TR NG I H C Y KHOA HU

B MN DI TRUYN Y HC


















GIO TRNH

SINH HC PHN T













2007
CHNG 2: SINH HC PHN T


1. DNA (desoxyribonuclic acid) 1
Nucleic acid 1
DNA (desoxyribonuclic acid) 2
Cu trc 2
Cc dng cu trc khng gian ca DNA 3
S bin tnh (denaturation) v hi tnh
(renaturation) ca DNA 4
DNA trong t bo 5


B gene ca prokaryote 5
B gene ca eukaryote 6
Phn loi DNA 6
Gene nhy 8
Gene 10
Cu hi n tp 11
Cu hi trc nghim 11


2. CHC NNG CA DNA 13
Bo qun v truyn t thng tin di truyn
13
M di truyn 13
Kh nng nhn i chnh xc 14


Tim nng t sa cha 14
Kh nng t bin 14
Cu hi n tp 15
Cu hi trc nghim 15


3. C CH T NHN I CA DNA 16
T nhn i prokaryote 16
Tch ri hai mch n ca phn t DNA
16
Tng hp on mi (primer) rna 17
Tng hp cc mch mi trn khun DNA
17
Hon chnh chui polynucleotide mi tng
hp 18


4. RNA (ribonuclic acid) 28
Cc loi RNA 28
rRNA (RNA ribosome) 28
tRNA (RNA vn chuyn) 28
mRNA (RNA thng tin) 30


Qu trnh t nhn i ca DNA
eukaryote 21
C ch sa sai DNA 23
C ch sa sai DNA trong qu trnh t
nhn i 23
C ch sa sai DNA ngoi qu trnh t
nhn i 24
Cu hi n tp 24
Cu hi trc nghim 24



Ribozyme v kh nng t ct (self-
Splicing) 30
Cu hi n tp 30
Cu hi trc nghim 30


5. QU TRNH PHIN M (TRANSCRIPTION) 33
Qu trnh phin m prokaryote 33
Giai on m u 33
Giai on ko di 34
Giai on kt thc 35
Qu trnh phin m eukaryote 37

Giai on m u 37
Giai on ko di 40
Giai on kt thc 40
Qu trnh trng thnh ca cc tin mRNA
40
Gn m chp (capping) 40


a
Gn ui polyA 41
Ct ni (splicing) 42
Cu hi n tp 43
Cu hi trc nghim 44


6. CU TRC V CHC NNG CA PROTEIN 45
Cu trc ca protein 45
Cu trc bc 1 45
Cu trc bc 2 46
Cu trc bc ba 47

Cu trc bc bn 47
Chc nng sinh hc 47
Cu hi n tp 49
Cu hi trc nghim 49


7. QU TRNH DCH M V IU HO SINH TNG HP PROTEIN 51
Qu trnh dch m 51
Gn amino acid vo tRNA 51
Ribosome 51
Cc giai on ca qu trnh dch m 52
Polysomes 55
iu ho s biu hin ca gene 56
iu ho s biu hin ca gene
prokaryote 56
iu ho s biu hin ca gene
eukaryote 58
Kim sot qu trnh phin m 59

Kim sot s biu hin ca gene thng qua
qu trnh hon thin mRNA 60
Kim sot s biu hin ca gene thng qua
s hng nh ca RNA 60
Qu trnh lm im lng RNA (RNA
silencing) 60
Kim sot qu trnh dch m v sau dch
m 61
Cu hi n tp 62
Cu hi trc nghim 62


































b
1. DNA (desoxyribonuclic acid)



Mc tiu

1. Trnh by c:

- Cu trc c bn ca mt nucleotide

- Cu trc ca DNA, cc loi DNA, gen nhy

- Cu trc c bn ca gene prokaryote v eukaryote

2. Phn bit c b gene ca prokaryote v eukaryote



NUCLEIC ACID
Nucleic acid l vt cht mang thng tin di truyn ca cc h thng sng c cu trc a
phn hnh thnh t cc n phn l nucleotide. Nucleic acid gm hai loi phn t c
cu to kh ging nhau l desoxyribonucleic (DNA) v ribonucleic acid (RNA).





Hnh 1: (a) Cc cu phn ca nucleotide; (b) Nucleotide; (c) Chui polynucleotide

Mi nucleotide c cu to gm ba thnh phn (hnh 1)

1. Nhm phosphate

2. ng Pentose (ng 5 Carbon). DNA ng ny l deoxyribose cn
RNA l ribose


1
3. Mt base nitric, base ny gm c hai nhm:

- Purine gm adenine (A), guanine (G)

- Pyrimidine gm thymine (T), cytosine (C) v uracyl (U)

Cc nucleotide c ni vi nhau bng lin kt phosphodiester to thnh chui
polynucleotide.

DNA (desoxyribonuclic acid)
CU TRC























Hnh 2 : Nguyn tc b sung gia cac base
purine va pyrimidine.


xon di 34 . (hnh 2 v 3)

Phn t ADN l mt chui xon
kp gm hai mch n, mi mch n l
mt chui polynucleotide. Mi
nucleotide gm (1) nhm phosphate, (2)
ng deoxyribose (C5H10O4) v (3)
mt trong bn loi base (A, T, G v
C)(hnh 2).

Hai chui polynucleotide kt hp
vi nhau nh cc lin kt hydro hnh
thnh gia cc base ca hai mch theo
nguyn tc b sung gia mt bn l
purine (A v G) v mt bn l pyrimidine
(C v T) trong A b sung vi T bng
hai lin kt hydro, G b sung vi C bng
ba lin kt hydro to nn mt cu trc
gm hai chui polynucleotide xon
quanh mt trc thnh hnh l xo vi
ng knh khong 2nm v mi bc



Hnh 3: Cu trc ca mt on DNA

Nguyn tc b sung m bo khong cch u n gia hai mch n ca DNA
(tng chiu di ca mt base purine v mt pyrimidine), cc phn t ng v cc
nhm phosphate nm pha ngoi to nn trc ng-phosphate hnh thnh lin kt
vi cc phn t nc, cc base quay vo pha trong hn ch s tip xc gia chng
vi cc phn t nc gi cho phn t DNA n nh.

Mi mch n l mt trnh t nucleotide c nh hng vi mt u l u 5'
phosphate t do (nhm phosphate t do gn vo C5 ca ng desoxyribose) k hiu


2
l 5'P, u kia c nhm OH v tr C3 nn gi l u 3' hydroxyl t do, k hiu l
3'OH. Hng quy c theo chiu t 5' n 3'. Hai mch n trong cu trc ca phn t
DNA i ngc chiu nhau gi l i song (anti paralell).

5'P 3'OH

3'OH 5"P

Nh vy u 5' P ca mch ny i din vi u 3' OH ca mch b sung.
Thng qua cu trc xon kp ca DNA c th nhn thy:
(1) Trnh t cc base ca mch ny khc vi trnh t base ca mch b sung vi
n.


(2) Hai mch ca DNA gn vi nhau bng cc lin kt hydro yu to nn tnh
linh hot cho phn t DNA khi thc hin cc hot ng chc nng nhn i hoc
tng hp RNA.

CC DNG CU TRC KHNG GIAN CA DNA

M hnh trn ca DNA c m t trn do Watson v Crick a ra nm 1953
v c cng nhn nh l mt cu trc duy nht ca DNA trong mt thi gian di.
Vo thp nin 70, vi s h tr ca cc k thut phn tch chnh xc nhiu dng DNA
c pht hin. Vic phn nh cc dng DNA c thc hin da trn 2 ch s :

- h: chiu cao gia hai nuceotide k nhau

- n: S cp nucleotide trong mt vng xon

Mt s dng DNA c lit k trong bng di y:



Dng
S cp base ca
mt vng xon
(n)
Gc xon so
vi mt phng
ca base ()
khong cch
h gia 2 base
k nhau (A
o
)
ng knh
ca chui
xon kp (A
o
)

A

11

+ 32,7 2,56

23

B

19

+ 36,0 3,38

19

C

9 1/3

+ 38,6 3,32

19

Z

12

- 30,0 3,71

18

Dng DNA m Watson v Crick m t l dng B, dng ph bin nht, tn ti
trong iu kin sinh l bnh thng. Cc dng khc xut hin nhng iu kin m
v ion khc nhau. Dng Z c chiu xon ngc v pha bn tri theo hnh zigzag nn
gi l dng Z. Cc dng DNA khc nhau cho thy DNA trong t bo sng khng tn
ti vi mt dng duy nht m tu trng thi sinh l c th di dng ny hay dng
khc.(hnh 4)





3

Hnh 4: Cc dng DNA

S BIN TNH (DENATURATION) V HI TNH (RENATURATION) CA
DNA

Hai mch n ca phn t DNA gn vi nhau thng qua cc lin kt hydro yu.
Cc tc nhn lm t gy cc lin kt ny nh khi un nng phn t DNA nhit
khong 80 - 95
o
C s lm cho hai mch tch ri nhau. Hin tng ny c gi l hin
tng bin tnh (denaturation) ca DNA. (hnh 5)

















Hnh 5: Hin tng bin tnh ca DNA

Nhit lm hai mch DNA tch ri nhau c gi l im chy (melting point)
ca DNA. im ny c trung cho mi loi DNA, ph thuc vo s lng cc lin kt
hydro trong cu trc. T l cp G-C cng cao th im chy cng ln. Mt s cht nh
formanide c kh nng lm h thp im chy nn c dng trong k thut lai phn
t lm gim nhit lai.

