+
. V vy kha cnh ny phn t c tnh acid v ngi ta gi l nucleic acid.
Trnh t ca cc base nit ring l v thnh phn ca n trong ton b phn t l
nhng tiu chun quan trng nht ca nucleic acid. DNA cha ch 4 base nit khc
nhau: adenine, thymine, guanine v cytosine. Trnh t ca chng trong phn t DNA
gn lin vi ton b yu t di truyn ca sinh vt. Sinh vt cng pht trin cao th si
DNA ca chng cng di v trng lng cng ln. Trng lng phn t ca DNA ca
vi sinh vt khong 10
9
, ca ngi khong 3,8. 10
10
. C khong hng nghn nucleotide
trn mt phn t. Phn t di nht c bit c 10
8
nucleotide, tng ng vi di
l 3mm.
Bnh thng th phn t DNA gm 2 si chy ngc chiu nhau v qun vo nhau.
Hai si chy i song v ngc chiu nhau vi theo hng t C
5
n C
3
. Nhng base
nit hng v trc helix c bo v trc nhng nh hng bn ngoi nh chui
ribosephosphate.
Tng t polypeptide cc phn t DNA c cu trc bc mt l trnh t cc
nucleotide v cu trc bc hai l s xon ca mch kp. Trn mt vng xon c 10 base
nit. y mi base nit c mt base i din xc nh. Adenine i din vi thymine,
guanine vi cytosine. S i din ny l do cc cu hydro c to nn gia dn xut
pyrimidine (thymine, cytosine) v dn xut purine (adenine, guanine). Adenine ni vi
thymine bng 2 lin kt hydro v guanine vi cytosine l 3 lin kt ( hnh 6.2).
Watson v Crick a ra m hnh DNA gm 2 si b sung. M hnh ny c ngha
i vi cc qu trnh sinh hc c bn s c gii thch r hn di y. Lin kt gia
cc base b sung thc ra l khng bn vng nhng rt chn lc. Chng to iu kin
cho si b sung c th c tng hp t mt si ring l. Hai si ny ging nh nh v
phim.




(a) Cu trc ca DNA (b) Cu trc ha hc ca mt phn DNA
Lin kt hydrogen
0,34 nm
3,4 nm
1 nm
226
Hnh 6.2 Chui xon kp ca DNA




























Hnh 6.3 S cp i ca adenine v thymine bng 2 lin kt hydro v ca guanine v
cytosine bng 3 lin kt hydro


















Hnh 6.4 S biu din cu trc ba chiu ca desoxyribosephosphate c base nit l
thymine v adenine

Mch kp ca DNA c ng knh khong 2 nm, di c th n nhiu mm. DNA
ca Escherichia coli l mt si c 4,2 .10
6
cp base. Mch kp ca DNA l tay phi
Adenine


N
N
N
N


HNH

Thymine


C
H
3
C NH
N


O
d-Ribose d-Ribose
Guanine


C
HN
H N
N
N


O
d-Ribose
N


HNH
d-Ribose
=O

C
HN N
Cytosine

Adenine

Phosphate

Thymine

Phosphate

Desoxyribose

Desoxyribose

(a) Cu trc ca DNA (b) Cu trc ha hc ca mt phn DNA
Lin kt hydrogen
0,34 nm
3,4 nm
1 nm
227
ngha l nhng vng xon theo hng quay ca kim ng h, to nn rnh ln v nh.
Rnh ln hay nh l nhng im tc ng c bit ca enzyme. Enzyme
desoxyribonuclease I (DNAase I) kt hp tnh in vng xon nh, y c bit
nhm NH
2
tch in dng ca arginine v lysine ca enzyme kt hp vi nhm
phosphate tch in m ca DNA. DNAase I l enzyme thu phn, c kh nng thu
phn tch ra mt trong hai si. Nhng enzyme endonuclease kt hp vi mt trnh t
xc nh c 6 base nit, thu phn hai mch ca si kp. Nhng nuclease ny l nhng
enzyme gii hn, ng mt vai tr quan trng trong cng ngh sinh hc.



























Hnh 6.5 a) Mch kp DNA vi rnh ln v rnh nh
b) Xon "tay phi", mi tn ch hng xon theo kim ng h

Dng B-DNA c th chuyn sang dng A-DNA bng s kh nc, chng cng xon
theo hng kim ng h. Nhng nhm phosphate ca chng hp th nhng phn t
nc t hn dng B-DNA. Dng A-DNA rng hn v ngn hn dng B-DNA.

6.2 Phn loi nucleic acid

Bng 6.1 Gii thiu cc nucleic acid
K hiu Ngun Trng lng phn t S lng nucleotide

DNA nhn, ty th, lp th 10
9
-3,8 x 10
10
4 x 10
6
-10
8


mRNA nhn, t bo 10
8
4 x 10
3

Bt u
Rnh ln
Rnh nh
b
a
228
rRNA t bo cht, ribosome

10
8
4 x 10
3

tRNA t bo cht 25 000 80 -100

cRNA Nhn 10
6
-10
7
10
3


ribonucleic acid c 4 loi khc nhau (bng 6.1). mRNA chuyn thng tin di
truyn t DNA n trung tm tng hp protein ca t bo. Trng lng phn t khong
10
6
, bng 1/1000 n 1/10 000 trng lng ca DNA, v vy n c di ngn hn.
Nhng base nit quan trng nht l guanine, cytosine, adenine v uracil. Ngc li
DNA cha base nit thymine thay cho uracil v b sung vi adenine. mRNA c tng
hp trong nhn v mc nht nh c tng hp trong ty th v lp th.
rRNA l RNA ribosome, l ni tng hp protein ca t bo. Thnh phn ribosome
gm c cc phn t protein v cc rRNA. Cc rRNA c trng lng phn t khong 10
6

Da. Bn cnh adenine, uracil, guanine v cytosine rRNA cn cha nhng base nit khc
m phn ln c methyl ho. Nhng rRNA ca thc vt c methyl ho nhiu
hn rRNA ca ng vt. Khong 80% RNA tng s ca mt t bo l rRNA, phn cn
li l tRNA, trong khi mRNA v cRNA ch chim mt thnh phn nh.
tRNA l RNA vn chuyn aminoacid n trung tm tng hp protein. tRNA ch c
khong 80 nucleotide, v vy c trng lng phn t thp, ch khong 25 000 Da. Bn
cnh 4 base nit k trn chng cn cha nhng base nit khc c methyl ho.
V l do ny m n khng th tn ti trng thi mch kp hon chnh. Trong phn t
ca n c nhng on khng kt hp vi nhau, hnh thnh nn cu trc l c tam ip
(c ba nhnh), cu trc ny rt quan trng i vi chc nng sinh l ca chng (hnh
7.20).
RNA chromosome, ngi ta cn gi l hnRNA (heterogen nuclear RNA) ch c
sinh vt nhn chun. l dng tin cht ca cc loi RNA.

6.2 Sinh tng hp nucleic acid

6.2.1 Sinh tng hp nucleotide dng purine
Qu trnh sinh tng hp nucleotide dng purine c bt u t -D-ribose 5-P, sn
phm trung gian ca chu trnh pentosephosphate. Cc nguyn t ca base c ngun gc
t nhng hp cht khc nhau v c gn ln lt vo gc ribose.
















-D-Ribose-5-P
P-O-CH
2

HO
H
O
H
OH
H
H
OH
P-O-CH
2

HO
H
O
H
OH
H
H
O POP
-D-5-P-Ribosyl-PP (PRPP)
AMP
ATP
5-P- -D-Ribosylamine
P-O-CH
2

HO
H
O
NH
2
OH
H
H
H
AMP
ATP
P-O-CH
2

HO
H
O
H
2
C-NH
2


O=C-NH
OH
H
H
H
5-P Ribosyl Glycinamide
(GAR)
Glycine
Amino-P Ribosyl transferase
H2O
Pi
L-glutamine
L-glutamate
229














































P-O-CH
2

HO
H
O
H
2
C-NH
CHO
O=C-NH
OH
H
H
H
5-Phosphoribosyl
N-Formylglycinamide
(FGAG)
Phosphoribosyl-glycine amide
ormyltransferase
H
2
O
THF
CH
2
H
N
CH
N
H
CH
2


HC-N-R

O
10-Formyl-THF
P-O-CH
2

HO
H
O
H
2
C-NH
CHO
HN=C-NH
OH
H
H
H
5-Phosphoribosyl
N-Formylglycinamide
(FGAM)
Ribosyl-formyl-glycinamide synthase
H
2
O
ADP + P
i
ATP
L-glutamine
L-glutamate
ATP
ADP + P
i
P-O-CH
2

HO
H
O
OH
H
H
H
1(5-P Ribosyl)-5-Amino-
4- Carboxymidazole
HOOC
C N
CH
C N
H
2
N

P-O-CH
2

O

HC N
CH
C N
H
2
N

CO
2
230











































Phosphoribosyl-
formylglycine-
Amidinecycloligase
HO
H
OH
H
H
H
1(5-P Ribosyl)-
5-Aminoimidazole
(AIR)

P-O-CH
2

HO
H
O
OH
H
H
H
1(5-P Ribosyl)-
5-Amino-4-(N-
Succinocarboxamide)-
Imidazole
COOH
O
CH
2

C
HCN CN
H CH
COOH C N
H
2
N

P-O-CH
2

HO
H
O
OH
H
H
H
1(5-P Ribosyl)-
5-Imidazolecarboxamine
(AICAR)


O

C
H
2
N CN
CH
C N
H
2
N

Fumarate
Adenylo-
Succinate lyase
P-Ribosyl-
Aminoimidazole-
Succinocarboxamide
Synthase
ATP
Aspartate
Pi
CH
2
H
N
CH
N
H
CH
2


HC-N-R

O
P-O-CH
2

HO
H
O
OH
H
H
H
Inosine-5-phosphate


O

C
H
2
N CN
CH
HC CN
N

P-O-CH
2

HO
H
O
OH
H
H
H
5P-Ribosyl-5-Formamido
4-Imidazolecarboxamine


O

C
H
2
N CN
CH
O=C CN
N
H H

Cc
nucleotide
dng
purine
IMP cyclo-
hydrolase
H
2
O
P-Ribosyl-Amino-
midazolecarboxamine
Formyltransferase

10-Formyl-THF
THF
231

Hnh 6.6 Cc phn ng tng hp purine nucleotide

Ribose 5-P c hot ho bng vic tip nhn mt nhm pyrophosphate, to nn 5-
phosphoribosyl-1-1pyrophosphate (PRPP). Sau nhm pyrophosphate ny c thay
th bng mt nhm amin c ngun gc t glutamine, cho sn phm l 5-
phosphoribosylamine. Phn ng ny c thc hin do s thu phn pyrophosphate.
ng thi ribose dng chuyn sang dng . Enzyme ribose-P-pyrophosphokinase
v amido-P-ribosyltransferase xc tc cho hai phn ng, l nhng im tip cn ca cc
qu trnh iu khin.
Phn ng tip theo l gn thm glycine trc tip vo nhm NH
2
, c thc hin do thu
phn ATP. Sau N
10
-formyltetrahydrofolat gn vo mt nhm formyl. Trong mt
phn ng ph thuc vo ATP v glutamine nhm oxo C-2 c thay th bng mt
nhm imin. ATP cung cp nng lng cho vic ng vng v bin i vng imidazol
thnh vng thm 1(5-phosphoribosyl)-5-aminoimidazol (AIR). Mt phn ng carboxyl
ho xy ra khng c biotin, tip nhn CO
2
to nn dn xut carboxy. Phn t ny gn
vi aspartate bng mt lin kt amide. Sau khung aspartate- carbon c gim tr
li, gii phng fumarate. Aspartate a vo cht to thnh 1(5-phosphoribosyl)-5-
amino-4-imidazol-carboxamid (AICAR) ch nhm amine ca n. S formyl ho tip
theo bi N
10
-formyltetrahydrofolate cung cp thnh phn cui cng ca vng 6 cnh.
Vng c ng li bng cch loi H
2
O, xut hin inosin-5-phosphate. IMP khng tch
lu trong t bo, n c bin i nhanh thnh adenosine- v guanisine-
mononucleotide.

6.2.2 Sinh tng hp nucleotide dng pyrimidine

Sinh tng hp uridin-5phosphate (UMP)























H
2
N-C - OH

O



H
2
N-C - OP

O


Carbamoyl-P Carbamate
HCO
3
-
ATP

ADP + P
i
Bicarbonate Gluta-
mine
Gluta-
mate
ATP

ADP

P
i
N-Carbamoyl-L-Aspartate
CH
2

C
O=C
HO
C H
N
H
O

COOH
NH
2
COOH

CH
2


H-C-NH
2


COO H
L-Aspartate
232











































Hnh 6.7 S tng hp uridine-5phosphate (UMP)
Cc nucletide
dng pyrimidine
CH

C
O=C
N
CH
N
O

P-O-CH
2

HO
H
O
H
OH
H
Ortidine-5-P
Decarboxylase
Uridine-5-Phosphate
CO
2
(S)-4,5-Dihydro-Orotate
CH
2

C
O=C
HN
C H
N
H
O

H
2
O

COOH
Dihydro-Orotate
Dehydrogenase (mng)
CH


C
O=C
HN
C
N
O

UQH
2
COOH
Orotate
UQ

Chui h hp
-D-Ribose-5-P
P-O-CH
2

HO
H
O
H
OH
H
H
OH
P-O-CH
2

HO
H
O
H
OH
H
H
O POP
CH

C
O=C
N
C-COOH
N
O

P-O-CH
2

HO
H
O
H
OH
H
H
-D-5-P-Ribosyl-PP (PRPP)
AMP
ATP
Mg
++
PP
i
Ribosephosphate
Pyrophophokinase
Orotate P-Ribosyl
Transferase
Orotidin-5-Phosphatate
233

s tng hp purine thnh phn ca vng purine tng bc c gn vo nhm ribose,
cn s tng hp pyrimidine trc ht xut hin vng base, sau ribose c gn
vo. Nhng nguyn t ca vng c gn vo t aspartate v carbamylphosphate.
sinh vt nhn chun carbamylphosphate l mt thnh phn ca chu trnh ornithine,
nhng hai ng hng phn ng c tch ra trong khng gian. s tng hp
pyrimidine, carbamylphosphate c to nn trong t bo cht (nhng phn ng tip
theo cng xy ra y), phn ng tng ng ca chu trnh ornithine xy ra ty th.
Carbamyl-synthase tham gia vo hai qu trnh c nhng c im khc nhau: Synthase
II tham gia vo tng hp pyrimidine s dng glutamine l ngun nit v khng c
hot ho bin cu bi N-acetylglutamate, synthase ca chu trnh ornithine phn ng v
c hot ho bi NH
3
. vi khun ch c carbamyl-synthase typ II.
Phn ng tip theo c xc tc bi enzyme aspartate-carbamyl-transferase, l bc
xc nh ca sinh tng hp pyrimidine, y c tc ng ca c ch iu khin.
Bc tip theo l ng vng v gii phng H
2
O. S oxy ho tip theo to thnh orotate.
sinh vt nhn chun dihydroorotate-oxidase, nh v pha ngoi ca mng ty th,
chuyn cht kh n quinone. vi khun enzyme gn kt vi mng v phn ng vi
cc quinone. Phn ng vi 5-phosphoribosylpyrophosphate (PRPP) dn n s kt hp
vi ribose, PPi c gii phng ra. Bc cui cng ca sinh tng hp uridin-5-P l kh
carboxyl ho orotidine-5-P.
Pyrimidine-mononucleotide c bin i thnh di- v triphosphate bng s phosphoryl
ho. Uridin-5-triphosphate (UTP) c amine ho bng glutamine ( ng vt c v)
hoc NH
4
+
( E.coli) trong mt phn ng cn ATP, to thnh cytidin-5-triphosphate
(CTP). Phn ng amine ho ny tng t phn ng bin i ca xanthosin-5-
phosphate, tuy nhin Pi c gii phng thay v PPi.

6.3 Sinh tng hp DNA: DNA c kh nng t tng hp (t nhn i)

cho qu trnh sinh tng hp DNA xy ra cn phi c cc iu kin sau y
- C DNA lm khun
- C mt 4 loi desoxyribonucleotide di dng triphosphate
(Nguyn liu cho qu trnh tng hp DNA l nhng desoxynucleotidtriphosphate)
Desoxyadenosintriphosphate d ATP
Desoxyguanintriphosphate d GTP
Desoxythymidintriphosphate d TTP
Desoxycytosintriphosphate d CTP
Chng l nhng cht tng t ATP, GTP, TTP v CTP v cha desoxyribose thay
cho ribose.
- C cc enzyme DNA-polymerase xc tc
Nhng enzyme quan trng nht l DNA- polymerase, DNA-ligase v mt helicase.
Khun l hai si ca mch xon kp. S tng hp bt u bng vic thu phn lin kt
hydro ca mch kp nh mt enzyme nuclease, tch lin kt ester gia ribose v gc
phosphate ca hai si, nh vy t v tr ny mch kp c th c dui xon. V mt
nng lng y khng phi l qu trnh n gin v nh trn cp hai si c ni
vi nhau bng cc cu hydro. Trong ng nghim ngi ta ct lin kt ny khi a nhit
ln 90
0
C. dui xon cn mt phc h enzyme c bit l helicase, l mt phc h
nh v ch ch ba ca mch kp v lm dui xon DNA, trc khi c hai si dui
xon lm khun cho s tng nhng si DNA mi. nhng sinh vt khc nhau ngi ta
thy helicase c nhng c tnh khc nhau: E.coli, nm men, ng vt v thc vt bc
cao. V nguyn tc c ch tc ng ca n nh sau: si kp c dui xon v helicase
234
y ch ch ba tip tc dch chuyn. Qu trnh m xon thc cht vn cha c gii
thch cn k. Nhng lin kt hydro gia cc base nit phi c ct t. Rt c th mt
si DNA c ct ra bng thu phn lm cho xon c m ra, sau c ni li vi
nhau. Ton b qu trnh cn nng lng. Tu theo loi helicase m nng lng cn l
ATP hoc cc nucleosidtriphosphate khc, v cc desoxynucleosidtriphosphate cng
c s dng y ch ch ba dch chuyn. Nh c cc nucleosidtriphosphate m cu
hnh ca protein enzyme thay i do qa trnh phophoryl ho enzyme, qu trnh ny
tng t nh co tht c.





























Hnh 6.8 S biu din s sao chp. S m xon kp nh helicase v s tng hp
DNA hai si khun nh DNA-polymerase

Nh s 6.8 mt phn mch m xon c mt u l gc phosphate v mt u
l mt nhm OH. Nhng on tch ra ny l khun cho vic to nn mt si b sung.
Si mi ny chy theo hng ngc li vi si khun. Ngi ta phn bit gia chng 1
si leading strand l si trc v 1 si lagging strand l si sau. Si sau
c ct ra bng thu phn v sau c ni kt li v to thnh ng cong ln c
thc hin nh phc h helicase bao gm nhiu phn t protein theo hnh 6.9.
ng cong ny c tc dng lm cho si DNA mi to thnh t khun l si sau
ko di ra hng theo ch ba, tuy nhin hng ko di vn m bo t 5 3 . V vy
khi nim lagging c ngha l nhng base nit b sung cho si lagging strand c
c chm hn. dui mch xon kp th mt on ca si lagging strand c ct
ra, m xon v cc on c gi l Okazaki c tng hp. Trc khi nhng on
ny ni li vi nhau nh enzyme ligase chng phi c hot ho ngha l chng c
OH
DNA-polymerase
Helicase
DNA-polymerase
dATP
dGTP
dTTP
dCTP
P
P
P
P
HO
235
kt hp vi primer. Primer ny l mt RNA. Enzyme xc tc cho phn ng ny l
primase.
Phn ng ca primase xy ra sau khi dui xon, tip theo l phn ng polymerase,
ngha l lp rp cc nucleotide vo si mi v cui cng l phn ng ligase. Ch ch ba
chuyn dch v pha trc theo hng 5 3 ca si lagging strand. Vn tc dui
xon l 700 cp base trong 1 giy nhit 30
0
C.
Polymerase to lin kt ester gia OH v tr carbon th 3 ca si mi vi
phosphate ca d nucleosidtriphosphate mi i vo. y pyrophosphate c tch ra
v nhanh chng c thu phn to photphate v c.






























Hnh 6.9 Si trc v si sau phc h helicase-primase-polymerase

Nng lng cn to lin kt ester cha trong hai lin kt cao nng trong
d nuclotidtriphosphate. S kt hp vo khun theo nguyn tc b sung tin hnh theo
cch H ca nhm OH v tr carbon th 3 ca ribose c thay th bng 1 d nucleotide,
chnh xc hn nhm OH c thay th bi acyl ca 1 d nucleotide
(nucleosidphophoryl) (xem hnh 6.10). Chui cng ko di v tr 3 theo hng C-5
C-3 hoc t u cui phosphate n OH ca v tr carbon th 3.
S kt hp vo khun theo nguyn tc b sung, ngha l thymine s kt hp vi
adenine, guanine kt hp vi cytosine. Bng cch ny mt bn sao trung thnh ca si
khun c tng hp. Nu trong qu trnh kt hp m xut hin mt li, v d hai base
nit i din nhau khng b sung to nn khng gian m 1 d nucleotide mi vo khng
kt hp c vi nhm OH ca ribose. Polymerase c kh nng sa cha nhng li
Okazaki
Si trc
Si sau
Phc h helicase-primase-
polymerase
5 P
5 P
5 P
5 P
OH 3
OH 3
236
ny, thc cht l ct nucleotide lp sai ra v kt hp nucleotide chnh xc vo. C
ch sa cha quan trng ny lm cho t l li qu trnh nhn i l rt thp, thc ra
khong 10
9
cp base th mi c mt li. Nhng li trong qu trnh ny l nhng t bin
dn n nhng ri lon rt ln trong trao i cht. Mt khc n cng cn thit v to nn
tnh mm do thch nghi vi mi trng sng.




































Hnh 6.10 S kt hp mt nucleotide vo si DNA bng vic to lin kt ester.
Nucleosidphosphoryl thay th H OH ca d'ribose

Nhng si c to nn gm khong 1000 nucleotide, c kt hp nh cc ligase.
Enzyme ny xc tc to lin kt ester gia gc phosphate ca carbon th 5 v nhm OH
v tr carbon th 3 ca d ribose. Phn ng ny gn lin vi s phosphoryl ho, gc
phosphate do ATP cung cp. Nhng si mi c tng hp to xon vi si khun, nh
vy mch kp xut hin. Ngi ta gi s nhn i ny l bn bo tn v trong mt mch
Si DNA - CH
2

H O
Ad O
O O O

HOP~OP~OP~O

OH OH OH
CH
2

OH
O
Cyt
Si DNA - CH
2

Ad O
O

O=P~O

OH
CH
2

OH
O
Cyt
P - P
237
kp mi th mt si c t trc (bo th) v mt si mi to thnh. Hai mch kp mi
hon ton ging vi mch kp ban u. Thng tin di truyn tng ln gp i.
Vic sao chp thng tin di truyn l mt qu trnh c ngha sinh hc v trnh t cc
base trong DNA l trnh t cc ch ci, m vi nhng ch ci ny thng tin di truyn
c biu din. Trc khi t bo phn chia (mitose) DNA c nhn i, nh vy
nhng t bo con c h gen ging nh t bo m.

6.5 Sinh tng hp RNA (s phin m - transcription)


V nguyn tc tng t qu trnh tng hp DNA. y mt si DNA l khun v
cc nucleosidtriphosphate l nguyn liu v kt qu l mt phn t RNA c tng hp.
sao chp cn mt si DNA khun l si giu base pyrimidine, cc
nucleosidtriphosphate v cc enzyme, ch yu l RNA-polymerase. Nguyn liu l
ATP, GTP, CTP v UTP thay th cho TTP. Bas nit uracil b sung vi adenine trn
khun. nhm OH v tr C
3
ca ribose c kt hp vi nucleoside tip theo c s
tch ra ca pyrophosphate. Chui cng ko di u C
3
ca ribose. Phn ng do RNA-
polymerase xc tc tng t s tng hp DNA.
Nng lng cn cho s tng hp ny l s tch ra ca pyrophosphate. Trong hnh
6.11 ch ra s kt hp ca base b sung GTP vi cytosine trn mch khun vi s tch
ra ca pyrophosphate. Chui RNA ko di ngc hng vi si DNA khun. s tng
hp ny s to nn mt mch kp tm thi, gm DNA v RNA.
V vy ngi ta gi hin tng ny l lai, mch kp l phn t lai. V RNA c bn l
nh hn mch DNA, nn ch mt on tng i ngn ca DNA c c tng hp
RNA. Nh c tnh ny m mt on c c k tip nhau bi nhiu enzyme
polymerase. RNA mi c to thnh khng ch l 1 m c t 30 n 100 phn t.
RNA-polymerase trt trn on DNA v tng hp nn si RNA. bt u v kt
thc on DNA cn c nhng tn hiu ho hc, nhng tn hiu ny cho n nay cha
c bit nhiu. Tuy nhin nhng on ring bit ca DNA th hon ton xc nh, n
mang thng tin di truyn cho 1 phn t RNA xc nh.













Hnh 6.11 S tng hp RNA si giu pyrimidine

Nu si khun DNA c li, v d thiu 1 base nit, s c mt c ch sa cha.
C nhng enzyme lm nhim v nhn bit nhng v tr li trn si DNA. C ch ny rt
quan trng, nu khng th nhng thng tin v ngha c c.
sinh vt nhn s th mRNA c c trc tip t DNA. nhng on DNA m
ho cho rRNA v tRNA th trc ht nhng on RNA c tng hp nn, sau
T A C A G T
G U C A
P P P
OH
P
HO
P P ~ P
GTP
ATP
GTP
UTP
CTP
Si giu pyrimidine
238
chng s c bin i to rRNA v tRNA trng thnh. S bin i ny ch yu l
thay i cc base nit. Mt phn c methyl ho, nhng nhm OH c thay th bi
cc nhm methyl. Qua nh hng n kh nng to cu hydro ca cc base nit. S
methyl ho cc base nit xy ra sau khi sao chp c ngha i vi cu trc bc hai ca
rRNA v tRNA. DNA sinh vt nhn s nm t do trong t bo cht v ch yu dng
vng. trong ty th v lp th c cha mt lng nh DNA, DNA ny tng t DNA
ca sinh vt nhn s. Mt phn t DNA dng vng khng c im bt u v im kt
thc. c chng phi c 1 im kt hp ca polymerase. Vng bt u ny
(initiation sites) c c trng bi mt trnh t base c bit. S di y c nhng
ch in m l 4 im khc nhau cho s sao chp DNA t ty th ca ng.












sinh vt nhn chun s tng hp RNA khng thc hin trc tip trn DNA. a s
on sao chp tng i di, c gi l hnRNA hoc cRNA, sau chng c bin
i thnh mRNA, rRNA v tRNA. Qu trnh ny gi l processing (hinh 6.13). Khi
20 nucleotide u tin c lp rp tng hp mRNA th 1 m (cap) c gn vo
u si RNA v tr C
5
ca ribose. M ny gm guanine methyl ho, c kt hp
vi RNA nh ba gc phosphate. Qu trnh tng hp mRNA tip tc cho n khi to nn
trnh t base adenin-adenin-uracil-adenin-adenin. y l tn hiu cho s kt thc qu
trnh sao chp, c khong 20 base nm sau tn hiu kt thc. u cui ny mt si
polyadenine c gn vo, c t 150 -200 adenylnucleotide. Phn ny bo v cho
mRMA trc s tn cng ca enzyme trong t bo cht. M guanosine kch thch qu
trinh sao chp. Bn sao ny cha mt s on c ct ra. Nhng on c ct ri li
gn kt vi nhau theo th t. Bng cch ny mRNA c tng hp. Nhng trnh t trn
phn t mRNA c gi l exon, nhng on c ct ra gi l intron (hnh 6.12).
Qu trnh processing xy ra trc tip sau khi sao chp v ch yu l qu trnh
methyl ho. y nhm OH hoc nhm NH
2
ca base nit c thay th bi nhm
methyl. Bng cch ny kh nng to cu hydro b hn ch.
Qu trnh processing ca tRNA bn cnh methyl ho cn c nhng thay i cc
base nit, v vy m cc base nit mi nh dihydrouracil v pseudouridine xut hin.
rRNA v tRNA c tng hp trong nhn, RNA-polymerase II xc tc cho tng hp
mRNA, v RNA- polymerase III cho tRNA. Hu ht rRNA c tng hp trong hch
nhn (nucleolus) nh enzyme RNA polymerase I. C ba loi RNA u c tng hp
trong nhn t bo v c a ra t bo cht, ni m chng tham gia vo qu trnh tng
hp protein. Ba loi RNA trong t bo cht chim t l nh sau: rRNA chim 80%,
tRNA chim 15% v mRNA chim 5%.




