QU HAI (CH BIN) - NGUYN B LC

TRN THANH PHONG - CAO NG NGUYN

GIO TRNH

HA SINH

NH XUT BN I HC HU

Li ni u
Ha sinh hc l khoa hc nghin cu thnh phn ha hc ca c th
sng v nhng qu trnh chuyn ha ha hc ca cc cht v nng lng
trong qu trnh hot ng sng xy ra trong c th sinh vt. L ngnh khoa
hc c bn, ng thi cng l mt khoa hc ng dng, l ngnh khoa hc
c s cho cc khoa hc c bn quan trng khc nh cng ngh sinh hc,
sinh hc phn t; ha sinh hc tc ng ln hay ni rng hn l mi nhn
nh hng v gii quyt cc vn phc v cho nng, lm, ng
nghip, phc v cho y hc... Chnh v vy, ha sinh hc khng ch l cng
c ca sinh hc m cn l cng c ca cc chuyn ngnh khc lin quan
n sinh hc nh nng, lm, ng, y dc.
Gio trnh nhm cung cp cho sinh vin cc ngnh, chuyn ngnh
lin quan n sinh hc trong i hc Hu nhng kin thc c bn v cu
to v thnh phn ha hc ca cc sinh cht, mi lin quan gia cu trc
v chc nng, cc qu trnh chuyn ha ca chng v nng lng trong c
th sinh vt.
Cun sch c bin son theo chng trnh gio dc i hc
c i hc Hu ph duyt, bi tp th tc gi ca cc trng i hc
khoa hc, s phm thuc i hc Hu. Sch cng c th dng ti liu
tham kho cho sinh vin cc trng khc, nhng ngi chun b thi tuyn
sau i hc cng nh cc cn b nghin cu lin quan.
Cc tc gi xin cm n nhng ng nghip gp nhiu kin b
ch trong qu trnh bin son. c bit cc tc gi xin chn thnh cm n
GS.TSKH L Don Din - Gim c Trung tm T vn u t Nghin
cu pht trin Nng thn Vit Nam (INCEDA), Ch tch Hi Ha sinh
Vit Nam nhn phn bin v cho rt nhiu nhng li khuyn qu bu
nhm hon thin gio trnh.
Vi thi gian bin son v kinh nghim cn hn ch, cun sch cn
cha tht y v chc chn khng trnh khi nhiu thiu st. Chng ti
rt mong nhn c nhng kin ng gp ca cc bn ng nghip, sinh
vin v bn c ln xut bn sau s c hon thin hn.
Thay mt cc tc gi
Ch bin

PGS.TS Qu Hai

11

M u
1. Lch s, i tng, nhim v ca ho sinh
th k XIX, khi m ho hc pht trin nh v bo, th ranh
gii gia sinh hc v ho hc xut hin mt lnh vc khoa hc mi
nhm nghin cu thnh phn ho hc ca c th sng v nhng qu trnh
chuyn ho ho hc ca cc cht v ca nng lng trong qu trnh hot
ng sng xy ra trong c th ca chng. Lnh vc khoa hc ny c gi
l ho hc sinh vt hoc vn tt hn- ho sinh hc (biochemistry).
C th ni rng, ho sinh hc l mt phn lnh vc ca khoa hc
cuc sng. Nhim v ca chng nhm nghin cu cc hin tng sng
bng cc phng php ho hc.
y l mt khoa hc tr tui ca th k XX ang trn pht trin
mnh m v ang xm nhp vo nhiu lnh vc mi nhn ca sinh vt hc,
y hc v nng hc... Ho sinh hc mi tr thnh mt khoa hc c lp vo
na sau th k th XIX mc du ngay t thi thng c con ngi lm
quen vi nhiu qu trnh ho sinh hc trong cuc sng hng ngy ca h
(nu ru, nng bnh m, thuc da, lm dm, tng, nc mm...).
Tuy vy ch mi gn y, tt c mi qu trnh sinh hc ny mi
c nghin cu mt cch khoa hc v c gii thch mt cch y .
C th ni, s xut hin mn ho sinh hc l kt qu tt yu ca s
pht trin v phi hp gia ho hc v sinh vt hc.
Tnh cht v phng hng ca ho sinh hc l nghin cu trn c
th sng, tm ngha chc phn ca tt c mi thnh phn, mi sn phm
chuyn ho, trn c s , tm hiu su v:
- Mi lin quan gia qu trnh ho hc v sinh vt hc.
- Mi lin quan gia cu trc v chc nng sng ca cc c quan
trong c th.
- C ch iu ho ton b qu trnh sng.
Tu theo i tng nghin cu m ngi ta phn chia ho sinh hc
thnh ho sinh ng vt, ho sinh thc vt, ho sinh vi sinh vt v y ho
sinh.
Trn mi i tng, ho sinh nghin cu song song hai mt "tnh"
v "ng".
Vic nghin cu cc cht c trong thnh phn ca c th sinh vt
l nhim v ca tnh ho sinh. Tnh ho sinh gn lin rt mt thit vi ho
hc sinh hu c. y chnh l ha sinh m t.
Cn vic nghin cu cc chuyn ha ho hc xy ra trong qu
trnh hot ng sng ca c th ngha l nghin cu v mt ho hc ca s

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trao i cht trong mi lin quan vi ton b c th v mi trng xung
quanh l nhim v ca ng ho sinh.
Tnh ho sinh v ng ho sinh lin quan vi nhau rt cht ch vic nghin cu cc qu trnh ha sinh hc s v ngha nu khng c s
nghin cu cc cht tham gia trong cc qu trnh ny.
Ho sinh m t gn lin vi s pht trin ca ho hu c. Cui th
k XVIII, u th k XIX, hng lot cc hp cht hu c c tch ra
t thc vt v t cc t chc ng vt: citric acid, malic acid, tatric acid,
oxalic acid, urea v cc alkaloid. Ngi ta xc nhn rng trong thnh
phn ca tt c cc cht bo u cha glycerin. Trong thi gian ny,
Lavoisier cng gii thch c c ch ho hc ca s h hp v s
chy. ng chng minh rng, lng nhit do cc c th sng sn sinh ra
cng bng lng nhit thu c khi t chy cc cht dinh dng bn
ngoi c th (khi h hp trong c th, carbon v hydrogen b oxy ha t
t, qu trnh ny rt ging s chy bnh thng).
Tip theo cc cng trnh chit xut, tinh ch v phn tch cc cht
hu c l nhng cng trnh nghin cu tng hp cc cht hu c. Quan
im cho rng, vt cht sng khc vi vt cht khng sng b nh
hon ton khi m Whler vo nm 1828 iu ch c carbamid (urea)
bng phng php nhn to t cc cht v c. Pht minh ca Whler l
bng chng cho rng to nn cht sng khng cn thit bt k mt
"lc" sng no (vis vitalis) v nh vy m ng hng cho ho
sinh pht trin. (Thi k ny khoa hc i su vo s sng ang b khng
ch v km hm bi thuyt "hot lc" cho rng cc cht hu c tham gia
vo cu to ca c th sinh vt ch c th to ra bi mt "ng siu t
nhin").
Trong sut khong hai th k trc, s pht trin ca ho sinh hc
xy ra tng i chm. S bt u tht s ca ho sinh hc thng c
coi l vo nm 1866 khi Tbigen thnh lp b mn ho sinh u tin di
s lnh o ca Hoppe - Seyler ( nc c). S u tin ca tp ch
mang tnh cht ho sinh hc c ra mt vo nm 1877 (Hoppe - Seyler's
Zeitschrift fr physiologische Chemie). Tin ca n l s hot ng ca
Liebig c v trng phi ho hc hu c ca ng. Bng cch s dng
cc phng php nghin cu mi, Liebig xc nh thnh phn ca
nhiu thc phm, chia cc cht cha trong thc phm thnh protein,
glucid, lipid v xc nh hm lng nitrogen trong protein. Sau nhng
cng trnh nghin cu ca Pasteur v s ln men, cc nh khoa hc ch
nhiu n bn cht enzyme vn xc tc cho cc qu trnh khc nhau
trong c th sng. Nhiu cng trnh c gi tr khc, trong c cng trnh
ca Fischer i vo cu trc v tng hp glucid, lipid, amino acid v

13
protein, Pavlov trong thi gian ny nghin cu cc c ch enzyme v
Miescher th nghin cu nucleic acid. Thi gian ny ngi ta cng pht
hin ra vitamin.
Vic xc lp nn thnh phn ho hc ca thc vt, vic pht hin
ra cc enzyme v vic lm sng t vai tr ca chng trong s trao i cht,
s pht hin ra vitamin v hormon, s pht trin ca ho hc v amino
acid v protein, v glucid, lipid to iu kin cho vic hnh thnh ng
ho sinh v chnh nh s pht trin ca ng ho sinh m ngi ta xy
dng c nhng quan im thng nht v cc quy lut chung ca cc qu
trnh trao i cht v ca nhng chuyn ho nng lng trong c th sinh vt.
Ngy nay chng ta bit r rng, tt c cc cht xy dng nn t bo
sng, u thay i khng ngng. c trng ca s sng l s bin i ho hc.
Nh vy ho sinh hc hin i trc ht l ng ho sinh. Trc
tin l nhng hin tng trao i cht, l s chuyn ho v phn gii
cc cht dinh dng nhm thu nhn nng lng ho hc cng nh xy
dng vt cht ca t bo. Cc phn ng ho hc ny c xy ra nh tc
dng xc tc ca nhng enzyme, v vy vic nghin cu cc enzyme
chim mt phn quan trng trong ho sinh hc.
Th n thuc v ng ho sinh l iu ho ho hc. iu ny
c xy ra trn con ng cc sn phm trao i cht xc nh, thng l
cng vi s tham gia ca nhng cht c bit l hormon c to thnh
trong cc tuyn ni tit.
Th ba thuc v ng ho sinh l cc qu trnh ho hc c xy
ra gn lin vi cc cu trc v thc hin cc chc nng ca cc phn t
cu trc.
Vic xc nh trnh t ca amino acid trong protein v cu trc
khng gian ca protein (Sanger, Perutz v Kendrew) cng nh cu trc
ca nucleic acid (Chargaff, Watson v Crick) l bt u mt giai on mi
trong s pht trin ca ho sinh hc tc l thi k ca sinh hc phn t.
Mt iu r rng l khng th tch ring sinh hc phn t v ho sinh hc
ra khi nhau. V c bn m ni th "sinh hc phn t" ch l tn mi,
nhng l mt phn ngnh ring ca ho sinh. N c gng lm sng t cc
hin tng sng c bn trn c s phn t c ngha l trn c s ho sinh
hc. Chnh v vy theo quan nim hin i th ho sinh hc l khoa hc
nghin cu c s phn t ca s sng.
2. Thnh phn ho hc ca c th sng
Trong c th sng, nc l thnh phn quan trng nht. Tr mt s
m hoc loi t bo (cc ht thc vt, cc bo t m xng, m m),
lng nc ca chng khng t n 80%, cn lng nc ca mt s
sinh vt khc cng vt qu 90% (ton b lng nc ca c th con

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ngi khong 50-70%). iu cn lu l mt s dng sng bc thp, cc
virus, cc bo t sng qua trng thi kh ho hon ton, song trong trng
thi khng c nc th cc hot ng sng ca chng b hon li.
T cc c s trn chng ta c th ni rng cc qu trnh ho hc
c trng cho s sng c xem nh l nhng phn ng tin hnh trong
mi trng nc.
Ngoi ra ngi ta pht hin c hn 60 nguyn t c trong c
th sng. Cc nguyn t ny c trong c th vi nhng lng rt khc
nhau. Mt s c coi l nhng nguyn t cn thit xy dng c th v
phc v cho s pht trin bnh thng ca c th; mt s khc th chc
nng sinh hc ca chng cha c bit r; s cn li c coi nh do s
xm nhp ngu nhin. C H O N S P Cl Ca Mg K Na u l nhng nguyn
t rt cn thit i vi c th sng, 11 nguyn t ny chim gn 100%
khi lng ton phn ca thc vt v ng vt. Nhng nguyn t dng
vt c gi l yu t vi lng, vai tr quan trng ca chng i vi cuc
sng cng dn dn c sng t l nhng ion kim loi nng Co, Zn,
Mn, Mo; trong trng hp cc c th ng vt bc cao cn c I, cc cy
xanh th c B. Cn nh rng, tr I v Mo, cc nguyn t c k n
u nm trong s 30 nguyn t u ca h thng tun hon v hn mt
na cc nguyn t c s th t n 30 c vai tr sinh hc quan trng.
Nhiu hp cht hu c trc y c coi nh l ring bit thuc
gii sinh vt, nay cng c nghin cu, tng hp trong phng th
nghim. l cc protein, nucleic acid, glucid, lipid, enzyme v vitamin.
y l nhng cht chim v tr hng u trong sinh hc v cng chnh l
i tng nghin cu chnh ca cc chng trong gio trnh ny.
3. Mi quan h ca ho sinh vi cc ngnh khoa hc ca sinh
hc, nng nghip v y hc
Trong sinh hc c nhiu ngnh, nhng nghin cu sinh hc ngy
nay l nghin cu mc phn t. Ho sinh l khoa hc nghin cu s
sng mc phn t, cho nn c th ni bt c chuyn ngnh no ca
sinh hc nh ng vt hc, thc vt hc, vi khun hc, sinh l hc, t bo
hc, m phi hc... u cn phi trang b kin thc v k thut ho sinh
nghin cu khoa hc chuyn ngnh mnh. Do khi ni n cc chuyn
ngnh ca sinh hc hin i th trc ht phi ni n ho sinh trong
nhng nm gn y. Ngay c cng ngh gen, cng ngh enzyme cng
chnh l lnh a ca ho sinh. Chng hn i vi ng vt, thc vt, vi
sinh vt, ngy nay mun nghin cu phn loi chnh xc cc ging chng
cng phi dng cc ch tiu phn t mt s protein, enzyme hay nucleic
acid trong ty lp th. i vi nng nghip mun tng nng sut cy trng
phi ch n qu trnh ho sinh quang hp, qu trnh ho sinh ny mm,

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qu trnh ho sinh pht dc, qu trnh chuyn ho, sinh tng hp cc cht
to nn ht, qu, qu trnh tc ng ho sinh ca cc cytokinin n s sinh
trng v pht trin ca cy. i vi vt nui cng vy, mun c nng
sut tht, sa, trng cao, ngi chn nui phi hiu c cc qu trnh ho
sinh pht trin n tng giai on pht trin ca con vt, n tng b phn
c bp, bung trng ca chng c s tc ng mnh m. i vi y
dc hc, vn ch cht nghin cu bnh nguyn, bnh l, chn on v
iu tr bnh cng u lin quan cht ch n ho sinh, tc lin quan n
s thay i cc phn t bnh l xy ra trong c th v tm nhng cht hot
tnh sinh hc c tc dng phng chng hoc cha khi bnh.
V vy c th ni ho sinh l gc, l c bn gip hiu su sc
cc khoa hc khc ca sinh hc, nng nghip v y hc.
4. Mt s thnh tu ni bt ca ha sinh trong thi gian gn y
Trong qu trnh pht trin ca mnh, nhiu ngnh nh ca ho sinh
ra i.V ho sinh mt s chc phn h thng quan trng c ho sinh
min dch, ho sinh di truyn, c bit mt ngnh mi gn y xut
hin l cng ngh ho sinh. Cc lnh vc nh ca ho sinh ng gp
mt cch tch cc vo thnh tch chung ca ho sinh. Nhiu gii thng
Nobel ghi cng cc kt qu nghin cu quan trng, m ra nhiu cnh
ca mi cho s pht trin ca ho sinh nh ho sinh ca h thng min
dch ca Snell, Bena Cerraf v Dausset nm 1980. Cng nm y Paul Berg
cng c gii thng Nobel bi cng trnh nghin cu gn cc mu
DNA. Nm 1981-1982, thnh tu tng hp gen - interferon gm 514 i
base bi Leicester c thc hin. T n nay hng lot cng trnh
khc v nghin cu ho sinh c p dng trong lnh vc khoa hc.
Gn y, nm 1997 gii thng Nobel y hc trao cho Staley
Prusiner v cng trnh nghin cu prion, mt khi nim mi v "nhim
khun", gy bnh no th xp ngi v ng vt. Prion (PrP) l protein
tn ti hai dng ng phn alpha v bta. c th kho mnh th PrP c
dng alpha cn khi c th b bnh th dng alpha b dui ra v xp thnh
cc bng song song gi l PrP bta. Dng ny rt bn vi enzyme tiu ho
v khng b ph hu nhit cao (n 200oC). Do vy prion nh l tc
nhn gy bnh hon ton mi c b sung vo danh sch nhng tc nhn
gy bnh nh virus, vi khun, nm. Cng trnh ny a ra khi nim bnh
l phn t hon ton mi trong sinh hc v y hc. Cng trnh khng ch
pht hin ra tc nhn gy bnh xp no m cn t nn mng cho s tm
hiu c ch mt tr lin quan n bnh gi v bnh Alzheimer, cng nh
t nguyn tc cht ch cho vic ghp cc c quan ph tng ca ng vt
cho con ngi v thuc men ch t ng vt dng cho ngi.

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TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n (ch bin). 1979. Ha sinh i cng (tp I, II). NxB KH&KT.
H Ni.
2. i Duy Ban. 2005. Ha sinh hc v ha sinh y hc. Ha sinh hc. 1: 8-13.
3. Phm Th Trn Chu, Trn Th ng. 2000. Ha sinh hc. Nxb Gio dc. H
Ni.
4. L Don Din. 1975. Ha sinh thc vt. Nxb Nng nghip. H Ni
5. Nguyn Tin Thng, Nguyn nh Huyn. 1998. Gio trnh sinh ha hin
i. Nxb Gio dc. H Ni.
6. Nguyn Xun Thng, o Kim Chi, Phm Quang Tng, Nguyn Vn
ng, 2004. Ha sinh hc. Nxb Y hc, H Ni.

Ti liu ting nc ngoi

1. Karlson. P., 1972. Biokmia. Medicina Knyv Kiad Budapest.
2. Lehninger A.L., 2004. Principle of Biochemistry, 4th Edition. W.H Freeman.
3. Stryer L., 1981. Biochemistry. W.H.Freeman and company. San Francisco.
4. Straub .F. B. 1965. Biokmia. Medicina Knyv Kiad Budapest.

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Chng 1

Saccharide
L hp cht hu c c to nn t cc nguyn t: C, H, O c
cng thc cu to chung Cm(H2O)n, thng m = n. Do c cng thc cu
to nh trn nn saccharide thng c gi l carbohydrate - c ngha l
carbon ngm nc.
Tuy nhin c nhng saccharide c cng thc cu to khng ng
vi cng thc chung ni trn v d: deoxyribose (C5H10O4).
C nhng cht khng phi l saccharide nhng c cng thc cu
to ph hp vi cng thc chung trn v d : acetic acid (CH3COOH).
Saccharide l thnh phn quan trng trong mi sinh vt .
thc vt, saccharide chim t 80 - 90% trng lng kh,
saccharide tham gia vo thnh phn cc m nng , v d cellulose, hay
tch tr di dng thc phm d tr vi lng ln, v d tinh bt. ng vt,
hm lng saccharide thp hn nhiu, thng khng qu 2%, v d glycogen.

1.1. Monosaccharide
1.1.1. Cu to v danh php
L cht c cha nhiu nhm ru v mt nhm kh oxy (nhm kh l
nhm carbonyl l aldehyde hay ketone).
- Nhm kh l aldehyde ta c ng aldose v c cng thc tng qut:
CHO
(CHOH)n
CH2OH
- Nhm kh l ketone ta c ng ketose c cng thc tng qut:
CH2OH
C= O
(CHOH)n
CH2OH

18
CHO - CH2OH c xem nh l monosacchariden gin nht.
Trong thin nhin monosaccharide c cha t 2 n 7 carbon v c
gi tn theo s carbon (theo ting Hy Lp) + ose
V d: monosaccharide c 3C gi l triose. Tng t ta c tetrose,
pentose, hexose, heptose.
1.1.2. ng phn quang hc
Quy c Fischer: Fischer l ngi u tin nu ra nguyn tc biu
din cc monosaccharide bng cng thc hnh chiu ca chng. Theo :
hnh chiu ca cc nguyn t carbon bt i (C*) v cc nguyn t C khc
nm trn mt ng thng, nguyn t C c s th t nh nht c hnh chiu
nm trn cng. Cn cc nhm th c hnh chiu bn phi hay bn tri.
V d : glyceraldehyde.
V glyceraldehyde c 1 C* nn theo quy tc ca Vant Hoff c 2
ng phn (N = 2n)
1CHO
1CHO
HO- 2C* -H

H-2C*-OH

3CH2OH

D: -OH bn phi
L: -OH bn tri

3CH2OH

L glyceraldehyde D glyceraldehyde.
Khi phn t monosaccharide c nhiu C* th cng thc c dng D
hay L c cn c vo v tr nhm OH ca C* xa nhm carbonyl nht.
V d :
CHO
CHO
H-C-OH
HO-C-H
H-C-OH
H-C-OH
CH2OH
D glucose

H-C-OH
HO-C-H
H-C-OH
HO-C-H
CH2OH
L glucose

19
Ch : monosaccharide t triose tr ln u c C* tr dihydoxy aceton
CH2OH
C=O
CH2OH
1.1.3. Cng thc vng ca monosaccharide
Cng thc thng theo Fischer nh trnh by trn khng ph hp
vi mt s tnh cht ho hc ca chng nh: mt s phn ng ho hc
thng xy ra vi aldehyde khng xy ra i vi monosaccharide . V vy
c th ngh rng nhm -CHO trong monosaccharide cn tn ti di dng
cu to ring bit no .
Mt khc: monosaccharide c th to ether vi methanol to thnh
mt hn hp 2 ng phn c cng nhm methoxy (- OCH3). iu
chng t trong monosaccharide cn tn ti mt nhm -OH c bit.
Qua nghin cu Kolle cho thy: s ng phn thu c ca
monosaccharide thc t nhiu hn s ng phn tnh theo cng thc N=2n, do
gii thch cc hin tng trn, Kolle cho rng ngai dng thng
monosaccharide cn tn ti dng vng.
S to thnh dng vng xy ra do tc dng ca nhm -OH cng
phn t monosaccharide to thnh dng hemiacetal hay hemiketal.

20
V d : cu to vng ca glucose xy ra nh sau:

Do s to thnh hemiacetal vng m C1 tr nn C*, nhm -OH mi

c to ra C1 l -OH glucoside. Tng t vi ketose th C2 tr nn C*,
nhm -OH mi c to ra C2 l -OH glucoside khi to thnh
hemiketal.
Cch biu din cng thc vng nh trn da vo nguyn tc ca
Haworth: C v cu ni vi oxy nm trn mt mt phng , cc nhm th
cng thc thng nm bn phi th cng thc vng nm di mt phng v
ngc li. Ring cc nhm th ca C c nhm OH dng to cu ni oxy th
theo quy tc ngc li.
1.1.4. Hin tng h bin ca monosaccharide
Nh ta thy, khng th gii thch c tt c cc tnh cht ca
monosaccharide nu ta ch tha nhn mt dng cu to no ca
monosaccharide. Nn ngi ta cho rng cc dng cu to c th
chuyn ho ln nhau.
pyranose
pyranose
Dng thng
Furanose
Furanose

21
1.1.5. Tnh cht ca monosaccharide
1.1.5.1. L tnh
Cc monosaccharide tan trong nc, khng tan trong dung mi
hu c, c tnh quay cc tr biose v khng c C*.
1.1.5.2. Ho tnh
a. Monose l tc nhn kh
Trong mi trng kim, kh cc ion kim loi nng c ho tr cao
thnh ion c ha tr thp hay cc ion kim loi thnh kim loi.
Tnh kh ny do nhm aldehyde hay nhm ketone to ra v cc
monose bin thnh acid.
V d: Cu2+ b bin i thnh Cu+ trong phn ng vi thuc th
Fehling, Ag+ b bin i thnh Ag trong phn ng trng gng.
b. Phn ng vi cc cht oxy ho
Tu thuc vo cht oxy ho:
- Cht oxy ho nh nh nc brom ng aldose s thnh aldonic
acid, vi ketose phn ng khng xy ra.
- Cht oxy ho mnh nh HNO3 m c c s oxy ho xy ra 2
u cho ta di acid.
- Trng hp c bit nu ta bo v nhm -OH glucoside bng
cch methyl ha hay acetyl ho trc khi oxy ho bng nc brom, sn
phm to thnh l uronic acid.
c. Phn ng vi cht kh
D dng vng chim t l rt ln trong thnh phn, dng thng
chim t l nh nhng cho ta thy r tnh cht ca mt carbonyl tht
s. Khi b kh: monose s bin thnh polyalcohol.
d. Phn ng to furfural
Di tc dng ca acid m c, cc aldopentose to thnh furfural
v aldohexose bin thnh hydroxymethylfurfural. Cc sn phm ny khi
cho tc dng vi cc phenol cho mu c trng nh: naphthol cho vng
mu tm (Molisch). y l phn ng phn bit ng vi cc cht
khc. Nu ng 5C s cho mu xanh cm thch vi orcinol (Bial).
e. Phn ng ester ho
Cc gc ru ca monose c kh nng kt hp vi acid to thnh ester.
Cc ester phosphate thng gp l: Glucose-6-phosphate, fructose6-phosphate...

22
1.1.6. Cc monose quan trng
1.1.6.1. Pentose

1.1.6.2. Hexose
Cc hexose quan trng nh:
* Glucose: cn gi l dextrose v lm quay mt phng nh sng
phn cc v pha phi.
Ph bin rng ri trong thc vt nht l trong qu nho, nn cn gi l
ng nho, trong mu ngi c 0.8 - 1,1 g/l, nhng ngi b bnh i ng
c th n 2g/l. Cc disaccharide quan trng l saccharose, lactose, maltose v
cc polysaccharide quan trng l tinh bt, glycogen. Ngi ta s dng glucose
trong y hc nh cht tng lc.
* D - Mannose: t gp trng thi t do, thng gp trong
polysaccharride v glucoside
* D - Galactose: l thnh phn ca lactose c trong sa cn gi l
ng no tu. Chng l thnh phn cu to ca raffinose, hemicellulose.
pectine...
* D - Fructose cn gi l levulose v lm quay mt phng nh sng
phn cc v pha tri.
Fructose cn gi l ng qu, c trng thi t do trong tri cy
chn v mt ong. Chng l thnh phn ca disaccharide saccharose. Trong c
th ta cn thy dng ester vi phosphoric acid ng vai tr quan trng trong
trao i cht. Fructose c ngt rt ln, dng c ngt bng 1/3 dng .

23

24

1.2. Oligosaccharide
1.2.1. Disaccharide
S to thnh disaccharide l do s kt hp ca 2 monose cng loi
hay khc loi nh lin kt glucosidic. Lin kt glucosidic c th c to
thnh gia -OH glucoside ca monose ny vi -OH glucoside ca monose
kia, hay gia mt nhm -OH glucoside ca monose ny vi -OH ( khng
phi -OH glucoside) ca monose kia.
Disaccharide ch c tnh kh khi t nht mt trong 2 nhm -OH
glucoside trng thi t do. Ngha l disaccharide s khng c tnh kh
khi 2 nhm -OH glucoside lin kt vi nhau.
Cc disaccharide quan trng
* Maltose do 2 phn t - D-glucose lin kt vi nhau v tr C1 C4 to thnh. Cng thc cu to:

Maltose c nhm -OH glucoside trng thi t do nn c tnh kh.

Maltose c nhiu trong mm la v mch nha (maltum) nn gi n l maltose.
* Lactose (ng sa) do mt phn t D-galactose lin kt vi
mt phn t D- glucose v tr C1- C4.

* Saccharose do mt phn t D-glucose lin kt vi mt phn t

D-fructose v tr C1-C2. Do n khng c tnh kh, cn gi l ng
ma v c nhiu trong ma. D b thy phn khi un nng.

25

1.2.2. Trisaccharide
L oligosaccharide c cha 3 monosaccharide, ph bin trong
thin nhin l raffinose. Cng thc cu to nh sau: -D-galactopyranosyl
1-2 -D glucopyranosyl 1-2 -D fructofuranose. Do c cng thc nh trn
nn khng c tnh kh oxy. D b thy phn, di tc dng ca
fructofuranosidase s to thnh fructose v melobiose vi galactosidase
s to thnh galactose v saccharose.

1.3. Polysaccharide
Cn gi l glycan, ty thnh phn monose c trong polysaccharide
ngi ta chia chng ra lm: homopolysaccharide (ch cha mt lai
monosaccharide) v heteropolysaccharide (c t nht 2 lai
monosaccharide).

26
Polysaccharide ng vai tr quan trng trong i sng ng vt,
thc vt. Mt s polysaccharide thng gp nh tinh bt, glycogen,
cellulose...
1.3.1. Polysaccharide thc vt
1.3.1.1. Tinh bt
L polysaccharide d tr ca thc vt, do quang hp to thnh.
Trong c v ht c t 40 n 70% tinh bt, cc thnh phn khc ca cy
xanh c it hn v chim khong t 4 n 20%.
Tinh bt khng ha tan trong nc, un nng th ht tinh bt
phng ln rt nhanh to thnh dung dch keo gi l h tinh bt.
Tinh bt c cu to gm hai phn: amylose v amylopectin, ngoi
ra cn c khong 2% phospho di dng ester. T l amylopectin/amylose
cc i tng khc nhau l khng ging nhau, t l ny go np l ln
hn go t.
*Amylose
Chim 15 n 25% lng tinh bt, do nhiu gc D- glucose lin
kt vi nhau thng qua C1-C4 to thnh mch thng khng phn nhnh.
Trong khng gian n cun li thnh hnh xon c v c gi bn vng
nh cc lin kt hydro. Theo mt s ti liu trong amylose cn c cha
cc D- glucopyranose dng thuyn.

Amylose bt mu xanh vi iodine, mu ny mt i khi un nng,

hin mu tr li khi ngui. Mt c trng ha l khc cn ch l n b
kt ta bi ru butylic.

27

Ht tinh bt trong lc lp amylose

* Amylopectin
Cu to do cc phn t D- glucose lin kt vi nhau, nhng c
phn nhnh. Ch phn nhnh l lin kt C1-C6 glucosidic.

28

1.3.1.2. Cellulose
c cu to bi nhng phn t D-glucose lin kt vi nhau
bng lin kt 1-4 glucosidic.
Chng l thnh phn ch yu ca vch t bo thc vt. i vi
ngi th cellulose khng c gi tr dinh dng v cellulose khng b thy
phn trong ng tiu ha. Mt s nghin cu cho thy n c vai tr trong
iu ha tiu ho. ng vt n c thy phn c cellulose nh enzyme
cellulase.
Cellulose khng tan trong nc, tan trong dung dch Schweitzer.
Khi un nng vi H2SO4, cellulose s b thy phn thnh cc phn t
D-glucose.
Cellulose c dng hnh si di, nhiu si kt hp song song vi
nhau thnh chm nh cc lin kt hydro, mi chm (micelle) cha khang
60 phn t cellulose. Gia cc chm c nhng khong trng, khi ho g
khong trng ny cha y lignin v ta xem lp lignin ny nh l mt lp
cement. Lignin l cht trng hp ca coniferylic alcohol

Cc gc -OH ca cellulose c th to ester vi acid v d: to nitro

cellulose vi HNO3 , to acetyl cellulose vi CH3COOH.

29
1.3.1.3. Hemicellulose
Tn gi chung cho lp polysaccharide thng i theo vi cellulose
trong thc vt. Hemicellulose khng tan trong nc, tan trong dung dch
kim v thy phn bng acid d hn cellulose.
Khi b thy phn hemicellulose to thnh mt hn hp gm cc
hexose v pentose hay ch mt mnh hexose m thi. Trong hemicellulose
khi monose no chim a s th hemicellulose c tn tng ng vi
monose :
Xylose chim a s th hemicellulose c tn l Xylan,
Arabinose chim a s th hemicellulose c tn l Araban,
Galactose chim a s th hemicellulose c tn l Galactan...
Xylan c nhiu trong rm r, trong mt s c quan ca thc vt,
galactose c nhiu trong rm, g v cc loi ht.
1.3.1.4. Inulin
L polysacchride d tr ca thc vt c trng lng phn t
khong 5000-6000, do nhng phn t D- fructose lin kt vi nhau bng
lin kt 1-2 v tn cng bng mt phn t saccharose. Inulin c tm thy
trong c thc dc khong 40%. Ngi ta x dng inulin sn xut
fructose. xc nh inulin ngi ta thy phn n v xc nh bng phn
ng nh tnh Seliwanoff.
1.3.1.5. Pectin
L loi polysaccharide c nhiu trong qu , c v thn cy, thnh
phn chnh l galacturonic acid c nhm -COOH b methyl ha. Ngi ta
s dng rng ri pectin trong sn xut keo.
1.3.2. Polysaccharide ng vt
1.3.2.1. Glycogen
L polysaccharide d tr ng vt c tm thy trong gan v
c, hin nay cn tm thy trong mt s thc vt nh ng, nm.
C cu to ging amylopectin nhng phn nhnh nhiu hn, b
thu phn bi phosphorylase ( c coenzyme l pyrydoxal phosphate),
ct lin kt 1-6 cn enzyme debranching. Sn phm cui cng l cc phn
t glucose-1-P.

30
Pha ngoi glucose lin kt 1-6

Mch chnh

ht glycogen t bo gan

1.3.2.2. Hyaluronic acid

C cng thc cu to c lp li t n v sau:

Hyaluronic acid c trng lng phn t rt ln, c th ln n

nhiu triu, hyaluronic acid rt ph bin v l thnh phn quan trng ca
m lin kt, c tm thy trong dch khp xng, trong thy tinh th mt,
n tc dng nh mt lp cement bo v bn trong t bo chng li s
xm nhp ca vi khun cng nh cc cht l khc. khp xng n lm

31
cho dch c tnh trn gip c ng khi b au. Hyaluronic acid b thy
phn bi hyaluronidase, enzyme ny c tm thy trong vi khun gy
bnh, trong tinh trng. Hyaluronidase to d dng cho tinh trng i vo
non ca bun trng, mt khc n cng l yu t gip cho cc cht khc
v vi khun gy bnh i vo cc m trong c th.
1.3.2.3. Chondroitin
L heteropolysaccharide, thnh phn khng th thiu c m
xng sn.

1.3.2.4. Heparin
Heteropolysaccharide c tc dng chng li s ng mu v ngn
chn s bin i prothrombin thnh thrombin.

GlcA2S hoc IdoA2S

1.3.3. Mt s polysaccharide ph bin khc

1.3.3.1. Chitin
L homopolysaccharide, c v s, c, cc loi cn trng v
nm mc. N c cu to nh sau:

32

1.3.3.2. Dextran
c tm thy vi khun v nm men, cu to mch chnh l Dglucose1-6, nhnh l 1-3 v thnh thong c nhnh 1-2 hay 1-4. Do
c cu to 1-6 nn i vi ng vt, dextran khng b phn gii hay b
phn gii rt chm.

33
Dextran c di v hnh dng ging albumin, ngi ta thng
dng nhit thy phn khng hon ton dextran nhm thay th protein
ca huyt tng , dung dch 10% ca n hon ton trong sut. Trong cng
ngh ngi ta tng hp dextran v c gi l sephadex s dng trong
tch tng phn protein.

TI LIU THAM KHO

Ti liu ting Vit
1. Phm Th Trn Chu, Trn thi ng. 1999. Ho sinh hc, NXB Gio dc,
H Ni.
2. Qu Hai. 2004. Gio trnh Ha sinh i cng, Ti liu lu hnh ni
b Trng HKH Hu.
3. Trn Thanh Phong.2004. Gio trnh Ha sinh i cng, Ti liu lu
hnh ni b Trng HKH Hu.
4. L Ngc T (ch bin), L Vn Ch, ng Th Thu, Phm Quc Thng
Nguyn Th Thnh, Bi c Hi, Lu Dun, L Don Din, 2000. Ha sinh
Cng nghip, Nxb KH&KT, H Ni.

Ti liu ting Anh

1. Gilbert H. F. 1992. Basic concepts in biochemistry, Copyright by the
Mcgraw- Hill companies, Inc.
2. Lehninger A. L. 2004. Principles of Biochemistry, 4th Edition. W.H
Freeman.

34

Chng 2

Lipid
Cng nh saccharide, protein, lipid l cht hu c phc tp, ta c
th nh ngha nh sau:
* nh ngha rng: Lipid l cht tan c trong dung mi hu c,
khng tan trong nc, nh ngha ny khng phn nh ht tnh cht ca
cc lipid v:
- C lipid khng tan c trong dung mi hu c nh phospholipid
khng tan trong aceton.
- Nhng cng c cht khng phi lipid nhng tan c trong dung
mi hu c.
* nh ngha hp: Lipid l ester ca ru v acid bo. Tuy nhin
c nhng lipid do acid bo lin kt vi ru bng lin kt peptide.
* nh ngha dung ho: Lipid l nhng cht chuyn ho ca acid
bo v tan c trong dung mi hu c.
Lipid rt ph bin ng vt cng nh thc vt v tn ti di 2
dng m nguyn sinh cht (dng lin kt) v dng d tr (dng t do).
- M nguyn sinh cht: thnh phn ca mng t bo cng nh cc
bo quan khc v d: ty th, lp th... dng ny khng b bin i ngay c
khi con ngi b bnh bo ph hoc b i.
- Dng d tr (dng t do) c tc dng cung cp nng lng cho c
th, bo v cc ni quan, l dung mi cn thit cho mt s cht khc.
Cn c vo thnh phn nguyn t c mt, ngi ta chia lipid ra lm 2 loi
* Lipid n gin: trong phn t ch cha C, H, O.
* Lipid phc tp: ngoi C, H, O cn c mt s nguyn t khc
nh N, P, S.

2.1. Lipid n gin

2.1.1. Glycerid
Glycerid l ester ca ru glycerol v acid bo, l m d tr ph
bin ng vt v thc vt.

35

1- Stearoyl, 2- linoleoyl, 3-palmitoyl glycerol,

mt triacylglycerol hn tp

2.1.1.1. Glycerol
L triol khng mu, v ngt nhn. Khi t glycerol hay lipid c
cha glycerol vi cht ht nc s to acrolein c mi kht.
2.1.1.2. Acid bo
Acid bo thng gp l nhng acid bo c s carbon chn, mch
thng, c th no hay khng no v chui C xp theo hnh ch chi.
Tuy nhin cng c nhng acid bo ngoi nhm chc acid cn cha
nhng nhm chc khc nh ru, ketone, mch carbon c vng hay nhnh.
a. Acid bo chn, thng, no: CH3(CH2)nCOOH
C4 CH3 -(CH2)2 COOH butylic acid c nhiu trong c.
C6 CH3 -(CH2)4 -COOH caproic acid c trong b, sa d.
C8 CH3 - (CH2)6-COOH caprylic acid c trong b, sa d.
C10 CH3-(CH2)8 COOH capric acid
c trong b, sa d.