S bin tnh ny c tnh thun nghch. Sau khi DNA b bin tnh nu h nhit
t t tr li bnh thng th chng c th gn li vi nhau trn c s ca nguyn tc b
sung tr thnh mch kp. Hin tng ny c gi l hi tnh (renaturation).





4
DNA TRONG T BO
Tt c cc sinh vt c cu ti t bo, ty th, lp th u cha DNA mch kp. Cc
virus c b gene a dng gm RNA hoc DNA mch n hoc mch kp. Bng di
y gii thiu chiu di b gene n bi cn c theo s cp base ca cc nhm sinh
vt chnh nh sau:

Sinh vt S cp base
Virus 10
3
n 10
5


Vi khun E. Coli 4,5 x 10
6

Nm men 5 x 10
7


Rui gim Drosophila

ng vt c xng sng
1,5 x 10
8

10
8
n 10
10

Thc vt 10
10
n 10
11

Ngi 3 x 10
9



Qua bng trn c th nhn thy khng c s tng quan gia mc tin ho v
s lng cp base trong b gene. Gia b gene ca prokaryote v eukaryote c s
khc bit ng k v thnh phn cu to v cch t chc ca DNA trong t bo.

B GENE CA PROKARYOTE

B gene ca vi khun E. Coli v a s cc sinh vt prokaryote u l mt phn t
DNA c dng vng v khng gn vi protein to thnh nhim sc th nh
eukaryote. (tuy nhin khi nim nhim sc th hin nay cng c dng cho c vi
khun v c hiu l si DNA). (hnh 6)

prokaryote, DNA thng dng siu xon vi DNA mch kp xon vn thnh
hnh s 8. y l dng t nhin trong t bo vi khun, si DNA c cc phn t RNA
ni gip cun xon v lm cho chiu di phn t c rt ngn ng k. Tnh trng
siu xon ny c th thay i thun nghch dc tc ng ca cc enzyme DNase hoc
RNase. Ngoi ra DNA c th di dng vng trn hoc dng thng.



















Hnh 6: DNA prokaryote




5
B GENE CA EUKARYOTE

Cch t chc ca DNA trong nhn t bo
DNA ca eukaryote c kch thc rt ln, v d ngi tng chiu di ca tt c
DNA c trong nhn t bo c th di n 2m. c th nm gn trong nhn t bo,
DNA phi c cun xon nhiu mc cu trc khc nhau.

Nucleosome
Cc on DNA vi chiu
di tng ng vi khong t
140 n 150 cp base (base
pair: bp) cun quanh mt li
gm 8 phn t protein histone
(2 H2A, 2 H2B, 2 H3 v 2H4)
to thnh nucleosome vi
ng knh khong 11nm. Cc
nucleosome ni vi nhau bng
mt on DNA khong 20 - 60

bp vi mt phn t histone
trung gian (H1). Cc histone
Hnh 7: Nucleosome
lin kt vi phn t DNA nh cc lin kt ion hnh thnh gia cc nhnh bn mang
in tch m ca cc histone vi cc nhm phosphate mang din tch dng ca DNA.
(hnh 7)

Si v quai chromatin
Khong 6 nucleosome cun li thnh mt solenoid ng knh khong 30nm, cc
solenoid cun li thnh cc quai chromatin (chromatin loop) ng knh khong
300nm, mi quai chromatin c khong 100.000 bp (100 kb). Bng cch ny DNA c
th gim chiu di xung khong 1/10.000 ln so vi chiu di ca n trc khi cun
xon. (hnh 8)

Cht d nhim sc
Cc quai chromatin tip tc cun xon vi ng knh khong 700nm.

Nhim sc th
k gia ca qu trnh phn bo chromatin cun xon mc ti a to nn
NST vi ng knh khong 1.400nm.

PHN LOI DNA
eukaryote mc d DNA mang thng tin m ha cho cc protein nhng trong
thc t ch c mt t l rt thp DNA thc hin chc nng ny. ngi c t hn 5%
trong s 3 t cp nucleotide trong genome ca ngi thc s lm chc nng mang
thng tin, cn li phn ln vt liu di truyn cha c bit chc nng. DNA ca
ngi c chia thnh cc loi sau (hnh 9):








6

Hnh 8 . M hnh cun xon ca DNA



DNA c bn (single - copy DNA)
DNA c bn chim khong 45% genome v gm cc gene m ho cho cc
protein. Cc on DNA loi ny ch c thy mt ln duy nht (hoc vi ln) trong
genome. Tuy nhin phn m ha cho protein ch chim mt phn nh trn loi DNA
ny m thi, phn ln cn li l cc intron hoc l cc on DNA nm xen gia cc
gene.

DNA lp (repetitive DNA)
DNA lp chim 55% cn li ca genome, y l cc on DNA c lp i lp
li c th ln ti hng ngn ln trong genome, c 2 loi chnh:


7
DNA v tinh (satellite DNA)
Loi DNA ny chim khong 10% genome v tp trung mt s vng nht
nh trn NST, chng sp xp ni ui nhau, ci ny tip theo ci kia. DNA v
tinh c chia thnh 3 loi nh:

- DNA v tinh alpha: c kch thc 171 bp, lp i lp li nhiu ln vi
chiu di hng triu bp hoc hn. Loi ny c thy cnh tm ng ca
NST.

- DNA tiu v tinh (minisatellite DNA): c kch thc t 14 - 500 bp, lp
i lp li vi chiu di khong vi ngn bp.

- DNA vi v tinh (microsatellite DNA): c kch thc t 1 - 13 bp, lp i
lp li vi tng chiu di khng qu vi trm bp.

Hai loi DNA tiu v vi v tinh c s khc nhau rt ln trong chiu di gia
ngi ny vi ngi khc v iu ny lm chng tr nn rt hu ch trong vic lp bn
gene. DNA tiu v tinh v vi v tinh c gp vi tn s trung bnh l 1 trn mi
2 kb trong genome v chng chim khong 3% genome.

DNA lp li ri rc
Loi DNA ny chim khong 45% genome, gm 2 loi:
- Cc yu t ri rc c kch thc ngn (SINEs: short interspersed elements):
kch thc t 90 - 500bp.

- Cc yu t ri rc c kch thc di (LINEs: long interspersed elements):
kch thc 7.000 bp






GENE NHY
Hnh 9 . Cc loi DNA

c Barbara McClintock pht hin trn bp cch y hn 50 nm. Gene nhy
l mt on DNA c kh nng di chuyn t v tr ny sang v tr khc trn DNA, n c
th ri b hoc xen vo cu trc ca 1 gene. Hin tng ny c thy c vi sinh
vt, thc vt v ng vt vi tn s khong 10
-5
10
-7
t bo.






8
Cc trnh t gn xen (IS: insertion sequences)
Cc trnh t gn xen ny c thy vi khun E. Coli vi tnh cht di chuyn t
v tr ny sang v tr khc trn DNA. Cc IS c kch thc khong 1 - 2 kb vi trnh t
trung tm c trng cho tng loi IS, hai u l hai trnh t ngn vi khong 50 bp
ging nhau nhng c chiu ngc nhau gi l on lp nghch hng (inverted
repeat) pha trong v hai trnh t ngn ging nhau vi khong 5 - 11 bp nhng
cng chiu vi nhau nn c gi l on lp ng hng (direct repeat). Cc IS
khng m ho cho protein v ch c pht hin thng qua hu qu ca n trn vi
sinh vt. (hnh 10)


Hnh 10: S ca mt trnh t gn xen (IS: insertion sequence)



Transposon

Hnh 11: (a)Transposon c ngun gc t DNA; (b) Transposon c ngun gc t retrovirus

9
Gene nhy ng vt v thc vt chng c gi l transposon, c kch thc
ln hn IS. Transposon cng c hai u l hai trnh t ging nhau nhng ngc chiu
nhau. Phn trnh t trung tm ngoi cc gene s c gn xen cn c cc gene m ho
cho cc enzyme cn thit cho qu trnh ny.