1 2
3
Exon Intron
S ct ra ca cc exon
rrn 18 T G A A T T G A C A T A G A T A A A T C T T
rrn 26 A A G A A A A T C G T A T A A A A A T C A A
atp 1 A A A A G T A A C G T A T T A A A A A A C A
atp 2 C C C A T T T T C A T A G A G A A A G A T G
Ngc dng Xui dng
239















Hnh 6.12 S ct ra cc exon t bn sao v s gn li to thnh si mRNA


















Hnh 6.13 S biu din "processing" bn sao to mRNA, rRNA v tRNA


6.6 S chuyn ho nucleic acid
Trong qu trnh trnh trao i cht cc phn t DNA v RNA khng cn thit na s
b phn gii di tc dng ca cc nuclease thnh cc mononucleotide. Cc
mononucleotide c th c dng tng hp cc phn t anucleic acid mi, hay b
phn gii tip tc.
Nuclease xc tc cho s ct lin kt phosphodiester ca DNA (DNAse) ca RNA
(RNAse). DNAse ng mt vai tr quan trng trong s tng hp, sa cha v ti t hp
ca DNA hoc l h thng bo v, trong chng phn gii DNA ngoi lai. RNAse
tham gia vo qu trnh bin i tin mRNA thnh mRNA (RNA-processing), v iu
khin s phin m do s phn gii mRNA. Nuclease c th tch ra cc nucleotide u
cui (exonuclease) hoc ct bn trong phn t (endonuclease).

Exodesoxyribonuclease (Exo-DNAase)
U-
C-
G-
A-
U-
U-
-A
-G
-C
-T
-A
-A
hn-RNA
(nhn)
Methyl ha Phn u gn vi
GTP + polyadenyl
Methyl ha
dihydrouracil
Pseudouridine
X l
mRNA rRNA tRNA
240
Exo-DNAase phn bit hng phn gii, trong c cht l si n hoc si kp
v mono- hoc oligonucleotide (him hn) c tch ra. Mt s enzyme c th c nhiu
chc nng. Ty theo c ch phn ng m cc nuclease tch ra khi c cht hoc l gn
vi c cht qua nhiu chu k phn ng.
Enzyme exonuclease III ca E.coli c nhiu chc nng: ct c hiu nhng v tr
nht nh nh DNA-nuclease hoc l nh 3-phosphatase.

Endodesoxyribonuclease (Endo-DNAase)
Thng th hin ch yu i vi si DNA n v kp.
Endonuclease sa cha nhng v tr li, nhn bit DNA li v ct DNA mt pha
ca v tr li tch ra.
Mt c tnh c bit khc ca nhng enzyme ny l nhn bit cc trnh t base.
Mt v d l T
4
-endonuclease IV ct c hiu cytosine.
Enzyme ct gii hn (restriction endonuclease) tn ti s lng ln trong vi sinh vt.
Hn 2000 loi enzyme ny c pht hin, chng nhn bit trnh t c di t 4
n 8 nucleotide trn si kp vi chnh xc cao.
Ribonuclease (RNAse)
Ribonuclease cng phn bit hot tnh exo v endo, tc ng vo nhng u cui
nht nh (exo-enzyme) hoc nhng trnh t c hiu (endo-enzyme). Mt s nuclease
ct c DNA cng nh RNA.
Mt s phn ng dng RNAase c xc tc bng trnh t RNA (ribozyme).
Cc mononucleotide b thu phn bng cc enzyme (phosphatase, nucleosidase) to
nn cc sn phm cui cng l cc base purine v pyrimidine.
Cc cytosine c bin i thnh cc uridine tng ng bng phn ng kh amin
ho. S phn gii uracil v thymine bt u bng phn ng kh lin kt i gia v tr
th 5 v 6 v tip theo l s m vng bng phn ng thu phn. S kh amin ho v
kh carboxyl ho to nn -alanine cng nh 3-aminoisobutyrate.
Cc base purine b phn gii thnh uric acid c o thi ra ngoi c th. Qu trnh
ny in hnh cho cc ng vt c xng sng (ngi, vn ngi, chim v b st). i
vi cc ng vt khc sn phm phn gii cui cng ca cc base purine l allantoin.
Trong c th allantoin b phn gii thnh urea
Cc base pyrimidine thng b phn gii thnh urea v NH
3
.

Cu hi
1. Nhng im khc nhau v cu trc v thnh phn trong h gen ca sinh vt nhn
chun (eucaryote) v sinh vt tin nhn (procaryote)?
2. Sinh tng hp nucleic acid sinh vt nhn chun (eucaryote) v sinh vt tin
nhn (procaryote) v nhng im khc nhau gia chng?
3. Mi lin quan gia qu trnh sinh tng hp v phn gii nucleic acid vi cc qu
trnh trao i cht khc trong c th sng?


166
CHNG 7
Protein v s trao i Protein trong c
th thc vt

7.1 i cng v protein

Protein c to nn t cc gc aminoacid. Cc gc ny ni vi
nhau bng lin kt peptide. Lin kt peptide l lin kt gia nhm
carboxyl ca 1 aminoacid v nhm amin ca mt aminoacid tip theo, v
thc ra y 1 H ca nhm NH
2
c thay th bi acyl ca aminoacid.
Trong 1 dipeptide c hai, trong 1 tripeptide c ba v trong 1 oligopeptide
c t 3 cho n 10 gc aminoacid lin kt vi nhau bng lin kt peptide.
Polypeptide cha nhiu gc aminoacid.
Nhng hp cht gm nhiu gc aminoacid ni vi nhau bng lin
kt peptide c gi l polypeptide. Mi aminoacid cha 3 nhm nguyn
t (NH, CH v CO) cu to nn theo s nghim ngt sau:






Ba nhm nguyn t ny l lun lp li trong chui polypeptide, phn
cn li ca cc gc aminoacid (R) l chui bn.







Trong cc protein t nhin c mt 20 loi aminoacid khc nhau. Nu
mt chui ngn c 4 gc aminoacid (tetrapeptide) th t c 20
4
=
160 000 kh nng, ngha l c 160 000 tetrapeptide khc nhau. Nu di
1 chui gm 300 gc aminoacid, mt di m thng gp i vi nhiu
polypeptide, th ngi ta c mt s lng khng tng tng c l
H
N H
C
C
O
R
H
2
N
CH
R
CO
NH
CH
R
CO
NH
CH
COOH
R
n
Nhm amin
u cui
Chui polypeptide
Nhm carboxyl
u cui
167
10
300
kh nng. Nhng trong t nhin ch tn ti mt s lng nh cc kh
nng trn.
Nh trn nu, tt c polypeptide c cng mt trnh t CO-CH-
NH. c tnh ca tng polypeptide v ca protein c xc nh do c
tnh ho hc ca cc gc aminoacid, n c th tch in m, dng, a
nc hay k nc. Trong t nhin, tu thuc vo trnh t v thnh phn,
to nn rt nhiu i lc ho hc khc nhau, chng c ngha ln trong
cc phn ng ho hc v i vi s nhn bit trnh t aminoacid. Trong
c nhng vng gc aminoacid ch yu l acid hoc base, nhng vng khc
l nhng gc aminoacid a nc hay k nc. Nhng protein ca mng
thng c nhng gc k nc, nhng gc hng ra ngoi hoc nh vo t
bo cht l a nc. S hnh 7.1 l mt v d.


















7.2 Chc nng ca protein

Chc nng ca protein c chia ra cc nhm nh sau:
1. Chc nng enzyme
2. Chc nng cu trc
3. Chc nng d tr
T bo cht
Si polypeptide
Mng
NH
3
+

Bn ngoi
-
OOC
Thr
Al
Glu
Va
Leu
Ileu
Gly
Val
Gly
Al
His
168
4. Chc nng vn chuyn
5. Hormone
6. Chc nng bo v
7. Chc nng co rt
8. Cht c

V chc nng enzyme ca protein c cp nhiu ln, c bit
trong cu to v c ch tc ng ca enzyme. Protein l cht xy dng
ng mt vai tr c bit quan trng trong c th ng vt. Cht khung
c to nn t protein. Thuc loi ny gm c da, tc, sng, lng v
collagen l thnh phn c bn ca gn v c khong 18% tham gia vo cu
to ca xng.
Mt chc nng ph bin khc ca protein l cu to nn mng sinh
hc.
Mng sinh hc c chc nng l gii hn nhng vng trao i cht v
tham gia vo vic vn chuyn cc cht. Mng sinh hc cng c kh nng
chuyn i nhng tn hiu. Protein mng cng c th l cc enzyme. Chc
nng ny c th hin mng trong ca ty th v lp th. Mng sinh hc
bao gm lp kp lipid vi nhng protein phn b trong . Nhng thnh
phn lipid quan trng nht l phosphoglyceric, glycoglyceric, sterol v
sphingolipid. Lp kp lipid ca mng sinh hc l nhng phn t lipid c
nhng ui k nc quay li vi nhau, trong khi nhng gc ng v
phosphate hng ra pha ngoi hnh 7.2).
y thnh phn glycerine l trc vung gc vi mt phng ca
mng. Lp kp lipid khng i xng, ngha l hai pha ca mng c
nhng phn t lipid khc nhau. Nhng phn t protein nm trong lp kp
lipid lm bn vng mng v nh ra hai pha mng hnh 7.2 . Thnh
phn protein nm trong lp kp lipid, c c tnh ch yu l k nc,
thnh phn protein m nh ra pha ngoi, c c tnh ch yu l a nc.
Si protein nh ra pha t bo cht thng kt hp vi mt protein ngoi
vi, nhng phn t protein hng v pha mng t bo th thng kt hp
vi mt chui carbohydrate hnh 7.2 .






169














Hnh 7.2. S biu din mt on ct ca mng sinh hc

trong lp kp lipid th phn t protein c nhng lin kt k nc
v lin kt ion. Nhng lin kt ion nh lin kt gia nhm NH
3
ca
protein v ca gc phosphate ca phospholipid. Nhng lin kt ny nhy
cm vi pH. V vy mng sinh hc b nh hng, thm ch b ph hu
bi gi tr pH thi qu. Mc qunh c nhiu hay t ca mng sinh hc
b nh hng ln bi nhit . nhit thp mng sinh hc c cu trc
ging nh tinh th. Khi nhit tng ln n nhit chuyn pha cu trc
rn s chuyn sang cu trc tinh th lng. Nhit chuyn pha tng theo
di v no ca acid bo. Ca
2
lm n nh mng, n lin kt vi
nhng nhm phn u ch yu tch in m, v d gc phosphate ca
phospholipid. Ca
2+
c kh nng to lin kt vi hai nhm tch in m. V
vy Ca
2
s lm gim tnh thm ca mng, trong khi nhng ion ho tr
mt, c bit l H lm tng tnh thm. Tnh thm cng b nh hng bi
c tnh ca lipid. Thnh phn steroid v nhng gc acid bo no v c
mch carbon di lm cho mng st li, nhng gc acid bo cha no lm
cho mng lng lo ra, do vy lm tng tnh thm ca mng.
Khi ngi ta ni n tnh thm ca mng ch yu l ni n tnh
thm ca nhng cht a nc. Nhng cht ny thm qua mng rt t, trong
khi nhng phn t k nc c th thm qua hoc khuch tn qua mng.
Hu ht nhng cht trao i c c tnh a nc, i vi chng mng sinh
Lipid

Protein

Protein phn b trong mng
u a nc
u k nc
Oligosaccharide
Lp kp
phospholipid
Vng k nc
ca protein
Vng a nc
ca protein
170
hc l ci chn ng k v y chng cn mt c ch c bit hn
vn chuyn qua mng.
Protein d tr c tch lu trong m d tr v khi cn s dng th
c huy ng. Protein d tr in hnh ng vt l protein trng v
sa. l protein cn thit i vi ng vt cn non, tng t protein ca
ht v qu, l ngun dinh dng i vi cy con.








Hnh 7.3 S ct ngang ca ht ng cc

Cht lng protein c nh gi theo thnh phn cc aminoacid
khng thay th. l nhng aminoacid m c th ngi v ng vt
khng c kh nng tng hp c. Hm lng cc aminoacid khng thay
th ca cc loi protein khc nhau c trnh by bng 7.1 v 7.2

Bng 7.1 Nhng aminoacid khng thay th i vi ngi
--------------------------------------------------------------------
Valine Lysine Phenylalanine
Leucine Methionine Tryptophan
Isoleucine Threonine

Bng 7.2 Hm lng aminoacid khng thay th ca cc loi protein
khc nhau (%)
------------------------------------------------------------------------------------------
Protein trng 51 Protein u tng 40
Protein sa 50 Protein go 39
Protein m c 47 Nm 35
Collagen 17 Protein la m 33
Protein l cy 40 Protein ht lc 32
------------------------------------------------------------------------------------------

Ni nh
Lp aleuron
Phi
171
Nhng protein ng vt, v c protein ca l cy phn ln l giu
aminoacid khng thay th. Protein ca ht v ng cc cha t aminoacid
khng thay th. protein ng cc l lysine, protein ht v u cc loi
l methionine c hm lng thp v n gii hn gi tr sinh hc ca nhng
loi protein ny.
s gii c cc kim loi nng nh nhng polypeptide n gin
theo kiu phytochelatin c ngha quan trng. V d polypeptide n gin
c ngun gc t glutation v c cng thc chung nh sau:
( -glutamyl-cysteinyl)
n
-glycine
Do c nhiu nhm SH chng c kh nng kt hp cht vi cc kim
loi nng, lm cho nhng kim loi nng ny khng th gy ri lon trao
i cht. S tng hp phytochelatin c kch thch bi nhng kim loi
nng nh Cd, Cu, Ag, Bi v Au.
Protein bo v c mt vai tr ln trong sinh hc min dch. ng vt
c xng sng c mt c ch phc tp, pht trin cao, vi c ch ny
chng ngn nga nhng tc nhn vi sinh vt gy bnh (virus, vi khun,
nm, cht c vi khun). Chc nng ny c phn lin quan n c tnh
ca chui polypeptide. H thng t v ton b, sinh hc min dch l mt
lnh vc khoa hc pht trin c lp. Mt protein l (virus, vi khun, nm)
xm nhp vo mu hoc vo m th c ch t v c huy ng rt nhanh.
Protein l c gi l antigen. N c mt vng gm mt trt t xc nh
cc nguyn t, vi vng ny n kt hp vi t bo lympho v kch thch t
bo ny sn sinh ra khng th. Nhng t bo lympho tn ti trong h thng
min dch vi s lng 10
9
v c trn b mt ca n nhng vng nhn, ni
m antigen c kt hp vo. Nhng vng nhn ny rt khc nhau v
ph hp mi vng cho 1 antigen xc nh. Nhng tc nhn khc nhau c
nhng t bo lympho xc nh khc nhau vi nhng vng nhn ph hp.
Khi mt antigen kt hp vi t bo lympho th n bt u sn sinh khng
th c hiu i vi tc nhn gy bnh. Nhng t bo lympho khc khng
c kch thch cho vic sn sinh ra khng th. C sn mt s lng ln
cc t bo lympho khc nhau, chng c th tng hp c rt nhanh
nhng khng th khc nhau khi khng nguyn xut hin. Nhng loi
khng th khc nhau ny l xc nh, tn ti vi s lng khng m
c, c th mt vi triu, y mi mt loi c mt v tr kt hp duy
nht c trng. Kh nng ln khng th tng tng c ca h thng
min dch lm cho protein l, protein ca tc nhn gy bnh tr thnh
v hi. Nhng khng th ny c gi l globulin min dch. Chng
chim khong 20% protein tng s trong mu.
172
Globulin min dch Ig c chia lm 5 nhm khc nhau. Tuy nhin
thnh vin c bn Ig cc nhm u c dng ch Y. Cu trc ny gm 2 si
polypeptide ngn, xc nh, mi si c khong 220 gc aminoacid (hnh
7.4).






















Hnh 7.4: S biu din khng th v khng nguyn.
a) Khng th gm 4 chui polypeptide
b) Khng th kt hp vi khng nguyn
c) Kt hp gia khng nguyn v khng th

C ch m khng th nhn bit v tr khng nguyn ca protein l l
c bit th v. S kt hp gia khng th v khng nguyn l thun
nghch v c th so snh vi s kt hp gia enzyme v c cht.
NH
2
NH
2
NH
2
COOH
COOH
HOOC
HOOC
H
2
N

Ch kt hp vi khng
nguyn (antigen)
Ch kt hp vi t bo
phage (cytophage)
NH
2
NH
2 H
2
N

H
2
N

Khng nguyn
Khng nguyn
Khng th
a.

b.

c.

173
Lin kt ny khng phi ng ho tr m l nhng lin kt hydro v
lin kt ion. iu lm r rng trnh t aminoacid ca v tr kt hp xc
nh tnh c hiu ca lin kt. T rt ra rng chui polypeptide ngn
hay di ca khng th l mt on siu bin c t 20-30 gc aminoacid,
ngi ta cho rng, trong vng siu bin ny cha v tr lin kt c hiu
i vi khng th. Ch kt hp thc s ch gm 5-10 aminoacid. V tr kt
hp rt chn lc i vi khng nguyn.
V d protein vn chuyn l albumin v hemoglobin trong mu,
myoglobin trong m c v leghemoglobin trong mng ca Rhizobium.
Albumin vn chuyn acid bo. Hemoglobin, myoglobin, leghemoglobin
c kh nng kt hp lng lo vi O
2
heme ca chng. Chng ging nhau
ch u l phng tin vn chuyn O
2
. S kt hp ny ch xy ra
dng Fe kh (Fe
2+
), ngha l kt hp vi heme. Hemoglobin vn chuyn
O
2
t phi qua mu n nhng c quan v m rt khc nhau.
Leghemoglobin iu khin s cung cp O
2
cho chui enzyme h hp ca
vi khun. Myoglobin vn chuyn O
2
trong m c, c nhiu trong m c
ca ngi ln v l cht d tr O
2
. Trong khi ln (khi dng th) th O
2

c kt hp vi myoglobin, c gii phng ra ty th. Fe
2+
ca heme
c th kt hp vi CN
-
v CO lm cho v tr hp th O
2
b ng li, hu
qu l O
2
khng c vn chuyn, dn n cht ngt. Nng tng i
cao CO trong khng kh cng nh s t chy khng hon ton trong
ng c, v khi ht thuc l, cn tr s vn chuyn O
2
trong mu. Ngi
ta gi l ht b ng, l nguyn nhn gy hi n sc kho
V d v chc nng hormone ca protein l insulin, gm 1 chui A
vi 21 aminoacid v 1 chui B vi 30 aminoacid, hai chui ny c ni
vi nhau qua hai cu disulfide. Insulin iu khin nng ng glucose
trong mu. Khi khng insulin th s tip nhn ng trong t bo b
hn ch. V vy mc ng trong mu tng v dn n s thi ng
mnh m qua nc tiu (bnh i ng). V vy nhng t bo ny thiu
ng lm cho ton b c th suy yu.

Mt s peptide quan trng trong t nhin:
- Glutation (tripeptide): glutamyl-cysteinyl-alanine: l mt
tripeptide c chc nng sinh l quan trng. N c to nn t 1 gc
glutamyl, 1 gc cysteine v glycine.



174











Hai nguyn t glutatione c kh nng phn ng tch 2H to thnh
1 disulfide. y l h thng m sulfhydryl, chc nng quan trng nht
ca n l gi nhm SH ca enzyme v coenzyme trng thi kh.
Glutathione c trong tt c cc c th sng, tham gia vo nhiu phn ng
oxy ho kh, v d s gii c cc gc c. Ngoi ra trong thc vt
glutathione cn l dng d tr v dng vn chuyn lu hunh dng kh.
Trong nhng loi c c nhng tripeptide tng t nh glutatione v trong
cy h u 1 glutamyl-cysteinyl-alanine c tm thy.















COOH

H
2
N-CH

CH
2


CH
2


CO

-Glutamyl
NH CO CH

CH NH COOH

CH
2

SH

Glycine Cysteinyl
H

175

























Nhng dng tng t c chc nng nh glutatione. Cysteine v
cystine v nguyn tc to nn h thng oxy ho kh tng t glutatione,
biu din nh sau:







Glu NH CH CO Glycine

CH
2
- SH

Glutathion
dng kh
CH
2
- SH

Glu NH CH CO Glycine

Glu NH CH CO Glycine

CH
2
- S

CH
2
- S

Glu NH CH CO Glycine

C cht
C cht H
2
Glutathion
dng oxy ha
Cu disulfide
Glu NH CH CO Serine

CH
2
- SH

Glu NH CH CO Alanine

CH
2
- SH

COOH

H
2
N-CH

CH
2
S H
2x Cysteine
COOH

H
2
N-CH

H S CH
2

COOH

HN-CH

CH
2
S
COOH

H
2
N-CH

S CH
2

C cht
C cht H
2
Cystine
176
Oxytocin v vasopressin: Oxytocin v vasopressin l nhng
oligopeptide, chng c cu trc rt ging nhau, u bao gm 9 gc
aminoacid khc nhau, trong phn t c mt cu disulfide. Hormone
oxytocin kch thch phn c phng ca rut gi, bng quang, ti mt v t
cung. Hormone vasopressin nh hng n co rt ca phn c phng ca
mch mu, lm tng p huyt. Vasopressin c vai tr quan trng trong
vic lm c nc tiu. Nu nng thp th ng vt c v thi ra 1
lng ln nc tiu long, l mt bnh c gi l bnh tho nht.













7.3 Cc bc cu trc ca protein

Phn ln cc lin kt c trong chui polypeptide c th quay t do
v trc ca 1 chui polypeptide rt linh ng. Tu thuc vo lc tc ng
m chng c nhng dng rt khc nhau. S bin dng ca mt chui
peptide, s sp xp ca nhng phn si, s un cong v nhng np gp c
trong mt chui peptide, c gi l cu trc.
thun tin ngi ta thng phn bit cu trc ca phn t protein
thnh 4 bc nh sau: bc I, bc II, bc III v bc IV.
Cu trc bc I l trnh t sp xp cc gc aminocid trong chui
polypeptide. Cu trc ny c gi vng nh lin kt peptide (lin kt
cng ho tr).
V mi mt aminocid c gc khc nhau, cc gc ny c nhng c
tnh ho hc khc nhau, nn mt chui polypeptide cc iu kin khc
nhau c nhng c tnh ho hc rt khc nhau.
Gly - NH
2


Leu

Pro NH
2

Cys Asp

S Glu NH
2


S

Cys Ileu
Gly - NH
2


Arg

Pro NH
2

Cys Asp

S Glu NH
2


S

Cys Phe
Oxytocin Vasopressin
177
Tuy nhin v tng qut th tt c cc si polypeptide c xy dng
mt cch c h thng t cc nhm nguyn t CO, CH v NH. S xy dng
c h thng ny l c s to nn cu trc bc hai.
Cu trc bc II l tng tc khng gian gia cc gc aminoacid
gn nhau trong mch polypeptide. Cu trc c lm bn ch yu nh
lin kt hydro c to thnh gia cc lin kt peptide k gn nhau, cch
nhau nhng khong xc nh.
Cu trc bc hai ca phn t protein: xon ( -helix), phin l gp
v xon collagen. Loi -helix l si dng xon c, qun xung quanh
mt trc, mi vng xon c 3,6 gc aminoacid (hnh 7.5).
y do nhm CO v nhm NH ca aminoacid th t trn chui
gn nhau, gia hai nhm ny c th to nn mt cu hydro. Cc lin kt H
tng i lng lo v c th c ct t khi nhit cao. Phn t protein
mt cu trc bc hai th enzyme mt hot tnh xc tc.
-helix l cu trc ca hu ht nhng protein mch thng (protein
si). Dng si tn ti trong -keratin ca tc, lng, mng v da t chui
polypeptide c cu trc . Cu trc ny trong phn tch Rnghen l nhng
chu k 0,5-0,55 nm. Khi x l bng hi nc ngi ta c th ko di
chui, lc ny xut hin chu k l 0,7 nm.
Cu trc tn ti trong -keratin c gi l cu trc phin l gp.
Nh trong hnh 7.6 chng gm nhng si polypeptide i song, gn vi
nhau bng cc lin kt hydro. Khc vi -helix, nhng lin kt hydro
c to nn khng phi trong mt si m gia si ny vi mt si khc.
Nh vy c th nhiu si kt hp vi nhau to phin l gp. Cu trc
phin l gp tn ti trong si la, nhng cng c trong nhng vng ca
protein hnh cu.
178


Hnh 7.5 S biu din mt -Helix. Mi tn ch lin kt H gia CO v H

Thuc protein thng cn c collagen, gm 3 si polypeptide xon vo
nhau, chng li kt hp nhiu si li vi nhau bng lin kt ng ho tr
(hnh 7.7). i phn t ny ch c mt s cu hydro c to nn, v ngoi
glycine n c to nn ch yu t proline v hydroproline. Khi mt
aminoacid i vo to lin kt peptide, trong chui peptide cn li 1 N l
yu t cu to ch khng phi l -NH-, nh c ch ra trong s di .
y 1 H ca nhm NH c thay th bng 1 aminoacyl, ngha l thiu H
cho vic to lin kt hydro v v l do ny m si peptide khng th t xon
c.








179












Nhng si collagen chy song song to nn nhng b si dai ca
gn. Collagen cng c trong xng v trong cc m ni. Elastin l mt
protein, gm nhng si protein tng i ngn, gn kt vi nhau nh lin
kt ng ho tr (hnh 7.7). Nhng si polypeptide quay theo dng xon
c, t dui xon khi c p lc (hinh 7.8). Vt liu ny to nn mt khi
dng keo dnh, cho php quay trong khng gian ngoi t bo v s co rt
ca cc m.













Hnh 7.6 Nhng chui polypeptide chy i song trong cu trc -keratin
(phin l gp)
H
2
N CO

CH

R
COOH

H N CH

H
2
C CH
2

CH
2
COOH

N CH

H
2
C CH
2

CH
2
H
2
N CO

CH

R
Aminoacyl
Proline
CH
R
CH
R
CH
R
CH
R
CH
R
CH
R
R
CH

R
CH

R
CH

R
CH

R
CH

R
CH

N
H
N
H
N
H
N
H
N
H
N
H
C
O
C
O
C
O
C
O
O
C

O
C

O
C

O
C

O
C

O
C

C
O
C
O
H
N

H
N

H
N

H
N

H
N

H
N

180









Hnh 7.7 S biu din mt si collagen bao gm ba si ring l












Hnh 7.8 Cu trc Elastin khi khng c v c lc tc ng

Nhng v d ny ni ln rng polypeptide c nhiu c tnh v chc
nng khc nhau.
Vic xc nh cu trc bc I ca phn t protein c ngha quan trng:
- L bc u tin quan trng xc nh c s phn t hot tnh
sinh hc, tnh cht ho, l ca protein.
- L c s xc nh cu trc khng gian ca phn t protein da
vo cu trc khng gian ca cc phn t protein tng ng.
- Cu trc bc I l bn phin dch m di truyn. V vy cu trc
ny ni ln quan h h hng v lch s tin ho ca th gii sinh
vt.
- L yu t gp phn quan trng trong nghin cu bnh l phn
t. Kt qu nghin cu cho thy: khi thay i th t aminoacid,
thm ch ch mt gc aminocid trong phn t protein c th lm
thay i hot tnh sinh hc, gy nhng bnh c trng. V d
in hnh l bnh thiu mu hng cu hnh li lim, l do cu
Khng c lc
tc ng
Khi c lc
tc ng
181
trc bc I ca hemoglobin thay i: gc aminocid v tr th 6
trong chui ca hemoglobin A (bnh thng) b thay th bng
gc aminoacid valin.
- L c s tng hp nhn to protein bng phng php ho
hc hoc bng phng php cng ngh sinh hc. Frederick
Sanger (1953) ra phng php v s dng phng php
ny thnh cng xc nh trnh t sp xp cc aminocid trong
phn t insulin (polypeptide c hot tnh hormone). n nay rt
nhiu protein c xc nh cu trc bc I. Insulin l protein
u tin c tng hp bng phng php ho hc vo nm
1966. Ngy nay bng phng php cng ngh sinh hc ngi ta
s dng E.coli tng hp insulin.