36
C12
n=10 lauric acid c trong du da.
C14
n=12 myristic acid
c trong du da.
C16
n=14 palmitic acid
c trong du ng vt,thc vt.
n=16 stearic acid c trong du ng vt,thc vt.
C18
C20
n=18 arachidic acid
c trong du lc.
b. Acid bo chn, thng, khng no
- Cha mt ni i (C): 10
9

C 16 (9-10): CH3-(CH2)5 -CH = CH- (CH2)7COOH

Palmitoleic acid : Tm thy trong du thc vt.
C18 (9-10): CH3-(CH2)7 -CH = CH- (CH2)7 COOH
Oleic acid: acid ny c ba ng phn.
C18 (6-7): Petroselenic acid
C18 (11-12): Vaccenic acid.
C18 (12-13): Heparic acid
- Acid bo c 2 ni i (C):
C18 (9-10,12-13): Linoleic acid
CH3-(CH2)4 -CH = CH- CH2-CH=CH-(CH2)7-COOH
C th khng tng hp c acid ny m ly t ngoi vo. Ngy
xa ngi ta quan nim acid ny l vitamin v gi l vitamin S. Nhng
thc cht l mt acid bo m c th cn vi mt lng ln.
- Acid bo c cha 3 ni i (C):
C18((9-10,12-13,15-16): Linolenic acid, c th khng tng hp
c acid ny.
- Acid bo c 4 ni i (C):
C20 (5-6,8-9,11-12,14-15): Arachidonic acid.
Ngoi ra cn c cc acid bo c cha ni ba nhng khng quan trng.
c. Acid bo c cha chc ru
Thng gp trong lipid phc tp v cha nhm ru gn chc acid
nn c tn l - hydroxy...

R-CH-COOH
OH

37
V d: - hydroxy lynoceric acid CH3 -(CH2 )21- CH-COOH
OH
Ricinoleic acid
CH3-(CH2)5 - CH - CH2 - CH = CH- (CH2)7 - COOH
10

OH
d. Gc R trong phn t acid c nhnh v c s C l
Phocenic acid:
Undecylonic acid :
e. Acid bo c vng
* Chaulmoogric acid:

* Sferculic acid:

CH3
CH CH2 COOH
CH3
CH2 = CH - (CH2)8 - COOH
- (CH2)12 - COOH

CH3 - CH2 - C = C - (CH2)7 - COOH

CH2
2.1.1.3. Tnh cht ca acid bo v triglyceride
* Tnh cht vt l:
a. im tan chy
im tan chy ph thuc vo s C ca acid bo, acid bo c chui
C di th im tan chy cao v ngc li. Nhng acid bo c C l c im
tan chy thp hn acid bo c s C nh hn n 1 n v . Ngoi ra tan
chy cn ph thuc vo s ni i trong phn t acid bo, acid bo cha
nhiu ni i th im tan chy cng thp.
b. si
Acid bo c chui C di th si cng cao, thng p dng tnh
cht ny tch cc acid bo ra khi nhau.
c. Tnh ho tan
- Trong nc: acid bo c chui C ngn (4,6,8) d tan, C10 kh tan,
C12 khng tan. Nu acid bo dng mui th d ha tan hn.
- Trong dung mi hu khng phn cc nh benzen, ether, ether
du ho acid bo d tan.

38
- Trong dung mi hu c phn cc nh aceton, acid bo kh ho
tan hay ho tan rt t.
* Tnh cht ho hc:
a. S hydrogen ho
Acid bo cha no c th kt hp vi H2 to thnh acid bo no
R - (CH2)n - CH =CH- (CH2)n - COOH + H2
R - (CH2)n -CH2 - CH2 - (CH2)n - COOH
Ngi ta dng phn ng ny ch to thc phm nh margarin.
b. S halogen ho
Acid bo khng no kt hp vi cc nguyn t thuc h halogen (F,
Cl, Br, I) to thnh acid bo no.
R - (CH2)n - CH = CH- (CH2)n - COOH + I 2
R - (CH2)n - CH - CH- (CH2)n - COOH
I
I
C th dng phn ng ny xc nh s ni i trong phn t
acid bo. Phn ng d dng hay kh xy ra tu thuc vo v tr ni i i
vi nhm carboxyl, ni i cng gn nhm carboxyl phn ng cng kh
xy ra.
xc nh s ni i ngi ta cn c vo ch s Iod.
Ch s Iod: L s gam Iod cn thit tc dng ln 100gam cht
bo. Do ch s iod cng ln th s ni i cng nhiu.
c. S thu phn:
Ester nn khi thu phn s to thnh ru glycerol v acid bo.
Tc nhn thy phn l acid, kim, nc hay enzyme.
* Thy phn bng nc cn nhit v p sut cao.
* Thy phn bng kim: NaOH hay KOH
Ch s x phng ho: s mg KOH cn thit trung ho 1g cht bo
Do ch s x phng cng ln th di mch cng ngn, nn
c dng xc nh di ca mch C.
xc nh tnh cht ca cht bo ngi ta cn cn c vo mt s
ch s khc nh ch s acid.
Ch s acid: s mg KOH dng trung ho tt c acid bo t do
c trong 1g cht bo.

39
* Thu phn bng enzyme: trong c th lipid b thu phn bng
enzyme lipase.
- Lipase dch trng tc dng vo v tr .
- Lipase ty tng tc dng vo v tr v .
d. S i ha:
Du m lu c mi v v kh chu gi l s i ha, mt trong
nhng nguyn nhn gy ra l do oxy khng kh kt hp vo ni i to
thnh peroxide. Nu oxy kt hp vo nguyn t carbon ng cnh lin kt
i th s to thnh hydrogen peroxide. Sau peroxide v hydrogen
peroxide s b phn gii to thnh aldehyde v ketone. Cc aldehyde v
ketone ny u l nhng cht c mi v v kh chu.
2.1.2. Cerid
Cng l ester ca ru v acid bo, nhit thng th rn, c
ng thc vt, thc vt n thng to thnh mt lp mng ph ln l,
thn, qu ca cy. Cng thc tng qut:
R O CO R
Ru trong cerid l ru cao phn t, ch cha mt nhm OH ,
mnh C khng phn nhnh, rt t khi mch C c vng V d: Ru
cetol:CH3 - (CH2)14-CH2OH.
Sp ong, sp c voi (spermaceti) l ester ca ru cetol v
palmitic acid.

Ngoi ra trong sp ong v sp c voi cn c ru t do, acid bo

t do v hydrocarbon.
2.1.3. Sterid
L ester ca ru sterol v acid bo. Ru sterol c vng v trng
lng phn t rt ln, sterol tiu biu l cholesterol, acid mt. Acid bo
thng l palmitic, oleic, ricinoleic.

40
2.1.3.1.Cholesterol
Cholesterol bao gm nhn phenanthrene kt hp vi cyclopentan
to thnh cyclopentanoperhydrophenanthrene. Cholesterol c mang nhm
ru C3, ni i C5 - C6 v 2 gc CH3 C10, C13 v mt nhnh
isooctan C17.
Cholesterol ch c ng vt, trong mu c khong 2.10-3, c
nhiu trong c, nhng m l lch, gan, da cng c cha cholesterol hay
cc cht chuyn ho ca n. Cholesterol c tm thy u tin sn mt,
sn mt l do s dn mt n rut non b nghn, mt cha nhiu
cholesterol nn kt ta li thng sn mt. Cholesterol l cht quan trng
trong s sinh tng hp acid mt, vitamin D v nhiu cht khc.
Cholesterol + acid bo
cholesterid

Trong thin nhin, cc sterol trng thi t do nhiu hn trng

thi sterid. c th ngi, ch 10% sterol b ester ha to thnh sterid. T
l sterol v sterid cc m khc nhau l khng ging nhau.
* L tnh ca cholesterol: kt tinh di dng vy ng nh nh x
c, dng kt tinh cng khc nhau tu theo mi trng kt tinh.
* Ho tnh:
- Phn ng vi acid bo do nhm -OH C3.
- B hydrogen ha hay halogen ho C5 - C6.

41
- Phn ng mu:
+ Phn ng Liebermann: Cholesterol cho mu xanh lc, mu ny
rt bn trong nhiu gi, phn ng ny c dng xc nh cholesterol
bnh vin.
+ Phn ng Salkowski: Cholesterol cho vnh mu .
2.1.3.2. Acid mt:
Acid mt c tm thy trong ng vt c v gm 3 dng sau:
cholic acid, deoxycholic v chenodeoxycholic acid.
Acid mt l cht c i vi ngi. V vy trong mt, acid mt lin kt
vi acetamin to thnh mt cht t c hn.
Ngoi cholesterol v acid mt cn c cc sterol khc cng c
ngun gc ng vt nh hormone nang thng thn, hormone tuyn sinh
dc, cc sterol c ngun gc thc vt nh ergosterol, stigmasterol...

2.2. Lipid phc tp

Khc vi lipid t do c nhim v cung cp nng lng , hm lng
lun thay i. Lipid phc tp c nhim v tham gia xy dng cc cu t
ca t bo, hm lng khng thay i hay rt t thay i.
2.2.1. Glycerophospholipid (phosphatid)
Chng ta c th hnh dung cu to chung ca glycerophospholipid nh sau:

Glycerophospholipid l diester ca phosphoric acid. Mt pha

phosphoric acid lin kt vi glycerol, pha kia lin kt vi X. Ty cu to
ca X ta c cc loi glycerophospholipid khc nhau:

42

Lecithin: Lc X l choline nn lecithin cn c gi l choline

phosphatid

Lecithin c nhiu trong lng trng g, trong u nnh, trong

mu, trong cc dy thn kinh. Qua cu to ta nhn thy n gm 2 phn
- Phn phn cc bao gm phosphoric acid v base nitrogen a nc.

43
- Phn khng phn cc bao gm cc gc acid bo, ru glycerol
ght nc.
Do c cu to nh trn nn lecithin trong nc s to thnh dung
dch gi l dung dch gi.
Nh c tnh va a nc, va ght nc m hnh nh
phospholipid tham gia trong vic bo m tnh thm mt chiu ca cc
mng cu trc di t bo.
Lecithin c th b thu phn bng acid, kim hay enzyme:
* Thu phn bng acid: tt c lin kt ester u b ct t.
* Thu phn bng kim: ta c acid bo dng mui,
glycerophosphate v choline. Nhng choline b phn hy cho
trimetylamin. Vi kim nh ch c th ct lin kt ester gia ru v acid
bo.
* Thu phn bng enzyme: c 4 loi enzyme lecithinase A, B, C v
D tc ng ln cc lin kt ester khc nhau:
B
CH2O - CO - R1
A
CHO - CO - R2
C
OH
CH2O - PO

Lecithinase A ct lin kt v tr ca lecithin cho acid bo v

lisolecithin.
Cephalin: Trong cu to ca cephalin X l colamine.
CH2O-CO-R1
CHO-CO- R2
O
CH2O-P = O
O-CH2- CH2 -H3N+

44

Tng t lecithin, cephalin (X l ethanolamine) c cu tao gm hai

phn a nc v ght nc, l thnh phn ca dy thn kinh v c nhiu trong no.
Lisocephalin c to thnh khi ct lin kt ester v tr , cng
c tnh cht ph hy hng cu nh lisolecithin.
Serinphosphatid: Gi l serinphosphatid khi X l serine.
Trong c th: lecithin, cephalin, serinphosphatid thng gp
dng hn hp bi c s bin i tng h gia serine, choline v
colamine.
2.2.2. Sphingophospholipid
y l lipid phc tp, trong ru a nguyn t l sphingosine.
Acid bo lin kt vi ru sphingosine bng lin kt peptid. Ty theo X ta
c cc loi sphingophospholipid khc nhau
Acid bo

Sphingophospholipid quan trng nht l sphingomyelin, y X

l: phosphocholine. Acid bo trong sphingomyelin l lignoceric, palmitic,

45
stearic hay nervonic. Sphingophospholipid l diaminophospholipid, khc
vi phosphatid l monoaminophospholipid.
Sphingophospholipid khng tan trong ethylic ether, da vo tnh
cht ny tch chng ra khi hn hp lipid
2.2.3.Glycolipid
Glycolipid l lipid phc tp khng cha phospho, trong thnh phn
ca chng c cha hexose, thng l galactose hay cc dn xut ca
galactose, i khi l glucose. Thuc nhm ny c MGDG, DGDG v
sulfolipid kh ph bin trong lc lp v cc thnh phn khc ca t bo l.

6-Sulfo-6-deoxy--D-glucopyranosyldiacyglycerrol
(sulfolipid)

2.2.4. Sphingolipid
Cerebroside: trong phn t cerebroside ru sphingosine lin kt
vi acid bo bng lin kt peptide, vi galactose (X) bng lin kt
glucosidic.

46
Cc cerebroside khc nhau v thnh phn acid bo, c nhiu trong
m thn kinh, hng cu, bch cu, tinh trng
Acid bo

Ganglyoside: cu to ging cerebroside nhng X l phc hp

oligosaccharide

Phc hp
ologosaccharide

TI LIU THAM KHO

Ti liu ting Vit
1. Phm Th Trn Chu, Trn thi ng. 1999. Ho sinh hc, NXB Gio dc,
H Ni.
2. Qu Hai.2004. Gio trnh Ha sinh i cng, Ti liu lu hnh ni
b Trng HKH Hu.
3. Trn Thanh Phong.2004. Gio trnh Ha sinh i cng, Ti liu lu
hnh ni b Trng HKH Hu.
4. L Ngc T (ch bin), L Vn Ch, ng Th Thu, Phm Quc Thng
Nguyn Th Thnh, Bi c Hi, Lu Dun, L Don Din, 2000. Ha sinh
Cng nghip, Nxb KH&KT, H Ni.

Ti liu ting Anh

1. LehningerA.L. 2004. Principles of Biochemistry, 4th Edition. W.H
Freeman.
2. Mead, Alfin-Slater, Howton & Popjk. 1986. Lipids: Chemistry,
biochemistry and nutrion, Plenum, New York.

47

Chng 3

Protein
Protein l hp cht hu c c ngha quan trng bc nht trong c
th sng, v mt s lng, protein chim khng di 50% trng lng kh
ca t bo; v thnh phn cu trc, protein c to thnh ch yu t cc
amino acid vn c ni vi nhau bng lin kt peptide. Cho n nay
ngi ta thu c nhiu loi protein dng tinh th v t lu cng
nghin cu k thnh phn cc nguyn t ho hc v pht hin c
rng thng thng trong cu trc ca protein gm bn nguyn t chnh l
C, H, O, N vi t l C 50%, H 7%, O 23% v N 16%. c bit t
l N trong protein kh n nh (li dng tnh cht ny nh lng
protein theo phng php Kjeldahl bng cch tnh lng N ri nhn vi
6,25). Ngoi ra trong protein cn gp mt s nguyn t khc nh S 0-3%
v P, Fe, Zn, Cu...
Phn t protein c cu trc, hnh dng v kch thc rt a dng,
khi lng phn t (MW) c tnh bng Dalton (1Dalton = 1/1000 kDa,
c l kiloDalton) ca cc loi protein thay i trong nhng gii hn rt
rng, thng thng t hng trm cho n hng triu. v d: insulin c khi
lng phn t bng 5.733; glutamat-dehydrogenase trong gan b c khi
lng phn t bng 1.000.000, v.v...
T lu ngi ta bit rng protein tham gia mi hot ng sng
trong c th sinh vt, t vic tham gia xy dng t bo, m, tham gia hot
ng xc tc v nhiu chc nng sinh hc khc. Ngy nay, khi hiu r vai
tr to ln ca protein i vi c th sng, ngi ta cng thy r tnh cht
duy vt v ngha ca nh ngha thin ti ca Engels P. Sng l phng
thc tn ti ca nhng th protein. Vi s pht trin ca khoa hc, vai tr
v ngha ca protein i vi s sng cng c khng nh. Cng vi
nucleic acid, protein l c s vt cht ca s sng.

3.1. Amino acid

3.1.1 Cu to chung
Amino acid l cht hu c m phn t cha t nht mt nhm
carboxyl (COOH) v t nht mt nhm amine (NH2), tr proline ch c
nhm NH (thc cht l mt imino acid).

48

Trong phn t amino acid u c cc nhm COOH v NH2 gn vi

carbon v tr . Hu ht cc amino acid thu nhn c khi thu phn
protein u dng L- amino acid. Nh vy cc protein ch khc nhau
mch nhnh (thng c k hiu: R).

Hnh: 3.1. Cng thc cu to chung ca cc amino acid

3.1.2. Phn loi amino acid

Hin nay ngi ta phn loi amino acid theo nhiu kiu khc nhau,
mi kiu phn loi u c ngha v mc ch ring. Tuy nhin, h u
da trn cu to ho hc hoc mt s tnh cht ca gc R. V d c ngi
chia cc amino acid thnh 2 nhm chnh l nhm mch thng v nhm
mch vng.
Trong nhm mch thng li tu theo s c mt ca s nhm
carboxyl hay s nhm amine m chia ra thnh cc nhm nh, nhm amino
acid trung tnh (cha mt nhm COOH v mt nhm NH2); nhm amino
acid kim (cha mt nhm COOH v hai nhm NH2); nhm amino acid
acid (cha hai nhm COOH v mt nhm NH2).
Trong nhm mch vng li chia ra thnh nhm ng vng hay d
vng v.v...
C ngi li da vo tnh phn cc ca gc R chia cc amino acid
thnh 4 nhm: nhm khng phn cc hoc k nc, nhm phn cc
nhng khng tch in, nhm tch in dng v nhm tch in m.
Tuy nhin, hin nay cch phn loi cc amino acid ang c
nhiu ngi s dng nht l da vo gc R ca amino acid v c chia
lm 5 nhm:
Nhm I. Gm 7 amino acid c R khng phn cc, k nc, l:
glycine, alanine, proline, valine, leucine, isoleucine v methionine. (Hnh 3.2)

49

Hnh 3.2. Cng thc cu to ca cc amino acid nhm I

Nhm II. Gm 3 amino acid c gc R cha nhn thm, l

phenylalanine, tyrosine v tryptophan (Hnh 3.3.)

Hnh 3.3. Cng thc cu to ca cc amino acid nhm II

50

Nhm III. Gm 5 amino acid c gc R phn cc, khng tch in,

l serine, theonine, cysteine, aspargine v glutamine (Hnh 3.4)

Hnh: 3.4. Cng thc cu to ca cc amino acid nhm III

Nhm IV. Gm 3 amino acid c R tch in dng, l lysine,

histidine v arginine, trong phn t cha nhiu nhm amin (hnh 3.5).

Hnh 3.5. Cng thc cu to cc amino acid nhm IV

51

Nhm V. Gm 2 amino acid c gc R tch in m, l aspartate

v glutamate, trong phn t cha hai hm carboxyl (hnh 3.6).

Hnh 3.6. Cng thc cu to ca cc amino acid nhm V

3.1.3. Cc amino acid thng gp

Cc amino acid thng gp l nhng amino acid thng c mt
trong thnh phn ca cc loi protein. Chng c khong 20 loi v c
thu nhn khi thu phn protein. Cc loi amino acid ny c tn gi, khi
lng phn t v k hiu c trnh by trn bng 3.1.
3.1.4. Cc amino acid khng th thay th
Cc amino acid c hnh thnh bng nhiu con ng khc nhau.
Nh bit, trong phn t protein c khong 20 loi amino acid, tuy nhin
trong c th ngi v ng vt khng tng hp c tt c cc loi m
phi a t ngoi vo qua thc n. Nhng amino acid phi a t ngoi
vo c gi l cc amino acid khng th thay th. Ngi ta bit c c
khong 8-10 loi amino acid khng th thay th bao gm: Met, Val, Leu,
Ile, Thr,Phe, Trp, Lys, Arg v His v ngy nay ngi ta cn xem Cys cng
l mt amino acid khng th thay th.
3.1.5. Cc amino acid t gp
Ngoi cc amino acid thng gp trn, trong phn t protein i
khi cn c mt s amino acid khc, l nhng loi t gp. Cc amino
acid ny l dn xut ca nhng amino acid thng gp nh: trong phn t
collagen c cha 4-hydroxyproline l dn xut ca proline, 5hydroxylysine l dn xut ca lysine v.v...Mt khc, mc d khng c
trong cu trc protein, nhng c hng trm loi amino acid khc cng c
th tn ti dng t do hoc lin kt vi hp cht khc trong cc m v t

52

bo, chng c th l cht tin thn hay l cc sn phm trung gian ca qu

trnh chuyn ho trong c th.
Bng 3.1. Cc amino acid thng gp
Tn amino
acid
Glycine
Alanine

Tn amino acid gi theo danh

php ho hc
-aminoacetic acid
-aminopropionic acid

Tn
vit tt
Gly
Ala

K
hiu
G
A

Khi lng
(MW)
75
89

Proline

-pyrolydilcarboxylic acid

Pro

115

Valine

-aminoisovaleric acid

Val

117

Leucine

-aminoisocaproic acid

Leu

131

Isoleucine

-amino--metylvaleric acid

Ile

131

Methionine

-amino--metylthiobutyric acid

Met

149

Phe

165

Phenylalanine -amino--phenylpropionic acid

Tyr

181

Tryptophan

-amino-hydroxyphenylpropionic acid
-amino--indolylpropionic acid

Trp

204

Serine

-amino--hydoxypropionic acid

Ser

105

Threonine

-amino--hydroxybutiric acid

Thr

119

Cysteine

-amino--thiopropionic acid

Cys

121

Aspargine

amid ca aspartate

Asn

132

Glutamine

amid ca glutamate

Gln

146

Lysine

, diaminocaproic acid

Lys

146

Histidine

-amino--imidazolpropionic acid

His

155

Arginine

-amino--guanidinvaleric acid

Arg

174

Aspartate

-aminosuccinic acid

Asp

133

Glutamate

-aminoglutarate

Glu

147

Tyrosine

3.1.6. Mt s tnh cht ca amino acid

3.1.6.1. Mu sc v mi v ca amino acid
Cc amino acid thng khng mu, nhiu loi c v ngt kiu
ng nh glycine, alanine, valine, serine, histidine, tryptophan; mt s
loi c v ng nh isoleucine, arginine hoc khng c v nh leucine. Bt

53

ngt hay cn gi l m chnh l mui ca natri vi glutamic acid

(monosodium glutamate).
3.1.6.2. Tnh tan ca amino acid
Cc amino acid thng d tan trong nc, cc amino acid u kh
tan trong alcohol v ether (tr proline v hydroxyproline), chng cng d
ho tan trong acid v kim long (tr tyrosine).
3.1.6.3. Biu hin tnh quang hc ca amino acid

Hnh 3.7. ng phn lp th ca alanine

Cc amino acid trong phn t protein u c t nht mt carbon bt

i (tr glycine) v th n u c biu hin hot tnh quang hc, ngha
l c th lm quay mt phng ca nh sng phn cc sang phi hoc
sang tri. Quay phi c k hiu bng du (+), quay tri c k
hiu bng du (-). Gc quay c hiu ca amino acid ph thuc vo
pH ca mi trng.
Tu theo s sp xp trong cu trc phn t ca cc nhm lin kt
vi carbon bt i m cc amino acid c cu trc dng D hay L (hnh 3.7)
gi l ng phn lp th. S ng phn lp th c tnh theo 2n (n l s
carbon bt i)
Hu ht cc amino acid khc hp th tia cc tm bc sng ()
khong t 220 - 280 nm. c bit cng nng 10-3M, trong bc sng
khong 280 nm, tryptophan hp th nh sng cc tm mnh nht, gp 4 ln
kh nng hp th ca tyrosine (hnh 3.8) v phenylalanine l yu nht.

54

Phn ln cc protein u cha tyrosine nn ngi ta s dng tnh cht ny

nh lng protein
hp ph

- Bc sng(nm)
Hnh 3.8. Ph hp th nh sng cc tm ca tryptophan v tyrosine

3.1.6.4. Tnh lng tnh ca amino acid

Trong phn t amino acid c nhm carboxyl -COOH nn c kh
nng nhng proton (H+) th hin tnh acid, mt khc c nhm amin- NH2
nn c kh nng nhn proton nn th hin tnh base. V vy amino acid c
tnh cht lng tnh.
Trong mi trng acid, amino acid dng cation (tch in
dng), nu tng dn pH amino acid ln lt nhng proton th nht
chuyn qua dng lng cc (trung ho v in), v tip tc tng pH amino
acid s nhng proton th hai chuyn thnh dang anion (tch in m). V
vy i khi ngi ta coi n nh mt di-acid.

cation

lng cc

anion

Hnh 3.9. Tnh lng tnh ca amino acid

55

Tng ng vi phn ly H+ ca cc nhm COOH v NH3+ c cc

tr s pK1 v pK2 (biu th phn ly ca cc nhm c 1/2). T trc
tin chuyn sang dng lng tnh v sau cng chuyn thnh dng anion.

phn ly ca H+
Hnh 3.10. ng cong chun ca glycine nng 1 M 25OC

Ngi ta xc nh c pHi (pI= pH ng in) = pK1 + pK2 / 2. V

d khi ho tan glycine vo mi trng acid mnh th hu nh glycine u
dng cation. Nu tng dn lng kim, thu c ng cong chun .
Trn ng cong chun thy rng glycine ln lt nhng 2 proton
trc Tng ng phn ly ca nhm COOH c mt na c tr s
pK1= 2,34 v phn ly ca NH3+ c mt na c tr s pK2= 9,60. Nh
vy ta c
2,34 + 9,60
pHi =
= 5,97
2

56

Bng: 3.2 Cc tr s pK ca cc amino acid thng gp

Tn cc
amino acid
Glycine
Alanine
Proline
Valine
Leucine
Isoleucine
Methionine
Phenylalanine
Tyrosine
Tryptophan
Serine
Theonine
Cysteine
Aspargine
Glutamine
Lysine
Histidine
Arginine
Aspartate
Glutamate

pI

Cc tr s pK
pK1(ca COOH) pK2(ca NH+3) pKR(ca R)
2,34
2,34
1,99
2,32
2,36
2,36
2,28
1,83
2,20
2,38
2,21
2,11
1,96
2,02
2,17
2,18
1,83
2,17
1,88
2,19

9,60
9,60
10,96
9,62
9,60
9,68
9,21
9,13
9,11
9,39
9,15
9,62
10,28
8,80
9,13
8,95
9,17
9,04
9,60
9,67

10,07

8,18

10,53
6,00
12,48
3,65
4,25

5,97
6,01
6,48
5,97
5,98
6,02
5,74
5,48
5,66
5,89
5,68
5,87
5,07
5,41
5,65
9,74
7,59
10,76
2,77
3,22

Mt khc ti pK1 + 2 s phn ly H+ ca nhm COO- glycine l

99%, ch 1% dng COOH v pK2 -2 dng NH3+ l 99%, ch 1% dng
NH2. Nh vy trong vng pH t pK1 + 2 n pK2 -2, phn t glycine ch
yu dng lng tnh v kt qu ta c mt vng ng in. Ngoi ra cc
amino acid trong gc R c thm nhm COOH hay NH2 s phn ly ca
chng s c thm mt tr s phn ly na-pKR (xem bng 3.2).
3.1.7. Cc phn ng ho hc ca amino acid
Cc amino acid u c nhm NH2 v COOH lin kt vi C, v vy
chng c nhng tnh cht ho hc chung. Mt khc cc amino acid khc
nhau bi gc R, v vy chng c nhng phn ng ring bit. Ngi ta chia
cc phn ng ho hc ca amino acid thnh 3 nhm:

57

- Phn ng ca gc R
Do cc amino acid c cu to gc R khc nhau, nn ngi ta c th
dng xc nh tng amino acid ring r nh phn ng c trng ca n,
v d phn ng oxy ho kh do nhm SH ca cysteine, phn ng to mui
do cc nhm COOH hay NH2 ca glutamate hay lysine, phn ng to
ester do nhm OH ca tyrosine v.v...
- Phn ng chung
L phn ng c s tham gia ca c hai nhm - COOH v - NH2.
Tt c cc amino acid trong phn t protein u phn ng vi hp cht
ninhydrin to thnh phc cht mu xanh tm, ring imino acid nh proline to
thnh mu vng. Phn ng c thc hin qua mt s bc nh sau:
Di tc dng ca ninhydrin nhit cao, amino acid to thnh
NH3, CO2 v aldehide, mch polypeptide ngn i mt carbon; ng thi
ninhydrin chuyn thnh diceto oxy hindriden. Diceto oxy hindriden, NH3 mi
to thnh tip tc phn ng vi mt phn t ninhydrin khc to thnh phc
cht mu xanh tm (hnh 3.11)

Hnh 3.11. Phn ng ca protein vi ninhydrin

- Phn ng ring bit

C th chia cc phn ng ring bit theo hai nhm - COOH v - NH2
+ Cc phn ng ca nhm - COOH. Ngoi cc phn ng ca
nhm COOH thng thng to ester, to amid, to mui ...th n cn c
nhng phn ng c trng khc nh c th b kh thnh hp cht ru
amino di s xc tc ca NaBH4.
R-NH2CH-COOH
R-NH2CH-CH2OH
Nhm COOH c th to thnh phc aminoacyl-adenylate trong
phn ng hot ho amino acid tng hp protein, hay c th loi CO2
vn gp rt nhiu trong qu trnh thoi ho amino acid.

58

+ Cc phn ng ca nhm - NH2. Nhiu phn ng ca nhm amine

c dng nh tnh v nh lng cc ch tiu ca amino acid nh:
nh lng nitrogen ca amino acid ngi ta cho phn ng vi
HNO2 gii phng N2.
R-CH-COOH
+ HNO2
NH2

R-CH-COOH
OH + N2 + H2O

nh lng amino acid ngi ta cho phn ng vi aldehyde to

thnh base schiff.
xc nh amino acid u N-tn cng ngi ta cho tc dng vi
2-4 dinitrofluobenzen (phn ng Sanger) hay phenyliothiocyanate (phn
ng Edman).

3.2. Peptide
3.2.1. Khi nim chung
Peptide l nhng protein thng c cu trc on ngn khong t hai
n vi chc amino acid ni vi nhau, c khi lng phn t thng di
6.000. Chng c th c tng hp trong t nhin hoc c hnh thnh do s
thoi ho protein. Trong cc peptide cc amino acid c lin kt vi nhau
thng qua lin kt peptide (hnh 3.12).

Hnh 3.12. S to thnh lin kt peptide

Lin kt peptide c bn cao bi cu trc ca n c 4 e', 2e'

thuc v lin kt C=O cn 2e' thuc v b i e' t do ca nguyn t N.
Lin kt gia C-N l lin kt phc tp, n c th chuyn t dng n

59

dng lai (trung gian) th b mt phn ghp i ca lin kt (hnh 3.13).

Ngi ta cho rng t l ca lin kt kp ny l khong 30% i vi lin
kt C-N v 70% vi lin kt gia C v O. Nh vy u ca mt chui
peptide l amino acid c nhm -amine (-NH2) t do c gi l u Ntn cng v u kia c nhm - carboxyl ( -COOH) t do c gi l
u C tn cng. Lin kt peptide to nn b khung chnh ca chui
polypeptide, cn cc gc R to nn mch bn ca chui (hnh 3.14)
Dng cng ho tr

Dng ion +

Dng lai (hybrid)

Hnh 3.13. S tn ti cc dng ca lin kt peptide

Mch chnh

Mch bn
Hnh 3.14. Mch bn v khung ca mt chui polypeptide

3.2.2. Cc phng php xc nh peptide

Ngoi phn ng ca nhm NH2 v COOH u tn cng, cc gc R
ca peptide cng cho nhng phn ng mu c trng ca cc amino acid
t do tng ng. Mt trong nhng phn ng mu c trng nht dnh cho
lin kt peptide l phn ng Biure, phn ng ny khng xy ra vi

60

amino acid t do v vi dipeptide. Trong mi trng kim mnh, lin kt

peptide phn ng vi CuSO4 to thnh phc cht mu tm (hnh 3.15.)
v c kh nng hp th cc i bc sng 540 nm.
y l phn ng c s dng rng ri nh lng protein.
Phng php xc nh protein theo Lowry cng da trn nguyn tc ca
phn ng ny bng cch thm thuc th Folin-Ciocalteau lm tng
nhy ca phn ng sau khi thc hin phn ng biure, ng thi da vo
cc gc Tyr, Try nh thuc th to phc mu xanh da tri.
O-

O-

C =NH
HN

HN = C
Cu

NH
C

NH

HN

Hnh 3.15. Phc mu tm trong phn ng Biure

C mt s phng php tch phn lp v xc nh thnh phn, s

lng v trnh t amino acid trong peptide.
V nguyn tc chung cc phng php tch phn lp v xc nh
peptide v c bn cng nh i vi protein. Tuy nhin peptide l nhng
on ngn ca chui polypeptide, v th c th b qua giai on ct chui
polypeptide thnh cc peptide nh m c th tch, phn lp ngay bng
phng php in di hay sc k tch ring tng peptide.
Sau khi tch ring cc peptide, tin hnh thu phn hon ton
thnh cc amio acid t do, xc nh cc amino acid , amino acid u N-tn
cng v amino acid u C-tn cng. Cc d liu thu c qua s phn tch
ny s c so snh i chiu v tng hp li.
V d, Puppy v Bodo phn tch mt peptide ca dch khi thu
phn Cytocrom C thu c cc d kin sau y:
- Thnh phn amino acid ca peptide sau khi c thu phn hon
ton v tin hnh sc k l 2Cys, 1 Ala, 2 Glu, 1His, 1Thr, 1Val,v 1Lys.

61

- Dng phng php Sanger xc nh c amino acid u N-tn

cng l Cys v phng php carboxypeptidase xc nh c amino acid
u C - tn cng l Lys.
- Cu to ca peptide nh (bng cch thu phn tng phn ban u
v xc nh cc amino acid, amino acid u N - tn cng v amino acid
u C - tn cng ca mi peptide nh):
Cys- Ala

Glu- Cys

Cys-(Ala,Glu)

Cys- His

Ala- Glu

Glu (Cys, His)

(Val- Glu)
Thr (Val, Glu)
Glu- Lys
Thr (Val, Glu, Lys)

Tng hp cc d kin trn, h xc nh c trnh t cc amino

acid ca peptide nghin cu l:
H2N-Cys-Ala-Glu-Cys-His-Thr-Val-Glu-Lys-COOH.
y l nguyn tc chung xc nh mt trnh t trong peptide.
Tuy nhin i vi nhng peptide di, vic xc nh rt phc tp.
3.2.3. Cc peptide thng gp trong thin nhin
Trong t nhin tn ti nhiu dng peptide c chc phn quan trong
lin quan n hot ng sng ca c th nh l cc hormon, cc cht
khng sinh hay nhng cht tin thn ca t bo vi khun v.v... Bn cnh
cng c nhng peptide chc phn cha r rng, c nhng peptide l
sn phm thu phn ang cn dang d ca protein. Trong phm vi ca
gio trnh ny xin c gii thiu mt s peptide quan trng,c nhiu
ngha i vi cho hot ng sng ca sinh vt.
3.2.3.1. Glutathion v cc cht tng t
Glutathion l mt tripeptide -glutamyl-cysteyl-glycine c cng
thc cu to nh sau:
NH2

CH2 SH

HOOC-CH-CH2-CH2-CO-NH-CH-CO-NH-CH2-COOH
Trong cu trc ca glutathion, nhm SH ca cysteine l nhm hot
ng, v vy ngi ta thng vit tt ch glutathion l G-SH. y l
glutathion dng kh. Trong mi trng hot ng glutathion c th

62

nhng hydrogen (H) thnh dng oxy ho (GSSG) v ngc li c th

nhn H thnh dng kh:
2GSH

-2H
+2H

G-S-S-G

Nh phn ng trn, glutathion ng vai tr ca mt h thng oxy

ho kh (vn chuyn hydrogen). Glutathion l mt trong nhng peptide
ni bo ph bin nht, n phn b nhiu trong cc m v cc c quan nh:
gan, thn, lch, tim, phi, hng cu v.v...
3.2.3.2. Cc hormone sinh trng (HGH)
Hormone sinh trng ca ngi (HGH-human growth hormone)
cn c tn gi STH (somatotropin hormone) l mt chui polypeptide bao
gm 191 amino acid c khi lng phn t 20.000. Trong cu trc c hai
cu disulfua c to thnh gia amino acid 53 -165 v gia amino acid
182-189. Hot ng sinh hc ca HGH l chui gm 134 amino
acid. HGH c cu to rt ging vi hormon lactogen ca rau thai
(85% amino acid ging nhau) v gn ging prolactin ca ngi (32%
amino acid ging nhau).
Hormon sinh trng c tc dng i vi s tng trng ni chung,
kch thch s to sn hn l to xng, n cng l mt hormon chuyn
ho. Hormon sinh trng kch thch s tng hp protein t nhng amino
acid c vn chuyn d dng vo trong t bo nh chng, v l
hormon gy tng ng huyt, sinh i tho ng, ng thi kch thch
s thoi ho lipid m bo nhu cu v nng lng cho c th, gy tng
acid bo t do trong huyt tng.
S thiu ht HGH nu xy ra trc tui dy th s dn n chng
ngi ln, s d tha HGH nu xy ra trc tui dy th s dn n chng
ngi khng l, nu xy ra sau tui dy th s dn n chng ngi b to
cc (pht trin chiu dy ca u, xng v mt).

63

3.2.3.3. Insulin
T 1953, Sanger (gii thng
Nobel 1958) nghin cu, tinh ch
v xc nh hon ton cu trc ca
phn t insulin. Phn t insulin bao gm
51 amino acid, c cu trc gm 2 chui
polypeptide, vi khi lng phn t 5.700
chui A c 21 amino acid, chui B c 30
amino acid. Hai chui c ni vi nhau
bng 2 cu disulfua. Trong chui A cng
hnh thnh 1 cu disulfua gia amino acid
th 6 v amino acid th 11. Phn c hiu
(c trng ca mt loi) ch tp trung vo
cc amino acid th 8-9-10, 12-14 ca
chui A v c bit l amino acid th 30
ca chui B (hnh 3.16).
Ngi ta cng xc nh c
cu trc ba chiu ca insulin v thy rng
cu trc phn t insulin c gi vng
bi nhiu lin kt mui, lin kt hydro
v lin kt cu disulphate gia chui A
v chui B.
Insulin c tc dng r nht trong tt c
cc hormon ca tuyn tu, c bit i
vi qu trnh chuyn ho glucid, n c
tc dng h ng huyt. Insulin cn
kch thch qu trnh tng hp v c ch
qu trnh thoi ho glycogen c, gan
v m m. c bit, insulin tng cng
tng hp acid bo, protein v kch
thch s ng phn. Tc dng quan
trng nht ca insulin l kch thch s
thm nhp glucose, mt s ose, amino
acid trogn t bo c v m. Do vy
insulin lm gim lng glucose trong
mu. Ngoi ra insulin cng lm gim
s tn to glucose do lm gim nng
enzyme nh pyruvat carboxylase v
fructose 1-6 diphosphatase.