Phn ln cc transposon ca eukaryote c ngun gc t RNA. Cc RNA c
gn xen vo b gene theo cch ca cc retrovirus v vy chng cn c gi l
retroposon. Cc retroposon gm hai loi chnh cn c vo ngun gc: (1) Nhm c
ngun gc t cc DNA ca t bo; (2) Nhm c ngun gc t retrovirus. (hnh 11)

GENE
C th ni gene l mt on ca phn t DNA c chc nng di truyn. Tuy nhin
gia prokaryote v eukaryote c mt s khc bit ln

prokaryote
Trinh t ca cc nucleotide trn gene s m ho cho trnh t ca cc amino acid

eukaryote
Trong cu trc ca gene tn ti nhng vng khng mang m gi l intron v
nhng vng mang m gi l exon. y l mt c im quan trng phn bit gia
DNA ca sinh vt eukaryote v prokaryote.

Cc intron chim phn ln trong cu trc hu ht cc gene. Cc on intron ny
s c cc enzyme ct mt cch chnh xc ra khi cc phn t mRNA trc khi
chng i ra khi nhn t bo. V tr ct ca cc enzyme c xc nh bi cc on
DNA c gi l cc on ng nht (consensus sequences) (on ny c tn nh vy
v c thy ph bin tt c cc c th eukaryote) nm cnh mi on exon.

Hnh 12 : Cu trc chi tit ca mt gene Eularyotae

V hu ht cc gene ca c th eukaryote gm ch yu l cc on intron nn cho
php ngi ta ngh n vic cc on ny c th c mt chc nng no . Hin nay
chc nng ca cc intron vn cn ang c nghin cu. Ngi ta cho rng nhng
on intron do lm tng thm chiu di ca gene s to thun li cho s ti sp xp ca
cc gene trn cp NST tng ng qua qu trnh trao i cho trong gim phn hoc
nh hng n thi gian nhn i v phin m ca DNA. (hnh 12)

Cu trc ph bin ca mt gene eukaryote gm c ba vng:

1. Vng 5' khng phin m, mang cc trnh t c nhim v iu ho biu hin
ca gene v cc trnh t hot ho hot ng phin m.

2. Vng s phin m gm cc on intron v exon.

3. Vng 3' khng phin m c chc nng cha r.


10
CU HI N TP

1. M t cu trc ca mt n phn ca nucleic acid ?

2. M t cu trc ho hc v khng gian ca DNA ?

3. Th no l s bin tnh, hi tnh ca DNA ?

4. M t cc c im c bn ca DNA prokaryote ?

5. M t cc c im c bn ca DNA eukaryote ?

6. eukaryote, lm th no cc phn t DNA c th nm gn trong nhn t bo
k trung gian ?

7. ngi DNA c phn loi nh th no ?

8. M t cc loi DNA v tinh trong cu trc DNA ca ngi ?

9. Gen nhy l g ?

10. Nu cc c im c bn ca gene eukaryote ?



CU HI TRC NGHIM

1. Nucleic acid l mt a phn gm c:
A. Hai loi phn t l desoxyribonucleic acid (DNA) v ribonucleic acid (RNA)

B. Hai loi phn t l ribonucleic acid thng tin (mRNA) v ribonucleic acid vn
chuyn (tRNA)

C. Hai loi phn t l ribonucleic acid thng tin (mRNA) v ribonucleic acid vn
chuyn (tRNA)

D. Hai loi phn t l ribonucleic acid thng tin (mRNA) v ribonucleic acid
ribosome (rRNA)

E. Hai loi phn t l deoxyribonucleotide v ribonucleiotide

2. Base nitric Purine gm c:
A. Thymine (T), cytosine (C) v uracyl (U) B. Adenine (A) v uracyl (U)

C. Guanine (G) v cytosine (C) D. Adenine (A) v guanine (G)

E. Adenine (A)v thymine (T)

3. Hai chui polynucleotide kt hp vi nhau nh cc lin kt ...... (C: cng ha tr,
H: hydro; P: phosphodiester) hnh thnh gia cc base ca hai mch theo nguyn
tc b sung trong A b sung vi T bng ....... (2: hai lin kt hydro; 3: ba lin kt
hydro), G b sung vi C bng ....... (2: hai lin kt hydro; 3: ba lin kt hydro) to
nn mt cu trc gm hai chui polynucleotide xon quanh mt trc.

A. H, 3,2 B. P, 2, 3 C. C, 2, 3 D. H, 2, 3 E. P, 3, 2

4. Tng chiu di ca ton b phn t DNA c trong mt t bo lng bi ca ngi
l bao nhiu ?

A. 1 mt B. 2 mt C. 3 mt D.4 mt E. 5 mt

11
5. Mi mch n l mt trnh t nucleotide c nh hng vi mt u l u ........
( 5'; 3') phosphate t do, u kia c nhm OH v tr ........ (C5 ; C3) gi l u
........ (5', 3') hydroxyl t do.

A. 3'; C3,5' B. 5'; C5,3' C. 3'; C5,5' D. 5'; C3,3' E. 5'; C3,5'

6. Hai mch n ca phn t DNA gn vi nhau thng qua cc lin kt ......... (P:
phosphodiester; H: hydro). Cc tc nhn lm t gy cc lin kt ny nh khi un
nng phn t DNA nhit khong 80 - 95o C s lm cho hai mch tch ri
nhau. Hin tng ny c gi l hin tng ..........(B: bin tnh; T: thoi ho) ca
DNA. S bin tnh ny.........(C: c tnh thun nghch; K: khng c tnh thun
nghch)

A. P, B, C B. H, T, K C. H, B, C D. H, T, C E. P, B, K

7. B gene ca vi khun E. Coli v a s cc sinh vt .........( E: Eukaryotae; P:
Prokaryotae) u l mt phn t DNA xon kp c dng ...........(V:vng; K: khng
to vng) v .........(G: gn, KG: khng gn) vi protein to thnh nhim sc th.

A. P, V, G B. P, V, KG C. E, K, G D. E, K, KG E. P, K, G

8. M t no di y v nucleosome l khng ng:
A. on DNA trong cu trc c chiu di tng ng vi khong t 140 n 150
cp base

B. Mt li gm 8 phn t protein histone (2 H2A, 2 H2B, 2 H3 v 2H4)

C. Cc nucleosome ni vi nhau bng mt on DNA khong 20 - 60 bp vi mt
phn t histone trung gian.

D. Khong 6 solenoid cun li thnh mt nucleosome ng knh khong 30nm

E. Cc histone lin kt vi phn t DNA nh cc lin kt ion hnh thnh gia cc
nhnh bn mang in tch m ca cc histone vi cc nhm phosphate mang
din tch dng ca DNA.

9. M t no di y v DNA c bn l ng:
A. Chim khong 45% genome

B. Gm cc gene m ho cho cc protein. Phn m ha cho protein ch chim
mt phn nh trn loi DNA ny phn ln cn li l cc intron hoc l cc
on DNA nm xen gia cc gene.

C. Cc on DNA ny ch c thy mt ln duy nht (hoc vi ln) trong
genome

D. A v C ng E. A, B v C u ng



P N

1. A 2. D 3. D 4. B 5. D 6. C 7. B 8. D 9. E






12
2. CHC NNG CA DNA


Mc tiu
Trnh by c:

- Cc chc nng c bn ca DNA

- Cch thc DNA m ho thng tin di truyn



BO QUN V TRUYN T THNG TIN DI TRUYN
Tu thuc vo s lng, thnh phn v trt t sp xp ca 4 loi nucleotide m s
c v s loi DNA khc nhau to nn tnh a dng v c th cho mi DNA. Gi s c
mt chui polynucleotide gm 10 nucleotide th s c ti 4
10
= 1.048.576 loi khc
nhau. Vi c tnh ny cho php phn t DNA tr thnh vt cht mang thng tin di
truyn cho gn nh ton b sinh gii.