Cu trc bc III l tng tc khng gian gia cc gc aminacid xa
nhau trong chui polypeptide, l dng cun li trong khng gian ca ton
mch polypeptide.
Nhiu si polypeptide trong c th sng tn ti khng phi dng
thng m gp khc v qua m to nn cu trc khng gian ba chiu.
Tuy nhin cu trc ny hon ton xc nh, ch yu l do trnh t cc
aminoacid v mi trng. Khi mt si polypeptide tch ra khi ribosome
sau khi tng hp v c thi ra trong t bo cht nh l mi trng to
hnh th n hnh thnh nn cu trc t nhin rt nhanh, c bit i vi cu
trc hnh cu, mang li cho protein nhng c tnh sinh l quan trng. C
th do chuyn ng nhit ca cc si polypeptide m cc nhm ca cc
gc aminoacid tip xc vi nhau, dn n c th kt hp vi nhau. Trong
nhiu protein hnh cu c cha cc gc cysteine, s to thnh cc lin kt
disulfide gia cc gc cysteine xa nhau trong chui polypeptide, lm cho
mch b cun li ng k. Cc lin kt khc, nh lin kt Val de Waal, lin
kt tnh in, phn cc, k nc v hydro gia cc mch bn ca cc gc
aminoacid u tham gia lm bn cu trc bc III, nh protein hnh cu
biu din hnh 7.9.







182











Hnh 7.9 S biu din mt protein hnh cu cha nhng on c cu trc
-helix v phin l gp
Cu trc hnh cu ca protein c gi l cu trc bc ba, l cu trc
ca enzyme. chng mt nhm prosthetic c th kt hp ng ho tr, v
d heme. Nhng nhng nhm gc aminoacid ring l cng c th l cc
nhm hot ng trong phn ng enzyme. S ho tan ca protein c xc
nh mt mc nht nh nh cu trc bc ba. nhiu protein hnh
cu cc nhm k nc nh hng vo bn trong, nhng nhm a nc
hng ra ngoi. Nhng nhm a nc ny kt hp vi nc nh l cht
ho tan bng cc cu hydro, phc h ny tn ti trong dung dch. Trnh t
aminoacid ca protein ca nhng enzyme cng chc nng cho bit mc
quan h h hng ca cc loi. Mc quan h h hng cng gn, th mc
tng ng v trnh t aminacid cng ln. iu rt th v y l nhng
on trnh t quan trng nht, v d nhng vng phn ng, trong qu trnh
tin ho hu nh khng thay i, c bo tn v nh vy cu trc bc
hai, bc ba c duy tr.
Cu trc bc IV: Cu trc ny c hnh thnh cc phn t protein
gm 2 hay nhiu chui polypeptide hnh cu. Tng tc khng gian gia
cc chui ny trong phn t gi l cu trc bc IV. Mi chui polypeptide
ny gi l phn di n v (subunit). S kt hp gia cc phn t ny
lng lo v ch yu l do lin kt hydro v k nc. Bng cch ny hai
phn t xc nh c th kt hp vi nhau to thnh 1 dimer. Mt th d v
s kt hp ny l hemoglobin, c to nn t 2 chui vi mi chui c
141 v 2 chui vi mi chui l 146 gc aminoacid. i phn t dng
ny bn cnh protein cn l thnh phn cu to ca nucleic acid, cc
ribosome. Cc ribosome ca sinh vt tin nhn gm 55 si polypeptide
Phin l gp
NH
2
HOOC
Xon
183
khc nhau v 3 nucleic acid khc nhau kt hp li. Nhiu virus c lp v
bn ngoi, c cu to t nhiu phn t protein xc nh v bao quanh
nucleic acid xon c bn trong. Cc i phn t trn kt hp vi nhau t
ng trong mi trng ph hp thnh dng tn ti trong t nhin.
c tnh vt l ca protein ph thuc vo c tnh ho hc ca cc
gc aminoacid protein hnh cu cng nh s gp khc ca chng.
Chim u th trong s kt hp ca cc aminoacid c tnh acid th protein
c tnh acid. Ch yu l aminoacid c tnh kim l protein kim. Nhng
si protein c nhiu nhm a nc l do nhng aminoacid phn cc v
hydroxyacid th c ho tan ln. Ngc li, thnh phn ca gc k nc
(valine, leucine, isoleucine) th ho tan ca nhng protein ny thp.
Trng lng phn t protein ph thuc vo di ca tng chui
polypeptide v s lng cc si polypeptide cu to nn 1 phn t protein.
V kha cnh ny bng 7.3 ch ra s khc nhau ng k gia cc phn t
protein ring l.

Bng 7.3 Trng lng phn t v s lng si polypeptide ca mt s
phn t protein
MG S lng S lng
gc aminoacid chui polypeptide
Insulin 5 733 51 2
Ribonuclease 12 640 124 1
Myoglobin 16 890 153 1
Hemoglobin 64 500 574 4
Glutamate dehydrogenase 10
6
8 300 40
Synthetase tng hp acid bo 2,3 x 10
6
20 000 21
------------------------------------------------------------------------------------------
Cu trc ca mt hoc nhiu chui polypeptide c ngha quan
trng i vi ho tan v chc nng ca chng. Cu trc protein c
hiu l s sp xp ca nhng si ring l hoc nhiu si. Chng ph thuc
nhiu vo pH ca mi trng. Protein v chui polypeptide ho tan tt
khi nhng nhm a nc hng ra pha ngoi, nhm k nc hng vo
bn trong. Khi mt protein thay i cu trc th nhng nhm k nc quay
ra ngoi, protein mt kh nng ho tan trong nc, v d trng hp kt
ta khng dng tinh th ca protein sa trong mi trng chua. Acid
lactic c sn sinh do vi khun lm gim pH sa, lm thay i protein
sa. Nhiu nhm k nc c hng ra bn ngoi, protein mt kh nng
tan trong nc. V v s thng xuyn duy tr gi tr pH trong t bo cht
184
rt quan trng, v ch c nh vy chc nng hot ng ca cc enzyme
trong t bo cht mi c m bo.
Bn cnh H
+
cn c nhng cation khc c ngha i vi cc phn
ng enzyme v d K
+
. Nng ca n trong t bo cht cao hn so vi
cc loi cation khc v nm trong khong 100 n 150 mM. Nng K
+

cao c ngha i vi s tng hp protein.
Nng Ca
2+
trong t bo cht rt thp, tuy nhin n c vai tr quan
trng i vi cu trc ca cc protein khc nhau. Calmodulin l mt
polypeptide kt hp vi Ca
2+
lm cho cc vng k nc c hng ra
ngoi, cc vng ny kt hp vi cc enzyme v hot ho cc enzyme nh
phosphodiesterase, adenylate-cyclase, photoarylase v ATPase. Bng cch
tng t troponin C v paralbumin hot ho nhng enzyme khc bng
vic thay i cu trc khi kt hp vi Ca
2+
. Nhng Ca-protein ny thay i
phn ng enzyme. Nhng lin kt thc hin nhng v tr hon ton xc
nh, thc cht l helix-di ni-helix (hnh 7.10).











Hnh 7.10 S biu din lin kt Ca trong chui polypeptide ni hai cu
trc helix li vi nhau

Dng ny ca mt lin kt Ca tn ti tng i v c trng i vi
calmodulin, troponin C v paralbumin. Cu trc ny bao gm mt chui
polypeptide c 12 gc aminoacid, trong 7 nhm carboxyl c nguyn t
O kt hp vi Ca
2+
. Cu trc ny tng ng vi mt hnh thp i 5 mt,
ngha l mt din tch c bn vi 5 mt, pha trn v pha di cn c
mt CO. trung tm cu trc khng gian ba chiu ny c nguyn t Ca
2+
,
mi mt gc c 1 nhm CO (hnh 7.11).
Lin kt vi Ca
2+
trung tm lm thay i cu trc ca protein.
Chui xon (helix)
Chui xon (helix)
Di ni
Ca
++
185







Hnh 7.11 Mt phng c hnh 5 gc ca mt khi chp i vi Ca
2+
trung
tm

Troponin C c tng s 4 v tr kt hp vi Ca
2+
, trong 2 lin kt
c i lc mnh v 2 lin kt c i lc yu. S bo ho Ca
2+
ph thuc vo
nng Ca
2+
trong h thng. Cng nh vy i vi calmodulin. Nng
Ca
2+
cng nh hng n nhng qu trnh enzyme bng nhng Ca-protein.
Vai tr quan trng nht ca troponin C l iu khin s co rt ca m c.
Paralbumin cng c mt trong m c, c ngha i vi th gin m c.
V tr kt hp ca n vi Ca
2+
cng c i lc cao vi Mg
2+
.

7.4 Aminoacid

Phn ln nht ca nhng cht nit hu c d ho tan trong thc vt
l aminoacid, amide v amine. Trong trao i cht th aminocid c vai tr
rt quan trng. Chng l vt liu to nn protein. Cng thc chung ca
aminoacid:
COOH

H
2
N -CH

R
Cng thc tng qut ca L- -aminoacid

Nhm amine nh nguyn t C
2
, theo tn c l nguyn t C . V
vy ngi ta gi l nhm -amine. Cc aminoacid tn ti ch yu trong t
nhin c nhm amine ng bn tri trc, c gi l aminoacid dng L.
Dng D-aminoacid tn ti ch ring bit, v d trong thnh t bo vi khun.
Mi mt aminoacid t nht c c trng bi 1 nhm carboxyl v 1
nhm amine. Hn 100 loi aminoacid khc nhau m ngi ta bit, c
khong 20 loi l thnh phn ca protein. Di y l nhng aminoacid
quan trng v cng thc ho hc ca chng. n gin v theo thi
quen nhm carboxyl ca aminoacid proton ho. Tuy nhin dng ny
C
O

CO Ca
2+
CO

O O
186
khng c aminoacid no tn ti, v nh nu trn, khi nhm carboxyl
phn ly, th nhm NH
2
phi phn ly. Proton ho hay kh proton ho lun
lun ph thuc vo pH ca mi trng.
Cc aminoacid ring bit c nhng c tnh khc nhau l do phn
gc aminoacid. nhng aminoacid trung tnh c tnh trung tnh. Khi
chiu di chui tng th c c tnh k nc. Protein c cha nhiu
aminoacid nh valine, leucine, isoleucine c c tnh k nc l ch yu.
Nhng aminoacid c tnh acid trong phn gc c 1 nhm carboxyl. Protein
cha nhiu aminoacid c tnh acid l nhng protein acid. Tng t nh
vy i vi protein, ch yu c to nn t nhng aminoacid c tnh
kim l nhng protein kim. Phn gc ca aminoacid c ngha quyt
nh i vi c tnh ca protein m chng to nn. iu ny khng
nhng c ngha i vi tnh cht ho hc m c cu trc ca protein.

7.4.1 Phn loi aminoacid
Thy phn hon ton protein, thu c ch yu cc L- -aminoacid.
Mc d protein rt a dng nhng hu ht chng u c cu to t 20
L- -aminoacid. Da vo c tnh ca mch bn (gc R) aminoacid c
chia lm 7 nhm chnh sau y:
1. Aminoacid trung tnh mch thng
Nhm ny gm 5 aminoacid: glycine, alanine, valine, leucine,
isoleucine. Cc aminoacid ny u c 1 nhm amine v 1 nhm
carboxyl.
Glycine l axit amine n gin nht, nhm R ca n l nguyn t H, l
aminoacid duy nht khng cha carbon bt i. Bn aminoacid cn li
c mch bn khng phn cc.

Aminoacid trung tnh











COOH

H
2
N-CH

CH
3

Alanine
(Ala)
COOH

H
2
N-CH

H
Glycine
(Gly)
COOH

H
2
N-CH

CH

CH
3
CH
3
Valine
(Val)
COOH

H
2
N-CH

CH
2


CH

CH
3
CH
3
Leucine
(Leu)
COOH

H
2
N-CH

CH

CH
3
CH
2



CH
3
Isoleucine
(ILeu)
187

2. Cc hydroxyl aminoacid mch thng
Thuc nhm ny c 2 aminoacid l serine v threonine. Chng ging
vi nhm trn ch ch c 1 nhm amine, 1 nhm carboxyl v cng l
mch thng nhng c cha 1 nhm OH.

Aminoacid c nhm hydroxyl










3. Aminoacid cha lu hunh mch thng
Nhm ny c 2 aminoacid l cysteine v methionine. Khi oxy ho 2 nhm
-SH ca 2 phn t cysteine to thnh cystine c cha cu (-S-S-).

Aminoacid cha lu hunh











4. Cc aminoacid acid v cc amide
Thuc nhm ny c aspartic acid v glutamic acid . Trong phn t ca
chng c 1 nhm amine v 2 nhm carboxyl. pH sinh l (6-7) cc
aminoacid ny tch in m.
Amine ho nhm carboxyl mch bn ca aspartate v glutamate to
thnh cc amide tng ng l asparagine v glutamine.
COOH

H
2
N-CH

CH
2
-OH
Serine
(Ser)
COOH

H
2
N-CH

CH
2
-OH

CH
3


Threonine
(Thr)
COOH

H
2
N-CH

CH
2
-SH
Cystein
(Cys)
COOH

H
2
N-CH

CH
2


CH
2
-S-CH
3

Methionine
(Met)
188
s tng hp glutamic acid mt nhm amine t 1 cht cho c
chuyn n cetoacid. Ngi ta gi qu trnh ny l chuyn amine ho do
enzyme transaminase xc tc. Enzyme ny c nhm prosthetic l
pyridoxalphosphate.

Aminoacid c tnh acid Cc amide











5. Cc aminoacid kim
Trong nhm ny lysine v arginine tch in dng ( pH =7), cn
histidine cha nhm imidazol c tnh base yu pH = 7

Aminoacid c tnh base















COOH

H
2
N-CH

CH
2


COOH

Aspartic acid
(Asp)
COOH

H
2
N-CH

CH
2


CH
2


COOH

Glutamic acid
(Glu)
COOH

H
2
N-CH

CH
2


C=O

NH
2

Asparagine
(Asp-NH
2
)

COOH

H
2
N-CH

CH
2


CH
2


C=O

NH
2

Glutamine
(Glu-NH
2
)

COOH

H
2
N-CH


CH
2


CH
2


CH
2


CH
2
NH
2
COOH

H
2
N-CH


CH
2


CH
2
NH
2

CH
2
C=NH



NH

COOH

H
2
N-CH


CH
2


C CH


N

NH



CH

Lysine
(Lys)
Arginine
(Arg)
Histidine
(His)
189
6. Iminoacid (proline)
Prolin khc vi tt c cc aminoacid khc ch nhm amine bc 1
C kt hp vi mch bn to thnh vng pyrrolidine. Do proline l
mt iminoacid cha nhm amine bc 2.












7. Cc aminoacid thm v d vng
Cc aminoacid thuc nhm ny l phenylalanine, tyrosine v
tryptophan. Do c cha vng thm nn cc aminoacid ny c mt s
phn ng c trng.
Nhng aminoacid thm l nhng tin cht ca nhiu alcaloid,
phytohormone nh indolacetic acid.

Aminoacid cha nhn thm
















COOH

HN CH


H
2
C CH
2


CH
2
Proline
(Pro)
COOH

HN CH


H
2
C CH
2


CH

OH

Hydroproline
(Hypro)
COOH

H
2
N-CH

CH
2

Phenylalanine
(Phe)
COOH

H
2
N-CH

CH
2


OH
N
H
COOH

H
2
N-CH

CH
2

Tyrosine
(Tyr)
Tryptophan
(Try)
190
Nhng gc aminoacid ca histidine, lysin v cysteine l nhng nhm
hot ho enzyme. Nhng aminoacid thm tyrosin c h hng gn vi
thyronine, y l tin cht ca hormone tuyn gip trng thyrocine.
Thyrocine c to nn trong tuyn gip trng t thyronin nh s kt hp
vi iod.


















7.4.2 Tnh cht ca aminoacid
Tnh cht lng tnh ca aminoacid
Trong cu to ca aminoacid va c nhm amine (-NH
2
) th hin
tnh base, va c nhm carboxyl (-COOH) th hin tnh acid, ngha l
chng mang tnh lng tnh v c th phn ly thnh ion H
+
v OH
-
. Trong
dung dch nc, cc aminoacid c th ng thi phn ly nh sau:
R - CH - COOH + HOH R - CH - COOH R - CH - COO
-

+ H
+

NH
3
OH - HOH NH
2
NH
2

Nhng cng trnh nghin cu v quang ph khuch tn xc nhn
rng: phn t aminoacid khng c nhm carboxyl t do, cng nh khng
c nhm amine t do v chng u l mui ni c dng nh sau:
R - CH - COO
-
R - CH - COO
-


NH
3
+
NH
3
+


COOH

H
2
N-CH

CH
2


OH
Tyrosine
COO
-

H
3
N
+
-CH

CH
2


O

Thyronin

OH
COO
-

H
3
N
+
-CH

CH
2



O

Thyrocine

OH
J J
J J
191
Do c tnh lng tnh nn aminoacid l nhng cht m tt. V
aminoacid c tnh lng tnh trong dung dch nc nn tu theo pH ca
dung dch m s phn ly ca nhm carboxyl hoc nhm amine s b km
hm v aminoacid th hin tnh kim hoc tnh acid.
Trong mi trng acid, khi nng H
+
cao, s phn ly ca nhm
carboxyl s b km hm v phn t aminoacid tch in dng, trong in
trng n s dch chuyn v pha cc m.

R - CH - COO
-
R - CH - COOH
+ H
+

NH
3
+
NH
3
+


Trong mi trng kim, di tc dng ca cc ion OH
-
s phn ly
ca nhm amine s b km hm, nhm carboxyl s b phn ly, phn t
aminoacid tch in tch m v dch chuyn v cc dng trong in
trng.
R - CH - COO
-
R - CH - COOH R - CH - COO
-

+ OH
-
+ H
2
O
NH
3
+
NH
3
OH NH
2


im ng in pI (isoelectric point) phn t aminoacid trung ho
v in v n khng dch chuyn trong in trng.
Tu theo s nhm carboxyl v nhm amine trong phn t aminoacid
cng nh tu theo hng s phn ly ca cc nhm ny m im ng in
pI ca cc aminoacid s khc nhau.
Cc aminoacid monoamine dicarboxylic (aspartic acid, glutamic
acid) c im ng in trong mi trng acid, cc aminoacid diamine
monocarboxylic (acginine, histidine, lysine) c im ng in trong mi
trng kim.
Bng 7.4 im ng in ca cc aminocid
Tn aminoacid im ng in
(pI)
Tn aminoacid im ng
in (pI)
Alanine
Arginine
Asparagine
6,00
10,76
5,41
Lysine
Methionine
Hydroproline
9,74
5,74
5,83

ng dng tnh lng tnh ca aminoacid
192
Nh c tnh lng tnh m aminoacid c vai tr l cht m trong t
bo v trong trao i khong ca c th thc vt.
Da vo tnh cht ny nh tnh v nh lng cc aminoacid
trong dung dch nghin cu bng phng php in di, sc k trn giy,
sc k ct...
Cng do tnh cht lng tnh m cc aminoacid c th to mui ni
ng thi vi acid cng nh base:

R - CH - COO
-
R - CH - COOH
+ HCl
NH
3
+
NH
3
+
Cl
-




R - CH - COO
-
R - CH - COONa
+ NaOH + H
2
O
NH
3
+
NH
2


Ngoi ra phn t aminoacid cn to mui ni vi cc ion kim loi
nng nh Fe
2+
, Cu
2+
, Zn
2+
,... c bit vi Cu
2+
. Cu hnh ca mui
phc ny c th biu din nh sau:


R R - CH - NH
2
O - C = O
Cu
2 CH- NH
2
+ Cu
2+
O = C - O H
2
N - CH - R

COOH

Phn ng to ester
Vi ru, aminoacid c th to thnh ester

R - CH - COOH R - CH - COOC
2
H
5

+ C
2
H
5
OH + H
2
O
NH
2
NH
2


Cc ester ca aminoacid u l nhng cht lng d bay hi, c tnh
kim, d iu ch bng php ct chn khng.
193
Phn ng ca gc R
Gc R l nhm nguyn t trong phn t aminoacid vn lin kt vi
nguyn t C v khng tham gia trong vic hnh thnh lin kt peptide.
Bn cht ho hc ca gc cho php aminoacid thc hin nhiu phn
ng khc nhau:
- Phn ng to mui vi acid, base v kim loi nng (do nhm -
NH
2
v nhm -COOH c trong gc R) nh trnh by trn.
- Phn ng oxy ho kh (do nhm -SH v nhm -S-S-)

SH CH
2
- S - S - CH
2


CH
2
CH - NH
2
CH - NH
2

- 2H
2 CH - NH
2
COOH COOH

COOH + 2H
Cysteine v cystine ng vai tr quan trng trong vic vn chuyn e
-

v H
+
ca qu trnh oxy ho kh ni ring v qu trnh trao i cht ni
chung
- Phn ng to amide (do nhm -COOH)
COOH CO - NH
2


CH
2
CH2

CH - NH
2
+ NH
3
+ ATP CH - NH
2
+ H
3
PO
4
+ ADP

COOH COOH
Phn ng xy ra tng t vi glutamic acid.
- Phn ng kh amine ho bi HNO
2
(do nhm -NH
2
)
R - CH - COOH R - CH - COOH
+ HNO
2
+ N
2
+ H
2
O
NH
2
OH
Ngoi ra vng thm ca aminoacid tham gia phn ng nitro ho,
halogen ho v mt s phn ng c trng ca aminoacid: phn ng vi
aldehyd formic, vi ninhydrin...

7.5 Sinh tng hp aminoacid
194
vi khun v lc lp ca thc vt con ng quan trng nht
tng hp glutamate l t -cetoglutarate v glutamine. Phn ng ny c
xc tc bi enzym glutamat-synthase.
ng vt glutamate xut hin ch yu bng phn ng chuyn
amine gia -cetoglutarate v aminoacid. Phn ng ny xy ra ch yu
gan, l c quan trung tm ca s bin i aminoacid.
s chuyn amine ho glutamate v aspartate l nhng cht cho
nhm NH
2
. Mt s vn chuyn amine t glutamate n pyruvate c ch
ra nh sau:










Enzyme xc tc l glutamat-pyruvat-aminotransferase. -
cetoglutaric acid c to nn li tr thnh cht nhn NH
3
. Nh s chuyn
amine m nhiu aminoacid c to thnh. Nhng v d quan trng nht
l:

-cetoglutaric acid glutamic acid
oxaloacetic acid aspartic acid
glyoxylic acid glycine
pyruvic acid alanine
hydroxy pyruvic acid serine
-ceto hydroxybutyric acid threonine
Glutaminsemialdehyde ornithin

Tuy nhin khng phi tt c cc aminoacid c trong t nhin c
tng hp bng con ng chuyn amine ho. Cn c nhng con ng
phn ng quan trng khc tng hp nn cc aminoacid. Ngi ta bit
HOOC CH
2
CH
2
CH COOH

NH
2
Glutamate
HOOC CH
2
CH
2
CO COOH
CH
3
CO COOH

CH
3
CH COOH

NH
2
Pyruvate
-cetoglutarate
195
cc con ng phn ng tng ng cho 5 h aminoacid khc nhau (bng
7.5). V d hnh 7.12 ch ra con ng tng hp ca h glutamic acid.
1. Glutamic acid c phosphoryl ho nh ATP nhm -carboxyl
to thnh glutamylphosphate
2. Cht ny c kh nh NADPH to thnh aldehydglutamic acid
ng thi tch ra phosphate.
3. Cht ny c amine ho nh enzyme transaminase v tr -
carbon, to nn ornithine.
4. Cht ny c kh amine ho nh 1 enzyme transaminase v tr
-carbon lm xut hin -ceto- -amino-valeric acid.
5. Qua s ngng t mt vng 5 cnh c to nn (pyrrolidincarbonic
acid)
6. Cht ny c kh thnh proline nh NADPH.


























Hnh 7.12 Cc con ng phn ng ca h glutamic
COOH

H
2
N-CH

CH
2


CH
2


COOH

Glutamic acid
COOH

H
2
N-CH

CH
2


CH
2

O
C
O P

Glutamylphosphate
COOH

H
2
N-CH

CH
2


CH
2

O
C
H

Semialdehyd-
glutamic acid
COOH

H
2
N-CH

CH
2


CH
2


H
2
N - CH
2

Ornithine
ATP
ADP
NADPH+H
+
NADP
+
COOH

H
2
N-CH

R
COOH

C = O

R
COOH

O = CH

CH
2

H
2


N CH
2



CH
2

COOH

N CH
2



CH
2
CH
2

CH
2

COOH

HN CH
2



CH
2
CH
2

CH
2

NADPH+H
+
NADP
+
H
2
O
COOH

C = O

R
COOH

H
2
N-CH

R
1 2 3
4
5 6
Proline Pyrolidin
carbonic acid
-ceto- -amino
valeric acid
196

Bng 7.5 Nhng aminoacid quan trng nht ca cc h aminoacid

H glutamic H aspartate H
pyruvate
H serine H aminoacid
thm
Glutamate Aspartate Alanine Serine Phenylalanine
Ornithine Lysine Valine Glycine Tyrosine
Arginine Methionine Leucine Cysteine Tryptophan
Proline Threonine
Hydroxyproline Isoleucine

Arginine cng thuc vo h glutamic acid c th c tng hp nn
t ornithine. Hydroxyproline c tng hp t proline nh 1 oxidase c
chc nng hn hp. Cht cho H l ascorbic acid (vitamin C). Nh s
di y nhm OH nh v C
3
ca ascorbic acid c kh nng phn ly 1
H
+
. Phn t ny c c tnh acid. Gi tr pK ca n l 4,2. Ascorbic acid
tham gia vo phn ng vi proline khi tch ra phn t O
2
, trong n
cung cp nguyn t H cn thit cho s to H
2
O tng ng.









S hydroxyl ho proline xy ra khng phi nhng phn t proline
ring bit, m xy ra sau khi phn t proline tham gia cu to protein.
dng ny hydroxylproline l thnh phn quan trng trong collagen v
elastin. Nu thiu aminoacid ny s dn n s ri lon trong cc m lin
kt, m chng (xng, rng) v gn. Khi thiu vitamin C th vic xy
dng cc m ny b tn thng (bnh hoi huyt). S ri lon ny c
gii thch l thiu cht cho H, ascorbic acid, tng hp hydroxylproline
t proline.


CH
2
OH

HOCH
OH OH
O
Ascorbic acid
C = O
CH
2
OH

HOCH
O
-
OH
O
Ascorbate
C = O + H
+
197


















Proline v hydroxylproline l nhng aminoacid d vng. y
nguyn t N dng NH. Mc d vy ngi ta khng gi n l iminoacid,
v n th hin nh aminoacid. Thc t nguyn t H b thiu c thay th
bi 1 cht khc. iminoacid c nhm imine, y nguyn t N kt hp
vi C bng 1 lin kt i (HN=C).
S tng hp cc aminoacid thm i t phosphoenolpyruvate v
erythrosophosphate. Con ng tng hp phc tp ny khng nu ln
y. Mt sn phm trung gian quan trng ca qu trnh ny l shikimic
acid.
tng hp cysteine, mt aminoacid cha S, cn kh sulfate. Qu trnh
ny ch xy ra thc vt v cc vi sinh vt khc nhau, tng t qu trnh
kh nitrate, khng xy ra ng vt.
Sn phm u tin ca s kh sulfate l aminoacid cha S, cysteine.
N cung cp S cho tng hp nhng aminoacid quan trng cha S khc:
methionine. Qu trnh phn ng ch ra hnh 7.13.
1. Phosphohomoserine, mt hp cht tng t nh serine, phn ng
vi cysteine to thnh cystathionine v tch ra phosphate v c
CH
2
OH

HOCH
O H O H
O
Ascorbic acid
C = O
CH
2
OH

HOCH

O
O
Dehydro
Ascorbic acid
C = O

COOH

HN CH
2



CH
2
CH
2

CH
2

Proline
+ O
2
COOH

HN CH
2



CH
2
CH
2

CH


OH

Hydroxyproline

O
H
2
O
198
2. Cystathionine c tch ra thnh serine v homocysteine bng
phn ng thu phn
3. Nh c cht cho nhm CH
3
nguyn t H ca nhm SH ca
homocysteine (cht ny ging cysteine) c thay th bng 1
nhm methyl. Methionine c to thnh.






























Hnh 7.13 Tng hp methionine

7.6 Qu trnh kh nitrate

COOH

H
2
N-CH

CH
2
-S H
Cystein
COOH

H
2
N-CH

CH
2


P O CH
2

Phospho-
homoserin
P
i
COOH

H
2
N-CH

CH
2


CH
2

COOH

H
2
N-CH

CH
2
S
Cystathionine
COOH

H
2
N-CH

CH
2


HS CH
2

COOH

H
2
N-CH

CH
2
-OH
Serine Homoserin
COOH

H
2
N-CH

CH
2


CH
2
-S-CH
3

Methionine
H
2
O
CH
3
cht cho
Cht nhn H
+
1
2
3
199
Nhng nguyn t c bn ca carbohydrate v lipid l C, H v O.
Ngoi ra cn c cc nguyn t S v P, v d trong phospholipid v
sulfolipid, nhng chng khng phi l cc nguyn t c bn. Trong
protein v nucleic acid, N l nguyn t khng th thiu c. Trc khi
cp k hn trao i cht ca chng, chng ta cn tm hiu cc i phn t,
cc cht trao i, cc cht hu c c trng lng phn t thp c to
nn t cc cht v c nh th no. S sau y a ra 4 con ng
phn ng: tng hp polysaccharid, lipid, protein v nucleic acid.
Polysaccharid, lipid, protein v nucleic acid c kh nng to nn nhng
phc hp ln l nhng thnh phn ca t bo nh thnh t bo, mng t
bo, enzyme v ribosome.
Thc vt v mt s sinh vt tin nhn c kh nng tng hp nhng
phn t hu c t nhng cht v c ngho nng lng. Cng nh vy i
vi s c nh nit v c.