Hnh 3.16. Cc amino acid ca

chui A v B insulin b

64
3.2.3.4. Oxytocin, Vasopressin Vasotocin
Oxytocin l mt peptide c 9 amino acid. ng vt c v,
oxytocin ch khc s thay i ca 2 amino acid l: amino acid v tr
th ba l isoleucine v amino acid v tr th tm l leucine (bng 3.3).
Vasopressin ca loi ch nhi c cu trc trung gian gia vasopresin v
oxytocin ca ng vt c v (amino acid th ba l isoleucin v amino acid
th tm l arginine v c tn l vasotocin). Vasopressin l mt peptide c
cu trc gm 9 amino acid. .
Bng 3.3. So snh cu trc ho hc gia oxytocin v vasopressin ca mt s
loi ng vt
Va-

Lysine
Vasopressin

1
2 3 4 5
6 7 8 9
Cys-Tyr-Phe-Glu-Asn-Cys-Pro-Lys-Gly-NH2

So2 3 4 5 6 7 8 9
Arginine 1
vasopressin Cys-Tyr-Phe-Glu-Asn-Cys-Pro-Arg-Gly-NH2

Ln,
H m
Phn ln
ng vt c
v

pres-

sin

Vasotocin

Oxytocin

1
2 3 4 5 6 7 8 9
Cys-Tyr-Ile-Glu-Asn-Cys-Pro-Arg-Gly-NH2
1
2 3 4 5 6 7 8 9
Cys-Tyr-Ile-Glu-Asn-Cys-Pro-Leu-Gly-NH2

ng vt c
xng sng,
khng c v
ng vt c
xng sng
c v, chim

Phn ln ng vt c v amino acid th 8 ca vasopressin l

arginine (arg-vasopressin), tr ln v h m, amino acid th 8 l lysine (lysvasopressin). Oxytocin c tc dng trn c trn ca t cung v tuyn v, gy
co khi t cung sinh con v kch thch s tit sa khi cho con b.
Vasopressin c tc dng chng li niu, tng cng ti hp thu
nc thn, ng thi lm co mch, do c tc dng tng huyt p.

3.3. Protein
3.3.1. Cu trc phn t bc 1, 2, 3, 4
V mt cu trc ngi ta phn bit protein gm bn bc: bc I, bc II,
bc III v bc IV (Hinh. 3.17)

65
Bc I

Bc II

Bc III

Bc IV

Hnh 3.17. S cc bc cu trc ca protein

3.3.1.1. Cu trc bc I
Cu trc bc I biu th trnh t cc gc amino acid trong chui
polypeptide, cu trc ny c gi vng bng lin kt peptide (lin kt cng
ha tr). Cu trc bc I l phin bn ca m di truyn, vic xc nh c cu
trc bc I l c s tng hp nhn to protein bng phng php ho hc
hoc bng k thut ca cng ngh sinh hc.
Hin nay nhiu loi protein bit c trnh t cc amino acid
trong chui polypeptide nh: ribonuclease l mt protein c 124 amino
acid c ni vi nhau thnh mt chui, c 4 cu disulfua (hnh 3.18);
hemoglobin l protein c 4 chui polypeptide, 2 chui ( mi chui 141
amino acid) v 2 chui (mi chui 146 amino acid); trypsinogen b (229
amino acid); kimotrypsin b (229 amino acid); alcol dehydrogenase nga
(374 amino acid); glutamate dehydrogenase b (500 amino acid) v.v..

66

Hnh 3.18. Cu trc bc nht ca ribonuclesae ca b

3.3.1.2. Cu trc bc II
Biu th s xon ca chui polypeptide, l tng tc khng gian gia
cc gc amino acid gn nhau trong mch polypeptide.

Xon

Lin kt
hydrogen
Xon
Hnh 3.19. Cc kiu xun trong cu trc bc II ca protein

Ni cch khc, cu trc bc II l dng khng gian cc b ca tng

phn trong mch polypeptide. Cu trc ny c lm bn nh cc lin kt

67
hydrogen c to thnh gia lin kt peptide k gn nhau, cch nhau
nhng khong xc nh. Theo Pauling v Cori (1951) cu trc bc II ca
protein bao gm 2 kiu chnh l xon v phin gp .
trong tc ngi ta tm thy keratin l loi protein c hai dng cu
trc: dng bnh thng v dng dui thng.; cu trc phin gp tm thy
trong fibroin ca t. Cu trc xun hin nay c tm thy trong nhiu loi
protein khc nhau Mt khc t l % xon trong cc protein khc nhau cng
thay i kh nhiu. V d trong hemoglobin v mioglobin l 75%; lysosyme l
35%; ribonuclease l 17% ..
Bng 3.4. S lng xon v phin gp trong chui n mt s protein
S gc (%)
Protein (s gc)
Chymotrypsin (247)
Ribonuclease (124)
Carboxypeptidase (397)
Cytochrom C (104)
Lysosyme (129)
Myoglobin (153)

Xon

Phin gp

14
26
38
39
40
78

45
35
17
0
12
0

Ngoi ra cn c kiu xon collagen c tm thy trong phn t

collagen (hnh 3.20).

Hnh 3.20. Cu trc kiu xun collagen

68
n v cu trc ca n l tropocollagen bao gm 3 mch
polypeptide bn vo nhau thnh mt dy cp siu xon (v mi mch n
c cu trc xon, chiu cao ca mi gc xon trn trc siu xon ny l
2,9 anstron, mt vng xon l 3,3 gc amino acid. Ba chui polypeptide
trong dy cp ni vi nhau bng cc lin kt hydrogen.
3.3.1.3. Cu trc bc III
Biu th s xon v cun khc ca chui polypeptide thnh khi,
c trng cho potein cu, l tng tc khng gian gia cc gc amino acid
xa nhau trong chui polypeptide. Trong nhiu protein hnh cu c cha
cc gc Cys to nn lin kt disulfua gia cc gc Cys xa nhau trong
chui polypeptide lm cho chui b cun li (xem myoglobin hnh 3.21).
Ngoi ra cu trc bc III cn c gi vng bng cc loi lin kt khc
nh Van der Waals, lin kt hydrogen, lin kt tnh in gia cc gc
amino acid v.v...
3.3.1.4. Cu trc bc IV
Biu th s kt hp ca cc chui c cu trc bc III trong phn t
protein. Hay ni cch khc, nhng phn t protein c cu trc t 2 hay
nhiu chui protein hnh cu, tng tc vi nhau trong khng gian to nn
cu trc bc IV. Mi mt chui polypeptide c gi l mt tiu n v
(subunit), chng gn vi nhau nh cc lin kt hydrogen, tng tc Van
der Waals gia cc nhm phn b trn b mt ca cc tiu n v lm
bn cu trc bc IV.

Myoglobin

Hemoglobin

Hnh 3.21. Cu trc bc III ca myoglobin v bc IV ca hemoglobin

(hemoglobin l protein c 4 chui polypeptide2 chui v 2 chui ;

myoglobin ch gm mt chui polypeptide)

69
3.3.2. Mt vi tnh cht ca protein
3.3.2.1. Tnh cht l-ho ca protein
- Tnh tan ca protein
Cc loi protein khc nhau c kh nng ho tan d dng trong mt
s loi dung mi nht nh, chng hn nh albumin d tan trong nc;
globulin d tan trong mui long; prolamin tan trong ethanol, glutelin ch
tan trong dung dch kim hoc acid long v.v...
- Tnh ngm nc ca protein
Trong mi trng nc, protein kt hp vi nc trng ln tr
thnh dng keo, hay ni cch khc protein trng thi hydrate ho, cc
phn t nc bm vo cc nhm a nc trong phn t protein nh -NH2,
-COOH..., lp o nc bao quanh phn t protein l mt trong cc yu t
lm bn vng cu trc, ngn cch cc phn t protein khng cho chng
dnh vo nhau thnh ta.
- nht ca dung dch protein
Khi protein ho tan trong dung dch, mi loi dung dch ca nhng
protein khc nhau c nht khc nhau (bng 3.5). Ngi ta c th li
dng tnh cht ny xc nh khi lng phn t ca protein ( nht
cng cao th khi lng phn t cng cao).
Bng 3.5. nht ca mt s protein
Protein
Gelatin
Albumin trng
Gelatin
Albumin trng

Nng %
(trong nc)
3,0
3,0
8,0
8,0

nht tng i
(ca nc =1)
4,54
1,20
14,2
1,57

- Hng s in mi ca dung dch protein

Khi thm cc dung mi hu c trung tnh nh ethanol, aceton vo
dung dch protein trong nc th tan ca protein gim v protein s kt
ta do vic gim mc hydrate ho ca cc nhm ion ho ca protein,
lp o mt nc, cc phn t protein kt hp vi nhau thnh ta. Nh
vy, hng s in mi ca dung mi lm ngn cn lc tnh in gia
cc nhm tch in ca protein v nc. Mi lin h c c
trng bi biu thc:

70
L1 - l2
F=
Dr2
Trong :

D - hng s in mi ca dung dch

F- lc tnh in gia cc ion tch in
L1 , l2 - in tch cc ion, r - khong cch gia cc ion
- Tnh cht in li ca protein
Cng nh cc amino acid, protein l cht in li lng tnh v trong
phn t protein c nhiu nhm phn cc mnh (bn gc R) ca amino acid
v d: nhm COOH th hai ca Asp, Glu; nhm NH2 ca Lys; nhm OH
ca Ser, Thr, Tyr v.v...Trng thi tch in ca cc nhm ny ph thuc
vo pH ca mi trng. mt pH no m tng in tch (+) v in
tch (-) ca phn t protein bng khng, phn t protein khng di chuyn
trong in trng gi l pHi (isoelectric - im ng in) ca protein.
Nh vy protein cha nhiu Asp, Glu (amino acid c tnh acid mnh) th
pHi trong vng acid, ngc li nhiu amino acid kim nh Lys, Arg, His
th pHi trong vng kim.
mi trng c pH < pHi , protein a s l mt cation, s in
tch dng ln hn s in tch m. pH > pHi phn t protein th hin
tnh acid, cho ion H+, do s in tch m ln hn s in tch dng,
protein l mt a anion, tch in m.
Bng 3.5. Gi tr pHi ca mt s protein
Protein
Pepsin
Albumin trng
Casein
Albumin huyt thanh
Gelatin

pHi
1,0
4,6
4,7
4,9
4,9

Protein
Globulin sa
Hemoglobin
Ribonuclease
Trypsin
Cytochrom C
Prolamin

pHi
5,2
6,8
7,8
10,5
10,6
12,0

Trong mi trng c pH = pHi ca protein, protein d dng kt t li

vi nhau v th ngi ta li dng tnh cht ny xc nh pHi ca protein
cng nh kt ta protein. Mt khc do s sai khc nhau v pHi gia cc
protein m c th iu chnh pH ca mi trng tch ring cc protein ra
khi hn hp ca chng.

71
- S kt ta bng mui ca dung dch protein
Mui trung tnh c nh hng r ti ho tan ca protein hnh cu:
vi nng thp chng lm ho tan nhiu protein. Tc dng khng ph
thuc vo bn cht ca mui trung tnh, m ph thuc vo nng mui v s
in tch ca mi ion trong dung dch, tc l ph thuc vo lc ion ca dung
dch ( = 1/2 C1 Z1, 2 trong l k hiu ca tng, C1 l nng ca mi
ion, Z1 l in tch ca mi ion). Cc mui c ion ho tr 2 (MgCl2, MgSO4...)
lm tng ng k tan ca protein hn cc mui c ion ho tr 1 (NaCl, NH4Cl,
KCl...). Khi tng ng k nng mui trung tnh th tan ca protein bt u
gim v nng mui rt cao, protein c th b kt ta hon ton.
Cc protein khc nhau b kt ta nhng nng mui trung tnh
khc nhau. Ngi ta s dng tnh cht ny chit xut v tch ring protein
khi hn hp. l phng php dim tch (kt ta protein bng mui). Th
d dng mui amonium sulfate 50% bo ho kt ta globulin v dung dch
amonium sulfate bo ho kt ta albumin t huyt thanh.
- Biu hin quang hc ca protein
Cng nh nhiu cht ho hc khc, protein c kh nng hp th v
bc x nh sng di dng lng t h. V vy c th o cng hp th
ca protein trong dung dch hay cn gi l mt quang thng k hiu bng
ch OD (Optical Density). Da trn tnh cht ngi ta sn xut ra cc
loi my quang ph hp th phn tch protein. Nhn chung protein u c
kh nng hp th nh sng trong vng kh kin (t 350nm- 800nm) v vng
t ngoi (t 320nm xung ti 180nm).
Trong vng nh sng kh kin protein kt hp vi thuc th hp th
mnh nht vng nh sng 750nm (nh lng protein theo Lowry).
i vi vng t ngoi dung dch protein c kh nng hp th nh sng
t ngoi hai vng bc sng khc nhau: 180nm-220nm v 250nm - 300nm.
bc sng t 180nm-220nm l vng hp th ca lin kt peptide
trong protein, cc i hp th 190nm. Do lin kt peptide c nhiu trong
phn t protein nn hp th kh cao, cho php nh lng tt c cc loi
protein vi nng thp. Tuy nhin vng hp th ny ca cc lin kt peptide
trong protein c th b dch v pha c bc sng di hn khi c mt s tp
cht ln trong dung dch protein. Mt khc chnh cc tp cht ny cng hp th
nh sng t ngoi vng bc sng 180nm-220nm. V th trong thc t
thng o hp th ca dung dch protein bc sng 220nm-240nm.
bc sng t 250nm-300nm l vng hp th cc amino acid thm
(Phe, Tyr, Trp) c trongphn t protein hp th cc i 280nm. C th s
dng phng php o hp th ca dung dch protein bc sng 280nm

72
nh tnh v nh lng cc protein c cha cc amino acid thm. Hm
lng cc amino acid thm trong cc protein khc nhau thay i kh nhiu, do
dung dch ca cc protein khc nhau c nng ging nhau c th khc
nhau v hp th bc sng 280nm. Ngoi ra c nhiu cht khc trong
dung dch cng c nh hng n hp th protein. V vy, cc phng
php o hp th vng nh sng t ngoi thng c dung nh lng
protein c tinh sch hoc xc nh protein trong cc phn on nhn
c khi sc k tch cc protein qua ct.
- Kt ta thun nghch v khng thun nghch protein
Khi protein b kt ta n thun bng dung dch mui trung tnh c
nng khc nhau hoc bng alcohol, acetone nhit thp th protein vn
gi nguyn c mi tnh cht ca n k c tnh cht sinh hc v c th ho
tan tr li gi l kt ta thun nghch. Cc yu t kt ta thun nghch c
dng thu nhn ch phm protein. Trong qu trnh kt ta thun nghch mui
trung tnh va lm trung ho in va loi b lp v hydrate ho ca protein,
cn dung mi hu c vn ho nc s ph hy lp v hydrate nhanh chng.
Trong ch phm protein nhn c cn ln cc cht dng kt ta, cn s
dng phng php thch hp loi b cc cht ny. V d c th dng
phng php thm tch loi b mui.
Ngc li kt ta khng thun nghch l protein sau khi b kt ta
khng th phc hi li trng thi ban u. S kt ta ny thng c s dng
loi b protein ra khi dung dch, lm ngng phn ng ca enzyme. Mt
trong nhng yu t gy kt ta khng thun nghch n gin nht l un si
dung dch protein (s ni k hn trong phn bin tnh protein sau).
- Cc phn ng ho hc ca protein
Cng nh cc amino acid v peptide, protein c cc phn ng ho hc
tng t, l: phn ng ca cc nhm -COOH, -NH2, gc R v phn ng to
mu c trng ca lin kt peptide nh phn ng biure (xem chng 2 v 3.
y xin c gii thiu thm mt s phn ng mu c trng khc, c ngha
quan trng trong pht hin protein v cc gc amio acid trong chui
polypeptide.
+ Phn ng vi thuc th Folin-Ciocateau
Thuc th Folin-Ciocateau c cha phosphomolipdic acid v
phosphovolframic acid cc cht ny lm tng nhy ca phn ng biure, mt
khc phn ng vi gc Tyr v Trp trong phn t protein. Cc gc amino acid
ny tham gia trong qu trnh to phc cht mu xanh da tri.
+ Cc phn ng mu c trng khc ca protein

73
Nhng phn ng ny c c l do s c mt ca cc nhm nh chc
ha hc xc nh trong phn t protein. C th s dng chng pht hin
amino acid, protein trong dung dch.
Phn ng xanthproteic: cc gc amino acid Tyr, Trp, Phe trong
protein tc dng vi HNO3 c to thnh mu vng v sau khi thm kim
s chuyn thnh da cam.
Phn ng Pauli: cc gc Tyr, His trong protein tc dng vi
diazobenzosulfonic acid to thnh mu anh o.
Phn ng Millon: gc Tyr tc dng vi thu ngn nitrate trong
HNO3 c to thnh kt ta mu nu t.
Phn ng Saccaguichi: gc Arg tc dng vi dung dch kim ca
-naphtolv hypobromitecho mu anh o.
Phn ng Adamkievich: gc Trp tc dng vi glyoxylic acid v
H2SO4 c to thnh vng tm mt phn cch.
3.3.2.2. Tnh cht v chc nng sinh hc ca protein
Protein l hp cht hu c c ngha quan trng bc nht trong c
th sng, v mt s lng, n chim khng di 50% trng lng kh ca
t bo. T lu, bit rng protein tham gia mi hot ng sng trong c
th sinh vt, ngoi vai tr l thnh phn chnh trong cu trc ca t bo v
m, protein cn c nhiu chc nng phong ph khc quyt nh nhng c
im c bn ca s sng nh s truyn t thng tin di truyn, s chuyn
ho cc cht do cc enzyme, cc khng th chng li bnh tt, cc hormon
dn truyn cc tn hiu trong t bo v.v... u c bn cht l cc protein.
Trong c th protein c mt s vai tr sau y:
- Chc nng to hnh
Ngoi cc protein lm nhim v cu trc nh v virus, mng t
bo, cn gp nhng protein thng c dng si nh: sclerotin c trong lp
v ngoi ca su b; fibroin ca t tm, nhn; collagen, elastin ca m
lin kt, m xng. Collagen m bo cho bn v tnh mm do ca
m lin kt.
- Chc nng xc tc
Hu ht cc phn ng xy ra trong c th u do cc protein c
bit ng vai tr xc tc, nhng protein c gi l cc enzyme. Mc
d gn y ngi ta pht hin c mt loi RNA c kh nng xc tc
qu trnh chuyn ho tin RNA thng tin (pre-mRNA) thnh RNA thng
tin (mRNA), ngha l enzyme khng nht thit phi l protein. Nhng nh
ngha c tnh cht kinh in: enzyme l nhng protein c kh nng xc tc

74
c hiu cho cc phn ng ho hc, l cht xc tc sinh hc vn c ngha
c bit quan trng. Hin nay ngi ta bit c khong 3.500 enzyme khc
nhau, nhiu enzyme c tinh sch, kt tinh v nghin cu cu trc.
- Chc nng bo v
Ngoi vai tr l thnh phn chnh trong xc tc, cu trc ca t bo
v m, protein cn c chc nng chng li bnh tt bo v c th. l
cc protein tham gia vo h thng min dch, c bit nhiu loi protein
thc hin cc chc nng ring bit to nn hiu qu min dch c hiu v
khng c hiu. Cc protein min dch c nhc n nhiu hn c l cc
khng th, b th v cc cytokine.
Ngoi ra protein cn tham gia vo qu trnh ng mu chng
mt mu cho c th hoc mt s loi c th sn xut ra nhng c t c
bn cht l protein nh enzyme nc rn, lectin v.v...,c kh nng tiu dit
k th bo v c th.
- Chc nng vn chuyn
Trong c th c nhng protein lm nhim v vn chuyn nh
hemoglobin, mioglobin, hemocyanin vn chuyn O2, CO2 v H+ i khp
cc m, cc c quan trong c th. Ngoi ra cn c nhiu protein khc nh
lipoprotein vn chuyn lipid, ceruloplasmin vn chuyn ng (Cu) trong
mu v.v...Mt trong nhng protein lm nhim v vn chuyn c nhc
n nhiu nht l hemoglobin.
- Chc nng vn ng.
Nhiu protein lm nhim v vn ng co rt nh myosin, actin
si c, chuyn v tr ca nhim sc th trong qu trnh phn bo v.v...
-Chc nng d tr v dinh dng
Cc protein lm nhim v d tr nh casein ca sa, ovalbumin ca
trng, feritin ca lch (d tr st) v.v...Protein d tr ny chnh l ngun cung
cp dinh dng quan trng cho cc t chc m, phi pht trin.
- Chc nng dn truyn tn hiu thn kinh
Nhiu loi protein tham gia vo vic dn truyn tn hiu thn kinh i
vi cc kch thch c hiu nh: sc t th gic rodopsin mng li mt.
- Chc nng iu ho
Cc protein lm nhim v iu ho qu trnh trao i cht thng
qua tc ng ln b my thng tin di truyn nh cc hormon, cc protein
c ch c hiu enzyme u c chc nng iu ho nhiu qu trnh trao
i cht, chng hn cc protein repressor vi khun c th lm ngng qu
trnh sinh tng hp enzyme t cc gen tng ng.

75
- Chc nng cung cp nng lng
Protein cng l ngun cung cp nng lng quan trng cho mi
hot ng sng trong c th. Trong c th cc protein b phn gii thnh
cc amino acid, t tip tc to thnh hng lot cc sn phm, trong
c cc ceto acid, aldehyde v carboxylic acid. Cc cht ny u b oxy ho
dn dn to thnh CO2 v H2O ng thi gii phng ra nng lng.
3.3.3. Phn loi protein
Protein l hp cht a dng v thnh phn cu trc, chc nng;
khi lng phn t ca chng cng rt khc nhau (bng 3.6). Do s
phn loi protein c th da vo nhng mc ch khc nhau. C hai cch
phn loi thng thng nht:
3.3.3.1. Phn loi theo hnh dng
- Protein dng si
C hnh dng di, thng l hnh si, chiu di ca phn t protein
si ln hn ng knh ca n hng trm ln. Protein si tng i bn
vng, khng tan trong nc v dung dch mui long, cc chui
polypeptide ca protein si nm dc theo mt trc thnh nhng si di.
Protein si l yu t cu trc c bn ca m lin kt ng vt cao cp,
th d collagen gn v m xng,elastin m lin kt n hi, -keratin
tc da v.v...
- Protein dng cu
C dng gn nh hnh cu hoc hinh bu dc, chiu di ca phn t
protein cu c th ln hn ung knh ca n t 3 n 10 ln. Protein cu khng
bn vng bng protein si, a s tan trong dung dch nc v d khuych tn,
thng c chc nng hot ng sng ca t bo nh cc enzyme, cc hormon, cc
protein vn chuyn nh albumin huyt thanh, hemoglobin v.v...
- Protein dng trung gian
Mt s protein dng trung gian, va c c im ca protein si,
va c c im ca protein cu. Th d myosin (yu t cu trc v chc
nng quan trong ca c) c cu trc hnh que di l c im ca protein
si, nhng li tan trong dung dch mui l c im ca protein cu.
Ngoi ra, trong nhm trung gian ny cn c cht tin thn ca fibrin l
fibrinogen.
3.3.3.2. Phn loi theo thnh phn ho hc
Protein gm hng trm, hng ngn, thm ch hng vn amino acid
ni vi nhau bng lin kt peptide to nn mt hay nhiu chui
polypeptide c cu trc rt phc tp.

76
Cn c s c hay vng mt ca mt s thnh phn c bn cht
khng phi protein m ngi ta chia protein thnh hai nhm:
Bng: 3.6 Khi lng (MW) v cu trc phn t ca mt s protein
Protein

Khi lng
(Dalton)

S gc
amino acid

S chui
polypeptide

Glucagon
Insulin
Ribonuclease (ty b)
Lysozyme (lng trng trng)
Myoglobin (tim nga)
Chymotripsin (ty b)
Hemoglobin (ngi)
Albumin (huyt thanh ngi)
Hexokinase (men bia)
Tryptophan-synthetase (E.coli)
-globulin (nga)
Glycogen-phosphorylase (c th)
Glutamate-dehydrogenase (gan b)
Synthetase ca acid bo (men bia)
Virus khm thuc l

3482
5733
12.640
13.930
16.890
22.600
64.500
68.500
96.000
117.000
149.000
495.000
1.000.000
2.300.000
40.000.000

29
51
124
129
153
241
574
550
800
975
1.250
4.100
8.300
20.000
336.500

1
2
1
1
1
3
4
1
4
4
4
4
40
21
2.130

- Protein n gin
Protein n gin l nhng protein m phn t ca chng gm ton
amino acid. Th d mt s enzyme ca tu b nh ribonuclease gm
ton amino acid ni vi nhau thnh mt chui polypeptide duy nht
(c 124 gc amino acid, khi lng phn t 12.640), chymotrypsin gm ton
amino acid ni vi nhau thnh ba chui polypeptide (c 241 gc amino acid,
khi lng phn t 22.600) v.v...Da theo kh nng ho tan trong nc hoc
trong dung dch ngi ta c th chia cc protein n gin ra mt s nhm nh
nh:
- Albumin: tan trong nc, b kt ta nng mui (NH4)2SO4 kh
cao (70-100%).
- Globulin: khng tan hoc tan t trong nc, tan trong dung dch long
ca mt s mui trung tnh nh NaCl, KCl, Na2SO4..., v b kt ta nng
mui (NH4)2SO4 bn bo ho.
- Prolamin: khng tan trong nc hoc dung dch mui long, tan
trong ethanol, isopropanol 70-80%.

77
- Glutein: ch tan trong dung dch kim hoc acid long.
- Histon: l protein c tnh kim d tan trong nc, khng tan trong
dung dch amoniac long.
- Protein phc tp
Protein phc tp l nhng protein m phn t ca chng ngoi cc
amino acid nh protein n gin cn c thm thnh phn khc c bn cht
khng phi l protein cn gi l nhm thm (nhm ngoi). Tu thuc vo
bn cht ca nhm ngoi m ngi ta chia cc protein phc tp ra cc
nhm nh v thng gi tn cc protein bt u bng tip u ng ch
bn cht nhm ngoi:
- Lipoprotein: nhm ngoi l lipid.
- Nucleoprotein: nhm ngoi l nucleic acid.
- Glucoprotein: nhm ngoi l carbohydrate v dn xut ca n.
- Phosphoprotein: nhm ngoi l phosphoric acid.
- Chromoprotein: nhm ngoi l hp cht c mu. Tu theo tnh
cht ca tng nhm ngoi m c nhng mu sc khc nhau nh (
hemoglobin), vng ( flavoprotein)...

78

TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n, i Duy Ban, Nguyn Hu Chn, nh H, L c
Trnh. 1980. Ho sinh hc. NXB Y hc, H Ni
2.Phm Th Trn Chu, Trn Th ng. 1999. Ho sinh hc. NXB Gio
dc, H Ni.
3. Nguyn Ln Dng, Phm Vn Ty, Nguyn nh Quyn. 1999. Vi sinh
vt hc. NXB Gio dc,H Ni.
4. L c Trnh. 1998. Hormon. NXB Y hc. H ni

Ti liu ting Anh

1. Dennison C. 2002. A Guide To Protein Isolation. Kluwer Academic
Publishers. New York, Boston, Dordrecht, Lodon, Moscow.
2. Fersht A., 1998, Structure and Mechanism in Protein Science, W. H.
Freeman, 3rd Rev Edit.
3. Lehninger A. L. 2004. Principle of Biochemistry, 4th Edition. W.H
Freeman, 2004
4. Liebler D. C. 2002. Introduction to proteomics. Humana Press Inc.
Totuwa, New Jersey.
5. Lodish H, 2003. Molecular Cell Biology. 5th ed, W.H Freeman, 2003
6. Walker J. M. 1996. The Protein Protocols Hand book. 2nd ed. Humana
Press Inc. Totuwa, New Jersey.

79

Chng 4

Nucleic acid
4.1. Thnh phn ho hc ca nucleic acid
Nucleic acid, vt cht mang thng tin di truyn ca cc h thng
sng, l mt polymer c hnh thnh t cc monomer l nucleotide.
Trong nucleic acid c cha cc nguyn t C, H, O, N v P. Hm lng P
t 8- 10% Mi nucleotide gm 3 thnh phn kt hp vi nhau theo t l
1:1:1, bao gm: nhm phosphate, ng pentose (l ng 5 carbon) v
mt base nit (nitrogen).
4.1.1 Base nit (Nitrogen)
Cc base nit (nitrogen) thuc phn t nucleic acid u l dn xut
ca base purine hoc pyrimidine. Cc base purine gm adenine (6-amino
purine) v guanine (2-amino, 6-aminopurine), cc base nit pyrimidine
gm thymine (2,6-dioxy, 5-methylpyrimidine), cytosine(2-oxy,6aminopyrimidine) v uracil (2,6 dioxypyrimidine).( Hnh 4.1.)

Hnh 4.1 Cng thc cu to ca cc base nit (nitrogen) trong nucleic acid

4.1.2 ng pentose
ng pentose trong nucleic acid gm c hai loi l ng deoxyribose
v ribose. S c mt ca 2 loi ng ny l mt trong nhng c im phn
bit DNA v RNA.
4.1.3 Phosphoric acid
L mt acid v c - H3PO4

80

Hnh 4. 2. Cng thc cu to ca hai loi ng pentose trong nucleic acid

4.1.4. S to thnh nucleoside

Nucleoside l sn phm thu phn khng hon ton ca nucleic
acid. Nucleoside gm c hai thnh phn l ng pentose v mt base nit
(nitrogen, thuc purine hay pyrimidine).

81

Hnh 4. 3. Cu to ho hc ca cc nucleoside v nucleotide

4.1.5. S to thnh nucleotide

Nucleotide cng l sn phm thu phn khng hon ton ca
nucleic acid. Nucleotide gm c ba thnh phn: ng pentose, mt base
nit (nitrogen) v phosphoric acid (Hnh 4.3).
4.1.6. S to thnh nucleic acid
Cc nucleotide c ni vi nhau bng lin kt phosphodiester
Acide thng qua cc nhm OH v tr C3 v C5 ca ng pentose
to thnh mt chui di gi l polynucleotide. Do lin kt phosphodiester
c tao thnh v tr C3 v C5 nn chui polypeptide c tnh phn cc:
u 5 thng c gc phosphate v u 3 thng c OH t do. Nucleic
acid gm hai loi phn t c cu to rt ging nhau l DNA
(desoxyribonucleic acid) v RNA (ribonucleic acid).
4.1.7. Mt s nucleotide quan trng khng tham gia cu to nucleic acid
- ADP v ATP

82
ADP (adenosindiphosphate) v ATP (adenosintriphosphate) l
nhng dn xut ca adenine, chng tham gia qu trnh phosphoryl ho-oxy
ho. ATP c coi l ngun phosphate cao nng trong t bo (Hnh 4.4).

Hnh 4.4. Cu to ho hc ca ADP v ATP

- cAMP(AMP vng)
Adenosinemonophosphate vng c hnh thnh t ATP, cAMP ch
tm thy t bo ng vt v vi khun, n thng lin kt vi mng bo
tng ca t bo v tham gia nhiu qu trnh chuyn ho. cAMP c th
c sinh ra nh mt s hormone hot ho adenylcyclase (Hnh 4.5.)
- UDP v UTP
UDP (uridinediphosphate) v UTP (uridinetriphosphate) u l
nhng dn xut ca uracil l nhng coenzyme quan trng trong cc phn ng
trung gian chuyn ho glucose v galactose. Ngoi ra, chng cn tham gia
trong vic hnh thnh nhng hp cht phosphate giu nng lng.

Hnh 4.5. Cu to ho hc ca AMP vng (cAMP)

- CDP v CTP
CDP (cytidindiphosphate) v CTP (cytidinetriphosphate) l nhng
dn xut ca cytidine. CTP cng l hp cht giu nng lng v co th
tham gia nhiu phn ng khc nhau nh: phosphoryl ho ethanolamine

83
dn n s sinh tng hp cephaline hoc phn ng vi phosphate choline
hnh thnh cytidinediphosphate-choline (CDP-Choline, hnh 4.6).

Hnh 4.6. Cu to ho hc ca CDP- cholin

4.1.8. Cc coenzyme nucleotide

Hin nay ngi ta bit c mt s nucleotide tham gia cu to nn
cc coenzyme quan trng nh vitamin B5 (pantothenic acid) trong
coezyme A (SH-CoA), vitamin B2 (riboflavine) trong coenzyme flavin
adenine inucleotide (FAD) v vitamin PP (nicotinamide) trong coenzyme
nicotinamide adenine dinucleotide (NAD) v.v...Chng c phosphoryl
ho khi lm chc phn nhm ngoi ca cc enzyme trong chuyn ho
trung gian, (hnh 4.7)

84

FAD

Hnh 4.7. Cu to ho hc ca mt s coenzyme

4.2. Cu trc ca nucleic acid

4.2.1 DNA (Desoxyribonucleic acid)
Phn t DNA l mt chui xon kp gm hai chui n. Mi chui
n l mt chui nucleotide. Mi nucleotide gm ba thnh phn: nhm
phosphate, ng desoxyribose v mt trong bn base v thng c k
hiu bng ch ci u tin ca cc base (A-adenine, C- cytosine, Gguanine v T- thymine). Hai chui n kt hp vi nhau nh cc lin kt
hydrogen hnh thnh gia cc base b sung nm trn hai chui: A b sung
cho T v C b sung cho G. Mi chui n c mt trnh t nh hng vi
mt u 5phosphate t do, u kia l 3 hydroxyl t do (quy c l 5
3. Hng ca hai chui n trong chui xon kp ngc nhau, nn c
gi l hai chui i song. Nhng phn tch cu trc hin i cho thy

85
cu trc ca DNA khng phi lun lun tng ng vi dng c gi l B
m Watson v Crick a ra. Do s tc ng ca cc hp cht c trng
lng nh hoc protein dng B c th chuyn sang dng A (nn nhiu
hn) hoc l dng Z (xon tri). Chng c th t gp li (DNA) hoc xon
mnh, v d mt chui kp DNA c di l 20 cm c nn trong mt
chromosome c kch thc l 5 m.
Lin kt hydrogen

1 nm

3,4 nm

0,34 nm
(a) Cu trc ca DNA

(b) Cu trc ha hc ca mt phn DNA

Hnh 4.8. Chui xon kp ca DNA

Phn t DNA trong nhim sc th ca sinh vt eukaryote dng

thng, cn phn ln t bo prokaryote (vi khun) phn t DNA c dng vng.
D dng no th cc phn t DNA u tn ti di dng cun cht. Trong t
bo eukaryote, DNA kt hp cht ch vi cc protein l histone.

86
Trc ng-phosphate

Cc base

Thymine (T)

Adenine (A)

Cytosine (C)

DNA nucleotide
Phosphate
ng (deoxyribose)

Guanine (G)

Hnh 4.9. Cu trc ca cc nucleotide in hnh.

DNA ca eukaryote c kch thc rt ln (v d DNA ngi c th

di n 1 m) nn cu hi t ra l phn t ny phi c nn nh th no
vo th tch rt hn ch ca nhn. Vic nn c thc hin nhiu mc
, mc thp nht l nucleosome v mc cao nht l cuo trc nhim
sc cht. Tht vy, ng knh ca chui xon DNA ch l 20 A , trong khi
si nhim
sc cht quan
st di knh hin vi in t c ng knh
o
o
100 A , i khi t 300 A . iu ny chng t phn t DNA tham gia hnh
o
thnh nhng cu trc phc tp hn (Hnh 4.10). Si c ng knh 100 A
l mt chui nhiu nucleosome. l nhng cu trc hnh thnh t mt

87
o

chui DNA qun quanh mt li gm 8 phn t histon. Si 100 A ny

o
c t chc thnh cu trc phc tp hn l si c ng knh 300 A .
Trong nhn t bo, cc si va k trn kt hp cht ch vi nhiu protein
khc nhau v c vi cc RNA to hnh nhim sc cht, mc t chc
cao nht ca DNA
Chui xon kp DNA

2 nm

Dng xu chui ca
nhim sc th

11 nm

Si nhim sc cht
ng knh 30 nm gm
cc nucleosome c
ng gi

30 nm

Vng nhim sc th
dng lng lo

300 nm

Vng nn cht ca
nhim sc th trung k

700 nm

Nhim sc th trung k

1400 nm

Hnh 4.10. Cu trc nucleosome v nhim sc th.

Phn t DNA c sp xp trn nhim sc th lm cho chiu di

ngn li hn 50.000 ln.
Cc DNA eukaryote c c im khc vi DNA prokaryote. Ton
b phn t DNA prokaryote u mang thng tin m ha cho cc protein
trong khi DNA eukaryote bao gm nhng trnh t m ho (cc exon)

88
xen k vi nhng trnh t khng m ho (intron). Cc trnh t m ho
eukaryote chm ngp trong mt khi ln DNA m cho n nay vn cha
r tc dng. Ty theo mc hin din ca chng trong nhn, cc trnh t
DNA c chia lm ba loi:
- Cc trnh t lp li nhiu ln. V d: ng vt c v cc trnh t
ny chim 10-15% genome (h gen). l nhng trnh t DNA ngn (10200 kb), khng m ho, thng tp trung nhng vng chuyn bit trn
nhim sc th nh vng tm ng (trnh t CEN) hay u cc nhim
sc th (trnh t TEL). Chc nng ca cc trnh t ny cha r, c th
chng tham gia vo qu trnh di chuyn DNA trn thoi v sc (trnh t
CEN) hoc vo qu trnh sao chp ton vn ca phn DNA nm u mt
nhim sc th (trnh t TEL).
- Cc trnh t c s ln lp li trung bnh. V d: genome ngi
cc trnh t ny chim 25-40 %. Chng a dng hn v c kch thc ln
hn (100-1.000 kb) cc trnh t lp li nhiu ln. Cc trnh t ny phn b
trn ton b b gen. Chng c th l nhng trnh t khng m ha m
cng c th l nhng trnh t m ha cho rRNA, tRNA v RNA 5S.
- Cc trnh t duy nht: l cc gen m ha cho cc protein, c trnh
t c trng cho tng gen.
Mt c im ca phn t DNA c ngha rt quan trng c s
dng vo phng php lai phn t. l kh nng bin tnh v hi tnh.
Bin tnh l hin tng hai si n ca phn t DNA tch ri nhau khi cc
lin kt hydrogen gia cc base b sung nm trn hai si b t do cc tc
nhn ha hc (dung dch kim, formamide, urea) hay do tc nhn vt l
(nhit). Sau , nu iu chnh nhit v nng mui thch hp, cc
si n c th bt cp tr li theo nguyn tc b sung, hnh thnh phn
t DNA ban u, l s hi tnh.
4.2.2 RNA (Ribonucleic acid )
Phn t RNA c cu to tng t DNA vi ba im khc bit sau:
- Phn t RNA l chui n.
- ng pentose ca phn t DNA l deoxyribose c thay bng ribose.
- Thymine, mt trong bn loi base hnh thnh nn phn t DNA,
c thay th bng uracil trong phn t RNA.
Cu trc v chc nng ca RNA c s bin i r rt. V c bn
RNA ch l cht mang thng tin di truyn virus, sau ngi ta chng
minh rng n khng nhng ng vai tr c bn vic chuyn thng tin di
truyn m cn c vai tr cu trc khi to nn phc h RNA-protein.