Thng tin di truyn cho vic tng hp mt phn t protein c m ho trn
gene, mt on ca DNA, di dng m b ba.

M DI TRUYN

Mi protein c cu to t 1 hoc nhiu chui polypeptide. Mi chui
polypeptid c cu to t cc n v cu trc c bn l cc acid amine. C th c 20
loi acid amine khc nhau, trnh t ca cc acid amine ny trong chui polypeptide
c DNA quy nh.

Mi acid amine c m ha bi ba nucleotide k nhau trn DNA, ba nucleotide
ny c gi l mt codon. Vi 4 loi nucleotide khc nhau s c 64 codon khc nhau
bi thnh phn v trt t ca cc nucleotide (do protein c tng hp trc tip trn
RNA thng tin, do cc nucleotide A, U, G v C c s dng minh ho cc m
b ba) , trong s ny c 3 codon kt thc (stop codon) UAA, UAG v UGA c nhim
v bo hiu chm dt vic tng hp chui polypeptide. Trong s 61 m cn li c
nhiu codon cng m ha cho 1 acid amine, hin tng ny c goi l hin tng
thoi ha m (degeneration) (bng 1).

M di truyn c tnh ng nht cho ton b sinh gii tr mt s ngoi l i vi
cc codon ti th. DNA ca bo quan ny c mt s codon m cho cc acid amine
khc vi ngha ca cc codon ny trn DNA trong nhn.

ty th:
- UGA m cho tryptophan thay v bo hiu chm dt vic tng hp protein.

- AGA v AGG khng m cho arginine m bo hiu chm dt tng hp
protein.

- AUA m cho methionine thay v m cho isoleucine.







13
U Phe Ser Tyr Cys
U P Ser Tyr Cys
U L

Ser STOP STOP
U L Ser STOP Trp
C L Pro His Arg
C L

Pro His Arg

C L


Pro

Gln

Arg
C L Pro Gln Arg
A Il Thr Asn Ser
A Il Thr Asn Ser
A Il Thr Lys Arg
A M Thr Lys Arg
G V Ala Asp Gly
G V Ala Asp Gly
G V

Ala Glu Gly
KH NNG NHN I CHNH XC
M hnh DNA cho php phn t DNA nhn i mt cch chnh xc da trn
nguyn tc b sung theo kiu bn bo tn.



V TR 1 V TR 2 V TR 3

(u 5) U C A G (u 3)

U

C

A

G

U

C


A

G

U

C

A

G

U

C

A



Bng 1. Bng m di truyn

Ala: Alanine; Arg: arginine; Asn: asparagine; Asp: aspartic acid; Cys: cysteine; Gln: glutamine;
Gla: glutamic acid; Gly: glycine; His: histidine; Ile: isoleucine; Leu: leucine; Lys: lysine; Met:
methionine; Phe: phenylalanine; Pro: proline; Ser: serine; Thr: threonine; Trp: tryptophan;
Tyr: tyrosine; Val: valine

TIM NNG T SA CHA
Cu trc ca DNA to cho DNA c tim nng sa cha cc sai st trong cu trc.
Nhng sai st xy ra trn DNA trong qu trnh t nhn i hoc khi khng nhn i
u c ct b v thay th m bo tnh n nh v c trng ca phn t DNA.

KH NNG T BIN
Bn cnh tnh n nh, phn t DNA vn c th xy ra bin i gy ra cc bin d
di truyn, nhng bin i ny s lm thay i tnh cht ca gene v gp phn thc y
s tin ho.






14
CU HI N TP
1. Thng tin di truyn c m ho nh th no trn gene ?

2. Hy nu cc c im ca m di truyn ?

3. Th no l hin tng thoi ho m ?

4. Nu nhng im khc nhau c bn trong m di truyn trn DNA trong nhn v
trong ti th.

5. Nu cc chc nng c bn ca DNA ?



CU HI TRC NGHIM
1. Hin tng thoi ha m l hin tng:
A. Mt codon cng m ha cho nhiu acid amine

B. 3 codon UAA, UAG v UGA khng mang m m c nhim v bo hiu chm
dt vic tng hp chui polypeptide.

C. Mi codon ch m ha cho 1 acid amine

D. Nhiu codon cng m ha cho mt acid amine

E. Ton b sinh gii c cng mt loi m di truyn

2. Tng hp mt chui polynucleotide nhn to t ba loi nucleotide A, G, v T. trn
chui polynucleotide ny s c ti a bao nhiu loi codon khc nhau ?
A. 64 B. 8 C. 27 D. 9 E. 32

3. Mt chui polynucleotide gm c 10 nucleotide, s c ti a bao nhiu kiu sp xp
khc nhau trong trnh t ca cc nucleotide ?
A. 10
4
B. 10
10
C. 4
10
D. 40 E. 4. 10
2


4. Acid amine no di y ch c m ho bi mt codon duy nht:
A. Methionine B. Tryptophan C. Lysine
D. Arginine E. Valine




P N

1. D 2. C 3. C 4. A
















15
3. C CH T NHN I CA DNA


Mc tiu
Trnh by c:
- C ch t nhn i DNA prokaryote v eukaryote mc c bn

- C ch t sa sai DNA



T NHN I PROKARYOTE
Qu trnh t nhn i ca DNA l mt qu trnh phc tp. prokaryote qu trnh
ny gm nhng bc sau:

TCH RI HAI MCH N CA PHN T DNA

Qu trnh t nhn i bt
u t mt im xut pht gi
l im ori (origine) v trin
khai v c hai pha. Ton b
DNA dng vng
ca prokaryote l mt
n v sao chp duy nht.
Mi n v sao chp c gi
l replicon v do
b gene ca
prokaryote ch gm c mt
replicon duy nht.

u tin cc phn t
protein c gi l cc protein
m u (iniator protein) s gn
vo im Ori. Qu trnh gn
kt ny s lm cho 1 on
DNA tho xon to thun li
cho s tc ng ca enzyme
helicase v cc protein SSB
(Single Strand Binding
protein: protein lin kt vi
mch n ca DNA).







Hnh 1: Bt u qu trnh t
nhn i ca DNA




16
Hai mch ca phn t DNA s c tch ri nh enzyme helicase qua vic ph
v cc lin kt hydro gia cc base vi nng lng gii phng t NTP (nucleosid 5'
triphosphate). Cc mch n sau khi tch ri nhau ra s c duy tr n nh di
dng mch n nh cc protein SSB. Cc protein SSB gn ln trn mch n lm cho
hai mch khng ti kt hp tr li c. (hnh 1)

Di tc dng ca enzyme helicase trn DNA m s hnh thnh mt ch nhn i
(replication fork) tin hnh qu trnh sao chp trn hai mch khun ca DNA.

Mt loi enzyme protein khc cn thit cho vic tho xon ca DNA l DNA
gyrase mt loi enzyme topoisomerase. Enzyme ny s chy trc ch ba sao m
gip tho cu trc xon ca DNA. (hnh 2)





Hnh 2: Tch ri hai mch ca chui xon kp DNA

TNG HP ON MI (PRIMER) RNA

enzyme DNA polymerase c th tng hp DNA, trc tin phi c mt on
mi RNA ngn c tng hp nh mt phc hp protein gi l primosome.
Primosome bao gm nhiu protein v mt enzyme tng hp RNA da trn khun
DNA gi l primase. Nh s c mt ca on mi RNA ny m enzyme DNA
polymerase mi c th tip tc ni di chui polynucleotide phc v cho qu trnh
nhn i ca DNA. (hnh 3)

TNG HP CC MCH MI TRN KHUN DNA

Trong qu trnh tng hp DNA, enzyme DNA polymerase lun lun di chuyn
t u 3' n u 5' v mch mi lun lun c tng hp theo chiu t 5' n
3'. Enzyme DNA polymerase xc tc n u th cc protein SSB s c gii
phng khi mch khun DNA n . C hai loi DNA polymerase tham gia: (1)
DNA polymerase III tng hp DNA mi da trn nguyn tc b sung; (2) DNA
polymerase I ct b on mi RNA. Do hai mch khun ca DNA ngc chiu nhau
nn vic tng hp mch mi trn hai mch khun din ra khng ging nhau.