H
2
O
-
Phospho-
glyxerate
Oxaloacetate

Mono-
saccharide

Poly-
saccharide

Thnh t bo
CO
2

Acetate
Malonate


Glycerine
Acid bo

Lipid

Mng sinh hc
N
2

Axit -
oxoacid

Aminoacid
Protein Enzyme T
bo NH
3
NO
3
-
Ribose
Carbamyl-
phosphate

Nucleotide

Nucleic acid

Ribosome
Cc hp
cht v c
Hp cht
trung gian
n phn Hp cht
cao phn t
Cc kt hp
i phn t

MG: 50 - 250 MG: 100-300 MG: 10
3
-
10
8
MG: 10
6
- 10
9


Nit dng NH
3
, NO
3
-
v mt phn dng N
2
c th c ng
ho. S ng ho trc tip thc cht l s kt hp NH
3
vi mt phn t
hu c. Nitrate v N
2
trc ht c chuyn sang NH
3
. Hai qu trnh ny
s c cp phn sau y.
Qu trnh kh nitrate v nitrite l chuyn NO
3
-
thnh NH
3
qua NO
2
-
,
c xc tc bi 2 enzyme khc nhau l nitratreductase v nitritreductase.
Nitratreductase l 1 enzyme ph bin trong vi khun, nm, thc vt
bc cao v thc vt bc thp. nhng nhm sinh vt khc nhau th
200
enzyme ny c nhng c tnh khc nhau. Chc nng ch yu l kh NO
3
-

thnh NO
2
-
. Nhm prosthetic ca enzyme ny thc vt bc cao l FAD,
cytochrome b v Mo. Trnh t phn ng c phn tng t chui enzyme
h hp, c biu din trong hnh 7.14. NADH, mt s loi sinh vt l
NADPH chuyn e
-
n FAD, to thnh FADH
2
, sau in t ny c
chuyn n cytochrome b, cytochrome b trng thi kh, kh Mo
6+
thnh
Mo
4+
. Mo
4+
c kh nng ko 1 nguyn t O ca NO
3
-
v kh to thnh
NO
2
-
. Oxy dng kh phn ng trc tip vi proton to thnh H
2
O.
Trong s 2H
+
c gii phng ra trong qu trnh oxy ho kh gia
FADH
2
v cytochrome b (hnh 7.14).








Tng qut: NAD(P)H + H + NO
3
-
NO
2
-
+ H
2
O + NAD(P)
+
Hnh 7.14 Trnh t phn ng ca nitratreductase

Enzyme ny nh v trong t bo cht v dng kh ca n c to
nn t qu trnh ng phn (kh aldehydphosphoglyxeric thnh
phosphoglyxeric acid), thc vt C
4
bng s oxy ho malate thnh
oxaloacetate. Hot tnh ca enzyme nitratreductase b nh hng mnh bi
nh sng, nguyn nhn l do cht kh ca nitratreductase c gii phng
trc tip t lc lp. y lc lp tip nhn oxaloacetate hoc
phosphoglyxerate, trong qu trnh quang hp c kh thnh malate v
aldehydphosphoglyxeric. Hai cht ny c a ra li t bo cht. y
chng b oxy ho c s tham gia ca NAD
+
to thnh NADH. C ch
ny c bit c ngha thc vt C
4
, v chng mt lng ng k cht
kh c to thnh t malate i ra t lc lp.
Hot tnh ca enzyme nitratreductase b nh hng bi cc cht khc
nhau. Nitrate hot ho mnh m, ngc li NH
4
+
v cc aminoacid km
hm enzyme ny. Enzyme ny c na thi gian tn ti l mt s gi, v
vy n phi lun lun c tng hp mi. S tng hp ny ph thuc vo
cng nh sng.
NAD(P)
+

NAD(P)H + H
+

FADH
2
FAD

2
x
Cytchr.b Fe
III

2
x
Cytchr.b Fe
II
Mo
VI
Mo
IV
NO
3
-

NO
2
-

O
2
-

H
2
O
2H
+

201
Bn cnh nitratreductase ng ho nh nu trn, ngi ta cn bit
mt loi nitratreductase h hp. Loi ny c mt ch yu trong vi sinh vt
v c to nn c bit trong iu kin ym kh. Chc nng ca n l khi
khng c O
2
, in t bt ngun t chui enzyme h hp kt hp vi
nguyn t O ca NO
3
-
. Trong trng hp ny NO
3
-
thay th nguyn t oxy
trong chui h hp. Nitratreductase h hp ny gn cht vi mng v cng
tn ti vi cytochrome, b c ch bi O
2
. Ngc vi nitratreductase ng
ho khng kt hp v mn cm vi O
2
, nitratreductase h hp l 1 enzyme
xc tc qu trnh phn nitrate ho . Phn nitrate ho c hiu l kh NO
3
-

cho ti N
2
O v N
2
. Qu trnh ny xy ra ch yu trong t, nc trong
iu kin ym kh v c vai tr quan trng trong t nhin. S kh NO
3
-

theo kiu h hp to thnh NO
2
-
xy ra trong h thng tiu ho ngi v
ng vt.
NO
2
-
c to ra nh enzyme nitratreductase trong t bo cht ca
t bo thc vt l 1 anion ca 1 acid yu. V vy mt phn NO
2
-
to nn
c proton ho v th hin cn bng phng trnh sau:

NO
2
-
+ HOH HNO
2
+ OH
-


NO
2
-
kh i qua mng ngoi ca ty th v lp th, tuy nhin d dng
i vi HNO
2
. Lc lp v ty th tip nhn ch yu HNO
2
v nh vy
lm cho t bo cht b kim ho. S kim ho ny lm tng s tng hp
cc anion hu c.
Trong ty th xy ra s kh HNO
2
thnh NH
3
. Enzyme nitritreductase
gm 1 chui polypeptide vi 1 siroheme (cng tn ti Fe v S) v mt
phc h 4 nguyn t Fe-S.
Trnh t phn ng ca nitritreductase c ch ra trong hnh 7.15.
Trong lc lp cht kh trc tip l ferredoxin. y c mt s kt hp
cht ch gia quang hp v kh nitrate.
Phn ng xy ra nh sau:

HNO
2
+ 2e
-
+ 2H
+
NH
3
+ O
2


Trong nhng c quan thc vt khng c mu xanh, v d nh r cy,
NO
2
-
cng c kh n NH
3
. Mt thi gian di qu trnh ny khng
c gii thch, v nhng b phn khng c mu xanh, c bit l r cy
khng c ferredoxin, cho n khi nhng kt qu mi ch ra rng, trong ty
th ca r c 1 ferredoxin tng t h thng oxy ho kh c kh nng kh
202
HNO
2
thnh NH
3
. H thng kh ny cha cht kh l NADH hoc
NADPH.









Tng qut: HNO
2
+ 2e
-
+ 2H
+
NH
3
+ O
2
Hnh 7.15 Trnh t phn ng ca nitratreductase

Hot tnh ca nitritreductase c kch thch bi nitrite v nitrate.
Nng nitritreductase cao hn nng nitratreductase, l nguyn
nhn ti sao khi cung cp nhiu nitrate cho cy dn n s tch lu nitrate
m khng tch lu nitrite. S tch lu nitrate xy ra khi thc vt c bn
qu nhiu nitrate hoc khi nh sng cho quang hp yu.
Trong trng hp nh sng yu dn n s thiu cht kh. l
nguyn nhn ti sao i vi ging rau ba lng nhng gi bui sng th c
nng nitrate cao hn bui chiu. Ban m ch yu l phosphoglyxerate
v mt t triosephosphate i ra khi lc lp. Ban ngy th ngc li.
Triosephosphate c kh nng kh NAD
+
thnh NADH khi phn gii theo
qu trnh ng phn cung cp cht kh cho qu trnh kh nitrate.
Ngc li phosphoglyxerate khng th kh NAD
+
v nh vy khng cung
cp NADH cho qu trnh kh nitrate.
Nng nitrate cao thng gp trong nhng b phn sinh trng
dinh dng v y c th nh hng n sinh l dinh dng. c bit
i vi nhng sn phm thc vt c dng n sng nh x lch, bp
ci, c rt. Thc ra nitrate khng c hi nhng trong qu trnh bo qun vi
sinh vt bin i thnh nitrite v trong c th ngi, c bit trong tuyn
nc bt nitrate c kh thnh nitrite. Trong mi trng acid to thnh
HNO
2
, n c th c tch thnh nitrosyl v hydroxyl.

HNO
2
HO N O OH
-
+ NO
+


T bo cht
NO
2
-


+ H
2
O


HNO
2
+ OH
-

Lc lp
Ferredoxin
Phc h
Nitratreductase
Siroheme
4 x Fe-S
2e
-
NH
3
+ O
2
2H
+
2e
-
2e
-
203
Cation nitrosyl (NO
+
) c kh nng oxy ho Fe
2+
ca hemoglobin
bng cch ly i 1e
-
ca Fe

Fe
2
-heme + NO
+
Fe
3
-heme + NO

Hemoglobin dng oxy ho khng th kt hp v vn chuyn O
2
,
dn n s ngt th. Hin tng ny gp ngi trong mt s thng u
v sau c th pht trin enzyme c kh nng kh hemoglobin dng
oxy ho.

7.7 Qu trnh c nh nit phn t
Ch sinh vt tin nhn mi c kh nng c nh N
2
, l 11 trong 47
h vi khun v 6 trong 8 h vi khun lu hunh. Mt s sng t do, mt
s sng cng sinh vi thc vt bc cao. Nhng vi sinh vt c nh m
quan trng nht l:
- Rhizobium, sng cng sinh vi cy h u.
- Bradyrhizobium, cng sng cng sinh vi cy h u, chng khc
vi rhizobium ch l sinh trng chm hn.
- Loi frankia, vi khun sng trong t.
- Vi khun lu hunh heterocysten sng cng sinh vi cc loi thc
vt khc nhau nh vi bo hoa du (azolla) l.
Nhng loi vi sinh vt sng cng sinh v t do c cha gen Nif
trong h gen (Ni l vit tt ca nitrogen v f l fixing). Gen ny m ho
cho enzyme nitrogenase, l mt a enzyme, xc tc cho phn ng c nh
N
2
, kh N
2
thnh NH
3
. Nitrogenase gm 2 protein cha Fe-S, protein nh
hn c trng lng phn t 60 kDa, cha 1 chm Fe-S c 4 nguyn t,
phn t protein ln hn c trng lng phn t 220 kDa v cha nguyn
t S linh ng, 36 nguyn t Fe v 2 nguyn t Mo. Phn t protein nh
hn c chc nng vn chuyn e
-
, trong e
-
ca ferredoxin hoc
flavodoxin vn chuyn ln phc h Fe-Mo (hnh 7.16).
S vn chuyn e
-
ny cn nng lng, do ATP cung cp. ATP kt
hp vi protein cha Fe qua 1 cu Mg v sau c thu phn thnh
ADP v Pi. S thu phn lm thay i hnh dng ca protein enzyme,
e- c vn chuyn ln phc h Fe-Mo. vn chuyn 1e
-
cn 2 ATP.
Phc h Fe-Mo thc cht l 1 N
2
-reductase. Phc h nh hn c gch
cho l 1 protein cha Fe, phc ln hn l protein cha Fe/Mo, phc ln
ny thc cht l dinitrogen-reductase (hnh 7.16).

204
















Hnh 7.16 Cc phn ng xy ra bacteroid ca vi khun rhizobium

vn chuyn e
-
t phc Fe/Mo n N
2
th s thay i ho tr Mo
c mt vai tr quan trng, nh hnh 7.17.
1. 2e
-
v 2H
+
c kt hp vi nguyn t Mo
2. Nguyn t H
2
ang gn vi enzyme b y ra bi N
2

3. N
2
c kh thnh -N=N do tip nhn H
+
+ e
-

4. -N=N c kh thnh N-NH
2
do tip nhn H
+
+ e
-

5. N-NH
2
c kh thnh NH
3
do tip nhn H
+
+ e
-
, NH
3
c tch
ra khi phc h enzyme
6. Nguyn t N ang gn vi enzyme c kh thnh NH
3
do tip
nhn 3e
-
v 3H
+
v sau NH
3
c tch ra khi phc h.
S gn N
2
vi phc h Fe-Mo cn 1 lin kt trc ca phc h
ny vi H
2
. V vy kt qu l bn cnh to ra NH
3
ngoi ra cn xut hin
H
2
, phn t ny c th c tch ra thnh H
+
v e
-
v e
-
ny c a tr
li h thng (hnh 7.16).



Ferredooxin
Flavodoxin
Chu
trnh
Krebs
NADH
ATP
ATP + P
i
2H
+
+ 2e
-
H
2

2H
+
2NH
3
N
2

Hydrogenase
Leghemoglobin
Chui
enzyme
h hp
Cht ng ha
O
2
Nitrogenase
e
-
H
+
205
















Tng qut: 8H
+
+ 8e
-
+ N
2
H
2
+ 2NH
3
Hnh 7.17 S kh N
2
thnh NH
3
phc h Fe-Mo ca nitrogenase
rhizobium v bradyrhizobium h thng nitrogenase c biu
din hnh 7.15, h thng ny nh v bacteroid c tm thy trong
khng bo nhim ca nhng t bo nt sn. Bacteroid tng t mt c
quan v c th so snh vi ty th. N c cung cp cc hp cht carbon
hu c, ch yu l anion hu c (succinate, malate, pyruvate) bi t bo
cht ca k ch, nhng cht ny c tip tc phn gii trong chu trnh
Krebs. y xut hin NADH, c s dng mt phn l cht cho in t
ferredoxin v flavodoxin, mt phn c s dng lm nguyn liu ca h
hp. H thng oxy ho kh ca chui h hp cu to nn mng. Chng
cung cp ATP v ATP cn cho s hot ho enzyme nitrogenase. Chui h
hp c cung cp O
2
bi leghemoglobin. Phc h protein cha heme c
trong khng bo nhim ny c th so snh vi myoglobin trong m c
hoc hemoglobin trong mu. N cung cp O
2
cho chui enzyme h hp v
n kim sot O
2
i vo bacteroid. S kim sot ny rt quan trng, v
enzyme ny b c ch bi O
2
.
NH
3
c to nn bi nitrogenase c a n t bo cht ca t
bo ch v y c s dng tng hp nn aminoacid. S tng hp
enzyme nitrogenase c iu khin bi enzyme glutamate synthetase,
N
2
H
2
Fe Mo = N N

Fe Mo = N = NH

Fe Mo = N NH
2
Fe Mo N

Fe Mo

e
-
e
-
e
-
3e
-
2e
-
H
+
H
+
2H
+
3H
+
NH
3
Fe Mo H
2
1
2
3
4
5
6
NH
3
H
+
206
xc tc cho tng hp glutamin t NH
3
. Nu trong h thng c t NH
3
th
glutamate synthetase kch thch tng hp nitrogenase, nng NH
3
cao th
c ch s tng hp nitrogenase. C ch iu khin ny rt c ngha, tuy
nhin khi cung cp NH
3
d tha th nitrogenase khng c s dng, v
c NH
3
. V vy bn phn m c ch s to thnh nt sn ca cy h
u. vi khun rhizobium sng cng sinh, NH
3
to thnh c a nhanh
n t bo ch, nn s tng hp nitrogenase hu nh khng b km hm.
Hn na vi khun trong t bo ch lun c cung cp bi nhng anion
hu c. l nhng nguyn nhn quan trng nht, ti sao s cng sinh vi
khun/cy h u c t l c nh N
2
cao. V d cy c (trifolium
pratence) sng cng sinh vi vi khun trong iu kin sinh trng tt c
th c nh 300-400 kg N/ha/nm. Vi khun sng t do t l c nh nit
b nh hng do s tch lu NH
3
trong t bo vi khun v s cung cp
khng y carbon hu c. Loi vi khun c nh nit sng t do cn
cho s trao i cht ca chng nhng hp cht hu c d phn gii, nh
ng v cc acid hu c, thng c trong t vi lng khng . V
vy kh nng c nh m ca vi khun sng t do c bn l thp. Nhng
vi khun lu hunh t dng thuc loi anabaena, nostoc v rivularia t
ng ho CO
2
bng quang hp v c nh N
2
. Kh nng c nh N
2
b nh
hng rt nhiu khi nng O
2
cao.

7.8 Cc ng hng bin i NH
3
- chu trnh ornithine
S kh nitrate-nitrite v h thng nitrogenase sn sinh NH
3
, sau
NH
3
kt hp vi mt phn t hu c, thng l -cetoglutaric acid
hoc glutamic acid. Xc tc cho phn ng gn NH
3
ny thc vt bc cao
v bc thp l glutmate synthetase (GS), enzyme ny thng hot ng
trong mt nhm. Glutmate synthetase l mt protein c 8 tiu n v vi
trng lng phn t t 350 n 400 kDa. Ngi ta bit c 2 loi enzyme
ng phn, mt loi c trong t bo cht (GS1) v mt loi c trong lc
lp. Phn ng do glutmate synthetase xc tc nh sau:








207












Cht nhn NH
3
l glutamate. Trong phn ng, enzyme c
phosphoryl ho (kinase) v sau c kh phosphoryl ho
(phosphatase). C hai phn ng v tng quan l s thu phn ATP. Sn
phm phn ng l glutamine, mt amide. Mt phn t tng t glutamine
l asparagine, c to nn tng t glutamine. Tuy nhin n khng phi
l cht nhn NH
3
quan trng.
Vai tr trung tm ca glutmate synthetase i vi s ng ho NH
3

c gii thch glutamine l cht cho nhm NH
2
khi tng hp glutamate.
y -cetoglutaric acid l cht nhn, c ch ra nh sau:










Cc cht tham gia phn ng trao i 1 nhm NH
2
vi nguyn t O.
mi sn phm to thnh thiu 1 nguyn t H, c ferredoxin hoc
NAD(P)H cung cp. Bng cch ny t glutamine v -cetoglutaric acid 2
COOH

H
2
N-CH

CH
2


CH
2


C = O

OH H NH
2

Glutamate
COOH

H
2
N-CH

CH
2


CH
2


C=O

NH
2

Glutamine
ATP
ADP + P
i
COOH

H
2
N-CH

CH
2
+

CH
2


C=O

NH
2

-cetoglutarate
2e
-
2H
+
COOH

H
2
N-CH

CH
2


CH
2


COOH

Glutamic acid
COOH

H
2
N-CH

+ CH
2


CH
2


COOH

COOH

O = C

CH
2


CH
2


COOH

208
phn t glutamate c to nn. Qu trnh ny gm s chuyn nhm
amine v s kh v phn ng c xc tc bi glutamate synthetase.
Glutamate synthetase c hai loi isoenzyme c cu to khc nhau.
Glutamate synthetase ph thuc vo ferredoxin l 1 flavoprotein cha Fe-S
c trng lng phn t 140-230 kDa, enzyme ny c trong lc lp, nhn
cht kh qua ferredoxin trc tip t quang hp. Glutamate synthetase c
trong t bo cht c trng lng phn t khong 240 kDa. Cht kh ca n
l NADH (t ng phn) hoc NADPH (t oxy ho trc tip glucose). C
2 enzyme glutamate synthetase v glutamine synthetase hot ng trong 1
nhm, lm cho qu trnh phn ng thnh 1 vng trn, nh hnh 7.18.
Tng qut chu trnh ch ra rng -cetoglutaric acid c amine ho
nh NH
3
vi s tiu tn 1 ATP v 2e
-
. -cetoglutaric acid ( -
cetoglutarate) c ngun gc t chu trnh Krebs. Sn phm ca phn ng l
glutamate, c ngha quan trng, v nh chuyn amine ho m nhm
amine c chuyn n mt cht khc (cetoacid, aldehyd). Hu ht nit c
trong thc vt xy ra qua phn ng ny, mt ln c trong nhm amine ca
glutamate. Glutamate synthetase l 1 glutamine- -cetoglutarate-
aminotransferase, c gi tt l GOGAT. V vy ngi ta gi chu trnh
tng hp glutamate l con ng phn ng GOAGT.
Nhng enzyme ca chu trnh tng hp glutamate c bit c nhiu
trong nhng t bo ca nt sn. y nhng phn t NH
3
c cung cp
bi bateriod ca vi khun nt sn c ng ho qua chu trnh ny. Con
ng GOAGT cng c ngha trong lc lp ca thc vt C
3
. NH
3
c
gii phng ra mt lng ln trong h hp sng, nu NH
3
ny bay ra
khng kh th y l mt s mt mt ln i vi thc vt. NH
3
gii phng
ra c tip nhn bi chu trnh tng hp glutamate v nit ca n c s
dng li trong trao i cht ca thc vt dng aminoacid.










209








Tng qut: NH
3
+ ATP +2e
-
+ 2H
+
+ -cetoglutarate
Glutamate + ADP +Pi
Hnh 7.18 Chu trnh tng hp glutamate (con ng GOGAT)

lu ngi ta cho rng enzyme glutamate dehydrogenase (GDH)
cng c ngha i vi s ng ho NH
3
trong thc vt. Glutamate
dehydrogenase xc tc cho phn ng sau:











y l s kh amine ho oxy ho, l mt qu trnh gii phng NH
3
,
ngc vi qu trnh ng ho NH
3
. Tuy nhin khi nng NH
3
cao th n
s gy c cho thc vt. i vi nhiu loi vi sinh vt s ng ho NH
3

nh GDH c ngha. Trong thc vt GDH c ch yu trong ty th.
NADH m n to ra c th i vo chui h hp, NH
3
c th c s dng
cho tng hp asparagine nh enzyme asparaginsynthetase theo phng
trnh sau:

Glutamate
Glutamine
-cetoglutarate
Ferredoxin
Glutamate
(chuyn amine ha)
e
-
NAD(P)H

ADP + P
i
NH
3
+ ATP
-cetoglutarate
NAD
+
COOH

H
2
N-CH

CH
2


CH
2


COOH

Glutamate
COOH

CO

CH
2


CH
2


COOH

COOH

HN = C

CH
2


CH
2


COOH

Iminoglutarate
NADH + H
+
NH
3
H
2
O
210











Cht nhn NH
3
l aspartate. Phn ng ny tng t phn ng tng
hp glutamine. Glutamate dehydrogenase c tng i nhiu trong m r
v nhng m gi. V vy ngi ta cho rng, y n sn sinh NH
3
v
l iu kin tng hp asparagine, l dng nit vn chuyn v c vn
chuyn ra khi nhng m ny.

7.9 Cc ng hng tng qut ca s bin i aminoacid

Cc aminoacid v c bn c s dng tng hp protein song
cng c th chu nhng s bin i khc. l phn ng theo nhm
amine carboxyl v phn ng theo gc R ca aminoacid.
- Cc phn ng theo nhm -amine: l cc phn ng phn gii
aminoacid thnh NH
3
v cc sn phm tng ng theo 4 hng c
bn sau:

a. Kh amine ho bng cch thu phn

R - CH - COOH R - CH -COOH
+ H
2
O + NH
3

NH
2
OH
Aminoacid Hydroxyacid

b. Kh amine ho bng cch kh

R - CH -COOH R - CH
2
- COOH + NH
3

NADH
2
NAD
COOH

H
2
N-CH

CH
2


C = O

OH H NH
2

Aspartate
COOH

H
2
N-CH

CH
2


C = O

NH
2

Asparagine
ADP + P
i
ATP
211
NH
2
NADPH
2
NADP
Aminoacid Acid hu c

c. Kh amine ho bng cch oxy ho
R - CH - COOH R - C -COOH
+ 1/2 O
2
+ NH
3

NH
2
O
Aminoacid Cetoacid
Phn ng ny tin hnh qua 2 giai on: trc ht aminoacid b loi
hydro to thnh iminoacid tng ng vi s tham gia ca enzyme
dehydrogenase
R - CH -COOH R - C - COOH
NAD NADH
2

NH
2
NADP NADPH
2
NH
Aminoacid Iminoacid

Tip iminoacid kt hp vi phn t H
2
O, NH
3
b loi tr v gii phng
cetoacid

R - CH - COOH R - C -COOH
+

HOH + NH
3

NH O
Aminoacid Cetoacid


d. Kh amine ho bng cch chuyn ho ni phn t.

R - CH
2
- CH -COOH
R - CH = CH - COOH + NH
3

NH
2

Aminoacid Acid cha no

Trong cc phn ng trn th phn ng amine ho bng cch oxy ho
l ph bin trong thc vt so vi ng hng khc. Cc -cetoacid u
l sn phm ch yu ca s kh amine ho cc aminoacid.
- Cc phn ng theo nhm carboxyl: din ra ch yu theo hai qu
trnh: s kh carboxyl ho v s to thnh aminoacyl adenylate

a. Qu trnh kh carbocyl ho cc aminoacid
212
Qu trnh ny c thc hin trong cc ng vt, thc vt v c bit
l ph bin trong c th sinh vt. S kh carboxyl tin hnh theo s
sau:

R - CH - COOH Decarboxylase
R - CH
2
- NH
2

NH
2
CO
2
Amine


Sn phm phn ng l cc amine. Mi mt aminoacid ring bit c
xc tc bi mt enzyme decarboxylase tng ng.


Sn phm ca s kh carboxyl ho cc aminoacid l amine, diamine,
CO
2
, NH
3
v cc hp cht khc. Tr CO
2
, NH
3
, tt c cc hp cht
khc u b phn gii tip tc. Nu trong iu kin bt li th cc
amine v diamine tch t li gy c cho cy, trong iu kin bnh
thng chng s b oxy ho di tc dng ca enzyme oxydase to
thnh NH
3
v cc aldehyd.
Monoaminoxydase
R - CH
2
-NH
2
+ H
2
O + O
2
R - CHO + NH
3
+ H
2
O
2
Aldehyd dehydrogenase
R - CHO + H
2
O R -COOH

Cc acid v aldehyd li b oxy ho tip tc to thnh CO
2
, H
2
O v
NH
3
. Cc sn phm ny chnh l sn phm cui cng ca s phn gii
aminoacid.
b. Phn ng theo nhm carbocyl l qu trnh to thnh phc hp
aminoacyl-adenylate. Chi tit c ch ca phn ng ny trong phn
sinh tng hp protein.
- Nhng chuyn ho ca phn t aminoacid c lin quan n gc R
trong phn t.
Do tnh mun mu mun v ca gc R nn cc phn ng ho hc
ca chng v cng phong ph v a dng. Nhiu phn ng c thc hin
trong qu trnh trao i aminoacid, trong quan trng nht l cc phn
ng bin i t mt aminoacid ny thnh mt aminoacid khc. Nh ng
hng ny m kh nng tng hp cc aminoacid tng ln rt nhiu.
Sau y ch nu ln mt s v d:
Khi oxy ho phenylalanine s to thnh tyrosine
213
Thu phn arginine th ornithine c to thnh v t ornithine s
thu c proline hoc glutamic acid.

H
2
N - C = NH

NH NH
2
NH
2

Arginase
(CH
2
)
3
(CH
2
)
3
+ O = C
+ HOH
CH - NH
2
CH - NH
2
NH
2


COOH COOH

Arginine Ornithine Ur

Khi oxy ho nhm -SH ca cysteine s thu c cystine
SH
NAD NADH
2
CH
2
- S -S - CH
2

CH
2
Cystine reductase
CH-NH
2
CH-NH
2

CH - NH
2

NAD NADH
2
COOH COOH
COOH
Cysteine Cystine

T threonine nu loi tr gc acetaldehyd s thu c glycine.

CH
3


CHOH CH
3
CH
2
-NH
2

+
CH - NH
2
CHO COOH

COOH
Threonine Acetaldehyd Glycine

7.10 Sinh tng hp protein

214
Tham gia vo s tng hp polypeptide v protein c mRNA, tRNA,
rRNA, trong rRNA l thnh phn ca ribosome. Vi s tng
polypeptite thng tin di truyn c chuyn t DNA n polypeptide.
Trc khi nghin cu qu trnh tng hp protein cn phi cp n tng
nucleic acid v ribosome.




