89
Theo mt l thuyt tin ha m i din l Manfred Eigen, RNA l
cht mang thng tin di truyn, thnh vin trung gian ca s biu hin gen,
thnh phn cu to v l cht xc tc. Nhm OH ru v tr th hai ca
ribose cn thit cho a chc nng lm nhiu lon s to thnh chui kp,
qua lm tng khng bn vng ca lin kt photphodieste.
Trong t bo c ba loi RNA chnh, c cc chc nng khc nhau:
- Cc RNA thng tin (mRNA)
mRNA l bn sao ca nhng trnh t nht nh trn phn t DNA,
c vai tr trung tm l chuyn thng tin m ha trn phn t DNA n b
my gii m thnh phn t protein tng ng. Cc RNA c cu trc a dng,
kch thc nh hn so vi DNA v ch cha thng tin m ha cho mt hoc vi
protein v ch chim khong 2-5% tng s RNA trong t bo.
Qu trnh chuyn thng tin c th hin nh sau:
Phin m
DNA

Dch m
RNA

Protein

Escherichia coli, kch thc trung bnh ca mt phn t mRNA

khong 1,2 kb.
- RNA vn chuyn (tRNA)
tRNA lm nhim v vn chuyn cc amino acid hot ha n
ribosome tng hp protein t cc mRNA tng ng. C t nht mt loi
tRNA cho mt loi amino acid. tRNA vn chuyn cha khong 75
nucleotide (c trng lng khong 25 kDa), l phn t RNA nh nht. Cc
tRNA c cu trc dng c ba l (Hnh 7). Cu trc ny c n nh nh
cc lin kt b sung hin din nhiu vng ca phn t tRNA. Hai v tr
khng c lin kt b sung ng vai tr c bit quan trng i vi chc
nng ca tRNA:
- Trnh t anticodon gm ba nucleotide.
- Trnh t CCA, c kh nng lin kt cng ha tr vi mt amino
acid c trng.
- RNA ribosome (rRNA)
rRNA l thnh phn c bn ca ribosome, va ng vai tr xc tc
v cu trc trong s tng hp protein.
Ty theo h s lng rRNA c chia thnh nhiu loi: eukaryote
c rRNA 28S, 18S, 5,8S v 5S, cn cc rRNA E. coli c ba loi: 23S,

90
16S v 5S. rRNA chim nhiu nht trong ba loi RNA (80% tng s RNA
t bo), tip n l tRNA v mRNA ch chim 5%.
Ribosome ca mi t bo u gm mt tiu n v nh v mt tiu
n v ln. Mi tiu n v c mang nhiu protein v rRNA c kch thc
khc nhau.
Amino acid

anticodon

.
Hnh 4.11. M hnh cu trc ca mt tRNA

- Mt s loi RNA khc.

T bo sinh vt nhn chun cn cha mt s loi RNA khc, chng
u c vai tr nht nh trong b my tng hp protein nh:
+ snRNA (small nuclear) l nhng phn t RNA nh tham gia vo
vic ghp ni cc exon.
+ hn RNA( heterogenous nuclear) l nhng RNA khng ng nht
nhn t bo.
+ scRNA (small cytoplasmic) l nhng RNA nh ca t bo cht.

91
Bng 4.1. Cc phn t RNA trong E. coli
Loi

Tng s tng
i (%)

rRNA

80

tRNA
mRNA

15
5

H s
lng (S)

Khi lng phn

t (kDa)

S lng
nucleotide

23
16
5
4

1,2 103
0,55 103
3,6 101
2,5 101
Khng ng nht

3700
1700
120
75

Nhn chung tt c RNA trong t bo u c tng hp nh enzyme

RNA polymerase. Enzyme ny i hi nhng thnh phn sau y:
a) Mt khun mu, thng l DNA chui i.
b) Tin cht hot ha: Bn ribonucleoside triphosphate: ATP, GTP,
UTP v CTP.
Sinh tng hp RNA ging DNA mt s im, th nht hng tng
hp l 5 3, th hai l c ch ko di ging nhau: nhm 3-OH u
cui ca chui tng hp l v tr gn kt ca nucleoside triphosphate tip
theo. Th ba, s tng hp xy ra do thy phn pyrophosphate.
Tuy nhin, khc vi DNA l RNA khng i hi mi (primer).
Ngoi ra RNA polymerase khng c hot tnh nuclease sa cha khi
cc nucleotide b gn nhm.
C ba loi RNA trong t bo c tng hp trong E. coli nh mt
loi RNA polymerase. ng vt c v cc RNA khc nhau c tng
hp bng cc loi RNA polymerase khc nhau.

4.3. Mt s tnh cht ca nucleic acid

Dung dch nucleic acid c nht cao, c hot tnh quang hc (lm
quay mt phng nh sang phn cc).
Nucleic acid hp th mnh vng nh sng t ngoi c bc sng
250-280 nm, cc i hp th 260 nm. Tnh cht ny c s dng
dnh lng nucleic acid v xc nh sch ca nucleic acid.
Khi un nng dung dch nucleic acid nhit cao, thm acd hoc
kim ion ho cc base ca n, nucleic acid b bin tnh. Phn t DNA
xon kp b tho ri, hp th bc song 260 nm tng ln. S tng
hp th ny gi l hin tng tng sc t (hyperchromism). Nhit lm
mt mt na cu trc xon kp cu phn t DNA c gi l nhit

92
nng chy (melting temperature) vit tt l Tm. Cc DNA giu cc base G
v C c nhit nng chy cao.
DNA phn ng vi thuc th fucsin to thnh mu (phn ng
Feulgen), phn ng ny thng s dng trong ho t bo.
phn bit DNA v RNA ngi ta dng cc phn ng c trng
vi thuc th orcine to thnh mu xanh lc bn, desoxyribose ca DNA
phn ng vi diphenylamine to thnh mu xanh da tri bn.

TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n, i Duy Ban, Nguyn Hu Chn, nh H, L c
Trnh. 1980. Ho sinh hc. NXB Y hc, H Ni
2. Phm Th Trn Chu, Trn Th ng. 1999. Ho sinh hc. NXB Gio
dc, H Ni.
3. H Hunh Thu Dng. 1998. Sinh hc phn t. NXB Gio dc, H
Ni.
4. Phm Thnh H. 2001. Di truyn hc, (ti bn ln th 3) NXB Gio
dc, H Ni.

Ti liu ting Anh

1. Lehninger A.L. , 2004. Principle of Biochemistry , 4th Ed., E-Book
2. Lodish H., 2003. Molecular Cell Biology. 5th Ed. E-book

92

Chng 5

Vitamin
Vitamin l mt nhm cht hu c c cc tnh cht l, ho hc rt
khc nhau. Tc dng ca chng trn cc c th sinh vt cng rt khc nhau
nhng u rt cn thit cho s sng ca sinh vt, nht l i vi ngi v
ng vt. Khi thiu mt loi vitamin no s dn n nhng ri lon v
hot ng sinh l bnh thng ca c th.
Vitamin c tng hp ch yu thc vt v vi sinh vt. ngi
v ng vt cng c th tng hp c mt s Vitamin nhng rt t nn
khng tho mn nhu cu ca c th m phi tip nhn thm ngoi vo
bng con ng thc n.
C nhiu loi Vitamin khc nhau. Tn Vitamin c gi theo
nhiu cch nh gi theo ch ci, gi theo danh php ho hc, gi theo
chc nng. V d Vitamin B1 cn c tn ha hc l Thiamin, ng thi
theo chc nng ca n cn c tn antinevrit.
C nhiu kiu phn loi Vitamin, nhng kiu phn loi c s
dng ph bin nht l da vo kh nng ho tan ca Vitamin vo cc dung
mi. Ngi ta chia Vitamin ra 2 nhm: Vitamin tan trong nc v Vitamin
tan trong m.
Vitamin tan trong nc ch yu tham gia vo cc qu trnh lin
quan ti s gii phng nng lng nh qu trnh oxi ho kh, qu trnh
phn gii cc hp cht hu c...
Vitamin tan trong m tham gia vo cc phn ng to nn cc cht
c chc nng cu trc cc m, cc c quan.

5.1. Vitamin tan trong nc

5.1.1. Vitamin B1 (Thiamin)
Vitamin B1 l loi Vitamin rt ph bin trong thin nhin, c bit trong
nm men, cm go, mm la m ... Trong cm go c hm lng
Vitamin B1cao nht. Vitamin B1 c tch ra dng tinh th vo nm
1912 v xc nh c cu trc ho hc ca n.

93

NH2
CH2

N
H3C

CH3

CH2 - CH2OH
S

Vitamin B1 bn trong mi trng acid, cn trong mi trng kim

n rt d b phn hu khi un nng. Trong c th B1 c th tn ti trng
thi t do hay dng Thiamin pyrophosphate. Thiamin pyrophosphate l
dng B1 lin kt vi H3PO4 v c vai tr quan trng trong qu trnh trao
i cht ca c th. Thiamin pyrophosphate l coenzyme xc tc cho qu
trnh phn gii cc ceto acid nh pyruvic acid, oxaloacetic acid.... V vy
khi thiu Vitamin B1 s chuyn ho cc ceto acid b ngng tr lm cho c
th tch lu mt lng ln cc ceto acid lm ri lon trao i cht v gy
nn cc trng thi bnh l nguy him.
Vitamin B1 ho tan tt trong mi trng nc v chu nhit kh
nn khng b phn hu khi nu nng. B1 c tng hp ch yu thc
vt v mt s vi sinh vt. Ngi v ng vt khng tng hp c B1 m
phi nhn t ngun thc n. Ngun cha nhiu Vitamin B1 l cm go,
ng, la m, gan, thn, tim, no, nht l nm men.
Khi thiu B1 c th pht sinh bnh beri-beri, cn gi l bnh t
ph, do qu trnh trao i cht b ri lon. Nhu cu Vitamin B1 ph thuc
vo iu kin ngh nghip, vo trng thi sinh l ca c th, vo la tui.
Nhu cu hng ngy ca ngi ln l 1-3mg, ca tr em 0,5-2mg.
5.1.2. Vitamin B2 (Riboflavin)
Vitamin B2 l dn xut ca vng Isoalloxazin, thuc nhm flavin. Trong
c th B2 lin kt vi H3PO4 to nn coenzyme FMN v FAD l nhng
coenzyme ca h enzyme dehydrogenase hiu kh.
trng thi kh Vitamin B2 bn vi nhit v acid.
Vitamin B2 c nhiu trong nm men, u, tht, sa, gan, trng. Khi
thiu Vitamin B2 s tng hp cc enzyme oxi ho kh b ngng tr lm
nh hng n qu trnh oxi ho kh to nng lng cho c th. ng
thi khi thiu Vitamin B2 vic sn sinh ra cc t bo ca biu b rut cng

94
b nh hng gy nn s chy mu rut hay ri lon hot ng ca d dy,
rut. Vitamin B2 cn gip c th khng khun tt hn.
Nhu cu Vitamin B2 hng ngy ca mt ngi khong 2-3mg.
CH2 - (CHOH)3-CH2OH

H3C

H3C

NH

N
O
Riboflavin (Vitamin B2)

5.1.3. Vitamin PP (Nicotinic acid, nicotinamid)

Vitamin PP l nicotinic acid v amid ca n l nicotinamid.
O
C

COOH

NH2
N

N
Nicotinic acid

Nicotinamid

Vitamin PP l thnh phn ca coenzyme NAD, NADP c trong cc

enzyme thuc nhm dehydrogenase k kh.
Vitamin PP gip c th chng li bnh pellagra (bnh da sn si).
Khi mc bnh pellagra s dn n sng mng nhy d dy, rut, sau
sng ngoi da.
Vitamin PP dng nicotinic acid bn vi nhit, acid v c kim nn
kh b phn hu, cn dng nicotinamid li km bn vi acid v kim.

95
Vitamin PP khng b bin i khi nu nng nn thc n gi c hm
lng PP qua x l.
Vitamin PP c nhiu trong gan, tht nc, tim, c bit l nm men.
Nu c th thiu Vitamin PP s nh hng n cc qu trnh oxi ho kh.
Vitamin PP c tc dng ngn nga bnh ngoi da, sng mng nhy rut,
d dy.
Hng ngy nhu cu ca mt ngi khong 15-25mg Vitamin PP.
5.1.4. Vitamin B6 (Pyridoxin)
Vitamin B6 tn ti trong c th 3 dng khc nhau: Piridoxol,
Pyridoxal, Pyridoxamine. Ba dng ny c th chuyn ho ln nhau
B

CH2OH
HO
H3C

CH2OH

N
Pyridoxol

Vitamin B6 l thnh phn coenzyme ca nhiu enzyme xc tc cho

qu trnh chuyn ho amino acid, l thnh phn cu to ca
phosphorylase...
Vitamin B6 c nhiu trong nm men, trng, gan, ht ng cc, rau
qu ...
Nu thiu Vitamin B6 s dn n cc bnh ngoi da, bnh thn
kinh nh au u, bnh rng tc, rng lng ...
Hng ngy mi ngi ln cn 1,5-2,8 mg, vi tr em cn 0,5-2mg
Vitamin B6.
5.1.5. Vitamin C (Ascorbic acid)
Vitamin C l ascorbic acid. Trong c th Vitamin C tn ti 2
dng: dng kh l ascobic acid v dng oxy ho l dehydro ascobic.

96
CH2OH

CH2OH

CHOH

CHOH
O

OH

OH

Ascorbic acid

Dehydro ascorbic acid

Vitamin C tham gia nhiu qu trnh sinh l quan trng trong c th:
- Qu trnh hydroxyl ho do hydroxylase xc tc.
- Duy tr cn bng gia cc dng ion Fe+2/Fe+3, Cu+1/Cu+2.
- Vn chuyn H2 trong chui h hp ph.
- Lm tng tnh khng ca c th i vi nhng iu kin khng
thun li ca mi trng, cc c t ca bnh nhim trng, lm gim cc
triu chng bnh l do tc dng ca phng x.
Ngoi ra Vitamin C cn tham gia vo nhiu qu trnh khc c vai
tr quan trng trong c th.
Vitamin C c nhiu trong cc loi rau qu ti, nht l trong cc
loi qu c mi nh cam, chanh, bi... Nhu cu hng ngy cn 7080mg/ngi. Nu thiu Vitamin C s dn n bnh hoi huyt, gim sc
khng ca c th, b bnh chy mu rng, li hay ni quan (bnh scorbutus).
5.1.6. Vitamin B12 (Cyanocobalamin)
Vitamin B12 c cu to phc tp, trong thnh phn c cha nhm
CN, CO, amin. Thnh phn chnh ca Vitamin B12 l nhm porphyrin.
Vitamin B12 gip cho vic to huyt cu t v hng cu. B12 tham
gia cc qu trnh tng hp nucleotide nh xc tc cc phn ng metyl ho
cc base Nit. Thiu B12 s gy bnh thiu mu c tnh.
Ngoi cc loi Vitamin trn, trong nhm Vitamin tan trong nc
cn mt s Vitamin khc nh Vitamin B5, Vitamin Bc, Vitamin H...
B

97

5.2. Vitamin tan trong cht bo

5.2.1. Vitamin A (retinol)
Vitamin A c 2 dng quan trng l A1 v A2. Vitamin A c hnh
thnh t .caroten l tin Vitamin A. T .caroten to thnh 2 phn
t Vitamin A.
CH3

CH3

CH3

CH3

CH2OH

CH3

Vitamin A1

Vitamin A c nhiu trong du c, lng trng. Trong thc vt c

nhiu tin Vitamin A (.caroten) nht l trong c c rt, qu c chua, qu
gc,... qu u .
Vitamin A c vai tr quan trng trong c ch tip nhn nh sng
ca mt, tham gia vo qu trnh trao i protein, lipid, saccharide. Thiu
Vitamin A s b bnh qung g, kh mt, chm ln, st cn, gim kh
nng khng ca c th i vi cc bnh nhim trng.
Nhu cu Vitamin A hng ngy i vi ngi ln 1-2mg, tr em
di 1 tui 0,5-1mg.
5.2.2. Vitamin D
Trong c th tn ti nhiu loi Vitamin D, trong quan trng
nht l dng D2 v D3. Cc Vitamin D l dn xut ca cc sterol. Trong c
th Vitamin D c to ra t tin Vitamin D c sn di da nh nh sng
mt tri c tia t ngoi.
Thiu hoc tha Vitamin D u nh hng n nng photpho
v canxi trong mu. Thiu Vitamin D tr em d b bnh ci xng,
ngi ln b bnh long xng.
Vitamin D c nhiu trong du c, m b, lng trng. Tin
Vitamin D c sn trong m ng vt. Hng ngy mi ngi cn khong
10-20mg, tr em di 30 thng cn nhiu hn: 20-40mg.

98
5.2.3. Vitamin E (Tocopherol)
Vitamin E c nhiu dng khc nhau. l cc dng , , , ...
tocopherol. Cc dng khc nhau ny c phn bit bi s lng v v tr
ca cc nhm metyl gn vo vng thm ca phn t. Trong cc loi
Vitamin E, dng -tocopherol c hot tnh cao nht:
CH3
O

CH3

H3C

CH3

CH3

CH3

(CH2)3 - CH - (CH2)3 - CH - (CH2)3- CH - CH3

HO
CH3

- Tocopherol

Vitamin E c nhiu cc loi rau xanh, nht l x lch, ht ng

cc, du thc vt, gan b, lng trng, mm ht ho tho ...
Vitamin E c tc dng nh cht chng oxi ho nn c tc dng bo
v cc cht d b oxi ho trong t bo. Vitamin E cn c vai tr quan trng
trong sinh sn. Nhu cu Vitamin E hng ngy khong 20mg cho mt
ngi ln.
5.2.4. Vitamin K
C nhiu loi Vitamin K, vi cng thc tng qut l
O
CH3
R
O
Vitamin K cn cho qu trnh sinh tng hp cc yu t lm ng
mu (prothrombin) cho nn Vitamin K l Vitamin chng chy mu, thiu
Vitamin K tc ng mu gim, mu kh ng.

99
Vitamin K c nhiu trong c linh lng, bp ci, rau m, c chua,
u, ng cc, lng trng, tht b ...
Thng ngi kho mnh, vi khun ng rut c kh nng
cung cp Vitamin K cho nhu cu ca c th, ch cn b sung thm
khong 0,2-0,3mg/ngy/ngi.
5.2.5. Vitamin F
Vitamin F l cc acid bo khng no nh linoleic acid, linolenic
acid, arachidonic acid ...
Vitamin F c tc dng nui da, tiu m. Thiu Vitamin F ng vt chm
ln, vim da, rng lng, hoi t ui.
Vitamin F c nhiu trong cc loi du thc vt.
5.2.6. Vitamin Q (Ubiquinon)
Vitamin Q ln u tin c tch ra t m ng vt vo nm 1955.
Cu trc v chc nng ca Vitamin Q gn tng t nh Vitamin K v F.
Vitamin Q tham gia vo cc qu trnh oxi ho-kh ca c th vi
chc nng thnh vin ca chui vn chuyn in t ca ty th.
Vitamin Q c trong nhiu i tng nh vi sinh vt, thc vt, ng
vt ....
5.2.7. Vitamin P (Rutin)
Vitamin P l nhm Vitamin c cu trc t flavon. Hin nay ngi
ta pht hin khong 150 loi cht flavonoid khc nhau c tc dng nh
Vitamin P, trong c 10 cht c nghin cu k.
Thiu Vitamin P xy ra hin tng tng tnh thm ca mao qun,
chy mu bt thng, mi mt, suy nhc c th. Vitamin P c tc dng
lm gim tnh thm ca thnh mao qun. Vitamin P cn c th tham gia
vo qu trnh oxi ho kh ca c th nh chc nng ca Vitamin C.
Nhu cu Vitamin P khong bng 50% nhu cu Vitamin C.

100

TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n (ch bin). 1979. Ha sinh i cng (tp I, II). NxB
KH&KT, H Ni.
2. Phm Th Trn Chu, Trn Th ng. 2000. Ha sinh hc. Nxb Gio
dc, H Ni.
3. Nguyn B Lc. 1997. Ha sinh. Nxb Gio dc, H Ni

Ti liu dch
1. Musil J.G., Kurz .K., Novakava .O. 1982
Sinh ha hc hin i theo s . Nxb Y hc, H Ni.

Ti liu ting Anh

1. Farkas G. 1984. Nvnyi anyagcserelettan. Akadmiai Kiad
Budapest.
2. Lehninger A. L., 2004. Principle of Biochemistry, 4th Edition. W.H
Freeman.

101

Chng 6

Enzyme
Enzyme l protein c kh nng xc tc c hiu cho cc phn ng
ha hc. Chng thc y mt phn ng xy ra m khng c mt trong sn
phm cui cng. Enzyme c trong nhiu i tng sinh hc nh thc vt,
ng vt v mi trng nui cy vi sinh vt.Hin nay ngi ta thu c
nhiu loi ch phm enzyme khc nhau v s dng rng ri trong nhiu
lnh vc nh y hc , nng nghip, cng nghip

6.1. Bn cht ha hc ca enzyme

Ngoi tr mt nhm nh RNA c tnh xc tc, tt c enzyme u
l protein. Tnh cht xc tc ph thuc vo cu to ca protein. Nu mt
enzyme b bin tnh hay phn tch thnh nhng tiu n v th hot tnh
xc tc thng b mt i, tng t khi bn thn protein enzyme b phn
ct thnh nhng amino acid. V vy cu trc bc 1, 2, 3, 4 ca protein
enzyme l cn thit cho hot tnh xc tc ca chng.
Enzyme, cng nh nhng protein khc, c trng lng phn t
khong 12.000 n hn 1000.000.Mt s enzyme cu to gm ton nhng
phn t L amino acid lin kt vi nhau to thnh, gi l enzyme mt thnh
phn. a s enzyme l nhng protein phc tp gi l enzyme hai thnh
phn. Phn khng phi protein gi l nhm ngoi hay coenzyme. Mt
coenzyme khi kt hp vi cc apoenzyme khc nhau (phn protein) th
xc tc cho qu trnh chuyn ha cc cht khc nhau nhng chng ging
nhau v kiu phn ng.
Mt s enzyme cn ion kim loi cho hot ng nh:
Cu2+
Cytochrome oxidase
2+
3+
Cytochrome oxidase, catalase, peroxidase
Fe hoc Fe
+
K
Pyruvate kinase
2+
Mg
Hexokinase,
glucose
6-phosphatase,
pyruvate kinase
Mn2+
Arginase, ribonucleotide reductase
Mo
Dinitrogenase
Ni2+
Urease
Se
Glutathione peroxidase
2+
Zn
Carbonic anhydrase , alcohol dehydrogenase,
cc carboxypeptidase A v B

102
Mt s coenzyme v chc nng vn chuyn nhm tng ng ca
chng nh sau:
Biocytin
Coenzyme A
5- Deoxyadenosylcobalamin
(coenzyme B12)
Flavin adenine dinucleotide
Lipoate
Nicotinamide adenine dinucleotide
Pyridoxal phosphate
Tetrahydrofolate
Thiamine pyrophosphate

CO2
Nhm Acyl
Nguyn t H v nhm alkyl
in t
in t v nhm acyl
Ion Hydride (:H-)
Nhm Amino
Nhm 1 Carbon
Aldehyde

6.2. C ch tc dng
Nhng quan im hin nay nhm gii thch c ch tc dng ca
enzyme u cho rng khi enzyme (E) tong tc vi c cht (S) s lm
gim nng lng hot ha cc phn ng ha sinh. Mun lm gim nng
lng hot ha cc phn ng enzyme cn tri qua nhiu giai on trung
gian v to thnh phc cht nht nh gia E v S.
Khi kt hp vi phn t enzyme, do kt qu ca s cc ha, s
chuyn dch ca cc electron v s bin dng ca cc mi lin kt tham
gia trc tip vo phn ng s lm thay i ng nng v th nng nn
phn t c cht tr nn hot ng v d dng tham gia phn ng.
Vic to thnh phc hp E-S giai on u xy ra rt nhanh v rt
khng bn. Do sau mt thi gian di mi c chng minh bng thc
nghim. Bng chng r rng nht v s tn ti ca phc hp E-S l thnh
cng ca hai nh ha sinh Nht Bn K. Iaglu v T. Ozava l tch c
phc E-S trong phn ng kh amin bng cch oxy ha (loi tr nhm
amine) mt amino acid dy D do oxydase xc tc.
Nhn chung ta c th hnh dung c ch tc dng ca enzyme ln c
cht to sn phm bng phng trnh tng qut nh sau:
E+S ' ES  P+E
Giai on 1: E kt hp vi S to thnh E-S. Giai on ny xy
ra rt nhanh, nh cc lin kt khng bn nh lin kt hydro, tng tc tnh

103
din, tng tc Van der Waals Mi loi lin kt i hi nhng iu kin
khc nhau v chu nh hng khc nhau khi c nc.
Giai on 2: Sau khi to phc, c cht c nhng bin i nht
nh v mt in t, cu hnh lm c cht tr nn hot ng hn, phn
ng c d dng to thnh sn phm P.
Trong nhiu phn ng do enzyme xc tc c 2 hay nhiu lai c
cht, v d hexokinase xc tc phn ng:
ATP + glucose hexokinase
ADP + glucose 6 phosphate
C ch enzyme xc tc cho phn ng 2 c cht c th nh sau:
a/ C ch to phc 3 thnh phn
S2

b/ C ch khng to phc 3 thnh phn

y l trng hp c cht th 2(S2) ch kt hp vo enzyme (

trng thi E) sau khi P1 c to thnh.

6.3. Trung tm hot ng (TTH) ca enzyme

T kt qu nghin cu v bn cht ho hc, v cu trc trung tm
hot ng , c ch tc ng, v trung tm hot ng chng ta c th c
mt s nhn xt chung v trung tm hot ng nh sau:
- L b phn dng lin kt vi c cht.
- Ch chim t l rt b so vi th tch ton b ca enzyme.
- Gm cc nhm chc ca amino acid ngoi ra c th c c cc ion
kim loi v cc nhm chc ca cc coenzyme.

104

i vi E mt thnh phn: TTH ch gm nhng nhm chc ca

cc amino acid nh nhm hydroxy ca serin, carboxy ca glutamic, vng
imidazol Cc nhm chc ca cc amino acid c th xa nhau trong chui
polypeptide nhng nh cu trc khng gian nn n gn nhau v mt khng gian.
i vi E hai thnh phn: TTH cng nh trn, cc nhm chc
ca cc amino acid tham gia to thnh TTH lin kt vi nhau bng cc
lin kt hydro. Ngoi ra trong TTH ca loi ny cn c s tham gia ca
coenzyme v c th c ion kim loi.
Theo Fisher TTH c cu trc c nh, khi kt hp vi c cht
to phc E-S ta c th hnh dung ging nh cha kha v kha. Ngy
nay ngi ta chng minh c rng: TTH ca enzyme ch c cu to
hon chnh khi c s tng tc vi c cht (thuyt tip xc cm ng ca
Koshland).

6.4. Tnh c hiu ca enzyme

Ngi ta chia tnh c hiu ra lm 3 kiu:
+ c hiu phn ng
+ c hiu c cht
+ c hiu khng gian
a/ c hiu phn ng: l biu hin ca mt enzyme ch
thng xuyn xc tc cho mt kiu phn ng nht nh, v d vn chuyn
hydro t cht cho (ru bc nht hay ru bc hai) n cht nhn (NAD+
hay NADP+) hay chuyn nhm amin t mt amino acid n mt ceto
acid. Cc phn ng loi th nht do dehydrogenase xc tc, cn phn ng
loi th hai do aminotransferase xc tc.

105
b/ c hiu c cht: Tu mc ngi ta chia thnh: c hiu
tng i v c hiu tuyt i
+ c hiu tuyt i: Enzyme ch tc dng ln mt c cht nht
nh, mt v d c tnh cht kinh in v chuyn ho tuyt i l urease,
enzyme ch phn gii ure:
Hng trm th nghim trn cc dn xut ca ure u cho thy
chng khng b phn gii di tc ng ca urease. Thc ra ngi ta
pht hin kh nng phn gii c cht hydroxyure nhng vi tc b hn
khong 120 ln.
+ c hiu nhm tuyt i: Cc enzyme ny ch tc dng ln
nhng cht c cng mt kiu cu trc phn t, mt lin kt v c nhng
yu cu xc nh i vi nhm nguyn t i vi nhm nguyn t gn
lin kt chu tc dng. v d : maltase ch phn gii lin kt glucosidic
c to thnh t glucoside ca glucose vi -OH ca monose khc.
+ c hiu nhm tng i: Cc enzyme khng c nhng yu cu i
vi nhm chc gn lin kt chu tc dng. v d lipase thu phn lipid.
c/ c hiu khng gian: Cc enzyme ch xc tc cho mt dng
ng phn no nh dng L hay dng D, dng cis hay trans m thi.

6.5. Cc yu t nh hng n tc ca phn ng enzyme

6.5.1. nh hng ca nng enzyme
Trong iu kin d tha c cht, ngha l [S] >>>[E] th tc
phn ng ph thuc vo [S], v= K[E] c dng y = ax. Nh ngi ta
o [E] bng cch o vn tc phn ng do enzyme xc tc.
C nhiu trng hp trong mi trng c cha cht km hm hay
hot ho th vn tc phn ng do enzyme xc tc khng ph thuc tuyn
tnh vi [E] .
v

[E]
Hnh 6.1: S ph thuc ca vn tc phn ng vo [E]

6.5.2 nh hng ca nng c cht [S]

Ta kho st trng hp n gin nht : ch mt c cht.

106

E v S

(1)
Gi v1 l vn tc ca phn ng to thnh phc cht ES.
Gi v-1 l vn tc ca phn ng phn ly phc cht ES to thnh
Gi v2 l vn tc ca phn ng to thnh E v P (sn phm).
v1 = k1[E][S]
v-1 = k-1[ES]
v2 = k2[ES]
Khi h thng t trng thi cn bng ta c:
k-1[ES]+k2[ES] = k1[E][S]
(k-1+k2)[ES] = k+1[E][S] (2)
Gi E0 l nng ban u:
[E0]=[E]+[ES]=>[E]=[E0]-[ES] (3)
Thay tr s [E] t (3) vo (2) ta c:
(k-1+k2)[ES] = k1([E0]-[ES]) [S]
k1 [E0] [S]
[ES] = -------------k-1+ k2+k1[S]
Nu t Km= k-1+k2/ k1
(Km: gi l hng s Michaelis Menten).
Ta c : [ES] = [E0][S]/ Km+[S]
Mt khc vn tc phn ng to thnh sn phm P l:
V = k2[ES]
Thay [ES] bng gi tr trn ta thu c:
k2[E0] [S]
v = ----------------- (4)
Km + [S]

107
Qua y ta thy nng enzyme cng cao th vn tc phn ng
enzyme cng ln. Vn tc t cc i khi ton b enzyme lin kt vi c
cht, ngha l:
Vmax= k2[E0]
Thay vo phng trnh (4) ta c:
[S]
v = Vmax
---------(5)
Km+ [S]
Phng trnh (5) gi l phng trnh Michaelis Menten.
Km gi l hng s Michaelis Menten c trng cho mi enzyme
Km c trng cho i lc ca enzyme vi c cht, Km c tr s
cng nh th i lc ca enzyme vi c cht cng ln, ngha l vn tc ca
phn ng do enzyme xc tc cng ln.

[S]
Hnh 6.2. Bin thin vn tc phn ng theo nng c cht.

Khi tng [S] th v phn ng tng, tng [S] n mt gi tr no th v

t n gi tr vmax v s khng tng na nu ta vn tip tc tng [S].
Khi Km=[S] th v =1/2 Vmax
Nm 1934. Lineweaver v Burk, trn c s ca phng trnh (5)
nghch o bin thnh dng ng thng y=ax+b, n c ngha ln i
vi vic nghin cu km hm enzyme.

108

1/v

1/Vmax
-1/Km

1/[S]

Hnh 6.3: S ph thuc ca tc phn ng vo nng c cht theo

Lineweaver-Burk

6.5.3. nh hng ca cht km hm (inhibitor)

L cht c tc dng lm gim hot hay lm enzyme khng cn
kh nng xc tc bin c cht thnh sn phm. Km hm enzyme c th
thc hin bng nhiu cch khc nhau (thun nghch hay khng thun
nghch). Thun nghch c:
Cch 1: Km hm cnh tranh (competitive inhibition)
Trong trng hp km hm cnh tranh l c cht v cht km hm
u tc dung ln trung tm hot ng ca enzyme, Cht km hm chon
ch ca c cht enzyme.

Hnh 6.4. Kiu km hm cnh tranh

Khi c cht d tha, nng cht km hm thp th c th loi

b tc dng ca cht km hm, cn nng c cht thp v nng cht
km hm cao th li c tc dng km hm hon ton.

1/v= (Km/Vmax) 1/S +1/Vmax

= 1+[I]/KI

109
1/v

[I]

1/Vmax

Khng c cht km hm

1/[S]
Hnh 6. 5. S ph thuc ca tc phn ng vo nng c cht theo
Lineweaver-Burk khi c km hm cnh tranh

Ngi ta thy km hm nh vy phn ln xy ra gia cht km

hm v c cht c s tng ng v mt ho hc. v d: malic acid c cu
trc gn ging vi succinic acid nn km hm cnh tranh enzyme
succinatedehydrogenase, l enzyme xc tc cho s bin i succinic acid
thnh acid fumaric acid.
Trng hp c bit ca km hm cnh tranh l km hm bng sn
phm. Trng hp ny xy ra khi mt sn phm phn ng tc dng tr li
enzyme v chon v tr hot ng phn t enzyme.
ng thng c cht km hm th c xin ln hn v ct trc
tung mt im l 1/Vmax
Cch 2: Km hm phi cnh tranh (uncompetitive inhibition)
c trng ca kiu km hm ny l cht km hm ch lin kt vi
phc hp ES, m khng lin kt vi enzyme t do.

110

Hnh 6.6. Kiu km hm phi cnh tranh

1/v=(Km/Vmax)1/[S] + /Vmax
1/v
[I]

-1/Km

khng c cht km hm

1/[S]
Hnh 6.7. S ph thuc ca tc phn ng vo nng c cht theo
Lineweaver-Burk khi c km hm phi cnh tranh

111

Cch 3: Km hm hn tp( mixed inhibition )

Hnh 6.8. Kiu km hm hn tp

Trong , cht km hm khng nhng lin kt vi enzmye t do

m cn lin kt vi c phc hp ES to thnh phc hp EIS khng to
c sn phm P.
Tng t nh trn ta c phng trnh :

1/v= (Km/Vmax)1/[S] +/vmax

1/v
[I]

1/Vmax

khng c cht km hm

1/[S]
Hnh 6.9. S ph thuc ca tc phn ng vo nng c cht theo
Lineweaver-Burk khi c km hm hn tp

112
Cc gi tr , c nh ngha nh trn. Trng hp = gi
l km hm khng cnh tranh (noncompetitive inhibition).
Mt trng hp km hm cn gp na l km hm enzyme bng
nng cao ca c cht gi l km hm c cht nh km hm urease khi
nng ure cao, ngoi ra cn c cc enzyme khc nh
lactatdehydrogenase,
carboxypeptidase,
lipase,
pyrophotphatase,
photphofructokinase (i vi ATP). Nguyn nhn ca nhng hin tng
ny cn cha c bit r. c th l:
+ Tn ti nhiu trung tm lin kt vi c cht bng cc i lc khc
nhau. Khi nng c cht thp th enzyme c th ch lin kt vi mt
phn t c cht, cn khi nng c cht cao n lin kt vi nhiu c
cht dn n hnh thnh phc hp ES khng hot ng.
+ C cht cng c th c lin kt nh nhng v tr c bit ca
enzyme. l mt nhm enzyme quan trng (enzyme d lp th) bn cnh
trung tm xc tc cn c trung tm iu chnh.
+ C cht c th lin kt vi mt cht hot ho v bng cch ny
n tch khi E.
+ C cht c th chon ch (ngn cn) mt cofactor ( ng yu t )
hay mt coenzyme.
+ C cht c th nh hng n ion lc ca mi trng v qua
lm mt i tnh chuyn ho ca enzyme.
6.5.4. nh hng ca cht hot ha (activator)
L cht lm tng kh nng xc tc nhm chuyn ha c cht thnh
sn phm. Thng thng l nhng cation kim loi hay nhng hp cht hu
c nh cc vitamin tan trong nc.
V d: Mg++ hot ha cc enzyme m c cht c phosphoryl
ha
nh
pyrophosphatase
(c
cht
l
pyrophosphate),
adenosinetriphosphatase (c cht l ATP). Cc cation kim loi c th c
tnh c hiu, tnh i khng v tc dng cn tu thuc vo nng .
6.5.5. nh hng cu nhit
Ta c th tng vn tc ca mt phn ng ha hc bng cch tng
nhit mi trng, hin tng ny tun theo quy lut Vant-Hoff. iu
ny c ngha khi tng nhit ln 100C th tc phn ng tng ln
khang 2 ln.
i vi phn ng do enzyme xc tc cng c th p dng c
quy lut ny nhng ch trong mt phm vi nht nh,v bn cht enzyme l
protein.Khi ta tng nhit ln trn 40-500C xy ra qu trnh ph hu cht

113
xc tc. Sau nhit ti u tc phn ng do enzyme xc tc s gim.
Nh tn ti nhit ti u ngi ta phn bit phn ng ho sinh vi cc
phn ng v c thng thng.
Mi enzyme c mt nhit ti u khc nhau, phn ln ph thuc
ngun cung cp enzyme, thng thng trong khong t 40-600C , cng
c enzyme c nhit ti u rt cao nh cc enzyme ca nhng chng a
nhit. Cc chng vi sinh vt a nhit, c bit cc vi khun chu nhit c
cha enzyme chu nhit cao.
Hat

nhit
Hnh 6.10. nh hng ca nhit ln hat enzyme

6.5.6. nh hng ca pH
S phn li khc nhau ca mt phn t protein cc gi tr pH khc
nhau lm thay i tnh cht ca trung tm lin kt vi c cht v tnh cht
hot ng ca phn t enzyme.iu ny dn n gi tr xc tc khc nhau
ph thuc vo gi tr pH. Nh bit mi enzyme c mt pH ti u,mi
enzyme c ng biu din nh hng pH ln vn tc phn ng do chng
xc tc. ng biu din c dng nh hnh sau:
Hat

Hnh 6.10. nh hng ca pH ln hat enzyme

114
nh hng ca gi tr pH n tc dng enzyme c th do cc c s sau:
a/ Enzyme c s thay i khng thun nghch phm vi pH cc hp.
b/ hai sn ca pH ti u c th xy ra s phn ly nhm
prosthetic hay coenzyme.
c/ Lm thay i mc ion ho hay phn ly c cht.
d/ Lm thay i mc ion ho nhm chc nht nh trn phn t
enzyme dn n lm thay i i lc lin kt ca enzyme vi c cht v
thay i hot tnh cc i.
Nh xc nh Vmax v Km ph thuc gi tr pH cho php nhn
nh li bn cht ca cc nhm tham gia vo lin kt c cht v qu trnh
t xc tc.
6.5.7. Cc yu t khc
+ nh sng: C nh hng khc nhau n tng loi enzyme, cc
bc sng khc nhau c nh hng khc nhau, thng nh sng trng c
tc ng mnh nht, nh sng c tc ng yu nht.
nh sng vng t ngoi cng c th gy nn nhng bt li,
enzyme trng thi dung dch bn hn khi c kt tinh dng tinh th,
nng enzyme trong dung dch cng thp th cng km bn, tc ng ca
tia t ngoi s tng ln khi nhit cao. V d di tc ng ca tia t
ngoi nhit cao enzyme amylase nhanh chng mt hot tnh.
+ S chiu in: in chiu vi cng cng cao th tc ng
ph hu cng mnh. Tc ng s mnh hn i vi dch enzyme c nng
thp. C th do to thnh nhng gc t do, t tn cng vo phn
ng enzyme.
+ Sng siu m: Tc ng rt khc nhau i vi tng loai enzyme,
c enzyme b mt hot tnh, c enzyme li khng chu nh hng.
Nhn xt chung: bn ph thuc vo trang thi tn ti ca
enzyme, cng tinh khit th enzyme cng km bn, dch cng long th
bn cng km, tc ng ca mt s ion kim loi trong dch vi nng
khong 10-3M nh Ca++ lm tng tnh bn.
Enzyme allosteric ( Enzyme d lp th, d khng gian)
Cho n nay, ngi ta m t enzyme m hat tnh enzyme ph
thuc nng c cht khng c dng hyperbol m c dng sigmoid l
enzyme allosteric (Hnh 6.11 ):

115

Hnh 6.11. Bin thin vn tc phn ng theo nng c cht

i vi enzyme ny, khi nng c cht thp th tc phn ng

tng chm, sau tip tc tng nng th tc nhanh chng t gi tr
cc i.
Nh ta bit, enzyme tun theo ng hc Michaelis-Menten th 1
hay nhiu c cht cng ch lin kt vo 1 v tr trn phn t enzyme, iu
ny s dn n enzyme bo ha c cht. Cn enzyme c ng cong tc
sigmoid ch xut hin khi enzyme l mt oligomer, nn c th lin kt
vi nhiu phn t c cht. iu ny c ngha trn enzyme allosteric c
nhiu trung tm lin kt, mi monomer c 1 trung tm lin kt.
Ngi ta cho rng, trong trng hp ny c tnh hp tc gia cc
v tr lin kt c cht trong phn t enzyme oligomer.
Cc enzyme oligomer ny c Monod gi l allosteric enzyme d
lp th (allosteric enzyme). ng cong tc sigmoid c th b cht iu
ha (modulator) y v pha tri hay phi. Cht iu ha dng tc lm
tng i lc ca enzyme allosteric vi c cht, ngc li l cht iu ha
m. Cc cht iu ha c th lm nh hng khc nhau n cc thng s
ng hc, lm thay i gi tr ring l mt trong hai gi tr Km hay Vmax.