17
Trn mch khun 3' 5'
Do DNA polymerase III xc tc theo hng t 3' 5' nn mch mi s c
tng hp theo hng 5' 3', cng hng vi hng tho xon v c tng hp mt
cch lin tc, mch ny c tng hp sm hn mch kia nn tm gi l mch
trc (leading strand). (hnh 4)


Hnh 3: Tng hp on mi RNA vo ko di chui polynucleotide

Trn mch khun 5' 3'
Do DNA polymerase i ngc vi hng tho xon nn mch mi khng c
tng hp mt cch lin tc m di dng nhng on ngn theo chiu t 5' 3' gi l
on Okasaki vi kch thc khong t 1.000 n 2.000 base, cc on Okasaki sau
mi c ni li vi nhau nh enzyme ligase. Do mch ny c tng hp mun
hn nn c gi l mch sau (lagging strand).

HON CHNH CHUI POLYNUCLEOTIDE MI TNG HP

Khi mch kp tch ra, enzyme primase s tng hp on mi RNA khong 10
nucleotide c trnh t cc base b sung vi cc base trn mch khun theo nguyn tc
b sung. (hnh 5)

Trong qu trnh tng hp cc loi nucleosid triphosphat ATP, GTP, TTP, CTP
giu nng lng s n bt cp vi cc nucleotide trn mch khun DNA theo nguyn
tc b sung di tc dng ca DNA polymerase III. Cc nucleotide mi s c
enzyme ny gn vi u 3' OH ca mch ang c tng hp.

DNA polymerase s tng hp mch mi bng cch trt trn mch khun cho
n khi gp on mi RNA pha trc th dng li. DNA polymerase I nh hot tnh
exonuclease 5' 3' s ct b on mi RNA v lp cc nucleotide ca DNA vo ch
trng v polymer ho theo hng 5' 3' to nn mt on DNA ngn vi 10
nucleotide, on ny h hai u v s c ni vi mch trc v mch sau n bng


18
enzyme ligase. (hnh 5)


























.






















Hnh 4:

Qua trnht nhn i
trn hai mach khun
cua DNA vi mt
mach c ntg hp
lin tuc (mach trc) va
mt mach c tng
hp thannhg t
oan (mach sau).






19









































Hnh 5:

Qua trnh


ct oan
mi RNA va gn cachui
DNA polynucleotide lai vi
nhau nh enzyme DNA
ligase.








Trong hai phn t DNA mi c tng hp t DNA m, mi DNA ch c mt
mch mi c tng hp cn mch kia l ca DNA m ng vai tr khun tng hp.
Do s t nhn i ca DNA c thc hin theo kiu bn bo tn (semi
conservation).





20
vi khun E. Coli qu trnh t nhn i din ra rt nhanh, c th t n tc
50.000 nucleotide/pht. (hnh 6)
























Hnh 6: Qu trnh t nhn i ca DNA prokaryote

QU TRNH T NHN I CA DNA EUKARYOTE
Do t bo eukaryote c b gene ln hn nhiu v DNA kt hp vi protein to
thnh nhim sc th nn qu trnh t nhn i ca DNA eukaryote din ra phc tp
hn v chm hn (khong 50 nucleotide/giy). (hnh 7)

Qu trnh nhn i c thc hin ti nhiu replicon ngha l c nhiu im ori
cng lc tham gia vo qu trnh ny v s nhn i cng lan ra c 2 pha nh
prokaryote. T bo kim sot c ch ny mt cch cht ch, mi im ori ch c
php thc hin mt ln t nhn i.

Cc DNA polymerase eukaryote gm c:

- Polymerase /primase c nhim v tng hp mi RNA cho mch sau. Do
n khng c hot tnh exonuclease nn khng c kh nng sa sai.

- Polymerase c chc nng ging DNA polymerase I, c nhim v tng hp
chui polynucleotide, sa sai v hon chnh mch mi sau khi loi b on
mi RNA.

- Polymerase c chc nng ging DNA polymerase III

Ngoi ra cn c cc polymerase , polymerase vi chc nng cha r.

Ngoi cc polymerase ni trn, qu trnh t nhn i ca DNA cn c s tham
gia ca nhiu loi protein c hiu nh:

- PCNA (Proliferating Cell Nuclear Antigen: khng nguyn nhn t bo
ang phn chia) ng vai tr hot ho polymerase v .

- Cc RF- A v C (Replication Factor: yu t nhn i; RF-A; RF-C) cn thit
cho hot ng ca polymerase v .



21























































Hnh 7: Qu trnh t nhn i DNA eukaryote



Qu trnh t nhn i eukaryote c th tm tt nh sau:

1. DNA u tin c enzyme topoisomerase v RF-A tho xon

2. Trn mch sau:

- Phc hp polymerase / primase phi hp vi RF-A tng hp on
RNA mi di khong 10 nucleotide.


22
- Phc hp polymerase kt hp vi RF-C ko di on mi thm
khong 20 deoxynucleotide.

- Khi phc hp PCNA - ATP s nh ch hot ng ca polymerase
li v gip polymerase gn vo tng hp on Okazaki.

- Polymerase c gii phng s chuyn ln mch i din (mch
trc) v tng hp lin tc mch mi.

3. Cc phn t DNA sau khi c tng hp s t chc li thnh nucleosome
trong vng vi pht.

Nhn chung c ch t nhn i DNA ca eukaryote vn cha c hiu bit mt
cch y .

C CH SA SAI DNA
Di tc ng ca cc nhn t trong mi trng, DNA c th b bin i trong
cu trc, nhng bin i ny s gy ra nhng hu qu nghim trng cho c th sinh
vt, v vy trong t bo c nhiu c ch khc nhau m bo phn t DNA khng b
thay i di tc ng ca cc tc nhn i (hnh 8)























Hnh 8: C ch sa sai DNA trong qu trnh t nhn i v ngoi qu trnh
t nhn i.

C CH SA SAI DNA TRONG QU TRNH T NHN I

S chnh xc trong qu trnh t nhn i c mt ngha rt quan trng i vi
hot ng bnh thng ca t bo v ca c th, tuy nhin trong qu trnh t nhn i
ca DNA nguy c xy ra nhm ln l rt ln. Th nhim in vitro cho thy kh nng
sao chp sai l 1.10
-5
tc l trong 100.000 nucleotide s c 1 nucleotide b lp sai.
Trong thc t t l sai st thp hn rt nhiu, ngi c tnh tn s sai st l 1.10
-9

tc l 1 sai st xy ra trong 1 t nucleotide. S chnh xc ny c m bo thng qua
vai tr ca mt s enzyme DNA polymerase.





23
prokaryote, enzyme DNA polymerase I v III va c kh nng polymer ho,
va c hot tnh exonuclease 5' 3' v 3' 5', do trong qu trnh xc tc cho
hot ng t nhn i ca DNA nu gp nucleotide lp sai, enzyme ny s li li, ct
b nucleotide sai theo hng 3' 5'.

C CH SA SAI DNA NGOI QU TRNH T NHN I

Ngoi qu trnh t nhn i phn t DNA cng c th b bin i, nhng bin i
ny xy ra vi tn s khng nh nhng nh c ch sa sai nn nhng bin i ny
c duy tr mc thp.

Nguyn tc sa sai c tm tt nh sau: Cc enzyme c hiu c nhim v pht
hin trnh t sai, gn vo v ct on ny ra khi DNA ri s dng mch ng
lm khun tng hp li on va b ct cho ng. C khong 50 enzyme thuc loi
ny.



CU HI N TP
1. Hy nu nhng im c bn trong qu trnh nhn i ca prokaryote ?