Hnh 7.19 S tng hp protein

1. Nguyn liu tng hp polypeptide l cc aminoacid. tham
gia vo phn ng chng phi c hot ho nh ATP, to nn AMP-
aminoacyl aminoacid hot ho
2. Aminoacyl c chuyn ln tRNA.
3. tRNA vn chuyn aminoacyl n ribosome. y aminoacyl c
th to lin kt peptide.
P - P
Nhn t bo
Ribosome
mRNA
tRNA - aminoacyl
tRNA
AMP - aminoacyl
AMP
Aminoacyl
P
i
ATP
1
2
3
4
215
4. mRNA c tng hp trong nhn t bo ri i n ribosome, l
ni tng hp polypeptide. mRNA mang thng tin cho chui polypeptide
c tng hp, xc nh trnh t ca aminoacid trong chui polypeptide.
Thng tin di truyn trong mRNA th hin trnh t base ca chng v
ba nucleotide ng cnh nhau m ho cho mt aminoacid xc nh. Cc
b ba ny c gi l codon. Di y l cc codon ca mt on mRNA
m ho cho alanine, aspartic acid, cysteine v serine.






Hnh 7.20 Mt on DNA vi nhng codon m ho cho alanine, aspartic
acid, cysteine v serine

Bng 7.6. Nhng codon trn mRNA ca cc aminoacid

Base th
nht
Base th hai Base th
ba U C A G
U
Phe
Phe
Leu
Leu
Ser
Ser
Ser
Ser
Tyr
Tyr
Term
Term
Cys
Cys
Term
Trp
U
C
A
G
C
Leu
Leu
Leu
Leu
Pro
Pro
Pro
Pro
His
His
Glu-NH
2
Glu-NH
2

Arg
Arg
Arg
Arg
U
C
A
G
A
Ileu
Ileu
Ileu
Met
Thr
Thr
Thr
Thr
Asp-NH
2
Asp-NH
2
Lys
Lys
Ser
Ser
Arg
Arg
U
C
A
G
G
Val
Val
Val
Val
Ala
Ala
Ala
Ala
Asp
Asp
Glu
Glu
Gly
Gly
Gly
Gly
U
C
A
G
Term: terminal: l nhng codon kt thc chui polypeptide.


Ngy nay nhng codon m ho cho cc aminoacid cu to nn
protein c bit. iu rt th v l cc m di truyn cho tt c mi
sinh vt t sinh vt nhn s n gin nht cho n nhng ng vt c v l
G C U G A U U G C U C A

Ala Asp Cys Ser
216
nh nhau. Tuy nhin cng c mt s ngoi l nh ty th ngi. Qua
bng m di truyn ta thy phn ln cc aminoacid c m ho bi hai
codon, v cc codon c hai base nit u tin ging nhau.
Ribosome c to nn t rRNA v protein. s kt hp ny c s
tham gia ca Mg
+
. Ribosome c to nn t 1 tiu phn ln v 1 tiu
phn nh. ln ca cc tiu phn c c trng bi h s lng S ( h
s Svedberg).
dx/dt
S ---------

2
x
X l khong cch ca trc ly tm
vn tc gc
t l thi gian
dx/dt, khong cch m tiu phn di chuyn trong n v thi gian.
Khong cch ny cng di khi tiu phn cng nng.
H s lng S i vi protein c ln t 1 x 10
-13
n 200 x 10
-13
.
S bin i n v h s lng ca Svedberg l nhn vi 10
13
, tng ng h
s lng ca protein l 1-200. S cng ln th tiu phn cng ln (cng
nng). Bng 7.7 ch h s lng ca cc tiu phn ribosome khc nhau v
ca nucleic acid.
Bng 7.7 H s lng ca cc phn t khc nhau v cc kt hp (S)
-------------------------------------------------------------------------------------
tRNA 4
mRNA 6-25
rRNA 5-23
Tiu n v nh ribosome 30 (sinh vt nhn s)
Tiu n v ln ribosome 50 (sinh vt nhn s)
Ribosome 70 (sinh vt nhn s)
Tiu n v nh ribosome 40 (sinh vt nhn chun)
Tiu n v ln ribosome 60 (sinh vt nhn chun)
Ribosome 80 (sinh vt nhn chun)
-------------------------------------------------------------------------------------

Bng 7.8 Cc rRNA ca ribosome ca tiu phn ln v nh ca sinh
vt nhn chun v nhn s:
-------------------------------------------------------------------------------------
Sinh vt nhn s Sinh vt nhn chun
-------------------------------------------------------------------------------------
Tiu n v nh 16S rRNA Tiu n v nh 18S rRNA
217
Tiu n v ln 23S rRNA Tiu n v ln 5,8S rRNA
Tiu n v ln 5S rRNA Tiu n v ln 28S rRNA
Tiu n v ln 5S rRNA
--------------------------------------------------------------------------------------

Bng 7.8 cho bit ln ca ribosome v cc tiu n v ribosome.
Da trn n v S ny ta thy ribosome ca sinh vt tin nhn nh hn
sinh vt nhn chun.
Ribosome trong ty th v lp th tng t v ln v chc nng
nh ribosome sinh vt tin nhn. Hai tiu phn ca ribosome c gn
vi mRNA, ngha l khi khng c mRNA th hai tiu phn khng kt hp
vi nhau m trng thi ring l.
Cu trc khng gian hai chiu tRNA tng t l c tam ip. Tuy
nhin tRNA c cu trc khng gian ba chiu, v vy phc tp hn v c
biu din theo s hnh 7.21.
Trong tRNA c mt s base nit l nh cytosine v guanine
methyl ha l pseudouridine, dihydrouracil v hypoxanthine.















Pseudouridine lin kt vi ribose khng phi N-glycosid m l C-
glycosid ( v tr C
6
ca uracil). V y l nhng base nit l nn cc v
tr khc nhau ca tRNA khng c s cp i cc base. ngha c bit
Uracil


O =
N
N
H
OH

HN
N
N
N
H
O


Hypoxanthine

Dihydrouracil


O =
N
N
H
OH

Ribose


O =
N
N
H
OH

Pseudouridine ( )

218
hn v chc nng ca tRNA l c v tr anticodon v ch kt hp ca
aminoacyl.



















Hnh 7.21 Cu trc ca tRNA vi anticodon, ch kt hp vi aminoacyl,
dihydrouridine v pseudouridine

Ch kt hp ca aminoacyl ca tt c cc phn t tRNA l nh
nhau, l trnh t cytosine-cytosine-adenine. Aminoacyl c kt hp
vo ribose ca adenine. Anticodon c trng cho tng loi aminoacid, l
nhng base b sung cho codon trn mRNA. Anticodon c c t phi
sang tri. c tnh ca anticodon l hai base u tin c hiu i vi
aminoacid. Base th ba c th l khc nhau, n c th linh ng, c
ngha l khng to cu hydro vi coon. bng sau anticodon C-A
(codon G-U) m ho cho aminoacid valine.
Inosine l mt nucleoside c to nn t hypoxanthine v ribose.
Mt aminoacid c nhiu tRNA c hiu. Trnh t ca mt tRNA c hiu
Anticodon

Vng pseudouridine
Vng dihydrouridine
.. lin kt hydro
Ch kt hp ca aminoacid
P
u cui c phosphoryl ha
219
cho mt aminoacid nht nh c th l khc nhau. S khc bit trong cu
trc bc mt ca tRNA gia nm men v chut c khng nh.













Trc khi aminoacid c s dng tng hp protein chng phi
c hot ho nh ATP, ATP kt hp vi aminoacyl v pyrophosphate
c tch ra (hnh 7.22).
y xut hin mt adenosinphosphataminoacyl (aminoacid
hot ho). ATP cung cp nng lng cn thit cho qu trnh tng hp.
Adenosinphosphataminoacyl mang gc acyl ribose cui cng ca tRNA.
N kt hp vi aminoacyl v tr carbon th 3 ca ribose ng thi gii
phng AMP. Nh vy aminoacyl lin kt vi cht mang c hiu v vn
chuyn chng n v tr tng hp protein. Ton b qu trnh hot ho
aminoacid v kt hp aminoacyl vi tRNA c xc tc bi mt enzyme
aminoacyl-tRNA-synthetase. Ribosome, mRNA v tRNA-aminoacyl to
iu kin cho vic to lin kt peptide. Theo kt qu th nghim vi
khun, c bit l E.coli th qu trnh xy ra nh sau: Tiu n v nh ca
ribosome kt hp vi gc phosphate ca mRNA cng vi mt tRNA-
aminoacyl. Chng c gi l phc h khi u v sau mi c kh
nng kt hp vi tiu phn ln ca ribosome, to ra b my tng hp
polypeptide. Tip theo mt phn t tRNA-aminoacyl th hai kt hp vi
ribosome v m bo tRNA-aminoacyl ng v tr ca mnh (hnh 7.23).
V tr chnh xc l base nit ca codon c kt hp b sung vi base nit
anticodon.


N
N
N
N
OH
HOCH
2

OH OH
O
Hypoxanthine
Inosine
220























Hnh 7.22 Hot ho aminoacid v s kt hp ca aminoacyl vo tRNA

- Amino acid c hot ho bng cch gn vi tRNA ring ca n. Qu
trnh ny gm 2 phn ng, c xc tc bi cng mt enzym c hiu i
vi mi amino acid, l cc amino acid-tRNA-synthetase. Nh vy c
20 amino acid th cng c 20 loi enzyme khc nhau.
Trong phn ng th nht amino acid phn ng vi ATP to nn AMP-
aminoacyl v 1 pyrophosphate (hnh 7.22, phn ng 1). Hp cht c to
nn bi nhm carboxyl ca amino acid v mt gc phosphoric acid v gn
vi enzyme nn c kh nng phn ng cao.
Trong phn ng th hai, n s phn li, cho php aminoacyl t AMP-
aminoacyl c vn chuyn n tRNA. y aminoacyl thay th H ca
nhm OH v tr carbon th 3 to thnh aminoacyl-tRNA.
O O O

AdenosinOP~O~PO~POH

OH OH OH
O

AdenosinOPOCOCHR

OH NH
2
RCHCOOH

NH
2
P - P
1
tRNA-O-CH
2
O H
O
Adenine
OH
tRNA-O-CH
2
O

O
Adenine
OH
COCHR

NH
2
2
AMP
221

- giai on khi u tng hp chui polypeptide cn c hai iu kin:
Mt l trn mRNA c mt khu vc khng m ho, l du hiu kt hp
vi ribosome m u cho vng m ho. Hai l c b m khi u AUG
lm im xut pht. vi khun i khi thy m khi u GUG thay cho
AUG.
Nh ta bit b m AUG m ho cho methionine. vi khun, ngi ta
thy c hai loi tRNA i vi methionine: mt l tRNA nhn gc
methionine avo thnh phn ca chui polypeptide Met-tRNA, hai l
tRNA nhn gc formyl-methionine (fMet-tRNA) c vai tr quan trng
trong vic khi u tng hp chui polypeptide. sinh vt nhn s, nhm
amine ca methionine c th c formyl ho bi N
10
-formyl-
tetrahydrofolic acid, nh enzyme transformylase c hiu. Do nhm amine
b gc formyl bao vy nn khng cho php n tham gia vo qu trnh
ko di chui, m ch tham gia vo qu trnh khi u tng hp protein.
sinh vt nhn s anticodon c nh v gia ca 16SrRNA. Base
b sung kt hp vi base ca codon mRNA khong cch t 0,3 n 0,4
nm, nh vy s xut hin lc hp dn Van der Waal. Bng cch ny trc
ht tRNA-aminoacyl kt hp vi tiu phn nh ca ribosome v thc cht
l vi mRNA. Phc h ny phn ng vi nhn t ko di. Nhn t ny
c trn tiu phn ln ca ribosome.
sinh vt nhn s, ngoi cc yu t tham gia khi u tng hp
protein nh fMet-tRNA, mRNA, cc tiu n v ribosome 30S v 50S,
GTP, cn c 3 protein na l cc yu t khi u: IF
1
(M 9000), IF
2
(M
65000-80000) v IF
3
(M 29000). Kt qu ca giai on ny l to thnh
phc hp khi u: fMet-tRNA-mRNA-ribosome 70S.
Qu trnh c bt u bng s kt hp gia yu t IF
3
vi tiu n v
30S. Nh yu t ny m tiu n v 30S kt hp vi mRNA v tr khi
u. Trong giai on hai fMet-tRNA gn vi phc IF
2
-GTP to thnh
IF
2
-GTP-fMet-tRNA (t hp 3). giai on ba v bn phc hp IF
3
-30S
kt hp vi t hp 3. Nh yu t IF
2
m fMet-tRNA c a vo khu
vc P. Ngi ta cha bit chnh xc vai tr ca IF
1
, c th n tham gia vo
vic i mi chu trnh bng cch gp phn gii phng yu t IF
2
ra khi
phc hp. Cui cng tiu n v 50S kt hp vi tiu n v 30S to ra
phc hp ribosome 70S, ng thi gii phng c 3 yu t khi u cng
nh gii phng c GDP v Pi.
sinh vt nhn chun, qu trnh tng hp polypeptide trong t bo
cht c tRNA khi u cng mang methionine nhng khng c formyl
ho. y cng c cc phn ng vi cc yu t khi u eIF
1
, eIF
2
v
eIF
3
tng t nh sinh vt nhn s.
222
Trn tiu phn ln c ba vng quan trng: vng A: l ni tRNA-
aminoacyl c kt hp, vng P: ni gn tRNA-peptidyl v vng E: thc
hin s ko di chui peptide (hnh 7.23).










Hnh 7.23 Nhng vng phn ng quan trng ca ribosome
A: v tr gn ca tRNA-aminoacyl, P: v tr gn ca peptidylaminoacyl

- giai on ko di chui polypeptide, amino acid th hai (do m tip
theo ca mRNA quy nh) gn vi tRNA ring ca n s c a vo
phc hp khi u vng A. Giai on ny cn GTP v cc yu t ko
di chui. sinh vt nhn s yu t ko di l EF-T (gm 2 b phn hp
thnh l EF-Tu v EF-Ts). Yu t ny sinh vt nhn chun l eEF-1. Cc
yu t ny l nhng protein: EF-Tu (M 43000), EF-Ts (M 74000). Chng
c vai tr a aminoacyl-tRNA vo vng A ca ribosome thng qua cc
bc sau:
EF-Tu + GTP EF-Tu. GTP (t hp hai)
EF-Tu.GTP + amonoacyl-tRNA aminoacyl-tRNA-EF-Tu-GTP
(t hp ba).
T hp ba ch kt hp vi vng A trong ribosome v vng P c fMet-
tRNA ( giai on sau l peptidyl-tRNA) chim ch.
y l giai on quyt nh bo m nh hng ng to ra lin kt
peptide gia aminoacyl-tRNA v peptidyl-tRNA.
Trc khi aminoacyl c gn kt vo peptidyl th n kt hp vi
GTP vng E ca ribosome to nn mt phc gm ba yu t. Sau khi
GTP thu phn th lin kt peptide c to thnh. Qu trnh ny c xc
tc bi enzyme peptidyltransferase, chuyn peptidyl n tRNA-aminoacyl
va kt hp vo (hnh 7.24). Enzyme ny c cu to ch yu t rRNA.
Phn phi protein 50S rRNA th hin y hot tnh ca
peptidyltransferase. Nucleic acid c hot tnh enzyme c gi l
ribozyme. Protein ca ribosome tham gia qu trnh enzyme l do s thay
Tiu n v nh
mRNA
Tiu n v ln
P A
E
223
i cu hnh. i vi hnh th ca tng thnh phn ribosome ion Mg
+
c
mt ngha quan trng.
Sau khi kt hp vi ribosome, GTP b thu phn ng thi gii
phng t hp EF-Tu-GDP ra khi ribosome. Phc hp EF-Tu-GDP khi tc
ng tng h vi EF-Ts v GTP s li bin thnh EF-Tu-GTP. Do n
c th lai tng tc vi aminoacyl-tRNA tip theo. Cc nghin cu cho
thy s lng yu t EF-Tu bng khong s lng phn t aminoacyl-
tRNA. T ngi ta cho rng phn ln cc aminoacyl-tRNA l nm
trong thnh phn t hp ba ch khng phi trng thi t do.
Hnh 7.24 biu din s hnh thnh lin kt peptit. V tr A
ribosome l vng nhn, v tr P l vng cho. vng nhn kt hp vi
tRNA-aminoacyl mi i vo (aminoacyl vi R
0
hnh 7.24). vng cho
th tRNA-aminoacyl kt hp vi ribosome cng nh tRNA-
polypeptidyl. tRNA-polypeptidyl mang mt chui polypeptide vi hai
aminoacid (R
1
v R
2
). v tr ny c hai thnh phn tRNA-aminoacyl v
tRNA-polypeptidyl tham gia phn ng: nguyn t H ca nhm amine ca
aminoacid gn kt vi polypeptidyl ribose ca tRNA bng lin kt ester.
Chui polypeptidyl c chuyn t cht cho n cht nhn. tRNA gn
vng cho by gi trng thi t do, ngha l ribose cui cng ca n c
nhm OH v tr C
3
t do. Lc ny chui polypeptide di thm mt n
v (aminoacyl) (hnh 7.24, bn phi). y l phn ng vn chuyn
peptidyl. Bng cch ny lin kt peptide c to nn v tRNA c gn
kt trc tr nn t do.
Phn ng ko di chui cn c s tham gia ca GTP v mt yu t
ko di khc l EF-G ( sinh vt nhn chun l eEF-2). Yu t EF-G l
mt trong nhng yu t c bn ca t bo, c khi lng phn t 72000.
Yu t ny khi kt hp vi ribosome gp phn to cho ribosome chuyn
dch mt khong cch bng mt b ba trn phn t mRNA, khi
peptidyl-tRNA chuyn t vng A sang vng P.
Sau khi kt hp ribosome dch chuyn n codon tip theo, tRNA
mang aminoacid tip theo c kt hp vo. Ribosome l h thng c ho
hc c kh nng chuyn ng tng t nh protein c.
- Giai on kt thc: Qu trnh tng hp si polypetide kt thc khi trn
mRNA c mt codon kt thc xut hin hoc l n u cui ca
mRNA. Nhng codon kt thc l UAA, UAG v UGA. Si polypeptide
tch khi ribosome v t ng hnh thnh cu trc bc bn. Sau
ribosome b phn ly thnh cc tiu n v 30S v 50S ri nhp vo kho d
tr ribosome. S nhn bit cc b m kt thc c thc hin bi cc yu
t kt thc. l cc protein, sinh vt nhn s l RF
1
(M 4000) nhn
224
bit b m UAA v UAG v RF
2
(M 47000) nhn bit m UAA v UGA.
sinh vt nhn chun c mt yu t kt thc l eRF. thc hin phn
ng gii phng chui polypeptide th peptidyl-tRNA phi nm vng P,
cn cc yu t kt thc tng tc vi vng A ca ribosome.






















Hnh 7.24 S to thnh lin kt peptide ribosome
a) tRNA-amioacyl kt hp vi tRNA-peptidyl
b) Aminoacyl c chuyn ln peptidyl, tRNA gn vng cho tr nn t do,
si peptide di hn 1 aminoacid ang vng cho

Bng cch ny hng tng hp ca si polypeptide c xc nh,
v aminoacid ch c th c kt hp vi nhm carbonyl m khng kt
hp vi nhm amine. sinh vt nhn chun s tng hp polypeptide cng
bt u bng methionine, khng c formyl ho. Methionine v
Nhn din codon
Hnh thnh lin
kt peptide
Ribosome chuyn
dch v tr
Ribosome nhn
tRNA-aminoacyl
tip theo
225
methionine c formyl ho s tch ra bng thu phn khi qu trnh tng
hp polypeptide kt thc.
Qu trnh tng hp protein xy ra ribosome ca t bo cht v
ty th, lp th. Chng c h thng tng hp protein ring v c nhng
enzyme cn thit cho qu trnh . Tuy nhin trong ty v lp th khng
c tt c cc loi enzyme.



























Hnh 7.25 Cc tiu n v ca ribosome, polysome v mRNA xuyn qua cc
ribosome
R-CHO
tRNA-methionyl
tRNA-O-CH
2

O
O
N

CO

H
2
N -CH

CH
2
-S-CH
3



OHC-
tRNA-O-CH
2

O
O
N

CO
H
N -CH

CH
2
-S-CH
3



R-H
Formyl-tRNA-methionyl
DNA

RNA polymerase
Hng dch m
Polyribosome
Ribosome
Polypeptide
mRNA
226
Trong nhiu trng hp mt si mRNA thc hin gii m trn nhiu
ribosome, nh vy cng mt si mRNA c t 20 n 40 protein c tng
hp. S tp hp mt lot ribosome trn mt si mRNA c gi l
polysome. hnh 7.25 biu din s sp xp ca cc ribosome. Trn s
cho thy nhng ribosome no trt xa nht trn si mRNA th si
polypeptide c to nn l di nht.
Mt s khng sinh c th nh hng n qu trnh sao chp v gii
m. Qu trnh sao chp b c ch bi actinomycin D v rifamycin. Cht
th nht phong to base nit guanine, cht th hai c ch RNA-
polymerase. Qu trnh gii m b c ch bi puromycin, streptomycin v
chloramphenicol v thc ra l chng kt hp vi vng phn ng ca
ribosome. Khng sinh cng km hm qa trnh tng hp protein ca vi
sinh vt.
T cc vn nu trn ta c nhn xt rng: qu trnh sinh tng hp protein
v cng phc tp, c s tham gia ca nhiu yu t. Ngi ta xc nhn
c ti hng trm yu t tham gia vo qu trnh sinh tng hp protein trong
t bo.
V mt nng lng, qu trnh sinh tng hp l mt qu trnh thu nng
lng. vi khun cng nh sinh vt nhn chun, nhu cu nng lng
cho qu trnh l rt ln. a mt amino acid vo chui polypeptide cn
chi ph 3 lin kt cao nng:
- 1 ATP cho hot ho amino acid giai on u
- 1 GTP a amonoacyl-tRNA (trong t hp ba) vo vng A ca
ribosome
- 1 GTP cn ribosome di chuyn mt b ba trn mRNA
Do giai on khi u cn cn 1 GTP tham gia, nn tng hp lin kt
peptide u tin phi s dng ti 4 lin kt cao nng.
Ngi ta cho rng phn t GTP tham gia vo qu trnh ny c hai chc
nng: mt l to cu hnh thch hp cn thit cho s tng tc tip theo,
hai l bo m nng lng ribosome di chuyn trn mRNA.
V mt tc , qu trnh tng hp protein xy ra rt nhanh v ph thuc
vo nhit . vi khun nhit 37
0
C trong mt giy chui polypeptide
tng t 12 n 17 aminoacid. V vy, tng hp 1 phn t protein trung
bnh c ln 300 aminoacid mt khong 20 giy. Trong t bo sinh vt
nhn chun, tc tng hp chm hn, v d trong t bo hng cu li
37
0
C tc ko di chui peptide l 2 aminoacid trong 1 giy. So vi tc
giai on ko di chui th giai on khi u v kt thc xy ra chm
chp hn.
7.11 S phn gii cc hp cht cha nit

227
S phn gii protein, nh trong h tiu ho ca ng vt, ht ny
mm hoc trong t khi phn gii cc hp cht hu c nh vi sinh vt,
bt u bng s thu phn. Cc lin kt peptide b ct ra do cc peptidase,
xut hin cc aminoacid t do. Tu thuc vo loi peptidase m chui
peptide c thu phn gia hoc mt u chui.













qu trnh ny mm cc aminoacid c to ra c s dng
tng hp nn cc protein mi. S phn gii protein nh vi sinh vt trong
t v trong h tiu ho c thc hin tip theo l cc aminoacid b kh
amine ho, gii phng ra NH
3
. Nhng vi sinh vt sng trong t c kh
nng thc hin phn ng ny v vy c gi l vi sinh vt to amn.











ATP

ADP + P
i
ATP

ADP

P
i
Aspartate

Arginosuccinic acid

Fumaric acid

Arginine

Urea

Ornithine

Citruline

Carbamylphosphate

Carbamic acid

CO
2
+ NH
3
Lin kt peptide
R

CH NH COOH

H
2
N CO CH

R

H
2
O
R

CH H
2
N COOH

H
2
N COOH CH

R
228
Hnh 7.26 Chu trnh Ornithine

Nguyn tc ca s kh amine ho l phn ng oxy ho v s tch
NH
3
(kh amine ho oxy ho). trong ng vt, c bit l glutamate
c s kh amine ho bng cch oxy ho. Enzyme xc tc cho phn ng
ny l glutamate dehydrogenase. Trong iu kin ym kh nhng
aminoacid b kh amine ho bng phn ng kh (xem s ), mt qu
trnh xy ra trong t ngp nc, nh t la.








Kh amin ha bng cch oxy ha









Kh amin ha bng cch kh

Trong c th ng vt c v urea c to ra t NH
3
qua chu trnh
c gi l ornithine (hnh 7.26). Trong chu trnh ny gc carbamyl v
guanidyl ng vai tr quan trng. C hai u xut pht t urea, nh cng
thc sau:



Aminoacid

COOH

C=NH

R



COOH

H
2
N -CH

R


Iminoacid

COOH

C=O

R


Cetoacid
FAD
FADH
2
H
2
O
NH
3
Aminoacid

COOH

H
2
N -CH

R


2e
-
NH
3

COOH

CH
2

R


Monocarboxylic acid
2H
+
229













Chu trnh bt u vi s tng hp carbamic acid (xem s ). y l
phn ng cn bng t nhin. Cht phn ng c cung cp do qu trnh
h hp (CO
2
) v qu trnh kh amine ho bng cch oxy ho (NH
3
).
Carbamic acid c phosphoryl ho nh ATP to thnh
carbamylphosphate:






Carbamylphosphat phn ng vi ornithine to citrulline v
phosphate v c c tch ra.








COOH

H
2
N-CH
NH
2

CH
2


C = O
CH
2

O
CH
2
- HN H P

Ornithine Citruline
P
i
COOH

H
2
N-CH

CH
2
NH
2

CH
2
C = O

CH
2
- NH

NH
2


C = O

NH
2

Urea
NH
2


C = NH


Gc guanidyl
NH
2


C = O


Gc Carbamyl
NH
2


C = O

OH
Carbamic acid


C = O

O
+ HNH
2
NH
2


C = O

OH
Carbamylphosphate
NH
2


C = O

O ~ P
ATP
ADP
230
y nguyn t H ca nhm amine cui cng c thay th bng
nhm carbamyl. Citrulline phn ng vi aspartic acid to thnh
argininosuccinic acid. Phn ng cn ATP.











Qua s bin i ni phn t cht ny c tch thnh arginine v
fumaric acid.











Arginine l 1 dn sut ca guanidyl. Nh arginase (hydrolase)
arginine c tch ra thnh ornithine v urea.





COOH

H
2
N-CH

CH
2
NH
2

CH
2
C = O

CH
2
- NH

COOH

H
2
N-CH

CH
2


COOH

Aspartic acid
COOH

H
2
N-CH

CH
2
NH
2
COOH

CH
2
C = N - CH

CH
2
- NH CH
2


COOH

Argininosuccinic acid
ATP
ADP + P
i
COOH

H
2
N-CH

CH
2
NH
2

CH
2
C =

CH
2
- NH

COOH

N - CH

HCH

COOH

Arginine
COOH

H
2
N-CH

CH
2
NH
2
COOH

CH
2
C = N + CH

CH
2
- NH CH

COOH

Fumaric acid
231









Sau khi to thnh ornithine chu trnh mi li bt u. Cht i vo
chu trnh: CO
2
, NH
3
, asparagine v ATP. Chu trnh to nn ur v
fumaric acid. Hai nguyn t N ca ur bt ngun t NH
3
v asparagine.
Nh vy NH
3
, dng c c bin i thnh dng khng c, v c
thi ra cng vi nc tiu. Fumaric acid l cht trao i ca chu trnh
Krebs v cng c tip tc bin i . Vic s dng ATP y ni
ln rng, vic gii c NH
3
cn nng lng. Khung carbon ca fumaric
acid bt ngun t asparagine. Nh vy mt aminocid trong qu trnh phn
gii ca chu trnh Krebs ni vi chu trnh ornithine.
Nhng enzyme ca chu trnh ornithine phn th nh v trong c cht
ca ty th l glutamate dehydrogenase v citrullinsynthetase, nhng
enzyme cn li ca chu trnh nh v trong t bo cht.
Trong thc vt bc cao cng c cc enzyme ca chu trnh ornithine.
Trong ht ny mm arginase v ornithin-transcarbamylase c tm thy.
Trong enzyme cui cng xc tc cho phn ng tng hp citrulline t
carbamylphosphate v ornithine. V vy ngi ta cho rng, nhng enzyme
k trn c vai tr quan trng i vi vic huy ng cc protein d tr.
Trong cy nho arginine l mt dng d tr nit, c tch lu vo ma thu
trong cnh (g).
Trong nhiu trng hp ur c tch thnh CO
2
v NH
3
trong t
nh urease, nh vy nit c bn dng ur c tip nhn dng
NH
4
+
hoc sau c bin i nh vi sinh vt sang dng NO
3
-
.