Hnh 6.12 Minh ha khi c modulator

116

6.6. Cch gi tn v phn loi enzyme

Nh ta bit mi enzyme xc tc cho mi kiu phn ng ho hc
duy nht (nh oxy ho mt kiu c cht nht nh, thu phn mt kiu lin
kt nht nh,vn chuyn mt nhm cht nht nh t mt cht cho n
mt cht nhn c a ch, trong c c vic bin i ch mt c cht duy
nht), mt khc cn c mt kiu phn ng ho sinh nht nh c th c
xc tc bng cc enzyme khc nhau.
Da vo tnh c hiu phn ng ca enzyme, nm 1961 tiu ban
enzyme hc quc t trnh by mt bo co, trong c ngh nhng
nguyn tc nh tn v phn loi enzyme. Ngi ta chia enzyme ra lm 6 lp:
1. Oxydoreductase: cc enzyme xc tc cho cc phn ng oxi ho-kh.
2. Transferase: cc enzyme xc tc cho cc phn ng chuyn v.
3. Hydrolase: cc enzyme xc tc cho cc phn ng thy phn.
4. Lyase: cc enzyme xc tc cho cc phn ng phn ct khng cn nc.
5. Isomerase: cc enzyme xc tc cho cc phn ng ng phn ho.
6. Ligase (synthetase): cc enzyme xc tc cho cc phn ng tng
hp c s dng lin kt giu nng lng ca ATP .v.v.
Mi lp chia thnh nhiu t (di lp), mi t chia thnh nhiu
nhm (siu lp).
Tn enzyme thng c gi: Tn c cht c hiu - loi phn ng
xc tc cng thm tip v ng ase.
ng trc tn enzyme thng c 4 con s: s th nht ch lp, s th
hai ch t, s th ba ch nhm, s th t ch s hng enzyme trong nhm.
V d: (2.6.1.1) L.aspartate: -cetoglutarate aminotransferase.
Enzyme ny xc tc cho phn ng chuyn nhm amine t
L.aspartate n -cetoglutarate.
L.aspartate +-cetoglutarate ' oxaloacetate + glutamate

6.7. Cc coenzyme quan trng

Nicotinamide adenine dinucleotide

117

Flavine adenine dinucleotide

Lipoic acid

Coenzyme A

Biotin

118

Thiamin diphosphate

(Nhm - amino protein, Schiff base)

Pyridoxal phosphate

Tetrahydrofolic acid

119

TI LIU THAM KHO

Ti liu ting Vit
1. Phm Th Trn Chu, Trn thi ng. 1999. Ho sinh hc, NXB Gio dc,
H Ni.
2. Qu Hai. 2004. Gio trnh Ha sinh i cng, Ti liu lu hnh ni
b Trng HKH Hu.
3. Trn Thanh Phong. 2004. Gio trnh Ha sinh i cng, Ti liu lu
hnh ni b Trng HKH Hu.
4. L Ngc T (ch bin), L Vn Ch, ng Th Thu, Phm Quc Thng
Nguyn Th Thnh, Bi c Hi, Lu Dun, L Don Din, 2000. Ha sinh
Cng nghip, Nxb KH&KT, H Ni.

Ti liu ting Anh

1. Bergmeyer H. U. 1968. Methods of enzymatic analysis, translated from
the third German edition, Acanamic, New York.
2. Copeland R. A. 2000. Enzymes, copyright by Wiley-VCH, Inc.
3. Lehninger A. L. 2004. Principles of Biochemistry, 4th Edition. W.H
Freeman.
4. Mikkelsen S. R. 2004. Bioanalytical chemistry, copyright by John
Wiley & Sons, Inc.

120

Chng 7

Hormone
7.1. C ch tc dng ca hormone
Hormone l nhng cht hu c c to thnh trong c th c tc
dng iu ho cc hot ng sng trong c th. Lng hormone trong c
th thng rt thp.
Hormone c c thc vt v ng vt. ng vt hormone c
sn xut ti cc tuyn ni tit v tc ng n cc m khc ni n c
to ra. Hormone t tuyn ni tit c tit trc tip vo mu v c mu
vn chuyn n cc m chu tc dng.
Hormone c tnh c hiu. Hormone c tc dng iu ho cc qu
trnh sinh l, ho sinh trong c th m khng tham gia trc tip vo cc
phn ng ca c th. Hormone c tc ng n tc sinh tng hp
protein, enzyme, nh hng n tc xc tc ca enzyme; thay i tnh
thm ca mng t bo, qua iu ho hot ng sng xy ra trong t bo.
Mt s hormone tc ng n c th thng qua cht trung gian
AMP vng. AMP vng l cht truyn tin th 2, cn hormone l cht
truyn tin th nht. Theo c ch ny tc dng ca hormone ln t bo ch
xy ra qua nhiu giai on kh phc tp.
- Trong mng nguyn sinh cht ca t bo c cha cht nhn
hormone, cht ny s kt hp c hiu vi hormone.
- S kt hp kch thch lm tng hot ca adenylatcyclase xc
tc cho phn ng chuyn ho ATP thnh AMP vng.
- Adenylatcyclase xc tc cho phn ng chuyn ho ATP thnh
AMP vng.
- AMP vng lm thay i vn tc ca cc qu trnh xy ra trong t
bo lin quan n hot ng ca hormone.
- Nh vy tc dng ca hormone theo c ch ny phi thng qua
AMP vng m khng tc ng trc tip vo t bo.
- Qu trnh hot ho adenylatcyclase bi phc hormone-cht nhn
c thc hin qua cht trung gian l protein G. Phn t protein ny c
kh nng kt hp vi GDP hay GTP. Dng phc protein G-GTP c tc
dng hot ho adenylatcyclase, cn protein G-GDP khng c tc dng
ny. Nh vy mun chuyn sang dng hot ng phi c s tham gia ca
GTP, nu l protein G-GDP cn c s thay th GDP bng GTP nh phc

121
hormone-cht nhn xc tc. Dng thng tin c truyn t cht nhn
hormone n protein G ri n adenylatcyclase.
- Protein G khng ch c vai tr trung gian mang thng tin t cht
nhn hormone n adenylatcyclase m cn c hot tnh ca GTPase, l
kh nng thu phn GTP. Nh kh nng nn n xc tc cho qu trnh
chuyn phc proteinG-GTP hot ng thnh dng proteinG-GDP khng
hot ng do thu phn GTP trong phc proteinGTP thnh GDP to nn
phc proteinG-GDP. Bng c ch protein G c vai tr quan trng trong
qu trnh hot ho hay phn hot ho adenylatcyclase. Khi lng
hormone gim adenylatcyclase tr thnh dng khng hot ng.
- Nhiu hormone c c ch tc ng thng qua vai tr trung gian ca
AMP vng. Nh vy AMP vng tham gia vo nhiu qu trnh khc nhau
trong c ch tc ng ca hormone. l do AMP vng c tc dng hot
ho proteinkinase l enzyme xc tc qu trnh photphoryl ho nhiu loi
protein khc nhau. Thng cc protein enzyme dng phosphoryl ho l
dng c hot tnh sinh hc.
- Cc hormone tc dng theo c ch qua AMP vng, tn hiu c
khuych i ln nhiu ln, do vy nng cc hormone trong mu rt
thp, ch khong 10-10M, nhng ch cn hot ho c mt phn t
adenylatcyclase c th to ra c nhiu phn t AMP vng nn nng
AMP vng trong t bo ch cao hn nhiu lng hormone trong mu.
Tc dng hot ho proteinkinase nh AMP vng li lm cho tn hiu c
khuych tn tip tc v nhiu phn t protein c hot ho nh
proteinkinase. iu gii thch c ti sao nng hormone trong mu
rt thp m tc dng kch thch ca n li rt mnh.
- Mt c ch tc ng th hai ca hormone l khng qua AMP
vng.Insulin l hormone tc ng n t bo ch khng qua bc trung
gian l lm tng lng AMP vng. Insulin lin kt cht ch vi cht nhn
c hiu ca n trn mng nguyn sinh cht ca t bo ch. Tng tc
gia Insulin v cht nhn bo m cho tc ng ca Insulin c th hin
nhanh chng. Insulin cn c tc dng phosphoryl ho protein tham gia vo
c ch kch thch qu trnh trao i glycogen.
- C ch tc dng ca cc hormone thc vt hon ton khc
hormone ng vt. Cc hormone thc vt tc ng ln hot tnh cc
enzyme bng cch lin kt vi enzyme to phc hot ng. Khi lin kt
vi hormone hot tnh ca enzyme c tng ln.
- Hormone thc vt cn lm thay i tnh cht ca mng
cellulose, mng nguyn sinh qua tc ng kch thch qu trnh
sinh trng ca t bo.

122
- Mt c ch tc ng quan trng na ca hormone thc vt l thay
i tnh cht ca nguyn sinh cht ca t bo, t nh hng n cc
hot ng sinh l, trao i cht ca t bo

7.2. Cc hormone quan trng

7.2.1. Hormone ng vt
- Hormone ng vt c nhiu loi vi cu to v chc nng rt khc
nhau. Da vo cu to ho hc c th chia hormone ng vt thnh 3
nhm:
- Hormone steroid l dn xut ca cholesterol.
- Hormone l dn xut ca amino acid.
- Hormone l peptide hay protein.
7.2.1.1. Hormone l steroid
- y l nhm hormone c s lng ln, c vai tr quan trng v a
dng. Ngi ta chia steroid thnh 5 nhm nh vi nhiu loi khc nhau:
T
T
1

Nhm
Progestagen

Glucocorticoid

i din

Ni to
thnh

Progesterol

-Th vng
-V thng
thn

Cortisol

3
4

V thng
thn

Vai tr
Hormone dng thai
gip trng pht trin
- Kch thch tng
glycogen v tch
glycogen gan.
- Kch thch phn
protein, lipid.
- Chng vim,
nc mui.

hp
lu
gii
tch

Mineral corticoid

Andosterol

V thng
thn

- Tng hp th Na+, Cl- Tng tch nc.

- Bi tit K+

Androgen

Testosterol

Tinh hon

Pht trin cc c im
ca nam gii.

Estrogen

Estron

- Pht trin cc c
im n gii.
Bung trng
- Pht trin nim mc
d con.

123
7.2.1.2. Hormone l dn xut amino acid
n nay ngi ta bit mt s hormone l dn xut amino acid
nh adrenaline, noradrenaline, thyroxine...
- Adrenaline v noradrenaline l cc hormone do tuyn thng
thn to ra. Cc hormone ny c tc dng kch thch s phn gii
glycogen, lm gim s tng hp glycogen nn lm tng hm lng
glucose trong mu.
.

OH

CHOH - CH2 - NH - CH3

HO

Adrenaline
OH
CHOH - CH2 - NH2

HO

Noradrenaline
Thiroxine l hormone do tuyn gip sn xut c tc dng tng cng
qu trnh trao i cht, gip cho c th pht trin bnh thng. Nu thiu
thyroxine gy nn trng thi thiu nng tuyn gip lm cho c th ln,
km pht trin, n n. Ngc li nu tha thyroxine cng gy bnh l
u nng tuyn gip lm cho ngi cao qu kh, khng cn i.
I
I

HO

CH2 - CH

NH2
COOH

O
I
Thyroxine (Tetraiodothyronine)

124
y l nhm hormone c vai tr quan trng trong qu trnh iu
ho trao i cht trong c th, c bit l iu ho lng ng trong
mu.
Mt s hormone l peptide:
STT

Hormone

Ni to ra

Vai tr

Tyrocalcitonin

Tuyn gip Gim hm lng Ca++ trong mu

Insulin

Tuyn ty

Gim lng ng trong mu

Glucagon

Tuyn ty

Tng lng ng trong mu

Oxytoxin (HGF)

Tuyn yn

Gy co d con, kch thch

Vasopressin (ADH)

Tuyn yn

Tng p, chng bi tit

Melanotropin (MSH)

Tuyn yn

Kch thch tng sc t da

Somatotropin (STH)

Tuyn yn

Kch thch tng trng, tng TC

Corticotropin (ACTH)

Tuyn yn

Kch thch tuyn trn thn

Thyreotropin (TSH)

Tuyn yn

Kch thch tuyn gip

10

Kch nang t (FSH)

Tuyn yn

Kch thch to estradiol

Sau y s cp n mt s hormone trong nhm ny:

- Insulin: Insulin c tit t t bo beta ca o Langhan ca
tuyn ty khi lng ng trong mu cao. Insulin kch thch cc qu trnh
tng hp, km hm cc qu trnh phn gii glycogen gan, m m. Insulin
cn kch thch s phn gii glucose. Nh insulin lm gim lng ng
trong mu, do chng li bnh i tho ng.
Insulin c khi lng phn t l 5800. Cu to insulin gm 2
chui polypeptide: chui A c 21 amino acid, chui B c 30 amino acid.
Hai chui lin kt vi nhau bng 2 lin kt disunfit.
Tin cht ca insulin l proinsulin v preproinsulin.T
preproinsulin bin i thnh proinsulin, sau insulin c to nn t
proinsulin.
- Glucagon l hormone peptide, c tc dng ngc vi insulin. Khi
lng ng trong mu gim qa mc cho php th tuyn tu sn sinh ra
glucagon c tc dng lm tng lng ng trong mu nh km hm qu
trnh tng hp glycogen.
Glucagon c khi lng phn t 3.500, bao gm 29 gc amino
acid to chui polypeptide mch thng.

125
7.2.2. Hormone thc vt
Hormone thc vt l cc cht c vai tr quan trng trong qu trnh
sinh trng, pht trin ca thc vt. C nhiu loi hormone khc nhau
trong c th thc vt. Cc loi hormone ny khc nhau v bn cht ho
hc, v vai tr i vi thc vt. C th chia hormone thc vt thnh 5
nhm:
- Auxin.
- Gibberellin.
- Cytokinin.
- Absisic acid.
- Ethylen.
7.2.2.1. Auxin
Auxin l nhm hormone quan trng, ph bin nht thc vt. C
nhiu loi auxin khc nhau vi cu trc ho hc khc nhau. Loi auxin
quan trng nht l -indol-acetic acid (IAA), ngoi ra mt s auxin khc
cng kh ph bin l napthalen-acetic acid (NAA), phenyl-acetic acid
(PAA) ...
CH2-COOH

CH2-COOH

CH2-COOH
N
H

IAA

NAA

PAA

Auxin c vai tr nhiu mt i vi thc vt:

- Kch thch s sinh trng t bo, t kch thch s sinh trng
cc c quan v ton c th.
- C vai tr quyt nh hin tng u th nh.
- C vai tr quyt nh cc c ng sinh trng nh hng sng,
hng trng lc.
- Kch thch qu trnh ny mm, rt ngn thi k ng ca ht, c.
- c ch s rng l, kch thch s to qu.

126
- Kch thch cc hot ng sinh l, cc qu trnh trao i cht v
nng lng ca c th.
7.2.2.2. Gibberellin
Gibberellin l nhm hormone quan trng th hai thc vt.
Gibberellin c cc nh khoa hc Nht pht hin ln u tin loi nm
gy bnh la von (Gibberellin fujcoroi). C nhiu loi Gibberellin khc
nhau, n nay tm thy hn 70 loi Gibberellin c mt thc vt, vi
sinh vt. Ngi ta t tn cc Gibberellin theo th t thi gian pht hin
GA1. GA2 .... GAn, trong quan trng nht c th k n l GA3. Cc
Gibberellin u l dn xut ca vng gibban.

HO

C=O

CH3

OH
COOH

CH2

Cu to GA3
Gibberellin c vai tr quan trng trong qu trnh sinh trng, pht
trin ca thc vt:
- Kch thch s sinh trng ca t bo, qua kch thch s sinh
trng ca cc c quan v c th.
- Kch thch qu trnh ny mm, ph trng thi ng ca ht, c.
- Kch thch s ra hoa ca cy ngy di.
- Kch thch cc hot ng sinh l, cc qu trnh trao i cht v
nng lng ca c th.
7.2.2.3. Cytokinin
Cytokinin l cc dn xut ca base Adenine. C nhiu loi
cytokinin khc nhau, quan trng nht l kinetin v zeatin.

127

HN - CH2

HN - CH2- CH = CH

CH2OH
CH3

N
N

N
N

N
H

Kinetin

Zeatin

Xitokinin tham gia v nhiu hot ng sng quan trng ca thc vt:
- Kch thch s phn bo qua kch thch s sinh trng ca t bo.
- Lm chm qu trnh ho gi ca t bo, m.
- Gip cho thc vt chng li cc stress ca mi trng c hiu qu.
- L thnh phn cu to ca nucleic acid (trong mt s loi RNA)
nn c vai tr trong qu trnh trao i nucleic acid v protein.
- Kch thch cc hot ng sinh l, cc qu trnh trao i cht v
nng lng ca c th.
7.2.2.4. Absisic acid
Acid absisic (ABA) l nhm cht c ch sinh trng c tc dng
ngc li 3 nhm cht trn. Absisic acid l dn xut ca triterpen.
CH3

CH3

CH3
OH

COOH
O

CH3
ABA

128
Tc dng ch yu ca ABA l c ch qu trnh sinh trng ca t bo,
gy hin tng rng l, rng qu. ABA ko di thi gian ng ca ht, c.
Do c ch s sinh trng ca thc vt nn ABA phi hp vi
nhm cht kch thch sinh trng iu ho qu trnh sinh trng ca
thc vt xy ra cn i.
7.2.2.5. Ethylen
Ethylen (CH2 = CH2) l nhm hormone thc vt c tc dng gn
ging ABA nn thuc nhm cht c ch sinh trng. Etylen thc y qu
trnh chn ca qu, qu trnh rng l.
Khc vi hormone ng vt, hormone thc vt c tng hp
trong cc phn khc nhau ca cy m khng c cc tuyn tit chuyn bit.
Cc hormone thc vt c tng hp cc vng khc nhau ca cy.
Auxin, gibberellin ch yu c tng hp ti cc phn non ca
cy, nht l vng sinh trng nh nh sinh trng, tng tng Sau khi
tng hp Auxin, gibberellin c vn chuyn trong cc m dn hay qua h
thng t bo sng a n cc vng tc dng. Hormone thc vt cng
khng c t bo ch chuyn bit nh ng vt m tc ng ln ton c th.
Cytokinetin c tng hp mnh phn r non, cn absisic acid ,
ethylen l c tng hp nhiu cc phn gi ca cy. Sau khi tng hp
cc hormone ny cng c vn chuyn n cc vng khc nhau trong c
th thc hin cc chc nng ca chng.

129

TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n (ch bin). 1979. Ha sinh i cng (tp I, II). NxB
KH&KT. H Ni.
2. Phm Th Trn Chu, Trn Th ng. 2000. Ha sinh hc. Nxb Gio
dc. H Ni.
3. Nguyn B Lc. 1997. Ha sinh. Nxb Gio dc. H Ni

Ti liu dch
1. Musil J.G., Kurz .K., Novakava .O. 1982
2. Sinh ha hc hin i theo s . Nxb Y hc. H Ni.

Ti liu ting nc ngoi

1. Farkas G. 1984. Nvnyi anyagcserelettan. Akadmiai Kiad Budapest.
2. Lehninger A. L., 2004. Principle of Biochemistry, 4th Edition. W.H
Freeman.

130

Chng 8

Khi nim v s trao i cht v

trao i nng lng
Trao i cht v trao i nng lng l bn cht ca hot ng sng
ca mi c th sinh vt, l biu hin tn ti s sng. S trao i cht ca
c th lun gn lin vi s trao i v chuyn ha nng lng. Chnh v
vy, trao i cht v trao i nng lng l hai mt ca mt qu trnh lin
quan cht ch vi nhau.

8.1. Khi nim chung v s trao i cht

C th sng tn ti, pht trin trong mi trng v khng ngng lin
h mt thit vi mi trng . N hp th cc cht khc nhau t mi
trng ngoi, lm bin i cc cht v mt mt to nn cc yu t cu
to ca bn thn c th sng, mt khc li thi vo mi trng ngoi cc
sn phm phn gii ca chnh c th cng nh cc sn phm hnh thnh
trong qu trnh sng ca c th. Qu trnh thc hin c l do cc bin
i ha hc lin tc xy ra trong c th. Ngi ta gi ton b cc bin i
ha hc l s trao i cht.
S trao i cht bao gm nhiu khu chuyn ha trung gian. Cc
qu trnh ny xy ra phc tp trong tng m, tng t bo bao gm 2 qu
trnh c bn l ng ha (tng hp) v d ha (phn gii) to nn chu k
trao i cht lin tc gia cht nguyn sinh v cht nhn vo.
Qu trnh ng ha l s hp th cc cht mi t mi trng bn
ngoi, bin i chng thnh sinh cht ca mnh; bin i cc cht n
gin thnh cht phc tp hn, s tch ly nng lng cao hn. y l qu
trnh bin i cc cht khng c hiu (cc cht hu c ca thc n nh
glucid, lipid, protein) t cc ngun khc nhau (thc vt, ng vt, vi sinh
vt) thnh cc cht hu c khc (glucid, lipid, protein) c hiu ca c th.
c im ca qu trnh ny l thu nng lng. Nng lng cn thit cung cp
cho cc phn ng tng hp trn ch yu dng lin kt cao nng ca ATP.
Qu trnh d ha l qu trnh ngc li ca qu trnh ng ha, l
s bin i cc cht phc tp thnh cc cht n gin v gii phng nng
lng cn thit cho hot ng sng. Nh vy y l qu trnh phn gii
cc cht d tr, cc cht c trng ca c th thnh cc sn phm phn t
nh khng c trng v cui cng thnh nhng cht thi (CO2, H2O,

131

NH3...) thi ra mi trng. Nng lng c tch tr trong ATP v

c s dng cho nhiu phn ng thu nng lng khc.
Hai qu trnh ng ha v d ha xy ra lin tc lin quan vi nhau
v khng tch ri nhau. Qu trnh ng ha l qu trnh i hi nng
lng cho nn ng thi phi xy ra qu trnh d ha cung cp nng
lng cho qu trnh ng ha. Do s trao i cht v trao i nng
lng l hai mt ca mt vn .
Ty theo kiu trao i cht, ngi ta chia sinh vt ra thnh hai
nhm: nhm sinh vt t dng v nhm sinh vt d dng.
Nhm sinh vt t dng bao gm tt c cc sinh vt t tng hp
cht dinh dng cn thit cho chng. tn ti v pht trin, nhm ny
ch cn H2O, CO2, mui v c v ngun nng lng. C hai hnh thc t
dng. l hnh thc t dng quang hp v hnh thc t dng ha
hp. Hnh thc u th hin cy xanh v vi khun ta, vi khun lu
hunh vn dng quang nng tng hp cht hu c. Hnh thc sau c
th hin mt s vi khun nhn nng lng trong qu trnh oxy ha cc
cht v c.
Nhm sinh vt d dng bao gm cc sinh vt khng c kh nng t
tng hp cht dinh dng t cc cht v c m phi sng nh vo cc cht
dinh dng ca nhm sinh vt t dng tng hp nn.
Nh vy, qu trnh trao i cht ca th gii sinh vt lin quan cht
ch vi nhau, to nn chu k trao i cht chung.
Glucid, lipid,
protein

nh sng

O2

Sinh vt

Sinh vt

t dng

d dng

CO2, H2O, mui

cha Nitrogen
Ngoi cch chia trn, cng theo kiu trao i cht, ngi ta chia sinh
vt thnh hai nhm ln: nhm hiu kh (aerob) v nhm k kh (anaerob).

132

Nhm hiu kh l kiu trao i cht m cc qu trnh oxy ha c s tham

gia ca oxy kh quyn. Nhm k kh l kiu trao i cht m cc qu trnh
oxy ha khng c s tham gia ca oxy kh quyn.
a s cc sinh vt thuc nhm hiu kh. Nhm k kh ch l mt
phn nh ca nhm sinh vt d dng bc thp. Tuy vy, gia cc c th
hiu kh v k kh khng c ranh gii r rng. Sinh vt hiu kh biu hin
r rng nht nh ngi chng hn cng c thc hin mt phn cc qu
trnh trao i cht theo con ng k kh (v d nh m c)
Qu trnh chuyn ha trong c th sng mang tnh thng nht v
ring bit. Cc con ng chuyn ha ln trong mi c th ng vt, thc
vt n bo, a bo u theo nhng giai on tng t nhau. Tuy vy, nu i
su vo tng m, c quan, c th tng loi th li c nhng nt ring bit.
Cc phn ng ha hc trong c th xy ra lin tc pH trung tnh,
370C, di tc dng xc tc ca enzyme.
ng vt, cc qu trnh chuyn ha c iu khin bi h thng
thn kinh

8.2. Khi nim chung v trao i nng lng v nng lng sinh hc
Trao i cht lun gn lin vi trao i nng lng. i vi c th
ngi, ng vt v phn ln vi sinh vt th ngun nng lng duy nht l nng
lng ha hc ca cc cht trong thc n. Trong c th, cc cht dinh dng
ch yu v quan trng l glucid, lipid v protein u b oxy ha. Lipid v glucid
i vo c th b t chy s sinh ra CO2, H2O v NH3, cht ny tc dng vi
CO2 chuyn thnh carbamid (ure).
Cc qu trnh oxy ha kh sinh hc thuc cc phn ng d ha c ngha
rt quan trng. Chng khng nhng ch l ngun nng lng quan trng dng
thc hin cc phn ng tng hp khc nhau m cn l ngun cung cp cc
hp cht trung gian dng lm nguyn liu cho cc phn ng tng hp v ng
vai tr ht sc quan trng trong vic lin hp cc qu trnh trao i cht.
tn ti v pht trin, c th cn phi c cung cp lin tc nng
lng. Trong hot ng sng ca mnh, c th bin i nng lng t dng ny
sang dng khc v s bin i nng lng trong c th sng cng tun theo cc
quy lut vt l nh s bin i nng lng gii v c.

133

So snh v nng lng sinh hc v nng lng k thut ta thy c nhng

c im sau: th nht, c th khng s dng nhit nng thnh cng c ch
c; th hai, s gii phng nng lng trong c th l dn dn, tng bc; th
ba, s gii phng nng lng i km theo s phosphoryl ha ngha l nng
lng gii phng c c nh li lin kt este vi phosphoric acid trong
phn t ATP vn c gi l lin kt cao nng. T dng nng lng trung gian
ny (ATP) m c th d tr v s dng nng lng vo cc hot ng sng; th
t, c th khng s dng c nng lng t do ca tt c cc loi phn ng pht
nhit m ngun nng lng duy nht c th s dng l ca cc qu trnh oxy ha.
8.2.1. S bin i nng lng t do
S thay i v i lng ca nng lng t do l mt ch tiu quan trng
nht ca hiu ng nng lng tc l h s ca tc dng hu hiu ca phn ng.
C th nh ngha nng lng t do l lng nng lng m mt nhit
nht nh no c th bin thnh cng.
T bo c th to ra v duy tr c cu trc trt t v phc tp ca mnh
nh chng lin tc tip nhn nng lng t do t mi trng dng quang
nng hoc ha nng v bin ha n thnh cc dng nng lng sinh hc
phc v cho cc qu trnh hot ng sng. S bin ha, tch ly v s dng
nng lng sinh hc xy ra song song vi s chuyn ha vt cht v tun th
cc nguyn tc ca nhit ng hc.
Nhng bin i nng lng t do ca h thng phn ng c k hiu
bng UG c gi tr l Kcal/mol. i lng ca UG l hiu s gia lng nng
lng t do ca trng thi cui (sau phn ng) G2 v nng lng t do ca
trng thi u (trc phn ng) G1. Nu UG<0 (c gi tr m), phn ng ta
nhit, c th xy ra mt cch t pht. V d cc phn ng thy phn u thuc
loi phn ng ny. Nu UG = 0, h thng trng thi cn bng. Nu UG>0
(c gi tr dng), phn ng thu nhit, mun thc hin phn ng cn phi cung
cp nng lng. Cc phn ng thu nhit ch c th c thc hin cng vi cc
phn ng ta nhit, ngha l vic tng nng lng t do ch c th c c do
cc phn ng lin hp khc tin hnh vi vic gim nng lng t do. Cc qu
trnh c bn gn lin vi hot ng sng ca c th, nhiu kiu lm vic ca t
bo, cc phn ng tng hp u l nhng phn ng thu nhit lun lun lin hp
vi cc phn ng ta nhit.
UG c tnh theo cng thc:

134

UG = UG 0 + RT lnK
trong UG0 l s bin i nng lng t do tiu chun ca phn ng
250C khi nng ca tt c cc cht phn ng l 1 mol v p sut l 101,3 KPa
(1atm), R l hng s kh, T l nhit tuyt i, K l hng s cn bng ca
phn ng bng [C]c. [D]d /[A]a[B]b tc l nng ca cc cht tham gia phn ng
A + B ' C + D; a, b, c, d l s lng phn t A, B, C, D tham gia phn ng.
Trong h thng sinh hc, khi tnh gi tr UG0 cn ch n pH, nng
H+ l 1 mol, pH=0. Trng thi ion ha ca nhiu hp cht sinh hc b bin i
khi pH thay i. V vy, thun tin cho vic tnh ton, xem trng thi chun
ca pH l 7 v k hiu s thay i nng lng t do chun pH 7,0 l UG0'.
8.2.2. Lin kt cao nng v vai tr ca ATP
Cc lin kt ha hc gia cc nguyn t u l nhng tc nhn mang ch
yu ca nng lng t do trong cc cht hu c. V vy, trong vic bin to ca
cc lin kt ha hc trong phn t, mc nng lng t do ca hp cht s thay
i. Xt v mt nng lng trong cc hp cht hu c c hai loi lin kt: Lin
kt thng v lin kt cao nng (lin kt giu nng lng). Lin kt thng l
lin kt m khi phn gii hoc to thnh n c s bin i nng lng vo
khong 3 Kcal trn mt phn t gam (V d nh lin kt este); cn i vi lin
kt cao nng s bin i ny ln hn nhiu t 7 12 kcal/mol. Trong cc hot
ng sng ca c th sinh vt, cc qu trnh tng hp cc cht phn t ln t
cc cht n gin, vn chuyn tch cc cc cht qua mng t bo, qu trnh vn
ng v.v. lun i hi nng lng t do. Trong h thng sng cn c cc cht,
cc h thng nhn nng lng t do t cc qu trnh ny chuyn n cho cc
qu trnh khc. ATP l cht ph bin gi vai tr ny, l cht c vai tr trung
tm trong trao i nng lng t bo v c th sng, l cht lin kt hoc c
th ni l mt xch gia h thng s dng nng lng v h thng sn sinh ra
nng lng.
Trong phn t ATP c 3 gc phosphate, 1 gc kt hp vi gc ribose qua
lin kt este, 2 lin kt gia 3 gc phosphate l lin kt anhydric. l cc lin
kt cao nng c k hiu bng du ~ . ATP ( Adenosine Tri Phosphate)
c biu th mt cch khi qut nh sau: Adenosine - P ~ P ~ P (trong
P l cc gc phosphoric acid ). Khi ct t cc lin kt cao nng ny, s
gii phng s nng lng ln gp hn 2 ln so vi lin kt este:
ATP + H2O ' ADP + H3PO4
UG0 = -7 Kcal/mol
( P )

135

UG0 = - 8,5 Kcal/mol

ATP + H2O ' AMP + H4P2O7

( P ~ P )

Nu tip tc thy phn lin kt este ca AMP to thnh adenosine v

phosphate v c, nng lng t do c gii phng ca phn ng ny thp hn nhiu.
S chuyn ha tng h gia ATP v ADP c vai tr c bit quan trng
trong qu trnh trao i nng lng ca h thng sng.
Trong a s trng hp thng thy phosphore hoc sulphure tham gia
to thnh lin kt cao nng (Bng 8.1).
Bng 8.1. Mt s dng lin kt cao nng thng gp

Dng lin kt

C trong cc cht

- Anhydrid phosphate
(pyro phosphate)

ATP, GTP.......

-7

~O ~P

ADP, GDP.....

-7

- Acyl phosphate
O~

UG0 ( Kcal/mol)

Kiu lin kt

O
RCO~ P

- 10
1,3Diphosphoglyceric
acid
Aminoacyl-AMP

-7

- Enol phosphate

Phosphoenol

- 12,8

RCO~ P

Pyruvic acid

CH2
N~ P

- Amid phosphate

Creatin phosphate

( phosphoguanidin)

Arginin phosphate

- 10,5

R C NH ~ P
NH
- Thioeste
C~S

Acetyl coenzyme A

Acyl coenzyme A

RC~SR

- 8,8

136

8.3. Qu trnh oxy ha kh sinh hc

C th nh ngha qu trnh oxy ha kh l qu trnh trao i in t. S
oxy ha l s tch mt hay nhiu in t, ngc li s kh oxy l s thu in
t. Tt c cc cht tham gia vo qu trnh oxy ha kh c th sng u c kh
nng nhng hoc thu in t.
chnh l kh nng oxy ha kh. Song song vi s oxy ha c s kh
oxy v in t c chuyn t cht b oxy ha sang cht b kh:
- 2e

V d: 2Fe

2+

+ Cl2 J 2Fe 3+ + 2Cl-

i lng c trng cho kh nng oxy ha kh ca mi cht gi l th

nng oxy ha kh. C th tnh c th nng oxy ha kh theo cng thc sau:
E'n= E'o +

RT
nF

ln

[dng oxy ha]

(1)

[ dng kh]

Trong : En l th nng oxy ha kh ca mt cht nht nh trong

nhng iu kin nht nh. E0 l th nng oxy ha kh cc iu kin tiu
chun ( nng ca hai dng bng nhau)
R l hng s kh, T l nhit tuyt i, F l tr s Faraday
Bng 8.2 trnh by E0, hiu in th oxy ha kh UE0 v nng lng t
do UGo ca mi h.
Th nng oxy ha kh cn dng tnh nng lng t do (Go) c
gii phng ra trong qa trnh oxy ha kh theo phng trnh:
Go = -nF.E'o (2)
(Cc k hiu c gii thch cng thc tnh th nng oxy ha kh
v lin quan n bng 8.2 trn)
* Tin trnh ca s oxy ha sinh hc:
S phn gii cht dinh dng v gii phng nng lng ca t bo (s d
ha) c th c chia thnh 3 giai on c bn:
giai on u: cc hp cht cao phn t b thy phn thnh cc cht
n gin c phn t nh hn: cc glucid (tinh bt, glucogen v.v...) thnh cc

137

monosaccharid (glucose), cc protein thnh cc amino acid, cc lipid thnh cc

acid bo.
giai on th hai: bin nhng cht n gin thnh nhng cht 2 carbon
l acetyl CoA (CH3 - COSCoA) (thiu). Acetyl CoA c coi l sn phm
thoi ha ca cc cht glucid, lipid v protein. N c hnh thnh do s -oxy
ha acid bo, do s oxy ha ca khong mt na s -amino acid cng nh do
s oxy ha hiu kh glucose.
Bng 8.2. Th nng oxy ha tiu chun ca mt s h thng

H thng oxy ha kh

Eo (volt)
pH7, 30oC

UE0
(volt)

UGo
(kcal/pH7,
30oC)

in cc hydro 2H+/ H2

-0,42

NAD+/ NADH + H+

-0,32

FAD/ FADH2

-0,10

+0,22

-10,1

Cytochrome b Fe3+/ Fe2+

+0,04

+0,14

-6,4

Cytochrome c1 Fe3+/ Fe2+

+0,23

+0,19

-8,7

Cytochrome c Fe3+/ Fe2+

+0,26

+0,03

-1,4

Cytochrome a Fe3+/ Fe2+

+0,29

+0,03

-1,4

Cytochrome a3 Fe3+/ Fe2+

+0,55

+0,26

-12,0

in cc oxy 1/2 O2 / O2-

+0,81

+0,26

-12,0

+1,13

-52,0

Phosphoryl
ha
ADP
ATP

giai on th ba: Acetyl CoA c hnh thnh giai on th hai s b

oxy ha hon ton trong chu trnh Szent-Gyrgyi-Krebs (chu trnh citrat)
hnh thnh CO2, H2O v gii phng nng lng. Phn ln nng lng c gii
phng ra giai on th ba ny (khong 2/3)
Trong giai on th hai v th ba khong 30-40% nng lng ha hc
c bin thnh nhit, hn 60% nng lng ny c s dng tng hp cc
hp cht cao nng.