2. Hy nu nhng im c bn trong qu trnh nhn i ca eukaryote ?

3. Nhng im khc nhau c bn trong qu trnh t nhn i trn hai mch ca DNA
prokaryote ?

4. DNA c sa sai nh th no trong qu trnh t nhn i v trong giai on
khng nhn i?



CU HI TRC NGHIM

1. Qu trnh t nhn i Prokaryotae bt u t mt v tr xut pht gi l:
A. im ori B. Replicon C. Codon
D. Ch nhn i E. Okazaki


2. Mi n v sao chp trong qu trnh t nhn i Prokaryotae c gi l:
A. im ori B. Replicon C. Codon
D. Ch nhn i E. Okazaki


3. Trong qu trnh t nhn i Prokaryotae, cc phn t protein c gi l cc
protein m u (iniator protein) s gn vo mt v tr trn DNA lm cho 1 on
DNA tho xon to thun li cho s tc ng ca enzyme helicase v cc protein
SSB, v tr l:
A. im ori B. Replicon C. Codon
D. Ch nhn i E. Okazaki


4. Trong qu trnh t nhn i Prokaryotae, enzyme helicase c nhim v:
A. Lm n nh cc mch n ca DNA sau khi tch nhau ra v lm cho hai
mch khng ti kt hp tr li c



24
B. Tho xon phn t DNA
C. Tng hp on mi RNA
D. Tip tc ni di chui polynucleotide

E. Ph v cc lin kt hydro gia cc base tch hai mch ca phn t DNA

5. Trong qu trnh t nhn i Prokaryotae, cc protein SSB c nhim v:
A. Lm n nh cc mch n ca DNA sau khi tch nhau ra v lm cho hai
mch khng ti kt hp tr li c

B. Tho xon phn t DNA
C. Tng hp on mi RNA
D. Tip tc ni di chui polynucleotide

E. Ph v cc lin kt hydro gia cc base tch hai mch ca phn t DNA

6. Trong qu trnh t nhn i Prokaryotae, enzyme DNA gyrase c nhim v:
A. Lm n nh cc mch n ca DNA sau khi tch nhau ra v lm cho hai
mch khng ti kt hp tr li c

B. Tho xon phn t DNA
C. Tng hp on mi RNA
D. Tip tc ni di chui polynucleotide

E. Ph v cc lin kt hydro gia cc base tch hai mch ca phn t DNA

7. Trong qu trnh t nhn i Prokaryotae, enzyme topoisomerase c nhim v:
A. Lm n nh cc mch n ca DNA sau khi tch nhau ra v lm cho hai
mch khng ti kt hp tr li c

B. Tho xon phn t DNA
C. Tng hp on mi RNA
D. Tip tc ni di chui polynucleotide

E. Ph v cc lin kt hydro gia cc base tch hai mch ca phn t DNA

8. Trong qu trnh t nhn i Prokaryotae, enzyme primase c nhim v:
A. Lm n nh cc mch n ca DNA sau khi tch nhau ra v lm cho hai
mch khng ti kt hp tr li c

B. Tho xon phn t DNA
C. Tng hp on mi RNA
D. Tip tc ni di chui polynucleotide

E. Ph v cc lin kt hydro gia cc base tch hai mch ca phn t DNA

9. Trong qu trnh t nhn i Prokaryotae, enzyme DNA polymerase c nhim v:
A. Lm n nh cc mch n ca DNA sau khi tch nhau ra v lm cho hai
mch khng ti kt hp tr li c


25
B. Tho xon phn t DNA
C. Tng hp on mi RNA
D. Tip tc ni di chui polynucleotide

E. Ph v cc lin kt hydro gia cc base tch hai mch ca phn t DNA

10. Trong qu trnh t nhn i Prokaryotae, enzyme no di y c nhim v hnh
thnh ch nhn i
A. Helicase B. Topoisomerase C. DNA gyrase
D. DNA polymerase E. Primase


11. Trong qu trnh t nhn i Prokaryotae, phc hp protein primosome c nhim
v:

A. Lm n nh cc mch n ca DNA sau khi tch nhau ra v lm cho hai
mch khng ti kt hp tr li c

B. Tho xon phn t DNA
C. Tng hp on mi RNA
D. Tip tc ni di chui polynucleotide

E. Ph v cc lin kt hydro gia cc base tch hai mch ca phn t DNA

12. Trong thc t t l sai st trong qu trnh t nhn i ngi c tnh c tn s:
A. 1.10
-5
tc l 1 sai st xy ra trong 100.000 nucleotide.
B. 1.10
-6
tc l 1 sai st xy ra trong 1 triu nucleotide.
C. 1.10
-7
tc l 1 sai st xy ra trong 10 triu nuclotide.
D. 1.10
-8
tc l 1 sai st xy ra trong 100 triu nucleotide.
E. 1.10
-9
tc l 1 sai st xy ra trong 1 t nucleotide.

13. C ch sa sai DNA trong qu trnh t nhn i c m bo thng qua vai tr
ca enzyme
A. Helicase B. Topoisomerase C. DNA gyrase
D. DNA polymerase E. Primase




P N

1. A 2. B 3. A 4. E 5. A 6. B 7. B 8.C 9. D 10. A 11. C

12. E 13. D












26
4. RNA (ribonuclic acid)


Mc tiu
Trnh by c:

- Cu trc chung ca phn t RNA

- Cu trc v chc nng ca cc loi RNA



Cc RNA c tng hp t cc gene tng ng trn DNA , ng vai tr trung
gian trong qu trnh sinh tng hp protein. Cc phn t RNA u c c im chung
nh sau v cu trc (hnh 1):

- Cu trc a phn dng mch n polynucleotide.

- Trong cu trc ca cc n phn nucleotide: ng pentose l ribose
C5H10O5, base Uracil (U) thay cho thymin (T).






































Hnh 1: (a) Cu trc bc 1 (mch thng) ca RNA; (b) Cu trc bc 2 ca RNA
(to xon khng hon ton)





27
Cn c trn chc nng c th chia ra lm 3 loi RNA c bn sau:

- RNA thng tin (mRNA: messenger RNA)

- RNA vn chuyn (tRNA: transfer RNA)

- RNA ribosome (rRNA: ribosomal RNA)

CC LOI RNA
rRNA (RNA ribosome)





Hnh 2: Cc rRNA c hon thin sau khi phin m. Lu l eukaryote khng xy ra qu
trnh ct ta nhng on nucleotide v phn t 5S rRNA c phin m
ring t gene rRNA nh (small rRNA)

Trong t bo rRNA gn vi cc protein thnh mt phc hp ribonucleoprotein
gi l ribosome tham gia vo qu trnh gii m. rRNA chim t 75% n 80% tng s
RNA ca t bo. Da trn h s lng S (S: sedimentation) trong qu trnh ly tm phn
tch m ngi ta chia thnh cc loi sau (hnh 22):

prokaryote cc ribosome c h s lng 70S, gm 2 n v:

- n v ln 50S c 1 rRNA 23S v 1 rRNA 5S

- n v nh 30S ch c 1 rRNA 16S

Eukaryota cc ribosome c h s lng 80S, gm 2 n v:
- n v ln 60S c 1 rRNA 28S v 1 rRNA 5,8S v 1 rRNA 5S.

- n v nh 40S ch c 1 rRNA 18S

rRNA c cu trc khng gian phc tp do c nhiu on bt cp vi nhau theo
nguyn tc b sung.

tRNA (RNA vn chuyn)

tRNA c nhim v gn vi amio acid c hiu a n ribosome trong qu
trnh gii m. Mi amino acid c t nht mt tRNA c hiu. Cc tRNA c cu trc


28
chung nh sau (hnh 3):

C khong t 73 - 93 nucleotide

Cun xon mt u da trn nguyn
tc b sung gia cc cp nucleotide AU
v GC nhng khng hon ton to nn
hnh l ch ba vi cc quai (loop) khng
to xon.

u 3' ca nhnh tip nhn (acceptor
arm) c trnh t kt thc l CCA, y l
u gn amino acid trong qu trnh vn
chuyn thng qua lin kt cng ho tr.
(hnh 4)

Nhnh TC c mang 3 base T,
pseudouracyl () v C trong quai

Mt u khng to xon mang b ba
i m (anticodon) b sung vi b ba m
sao (codon) trn phn t mRNA. Mt c
im ng ch ca tRNA l mt tRNA
c th kt hp vi hai codon khc nhau
cng m ho cho mt amino acid.



Hnh 3: Cu tcrucua mt tRNA

Vic gn amino acid vo
tRNA c hiu c thc hin
qua trung gian ca enzyme
amino acyl-tRNA synthetase.
C 20 loi enzyme ny tng
ng vi 20 loi amino acid khc
nhau. Qu trnh gn amino acid
vo tRNA gm hai giai on nh
sau:













Hnh 4: V tr gn amino acid trn tRNA

Giai on 1: enzyme nhn
bit v gn vi mt amino acyl
c hiu

Enzyme + amino acid +
ATP Enzyme-aminoacyl-
AMP +P-P

Giai on 2: Amino acid
chuyn t phc hp enzyme-
aminoacyl sang tRNA tng ng

Enzyme-aminoacyl-AMP + tRNA tRNA-aminoacyl +AMP + Enzyme







29
mRNA (RNA thng tin)

c sao ra t trnh t ncleotide trn DNA, c vai tr chuyn thng tin m ho
trn DNA n ribosome tng hp phn t protein tng ng. mRNA c cu trc
mch n. mRNA ca prokaryote c cu trc n gin, thi gian bn hu ngn, trung
bnh khong 2 pht, mRNA ca eukaryote c thi gian bn hu lu hn, khong t 30
pht n 24 gi.

mRNA c on u mang cc tn hiu cho ribosome nhn bit gn vo v bt
u thc hin qu trnh dch m v phn ui mang m kt thc bo hiu chm dt
qu trnh dch m.