NH
2


C = O

NH
2

Urease
2NH
3
+ CO
2
+ H
2
O
NH
2


C = O

NH
2

NH
2


C = NH

OH
COOH

H
2
N-CH NH
2

CH
2
C = NH

CH
2
OH

CH
2
- NH H

Urea
COOH

H
2
N-CH

CH
2


CH
2


CH
2
- NH
2

Ornithine
+
232
S bin i NH
3
trong t thnh NO
3
-
c thc hin nh vi khun,
chng oxy ho (t chy) NH
3
cng nh HNO
2
thu c nng lng
cho qu trnh trao i cht. Nhng loi vi khun quan trng nht oxy ho
NH
3
l loi nitrosomonas, nitrosocystus, nitrospira. S oxy ho cn c O
2

tng ng theo phng trnh sau:

NH
3
+ 3/2 O
2
HNO
2
+ H
2
O

S oxy ho HNO
2
thc hin nh loi nitrobacter v tng ng vi
phng trnh sau:

HNO
2
+ 1/2 O
2
HNO
3


qu trnh ny mt acid mnh, HNO
3
c to thnh. l
nguyn nhn thc t s phn gii cc cht hu c cha nit lm cho t
chua. Qu trnh to nitrite (HNO
2
-
) v nitrate trong t c gi l qu
trnh nitrate ho. Nhng loi vi khun cn cho qu trnh ny c nhiu
trong t. t trng thng thong vi pH trung tnh l iu kin l
tng i vi nhng vi khun ny.
Trong iu kin ym kh c th xy ra s kh NO
3
-
(kh nitrate h
hp hoc l d ho). Qu trnh ny c gi l phn nitrate ho. Ngy nay
ngi ta bit 23 loi khc nhau c kh nng ny. l Alcaligenes,
Agrobacterium, Azospirillum, Holobacterium, Pseudomonas v
Rhizobium. Chng c kh nng to ra nhng enzyme kh trong iu kin
thiu O
2
, s dng O trong NO
3
-
cho h hp ho kh, l cht nhn e
-
ca
chui h hp. qu trnh ny NO
3
-
b kh thnh N
2
O v N
2
. Trong qu
trnh phn nitrate ho cng xut hin NO. Trnh t phn ng c gii
thch nh sau:









2H
2
O
4e
-
+ 4H
+
2H
+
H
2
O

2NO
3
-
2NO
2
-
2HNO
2
NO
2
+ NO 2e
-
+ 2H
+




N
2
N
2
O 2NO H
2
O

H
2
O 2e
-
+ 2H
+
H
2
O 2e
-
+ 2H
+

233
qu trnh ny c 4 enzym kh tham gia: NO
3
-
-Reductase, NO
2
-
Reductase, NO-Reductase v N
2
O-Reductase. Enzyme quan trng nht l
NO
2
-
-Reductase, xc tc cho qu trnh chuyn ion NO
2
-
thnh dng cht
kh (NO). N
2
O-Reductase khng c trong tt c vi khun phn nitrate ho.
mt s vi khun qu trnh ny kt thc vi chui phn ng N
2
O. T
qu trnh phn ng rt ra rng, s phn nitrate ho to tnh kim, v chng
cn H
+
mi bc ca phn ng. S phn nitrate ho c mt vai tr ln
trong t nhin. T hp cht nit v c v hu c N
2
c gii phng li i
vo khng kh. S phn nitrate ho i lp vi s ng ho N
2
. Trong
qu trnh u tin sn sinh N
2
, qu trnh th hai cn N
2
. Nh vi khun c
nh m m hng nm t 100-200 x 10
6
t N t khng kh c a vo
t. Bng qu trnh phn nitrate ho m hng nm c khong 200-300 x
10
6
t N dng N
2
c gii phng ra trong khng kh. N
2
O c to ra
trong qu trnh phn nitrate ho l vn sinh thi kh khn. l cht
rt bn vng, bay ln n tng tnh kh v tham gia vo s phn gii
tng ozon.
Cu hi
1. Phn nhm amino acid v cho v d ?
2. Qu trnh kh nitrate v ngha ca qu trnh ?
3. Qu trnh c nh Nit phn t?
4. Cc ng hng gii c NH
3
thc vt ? C ch v ngha ca
chu trnh Ornithine?.
5. Mi lin quan gia chu trnh KREBS v chu trnh Ornithine ?
6. Cc ng hng tng hp amino acid trong thc vt ?
7. Phn gii amino acid?
8. Sinh tng hp v phn gii protein?


1
Chng VIII
Mi lin quan gia cc cht
trong trao i cht

Trn y chng ta nghin cu s trao i cht ca tng hp cht
c bn trong c th sng. Tuy nhin s phn chia nh vy ch gip lm
n gin ho nhng vn mun trnh by. Trong thc t gia cc cht c
nhiu mi lin h tng h. iu buc chng ta phi nghin cu s
trao i cht ca t bo trong mt tng th cc mi lin h.
Tht vy, trong c th khng th tm thy s trao i cht ca mt
cht no xy ra tch ri vi s trao i ca cc hp cht khc. Mi lin
quan tng h gia s trao i cc hp cht ny th hin trn hai mt c
bn: nguyn liu v nng lng.
Mi lin quan v mt nguyn liu l kh nng chuyn ho mt cht
ny thnh mt cht khc thng qua mt s sn phm trung gian. V d,
carbohydrate c th chuyn ho thnh aminoacid bng cch amine ho
mt s cetoacid. Ngc li mt s aminoacid c th chuyn thnh
carbohydrate bng cch loi nhm aminoacid thnh cc cetoacid, ri t
tng hp carbohydrate.
Mi lin quan v mt nng lng th hin ch: khi phn gii mt
hp cht no nng lng c tch lu trong ATP. Ngun ATP ny
c s dng cho cc phn ng tng hp. V d, ATP c to thnh
trong qu trnh ng phn, quang phosphoryl ho (quang hp) v ch
yu c to thnh trong qu trnh phosphoryl ho oxy ho (h hp). S
phosphoryl ho oxy ho qua chu trnh Krebs v chui vn chuyn in t
cho thy rng: bt k acetyl-CoA c ngun gc t carbohydrate, hoc acid
bo hay aminoacid cng u b oxy ho v tng hp ATP.
Nh kh nng chuyn ho tng h gia cc cht m c th sinh vt
thch ng vi mi trng. V d vo ma ng, cy trng xy ra s
chuyn ho tinh bt thnh ng v cht bo, nh kh nng chu rt
ca cy trng c nng cao. Hoc i vi mt s ng vt ng ng, do
d tr lipid ln, nn m bo cung cp nng lng v cc cht cn
thit cho c th s dng trong sut thi gian di ma ng.
nghin cu mi lin quan ny chng ta cn xem xt s trao i
ca tng cp hp cht sau:

2
8.1 Mi lin quan gia carbohydrate v lipid
Carbohydrate d dng chuyn thnh lipid thng qua hai hp cht
trung gian l dihydroxyacetonphosphate v acetyl-CoA
dihydroxyacetonphosphate glycerolphosphate
carbohydrate glucose

Glyceraldehydphosphate acetyl-CoA acid bo

- Glycerine hoc glyceraldehydphosphate c th bin i thnh
fructose-1,6- diphosphate (xem chng lipid).
- Acetyl-CoA thng qua chu trnh Krebs khng th tng hp c
carbohydrate v hai nguyn t carbon ca n b loi thnh CO
2

trc khi to ra hp cht oxaloacetic l cht c vai tr tng hp
mi carbohydrate (qu trnh gluconeogenese).
Khi m c b mt mi c hin tng l chu trnh Krebs khng th tip
nhn tt c cc phn t pyruvate to ra t qu trnh ng phn. Pyruvate
c kh thnh lactate, cht ny c tch lu v khi m c ngh ngi
chng li c oxy ho thnh pyruvate.









Sau pyruvate vo trong ty th v c carboxyl ho nh enzyme
pyruvatcacboxylase to thnh oxaloacetate. Pyruvate vo trong ty th v
c th tham gia vo 2 ng hng phn ng khc nhau (hnh 8.1). N c
th c bin i nh enzyme pyruvatdehydrogenase v sau i vo chu
trnh Krebs. Pyruvate cng c th c carboxyl ho to oxaloacetate
v chu s bin i theo con ng gluconeogenese. Theo ng hng
no l tu thuc vo nng acetyl-CoA, cht c kh nng hot ho
enzyme pyruvatcarboxylase theo c ch bin cu, trong khi ADP li c
ch enzyme ny.


NADH + H
+

NAD
+
COOH
CH
3

CO Pyruvate
COOH
CH
3

HCOH Lactate
3










Trong t bo cht malate c oxy ho thnh oxaloacetate nh
NAD
+











Oxaloacetate c bin i tip tc thnh photphoenolpyruvate nh
enzyme quan trng ca gluconeogenese l phosphoenolcarboxykinase.
Phn ng ny gm 1 phn ng kh carboxyl ho v 1 phn ng
phosphoryl (phn ng kinase). Phn ng cn 1GTP, tng t ATP.










COOH
COOH
CH
2
C=O
Oxaloacetate PEP
COOH
CH
2

C - O ~
P
GTP
GDP+CO
2
PEP-carboxykinase
CO
2
ATP

ADP + P
i

COOH
CH
3

C = O
Pyruvate
COOH
CH
2

Oxaloacetate
COOH
C = O
Pyruvat-carboxylase
NAD
+

NADH+H
+
COOH
CH
2

Malate
COOH
HCHO
Malat-dehydrogenase
COOH
CH
2

Oxaloacetate
COOH
C = O
4
PEP l 1 cht c to ra trong qu trnh ng phn. T cht ny i
ngc li nhng phn ng ring l ca qu trnh ng phn cho n
fructozo-1,6-diphotphate. Cht ny c kh phosphoryl ho v ng
phn ho cho n glucose cng nh glucose-1-phosphate. Cc nguyn t
C trong lactate c s dng tng hp nn glucose hoc glycogen nh
ATP v NADH. S phosphoryl ho trc tip pyruvate to thnh PEP l
khng thc hin c v l do nng lng, ngha l mc nng lng ca
pyruvate thp hn nhiu so vi mc nng lng ca PEP. Nh hnh 8.1
s bin i ny c thc hin qua oxaloacetate, nhm i quanh dc
ng gia pyruvate v PEP. Con ng ny i hi mt nng lng b
sung, l 1 ATP cho carboxyl ho pyruvate v 1 GTP cho to thnh PEP.
Hai ngun nng lng ny tng hp PEP, lin kt cao nng ca
chng gii phng 62 kJ/mol.























5





























Hnh 8.1 S Gluconeogenes

Nh trn nu enzyme pyruvatcarboxylase c iu khin theo
c ch bin cu. S iu khin ny l mt c ch c ngha. Khi nng
acetyl-CoA cao th khng cn to acetyl-CoA b sung theo con ng
kh carboxyl ho bng cch oxy ho. Trong trng hp ny
pyruvatcarboxylase c hot ho nh acetyl-CoA v pyruvate c s
dng to oxaloacetate. Ngc li khi nng ADP cao, ngha l nng
ATP thp th pyruvatcarboxylase b c ch. Pyruvate c thc hin
kh carboxyl ho bng cch oxy ho cung cp acetyl-CoA cho chu
trnh Krebs, l chu trnh cng vi chui enzyme h hp ti to ATP.
Glucose
1,6 Fructosediphosphate
Triosephosphate
Phosphoglycerate
PEP
Pyruvate
Acid bo
Pyruvate
Acetyl-CoA
Malate
Oxaloacetate
Glutamate
Aspartate Chu trnh Krebs
Ty th
T bo cht
GDP+CO
2
GTP
NADH+H
+

NAD
+
NAD
+
NADH+H
+
CO
2
+ATP
ADP+P
i
NH
3
NH
3
Oxaloacetate



-cetoglutarate
6
thc vt v mt s vi sinh vt c chu trnh glyoxilate xy ra
glyoxisome, cu trc ny cha enzyme -oxy ho peroxesome. y
acetyl-CoA c ngun gc acid bo s to thnh oxaloacetic acid, sau l
phosphoenolpyruvate, ri t to glucose.
C mi vng chu trnh glyoxilate (xem chng lipid) t 2 phn t
acetyl-CoA to ra c 1 phn t succinic acid. Acid ny b oxy ho
to ra oxaloacetic acid. Oxaloacetic acid s b kh carboxyl ho bin
thnh phosphoenolpyruvic acid. Cht ny s chuyn thnh glucose-6-
phosphate.
Nh vy t 4 phn t acetyl-CoA to ra 2 phn t succinic acid,
sau 2 phn t oxaloacetic acid. Cc bin i tip theo s cho ra 2 phn
t phosphoenolpyruvic acid. Cui cng thu c 1 phn t glucose.

8.2 Mi lin quan gia carbohydrate v protein
S phn gii carbohydrate to ra mt s -cetoacid, khi amine ho
chng s to thnh cc aminoacid tng ng. Cc -cetoacid l:
pyruvic acid alanine
-cetoglutaric acid glutamic acid
oxaloacetic acid aspartic acid
Cc aminoacid ny nh kh nng chuyn ho ring l s to thnh
cc aminoacid khc. Chng hn t glutamic acid c th tng hp thnh
proline (hnh 7.12). Aspartic acid c th to thnh lysine, threonine,
isoleucine v methionine.












7



















8.2 Cc ng hng tng hp aminoacid trong thc vt

Mt sn phm khc ca s phn gii carbohydrate l 3-
phosphoglycerate. Cht ny to thnh serine bng cch sau:

COOH NAD NADH +H
+
COOH
Glucose

NH
3

H - C - OH C =O
dehydrogenase transaminase
CH
2
O-P CH
2
O-P
3-phosphoglycerate 3-phosphohydroxypyruvate





Glucose
Fructose 1,6-bisphosphate
3-Phosphoglycerate
Phosphoenolpyruvate
Pyruvate
Acetyl-CoA
Citrate
Isocitrate
Oxalosuccinate
-cetoglutarate Succinate
Fumarate
Malatee
Oxaloacetate
Proline
Glutamate
Arginine
Glutamine
Histidin
Omithine
Citrulline
Cysteine
O-Acetylserine
Serin
e
Glycine
Tryptophan Tyrosine Phenylalanine
Chorismate
Shikimate
Prephenate
Anthranilate
Alanine
DAHP
Erythrose 4-phosphate
Asparagine
Aspartate
Lysine
Threonine
Methionine
Valine Isoleucine Leuxine
Aspartate
4-phosphate
Aspartate
4-semialdehyde
2,3-Dihydro-
dipicolinate
Homoserine
Homoserine
4-phosphate
DAP
Cysteine
Cystathionine
Pyruvate
Homocystein -ketobutyrate
Pyruvate
8
COOH H
2
O COOH

H-C-NH
2
H- C-NH
2

photphatase
CH
2
OP CH
2
OH
3- phosphoserine serine

T serine c th to thnh cysteine hoc glycine (nh phn ng loi b
nhm gc).
Ngc li mt s aminoacid (alanine, phenylalanine, tyrosine,
histidine, tryptophan, serine, cysteine , glutamic acid , proline, aspartic
acid) c coi l cc aminoacid to glucose. Trong qu trnh trao i cht
cc aminoacid ny to thnh pyruvic acid hoc mt hp cht trung gian
ca chu trnh Krebs nh -cetoglutarate, oxaloacetate. T oxaloacetate c
th to ra phosphoenolpyruvate, ri t s tng hp nn glucose.
Ngoi ra gia qu trinh d ho glucid (chu trnh Krebs) v qu trnh
d ho aminoacid (chu trnh ornithine) c nhng giai on to ra nhng
sn phm trung gian ging nhau, l aspartic acid, glutamic acid,
fumaric acid. iu chng t s trao i carbohydrate cng nh hng
qua li i vi s trao i protein.

8.3 Mi lin quan gia lipid v protein

Acid bo l tin cht ca mt s aminoacid thng qua cc qu trnh
sau: hoc qua chu trnh Krebs, mt s aminacid c tng hp t -
cetoglutaric acid, hoc qua chu trnh glyoxilate, to ra oxaloacetic acid.
Oxaloacetic acid b loi carboxyl to thnh pyruvate t 2 cht ny mt s
aminoacid cng c tng hp (hnh 8.2).
Ngc li mt s aminoacid (leucine, isoleucine, tryptophan) khi
phn gii s to thnh acetyl-CoA, t tng hp nn acid bo. Mt s
aminoacid khc (alanine cysteine, serine) b phn gii thnh pyruvic acid.
Theo con ng tng hp mi glucose, pyruvic acid s to thnh 3-
phosphoaldehydglyceric (ngc qu trnh ng phn).
S chuyn ho protein thnh lipid c th khi qut trong s sau:




9

Protein



aminoacid


leucine, alanine,
isoleucine, cysteine,
tryptophan serine


acetyl-CoA pyruvic acid


acid bo 3- phosphoaldehydglyceric


glycerine




Lipid


Cu hi
1. Ti sao trong ht ng cc khi ny mm hm lng carbohydrate
gim xung th hm lng lipid v protein li tng ln ?
2. Ti sao ht cy c du ny mm hm lng lipid gim xung th
hm lng carbohydrate v protein li tng ln?
3. Ti sao ni pyruvic acid l ng ba ng ca s trao i cht?






248
CHNG IX
CC CHT C NGUN GC TH CP

Ngoi cc cht carbohydrate, lipid, protein, nucleic acid, trong
thc vt cn tn ti nhng hp cht khc nhau do qu trnh trao i cc
cht ni trn to thnh. Mc du chng ch chim mt t l rt t trong cy,
nhng cc cht ny quy nh tnh c th ca s trao i cht trong tng
loi thc vt. Cc hp cht ny c gi l cc cht c ngun gc th cp.
Mt s trong chng v d nh cc acid hu c va mi c to
thnh c t bo s dng vo cc qu trnh tng hp khc nhau, do
vy chng khng c tch ly vi lng ln m l cc sn phm trung
gian ca qu trnh trao i cht.
Mt s cht khc v d nh cc hp cht phenol, tannin, alkaloid,
tinh du li c tch ly vi lng ln, do c tnh c th v trao i
cht.
Trong phm vi gio trnh ny, chng ti xin gii thiu mt s nhm
hp cht quan trng.
1. Tannin: Tn gi tannin c xut vo nm 1796 k hiu
nhm hp cht cha trong mt s cy vn dng thuc da ng vt.
T tan c ngun gc t ting latinh ca cy si, v t v cy si
ngi ta thu c cht thuc da.
Cc tannin c chia thnh 2 nhm: Cc tannin c th thy phn c
v cc tannin khng th thy phn c.
1.1. Cc tannin c th thy phn c: C c tnh ester. Chng
l dn xut ca gallic acid v protocatechic acid. Hai gallic acid kt hp
vi nhau to thnh digallic acid. Acid ny c vai tr trong vic to thnh
tannin.












OH


HO COOH


OH Gallic acid

OH


HO COOH


Protocatechic acid
OH OH


HO CO O COOH


OH OH Digallic acid




249
1.2. Tannin khng th thy phn c: Trong thnh phn c
catechin v gallocatechin. Hai cht ny khi phn ng vi gallic acid s cho
ta catechingallate v gallocatechingallate.
Theo Kursanov th tannin ca ch xanh c 12% gallic acid t do;
78% catechingallate v mt t catechin t do.
2. Cc acid hu c: c hnh thnh do qu trnh trao i cht,
chng tham gia vo chu trnh Krebs v nhng bin i trung gian khc.
Tuy nhin mt s thc vt cng tch ly mt s acid hu c khc nhau
cc c quan khc nhau nh l, qu, thn,chnh nhng acid ny gy
nn pH khc nhau ca t bo. C 2 nhm acid hu c:
- Nhm d bay hi:thng c mi hc nh acetic acid, butyric acid
- Nhm khng bay hi:thng l cc acid hu c cha nhm -OH
hoc nhm cetone nh pyruvic acid, glycolic acid, oxaloacetic acid, malic
acid,
Sau y l mt s acid hu c quan trng:
2.1. Citric acid: Acid ny c ph bin trong cy trng. Trong qu
cc cy thuc h cam, qut, citric acid l acid hu c ch yu. chanh
hm lng citric acid t ti 9% trng lng kh. Citric acid c s
dng rng ri trong cng nghip thc phm v nh l cht bo qun trong
truyn mu.
2.2. Malic acid (HOOC - CH
2
CHOH - COOH): Malic acid
chim u th qu to ty v qu cc cy c ht. Malic acid c ht v
l cc cy ng cc, cc cy u . Trong thuc l v thuc lo, hm
lng malic acid t 6,5%. Malic acid cha nhiu cc cy mng nc
(l bng, xng rng), c chua,
Malic acid c hng v d chu, khng c i vi c th ngi.
N c s dng rng ri sn xut nc qu v bnh ko.
2.3. Oxalic acid (HOOC - COOH): C ph bin trong cy trng
dng t do cng nh dng mui. qu, oxalic acid thng c lng
rt nh 0,005-0,06% nhng trong l mt s cy nh chua me, oxalic acid
li c mt hm lng ln.
2.4. Tartric acid (HOOC - CH(OH) - CH(OH) - COOH): Tartric
acid thng c trong cy di dng D-tartric acid, cng c th dng
L-tartric acid. N cha nhiu trong qu nho. Tartric acid v mui Kali ca
n c s dng trong sn xut nc qu, trong cng nghip ho cht v
cng nghip dt.
3. GLYCOSIDE: L nhng dn xut ca carbohydrate (thng l
dn xut ca monose) vi cc hp cht c bn cht ho hc rt khc nhau




250
khng phi l carbohydrate. Phn khng phi carbohydrate c gi l
aglycon (Aglycon c th l ru, cc hp cht thm, cc alkaloid, )
Ty thuc vo kiu lin kt gia phn carbohydrate v aglycon m
ngi ta chia ra cc nhm sau:
3.1. S-glycoside: Aglycon kt hp vi carbohydrate qua lu hunh.
V d: Sinigrin c ci ng, c mi hc






3.2. N-glycoside: Aglycon kt hp vi carbohydrate qua N.
V d: cc nucleotide
3.3. C-glycoside: Aglycon kt hp vi carbohydrate qua C.
3.4. O-glycoside: Aglycon kt hp vi carbohydrate qua Oxi (O)
V d: - Solanin: rt ph bin trong mm khoai ty, trong thn l
khoai ty.
- Saponin: khng cha nit trong phn t ca chng. Saponin c
kh nng ph hy hng cu, lm ho tan hng cu. Saponin c nhiu cy
Offininalis, hm lng Saponin t cc i thi k bt u ra hoa v cc
tiu thi k chn qu.
sn c loi glycoside sinh ra HCN. l linamarine v
lotostraloside. Cc glycoside ny tan trong nc. Sn phm ca s phn
hy ny l glucose v aglycon. Phn hy aglycon ny s to ra HCN:













S.C
6
H
11
O
5

CH
2
= CH CH
2
NC
OSO
2
OK
Sinigrin
C
6
H
11
O
5


O OH

R C C N + H
2
O C
6
H
12
O
6
+ R C C N
glucose Aglycon
CH
3
CH
3


R

Nu R l CH
3
- th l linamarine HCN + C = O
Nu R l C
2
H
5
-

th l lotostraloside
Acetone CH
3





251
4. Alkaloid: L nhng cht d vng cha nit, c c tnh kim v c
tc dng sinh l mnh; nhiu alkaloid l nhng cht c.
i b phn alkaloid tc ng n h thn kinh. liu lng thp
chng c tc dng kch thch, cn liu lng cao chng c tc dng c
ch.
V d: Cocaine c s dng trong y hc nh l cht gim au ti ch,
tc ng n nhng u cm gic ca h thn kinh ngoi bin. Trong qu
c c dc cha atropine l cht tc ng mnh n cc dy thn kinh
vn ng ca mt, lm cho ng t dn ra.
Sau y l mt s alkaloid in hnh:
4.1. Hordenine: C trong ht i mch lc ny mm vi mt lng
khong 0,2%. Tc dng ca hordenine l lm tng huyt p ca ngi v
ng vt. Trong thc vt, hordenine c to thnh t aminoacid l
tyrosine, theo s sau:















4.2. Ricinin: l alkaloid c cy thu du (trong ht cha khong
0,15%, trong l non gn 1%). Ricinin c c tnh v cha nhm CN. Do
hm lng ricinin c nhiu trong b kh thu du (n 0,2%) nn ngi ta
khng dng kh du thu du lm thc n cho gia sc.







O CH
3


CN


O
N CH
3

Ricinin
CH
3

H
2
N CH COOH CH
2
N

CH
2
CH
2
CH
3

CO
2
2CH
3
-


- 2H


OH OH
Tyrosine Tyramine Hordenine

H
2
N CH
2


CH
2







OH




252
4.3. Piperin: c nhiu trong t, trong ht h tiu (t 5 9%).
Piperin khng c, n ch gy cm gic cay nng tng b phn.












4.4. Atropin: C nhiu trong qu c c dc v trong ht cy h
c. Atropin c tc dng mnh ln h thn kinh hot ng ca mt. Atropin
c th c dng lm cht gii c khi b ng c bi nicotin. Cc liu
lng iu tr cao ca atropin l 0,001-0,003 gam. liu lng cao hn
th atropin l mt cht c.









4.5. Cocaine: l alkaloid ch yu ca loi cy coca min Nam
M. Cy ny hin nay cng c trng n v trn o Java. Hm
lng cocaine trong l t t 1-2%.









O O





N
Piperin
CO - CH = CH CH = CH




N CH
3
O CO CH C
6
H
5


CH
2
OH


Atropin


CO O CH
3




N - CH
3
O CO C
6
H
5



Cocaine







253
Cocaine lm t lit h thn kinh cm gic v c dng lm cht
gy t b phn. Ngoi ra n cng c tc dng ln h thn kinh trung ng
v gy nn cm gic say rt t trng. Khi ung nhiu ln cocaine, c th
s quen vi cocaine v s b bnh nghin cocaine. Liu lng c ca
cocaine i vi ngi lm l 0,2gam.
4.6. Morphin: Morphin v methylic ether ca n tc l codein c
ngha nht.













- Morphin l mt cht an thn l lm gim au. N c tc dng ln
h thn kinh trung ng v thn kinh ngoi vi.
4.7. Cafein: C trong ht c ph (1-3%), trong l ch (n 5%),
cn c trong cacao v mt s cy khc.












Cafein kch thch h thn kinh trung ng v hot ng ca tim.
Di tc dng ca cafein th huyt p s cao.
Cafein c tc dng thng tiu. Cafein c dng trong vic iu tr
cc bnh v tim v c dng lm thuc li tiu.

HO



O


N CH
3

HO Morphin





H
3
C O



O


N CH
3


HO Codein

O


H
3
C N N CH
3



O
N N
Cafein
CH
3





254
4.8. Nicotin: Trong thuc l (1 10%) trung bnh 4 %. Nicotin l
mt cht rt c. N c tc dng mnh ln h thn kinh trung ng v h
thn kinh ngoi vi. Di tc dng ca nicotin xy ra s co mch mu do
huyt p tng mnh. Con ngi c th cht do t lit h hp khi ung vo
c th mt lng khong 0,01 hay 0,04 gam nicotin. V c c tnh cao
nn trong y hc ngi ta khng dng nicotin. Tuy nhin n cng c ng
dng rng ri trong th y chng cc bnh ngoi da hoc ng dng trong
vic chng su b ph hoi ma mng.