138

Trong chu trnh citrat, cc hydrogen tch ra s c oxy ha qua chui h

hp to nn nng lng v H2O. Nng lng gii phng c tch tr cc
phn t ATP. Ton b qu trnh c th c minh ha bng s trn hnh 8.3.
Thc n
glucid

glucose

lipid

Acid bo

protein
Amino acid

Acetyl CoA
CO2

Chu trnh
citrat
ATP
1/2O2

H2NADFADCoQCytbCytc1CytcCytaCyta3
2H+
ADP+P

O
H2O

Hnh 8.3. Tin trnh oxy ha sinh hc

8.4. Chui h hp t bo v s phosphoryl ha oxy ha

8.4.1. Chui h hp t bo
Chui h hp t bo l mt h thng cc enzyme xc tc vn chuyn H+
v eletron t c cht n phn t oxygen to H2O. Trong t bo, oxygen l
cht oxy ha vn nng, cn cc phn t hu c khc nhau ng vai tr cht cho
in t. y, in t v ion hydrogen ca phn t c cht khng chuyn trc

139

tip cho oxygen khng kh m c chuyn dn qua mt chui phc tp nhiu

mt xch, bao gm cc h enzyme oxy ha kh, c th nng oxy ha kh nm
trong khong gia th nng oxy ha kh ca c cht v ca oxygen. Cc h
enzyme ny c sp t theo mt trt t tng dn th nng oxy ha kh to
thnh mt chui, gi l chui h hp hay chui vn chuyn in t ca t bo.
Vai tr ca chui h hp l oxy ha tng bc hydrogen ca c cht n H2O.
C ch hot ng ca chui h hp t bo c th tm lc nh sau:
Cht cho nguyn t hydrogen l NADH + H+ hoc trong mt s trng
hp l FADH2. Nguyn t hydrogen s c chuyn ti h coenzyme Q (CoQ)
thng qua h trung gian flavoprotein cha st v lu hunh. Tip theo hai in
t ca nguyn t hydrogen c tch ra v i vo h thng vn chuyn in t
theo trnh t cc cytochrome b-c1-a-cytochromeoxydase (a3), cui cng in t
c chuyn cho oxygen. Nguyn t oxygen b kh ( trng thi ion ha) s
kt hp vi 2H+ (proton) to ra phn t nc.
Qu trnh chuyn hydrogen v in t trong chui h hp c th phn
thnh 4 giai on:
- Giai on 1: Thng thng hydrogen c tch t c cht bi
dehydrogenase c coenzyme NAD+(hoc NADP +). Hydrogen ca c cht gn
vo NAD+, c cht t dng kh chuyn thnh dng oxy ha v NAD+ t dng
oxy ha bin sang dng kh. Mi c cht c mt dehydrogenase c hiu
tng ng:
AH2 + NAD+ A + NADH +H+
(Trong AH2 v A l c cht dng kh v dng oxy ha)
NADH khng th t oxy ha bi oxygen c, tc l khng th trc tip
chuyn hydrogen cho oxygen m phi chuyn sang cho dehydrogenase khc
c coenzyme l FMN hoc FAD.
- Giai on 2: NADH (hoc NADPH) b oxy ha bi dehydrogenase.
Enzyme ny l mt flavoprotein c coenzyme l FMN hoc FAD. Hai eletron
c chuyn t NADH + H+ ti FMN (hoc FAD) cho FMNH2 (hoc
FADH2):
NADH + H+ + FMN NAD+ + FMNH2
NADH dehydrogenase cng cha st, cht ny c l gi vai tr vn
chuyn eletron. st khng tham gia vo mt nhm hem no. NADH
dehydrogenase l mt protein cha st khng thuc hem.

140

- Giai on 3: H+ v eletron c chuyn t FMNH2 ti coenzyme Q l

mt dn xut quinone, cn c gi l ubiquinon (UQ). Coenzyme Q l mt
cht tc dng chuyn vn kh linh hot eletron gia flavoprotein v h thng
cytochrome. Ubiquinon c th nhn 1 hoc 2e- v to ra semiquinone (UQH-)
hoc ubiquinol (UQH2). c tnh ny cho php n lm cu ni vn chuyn e- t
cht cho 2e- sang cht nhn 1e-. Ngoi ra, v phc UQ nh v k nc, nn n
d dng di chuyn trong lp lipid i ca mng ty th lm con thoi vn chuyn
e- gia cc phc vn chuyn e- cng knh khc trong mng ty th.
- Giai on 4: Cc enzyme vn chuyn eletron t CoQH2 n oxygen.
l h thng cytochrome, n gi vai tr trung tm trong h hp t bo. Mi
cytochrome l mt protein enzyme vn chuyn electron c cha nhm ngoi
hem. cc phn t cytochrome, nguyn t st lin tc i t trng thi st hai
(Fe2+) - dng kh ti trng thi st ba (Fe3+) - dng oxy ha trong qu trnh
chuyn vn eletron. Nhm hem chuyn vn mt eletron; ngc li vi NADH,
flavin v coenzyme Q l nhng cht chuyn vn hai electron.
C 5 cytochrome gia CoQ v O2 trong chui chuyn vn electron. Th
nng Oxy ha kh ca chng tng theo th t: cytb, cytc1, cytc, cyta, cyta3.
Cu trc v tnh cht ca cc Cytochrome ny khc nhau. Nhm ph ca
Cytochrome b, c1, c l protoporphyrin c st, thng gi l hem. Cytochrome a
v a3 l nhng thnh phn cui ca chui h hp t bo, chng dng mt
phc cht gi l Cytochrome oxydase. Electron c chuyn ti phn
Cytochrome a ca phc cht, ri ti Cytochrome a3 c cha ng (Cu+) dng
kh trong qu trnh vn chuyn electron, c l n tham gia xc tc vn chuyn
electron t hem A ca Cytochrome a3 ti oxygen.
Qu trnh vn chuyn electron qua h thng Cytochrome c tm lc
nh sau:
2e-

+ 2 cytb Fe3+

2 cytb Fe2+

2 cytb Fe2+

+ 2 cytc1 Fe3+

2 cytb Fe3+

+ 2 cytc1 Fe2+

2 cytc1 Fe2+

+ 2 cytc Fe3+

2 cytc1 Fe3+

+ 2 cytc Fe2+

2 cytc Fe2+

+ 2 cyta Fe3+

2 cytc Fe3+

+ 2 cyta Fe2+

2 cyta Fe2+

+ 2 cyta3 Fe3+

2 cyta Fe3+

+ 2 cyta3 Fe2+

2 cyta3 Fe3+

+ 1/2 O2

2 cyta3 Fe3+

+ 1/2 O2-

141

Ton b chui h hp t bo t c cht dng kh AH2 ti oxygen phn

t qua NAD, flavoprotein, coenzyme Q, h thng Cytochrome c trnh by
hnh 8.4.
AH2

A
NADH + H+

NAD
ATP

ADP + P

FADH2 (FMNH2)

FAD (FMN)

CoQ
2Fe2+

CoQH2
2cytb

ATP

2Fe3+
ADP + P

2Fe3+

2cytc1 2Fe2+

2Fe2+

2cytc

2Fe3+

2Fe3+

2cyta

2Fe2+

ATP

2H+

ADP + P
2Fe2+

2cyta3 2Fe3+
H2O

1/2O2

1/2O2-

Hnh 8.4. Chui h hp t bo

Kt qu ca chui h hp t bo thng thng l H2O, nhng vn c

trng hp to thnh gc superoxyd (O2) v hydrogenperoxyd (H2O2). y l
cc cht c i vi t bo v chng tn cng cc acid bo khng no cu to
lipid mng t bo gy s bin cht ca cu trc mng. Theo cc s liu thc
nghim th v tr to thnh O2 chnh l vng CoQ - cytochrome b do qu trnh
t oxy ha ca cibi-semiquinone. Nh vy, thng xuyn c s r r 1 in t
trong ty th v ty th s dng khong 1 - 2% s lng electron vn chuyn
n cytochrome oxydase to thnh O2.

142

Superoxyd dismutase cha Mn (Mn.SOD) c mt trong matrix ch

chuyn c khong 80% O2 do s r r in t thnh H2O2. 20% O2 to
thnh c chuyn vo cytoplasme, y superoxyd dismutase ca
cytoplasme (SOD) cng hp tc vi cc h thng bo v khc s phn hy tip.
C th biu th cc qu trnh trn nh sau:
O2 + e- O2 (gc superoxyd )
2O2 + 2H+
2H2O2

C-ase

SOD

H2O2 + O2

2H2O + O2

SOD v C-ase l cc enzyme chng oxy ha (antioxydant enzymes), bo

v t bo chng li cc gc t do c hi.
Nh vy, qu trnh vn chuyn hydrogen n oxygen to ra H2O, thc
cht l mt qu trnh trao i electron (cho v nhn) mt cch lin tc. Bn cht
ca n l mt qu trnh oxy ha kh. V vy, ngi ta gi h hp t bo l oxy
ha kh sinh hc.
Mt iu cn lu thm l: chui h hp t bo trnh by l chui h
hp t bo bnh thng, nhng trong mt s trng hp, chui c th ko di
hoc ngn hn ph thuc vo th nng oxy ha kh ca c cht.
Quan nim hin i v h hp t bo cn b sung thm nhiu chi tit ca
qu trnh h hp t bo kinh in nh trnh by. Nhng dng di chuyn in
t v hydrogen cn ph thuc vo trng thi c cht n cc phc hp khc
nhau.
8.4.2. S phosphoryl ha oxy ha
Qu trnh tng hp ATP l qu trnh phosphoryl ha:
ADP + H3PO4 ATP
y l qu trnh cn nng lng. Nh chng ta bit, mi lin kt cao
nng trong ATP cha nng lng t do l 7Kcal/mol nn tng hp c
ATP t ADP theo phn ng trn cn cung cp nng lng tng ng
7Kcal/mol. Ngun nng lng cung cp cho qu trnh phosphoryl ha rt khc
nhau. S phosphoryl ha quang ha l qu trnh tng hp ATP lc lp th
nh nng lng nh sng xy ra trong quang hp. S phosphoryl ha oxy ha
l qu trnh tng hp ATP ty th nh nng lng thi ra trong cc phn ng
oxy ha kh.
Theo quan nim hin nay, s phosphoryl ha oxy ha l qu trnh hnh
thnh ATP bng cch chuyn electron v proton trong chui h hp t bo. S
to thnh ATP trong chui h hp t bo c th hin hnh 8.4. Theo

143

phng trnh (2) cn c s chnh lch th nng oxy ha kh gia cc cht tham gia
trong chui h hp t bo vo khong 0,152 volt to thnh mt phn t ATP
'

Eo =

G
nF

7
= 0,152volt
2.23,06

Trong chui h hp c 3 im tng hp gia s h hp vi s

phosphoryl ha: 1) gia NADH vi flavoprotein; 2) gia cytochrome b v c1;
3) gia cytochrome a v cytochrome oxydase (hnh 8.4.). iu c ngha l
proton v electron oc chuyn t NADH + H+ ti oxygen to c 3 im
phosphoryl ha, cn proton v electron c chuyn trong chui h hp t bo
t FADH2 ch c 2 im phosphoryl ha.
Mi tng quan P/O (t s P/O) l s phn t phosphate v c oc
chuyn thnh dng hu c i vi s tiu th mt nguyn t oxygen. T s ny
biu th s tng quan gia qu trnh phosphoryl ha v s oxy ha kh t bo,
c gi l ch s .
Nh vy c th ni rng s phosphoryl ha oxy ha qua h thng vn
chuyn in t ca chui enzyme h hp l con ng ch yu i vi cc sinh
vt hiu kh nhm khai thc nng lng ca cc hp cht hu c mt cch hu
hiu nht phc v cho cc hot ng sng ca mnh.

144

TI LIU THAM KHO

Ti liu ting Vit
1. Nguyn Hu Chn, 1983. Enzyme v xc tc Sinh hc. Nxb Y hc, H Ni.
2. Nguyn Hu Chn, Nguyn Th H, Nguyn Nghim Lut, Hong Bch
Ngc, V Th Phng, 2001. Ha sinh. Nxb Y hc, H Ni.
3. Phm Th Trn Chu, Trn Th ng, 2000. Ha sinh hc. Nxb Gio
dc, H Ni.
4. L Don Din, 1975. Ha sinh thc vt. Nxb Nng nghip, H Ni
5. Nguyn Tin Thng, Nguyn nh Huyn, 1998. Gio trnh sinh ha hin
i. Nxb Gio dc, H Ni
6. Nguyn Xun Thng, o Kim Chi, Phm Quang Tng, Nguyn Vn
ng, 2004. Ha sinh hc. Nxb Y hc, H Ni.
7. L Ngc T, La Vn Ch, Phm Trn Chu, Nguyn Ln Dng, 1982.
Enzyme vi sinh vt. Nxb KH&KT, H Ni.
8. L Ngc T (ch bin), L Vn Ch, ng Th Thu, Phm Quc Thng
Nguyn Th Thnh, Bi c Hi, Lu Dun, L Don Din, 2000. Ha sinh
Cng nghip, Nxb KH&KT, H Ni.

Ti liu ting nc ngoi

1. Farkas G. 1984. Nvnyi anyagcserelettan. Akadmiai Kiad Budapest.
2. Fehr J. - Verekei A., 1985. Szabad Gyk Reakcik Jelntsge az
orvostudomnyban. Medicina Knyv Kiad Budapest.
3. Karlson. P., 1972. Biokmia. Medicina Knyv Kiad Bydapest.
4. Lehninger A. L., 2004. Principle of Biochemistry, 4th Edition. W.H
Freeman, 2004.
5. Stryer L., 1981. Biochemistry. W.H.Freeman and company. San Francisco.

182

Chng 9

Cc sn phm chuyn ha
Mt trong nhng pht hin c ngha quan trng ca vi sinh vt hc
cng nghip l vic nhn thc c rng vi sinh vt c th c s dng
thc hin cc phn ng ho hc c bit vt ra ngoi kh nng ca
ho hc h c. Qu trnh s dng vi sinh vt cho mc ch ny c tn l
s chuyn ho sinh hc, n bao gm s sinh trng ca vi sinh vt trong
nhng ni ln men ln theo sau l s b sung ho cht cn c chuyn
ho ti mt thi im thch hp. Tip tc ln men thm mt thi gian na
v sinh vt tc ng ln ho cht ri tch chit dch ln men, v cui
cng sn phm mong mun c tinh khit. Mc d v ngnyn l chuyn
ho sinh hc c th c s dng cho nhiu qu trnh khc nhau song
trong thc t n ch c ng dng sn xut mt s hormone nht
nh.

I. S chuyn ha cc steroid
Vic s dng vi sinh vt thc hin nhng s chuyn ho steroid
c ngha rt ln trong cng nghip dc phm. Steroid hormones iu
chnh nhng trng thi trao i cht khc nhau ng vt k c ngi.
Mt trong nhng hormone , cortisone, c tc dng lm gim cn au c
lin quan n bnh vim khp. Cc dn xut cortisone khc lm du cc
triu chng lin quan n cc bnh d ng hoc vim.
Nhiu loi steroid hormones iu chnh hot ng gii tnh ngi
trong mt s c sn xut thnh dng thuc ung trnh th thai.
Cc c tnh sinh l ca mt steroid ph thuc vo bn cht v v tr chnh
xc ca cc thnh phn ho hc nm trn cu trc vng ca steroid gc.
Vo u nhng nm 1930, Kendall Trng i hc Tng hp
Basel tch c cortisone, mt steroid do tuyn thng thn tit ra.
Khong mt thp k sau Hench ch ra rng vic ung cortisone c th lm
du cn au cc bnh nhn b bnh vim khp.
Nhu cu thc t ca hormone tr nn cp bch v cc phng php
ho tng hp steroid c pht trin v th trng tim tng l rt ln. Tuy
nhin, ho tng hp kh phc tp yu cu ti 37 bc trong nhiu bc
xy ra di cc iu kin cc tr. Cortisone tng hp c theo con ng
ny tr gi 200 la mt gam.

183

Mt trong nhng iu phc tp ch yu gp trong ho tng hp

cortisone l vic phi a mt nguyn t oxygen vo mt v tr trong cu
trc steroid 4 vng c gi l v tr 11; y l bc quyt nh trong vic
to nn hot tnh sinh l ca phn t.
Vo nm 1952, Peterson v Murray thuc hng Upjohn pht hin
ra rng nm mc mc trn bnh m Rhizopus arrhizus c kh nng
hydroxyl ho progesterone, mt steroid khc, bng cch a mt
nguyn t oxygen vo v tr 11. Progesterone l mt tin cht trong ho
tng hp cortisone, v bng phng php hydroxyl ho nh vi sinh vt
(trong cng nghip thng dng cc chng h hng vi R. arrhizus) vic
tng hp c rt ngn t 37 xung 11 bc. Nh vy gi thnh gim
xung cn 6 la mt gam.
S hydroxyl ho progesterone mang li hiu qu kinh t do rt
ngn c s tng hp ho hc. S ln men c th thc hin 30oC vi
nc l dung mi v p sut ca kh quyn. Cc phn ng di cc iu
kin ny r hn nhiu so vi cc phn ng din ra di cc iu kin cc
tr v nhit v p sut v dung mi khng phi l nc nh trong ho
tng hp cortisone. n nay c mt s qu trnh khc ng dng vi sinh
vt tng hp cng nghip cc steroid. Nm Cunninghamella
blakesleana cng c kh nng hydroxyl ho steroid cortesolon to
thnh hydrocortisone nh vic gn oxygen vo v tr s 11. Nhng s
chuyn ho nhn steroid khc do vi sinh vt thc hin bao gm s hydro
ho, s loi hydro, s epoxygent ho, v s loi hoc thm cc chui bn
(hnh 9.1).
Cc steroid t c ngha thng mi l cc cocticosteroid nh
cortisone, hydrocortisone (hnh 9.2, hnh 9.3), prednison v dexametazon,
kch t tnh c testosteron v hormone ng dc estraiol (hai loi sau
dng cho cc loi thuc trnh th thai) v spironolacton (thuc li tiu).
Nguyn liu dng cho tt c cc qu trnh trn l cc loi ru phc
tp c tn l cc sterol. C hai ngun sterol thng thng : Sn xut du
u tng li mt cht thi giu stigmasterol v sitosterol; r ca cy
barbasco Mhic cha diosgenin. Ngoi ra, c th s dng ergosterin ly
t nm men hoc cc steroid sn xut thun tu bng con ng tng hp.
Nhiu vi sinh vt c kh nng thc hin cc phn ng chuyn ho
steroid. Tuy nhin iu quan trng l chng c th tin hnh phn ng vi
tc chuyn ho cao, hiu sut ln v khng to thnh sn phm ph hay
khng. Do yu cu ny m s chng c th s dng cho cng nghip b

184

thu hp li rt nhiu. hydroxyl ha ngi ta s dng x khun v nm

(c bit l Fusarium v cc loi Curvularia).
hydroxyl ho v tr 11-a ca progesterone, thay cho h si nm
ngi ta dng bo t trn ca Aspergillus olivaceus. Vic hydro ho c
tin hnh vi cc loi Saccharomyces, Streptomyces v Rhizopus. loi
hydro ngi ta dng cc vi khun nh Corynebacterium v nm
(Fusarium, Calonectria, Cylindrocarpon) cn ct vng c th dng
Penicillium chrysogenum hay Pseudomonas testosteroni cha mt
steroidizomerase c hiu.
Trong mt qu trnh chuyn ho steroid in hnh, vi sinh vt trc
ht c a vo mt mi trng thch hp khng cha steroid. Ngi ta
thng s dng cc ni ln men nh (10-50m3). Thng vo cui pha sinh
trng logarit th steroid mi c a vo vi nng 0,05-0,1%. V cc
steroid tan yu trong nc nn ngi ta dng cc dung mi hu c ho tan
c trong nc (nh methanol, propylenglycol, dimetylsulfoxygent) lm
cht ho tan trung gian.
Thi gian chuyn ho ko di t 6 n 48 gi, thng thng th qu
trnh c kt thc sau 20 gi bng cch tch t bo v chit sn phm.
Hiu sut t c l 60-95%. Cc sn phm ca phn ng nm bn ngoi
t bo.
Vic theo di phn tch s chuyn ho xc nh thi gian ln men
cc thch gi vai tr rt quan trng, v nu vt ra ngoi thi gian s
xy ra cc phn ng k tip khng mong mun. Nu khng ch tt th c
th tin hnh hai phn ng chuyn ho mong mun k tip nhau trong
cng mt ni ln men. Chng hn 6-fluo-21-acetoxygen-16-methyl-4pregnen-3,20-ion trc ht c hydroxyl ho v tr 11 nh
Aspergillus ochraceus v sau cng trong mi trng y nh b sung
Bacillus lentus m li c loi hydro v tr 1,2.
Vo nm 1980 gi cortisone M ch cn 46 cent mt gam, r
hn 400 ln so vi gi ban u. Vic tm c cc ng dng khc ca cc
steroid (dng cho trnh th thai, bnh thiu hormone, cc bnh v da,
vim v d ng) cng vi tnh hiu qu cao ca phng php sn xut
to ra mt nhu cu cp thit i vi cc loi dc phm ny. Doanh thu 4
loi steroid chnh (cortisone, aldosteron, prednison v prednisolon) trn th
trng th gii vo nm 1987 l 300 triu la.

185

Hnh 9.1: Sn xut steroid bng cc bin php ho hc kt hp vi s

chuyn ho vi sinh vt

186

Acid Deoxycolic

Cortisone

Hnh 9.2: Coctisone c tng hp t deoxycolic acide

Hnh 9.3: S sn xut 11 -hydroxyprogesterone v hydrocortisone

187

II. S to thnh phenyl-axetylcacbinol, tin cht ca epherin

Epherin l mt alcaloit c trong cy Ephedra vulgaris. Ging nh
arenalin, n c tc dng lm tng huyt p v c dng lm thuc
iu tr cc bnh suy nhc tun hon, hen, vim ph qun v.v. .. Cu to
ho hc ca n nh sau :

Tch cht ny t thc vt l mt cng vic tn km. Ho tng hp

n cng kh thc hin bi v do hai nguyn t C khng i xng ca phn
t m xut hin bn ng phn lp th trong ch c dng L l c dc
tnh.
Nh Saccharomyces cerevisiae m benzaldehyde c th c
chuyn ho thnh phenyl-axetylcacbinol, tin cht ca epherin, vi hiu
sut 50-60% khi c b sung vo mi trng cha mt nng t bo l
0,8- 1% .
Trong sn phm chuyn ho ny, nhm hydroxyl nm cng v tr
nh trong L-ephedrin. Trong phn ng ny, cc t bo nm men gn
"acetalehyde hot ng" c to thnh trong s ng phn vo
benzalehyde va b sung vo mi trng (phn ng cacboligase):

Benzaldehyde

''Acetaldehyde hot ng''

acetylcacbinol

L-phenyl-

Sau phenyl-acetylcacbinol c chuyn thnh L-epherin bng

con ng ho hc nh mt s kt hp hydro amin ho.
III. Sn phm t vi khun acetic
Trong s oxygen ho ethanol thnh acid acetic, chuyn ho thc
cht l mt s s dng c cht. y, NADPH2 xut hin c chuyn
qua chui h hp thu nhn nng lng, song c cht khng b phn gii

188

hon ton do vy qu trnh cng cn c gi l s oxygen ho khng

hon ton. Loi chuyn ho ny c th tri qua nhiu bc.
Cc i din ca vi khun acetic (Acetobacter, Gluconobacter)
oxygen ho khng ch ethanol m c mt ph rng cc ru bc mt v
ru bc hai cng nh cc polyol.
Cc loi vi khun m sn phm ca chng c gi li c gi l
cc loai oxygen ho thp (suboxygendant). Thuc v nhm ny c vi
khun acetic dng trong cng nghip l Acetobacter suboxygendans. Cc
loi ch tch lu acid acetic tm thi sau li oxygen ho tip c xp
vo nhm oxygen ho cao (peroxygendant), chng hn Acetobacter
peroxygendans, Acetobacter pasteurianum). Gia hai nhm ny c cc
dng chuyn tip.
1. Sn xut dm n
Cc phn ng c dng sn xut dm n nh Acetobacter
suboxygendans l :
CH3 CH2 OH

ase
alcohol
dehydrogen

CH3 CHO + 2H

Sau s hydrat ho axetaldehyde s din ra mt s loi hydro ln th hai :

Hydro c NADP nhn v qua cc xitocrom c chuyn n O2

l cht nhn in t cui cng. S to thnh acid acetic i hi cung cp
oxygen mnh. Khi thng kh khng y c th xy ra s ho hai ca
axetaldehyde thnh acid acetic v ethanol theo phng trnh sau y (phn
ng Canizzaro) :
Ethanol li i vo phn ng th nht v cui cng cng c oxygen
ho thnh acid acetic.
Ethanol hoc acid acetic khng th c dng lm ngun cacbon
duy nht v ngi ta khng tm thy cc enzyme cn thit ca chu trnh
glyoxylate Acetobacter xylinum. V vy sinh trng vi khun cn
c b sung cc ngun C khc nh glucose.

189

S cung cp oxygen c tnh cht quyt nh n k thut sn xut

dm. Trc kia qu trnh c tin hnh bng phng php nui b mt
nh mt lp vng ca Acetobacter xylinum (phng php Orlans), trong
vi khun c c nh trn cc v bo g gi. V bo ng trong mt
thng g hnh tr (generator) c thng kh t pha di v ti cc dung
dch cha ru (v d cc loi ru vang km phm cht) t pha trn
(hnh 9.4).
Ngy nay cc phng php chm ngy cng c ngha hn. Hin ti
ngi ta dng cc ni ln men c bit (acetator) c thng kh rt mnh
trong dch dinh dng c thay th tng phn bng dch dinh dng
b sung cng c thng kh y .
Sau khi ethanol c chuyn ho thnh acid acetic, phn ln dch
dinh dng trong ni c ly i v thay th vo l dch dinh dng
mi. Nh c th t c mt phng thc ging nh nui cy lin
tc.
Qu trnh ln men c tin hnh nh cc chng chn lc ca
Acetobacter suboxygendans trong mt dch dinh dng cha glucose vi
10-12% ethanol. Ethanol gn nh c chuyn ton b thnh acid acetic.
Nng acid acetic cao nht t c l 13%. Qu trnh din ra 2830oC v ko di khong 48 gi. Trong phng php c in Orlans, s
ln men ko di ti 5 tun l. Trong cc phng php hin i vi nng
ru hoc acid acetic cao l 12%, th mt s ngng thng kh t 10 n 20
giy s lm cht ti mt phn ba s vi khun. nhng nng c cht
thp hn, s ph thuc vo oxygen khng n ni kht khe ti nh vy.
Vi sinh vt oxygen ha ethanol thnh acid acetic thng c gi l
cc vi khun acetic, cc vi khun ny thc hin trao i cht pH mi
trng thp, iu ny phn bit chng vi cc vi khun khc.
Cc vi khun acid acetic l bn a hnh, t bo t hnh elip ti hnh
que thng hoc hi cong 0,5-0,8 0,9-4,2 m, ng mt mnh, thnh cp
hoc thnh chui c dng khng chuyn ng v dng chuyn ng vi
tin mao cc hoc vng quanh c th. Chng l bn hiu kh bt buc,
mt s to thnh sc t, mt s to thnh cellulose. Ngi u tin tm
cch phn loi cc vi khun acid acetic l Hansen (1894).
Ngy nay cc vi khun acid acetic mi phn lp c xp vo hai
chi chnh, Acetobacter, Gluconobacter. Cc loi ca Acetobacter (trn 60)
cha 5 c im: c mt catalase, oxygen ha ethanol qua acid acetic ti
CO2 v H2O, oxygen ha lactate thnh cacbonate, oxygen ho glycerol

190

thnh DHP v s sn sinh acid gluconic t glucose. Cc vi khun trong chi

Acetobacter thng c chia thnh bn nhm : oxygen ho mnh,
oxygen ho, oxygen ho trung bnh v oxygen ho yu.

Hnh 9.4: Thit b sn xut dm theo phng php c in

IV. Sn xut vitamin C ( acid L-ascocbic )

a s ng vt tng hp c ton b lng vitamin C cn thit cho
nhu cu ca mnh v do vy vitamin ny c tm thy trong cc m ca
chng (ch yu trong gan v thn vi nng 10-40 mg/100g). Tuy nhin,
ngi v mt s ng vt c xng sng cng nh cn trng li ph thuc
hon ton v ngun vitamin C t bn ngoi. Ch yu l rau (bp ci,
spinat, c chua, 30-150 mg/100g) v qu (cam, chanh, 40-50mg/100g).
cung cp cho con ngi lng vitamin C cn thit (45-70 mg/ngy).

191

Mt s vi sinh vt (nm, nm men, to) sn sinh mt lng rt nh

acid L-ascocbic cn cho cc qu trnh trao i cht ca chng. Cho n
nay cha tm thy acid ascorbic vi khun, hnh nh chng khng cn
acid ny.
ng vt c v (tr bn linh trng v mt s khc) acid Lascorbic c tng hp t D-glucose trong C1 ca glucose tr thnh C6
ca acid ascorbic v ngc li. S tng hp din ra t D-glucose ti acid
D-glucuronic v sau thnh lacton ca acid L-gulonic.
S oxy ho sau ca gulonolacton v tr C2, c xc tc bng
L-gulonolacton dehydrogenase, mt enzyme khng tm thy ngi, theo
sau l s enol ha s cho acid L-ascorbic (hnh 9.5). thc vt, acid Lascorbic c to thnh t D-glucose hay D-galactose qua mt s con
ng chuyn ha.
Mt trong nhng con ng ny gi khng lm cho trt t ca chui
cacbon b thay i, song cc sn phm trung gian ca con ng sinh tng
hp th cn cha bit r. C th con ng ny cng ging con ng
tng hp ng vt.

Hnh 9.5: Con ng sinh tng hp acid L-ascocbic

T trn 50 nm nay, cng nghip c th p ng nhu cu ngy

cng tng v vitamin C ca con ngi nh phng php bn tng hp t
glucose. Nhiu qu trnh ho hc v sinh ha k tip nhau tham gia vo
qu trnh ny v tt c u qua mt sn phm trung gian l acid 2-keto-L-

192

gulonic, t s thu c acid L-ascorbic nh con trng ho hc bng

cch lacton ho v ng phn ho (hnh 9.6).

Acid-keto-Lgulonic

Methyl-2keto-Lgulonate Acid L-ascobic

Hnh 9.6: S chuyn ho ho hc acid 2-keto-L-gulonic thnh acid Lascocbic

Cc quy trnh chun b acid 2-keto-L-gulonic c th c xp thnh

hai nhm: (1) nhm bt u bng s kh D-glucose v lm o ngc trt
t cacbon, v (2) nhm bt u bng s oxygen ha D-glucose trong
khng c s o ngc chui cacbon.
1.Cc quy trnh bt du bng s kh D-glucose
Quy trnh cng nghip ny c Reinstein v Gruessner ra t nm 1934
v n nay vn cn s dng. N bao gm 5 bc phn ng k tip nhau, tt c
u t sn lng ti 90-95% (hnh 9.7)

193

Hnh 9.7: S tng hp acid 2-keto-Lgulonic t D-glucose qua Dsocbitol v L-socboza

1. Kh D-glucose thnh D-sorbitol vi s c mt ca niken Raney,

sau loi ti mc ti a niken ha tan bng cch x l dung dch sorbitol
vi nha cation.
2. Oxygen ha D-sorbitol thnh L-sorbose nh vi sinh vt : c nhiu
chng Acetobacter chu c nng niken ti 10-20 mg/l, thc hin
nhanh chng phn ng ny trong cc mi trng cha nng sorbitol
cao. Qu trnh ln men din ra 30oC trong mt mi trng cha 200 g/l
sorbitol, 10 g/l cao ng, v 0,5 g/l CaCO3 vi mt chng Acetobacter
suboxygendans hon tt trong 24 gi v cho khong 180g sorbose trong
mt lit. Sau khi lc, loi ion v c c dch nui, sorbose c kt tinh (ti
tinh khit 99%)
3. To thnh diacetone-sorbose bo v cc nhm khng tham gia
vo giai on sau.
4. Oxygen ha ha hc diacetone-sorbose thnh acid diacetone-2keto-L-gulonic.

194

5. Gii phng acid diacetone-2-keto-L-gulonic nh thu phn.

Gia nhng nm 1960 v 1975 nhiu c s nghin cu ca M v
Nht tm cch gim s bc trong quy trnh Reinstein bng cch
oxygen ha trc tip D-sorbitol hay L-sorbose nh mt k thut vi sinh
vt thnh acid 2-keto-L-gulonic khi s dng cc chng Acetobacter v
Pseudomonas. Tt c nhng nghin cu ny u khng cho kt qu kh
quan, nng sut t khi vt qu 5 g/l trong cc iu kin tt nht vi mt
sn lng vo khng 10% so vi sn phm ban u. Vo nm 1981, cc
nh nghin cu Trung Quc cho bit sn xut c 37 g/l acid 2-ketoL-gulonic t 100 g sorbose khi s dng mt chng Gluconobacter
oxygendans. y l mt s ci thin ng k so vi cc kt qu trc .
2.Cc quy trnh bt u bng s oxygen ha D-glucose
Hai con ng c nghin cu : (1) ci gi l con ng acid Lionic bt u bng s oxygen ha glucose v tr cacbon s 5 ; (2) con
ng acid 2,5-diketo-D-gluconic.
a) Con ng acid L-idonic
Con ng ny gm ba bc k tip nhau : (1) oxygen ho sinh
ha hc D-glucose thnh acid 5-keto-D-gluconic qua acid D-gluconic ; (2)
kh ha hc (hoc sinh ha hc) acid 5-keto-D-gluconic thnh acid Lidonic ; (3) oxygen ha sinh ha hc acid L-ionic thnh acid 2-keto-Lgulonic (hnh 9.8).
Phn ng u tin d dng c thc hin bi Acetobacter
suboxygendans. Chng
ATCC 621 chuyn ha 100 g/l glucose trong 33 gi vi mt sn
lng l 90%. Mt s phng php c ra nhm kh acid 5-ketoD-gluconic. Trong thc t, ch c phng php ho hc do Grey xut
(1947) l c gi tri. Trong phng php ny, s kh t c nh s hiro
ho c xc tc di p lc v khng may l dn n vic to thnh hai
ng phn, acid L-idonic v acid D-gluconic theo nhng t l tng i
m trong cc iu kin tt nht th l 70 v 30%. Hai phng php loi
acid D-gluconic va to thnh c ch : (1) nh ln men vi
Acetobacter suboxygendans, n c th b oxygen ha tr li thnh acid 5keto-o-gluconic, cht ny sau s c quay vng, hay (2) hn hp c
th c ln men vi mt vi sinh vt (Pseudomonas fluorescens, P.
aeruginosa, Acetobacter melanogenus), bn ny s dng acid D-gluconic
sinh trng v phn gii n hon ton cng mt lc vi vic chuyn
ho acid L-idonic thnh acid 2-keto-L-gulonic. Bng phng ny ngi ta

195

thu c nhng kt qu tt : 67 g/l acid 2-keto-L-gulonic c to

ra t 100 g/l hn hp cha 70% acid L-idonic, song cn c nhng vn
k thut cha c chp nhn trn quy m cng nghip.

Hnh 9.8. S chuyn ho D-glucose thnh acid 2-keto-L-gulonic qua

acid L-inonic

b) Con ng acid 2,5-diketo-D-gluconic

Con ng ny , ch yu c nghin cu t nm 1970, gm hai
bc sinh ho hc : (1) oxygen ha D-glucose thnh acid 2,5-diketo-Dgluconic qua acid D-gluconic v acid 2-keto-D-gluconic ; (2) kh acid
2,5-diketo-D-gluconic thnh acid 2-keto-L-gulonic (hnh 9.9).
Bc mt khng gp tr ngi g nghim trng. Ngay t nm 1974
cc nh nghin cu Nht Bn cho bit v mt nng sut 220 g/l 2,5diketo-D-gluconat canxi t c t 200 g/l glucose sau 3 ngy ln men
nh Acetobacter fragi. Vo nm 1982 cc nh nghin c Shionogi
s dng : (1) cho bc mt, mt th t bin ca Erwinia c kh nng sn
xut 328 g/l 2,5-diketo-D-gluconat canxi trong 26 gi; (2) trong bc hai,
h s dng mt chng t bin ca Corynebacterium to ra 106 g/l 2keto-L-gluconat canxi trong 66 gi.

196

D-Glucose

Acid D-gluconic Acid 2-keto- Acid 2,5-diketo- Acid 2-ketoD-gluconic

D-gluconic
L-gluconic
Hnh 9.9: S chuyn ho D-glucose thnh acid 2keto-L-gulonic qua acid
2,5diketo-D-gluconic

Sau bc ln men u tin, mi trng nui Erwinia c kh trng

bng cch x l vi 0,25 g/l oecylsulfate natri 28oC trong 6 gi. Sau
khi lm lnh, ngi ta b sung glucose (0,2 kg/kg acid 2,5-diketo-Dgluconic) dng lm cht cho hiro trong phn ng kh, t sinh ra acid
2-keto-L-gulonic. Cht ny c a vo mi trng nui
Corynebacterium vo cui pha sinh trng. Sn lng chuyn ha tnh
theo phn t gam s l 94% v bc hai l 92% so vi 2,5-diketo-Dgluconat canxi s dng. Xt ton b, sn lng phn t gam ca acid 2keto-L-gulonic thu c vo cui bc ln men th hai so vi lng
glucose s dng trong bc ln men th nht l 86%.
y l quy trnh hiu qu nht i vi s sn xut acid 2-keto-Lgulonic k t khi quy trnh Reinstein ra i vo nm 1934 v n loi b tt
c cc bc ha hc. S lng cc bc c gim t nm xung ch
cn hai. Vic tch acid 2-keto-L-gulonic vo cui bc ln men th hai
t c vi mt sn lng l 90% bng cch x l dch nui vi
carbonate natri v bng cch c c ti lc 2-keto-L-gluconat natri kt
tinh.
c nhng cng b v kt qn nghin cu nhm thu nhn acid 2keto-L-gulonic t glucose ch bng mt ln ln men. Cc chuyn gia ca
hng Genetech vo nm 1985 t c iu ny nh vic nhn ln
chng Erwinia herbicola ATCC 2 1988, chng ny thc hin tt bc mt
trong quy trnh Shionogi, gen m ha cho vic tng hp acid 2,5
diketogluconic reductase tch ra t Corynebacterium sp. ATCC 31090.
Gen ny biu hin tt trong t bo ch song quy trnh cn lu mi c
th s dng cho sn xut cng nghip.
Nng glucose cc i c th ti t hp ny chp nhn cn
mc tng i thp v nng sut cn rt ngho nn (0,6 g/l acid 2-keto-L-

197

gluconic c to ra sau 57 gi trong mt mi trng cha 20 g/l

glucose). V rng s sinh tng hp acid L-ascorbic ng vt c bit
r, ngi ta hy vng mt ngy no s c th a cc gen m cho cc
enzyme c lin quan vo mt vi sinh vt thch hp cui cng c th thu
nhn vitamin C trc tip t glucose.