RIBOZYME V KH NNG T CT (SELF-SPLICING)

Mt s RNA c kh nng xc tc nh vai tr ca cc protein enzyme c gi l
ribozyme. nhng RNA ny qu trnh t ct c th xy ra trn RNA m khng
cn n vai tr ca cc enzyme protein. Hot ng ny lin quan n s bin i cu
trc ca on intron trn RNA lm cho on intron c hot tnh xc tc tng t
enzyme, t xc tc cho qu trnh t ct ra khi RNA.




CU HI N TP

1. RNA c cu trc ho hc nh th no ?

2. M t cu trc khng gian v chc nng ca rRNA, mRNA v tRNA ?

3. Phc h tRNA aminoacyl c hnh thnh nh th no ?



CU HI TRC NGHIM
1. Trn mRNA, b ba m ........ (A: AUG; B: UAA; C: UAG) m ho cho methionine
(Met) s bt u cho qu trnh dch m. Mc d ch c 1 codon m ho cho Met
nhng c....... ( 1. mt; 2. hai; 3. ba) loi tRNA mang Met n ribosome.
A. A, 2 B. A, 1 C. B,2 D.C, 3 E. B, 1

2. S khc bit c bn trong cu trc gia n phn ca DNA v RNA v tr:
A. H3PO4 B. ng C. Base nitric

D. B v C ng E. A, B v C u ng

3. Cc loi base nitric c th xut hin trong cu trc ca 1 phn t ribonuclotide:

A. A, T, G, X v mt s bin dng khc B. U, T, G, X

C. A, U, G, X v mt s bin dng khc D. A, T, U, G, X

E. A, G, X

4. Phn t ng c mt trong cu trc ca phn t RNA l:
A. Glucose B. Fructose C. Deoxyribose

D. Galactose E. Ribose


30
5. S khc bit c bn trong cu trc gia cc loi RNA do cc yu t sau quyt nh:

A. S lng, thnh phn cc loi ribonuclotide trong cu trc

B. S lng, thnh phn, trt t ca cc loi ribonuclotide v cu trc khng gian
ca ARN

C. Thnh phn v trt t ca cc loi ribonuclotide

D. Cu trc khng gian ca cc loi RNA
E. S lng cc loi ribonuclotide
6. M t no sau y v tRNA l ng:
A. tRNA l mt mch polyribonuclotide c s ribonuclotide tng ng vi s
nucltt trn mt mch ca gen cu trc

B. tRNA l mt mch polyribonuclotide gm t 80 n 100 ribonuclotide khng
to xon, mt u t do cn mt u mang axt amin

C. tRNA l mt mch polyribonuclotide gm t 80 n 100 ribonuclotide cun
xon mt u trn c s nguyn tc b sung thc hin gia tt c cc ribonuclotide
ca tARN, mt u mang axt amin v mt u mang b ba i m.

D. tRNA l mt mch polyribonuclotide gm t 80 n 100 ribonuclotide cun
xon mt u, c on cc cp base lin kt theo nguyn tc b sung v c nhng
on khng lin kt b sung, to nn nhng thy trn, mt u t do mang amino acid
c hiu v mt thy trn mang b ba i m

E. tRNA c dng mch n hay cun xon mt u vi s ribonuclotide t 160
n 13000, nhng tRNA ny s kt hp vi nhng protein c hiu to nn cc tiu
phn ribosome

7. Trong phn t ARN nguyn tc b sung c thc hin gia:
A. A v U bng 3 lin kt hydro; G v X bng 2 lin kt hydro

B. A v T bng 2 lin kt hydro; G v X bng 3 lin kt hydro

C. A v T bng 3 lin kt hydro; G v X bng 2 lin kt hydro

D. A v U bng 2 lin kt hydro; G v X bng 3 lin kt hydro

E. A v G bng 2 lin kt hydro; T v X bng 3 lin kt hydro



P N

1. A 2. D 3. C 4. E 5. B 6. D 7. D













31
5. QU TRNH PHIN M (TRANSCRIPTION)


Mc tiu
Trnh by c:

- Qu trnh phin m c bn ca eukaryote v prokaryote

- Qu trnh hnh thnh mRNA ca eukaryote





Qu trnh phin m (hinh 1) cho php
tng hp RNA trn khun DNA thng qua s
xc tc ca h enzyme RNA polymerase (tn
y l DNA-dependent-RNA polyme-
rase). Nh vy DNA khng nhng c kh
nng tng hp nn DNA m cn c kh nng
tng hp nn RNA, mt phn t khc. Kh
nng ny ca DNA c gi l kh nng d xc
tc (heterocatalysis).

Qu trnh phin m d prokaryote hoc
eukaryote u tun theo cc nguyn tc c
bn sau:



Hnh 1: Qua trnh phin ma trDnNA

- Ch mt trong hai mch ca DNA c dng lm khun tng hp RNA.

- Khng cn s tham gia ca on mi (primer)

- RNA polymerase bm vo DNA lm tch ri hai mch ca DNA v di
chuyn trn khun DNA theo hng t 3' n 5' v do mRNA c tng
hp theo hng t 5' n 3'.




Hnh 2: Qu trnh phin m prokaryote

Do enzyme RNA polymerase khng c kh nng sa sai nn chnh xc km
hn nhiu so vi qu trnh t nhn i, tuy nhin do RNA khng c sao chp nn
khng nh hng n vic truyn t thng tin qu cc th h.




32
QU TRNH PHIN M PROKARYOTE
prokaryote ch c mt loi RNA polymerase chu trch nhim tng hp tt c
cc loi RNA. RNA polymerase c cu trc bc 4 vi nm chui polypeptid '; ; ;
v . Cc chui ny ni vi nhau bng cc lin kt yu. Enzyme ny s bm vo v
tr khi ng (promotor) trn mch DNA dng lm khun bt u tng hp mRNA
(hnh 2, 3).



Hnh 3: Tm tt qu trnh phin m

Phn t mRNA c tng hp c th cha thng tin cho nhiu gene ni tip
nhau (polycistronic mRNA), cc protein do cc gene ny m ho thng lin quan vi
nhau trong mt qu trnh chuyn ho no .

Qu trnh phin m c th tm tt thnh 3 giai on nh sau:

GIAI ON M U


Hnh 4: V tr khi ng (promotor)

V tr khi ng trn DNA thng l mt trnh t gm 6 nuclotide, mt nm
cch im bt u tng hp RNA 10 bp (trnh t - 10) v mt cch im bt u
tng hp 35bp (trnh t -35).



33
Tiu n v gip RNA polymerase nhn bit v tr khi ng trn mt
mch ca DNA. u tin n s gn vo v tr trnh t -35, s gn kt ny sau s
lm cho trnh t -10 tho xon v mt si n DNA s c s dng lm khun
tng hp RNA. (hnh 3, 4)












































Hnh 5: Giai on khi u v ko di trong qu trnh phin m

GIAI ON KO DI

Khi phn t RNA ang tng hp c khong 8 nucleotide th tiu n v s tch
khi phc hp enzyme RNA polymerase v s gn vo mt promotor khc bt
u mt qu trnh phin m mi ng thi RNA polymerase s gn thm cc nhn t
ko di (elongation factors) s tch ri ca n v ny s gip RNA polymerase tip
tc trt trn mch khun tng hp RNA. (hnh 5)

Trong qu trnh tng hp RNA, si RNA ang tng hp s tch dn khi mch
khun ca DNA v on DNA sau khi phin m s c RNA polymerase ng xon
tr li. (hnh 3)




34
GIAI ON KT THC

Trn DNA ca vi khun c cc du hiu kt thc, khi RNA polymerase gp du
hiu ny th s ngng phin m, gii phng RNA v xc tc cho mt qu trnh phin
m mi. Phn t tRNA v rRNA sau khi gii phng s tip tc hon thin cu trc bc
2. (hnh 8)

vi khun c 2 kiu kt
thc:


(1) Cc du hiu kt thc
ph thuc vo yu t Rho
(Rho-dependent terminators),
du hiu ny s nh ch qu
trnh phin m khi c mt yu
t Rho ().