Nicotin c pht sinh t r. R pht trin tt th nicotin cng
nhiu. Trng tha, rung khng nc, ngt ngn sm th thuc l
hm lng nicotin cng cao. t tt, bn nhiu m, nicotin cng nhiu.
5. Terpen: L thnh phn ch yu ca tinh du. S c mt ca terpen
quy nh mi v ca nhiu thc vt (hoa hng, hoa nhi, l chanh, l s,
)
Mt s v d v terpen:
- Limonen: c trong tinh du thng, tinh du h chanh.
- Cimol c trong du khuynh dip, bch n.
- Terpinene, xivestren c trong du thng.
- Pinen c trong du thng.
- Menthol: c trong du bc h














N - CH
3

N Nicotin

CH
3







C = CH
2


CH
3

Limonen
CH
3







CH CH
3


CH
3

Cimol
CH
3







CH CH
3

CH
3

Terpinene




255












6. Cao su v nha kt (Gutta)
- Trong thc vt c to thnh t isopren:






n = 500 5000: gi l cao su
n = 100 : gi l gutta-percha
* Cao su l hp cht cao phn t polyisopren c hnh thnh trong
m cc cy thuc 300 tc khc nhau. Trong s nhng cy ny ch c cy
cao su Hevea Brasiliensis c s dng vi quy m cng nghip.
M cha cc ht cao su c tch ly trong cc t bo chuyn dng l
ng nha. cy cao su, m c hnh thnh v tch ly trong v cy v
cc ng nha vng. Nh c cc ng ni gia cc ng dn cnh nhau trong
cc ng nha vng m m c th chy ra t mt vng ln ca v cy trong
khi khai thc m.
* Nha kt (gutta) c hnh thnh trong cc cy tc Palaquium
(Sapotaceae) mc nhiu Malaysia. M cc cy ny khng chy ra d
dng nh m cao su, do vy thu hoch m phi cht cy. iu ny dn
n vic tiu dit dn ngun nha kt ban u Palaquium gutta.
Nha kt l cht do v nhit. Thut ng chuyn mn gi l cht nhit
do. nhit di 65
o
C n khng do v cng nhng nhit 65
o
C
th chuyn thnh dng mm, n hi nhng vn cha do.


CH
3





C = CH
2
CH
3

Xivestren




H
3
C
CH
3




Pinen
CH
3






OH

CH
3
CH
3

Menthol
CH
2
C = CH CH
2


CH
3

n





256
7. Cc cht iu ha sinh trng thc vt
Cc cht iu ho sinh trng thc vt l mt nhm cht c bn cht
ho hc khc nhau nhng u c mt tc dng iu ho qu trnh sinh
trng, pht trin ca cy v m bo mi lin h gia cc c quan, b
phn ca cy.
Cht iu ho sinh trng thc vt c chia ra lm 2 nhm:
a. Cc cht kch thch sinh trng.
b. Cc cht c ch sinh trng.
Hai nhm ny c tc dng i khng vi nhau v mt sinh l.
- Cc cht kch thch sinh trung bao gm cc cht m nng sinh
l c tc dng kch thch qu trnh sinh trng ca cy (gm cc nhm
cht auxin, gibberellin v cytokinin).
- Cc cht c ch sinh trng gy tc dng c ch ln qu trnh sinh
trng ca cy (gm cc cht nh abcisic acid (ABA), ethylene; cc cht
phenol, cc cht lm chm sinh trng, cc cht dit c, )
* Cc cht iu ho sinh trng do cy t tng hp gi l cht t
nhin, cn cc cht do con ngi tng hp bng con ng ho hc gi l
cht nhn to.

Bng phn loi cc cht iu ho sinh trng thc vt

Cht iu ho sinh trng t nhin Cht iu ho sinh trng nhn to

A. Cht kch thch sinh trng (Stimulator)


1. Auxin (IAA, IAN, PAA)

Auxin nhn to (IBA, -NAA; 2,4D; 2,4,5T,
MCPA)

2.Gibberellin (GA
1
, GA
2
, GA
3
,, GA
54
)



3. Cytokinin (Zeatin, diphenylurea)

Cytokinin nhn to (kinetin, BA, )
B. Cht c ch sinh trng (Inhibitor)

1. Abcisic acid (ABA)

Cht lm chm sinh trng (CCC, MH,
TIBA)

2. Ethylene

CEPA






257
3. Phenol


Cc danh php quc t:

IAA: -indol acetic acid.
IAN: -indol acetonitril.
PAA: Phenyl acetic acid.
IBA: -indol butyric acid.
NAA: -Naphtyl acetic acid.
2,4D: 2,4 dichlorophenoxyacetic acid.
2,4,5 T: 2,4,5 trichlorophenoxyacetic acid
MCPA: 4 chloro, 2 methyl phenoxyacetic acid.
ABA: Abcisic acid.
BA: Benzyladenin.
CCC: Chlorocholine chlorid.
MH: Malein hydrazide.
TIBA: Triiodobenzoic acid.
CEPA: Chloroethylenphosphoric acid.






258

A. CC CHT KCH THCH SINH TRNG THC VT
1. Auxin:
1.1. Lch s pht hin: Nm 1880, Dacwin pht hin ra hin
tng hng quang th gii thc vt.
Hng quang l kh nng hng v ngun nh sng chiu t mt
hng nh cc cy trng ca s lun lun vn ra ngoi ca s.
Hin tng hng quang c bit r rt nht i vi bao l mm
cc cy ho tho (Coleoptyl), v vy chng l i tng c dng nhiu
trong vic nghin cu tnh hng quang thc vt. Nu chiu sng t mt
hng n ngn bao l mm th s gy nn s un cong hng v ngun
sng, nhng nu chng trong ti hoc loi tr nh ngn ca bao l
mm th hin tng khng xy ra. ng cho rng nh ngn ca bao l
mm l ni tip nhn kch thch ca nh sng.
Sau Boyen Jensen pht hin ra rng nh ngn b loi tr
nu t tr li trn b mt ca bao l mm th n c kh nng gy phn
ng hng quang bnh thng nh trng hp cy nguyn vn. Nh vy
th mt cht gy hng quang no sn sinh trong nh ngn v vn
chuyn theo hng gc gy nn s sinh trng khc nhau ca cc m
pha di.
Paal (1919) ct ri nh bao l mm v t li ln b mt ct
nhng lch sang mt bn v trong ti. Hin tng un cong hng
ng xy ra mnh m nh trng hp c chiu sng mt hng. ng cho
rng nh ngn hnh thnh nn mt cht sinh trng no y, cn nh
sng xc nh s phn b ca cht v hai pha ca bao l mm.
Nm 1933 Kogh (H Lan) bo co rng ng tch c auxin
A v auxin B t nc tiu ngi. ng cng b trng lng phn t ca
auxin A l 328. Cc cht ny gy ra phn ng un cong mnh ca bao
l mm cy la mch. Nm 1934, Kogh v Kosterman tch c chng
t dch chit nm men v nm 1935 tch c chng t dch ca mi
trng nui cy nm Rhizopus.
Tip theo ngi ta xc nh bn cht ho hc ca auxin.
chnh l -indolylacetic acid (IAA). V lc u ngi ta cha xc nh
c s tn ti ca auxin trong cy nn ngi ta gi cht l
heteroauxin. Nhng nm v sau ngi ta ln lt tch c auxin t cc
i din ca thc vt thng ng khc nhau v chng minh rng y l
mt phytohormone quan trng nht tn ti trong ton b th gii thc vt.
ng thi ngi ta cng tng hp c rt nhiu cc hp cht khc
nhau nhng c hot tnh sinh l tng t nh IAA v thm ch nhiu cht
c hot tnh mnh hn nhiu so vi IAA v chng c s dng rng ri
trong sn xut nhm iu chnh sinh trng ca cy trng c li cho con





259

ngi. Chng thng c dng mch vng. l cc dn xut ca indol
nh -indol butyric acid (IBA); -indol propionic acid (IPA), Cc dn
xut cu naphtalen nh -naphtylacetic acid (-NAA); -naphtylacetic
acid (-NAA); Cc dn xut ca chlorophenoxyaxetic acid nh 2,4
dichlorophenoxyacetic acid (2,4D); 2,4,5 trichlorophenoxyacetic acid
(2,4,5T)






























1.2. Cc dng auxin trong cy:
Auxin c tng hp ch yu nh sinh trng ngn, t vn
chuyn n cc c quan khc nhau theo hng gc. Ngoi nh sinh

CH
2
- COOH


(IAA)
NH
CH
2
COOH






(-NAA)

CH
2
COOH



(-NAA)


O CH
2
COOH


Cl


(2,4D)

Cl
O CH
2
COOH


Cl


Cl (2,4,5T)

Cl


CH
2
CH
2
CH
2
COOH


(IBA)
NH





260

trng ngn, auxin cn c tng hp mt phn cc c quan cn non
nh l non, chi non, qu non.
Trong cy auxin tn ti hai dng: dng t do v dng lin kt.
* Auxin dng t do: l dng c hot tnh sinh hc. Chng tn ti
nh sinh trng ngn, sau n chi bn, u r v cc c quan non.
Cng xa nh sinh trng nng auxin t do cng gim dn.
Dng auxin t do ch yu trong cy l IAA. Auxin t do trong cy
chim mt t l rt thp. Nhiu ti liu cng b hm lng auxin t do
trong cy chim khong 5% auxin tng s.
* Auxin dng lin kt: l dng auxin ch yu trong cy. Hm lng
ca dng lin kt ny c th chim trn 90% lng auxin c trong cy.
IAA c th lin kt vi rt nhiu cc hp cht khc nhau nn chng khng
c hot tnh sinh hc hoc c hot tnh rt thp. IAA c th lin kt vi
ng; cc aminoacid v amide to nn cc auxin lin kt nh IAA-
glycoside, IAA-glucan, indolacetamide, indolacetylaspartate,
indolacetylglutamate, indolacetylinositol, indolacetylarabinose,








* Chc nng ca auxin lin kt trong cy ht sc quan trng. Trc
ht chng l ngun d tr auxin trong cy. S lin kt ca IAA vi ng,
vi cc aminoacid v cc cht khc lm cho chng bn vng vi cc tc
nhn c th lm ph hy auxin v l phng thc tt nht chng li s
tc ng ca cc peroxidase phn hy auxin. Ngi ta chng minh rng
auxin lin kt l dng auxin vn chuyn ch yu trong cy. V vy, s bin
i thun nghch ca IAA t do IAA lin kt di tc nhn kch thch
ca mi trng gp phn iu chnh s cn bng auxin trong cy v do
iu chnh qu trnh sinh trng ca cy.

1.3. S tng hp auxin trong cy:
C quan tng hp auxin l nh sinh trng ngn. T m phn
sinh nh, IAA c vn chuyn xung cc c quan pha di; auxin cn
c tng hp t hn trong cc c quan ang sinh trng nh l non,
qu non, r non,
COOH

CH
2
C = N CH

O H CH
2
COOH

NH Indolacetylaspartate





261

Ngi ta xc nhn cht tin thn tng hp nn IAA l
aminoacid tryptophan.
S tng hp auxin trong cy:































1.4. S phn gii auxin trong cy:
S phn gii auxin trong cy xy ra trong cc trng hp sau: Khi
c tha auxin do s tng hp qu nhiu auxin trong cy v sau khi auxin
gy hot tnh sinh l vi cy xong.
Vic lm mt hot tnh auxin c th xy ra bng hai con ng: S
oxy ho bng enzyme IAA-oxidase v s quang oxy ho.


CH
2
CH COOH

NH
2


NH Tryptophan


CH
2
C COOH

O

NH Indolpyruvic acid
NH
3

O
2



CO
2



CO
2


O
2
NH
3


CH
2
- CH
2


NH
2


NH
Tryptamine

CH
2
C = O

H

NH
Indolacetaldehyde

O
2


CH
2
COOH



NH (IAA)






262

* S oxy ho bng enzyme IAA-oxidase: y l enzyme oxy ho
IAA, enzyme ny hot ng rt mnh trong h thng r thc vt lm
mt hot tnh ca IAA vn chuyn xung l ni cui cng. Phn ng
oxy ho ca IAA xy ra nh sau:









* S mt hot tnh ca IAA c th xy ra do nh sng. Ngi ta
nhn thy rng nh sng t ngoi c th lm mt hot tnh ca IAA, bi v
cu trc vng ca phn t IAA hp thu nh sng t ngoi = 280 nm v
gy nn s phn gii IAA.

1.5. S vn chuyn ca IAA trong cy
Auxin c tng hp trong nh ngn v t y vn chuyn xung
cc c quan pha di. S vn chuyn ca auxin trong cy c tnh phn
cc r rt, theo hng gc, tc l t ngn xung r m khng vn chuyn
ngc li.
- C ch ca s vn chuyn phn cc ca IAA trong cy c
ngi ta a ra mt s da theo thuyt ho thm ca Goldsmith (1977)
nh sau:














IAA
o
Ngoi t bo
Trn



Bm H
+

H
+
H
+





Di

IAA
-
Cht mang IAA
-

H
+
(Protein)

IAA
o



IAA
o



IAA
-

Trong t bo


IAA
-



CH
2
COOH CH
2

IAA-oxidase
+ H
2
O
2
+ O
2

- H
2
O O
NH - CO
2
NH
IAA Methylenoximdol





263

Theo thuyt ho thm ca Goldsmith th:
- Ch c IAA
o
khng b oxi ho mi khuych tn qua mng t bo
pha trn v IAA
o
thm qua c mng lipit.
- Bn trong t bo, pH cao hn v c cc bm H
+
hot ng, do
IAA
o
b ion ho thnh IAA
-
. Nng IAA
-
tng ln v khng vn chuyn
ra khi t bo c v khng tan c trong lipid.
- Do nng ca IAA
o
trong t bo gim nn tn ti mt gradient
IAA
o
gia bn ngoi v bn trong t bo, IAA
o
s khuych tn vo t bo.
- IAA
-
c vn chuyn ra khi t bo nh cc cht mang protein
c hiu nm mng t bo pha di. Khi ra khi t bo, do pH thay i
nn IAA
-
IAA
o
v tip tc vn chuyn sang t bo di n.
Ngoi s vn chuyn phn cc ny, bng nguyn t nh du
ngi ta cn xc nhn c mt s vn chuyn ngc li (hng ngn)
nhng t l thp. S vn chuyn phn cc l qu trnh cn nng lng
ATP, cn s vn chuyn ngc li l qu trnh khuych tn.
Tc vn chuyn ca auxin trong cy rt thp, khong 1 cm/h
trong r v thn.
1.6. Hiu qu sinh l ca auxin i vi cy
Cc c quan khc nhau c hm lng auxin khc nhau. Hm lng
ny cn ph thuc vo tui cy, vo iu kin ngoi cnh. Cc c quan
cn non ang sinh trng c hm lng auxin cao hn cc c quan trng
thnh v c quan gi. Tuy nhin, phn ng sinh trng ca cc c quan
khc nhau ph thuc vo hm lng auxin l rt khc nhau. Ni chung
thn cy cm ng kch thch vi nng auxin cao hn chi v r.
Sau y l th minh ho mi quan h gia nng auxin v s
sinh trng:














%

+


200


100


0
10
-6
10
-4
10
-2
1 10
2
10
4
IAA

-100 R Chi Thn

-









264

a) Auxin v tnh hng quang, hng a:
* Hng quang (phototropism) l s sinh trng ca cy hng v
ngun sng kch thch mt hng.
* Tnh hng a (geotropism) l phn ng sinh trng ca cy
di tc ng ca lc ht tri t (trng lc).
T khi pht hin ra auxin ngi ta gii thch tnh hng quang
v hng a ca cy trn c s s phn b ca auxin hai pha ca c
quan khc nhau dn n s sinh trng khc nhau hai pha ca c quan
v kt qu l gy nn s un cong hng ng.
C s ca s phn b ca auxin khc nhau l do s khc nhau v in
th (in sinh hc).
Khi c chiu sng t mt hng th pha khut sng s tch in
dng v pha chiu sng s tch in m. Mt c quan nm ngang th mt
trn s tch in m v mt di s tch in dng. Ngi ta c th o
in th ca c quan trong cc trng hp .










trong t bo IAA thng dng IAA
-
v do m phn b v
pha mang in dng. Khi c nh sng chiu mt chiu th auxin phn b
v pha khut sng v hm lng ca IAA cao hn gy s sinh
trng mnh hn pha khut sng v lm cho cy sinh trng un cong
v pha chiu sng.
Trong trng hp hng a th vi c quan nm ngang, auxin s
phn b pha di nhiu hn, gy nn s sinh trng ca mt di
nhanh hn mt trn lm cho chi ngn sinh trng un cong ln trn. Tuy
nhin do r phn ng kch thch vi nng auxin thp hn nhiu so vi
thn, nn nng auxin cao s gy c ch v do m s sinh trng ca
mt di chm hn mt trn lm cho r m vo t.
b) Hin tng u th ngn v auxin:
Hin tng u th ngn l hin tng ph bin ca th gii thc
vt, khi m s sinh trng ca chi ngn, ca r chnh s c ch s sinh
trng ca chi bn v r ph. Nu loi tr chi ngn v r chnh th lp
_



+



+ nh sng







+





265

tc chi bn v r ph s sinh trng mnh. Trong thc t khi ngi ta
bm ngn, ta cnh, n cy, tc l loi b s c ch ca chi ngn,
to iu kin cho cc chi bn mc ra mnh m. y l c s ca k thut
to hnh cho cy n qu, cy cnh, cy cng nghip v ci to c qun
th cy trng.
Khi pht hin ra auxin, ngi ta cho rng auxin l tc nhn gy nn
hin tng u th ngn. Nu loi b chi ngn th lp tc chi bn mc ra,
nhng khi loi b chi ngn m x l IAA vt ct th chi bn cng
khng mc ln c, tng t nh trong trng hp c chi ngn tn ti.
C hai gi thuyt gii thch vai tr ca auxin i vi hin tng u
th ngn:
* Gi thuyt c ch trc tip ca IAA: Cho rng chi ngn l ni
tng hp IAA v do nng ca auxin trong chi ngn lun lun cao.
Trn con ng vn chuyn ca IAA xung di th gy nh hng c
ch ln s sinh trng ca chi bn v phn ng ca chi bn ln hm
lng auxin thp hn chi ngn. Trong trng hp ct b chi ngn tc l
lm gim hm lng IAA v do cc chi bn c gii phng khi
trng thi c ch.
* Gi thuyt c ch gin tip ca auxin: Cho rng auxin khng gy
nh hng trc tip nhng auxin gy nn s tng hp mt cht c ch
sinh trng no y nh ethylene chng hn; v n lt ethylene li gy
c ch sinh trng ca chi bn.
Tuy nhin d gi thuyt no th cng phi tha nhn vai tr ca
auxin ln hin tng u th ngn.
Thc ra, hin tng u th ngn c iu chnh bi s cn bng
hormone ca auxin/cytokinin. Auxin tng u th ngn, c ch chi bn,
cn cytokinin ngc li, lm gim u th ngn, kch thch chi bn. Auxin
c tng hp nh ngn v c vn chuyn xung r, nn cng xa
nh ngn th nh hng c ch ca auxin cng yu. Ngc li, cytokinin
c tng hp trong r v c vn chuyn t r ln ngn, nn cng xa r
nh hng kch thch ln chi bn cng yu dn. V vy, t ngn xung
gc th hin tng u th ngn cng gim dn, chi bn cng mc nhiu
hn.
c) Auxin kch thch s gin ca t bo:
Auxin lm tng kch thc ca t bo v do m lm cho cy
sinh trng c. Tc dng ca auxin c hai mt: Va gy s gin n ca
thnh t bo lm cho t bo phnh to ra v va hot ho cc qu trnh tng
hp cc vt cht cu to nn cht nguyn sinh. Auxin kch thch s gin n
ca t bo theo chiu ngang mnh hn l chiu dc.






266

d) Auxin vi s hnh thnh v s phn ha r :
Trong s hnh thnh r, c bit l r bt nh, auxin c tc dng
rt c trng. V vy ngi ta xem auxin nh l cht hnh thnh r.
S hnh thnh r bt nh ca cnh gim, cnh chit c th chia ra
lm 3 giai on:
- u tin l s phn phn ho t bo mnh m vng tin tng
tng, tr b, ni xut pht ca r bt nh thnh mt m t bo ln xn.
- Giai on tip theo l xut hin mm r.
- Giai on cui cng, r sinh trng, m thng v v ra ngoi
thnh cc r bt nh.
giai on u tin l giai on khi xng s hnh thnh r cn
hm lng auxin cao cho s phn phn ho t bo. Cc giai on sau cn
hm lng thp hn v i khi khng cn.
Ngun auxin c th l ngun ni sinh v cng c th b sung t
ngoi vo. Thng xc tin qu trnh hnh thnh r nhanh chng cnh
chit, cnh gim ngi ta thng x l auxin ngoi sinh.
Vai tr ca auxin vi s hnh thnh r c chng minh r trong
nui cy m t bo. Nu ch c mt ca auxin trong mi trng nui cy,
m ch pht sinh r m thi. to cy hon chnh th cn phi c chi
na, tc l phi b sung cytokinin vo mi trng nui cy.
e) Auxin vi s hnh thnh qu, s sinh trng v to qu khng ht:
Ht phn l ngun rt giu auxin, kch thch s ny mm ca ht
phn v s sinh trng ca ng phn. Ngoi ra nm nhy cng tit ra cc
cht kch thch s sinh trng ca ng phn v ng thi cng sn sinh ra
cc cht c ch s ny mm ca ht phn v s sinh trng ca ng phn
khc loi, gy nn s tuyt giao gia cc loi. Khi th tinh xong, hp t s
pht trin thnh phi v chnh phi cng l ngun tng hp auxin mnh
m ri khuych tn vo bu kch thch s ln ln ca bu thnh qu. V
vy kch thc v c hnh dng ca qu hon ton ph thuc vo hm
lng v s khuych tn ca auxin trong phi ht.
Hu ht cc trng hp s pht trin ca qu ch xy ra sau khi th
tinh xong, tc l hnh thnh phi v bt u sn sinh auxin ni sinh.
Nhng cng c trng hp qu c hnh thnh khng c th tinh v to
nn qu khng ht.
Nm 1934, Yasuda thnh cng trong vic gy nn qu khng
ht bu b bng cch x l dch chit ca ht phn ln hoa. Sau ngi
ta phn tch v thy trong dch chit ca ht phn c cha nhiu auxin. T
ngi ta x l trc tip auxin ngoi sinh cho hoa th cng c th loi
tr s th tinh v to nn qu khng ht v rng auxin trc tip khuych
tn vo bu, kch thch sinh trng ca bu thnh qu khng ht.





267

f) Auxin v s rng l, hoa, qu, :
Laibach (1933) ch ra rng trong dch chit ca ht phn phong
lan c cha mt cht c th km hm s rng. Cht l IAA.
Trc khi rng, c mt tng ri c hnh thnh cung ca c
quan thc vt. y l mt vi lp t bo nhu m c thnh mng, hon ton
thiu lignin v suberin. Khi hnh thnh tng ri th c 3 trng hp xy ra:
- Lp v gia ca t bo b phn hy, cn lp v s cp vn
gi nguyn.
- Tt c thnh b phn hy.
- Tt c t bo b phn hy.
Khi nghin cu hm lng auxin lin quan n s hnh thnh tng
ri cho thy rng l non c hm lng auxin cao hn l gi, bn l c
hm lng auxin cao hn cung l. Khi hm lng auxin cao s ngn s
hnh thnh tng ri. V vy, nu x l auxin ngoi sinh s lm tng hm
lng auxin trong l nn c th ngn nga c s rng.
Thc ra s rng ca c quan gy ra do s tch ly cc cht c ch
sinh trng (abcisic acid, ethylene). Do s cn bng gia auxin /
(abcisic acid+ethylene ) c nh hng n iu chnh s rng cy.
Auxin cn nh hng n qu trnh bin i sinh l-sinh ho trong
cy: S vn ng ca cht nguyn sinh, cc qu trnh trao i cht, tng
qu trnh quang hp v h hp, tng qu trnh vn chuyn vt cht trong
cy,

1.7. C ch tc ng ca auxin:
Hiu qu c trng nht ca auxin chnh l kch thch s gin ca
t bo. S gin ca t bo xy ra do hai hiu ng: s gin ra ca thnh t
bo v s tng th tch v khi lng ca cht nguyn sinh.
Khi x l auxin, khong sau 10 15 pht th phn ng sinh
trng xy ra. Vy th auxin ng vai tr nh th no trong s sinh trng
nhanh chng nh vy?
Trc ht, chng ta cp n mt hin tng c pht hin t
nm 1930 l hin tng sinh trng acid. Trong mi trng acid th s
sinh trng ca thc vt tr nn nhanh chng hn. Vy mi quan h gia
hiu qu acid v auxin ln sinh trng ca t bo l nh th no? Hai nh
hng c phn bit vi nhau l:
- nh hng ca acid l ln thnh t bo, cn nh hng ca auxin
l ln t bo.
- nh hng ca acid xy ra nhanh hn, cn nh hng ca auxin
th chm hn nhiu.





268

Hai nh hng c th lin kt vi nhau nh sau: Auxin gy nn
acid cho thnh t bo, cn acid th gy nn lng lo, khng bn
vng ca thnh t bo v chnh nh lng lo ca thnh t bo m
thnh t bo mi gin ra c.
Cng trong thi gian ngi ta pht hin ra nhng bm trao
i cht trong cht nguyn sinh. Mt trong nhng bm l bm ion H
+
.
Bm H
+
t cht nguyn sinh vo thnh t bo, lm gim pH trong
thnh t bo. IAA l tc nhn hot ho bm proton . Nh vy khi c
mt ca IAA th bm proton hot ng v pH ca thnh t bo gim
xung.
Ti sao pH thp li lm gin thnh t bo? tr li cu hi
chng ta cn xem xt cu trc ca thnh t bo: Thnh t bo c cu to
t cc si cellulose, gia chng c ni vi nhau bng cc cu ni
polysaccharide, to nn mt cu trc rt bn vng. cho cc si
cellulose lng lo, tch ri nhau th cn phi phn hy cc cu ni
polysaccharide. Enzyme phn hy cc cu ni ny l pectinmethylesterase
hot ng pH thp (= 4) v khi hot ng s methyl ho nhm
carbocyl v ngn chn s hnh thnh cu ni ion vi canxi to nn
pectatcanxi, nn cc si cellulose tch ri nhau, chng c th trt ln
nhau m gin ra, Nh kh nng ht nc thm thu ca t bo m to
nn sc trng tc ng ln thnh t bo, lm cho thnh t bo phi gin
n ra.
Tuy nhin, do t bo tng kch thc nhanh chng nh vy nn cn
c s tng hp mi v nhanh chng cc thnh phn cu to nn thnh t
bo cng nh cht nguyn sinh (cellulose, hemicellulose, pectin, protein,
nucleic acid, ). y, auxin ng vai tr hot ho cc gen tng hp
nn cc enzyme cn thit cho s tng hp nn cc vt cht .














DNA mRNA Protein-enzyme

Auxin
Sinh trng





269



S v c ch tc ng ca auxin ln s gin ca t bo:
























1.8. ng dng cc hp cht auxin trong trng trt.
a) Nhn ging v tnh cy trng:
Hin nay c hai phng php x l auxin cho cnh chit, cnh gim:
* Phng php x l nng hay phng php x l nhanh. Nng
auxin dao ng t 1000 n 10000 ppm (1ppm = 0,1 mg% hoc
= 1x10
-4
% tc 0,0001% hay = 1 mg/l). Vi cnh gim th nhng phn gc
vo trong dung dch trong 3-5 giy ri cm vo gi th. Cn vi cnh chit
th sau khi khoanh v, tm bng bng dung dch auxin c ri bi ln trn
khoanh v, ni xut hin r bt nh. Sau b bu bng t m. Phng
php ny hiu qu cao, n gin v ho cht tiu tn t hn.
* Phng php nng long-x l chm: Nng auxin t 20
n 200 ppm ty thuc vo loi v mc kh ra r ca cnh gim. i
DNA


mRNA
IAA










ATP
H
+

H
2
O
Protein
Enzyme h hp
Enzyme tng hp cc
cu t ca thnh t bo
v nguyn sinh cht
Cellulose, pectin,
Protein,
Tng th tch
t bo
T bo gin, th
tch, kch thc
t bo tng
Enzyme gin
thnh t bo
Thnh t bo
lng lo, gim
tnh bn vng
Bm H
+






270

vi cnh gim, ngm phn gc vo dung dch auxin trong 10 24 gi ri
cm vo gi th. Vi cnh chit th trn dung dch vi t b bu.
Cc auxin s dng y l IBA; -NAA; 2,4D;...trong hiu qu
ca IBA > -NAA > 2,4D.
b) ng dng auxin tng u qu, sinh trng ca qu v to qu
khng ht:
S hnh thnh qu l mt qu trnh din ra c lin quan n s th
phn v th tinh. S th phn l qu trnh m ht phn ri trn nm nhy.
Ht phn c cha nhiu auxin nn khi gp dch nm nhy, chng s ny
mm, ng phn mc v sinh trng ko di xung tn non. ng phn
mang cc tinh t xung th tinh cho t bo trng to nn hp t v sau
hp t s pht trin thnh phi, v sau phi pht trin thnh ht. Bu
hoa s sinh trng thnh qu.
Qu ch c hnh thnh sau khi c qu trnh th tinh xy ra, cn
nu khng th tinh th hoa s b rng. iu c gii thch rng phi
ht l ngun tng hp nn cc cht kch thch sinh trng trong c
auxin. Cc cht ny s c vn chuyn vo m ca bu kch thch bu
ln ln thnh qu. V vy, hnh dng v kch thc ca qu hon ton ph
thuc vo hm lng cc cht ni sinh t phi ht. Chnh v l do m ta
c th s dng auxin v gibberellin ngoi sinh thay th cho ngun ni
sinh.
Nu chng ta x l cho hoa cha xy ra th tinh th auxin ngoi
sinh s khuych tn vo bu v kch thch s ln ln ca bu thnh qu m
khng qua qu trnh th tinh. Trong trng hp ny, qu to nn khng
qua th tinh v do s khng c ht.
Mt s cy trng nh c chua, bu b, cam chanh,...ngi ta
thng x l auxin di dng -NAA (10 20 ppm); 2,4D (5 10
ppm). Cn cc cy trng khc nh nho, anh o,.... th x l gibberellin
(20 50 ppm) li c hiu qu hn.
c) Phng nga rng qu:
S rng l do s hnh thnh tng ri cung l, cung qu. Auxin
l cht km hm s rng. Chnh v vy m ngi ta x l auxin cho cy v
cho qu non c th km hm s rng ca chng. V d nh phun
-NAA (10 20 ppm) cho l hoc qu non c th ko di thi gian tn
ti ca chng trn cy.
d) Ko di s chn ca qu:
S chn ca qu c iu chnh bng t l auxin/ethylene. Mun
km hm s chn ta cn tng hm lng auxin trong m qu. Phun dung
dch auxin ln qu xanh hoc qu sp chn ang trn cy c th ko di
thi gian tn ti ca qu. Trc y ngi ta hay s dng 2,4D





271

(10 15 ppm). Hin nay ngi ta s dng -NAA(10 20 ppm) cng
c hiu qu tt m khng c hi.
e) Dit tr c di:
Khi s dng nng cao c tc dng dit tr c di hi cy trng.
Cc cht nh 2,4D; 2,4,5T trc y c s dng nhiu dit tr c
di, nhng ngy nay ngi ta to ra rt nhiu cht dit c mi c hiu
qu cao m khng c hi cho mi trng.