V. Sn xut destran
a s cc polysaccharide ngoi bo t vi sinh vt u l sn phm
ca s chuyn ha ni bo c cht thnh cc sn phm trung gian, v cui
cng thnh polime. Destran khc cc polysaccharide ny, c cht khng
thm nhp vo t bo vi sinh vt m n c chuyn ha bn ngoi t bo
thnh -D-glucan phn nhnh, tc l destran. Ch c saccharose c
dng lm c cht cho phn ng ny.
Nh vy, polysaccharide c th c sn xut bi cc t bo nguyn
vn trong mi trng nui hoc c th c to ra t cc ch phm phi t
bo cha phc h enzyme destransaccharase (-1,6-glucan : D-fructose 2glucosyltransferase). Enzyme ny gii phng fructose v chuyn gc
glucose ln mt phn t cht nhn cng c lin kt vi enzyme :
(1,6--D-glucosyl)n + saccharose (1,6--D-glucosyl)n+1 +
fructose
Nng lng t do ca ln kt glucoside trong phn t disaccaride
nm vo khong 23 kJ trong khi nng lng t do ca lin kt glucoside
bn trong destran thp hn mt cht (12-17 kJ). Do vy phn ng din ra
theo chiu t tri sang phi i km vi mt s gim nng lng t do.
Trong qu trnh polime ha chui destran ang di ra vn lin kt
cht ch vi enzyme, mc polime ho tng cho n khi phn t cht
nhn (trong trng hp n gin nht l mt phn t c cht) gii phng
chui polime khi enzyme.
Cc thnh vin thp nht ca cc oligosaccharide khng phi l cc
phn t cht nhn c hiu qu, i lc i vi enzyme tng dn theo di
chui. Tuy nhin, s hn ch v khuch tn cng tng theo trng lng
phn t.
Bi vy, di cc iu kin thng thng, cc destran c trng
lng phn t rt cao s trc to thnh, c bit cc nng c cht
thp.
sn xut cc destran c trng lng phn t trung bnh, cn phi
tin hnh thu phn cc sn phm c trng lng phn t cao (bng acid

198

hoc bng enzyme) v phi phn on sn phm di nhng iu kin

kim sot rt cn thn. Trong khi , cc polime trng lng phn t thp
v ng nht c th c ly ra dng lm nguyn liu ban u cho mt
phn ng khc sn xut cc destran c mc trng hp mong mun.
Cc nng saccharose cao v c th c maltose khi c dng lm
nguyn liu ban u s to nn cc iu kin thun li tng hp cc
destran mi trng lng phn t thp v ng nht kiu .
Destran c sn xut bi hng lot loi vi khun, nh
Streptobacterium destranicum, Streptococcus mutans v cc loi khc.
Streptococcus mutans gi mt vai tr quan trng trong bnh su rng. Sn
phm ph thuc c vo chng ln vo cc iu kin s dnng cho sinh
trng v tng hp polime. mt s chng, destransaccharase nm dng
ha tan trong khi cc chng khc, n lin kt mt phn hay hu nh
hon ton vi t bo. Sn xut destran cng nghip dng Leuconostoc
mesenteroides to ra mt polime cha khong 95% lin kt -1,6 (phn
cn li l lin kt -1,3) v mt trng lng phn t l 4-5 107 dalton.
Cc destran c s dng trong ngnh dc lm cht dn mu
(cht thay th huyt tng). Tuy nhin nhng destran ny phi c trng
lng phn t thp nm trong mt phm vi rt nh (75000 25000).
Cc sn phm vi trng lng phn t thp hn c loi khi h
tun hon qu nhanh nn khng c gi tr iu tr y trong khi cc sn
phm c trng lng phn t qu cao li tham gia vo s ngng kt mu.
Cc dung dch cha 6% loi destran ny c gi tr v nht v tp
tnh keo-thm thu rt ging vi huyt tng ca ngi. V cc destran
khng b phn gii bi cc amylase trong c th ngi nn kh nng ph
hoi gan thp hn so vi cc cht thay th huyt tng khc. Cc destran
c m c th hp th rt tt, do vy c th c s dng cho nhiu loi
dc phm, chng hn st trong khi iu tr bnh thiu mu.
Gn y, ngi ta cho rng destran c th c s dng to nn
mt lp a nc trn b mt cc vt bng rng v hp th cc cht tit
t cc vt thng.
Trong vic sn xut cc destran, vic duy tr cung cp oxygen gi
mt vai tr quan trng c bit v trong qu trnh ln men dung dch tr
nn rt nht.
Hin nay ngi ta cha c kh nng tnh ton mi tng quan gia
nng lng khuy trn a vo v tc chuyn ng ca oxygen trong

199

dung dch vi tp tnh phi Newton ca dch lng nn trong a s trng

hp ngi k s sinh hc ch cn bit da vo kinh nghim ca bn thn
mnh.
Mi trng ln men cha cc mui v c, 2% cao ng v
saccharose. St v mangan l nhng nguyn t vt rt quan trng. Dung
dch c kh trng bng sc nng, c chuyn sang b ln men v cy
ging sau khi lm lnh.
Ln men c tin hnh 25oC, trong qu trnh ny, ch yu do s
to thnh acid lactic, gi tr pH gim t 6,5-7,0 xung cn 4,0. Dung dch
ln men c khuy lin tc v thng kh.
Khi destran c to thnh, dung dch tr nn m c cho n khi
qu trnh kt thc sau 3 hay 4 ngy, mt khi dng keo s c to thnh.
Dch ln men cha ch yu l destran, fructose t do, acid lactic, etanol v
cc vi sinh vt. Destran sau s c kt ta bng methanol. Dch trong
c cht li v methanol c thu hi bng phng php ct.
Nu ln men dn n vic to thnh cc destran c trng lng phn
t cao th chng s c thu phn bng acid chlohydric 100-105oC.
S hon tt thu phn c xc nh bi nht. Sau mt giai on
phn ng nht nh, phn ng lc u c lm chm li bng cch lm
lnh sau dng li hon ton bng cch b sung hydroxygenate natri.
Dung dch c trn vi cc yu t hp ph v lc qua kizengua.
Cui cng destran c kt ta phn on vi methanol, cc phn on
mong mun s c ti ha tan, lc, c c v sy phun. V rng vic thu
phn bng acid kh kim sot v vic phn on thng khng thu c
cc phn c thnh phn mong mun, nn ngi ta tm cch tng hp
trc tip cc destran c trng lng phn t thp. lm iu ny cn c
kin thc chc chn v ng hc ca phn ng.
Trng lng phn t b nh hng bi : a) nng ca saccharose,
b) cht nhn glucose, c) nhit phn ng. Chng hn khi vng mt mt
cht nhn, mt dung dch saccharose 10%, s thu c mt sn phm vi
mt trng lng phn t ln hn 108 dalton. nng 70% saccharose,
cc phn t destran ch yu c trng lng phn t thp s c to
thnh. Nu cc destran trng lng phn t thp c a vo lm mi th
enzyme s c nhiu im khi u v cc polime tng i ng nht s
c to thnh. iu quan trng l cc t bo vi khun khng cha cc vt
ca cc destran c trng lng phn t cao. V vy nguyn liu cy phi
c ra vi ln bng dung dch mui loi cc destran bm vo.

200

Nh vy mt hn hp 2% (w/v) mt cht nhn c trng lng phn

t thp (1000-25000) vi mt nng ng l 10% 15oC v pH 5,0 s
cho mt sn phm m 50% c trng lng phn t trong phm vi mong
mun gia 50000 v 100000 dalton. Cc destran ny c s dng trong
bnh vin v khng b phn gii bng thy phn.
Theo tnh ton sn lng destran vo nm 1980 Ty u l 1000
tn, v trn th gii l 200.000 tn. Trong khi hu ht cc polysaccharide
ngoi bo c s dng di dng khng b bin i v mt ha hc th
destran dng cho ngnh dc v nhiu mc ch khc li c chuyn ha
thnh cc phn on c trng lng phn t tng i thp gia 40000 v
70000 dalton nh cc quy trnh thu phn thch hp (thu phn bng acid
long).
Th trng ch yu ca cc dn xut destran di dng cc ch
phm lin kt cho cha cu trc khng gian ba chiu m cc nhm chc
nng c gn vo nh cc lin kt ete vi cc gc glucose chnh l
cc phng th nghim sinh ha hc.
sn xut cc loi gel cho mc ch sinh ha, destran c trng
lng phn t tng i cao c dng tng hp cc sn phm c tnh
thu hi nc cao. Phn ng vi destran trong dung dch kim dng
epichlorohydrin thu nhn gel, gel ny sau c nghin v trung ha.
V phn ng gia epichlorohydrin v polysaccharide l phn ng ta
nhit, cn ch nhit ta ra c loi b khi h thng.
iu ny l c bit quan trng v gel c to thnh trong phn ng
c tnh dn nhit thp. V cc gc ng kh tn cng ca cc phn t
destran c tnh phn ng capo v d b phn gii trong phn ng polime
nn chng b kh thnh sorbitol v borohydride trong dung dch kim
trc khi b sung epichlorohydrin.
Mc du cc gel lin kt cho thuc loi sephaex bn thn chng
cc k c ch trong vic phn on cc nguyn liu hot ng v mt sinh
hc, song s chuyn ha chng thnh cc dn xut ete s lm tng gi tr
tim tng ca chng. Cc ete carboxymethyl hay dimethylaminoethyl l
nhng sn phm kiu . Nhng hp cht ny khc bit nhiu vi cc
destran lin kt cho v tp tnh trng ca chng trong dch lng. Cc sn
phm c lin kt cho khc nhau s cho ra cc cht hp ph c l khc
nhau v gii hn phn on khc nhau.
Vic a cc nhm ha hc khc vo cc phn t s cho ra cc cht
hp ph mang c im a lipit. Chng c th c dng phn on

201

cc lipit, v.v. v c th dng trong dung mi lng cng nh trong cc dung

mi hu c phn cc. Mt s destran khc c dng lm cc cht mang
trong cc quy trnh nui cy t bo. Di dng ny (c tn thng mi do
hng dc phm AB ca Thy in t ra l Sephaex), destran c ng
dng rt rng ri tch v thun khit cc phn t sinh hc khc nhau v
in tch, v kch thc phn t v.v.. Protein khng b bin tnh bi mng
li polime a nc v s hp ph khng c hiu l rt thp.

Cu hi n tp chng 9
1. Da vo s sau y, hy nu cc nguyn tc ca s chuyn ha
steroid:

2. Nu cc bc chnh trong qu trnh sn xut mt

steroid.

hormone

3. Phn bit cc vi khun acetic peroxydant v suboxydant

4. Nu s khc bit trong cc phng php sn xut dm (Orleans,
Spring, phng php chm).
5. Nu ng dng v nguyn tc ha hc ca s to thnh cc destran.

202

6. Mt trong nhng iu phc tp ch yu gp trong ha tng hp

cortisone l................
y l bc quyt nh to nn.............Trong vi sinh vt hc cng
nghip, c th dng cc vi sinh vt sau y thc hin bc phn ng
ny: ...........
7.Destran c sn xut cng nghip nh.......................Trong mi
trng cha saccharose, vi khun ny tit ra enzyme.......................c chc
nng .........................

175

Chng 10

Trao i Lipid
10.1. S phn gii lipid
10.1.1. Phn gii glycerid
Glycerid d dng b thy phn do s xc tc ca cc loi lipase.
H2COOCR1

lipase

R1COOH

H2COH

HCOOCR2 + 3H2O

HCOH

H2COOCR3

H2COH

R2COOH
R3COOH

ng vt s thy phn glycerid xy ra nhanh chng nh s tc

ng ca mui acid mt lm nh tng ha glycerid nn d b thy phn.
10.1.2. S oxi ha acid bo
Acid bo b phn gii bng nhiu con ng:
- oxi ha.
- oxi ha.
- oxi ha.
Trong con ng ph bin v quan trng nht l .oxi ha.
10.1.2.1. .oxi ha acid bo
S phn gii acid bo bng cch ct dn tng cp C, tc l ti v tr
C ca chui carbon. Cc acid bo c mch carbon chn v cc acid bo
c mch carbon l c c ch .oxi ha khc nhau giai on cui. Acid
bo no v acid bo khng no c s khc nhau giai on sau.
* i vi acid bo no c mch C chn:
Qu trnh .oxi ha xy ra qua nhiu phn ng phc tp:

176

R - CH2 CH2 CH2 COOH A

CoA-SH

(Acid bo)

ATP

AMP +(P-P)

R.CH2 CH2 CH2 CO SCoA (Acyl-CoA)

FAD
2
FADH2
R-CH2 CH = CH CO SCoA (Enoyl-CoA)
H2O
3
R-CH2 CHOH CH2 CO SCoA (-Hydroxy-acyl-CoA)
NAD
4
NADH2
R-CH2 CO CH2 CO SCoA (-Ceto-acyl-CoA)
CoASH

R - CH3- CO SCoA
(Acyl-CoA)

5
CH3CO SCoA
(Acetyl-CoA)

177
Cc enzyme tham gia xc tc cc phn ng trn l:
1. Acyl-CoA-Synthetase.
2. Acyl-CoA-Dehydrogenase.
3. Enoyl-CoA-Hydratase.
4. Hydroxy-acyl-Thiolase.
Qua mt chu k phn ct, phn t acid bo ngn bt i 2 carbon,
kt qu cui cng ca cc chu k phn ct .oxi ha ca acid bo l cc
phn t acetyl-CoA . Nu phn t acid bo c n nguyn t C th s to ra
n/2 phn t acetyl-CoA. Cc phn t acetyl-CoA tip tc b phn gii qua
chu trnh Krebs to CO2 v H2O.
V mt nng lng, qu trnh .oxi ha to nn ngun nng lng
ln cung cp cho cc hat ng sng ca t bo.
Mi ln phn ct bt 2C s to nn 1 NADH2, 1FADH2, qua chui
h hp s tng hp c 5 ATP. ng thi mi phn t Acetyl-CoA b
phn gii thng qua chu trnh Krebs s to ra c 12ATP. T ngi ta
tnh c tng s ATP c to ra do s phn gii phn t acid bo no,
mch cacbon chn c n nguyn t C l:
n n

5 2 1 1 + 2 . 12 ATP

* i vi acid bo no c mch C l
i vi cc acid bo no c mch C l, qu trnh phn gii theo
phng thc .oxi ha xy ra ging vi acid bo no c mch carbon chn
ni trn nhng sau ln phn ct cui cng khng phi to ra 2 phn t
Acetyl-CoA m cho ta 1 phn t Acetyl-CoA v 1 phn t propionyl-CoA.
T propionyl-CoA li tip tc bin i thm mt chu k .oxi ha
na to ra 1 phn t CO2 v 1 phn t Acetyl-CoA.

178

CH3 CH2 CO SCoA

FAD

FADH2
CH2 = CH COSCoA
H2O

CH2OH CH2 COSCoA

CoA-SH
CH2OH CH2 COOH
NAD
NADH2
CHO CH2 COOH
CoASH

CO2

NAD
NADH2

CH3CO SCoA
Cc enzyme xc tc ging nh chu trnh trc
* i vi acid bo khng no.

179
Vi acid bo khng no, qu trnh phn gii xy ra ty v tr ni i.
- Nu v tr ni i ng vo v tr th qu trnh xy ra ging nh i vi
acid bo no nhng khng xy ra phn ng 2.
- Nu v tr ni i v tr khc th trc khi phn gii, acid bo khng no
b kh thnh acid bo no tng ng ri tip tc phn gii theo con
ng .oxi ha.
Acid bo no

Acid bo khng no
FADH2

FAD

10.1.2.2. .oxi ha
Phng thc .oxi ha l s phn gii acid bo xy ra ti v tr C,
mi ln phn gii mch C b ct ngn i 1 nguyn t C v to ra CO2.
2

R CH2 CH2 - COOH

H2O2

CO2
3

R CH2 CHO
NAD

NADH2
3

R CH2 COOH

180
10.1.3. Phn gii glycerin
Sau khi gii phng khi lipid n gin, glycerin tip tc c bin
i bng nhiu cch to nn cc sn phm khc nhau
CH2OH

ATP

ADP

CH2OH

NAD

NADH2

CHO

CHOH

CHOH

CHOH

CH2OH

CH2O P

Glycerin

Glycerol-P

CH2OP
Aldehyde phospho
glycerol (ALPG)

T ALPG bin i thnh pyruvic acid nh trong qu trnh ng

phn, sau pyruvic acid b phn gii tip thng qua chu trnh Krebs
to CO2 v H2O.
Nh vy s phn gii glycerin xy ra qua qu trnh ng phn v
qua chu trnh Krebs to sn phm cui cng l CO2 v H2O. Nng
lng gii phng trong qu trnh phn gii ny c dng tng hp
ATP cung cp cho cc hat ng sng ca t bo.
10.1.4. Phn gii glycero phospho lipide
Cc glycero phospholipide b phn gii qua nhiu giai on to ra
nhiu sn phm trung gian tham gia vo nhiu qu trnh trao i cht. Qu
trnh phn gii ny do nhiu enzyme thy phn xc tc.
- Phospholipase A1 thu phn lin kt gia glycerin v acid bo
th nht.
- Phospholipase A2 thy phn lin kt gia glycerin v acid bo
th hai.
- Phospholipasse B thy phn c hai loi lin kt trn.
- Phospholipase C thy phn lin kt gia glycerin v H3PO4.
- Phospholipase D thy phn lin kt gia H3PO4 vi choline,
ethanolamine hay serine.
Phi hp tc dng ca tt c cc loi enzyme ni trn, phn t
glycero phospholipid s b phn gii thnh glycerin, acid bo, H3PO4 v
choline (hay ethanolamine, serine)

181

A1

R1CO

O CH2
A2

R2CO

O CH

H2 C O

CH2 CH2 N (CH3)3

OH

10.2. Tng hp lipid

10.2.1. Tng hp acid bo
Acid bo c tng hp t acetyl-CoA. S tng hp acid bo no
v khng no giai on u ging nhau. Trc ht acid bo no c tng
hp sau hnh thnh acid bo khng no bng cch oxi ha cc acid bo
tng ng.
Qu trnh tng hp acid bo t acetyl-CoA xy ra ngc vi qu
trnh .oxi ha. T cc acetyl-CoA c ni dn li vi nhau thnh chui
trung bnh ri dn n vic to thnh Stearic acid (c 18C) l loi acid
bo no ch yu ca cc m. T Stearic acid c th tip tc ko di thm
chui carbon to nn cc acid bo c mch C di hn.
Trc ht t acetyl-CoA v CO2 kt hp vi nhau to nn
malonyl-CoA. Qu trnh ny c xc tc bi acetyl-CoA-carboxylase.
H2O
CH3COCoA

Acetyl-CoA

COOH CH2 CO SCoA

CO2

ATP

ADP

Malonyl - CoA

tin hnh phn ng ngng t gia acetyl-CoA vi malonylCoA cn c s tham gia ca mt loi protein c vai tr vn chuyn nhm
acyl, l protein vn chuyn gc acyl-ACP.
COOH CH2 CO ~ SCoA

Malonyl - CoA

COOH CH2 CO ~ SACP

ACPSH

CoASH

Malonyl - ACP

182
Tip theo acetyl-SACP v malonyl-SACP ngng t vi nhau vi
s xc tc ca enzyme acyl-synthetase. Khi cc phn t acetyl-ACP v
malonyl-ACP tc dng vi enzyme acyl-synthetase s xy ra s chuyn
cc gc acetyl v malonyl t nhm SH ca ACP sang nhm SH ca
enzyme ng thi CO2 c gii phng.
COOH CH2 CO ~ SACP + COOH CO ~ SACP
CH3CO CH2 CO ~ SACP + CO2 + ACP - SH

T CH3COCH2CO ~ SACP (aceto acetyl-ACP) b kh thnh

.hydroxy-butyryl-ACP.
CH3CO CH2 CO ~ SACP
NADH2

CH3CHOHCH2CO ~ SACP

.hydroxy butyryl - ACP

NAD

Phn ng tip theo l .hydroxy butyryl-ACP b kh nc to

nn crotonyl-ACP.
CH3CHOHCH2CO ~ SACP

CH3CH = CHCO ~ SACP

(Crotonyl-ACP)

Crotonyl-ACP b kh to nn butyryl-ACP
CH3CH = CHCO ~ SACP

CH3CH2CH2CO ~ SACP

Butyryl-ACP

NADH2

NAD

T butyryl-ACP tip tc mt chu k mi ngng t vi malonylACP cho ta phnt c 6 nguyn t C. Qu trnh c tip din nh vy
cho n khi to ra s C cn thit ca acid bo, sau Acyl-ACP ny s
bin i tr li thnh Acyl-CoA v cui cng to ra acid bo no bng cch
ct b CoA-SH.

183
Nh vy acid bo c mch cacbon chn vi n nguyn t C th qu
n
trnh s din ra 1 chu k.
2
Nu acid bo c mch C l th trong cc ln ni di mch C ni
trn, ln u tin khng phi phn ng xy ra t 2 Acetyl CoA m xy ra
t Acetyl-CoA v propionyl-CoA to ra acyl-CoA c 5 nguyn t C.
T c mi chu k li ni thm 1 phn t Acetyl-CoA lm cho phn t
acid bo di thm 2 nguyn t cacbon cui cng to nn phn t acid
bo c s nguyn t cacbon l.
Cc acid bo khng no c to ra t cc acid bo no tng ng
bng cch b oxy ha bi FAD.
Acid bo no

Acid bo khng no
FAD

FADH2

10.2.2. Tng hp glycerin

Glycerin c tng hp bng nhiu con ng. Con ng ph
bin l t cc sn phm trung gian ca qu trnh trao i glucose l AlPG
v PDA bin i thnh
AlPG

Glycero-P
NADH2

NAD

PDA

Glycero-P
NADH2

Glycerin + H3PO4

Glycerin +
H3PO4

NAD

10.2.3. Tng hp glyceride

T cc acid bo v glycerin c tng hp s to thnh
glyceride theo cc phn ng sau y:

184
R1-CO-SCoA
Glycerin

Phosphatidic acid + 2CoA-SH

Glycero-P
ATP

ADP

R2-CO-SCoA

R3-CO-SCoA
Photphatidic
acid

Diglyceride

Triglyceride

H3PO4

10.2.4. Tng hp glycero phospholipid

T photphatidic acid s to nn cc loi phospho lipid khc nhau

185

Photphatic acid

CTP
CDP-diacyl-glyceride

Glycero - P

Serine

Phosphatidyl
serine

Phosphatidyl
inosid

Phosphatidyl
Glycero-P

CO2
H3PO4

Phosphatidyl ethanolamine
3CH3

Phosphatidyl
Glycerin

Phosphatidyl - choline

10.2.5. Tng hp sterid

Sterid c to nn bi sterol v acid bo. Nguyn liu tng
hp sterol l acetyl-CoA. Qu trnh sinh tng hp sterol c th chia lm 3
giai on vi nhiu phn ng rt phc tp.
- Giai on chuyn acetyl-CoA thnh mevalonic acid.
- Giai on tng hp squalen.
- Giai on chuyn squalen thnh cholesterol.

186

TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n (ch bin). 1979. Ha sinh i cng (tp I, II). NxB KH&KT.
H Ni.
2. Phm Th Trn Chu, Trn Th ng. 2000. Ha sinh hc. Nxb Gio dc. H
Ni.
3. Nguyn B Lc. 1997. Ha sinh. Nxb Gio dc. H Ni

Ti liu dch
1. Musil J.G., Kurz .K., Novakava .O. 1982
Sinh ha hc hin i theo s . Nxb Y hc. H Ni.

Ti liu ting nc ngoi

1. Farkas G. 1984. Nvnyi anyagcserelettan. Akadmiai Kiad Budapest.
2. Lehninger A. L., 2004. Principle of Biochemistry, 4th Edition. W.H
Freeman.

187

Chng 11

Trao i Protein
11.1. S phn gii protein v amino acid
11.1.1. Phn gii protein
Thy phn l con ng phn gii protein ph bin thc vt v
ng vt. Qu trnh thy phn protein xy ra ti lysosome, ni cha nhiu
enzyme thy phn protein l protease. Qu trnh thy phn xy ra qua 2
giai on
- Nh peptid-peptido hydrolase, protein b thy phn thnh cc on
peptid ngn.
- Nh peptid-hydrolase thy phn tip cc peptid thnh amino acid.
Kt qu chung l
protease

Protein

n.amino acid
+(n-1)H2O

ng vt c v, s phn gii protein u tin do tc ng ca

pepsin. T bo nim mc d dy tit ra pepsinogen. Nh pepsin v HCl
ca dch d dy, pepsinogen bin i thnh pepsin hat ng v pepsin
hat ng s thy phn protein thnh amino acid.
11.1.2. Phn gii amino acid
C nhiu con ng phn gii amino acid.
11.1.2.1. Kh amine
Bng nhiu con ng khc nhau, cc amino acid b kh nhm
amine to ra cc sn phm tng ng.
- Kh amin bng cc enzyme kh. Nh enzyme kh xc tc,
amino acid b kh thnh acid tng ng v gii phng NH3.
Dehydrogenase
R CH COOH
NH
+
+
2
- Kh
amin bng
con ng
oxi
ha.
NADH+H
NAD

R CH2 COOH + NH3

188
Nh amino acid oxydase, amino acid b oxi ha to ceto acid
tng ng v NH3
R CH COOH
+ O2

NH2

oxydase

R CO COOH + NH3

- Kh amine bng con ng thy phn.

Nh tc dng ca enzyme thy phn hydrolase, amino acid b thy
phn to oxiacid tng ng v NH3
hydrolase

R CH COOH
NH2

+ H2O

R CHOHCOOH + NH3

Ngoi cc con ng ra, aspartic acid cn b kh amin bng con

ng kh ni phn t nh enzyme dezaminase xc tc
COOH CH2 CH COOH

Dezaminase
COOH CH = CH - COOH + NH3

NH2
Sn phm ca con ng kh amine cc amino acid l cc loi acid
tng ng v NH3.
11.1.2.2. Kh carboxyl
S loi carboxyl ca amino acid l cch phn gii amino acid rt ph
bin nh decarboxylase xc tc
R CH COOH

Decarboxylase

R CH2 NH2 + CO2

NH2
Sn phm to ra l cc amine, l cc cht c hat tnh sinh hc
cao c vai tr trong qu trnh trao i cht, cc hot ng sinh l ca c
th nh histamine.
11.1.2.3. Chuyn v amine
Bng con ng chuyn v nhm amine sang cho mt cetoacid,
amino acid bin i thnh ceto acid tng ng, phn ng nh enzyme vn
chuyn nhm amin xc tc amino transferase

189

R1 CH COOH

R2 C COOH

NH2

Amino -

R1 C COOH

R2 CH COOH

Transferase

NH2

Phn ng ny thc hin 2 chc nng: va phn gii 1 amino acid

thnh ceto acid, ng thi tng hp mi amino acid khc t ceto acid
tng ng.
Tr threonine v lysine, tt c cc amino acid cn li u c th
tham gia vn chuyn nhm amine bin i thnh cc ceto acid tng
ng, v d:
CH3 CH COOH

+ COOH CH2 CO COOH

NH2
CH3 CO COOH + COOH CH3 CH COOH
NH2

11.1.2.4. S bin i cc sn phm ca qu trnh phn gii amino acid

Cc ng hng phn gii amino acid trnh by trn to ra
nhiu loi sn phm. Cc sn phm ny tip tc c bin i to sn
phm cui cng.
- Cc cht hu c tip tc bin i bng cch oxy ha nh qu trnh
phn gii acid bo to acetyl-CoA, t tham gia vo chu trnh Krebs
phn gii tip.
- Cc amine c bin i thnh cc acid tng ng sau tip tc
bin i nh cc acid khc
R CH2 NH2 + O2

R CH = NH + H2O

R CH = NH + H2O

R CHO + NH3

R CHO
-

+ O2

R COOH

NH3 tip tc bin i bng nhiu con ng gii c cho c th

v NH3 tch ly nhiu s gy c.
+ NH3 c dng lm nguyn liu tng hp tr li amino acid
bng con ng amine ha, amide ha (s trnh by trong phn tng hp
amino acid).

190
+ NH3 b bin i thnh ure qua chu trnh ornithine thi ra ngoi
qua con ng nc tiu ng vt. Chu trnh ornithine chia lm 3 giai
on
1) Tng hp carbamyl-phosphate
CO2 + NH3 + ATP

O=C

NH2

+ ADP

O~P
Phn ng c xc tc bi enzyme carbamyl phosphate synthetase.
2) Tng hp arginine.
T carbanyl-phosphate v ornithine s to thnh citrullin bng mt
phn ng ngng t. Sau citrullin kt hp vi aspactic acid nh
arginino-succinic-synthetase to arginino-succinic acid. Tip theo
arginino-succinic acid b phn gii thnh arginine v fumaric acid nh
arginino-succinate-ligase.
3)
Arginine b phn gii nh arginase to ornithine v ure. Ure
c thi ra ngoi cn ornithine tip tc tham gia vo chu trnh mi.
Trn y l nhng ng hng chung ca s phn gii amino acid.
Tuy nhin mi amino acid u c con ng phn gii ring. Cc amino
acid bin i theo cc ng hng trn u dn n vic to nn cc sn
phm tham gia vo chu trnh Krebs phn gii thnh CO2 v H2O.

11.2. Tng hp amino acid

11.2.1. Amine ha
Mt s acid bo khng no v ceto acid c th amine ha to nn
amino acid tng ng
COOH CH = CH COOH + NH3

Fumaric acid
COOH CH2 CH2 CO COOH + NH3

-cetoglutaric acid
COOH CH2 CO COOH + NH3

oxaloacetic acid

COOH CH CH2 COOH

NH2

aspartic acid
COOH CH2 CH2 CH COOH
NH2

glutamic acid

COOH CH2 CH COOH

NH2

aspartic acid

191
V nguyn tc, mi amino acid u c th c tng hp bng con
ng ny t cc acid tng ng. Nhng trong t bo ch c 2 enzyme l
glutamate dehydrogenase v pyruvate dehydrogenase c hot mnh
thc hin xc tc loi phn ng trn, cn cc enzyme khc khng c kh
nng xc tc cho nn trong thc t ch c glutamic acid v alanin l 2
amino acid c tng hp bng con ng ny.
11.2.2. Amide ha
T 2 loi amino acid l aspactic acid v glutamic acid do c 2 nhm
carboxyl nn c th c amide ha to amino acid mi, dng amide
ca aspactic acid v glutamic acid l asparagine v glutamine
COOH CH2 CH COOH
NH2

O = C CH2 CH COOH

+ NH3

NH2

NH2

Asparagine

Aspartic acid
COOH CH2 CH2 CH COOH
NH2

+ NH3

O = C CH2 CH2 CH COOH

NH2

NH2

Glutamine

Glutamic acid
11.2.3. Tng hp amino acid nh ATP
Qu trnh tng hp amino acid nh ATP xy ra qua 2 giai on
- NH3 + ATP AMP ~ NH2 + P - P
y l phn ng hat ha nhm NH2 nh ATP. AMP ~ NH2 thc
hin phn ng chuyn v amine cho ceto acid to amino acid tng ng
AMP ~ NH2 +

R C COOH
O

AMP +

R CH COOH
NH2

Thc cht ng hng ny cng l hnh thc amine ha cc ceto

acid nhng khng s dng cc dehydrogenase m s dng ATP.

192
11.2.4. Chuyn v amine
Nh trnh by trn (Mc 11.1.2.3) amino acid c th b phn
gii bng con ng chuyn v amine ng thi vi vic tng hp mt
amino acid khc.
Nh qu trnh ny m thnh phn cc amino acid lun c i
mi ph hp vi nhu cu ca c th trong qu trnh sng.
11.2.5. Oxim ha
mt s vi sinh vt v thc vt c kh nng c nh Nit t do
qu trnh c nh m. Qua qu trnh c nh N2, NH2OH c hnh thnh
lm nguyn liu cho qu trnh tng hp amino acid theo cch oxim ha.
Ngoi ra mt s vi sinh vt v thc vt cn c qu trnh kh
nitrat (NO3-) thnh ammoniac (NH3). Trong qu trnh bin i theo ng
hng ny NH2OH c to thnh trc khi to NH3. NH2OH lm
nguyn liu tng hp amino acid bng cch oxim ha.
Qu trnh oxim ha xy ra qua 2 giai on
- Cc ceto acid kt hp vi NH2OH to nn oxim tng ng
Oximase

R C COOH
+ NH2OH

R C COOH
NOH
Oxim

Ceto acid

+ H2O

- Cc oxim b kh to amino acid tng ng

Oxim dehydrogenase
R CH COOH

R C COOH
NOH

NADH+H+

NAD+

NH2

+ H2O

vi sinh vt v thc vt, y l con ng tng hp amino acid

quan trng.

11.3. Tng hp protein

Qu trnh tng hp protein l vn quan trng ca sinh hc phn
t. Qu trnh xy ra phc tp vi s tham gia ca nhiu thnh phn.

193
11.3.1. Cc thnh phn tham gia tng hp protein
11.3.1.1. Nucleic acid
Tham gia vo qu trnh tng hp protein c cc loi nucleic acid vi
cc chc nng khc nhau
- DNA: mang thng tin v cu trc phn t protein theo dng m
ha. Mi protein c m ha trn 1 on DNA, l gen.
- RNAm: lm nhim v truyn thng tin v cu trc phn t protein
t gen sang chui polypeptide.
- RNAt: lm nhim v vn chuyn cc amino acid t cc vng trong
t bo n ribosome tin hnh tng hp chui polypeptide ti . ng
thi nhn bit v tr b ba m ha amino acid trn RNAm t amino acid
vo ng v tr ca n trn chui polypeptide.
- RNAr: cng vi protein, RNAr cu to nn ribosome, ni thc hin
qu trnh tng hp chui polypeptide.
11.3.1.2. Cc enzyme
Tham gia xc tc qu trnh tng hp protein, c nhiu loi enzyme
- Aminoacyl-adenilat-synthetase l enzyme xc tc qu trnh hat
ha amino acid, phn ng gn amino acid vo RNAt.
- Transpeptidase: xc tc phn ng to lin kt peptide ni cc
amino acid li thnh chui polypeptide v chuyn dch chui polypeptide
trong ribosome t v tr P sang v tr A.
- Translocase: l enzyme xc tc qu trnh di chuyn ca ribosome
trn RNAm.
Ngoi cc enzyme chnh ny cn c enzyme ct amino acid m u
ra khi chui polypeptide, enzyme xc tc s to cc cu trc khng gian
ca protein
11.3.1.3. Nng lng
Qu trnh tng hp protein cn nng lng. Nng lng cung cp
cho qu trnh ny l ATP v GTP.
- ATP cung cp nng lng cho giai on hat ha amino acid.
- GTP cung cp nng lng cho giai on tng hp chui
polypeptide ribosome.
11.3.1.4. Nguyn liu
Nguyn liu tng hp protein l cc amino acid.
Trong s cc amino acid c loi amino acid m u l methionine
Eucariote v formyl methionine Procariote.

194
11.3.1.5. Ribosome
Ribosome l ni tin hnh tng hp chui polypeptide. Thnh
phn ribosome gm protein v RNAr. Cu trc ribosome gm 2 tiu th:
tiu th ln v tiu th b. Trong ribosome c 2 vng hat ng: vng A l
ni tip nhn cc amino acid mi cn vng P l ni to nn chui
polypeptide. tiu th b cha mt loi RNAr, trn phn t RNAr ny c
1 on c thnh phn cc nucleotide tng ng b sung vi on khng
m ha trn RNAm. Nh khi bt u qu trnh tng hp, RNAm n gn
vo ribosome v t ng b ba m u ca n vo v tr P nh s lin kt
gia on khng m ha trn RNAm vi on b sung trn RNAr.
11.3.1.6. Cc yu t tham gia tng hp protein
* Yu t m u. l nhng phn t protein vi chc nng tham
gia vo vic kch thch s m u trong qu trnh tng hp chui
poplypeptide.
Procariote

Eucariote

Yu t

Chc nng

Yu t

Chc nng

IF-1

Kch thch hat ng ca

IF2, IF3

eIF-1

Gn vi RNAm

IF-2

Lm d dng qu trnh kt
hp f.Met-RNAt vi tiu
th b 30S

eIF-2

Lm d dng s kt hp
Met-RNAt vi tiu th
b 40S

IF-3

Gn vi tiu th b 30S,
ngn khng kt hp vi
tiu th ln 50S

eIF-3

Kt hp vi tiu th b
40S

CBP-1

Kt hp vi m ca
RNAm

eIF-4a

Kt hp vi RNAm

eIF-5

Tch ri cc yu t khi
u khi 40S v kt hp
vi 60S

eIF-6

Tch ribosome
thnh 2 tiu th.

* Yu t ko di
Tham gia vo giai on ko di c cc yu t:

80S

195
- EF-Tu gip cho RNAtAa n gn vo v tr A ca ribosome.
- EF-Ts gip s gii phng GDP khi phc EF-Tu-GDP.
- EF-G xc tc s di chuyn ca ribosome trn RNAm theo chiu 5-3.
11.3.2. Tng hp chui polypeptide ti ribosome
11.3.2.1. Giai on hat ha amino acid
tham gia vo qu trnh tng hp protein cc amino acid phi
c hat ha v gn vo RNAt. Qu trnh ny xy ra hai phn ng, c
xc tc bi enzyme aminoacyl-adenylat-synthetase
[AMP ~ amino acid] E + P-P
Trong phn ng th nht ny amino acid kt hp vi ATP to ra
amino acid-AMP v gii phng pyrophosphat (P-P). Aminoacid-AMP
khng trng thi t do m gn vi enzyme to phc linh ng
Enzyme
Amino acid + ATP
[AMP ~ Amino acid] E + P-P
RNAt mang amino acid s di chuyn n ribosome thc hin qu
trnh tng hp chui polypeptide .
11.3.2.2. Giai on m u
Tham gia vo giai on m u c cc yu t m u. procariote
yu t m u l IF-1, IF-2, IF-3, cn Eucariote yu t m u l eIF-1,
eIF-2, eIF-3. Nng lng cung cp cho giai on m u l GTP. c bit
thc hin giai on tng hp nn amino acid m u cn c RNAt
mang amino acid m u.
procariote amino acid m u l mt loi methionine b bin
i thnh dng formyl methionine.
H2N CH COOH

CHO NH CH COOH
Formyl ha

(CH2)2

(CH2)2

CH3
Methionine

CH3

Formyl Methionine

Tham gia vn chuyn formyl methionine v methionine l 2 loi

RNAt c cng b ba i m l UAC tng ng b sung vi m m u

196
AUG trn RNAm. Nh vy, RNAt mang formyl methionine vo m u
qu trnh tng hp chui polypeptide, cn vic vn chuyn methionine
a vo thnh phn chui polypeptide ch xy ra khi trn RNAm c b ba
m ha methionine AUG.
Eucariote amino acid m u l methionine nn RNAt vn chuyn
methionine va lm nhim v m u nu gii m cho b ba m u AUG,
va lm nhim v a methionine vo tham gia thnh phn chui polypeptide
nu gii m cho b ba AUG nm cc v tr khc v tr m u.
Giai on m u c thc hin bi s tch ribosome thnh 2 tiu
n v ( procariote l 50S v 30S, cn Eucariote tng ng l 60S v
40S). Tip theo tiu n v b lin kt vi yu t m u IF3 to phc I
(IF3 -30S). ng thi RNAt mang amino acid m u (f.Met hay Met) gn
vi GTP, yu t m u IF2 to phc th II l (RNAtGTP-IF2). Tip theo
phc I v phc II kt hp li vi nhau ng thi RNAm n gn vo tiu
th b ca t hp trn. on khng m ha trn RNAm gn b sung vi
mt on trn RNAr ca tiu th b nh t b ba m u ca RNAm
vo ng v tr P ca tiu th ln khi tiu th b n gn vo phc trn.
Cui cng t hp (30S-IF3-RNAt-GTP-IF2-RNAm) gn vo tiu th ln,
khi phc li ribosome v gii phng cc yu t m u, GDP v H3PO4.
Kt qu ca giai on ny l to nn phc m u, trong RNAt
amino acid m u gn vo m m u ca RNAm nm v tr P ca
ribosome

197

Ribosome
80S khng
hot ng

RNAm

S ca giai on m u chui polypeptide Eucariota

11.3.2.3. Giai on ko di chui polypeptide
Sau khi phc m u c to nn, qu trnh tng hp chui
polypeptide bt u tin hnh. Tham gia giai on ko di chui c cc
yu t ko di (EF procariote, eEF Eucariote) GTP cung cp nng
lng, cc enzyme, cc amino acid-RNAt v phc h m u.
Qu trnh ko di chui polypeptide xy ra theo trt t cc b ba trn
RNAm k t sau b ba m u, theo chiu 5-3. ng vi cc b ba cc
RNAt tng ng mang cc amino acid ca n trong phc hp aminoacidRNAt n gn ng v tr bng cch nhn bit gia b ba m ha ca
RNAm vi b ba i m ca RNAt theo nguyn l b sung. Bng cch
t ng v tr cc amino acid trn chui polypeptide.
M u giai on ko di chui, amino acid-RNAt mang amino acid
u tin n gn vo v tr A ca ribosome ang b trng nh to lin kt
b sung gia b ba m ha trn RNAm vi b ba i m ca RNAt.
Sau khi phc hp amino acid-RNAt gn vo v tr A ca ribosome,
amino acid m u v tr P c tch khi RNAt ca n v chuyn sang
v tr A lin kt vi amino acid bng lin kt peptid. Qu trnh
c xc tc bi peptidyl Transferase. Nh vy v tr P ch cn RNAt
khng mang amino acid cn v tr A c RNAt mang 2 amino acid. Bc

198
tip theo l nh locaferase xc tc ribosome trt trn RNAm theo chiu
5-3 mt on 3 nucleotide. Kt qu l t hp RNAt mang 2 amino acid
chuyn sang v tr P cn v tr A ca ribosome li b trng nh phc h
m u v kt thc vic ni di thm 1 amino acid vo chui polypeptide.
Cc amino acid tip theo vo ni di chui cng c tin hnh qua
cc bc nh vi amino acid th nht trn. Th t cc b ba trn RNAm
quy nh trnh t cc amino acid tng ng vo tham gia qu trnh ni di
chui polypeptide. Nh vy trt t cc b ba trn RNAm quyt nh trt t
cc amino acid trn chui polypeptide.
RNAtAla + GTP+ EF

Trans peptidase

Trans locase
RNAt m u

S giai on ko di chui
11.3.2.4. Giai on kt thc s tng hp chui polypeptide
Qu trnh ko di chui s ngng khi gp tn hiu kt thc. Tn hiu
kt thc l b ba kt thc. Khi b ba kt thc ca RNAm (trn mt RNAm
c 1 trong 3 b ba UAG, UGA v UAA) nm vo v tr A ca ribosome.
S xut hin 1 trong 3 b ba trn, qu trnh tng hp chui polypeptide

199
c kt thc do cc b ba ny khng m ha amino acid nn qu trnh
ko di chui b ngt qung, chui polypeptide c tng hp b tch
khi RNAt cui cng m khng c RNAt tip gn vo nn c gii
phng ra khi ribosome v kt thc qu trnh tng hp. Tham gia vo qu
trnh kt thc c yu t gii phng RF lm nhim v nhn bit m kt thc
v gii phng chui polypeptide ra khi ribosome.
11.3.3. Hon thin phn t protein
Chui polypeptide c tng hp ti ribosome phi qua nhiu bin
i mi tr thnh phn t protein hon thin. Trc ht methionine u
chui b ct b nh peptidase xc tc. Sau t cc nhm chc trn cc
amino acid ca chui hnh thnh cc lin kt ni phn t to nn protein
c cc mc cu trc khc nhau. Trc ht t chui polypeptide lin kt
hydro c hnh thnh t cc nhm CO v NH ca cc amino acid to
nn cu trc bc II ca protein. T protein bc II cc lin kt disulfua, lin
kt ion, lin kt k nc to ra lm cho phn t protein c cu trc bc II
cun xon to nn phn t protein bc III. T mt s phn t protein
bc III cng chc nng c th lin kt vi nhau bng lin kt hydrogen,
tng tc Van der Waals to nn protein c cu trc bc IV.
Cc phn t protein c hon thin s c a n cc ni s
dng thc hin chc nng ca chng trong t bo.
11.3.4. iu ha tng hp protein
Qu trnh tng hp protein xy ra trong t bo c iu ha ph
hp vi nhu cu ca c th. Khi c th cn loi protein no th qu trnh
iu ha t iu chnh cho qu trnh tng hp protein xy ra, ngc li
khi khng cn mt loi protein no na th qu trnh tng hp protein
b c ch.
Cng trnh c gi tr u tin v c ch iu ha tng hp protein do
J.Monod v Jacob (1956) xut. Cc tc gi ny a ra thuyt Operon
gii thch c ch iu ha tng hp protein. Theo thuyt operon phn t
DNA cha nhiu loi gen:
- Gen cu trc-cistron,structural gene (S): gen ny m ha phn t
protein. Mi operon c th c nhiu gen cu trc m ha cho mt nhm
protein c chc nng lin quan nhau nh cc enzyme xc tc mt chui
phn ng.
- Gen tc ng operator (O).
- Gen khi ng promotor (P).
Bn cnh mi operon c gen iu ha Regulator (R) vai tr iu ha
hot ng ca operon, quyt nh s ng hay m ca operon.

200
Trt t cc gen trong operon nh sau
R

S1

S2

S3

S4

Mi operon chu trch nhim iu ha s tng hp mt nhm

protein-enzyme cng tham gia xc tc mt chui phn ng, trong mi
protein-enzyme do mt gen cu trc m ha.
C nhiu hnh thc iu ha tng hp protein theo operon nh iu
ha m tnh, iu ha dng tnh. Trong mi loi trn li c nhiu hnh
thc khc nhau nh iu ha bng cch cm ng, iu ha bng cch c
ch.
11.3.4.1. iu ha m tnh
iu ha m tnh l c ch iu ha m khi khng c phc h c
ch bm vo operon th operon m v tng hp protein xy ra. C 2 hnh
thc iu ha theo dng ny l iu ha cm ng m tnh v iu ha c
ch m tnh.
* iu ha cm ng m tnh. in hnh ca dng iu ha ny l
hot ng iu ha ca operon lac. Bn cht ca dng iu ha ny l khi
trong mi trng c cht cm ng th operon m , qu trnh tng hp
protein xy ra. Cht cm ng y l c cht ca cc enzyme do operon
kim sot, l lactose. Khi trong mi trng c lactose th operon m v
qu trnh tng hp nn cc enzyme tham gia phn gii lactose xy ra.
Ngc li khi trong mi trng khng c lactose th operon ng v qu
trnh tng hp protein khng xy ra. C ch qu trnh iu ha xy ra nh
sau: operon lac c gen khi ng, gen tc ng v 3 gen cu trc Z.Y.A
trong
- Gen Z: m ha enzyme .galactosidase.
- Gen Y: m ha enzyme permease .
- Gen A: m ha enzyme transacetylase.
Ba enzyme trn tham gia qu trnh phn gii lactose.
Gen iu ha tng hp protein c ch. Cht c ch ny nu trng
thi t do s c i lc vi operon v bm vo operon lm cho operon b
ng. Cn khi c cht cm ng do cht cm ng c i lc vi cht c ch
mnh hn operon nn lin kt vi cht c ch to phc khng hat ng.
Phc ny khng bm c vo operon nn operon m.
Nh vy trong t bo c lactose lm cht cm ng lin kt vi cht
c ch do gen iu ha tng hp ra lm cho cht c ch khng bm vo
operon, gen khi ng hat ng kch hot RNA polymerase. RNA

201
polymerase s xc tc cho qu trnh sao m cc gen cu trc thnh cc
RNAm, t cc RNAm s tng hp ra 3 loi enzyme trn. Ba enzyme xc
tc cho qu trnh phn gii lactose.
Khi lactose b phn gii ht, khng cn cht cm ng lin kt vi
cht c ch nn cht c ch bm vo operon, c ch hot ng ca RNA
polymerase, RNA polymerase khng hat ng sao m c nn qu
trnh tng hp protein cng khng xy ra.

RNAm

RNAm

* iu ha c ch m tnh. in hnh ca dng iu ha ny l hat

ng iu ha ca operon Tryp. Bn cht ca dng iu ha ny l khi
trong mi trng c cht ng c ch th operon ng, qu trnh tng hp
protein khng xy ra. Cn khi khng c cht ng c ch th operon m,
qu trnh tng hp protein xy ra. Cht ng c ch y l sn phm ca
cc enzyme do operon kim sot, l Tryptophan. Khi trong mi
trng khng c Tryptophan, operon m, qu trnh tng hp cc enzyme
tham gia tng hp Tryptophan xy ra. Ngc li, khi trong mi trng c
Tryptophan, operon ng, qu trnh tng hp cc enzyme khng xy ra.
C ch iu ha xy ra nh sau.
Operon Tryp c 5 gen cu trc m ha cho 5 enzyme tham gia qu
trnh tng hp Triptophan trong t bo: Tryp E, Tryp D, Tryp C, Tryp B
v Tryp A.
Gen iu ha tng hp protein c ch. Khi trong t bo c
Tryptophan ng vai tr cht ng c ch th cht c ch kt hp vi cht
ng c ch to phc hat ng. Phc ny c i lc vi operon nn bm
vo operon lm cho operon khng hot ng, RNA-polymerase khng sao

202
m nn khng c cc RNAm cho qu trnh tng hp protein, tng hp
protein-enzyme khng xy ra.
Khi trong t bo khng c Triptophan, cht c ch do gen iu ha
tng hp ra khng c i lc vi operon nn khng bm vo operon, RNApolymerase khng b c ch nn xc tc cho qu trnh sao m t cc gen
cu trc to ra cc RNAm tng ng. T cc RNAm tin hnh qu trnh
tng hp protein to nn cc protein-enzyme.

RNAm

RNAm

11.3.4.2. iu ha dng tnh

iu ha dng tnh l c ch iu ha tng hp protein m khi c
phc h c ch bm vo operon th operon m, qu trnh tng hp protein
xy ra. Khi khng c phc h c ch bm vo operon th operon ng, qu
trnh tng hp protein dng li. C bn ca qu trnh ny l c c ch iu
ha ngc hnh thc iu ha m tnh phn tch trn.

203

TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n (ch bin). 1979. Ha sinh i cng (tp I, II). NxB KH&KT.
H Ni.
2. Phm Th Trn Chu, Trn Th ng. 2000. Ha sinh hc. Nxb Gio dc. H
Ni.
3. Nguyn B Lc. 1997. Ha sinh. Nxb Gio dc. H Ni

Ti liu dch
1. Musil J.G., Kurz .K., Novakava .O. 1982
Sinh ha hc hin i theo s . Nxb Y hc. H Ni.

Ti liu ting nc ngoi

1. Farkas G. 1984. Nvnyi anyagcserelettan. Akadmiai Kiad Budapest.
2. Lehninger A. L., 2004. Principle of Biochemistry, 4th Edition. W.H
Freeman.

204

Chng 12

Trao i nucleic acid

12.1. S phn gii nucleic acid
12.1.1. Thy phn nucleic acid
S thy phn nucleic acid thnh mononucleotide c xc tc bi
cc enzmie thy phn tng ng.
DNA nh desoxyribonuclease xc tc bin i thnh cc
desoxyribonucleotide cn RNA do cc ribonuclease xc tc s b phn gii
thnh cc ribonucleotide.
12.1.2. Phn gii mononucleotide
Mononucleotide b phn gii bi tc dng ca cc phosphatase hoc
nucleotidase to nn cc nucleoside v H3PO4. Cc nucleoside li tip tc
b thy phn bi cc nucleosidase to base nit v pentose.
Cc sn phm ca qu trnh phn gii trn tip tc bin i
- H3PO4 tham gia vo cc qu trnh trao i saccharide hay cc qu
trnh trao i cht khc.
- Base Nit tip tc b phn gii to cc sn phm tham gia vo qu
trnh trao i cht ca t bo.
12.1.3. Phn gii base purine
Adenine v guanine bin i thnh xanthine, t xanthine qua mt s
phn ng tip theo to sn phm cui cng l ure v glyoxylic acid.
(1)

Adenine

(2)

Xanthin

Guanine

(3)

Allantoic acid
(4)

Ure +glyoxylic acid

Phn ng 1 v 2 do enzime desaminase xc tc, phn ng 3 do
xanthineoxydase v phn ng 4 do allantoicase xc tc.

205

12.1.4. Phn gii base pyrimidine

Cc base pyrimidine b phn gii to nn sn phm cui cng l
NH3, CO2, .amino isobutyric acid v alanine.
Cytosine

Uracil
H2O

Thymine

NH3

Dihydro Uracil
H2O + alanine

NADPH2 NADP
Dihydro Thymine

CO2 + NH3

NH3+CO2+ .aminoisobutyric acid

H2O

NADPH2 NADP

CO2, NH3 to ra trong cc qu trnh bin i trn c thi ra ngoi,

cn alanine v .aminoisobutyric acid tip tc bin i nh cc amino
acid khc.

12.2. Sinh tng hp nucleotide purine

Gc purine c to ra t nhiu thnh phn khc nhau: CO2,
aspartic acid, glycine, formate, glutamine.
CO2

aspartic acid

formate

glycine
formate

N-H

glutamine

Trong qu trnh tng hp khung purine s xy ra ng thi c qu

trnh tng hp nucleotide. Tm tt kt qu qu trnh nh sau:
Riboso5P + 2.glutamine + glycine + CO2 + 2 formate +
aspactic acid + H2O inosinic acid
T inosinic acid s to nn GMP v AMP
- Inozinic acid + aspactic acid + GTP AMP + fumaric acid GDP + Pv
- Inozinic acid + NAD + ATP + NH3 GMP + NADH2 + AMP + Pv

206

Ngoi ra, cc nucleotide purine cn c th c tng hp trc tip t

base purine v phosphoriboso-pyrophosphate (PRPP)
Adenine + PRPP AMP + P-P
Guanine + PRPP GMP + P-P

12.3. Sinh tng hp nucleotide pyrimidine

Khung pyrimidine c to ra t NH3, CO2 v aspartic acid

NH3

Asparic acid
CO2
N

Qu trnh tng hp nucleotide pyrimidine xy ra qua cc giai on sau:

CO2 + NH3 + ATP carbamyl-P
Carbamyl-P + Aspactic acid Orotic acid
Orotic acid + Riboso5P UMP CMP TMP.
Cc nucleotide pyrimidine cn c tng hp trc tip t base nit
pyrimidine vi PRPP
Uracil + PRPP UMP + P-P
Thymine + PRPP TMP + P-P
Cytosine + PRPP CMP + P-P

12.4. Tng hp DNA

Qu trnh tng hp DNA, hay cn gi l s ti bn, c ngha rt
quan trng trong i sng c th lin quan n c ch di truyn. y l
mt qu trnh phc tp c s tham gia ca nhiu yu t v xy ra nhiu
hnh thc.
C th chia qu trnh ti bn DNA thnh 3 kiu
- Ti bn bo th. L qu trnh tng hp DNA t 1 phn t DNA
gc to ra 2 phn t DNA con, trong c 1 phn t chnh l phn t
DNA gc cn 1 phn t c tng hp mi hon ton.

207

- Ti bn gin on. L qu trnh tng hp DNA t 1 phn t DNA

gc to ra 2 phn t DNA con c cc on mi tng hp v cc on c
ca DNA gc xen k.
Hai hnh thc ti bn trn t ph bin.
- Ti bn bn bo th. y l hnh thc tng hp DNA t 1 phn t
DNA gc to ra 2 phn t DNA con, trong mi phn t DNA con mt
chui ly t DNA gc v mt chui mi tng hp. Hnh thc ny c
Meselson v Stahl pht hin nm 1958 bng thc nghim nui cy E.coli.
Trc ht E.coli c nui cy trong mi trng ch cha 15N (Nit nng)
nn DNA c tng hp nn ch cha 15N s to nn phn t DNA c t
trng cao hn DNA thng. Sau chuyn E.coli vo mi trng cha
14
N (Nit thng) v theo di, phn tch cc th h DNA mi c to ra
bng phng php li tm phn on vi CSCl. Qua kt qu phn tch li
tm cho thy th h th nht 100% phn t DNA dng lai, mt chui
cha 15N v 1 chui cha 14N. th h th 2 c 50% dng lai v xut hin
50% dng 14N. iu chng t c ch ti bn DNA l dng bn bo th.
12.4.1. Cc yu t tham gia ti bn DNA
- DNA khun. tng hp phn t DNA mi cn c phn t DNA
lm khun. DNA va lm chc nng khun va tham gia trong sn phm
ca qu trnh tng hp.
- Nguyn liu. tng hp DNA mi cn c cc nguyn liu.
Nguyn liu l cc desoxy Ribonucleotide Triphosphate (dATP, dGTP,
dCTP, dTNP) dTMP va lm nguyn liu va cung cp nng lng cho
qu trnh tng hp DNA mi.
- Enzyme. Tham gia xc tc qu trnh ti bn DNA c nhiu loi
enzyme
+ DNA-polymerase,
+ Topoisomerase,
+ Helicase,
+ DNA-ligase.
- Protein. C nhiu loi protein tham gia vo qu trnh ti bn DNA
vi chc nng h tr, kch thch
+ Protein bm si n SBS,
+ Protein DnaA,
+ Protein DnaB,
+ Protein DnaG.

208

12.4.2. C ch ti bn DNA procariote

12.4.2.1. Giai on m u
- Protein DnaB lm nhim v m xon DNA bng cch phn hy cc
lin kt hydrogen gia 2 chui, tch 2 chui ra to nn chc ti bn.
- Protein SBS n gn vo chc ti bn.
- Primase xc tc s to RNA mi b sung vo chui khun 3-5.
12.4.2.2. Giai on ko di
* Tng hp chui sm
- Trn chui khun 3-5 sau khi to on RNA mi, cc nucleotide
tip tc n gn vo u 3-OH ca chui theo nguyn tc b sung vi
chui lm khun nh enzyme DNA-polymerace III xc tc.
- Chui sm c tng hp lin tc, tho xon n u cc
nucleotide t do trong mi trng t bo tng ng b sung vi cc
nucleotide trn chui khun ln lt n gn vo u 3-OH bng cch to
lin kt phosphodiester vi nucleotide cui cng u 3. ng thi
pirophosphate c tch ra.
* Tng hp chui mun
Trn chui khun 3-5 ca DNA khun, chiu tho xon v chiu
tng hp ngc nhau nn qu trnh tng hp khng din ra lin tc m to
ra cc on okazaki ngc chiu vi chiu pht trin ca chc ti bn.
Mi on okazaki c RNA mi ring c tng hp nh primase.
Mi c tng hp b sung vi chui khun 5-3 v ngc chiu tho
xon. Qu trnh tho xon xy ra c mt on khong 300 nucleotide
mi tng hp RNA mi theo chiu ngc li.
Xc tc cho qu trnh tng hp chui mun l phc hp protein c
tn l primosom. Primosom di chuyn trn chui khun 5-3 v tin hnh
tng hp on RNA mi nh primase sau tng hp tip on DNA b
sung vo chui khun nh DNA-polymerase to nn on Okazaki.
Sau khi on Okazaki hon chnh, RNA mi c tch ra nh DNApolymerase I sau thay vo v tr on RNA mi l on DNA tng
ng. Sau cng nh DNA-ligase ni 2 on Okazaki li vi nhau.
12.4.2.3. Giai on kt thc
Qu trnh ko di c tip din cho n khi ht phn t DNA khun.
Kt qu t 1 phn t DNA khun to ra 2 phn t DNA mi, trong mi
phn t DNA mi c 1 chui mi c tng hp t cc nucleotide trong
mi trng, cn mt chui l ca DNA khun.

209

12.4.3. Ti bn DNA Eucariote

Eucariote qu trnh ti bn DNA c bn ging procariote nhng
cng c mt s c trng ring.
- Trn mt phn t DNA khun qu trnh ti bn xy ra ng thi
nhiu im.
- Vn tc ti bn chm hn procariote
+ procariote vn tc 500 nucleotide/S.
+ Eucariote vn tc 50 nucleotide/S.
- Mt s enzyme khc procariote
+ DNA polymerase ,
+ DNA polymerase ,
+ DNA polymerase ,
+ DNA polymerase .

12.5. Tng hp RNA (sao m)

12.5.1. Cc yu t tham gia tng hp RNA
12.5.1.1. Khun
tng hp RNA cn c khun. Khun c th l DNA, cng c th
l RNA.
phn ln cc sinh vt RNA c tng hp t DNA, do DNA lm
khun. Phn t DNA lm khun ch s dng 1 on, tng ng 1 gen
tng hp nn 1 phn t RNA. Nh vy t 1 phn t DNA c th tng hp
ra nhiu RNA. ng thi trn 2 chui ca DNA, ch s dng chui 3-5
lm khun.
12.5.1.2. Nguyn liu
Cng nh tng hp DNA, trong qu trnh tng hp RNA cn cc
Ribonucleotide-Triphosphat lm nguyn liu v ngun nng lng.
12.5.1.3. Cc enzim v protein
* RNA-polymerase. C 2 loi RNA-polymerase, mt loi xc tc
qu trnh tng hp RNA t DNA mt loi xc tc qu trnh tng hp RNA
t RNA.
procariote RNA-polymerase c cu to phc tp. Phn t RNApolymerase gm 5 tiu n v , , , ,

210

Tiu n v

S lng

Chc nng

40.000

Nhn bit v tr m u

155.000

To lin kt P-diester

165.000

Gn DNA khun

95.000

Cha r

95.000

Nhn bit chui lm khun v

im m u

* Yu t Rho (): Rho l mt loi protein tham gia vo qu trnh kt

thc tng hp RNA.
12.5.2. C ch sao m
12.5.2.1. Giai on m u
Bc vo giai on m u RNA-polymerase tch yu t ra khi
enzyme.
Li enzyme tin hnh m xon DNA.
Yu t nhn bit bit chui lm khun v im m u nh cc tn
hiu trn promotor.
Hai chui DNA tch ra 1 on 30 nucleotide to nn vng sao m.
Chui n ca DNA (chui 3-5) nhn 1 nucleotide gn b sung
vo nucleotide m u trn DNA. Tip theo nucleotide th 2 n gn vi
nucleotide u bng lin kt phosphodiester v to lin kt b sung vi
nucleotide trn chui khun. Sau khi lin kt phosphodiester u tin ny
c to ra, yu t tch khi vng sao m v kt thc giai on m u.
12.5.2.2. Giai on ko di chui
Nh li enzyme cc nucleotide trong mi trng n ko di chui
theo nguyn tc b sung vi cc nucleotide trn chui khun DNA.
Qu trnh ko di chui xy ra rt phc tp gm nhiu phn ng lin
hon to ra s n nh ca vng m xon. Qu trnh xy ra theo trnh t sau:
- Tho xon trn DNA khun u 3 chui khun.
- Ko di thm 1 nucleotide trn chui RNA.
- Tho xon kp lai DNA-RNA u 5.
- ng xon trn DNA khun u 5.

211

Qu trnh c din ra theo chu k nh li enzyme xc tc cho n khi

gp tn hiu kt thc.
12.5.2.3. Giai on kt thc
C 2 kiu kt thc: kt thc nh yu t Rho v kt thc khng nh
yu t Rho.
- Kt thc nh yu t Rho: trn b mt ca mt s v tr kt thc c
loi protein Rho. Rho di chuyn trn RNA mi c tng hp v i ti
vng sao m, Rho lm nhim v tch xon lai DNA-RNA, gii phng
RNA v kt thc qu trnh sao m.
- Kt thc khng cn yu t Rho: Trn RNA c 1 on c cu trc
ngc chiu (palindrome) khi sao m to ra vng palindrome, vng ny s
to lin kt kp hnh thnh cu trc ci kp tc nn lm ngng qu trnh
sao m.
12.5.3. Qu trnh trng thnh ca RNA
Phn t RNA c sao t DNA l proRNA. T proRNA phi qua
qu trnh bin i phc tp mi to RNAm.
12.5.3.1. Gn m vo u 5
ProRNA cha c m nn trc ht cn gn thm m vo u 5 ca
Pro-RNA. M c tng hp sn trong nhn. M c gn vo u 5
bng lin kt anhydric acid vi nhm Triphosphate ca ProRNA ch
khng gn vo u 3 nh qu trnh ko di chui.
12.5.3.2. Gn ui vo u 3
Cng nh m, ui ca RNAm khng m ha trong gen m c
tng hp ring trong nhn. ProRNA cha c ui. ui c ni vo vi
ProRNA u 3 nh polyA-polymerase.
12.5.3.3. Ct b cc on Intron trn proRNA.
Trn Pro-RNA c cc on khng m ha amin acid (Intron I) cho
nn to ra RNAm cn ct b cc on I v ni cc on m ha (ExonE) li.
tn hiu di truyn c truyn t chnh xc, s ct ni cn c
chnh xc cao v ch cn ct sai 1 nucleotide s lm thay i ton b cc
m di truyn pha sau v tr ct.
Gia cc on E v I c cc trnh t nucleotide c trng ging nhau
mi pro-RNA.
- u 3 ca E pha trc lun l AG,
- u 5ca E pha sau lun l G,

212

- u 5 ca I lun l GU,
- u 3 ca I lun l G.
Trong Intron c mt on c vai tr quan trng trong c ch ct ni
ca pro-RNA. l v tr tch nhnh. Qua v tr ny, di tc ng ca
enzyme ct. Cc Intron b ct b ra v cc Intron ni li vi nhau.
Kt qu ca qu trnh bin i trn to nn phn t RNAm trng
thnh tham gia vo qu trnh dch m.

TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n (ch bin). 1979. Ha sinh i cng (tp I, II). NxB KH&KT.
H Ni.
2. Phm Th Trn Chu, Trn Th ng. 2000. Ha sinh hc. Nxb Gio dc. H
Ni.
3. Nguyn B Lc. 1997. Ha sinh. Nxb Gio dc. H Ni

Ti liu dch
1. Musil J.G., Kurz .K., Novakava .O. 1982
Sinh ha hc hin i theo s . Nxb Y hc. H Ni.

Ti liu ting nc ngoi

1. Farkas G. 1984. Nvnyi anyagcserelettan. Akadmiai Kiad Budapest.
2. Lehninger A. L., 2004. Principle of Biochemistry, 4th Edition. W.H
Freeman.

213

Chng 13

Mi lin quan gia cc qu trnh

trao i cht
Trong c th mi qu trnh bin i cc cht khng xy ra ring r
m c mi quan h qua li rt cht ch. Mi quan h th hin c trong qu
trnh bin i ca mt nhm cht ln trong mi quan h ca qu trnh bin
i cc nhm cht khc nhau.
Trong cng mt nhm cht mi quan h th hin qua qu trnh ng
ha v d ha. Mi quan h tng h ca s ng ha v d ha th rhin
c hai qu trnh xy ra u c cc cht trung gian chung. V d
phosphoglyceric aldehyde va l sn phm trung gian trong qu trnh phn
gii (d ha) va l sn phm trung gian ca qu trnh tng hp (ng ha)
saccharose.
Gia cc nhm cht mi quan h din ra phc tp hn, nhiu hnh
thc hn. Trc ht mi quan h c th hin qua cc cht trung gian.
Mt cht l sn phm phn gii ca nhm cht ny li l nguyn liu
tng hp cho nhm cht khc. V d Acetyl-CoA l sn phm ca qu
trnh phn gii glucose ng thi n l nguyn liu tng hp acid bo.
Mi lin quan tng h gia cc qu trnh trao i saccharide, lipid,
protein, nucleic acid c ngha quan trng trong s sng ca sinh vt. V
d vic chuyn ha tinh bt thnh ng sau to cc cht bo trong
nhng thng ma ng thc vt cng nh ng vt, c ngha quan
trng trong vic tng kh nng chu rt ca chng. Vic chuyn bin cht
bo thnh ng khi ht ny mm s m bo thc n cho phi.
Trong qu trnh hat ng sng ca c th, saccharide l cht d tr
quan trng; khi c th cn s phn gii thnh phosphoglyceric acid ri
thnh pyruvic acid lm nguyn liu cho s tng hp amino acid, acid bo
v cc cht khc. Cht bo d tr cng c vai tr tng t, khi phn gii
cht bo to nn acetyl-CoA t li tng hp amino acid, saccharide
Khi trong c th ht ngun d tr saccharide v cht bo th protein s
b phn gii to ra cc sn phm t c th tng hp lipid, saccharide.

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Qu trnh chuyn ha trn c vai tr quan trng c i vi sinh vt

t dng ln sinh vt d dng.
sinh vt t dng ngun cht hu c s cp do quang hp to ra
l glucid, trc ht l glucose. T ngun saccharide , nh s chuyn
ho tng h mi to ra c cc cht hu c m c th cn n nh
lipid, protein, nucleic acid.
sinh vt d dng s dng nhng cht hu c c sn trong thc
n to nn cc cht c trng cho c th. T cc cht protein, lipid c
trong thc n qua qu trnh bin i s kin to nn nhng cht c trng
cho c th. Do thnh phn thc n khng th c y i vi nhu cu
ca c th cho nn trong qu trnh ng ha vic chuyn ha nhm cht
ny thnh cht hu c khc rt cn thit cho c th.

13.1. Mi lin quan gia qu trnh trao i saccharide v trao

i lipid
Hai qu trnh trao i lipid v saccharide c mi quan h cht ch
vi nhau. Saccharide d dng bin i thnh lipid v ngc li thng qua
cc cht trung gian l AlPG, PDA v Acetyl-CoA
T glucose qua qu trnh ng phn s to nn pyruvic acid. T
Pyruvic acid b oxy ha s to nn acetyl-CoA. Acetyl-CoA l nguyn liu
tng hp acid bo. ng thi trong qu trnh ng phn cn to ra AlPG,
t AlPG bin i thnh glycero-P , t to nn glycerin. Nh vy t sn
phm phn gii ca saccharide to nn nguyn liu c bn tng hp
lipid l glycerin v acid bo.
Ngc li qua s phn gii lipid s to nn cc cht trung gian l
acetyl-CoA, glycerin. T acetyl-CoA, qua chu trnh ornithine s tng hp
tr li saccharide. T glycerin to nn glycero-P v t tng hp li
saccharide.
thc vt, vi khun, nm mc, chu trnh glyoxylic l con ng ni
trc tip qu trnh trao i lipid vi qu trnh trao i saccharide. Qua chu
trnh ny acid bo sau khi phn gii thnh acetyl-CoA s bin i thnh
oxalo acetic acid, t tng hp nn glucose. Ngc li t glucose s to
acetyl-CoA v t tng hp tr li lipid.

215

Mi lin quan gia s trao i saccharide v trao i lipid

Saccharide

AlPG

Pyruvic acid

Acetyl-CoA

Chu trnh glyoxylic

Glycero-P

Glycerin
Acid bo

Acetyl-CoA

Lipid

13.2. Mi lin quan gia s trao i saccharide v trao i

protein
Pyruvic acid l mt xch ch yu ni lin qu trnh trao i protein
v trao i saccharide.
Glucose qua qu trnh ng phn to ra pyruvic acid. Pyruvic
acid l nguyn liu tng hp mt s amino acid trong h alanine:
alanine, leucine, valine
Trong qu trnh tng hp protein cn nng lng ATP, m ATP l
sn phm quan trng ca qu trnh trao i saccharide.

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S mi lin quan gia trao i saccharide v trao i protein

Saccharide

Ribuloso5P

Erylozo4P
Erytroso4P
Shikimic acid

His

APEP

APG
Pyruvic acid

Ser

Phe, Tyr, Trip

Protein

Gly, Cys

Ala, val, leu

Asp

Tre, Ila, Arg

Met, Lys

Glu

Arg, Pro

13.3. Mi lin quan gia trao i saccharide v trao i nucleic acid

Trong qu trnh phn gii saccharide theo con ng
pentosophosphate, Ribozo5-phosphate c to nn. T Riboso-5phosphate hnh thnh nn phospho-ribosyl-pyrophosphate (PRPP) l
nguyn liu tng hp nn cc nucleotide purine v nucleotide pyrimidine.
Ngc li trong qu trnh phn gii nucleic acid Riboso-5P c to
thnh. T Riboso 5P s hnh thnh cc monosaccharide khc.
Kiu lin quan th 2 gia qu trnh trao i nucleic acid v trao i
saccharide l s lin quan cht ch gia qu trnh sinh tng hp cc
nucleotide diphosphate v nucleotide triphosphate vi mc phn gii
saccharide trong t bo v qu trnh phn gii ny gn lin vi qu trnh
phosphoryl ho oxy ho. S phn gii saccharide to nng lng tng
hp cc nucleotide diphosphate v nucleotide triphosphate.
Qua qu trnh tng hp saccharide li cn s tham gia ca cc sn
phm to ra trong qu trnh trao i nucleic acid, nh UTP tham gia vo
qu trnh tng hp polysaccharide.

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13.4. Mi lin quan gia s trao i protein v trao i lipid

Trao i protein v trao i lipid c mi lin quan cht ch thng
qua cc cht trung gian. S phn gii lipid to nn glycerin v acid bo v
mt s cht khc nh serine, choline, sphingosine, H3PO4
Trc ht acid bo b phn gii to ra acetyl-CoA lm nguyn liu
tng hp nn nhiu loi amino acid. Glycerin trong qu trnh phn gii
to ra phosphoglyceric acid, t lm nguyn liu tng hp nn nhiu
amino acid. Nhng mi quan h ny xy ra tng t nh mi lin quan
gia saccharide vi protein phn tch trn. Ngc li, khi phn gii
protein cng to nn cc hp cht trung gian, t tng hp nn lipid.
Cc amino acid do thoi ho protein to ra, b kh amine s to nn cc
acid nh pyruvic acid, oxalo acetic acid, -cetoglutaric acid.
Trong s cc acid va nu th pyruvic acid c vai tr quan trng
trong qu trnh tng hp lipid. T pyruvic acid, acetyl-CoA c to ra,
acetyl-CoA l nguyn liu tng hp nn cc acid bo ng thi t
pyruvic acid cng c th to ra glycerophosphate v t to thnh
glycerin. Glycerin v acid bo l nguyn liu tng hp lipid.
Trong qu trnh phn gii lipid s to thnh nn mt lng ln ATP
l ngun nng lng cho qu trnh trao i protein. Ngc li protein vi
chc nng enzyme c vai tr quyt nh i vi cc phn ng xy ra trong
trao i lipid cng nh cc cht khc v khng c enzyme th khng c cc
phn ng ho sinh xy ra tc l khng c trao i cht.

218

S mi lin quan gia trao i lipid v trao i protein

Lipid

Glycerin

Nhng cht khc

Acetyl-CoA

Glycero-P
AlP

Fru.1.6d

Pyruvic
acid
APEP

Rib5P
His

Acid bo

Ser

Ala, val,
leu

Gly, Xyl

Oxalo acetic
acid
AsP
Tre, Met,
Ile, Cys

-Ceto glutaric
acid
Glu

Arg,
Pro

Phe
Protein

13.5. Mi lin quan gia trao i protein v trao i nucleic acid

Gia qu trnh trao i protein v trao i nucleic acid c mi quan
h c bit quan trng m biu hin r nht l trong c ch truyn t
thng tin di truyn. DNA lm khun sao m thnh RNAm t tng
hp nn protein. Cu trc phn t protein c m ho trong DNA.
Qu trnh trao i nucleic acid li ph thuc vo s c mt ca cc
phn t protein-enzyme. ng thi mt s amino acid l nguyn liu cho
qu trnh tng hp nucleic acid nh aspartic acid l nguyn liu tng
hp nucleotide pyrimidine, aspartic acid, glutamin, glycine l nguyn liu
tng hp nucleotide purine.

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V mi quan h m nhiu nh nghin cu cho rng s tng

hp protein l s cp cn s tng hp nucleic acid l qu trnh th cp lm
nhim v tham gia vo qu trnh tng hp protein.
Trong qu trnh phn gii protein vi s tham gia ca ATP to
nn cc nucleotide-peptid. Cc phn t ny lai to nn nhng protein mi lm
i mi thnh phn cc protein trong c th. ng thi s tng hp protein
cng cn cc nucleotide triphosphat lm ngun nng lng (ATP, GTP).

13.5. Mi lin quan gia trao i lipid v trao i nucleic acid

Gia lipid v nucleic acid c t mi lin quan trc tip, ch yu l
lin quan gin tip qua s trao i saccharide v protein.
Tuy nhin cng c mi lin quan gia hai qu trnh trao i cht
ny. S .oxi ho acid bo l ngun duy tr y cho s tng hp cc
nucleotide diphosphat v nucleotide triphosphat qua qu trnh phosphoryl
ho. Ngoi ra trong qu trnh tng hp lipid nh tng hp phospholipid c
s tham gia ca CTP.

TI LIU THAM KHO

Ti liu ting Vit
1. Trn Th n (ch bin). 1979. Ha sinh i cng (tp I, II). NxB KH&KT.
H Ni.
2. Phm Th Trn Chu, Trn Th ng. 2000. Ha sinh hc. Nxb Gio dc. H
Ni.
3. Nguyn B Lc. 1997. Ha sinh. Nxb Gio dc. H Ni

Ti liu dch
1. Musil J.G., Kurz .K., Novakava .O. 1982
Sinh ha hc hin i theo s . Nxb Y hc. H Ni.

Ti liu ting nc ngoi

1. Farkas G. 1984. Nvnyi anyagcserelettan. Akadmiai Kiad Budapest.
2. Lehninger A. L., 2004. Principle of Biochemistry, 4th Edition. W.H
Freeman.