2. Gibberellin
2.1. Lch s pht hin: Vic pht hin ra gibberellin gn lin vi
nhng nghin cu v bnh la von m cc nh khoa hc Nht quan tm
t lu. Triu chng in hnh ca bnh l cy la tng trng chiu cao
qu mc, lm yu cy, gim nng sut trn 40%. Bnh la von l do nm
k sinh cy la c tn l Gibberela fujikuroi gy nn. Nm ny tit ra
mt cht no kch thch s sinh trng chiu cao ca cy la gy nn
bnh l.
Nm 1926, Kurosawa thnh cng trong vic gy bnh nhn to
cho cy la v cy ng. Nm 1935, Yabuta tch c hai hp cht di
dng tinh th t dch nui cy ca nm G.fujikuroi v ng gi l
gibberellin A v B nhng cha xc nh c bn cht ha hc ca chng.
Nm 1955 hai nhm nghin cu ca Anh v M ng thi
nghin cu vn ny. Brian (Anh) v Stodole (M) xc nh c
cng thc ha hc ca chng.
Sau cc nh khoa hc chit tch v xc nh GA t cc thc
vt bc cao khc nhau v xc nh gibberellin l mt phytohormone quan
trng v ph bin ca ton th gii thc vt.
Ngy nay ngi ta xc nh c trn 50 loi gibberellin khc
nhau v c k hiu l GA
1
; GA
2
; GA
3
, trong GA
3
(Gibberellic
acid) l c hot tnh mnh nht.












O



C = O GA
3
(Gibberellic acid)

HO OH



CH
3
COOH CH
2





272

2.2. S vn chuyn ca GA trong cy:
Khc vi auxin, gibberellin vn chuyn khng phn cc trong cy:
c th theo hung gc hoc hng ngn ty theo nhu cu v v tr ca c
quan cn GA.
2.3. S tng hp GA:
Cht tin thn ca s tng hp GA l acetyl-CoA:


Qu trnh tng hp GA cn c s tham gia ca ATP v NADPH
2
,
cng nh s xc tc ca Mg
+2
v Mn
+2
.
Qu trnh tng hp GA xy ra ch yu trong lc lp.
C th tm tt qu trnh tng hp GA nh sau:



























O

CH
3
C ~ S.CoA
2NADP CH
2
- COOH
2NADPH
2

3Acetyl-CoA C(OH) CH
3


3HS-CoA CH
2
CH
2
OH Mevalonic acid
2ATP



2CO
2


CH
3
C CH
2
CH
2
O (P) O (P)

CH
2
Isopentenylpyrophosphate


CH
2

CH
3





Kaurenic acid

CH
3
COOH

Gibberellin





273

Qu trnh tng hp GA dc xc tc bi cc enzyme c hiu. Tn
ti mt s t bin ln m c th thiu mt s gen nht nh iu khin s
tng hp nn cc enzyme xc tc cho s hnh thnh cc cht trung gian
no v kt qu l GA khng c hnh thnh, cy khng th sinh
trng chiu cao c. Vi cc t bin ny, nu x l GA ngoi sinh c
th ly li c dng bnh thng.
2.4. Hiu qu sinh l ca GA
a) Cc t bin ln ca thc vt:
Cc t bin ny do c s sai lch nht nh trong qu trnh tng
hp enzyme cn cho s tng hp nn GA; do GA khng c hnh
thnh nh bnh thng.
b) GA vi s gin v s sinh trng ca t bo:
Cng vi auxin, GA c nh hng quan trng ln s gin ca t
bo. GA kch thch gin ca t bo lm cho t bo gin v chiu dc. V
vy m t bo ko di rt nhanh chng. Kt qu kch thch s gin ca t
bo bi GA lm cho cy sinh trng nhanh v chiu cao. Mt s cy
cn chiu cao nh cy ay, cy gai, C th s dng GA ngoi sinh
iu khin chiu cao ca chng.
c) GA vi s ny mm:
Trong ht, ni nh ng vai tr th ng trong qu trnh ny mm,
n ch cha cht d tr. Khi ny mm th c cc qu trnh thy phn cc
cht d tr trong chng. S thy phn ny nh cc enzyme thy phn
trong amylase ng vai tr quan trng.
S hnh thnh nn cc enzyme thy phn ny do s hot ha ca
GA c tng hp hoc c gii phng t trng thi lin kt trong phi
ht. Chnh v vy m ngi ta c th kch thch cho ht ny mm bng
cch x l GA ngoi sinh.
Vi cc loi c: S ny mm ca chng cng c kch thch bi
GA c trong c. X l bng GA ngoi sinh cng ph b c trng thi
ng ngh ca c.
d) GA vi s sinh trng v to qu khng ht:
Hiu qu ca GA trong trng hp ny cng tng t nh ca
auxin. V vy nu x l GA ngoi cng c th thay th c GA ni sinh
trong phi ht v to nn c nhng qu khng c ht. Trong mt s
trng hp, auxin khng gy hiu qu th GA li c hiu qu tt nh
trng hp i vi nho, anh o, to,
e) GA vi s ra hoa v s phn ha gii tnh:
Trailakhian xem GA l mt trong hai thnh phn ca hormone
ra hoa (florigen) trong hc thuyt ra hoa ca ng. Khi x l GA cho cy





274

ngy di c th lm cho chng ra hoa trong iu kin ngy ngn v trong
iu kin ngy di s tng hp GA rt kh khn.
- GA cng c th thay th cho x l lnh v lm cho cy hai nm
ra hoa ngay trong nm u.
- GA cng c tc dng trong vic phn ha c quan sinh sn c
bit l s phn ha gii tnh c v ci. Trong trng hp ny th GA c
tc dng i khng vi cytokinin, tc l kch thch s biu hin v hnh
thnh gii tnh c: kch thch s hnh thnh ca hoa c v c ch s
hnh thnh hoa ci. Chnh v vy ngi ta s dng GA iu khin s
lng hoa c ca cc cy h bu b.
2.5. C ch tc ng ca GA
Mt trong nhng tc ng c trng ca GA l lm tng hot ng
thy phn ca ht, c, trong qu trnh ny mm. V d nh trong cc
ht la, ng, la m, phi ht l ni tng hp nn GA. GA c gii
phng ra khi phi v khuych tn vo lp t bo aleuron v ti y, GA
kch thch s hnh thnh v gii phng cc enzyme thy phn trong cc t
bo aleuron. Cc enzyme ny khuych tn vo t bo ni nh v ti y
chng xc tc cho s thy phn cc hp cht hu c phc tp thnh cc
cht n gin cn cho qu trnh ny mm.
GA ng vai tr l nhn t m gen m cc gen ny b c ch hon
ton trong qu trnh ng ngh ca ht, c. Trong qu trnh ng ngh, cc
gen ny b bao vy bi mt loi protein gi l histone.
GA l nhn t hot ha gen, gii phng gen ny khi trng thi c
ch v do cc enzyme thy phn mi c hnh thnh.
S tc ng ca GA ln s ny mm:














DNA mRNA Protein-enzyme

GA


Amylase
Tinh bt n.glucoso-(P)
n H
3
PO
4


H hp

T bo
ht nc Nguyn liu ATP



Sc trng
NY MM





275

* S gin ca t bo cng c th do c ch hot ha bm H
+
ca
GA v gy nn hiu ng mm t bo. S c mt ca GA cng gy nn s
tng cc cht hot ng thm thu v p sut thm thu ca t bo to iu
kin cho t bo ht nc, sc trng tng ln, t bo gin ra.
* Cc tc ng khc ca GA nh s ra hoa, s phn ha gii tnh:
c th c thc hin thng qua c ch hot ha gen ca GA.
2.6. ng dng ca GA
a) Kch thch tng chiu cao, tng sinh khi:
Vi cy ay, ngi ta phun GA (2 5 ppm) s lm chiu cao cy
tng ln nhiu, tng nng sut si m cht lng si m bo.
Vi cc cy rau nh bp ci, xu ho, x lch vic x l GA
(2 5 ppm) lm tng nng sut cht xanh.
b) Tng nng sut v to qu khng ht.
c) Ph b s ng ngh ca ht, c.
X l GA (2 ppm) c th ngm hoc phun s lm cho c khoai ty
ny mm nhanh.
d) iu chnh gii tnh:
X l GA cho bu, b, cc loi da, lm tng t l hoa c.
iu ny c ngha quan trng trong vic sn xut ht lai ca cc cy h
bu b. Ngi ta c th bin mt hng thnh cy mang hoa ci v xen mt
hng l cy mang hoa c. Chng th tinh cho nhau to nn ht lai.
e) ng dng trong sn xut malt bia:
kch thch sn xut amylase trong cng ngh sn xut bia ngi
ta x l GA (2 5ppm) cho ht i mch trong qu trnh ny mm s
tng nng sut malt v phm cht malt bia.

3. Cytokinin
3.1. Lch s pht hin:
T 1913, Haberlandt chng minh rng: Cc m thng tn c
kh nng sinh ra mt cht kch thch sinh trng v khuych tn vo m
khng thng tn kch thch s phn chia ca n.
Nm 1941, Overbeek chng minh l nc da c kh nng
tn ti mt cht kch thch s phn chia v phn ha t bo.
Nm 1955, Miller v Skoog tch c mt hp cht t vic hp
mu DNA ca tinh dch c mi, cht ny c kh nng kch thch s phn
chia t bo rt mnh trong nui cy m thuc l gi l Kinetin.
Cytokinin trong cy c chit tch t 1963(Letham v Muller)
dng kt tinh t ht ng non gi l Zeatin. Sau ngi ta chng minh
rng Zeatin c mt trong tt c th gii thc vt. Trc ngi ta cng





276

pht hin ra ngun cht giu c trong nc da c hot tnh cytokinin
l diphenylurea.
Cng thc cu to tng qut ca cytokinin nh sau:









Nu R l: th l Kinetin.


Nu R l th l Zeatin.













3.2. S trao i cytokinin trong cy:
R l c quan tng hp cytokinin. T r cytokinin c vn
chuyn ln cc b phn trn mt t. Cytokinin c tng hp mnh nht
l giai on cy con. Cytokinin phn b nhiu nht nhng ni c hot
ng trao i cht v sinh trng mnh nht. Do vy cy no sinh trng
mnh, chi non nhiu th hm lng cytokinin cao.
Cytokinin tn ti trong cy di dng t do c hot tnh sinh hc
v dng lin kt khng c hot tnh hoc hot tnh yu. Cytokinin c th
lin kt vi ng to nn glycoside. Ngoi ra cytokinin c mt trong
RNA vn chuyn cho serine, phenylalanine, tryptophan, tyrosine, cysteine.
NH - R

N N



N NH


CH
2


O


CH
2
CH = C CH
2
OH

CH
3


NH


C = O

NH
Diphenylurea






277

Cytokinin b phn gii nh h thng enzyme c hiu v sn phm
phn gii cui cng l ur.
3.3. Hiu qu sinh l ca cytokinin:
* Cytokinin v s gin n ca t bo
* Cytokinin v s phn ha chi.
* Cytokinin v s ha tr: Cytokinin l hormone ha tr. Chng c
tc dng ko di tui th ca l v ca c quan, ca ton cy.
Tc dng c trng nht ca cytokinin l km hm s ha gi ca
l. Khi cc l gi th chng c th rng i. Trong l s phn hy nhanh
protein trong bn l, tch cc cht phiprotein, lipid, nucleic acid qua mng
t bo gn l, cung l v dn theo s ph hy ca lc lp. Do vy vic x
l cytokinin c tc dng km hm s phn hy protein nn km hm c
s phn hy ca dip lc v ko di tui th ca l.
Trong thc t nu b r pht trin mnh, tr lu l v r l c quan
tng hp cytokinin. V vy mun ko di tui th ca c quan v ca ton
cy th cn phi iu khin s sinh trng ca b r tht tt. Ngc li
mun rt ngn thi gian sinh trng, nhanh chng ra hoa kt qu th phi
hn ch sinh trng b r, cht bt r.
* Cytokinin cng c nh hng n s ny mm ca ht. V vy
trong mt s trng hp nht nh, cytokinin c tc dng kch thch s ny
mm, ph b s ng ngh ca ht, c.

3.4. C ch tc ng ca cytokinin:
Cytokinin tc ng n s phn chia t bo, s phn ha c quan, s tr
ha c quan v ca ton cy. C ch tc ng ca cytokinin ln s sinh
trng ca cy c th tm tt nh sau:
- Khi thiu cytokinin th t bo khng phn chia c mc d
DNA vn tip tc c tng hp, nhng m s tng hp protein khng
xy ra. V vy c th cho rng cytokinin kim tra s tng hp protein
giai on gii m (translation):







- C th c c ch th hai l cytokinin c mt trong RNA vn
chuyn nn n nh hng n s tng hp protein. Cytokinin trong RNA
vn chuyn v tr lin kt vi anticodon trn ribosome. V vy chc nng

Transcription Translation
DNA mRNA Protein-Enzyme


Cytokinin
Sinh trng





278

iu chnh ca cytokinin c l bng c ch ngn chn s nhn mt sai lm
ca cc codon trong qu trnh tng hp protein.
- Hiu ng ca cytokinin trong vic ngn chn s ha gi ca cy
lin quan nhiu n kh nng ngn chn s phn hy protein, nucleic acid
v dip lc.
3.5. ng dng ca cytokinin
Cytokinin c ng dng ch yu trong vic nui cy m t bo
thc vt. Cc cht cytokinin nhn to nh benzyladenine (BA), kinetin
cng nh cytokinin t nhin trong nc da c ng dng rng ri
trong nui cy m. Vic c mt ca chng trong mi trng quyt nh s
phn chia v phn ha chi trong m nui cy.












B. CC CHT C CH SINH TRNG THC VT
1. Abcisic acid (ABA)
1.1. Lch s pht hin:
* 1961 - Luc Carn (M) tch c mt cht di dng tinh th
t qu bng gi v khi x l ln l bng non th gy nn hin tng rng
ca l v t tn l abcisin I.
1963 Ohkuma v Eddicott tch c mt cht khc t l bng
tr cng gy nn hin tng rng, h gi l abcisin II.
ng thi Waring v cng s bng phng php sc k tch
c mt cht c ch t l cy Betula v khi x l cho cy gy nn
hin tng ng ngh ca chi. H gi cht l dormin.
Nm 1966 ngi ta xc nh c bn cht ha hc ca cht .
Nm 1967 ngi ta gi cht l abcisic acid (ABA).






NH CH
2


N N

Benzyladenine (BA)

N NH





279









* ABA c tng hp hu ht cc b phn ca cy, tch ly
nhiu nht trong cc c quan sinh sn, c quan d tr, trong cc c quan
gi, c quan ang ng ngh.
- ABA gy nn s rng (l, hoa, qu, ), hot ha s hnh thnh
tng ri ch yu l s hnh thnh cc enzyme thy phn v t bo. ABA
c ch s sinh trng ca chi; ca cy. Gy nn s ng ngh.
- ABA l cht i khng ca auxin.
- ABA l hormone ha gi.
- ABA iu chnh s ng m kh khng: Vic pht hin ra vai tr
ca ABA trong vic iu chnh ny xut pht t vic pht hin ra mt t
bin ho c chua. Cy c chua lun lun b ho ny v kh khng ca n
lun lun m m khng ng c. Phn tch hm lng ca cc cht
trong t bin ny th thy thiu ABA. Khi x l ABA cho n th kh
khng ng li. t bin ny l do thiu gen tng hp ABA nn ABA
khng hnh thnh c. Vai tr ca ABA l c ch s hp thu K
+
bi t
bo kh khng. Mt dng vt cht khc c lin quan n s ng m ca
kh khng di tc ng ca ABA l malic acid. Acid ny c gii
phng cng vi K
+
khi c mt ca ABA v to nn mt s thiu ht K
+

t bo kh khng, thiu sc trng, t bo mt nc v kh khng ng li.
- ABA c th xem l hormone stress: Cc iu kin bt thun
ca mi trng nh hn, ng, nng, lnh, mn, su bnh, gi l iu
kin stress ca mi trng. Khi gp mt tc nhn stress bt c no th hm
lng ABA lp tc tng ln rt mnh. l phn ng thch nghi ca cy
chng li cc iu kin bt thun ca mi trng. Khi gp hn, hm lng
ABA trong l tng ln mnh (c l do s tng hp mi) v do kh
khng ng li, chng c s thot hi nc.
Ngi ta xc nhn rng ABA c tng hp trong lc lp v c
gi trong lc lp khi t bo nc. Khi t bo mt nc th tnh thm
ca mng lc lp tng ln v ABA c thm ra ngoi t bo cht, vn
chuyn vo t bo kh khng gy nn s ng ca kh khng. Khi ht
stress th tnh thm ca mng lc lp vi ABA gim xung v do ABA
CH
3
CH
3


CH = CH C = CH COOH
OH
CH
3

CH
3

O Abcisic acid (ABA)






280

c d tr trong lc lp. V vy, ABA c vai tr nht nh trong tnh
chng chu ca cy i vi cc iu kin bt thun ca mi trng.
1.2. C ch tc ng ca ABA
a) Tc ng ca ABA ln tnh thm ca mng:
ABA lm thay i in th qua mng v do m iu tit c
s tit ion K
+
qua mng. C ch ny lin quan n s ng m ca kh
khng.
C ch v vai tr ca ABA ln s ng m kh khng nh sau: Khi
thiu ABA, bm H
+
khng b c ch, malate v c K
+
vn chuyn vo
khng bo v do t bo ht nc kh khng m.
Khi x l ABA, bm H
+
b c ch nn H
+
khng vo thnh t bo;
malate v K
+
khng qua mng vo khng bo m vn chuyn th ng ra
khi t bo, lc t bo mt nc v kh khng ng li.













b) ABA tc ng ln c ch tng hp protein v nucleic acid:
Nm 1964, Dorothy v Bonner cho rng trong cc t bo ang ng
ngh th vt liu di truyn (DNA) hon ton b trn p v qu trnh tng
hp protein khng din ra; qu trnh sinh trng b ngng tr.
Nu x l cc cht i khng ca ABA nh GA chng hn lm
gim v c ch tc dng ca ABA ln h thng v do vy m qu trnh
sinh trng din ra.
Hiu qu c ch ca ABA l hiu qu ng gen (kha gen) do
qu trnh sao chp (Transcription) v gii m (Translation) khng din ra
c. Gibberellin (GA) s lm mt tc dng c ch gen ca ABA, lm cho
cc qu trnh din ra bnh thng. C l ABA tc ng ln giai on
transcription:



Transcription Translation
DNA mRNA Protein
(Enzyme, cu trc)
Sinh
trng
Thnh t bo Cht nguyn sinh Khng bo

Glucose

ABA c ch bm H
+
,
c ch ATP-ase CO
2
PEP
c ch thm K
+
v malate
vo khng bo
H
+
Malate
K
+
K
+
K
+






281

1.3. ng dng nhng hiu bit v ABA trong trng trt:
Trong cy ABA thng gy hiu qu c ch ln qu trnh sinh
trng pht trin khc nhau. loi b tc dng c ch ca ABA i vi
cy trng chng ta thng s dng cc tc nhn i khng vi ABA
lm thay i s cn bng hormone trong cy theo hng lm mt tc dng
ca ABA:
- ph b ng ngh, kch thch s ny mm th hoc l lm gim
lng ABA hoc lm tng gibberellin (GA) hay c hai.
- Km hm s rng ca l, qu bng cch tng hm lng auxin,
gim hm lng ABA.
- iu chnh s ha gi: T l quy nh gi
ha ca cy.

Mun chng s gi ha c th tin hnh bng cch phun cytokinin
ngoi sinh, cho cytokinin vo mi trng nui cy hoc bng cch tng
cng s pht trin ca b r tng hp nhiu cytokinin cung cp cho
cc c quan trn mt t ko di tui th ca chng.

2. Ethylene (CH
2
= CH
2
)
L sn phm ca s trao i cht, n l mt cht kh n gin.
Ethylene kch thch s chn ca qu. Ethylene cng tng tc vi ABA
gy nn s rng ca l, qu. Ethylene kch thch s ra hoa ca mt s thc
vt nh kch thch cho da ra hoa tri v. Trong ngh trng da vic s
dng cht to ra ethylene ( l ethrene) hoc t n (cho acetylene)
lm cho da ra thm mt v qu.
Ethylene l mt cht kh nn trong thc t khng s dng trc tip
ethylene m ngi ta s dng cc cht to ra ethylene nh CEPA:








- Trong vic phn ha gii tnh ca cc cy thuc h bu b th
ethylene c nh hng ln s hnh thnh gii tnh ci. X l ethrene s
lm cho chng pht trin ch yu hoa ci.


Cytokinin
ABA

O
H
2
O HCl H
3
PO
4

Cl(CH
2
)
2
P OH CH
2
= CH
2


OH Ethylene
CEPA (Chloroethylenephosphoric acid)





282

3. Cc cht phenol
L sn phm ca s trao i cht trong cy. Chng c tc dng c
ch hot ng trao i cht v c ch sinh trng. Cc phenol kch thch
hot ng ca enzyme phn hy auxin; km hm s gin ca t bo v lm
cho t bo ha g nhanh.
Mt s phenol thng gp nh:











4. Cc cht lm chm sinh trng (Retardant)
Retardant l mt nhm cht nhn to, c bn cht ha hc khc
nhau, chng gy nh hng c ch gin ca t bo, lm cy thp, m rng
phin l, gy s ra hoa sm.
Mt s v d in hnh:
4.1. CCC (Chlorocholine chlorid):
[(CH
3
)
3
N CH
2
CH
2
Cl]
+
Cl
-

- CCC l cht khng GA, n lm cho cy thp i. Lm tng s hnh
thnh dip lc. Khi x l CCC ln cy, tc thm vo cy rt nhanh, tn
ti vi tun ri b phn hy, mt hot tnh.
4.2. MH (Malein hydrazide)
MH l cht khng auxin v n lm tng hot tnh enzyme phn hy
auxin, l IAA-oxidase









O

NH


NH

O
O






O
Urgon



O
O

Cumarin



COOH

HO


Oxybenzoic acid





283

MH l cht c ch sinh trng, n c ch s ny mm v ko di
thi gian ng ngh; c ch s sinh trng chi bn ca cy nn c s
dng ngn chn s mc chi nch ca thuc l, trnh c vic nh
ta nhnh bng tay rt tn cng lao ng.

4.3. TIBA (Triiodobenzoic acid)






TIBA cng l cht khng auxin, n km hm s vn chuyn auxin
trong cy; lm gim s sinh trng ca chi ngn, hn ch chiu cao, xc
tin s phn cnh; TIBA cn xc tin s ra hoa v s hnh thnh c.

Cu hi:
1. Cc nhm hp cht th cp quan trng trong thc vt?
2. C ch tc ng ca cc nhm cht iu ha sinh sinh trng thc
vt v ng dng chng trong sn xut nng nghip?




















I
COOH


I I
M UC LUC

Chng 1 TRAO I CHT VA NNG LNG SINH HOC, 1
1. Khai nim chung v trao i cht,1
2. Nng lng sinh hoc, 1
2.1. c tnh nng lng cua s trao i cht,
11
2.2. Cac hp cht cao nng, 13

Chng 2: VITAMIN, 18
1. nh ngha vitamin,18
2. Cac vitamin hoa tan trong nc (cu tao, vai
tro), 18
3. Cac vitamin hoa tan trong cht beo (cu tao,
vai tro), 29

Chng 3 ENZYME VA S XUC TAC SINH HOC, 36
1. Khai nim chung v s xuc tac, 36
2. Enzyme la cht xuc tac sinh hoc, 37
3. Ban cht hoa hoc cua enzyme, 38
4. Cac yu t anh hng i vi hoat tnh xuc
tac cua enzyme, 42
5. C ch tac ng cua enzyme, 43
6. ng hoc cua enzyme, 44
7. Cach goi tn va phn loai enzyme
8. Cac nhom enzyme ring bit, 55

Chng 4 CARBOHYDRATE VA S TRAO I CARBOHYDRATE
TRONG C TH THC VT, 63
1. Cu tao, tnh cht va vai tro cua
carbohydrate, 63
2. Cac monose, 63
3. Cac polyose, 69
4. Hoa sinh quang hp, 73
Pha sang trong quang hp, 73
Pha ti trong quang hp (chu trnh Calvin, 81
H hp sang, 87
Chu trnh Hatch va Slack, 91
Chu trnh CAM, 98
Tng hp sucrose, 99
5. Hoa sinh h hp,103
5.1 nh ngha h hp, 103
5.2. Qua trnh ng phn, 109
5.3. Cac loai h hp, 113
5.4. H hp hao kh chu trnh Krebs, 117
5.5. Chu trnh pentosephosphate, 123

Chng 5 LIPID VA S TRAO I LIPID TRONG C TH THC
VT,128
1. Cu tao va tnh cht, 129
2. S phn giai triglycerid, 130
3. S phn giai cac acid beo cha bao hoa,
136
4. S phn giai cac acid beo co s nguyn
t carbon le, 136
5. Chu trnh glyoxilate, 138
6. Sinh tng hp triglycerid, 139
7. Sinh tng hp va phn giai phospholipid,
144

Chng 6 NUCLEIC ACID VA S TRAO I NUCLEIC ACID
TRONG C TH THC VT, 147
1. Cu truc cua nucleic acid, 147
2. Phn loai nucleic acid, 154
3 Sinh tng hp nucleic acid, 155

Chng 7 PROTEIN VA TRAO I PROTEIN TRONG C TH THC
VT
1. ai cng v protein
2. Vai tro cua protein
3. Cu truc cua protein
4. Aminoacid
Phn loa aminoacid
Tnh cht cua aminoacid
5 Sinh tng hp aminoacid
6 Qua trnh kh nitrate
7 Qua trnh c nh nit phn t
8 Cac ng hng bin i NH
3
- chu trnh
ornithine
9 Cac ng hng tng quat cua s bin
i aminoacid
10 Sinh tng hp protein
10.1 Cac yu t tham gia sinh tng hp protein
10.2 Cac giai oan sinh tng hp protein
10.3 C ch iu hoa sinh tng hp protein
11. Phn giai protein

Chng 8: MI LIN QUAN GIA CC CHT TRONG TRAO I
CHT
9.1 Mi lin quan gia carbohydrate va lipid
9.2 Mi lin quan gia carbohydrate va protein
9.3 Mi lin quan gia lipid va protein

Chng 9 CAC HP CHT CO NGUN GC TH CP
1. Khai nim chung v cac hp cht co ngun
gc th cp
2. Acid hu c
3. Isoprenoid va steroid
4. Cao su
5. Flavon va anthocyan
6. Cac dn sut cua benzen va cac hp cht co
vong ngng tu
7. Cac hp cht co ngun gc th cp cha nit
8. Cac cht kch thch v c ch sinh trng cua
thc vt










258


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