(2) Cc du hin kt thc
khng ph thuc vo yu t
Rho (Rho-independent
terminators) c th chm dt
qu trnh phin m khi khng
c yu t Rho.

C ch hot ng ca cc
du hin kt thc khng
ph thuc vo yu t Rho
(hnh 6)
Trn DNA mang cc an
lp o ngc (inverted
repeats), y l mt on c
trnh t nucleotide o ngc
trn DNA v c th b sung cho
nhau.. Khi on ny c phin
m chng s to thnh cu trc
dng kp tc (hairpin)

Pha sau on lp o
ngc th hai c mt trnh t
khong 6 nucleotide adenine,
c phin m thnh chui
polyuracyl pha sau cu trc
kp tc trn RNA
S c mt ca cu trc kp
tc s lm cho RNA polymera
di chuyn chm v dng li
dn n vic chm dt vic
phin m. S b sung gia A
v U pha sau cu trc kp tc



















































Hnh 6: Kt thc phin m bng cc du hin kt thc
khng ph thuc vo yu t Rho



35
tng i khng hng nh to thun li cho vic tch RNA ra khi mch khun.

C ch hot ng ca cc du hin kt thc ph thuc vo yu t Rho
(hnh 7,8)











































Hnh 7: Kt thc phin m bng cc du hin kt thc ph thuc vo yu t Rho

Trn DNA mang cc trnh t bo hiu chm dt hot ng phin m, gi l du
hiu kt thc.

Pha trc du hiu kt thc l mt trnh t DNA m cho mt on RNA khng
c kh nng to nn cu trc bc 2 gi l on RNA khng c cu trc
(unstructed RNA). Protein Rho s gn vo v tr ny trn RNA v di chuyn v pha
u 3' theo hng ca enzyme RNA polymerase.

RNA polymerase s dng li khi gp du hiu kt thc to iu kin cho yu t
Rho bt kp. Yu t ny c hot tnh nh enzyme helicase s tch cu trc kp RNA-
DNA , gii phng RNA v chm dt qu trnh phin m.







36


Hnh 8: Phn t RNA hon thin cu trc bc 2

QU TRNH PHIN M EUKARYOTE
Vic xc tc cho qu trnh phin m eukaryote do 3 loi RNA polymerase I. II.
III chu trch nhim trong RNA polymerase III phin m cho cc RNA nh nh
tRNA v loi RNA 5S, snRNA (small nuclear RNA: RNA nh nhn).

Phn t mRNA c tng hp ch cha thng tin ca 1 gene m thi
(monocistronic mRNA). Nhng khc vi prokaryote, mRNA sau khi phin m s
khng tham gia ngay vo qu trnh dch m m phi qua mt qu trnh ch bin
mi tr thnh mt mRNA trng thnh (hnh 30) tham gia dch m, qu trnh ny
c gi l qu trnh trng thnh (maturation). mRNA trc khi qua qu trnh trng
thnh c gi l tin mRNA (premessenger RNA). (hnh 9)



Hnh 9: Cu trc chung ca mt mRNA trng thnh eukaryote

Qu trnh phin m eukaryote tri qua cc giai on sau:

GIAI ON M U

Chu s kim sot ca mt trnh t c bit trn DNA gi l li khi ng (core
promoter) trong c trnh t lp gi l hp TATA (TATA box) nm cch v tr phin
m khong 25 - 35 nucleotide c vai tr quan trng trong qu trnh phin m
eukaryote.



37

Vai tr ca RNA polymerase II
(hnh 11)
RNA polymerase II bt u
phin m vi s phi hp vi cc
nhn t phin m (TF: transcription
factors) c bn cht l protein gm
c TFIID, TFIIB, TFIIA, TFIIE v
TFIIH. Ch TFII k hin cho loi
TF phi hp vi RNA polymerse
II, cc k t pha sau i din cho
cc loi TF khc nhau.

Hnh 10: TBP gn vi hp TATA trn DNA, be
cong DNA thanh hnh yn nga


Hnh 11: Giai on khi u qu trnh phin m eukaryote vi RNA polymerase II


38
TFIID gn vi hp TATA v v tr hot ng ca RNA polymerase II m bo
cho qu trnh phin m bt u mt cch chnh xc. TFIID c t nht chn chui
polypeptide, mt trong chng c gi l TBP (TATA-binding protein: protein gn
vi TATA) c kh nng ghi nhn v gn vo hp TATA trn DNA khun, b cong
on DNA thnh hnh yn nga to thun li cho hot ng phin m. (hnh 10)

Mt phc hp gm DNA polymerase, cc yu t phin m, cc yu t trung gian
khc n gn vi TFIID. TFIIA c vai tr duy tr mi quan h gia TFIID v DNA,
TFIIB gip chn la v tr bt u, TFIIH gip tho xon DNA, cc cht trung gian
ng vai tr trung gian gia h thng phin m v cc protein hot ha cho qu trnh
phin m.

Ngay pha trc li khi ng l mt trnh t lp gi l vng iu ha khi ng
(regulatory promoter). Vng ny s kt hp vi cc protein hot ha cho qu trnh
phin m v qua nh hng ti tc phin m.

Trn DNA c cc trnh t vi tc dng tng cng hot ng phin m ca cc
gene gi l vng tng cng (enhancer), vng ny c th nm pha trc hoc pha
sau gene v cch xa gene m n iu khin, i khi vng ny nm trong cc intron ca
gene. Mc d nm cch xa vng khi ng nhng thng qua cc protein hot ha cho
qu trnh phin m, DNA s to thnh mt quai DNA lm cho vng tng cng v
vng khi ng nm st nhau iu khin qu trnh phin m.

Vai tr ca RNA polymerse I (hnh 12)



























Hnh 12: Giai on khi u qu trnh phin m eukaryote vi RNA polymerase I

RNA polymerase I cn c 2 protein l SL1 v UBF bt u cho qu trnh
phin m. 2 protein ny s gn vo 2 vng chc nng trn DNA, mt vng nm quanh
v tr bt u phin m ko di t v tr -40 n +20 gi l yu t li (core element) cn
thit cho vic bt u phin m v mt vng pha trc vng yu t li v tr -180




39
n -107 gi l yu t kim sot
(control element) c vai tr tng
cng hiu qu hot ng ca vng
quanh v tr phin m. 2 vng ny
giu GC v c trnh t ging nhau

Nh vy c hai enzyme RNA
polymerase I v II u v vai tr
quan trng nh nhau trong vic bt
u qu trnh phin m.

Vai tr ca RNA polymersase III
(hnh 13)
Ch xc tc cho qu trnh phin
m cc RNA nh nh tRNA v loi
RNA 5S, snRNA. Vng tc ng ca
enzyme ny c th nm pha trc v
tr phin m (snRNA) hoc nm
trong vng phin m (t v rRNA).

GIAI ON KO DI

mRNA c tng hp trn
mch khun ca DNA nh nhn t
TFIIS

GIAI ON KT THC































Hnh 13: Cac v tr khi ng khac nhau trn ca
loai gene cua RNA polymerase III

Giai on ny eukaryote cha c bit mt cch tng tn. 3 loi RNA
polymerase s dng cc c ch khc nhau chm dt vic phin m.

RNA polymerase I cn mt yu t kt thc ging nh yu t Rho vi khun, yu
t ny gn vi mt trnh t DNA pha sau v tr kt thc.

RNA polymerase III kt thc phin m sau khi phin m mt trnh t kt thc to
ra mt chui gm cc uracyl trn phn t RNA, qu trnh ny khng cn s hnh thnh
cu trc "kp tc" pha trc chui uracyl nh vi khun.

i vi cc gene phin m di tc ng ca RNA polymerase II, qu trnh
phin m c th chm dt ti nhiu v tr nm trong khong t vi trm n vi ngn
cp base pha cui gene.

QU TRNH TRNG THNH CA CC TIN mRNA
Qu trnh trng thnh ca cc tin mRNA xy ra trong nhn t bo t trc
khi qu trnh phin m kt thc. Qu trinh ny gm cc bc sau:

GN M CHP (CAPPING)

Ngay sau khi bt u phin m, mt nhm phosphate ca nucleotide u 5' s
b tch i to iu kin cho mt guanine vi 1 phosphate gn vo u 5', sau base
guanine s gn thm nhm methyl vo v tr s 7 (7-methyl guanin), s mehtyl ha
ny c th xy ra v tr 2' trn phn t ng ca mt hoc mt s nucleotide tip
theo.


40
M chp c vai tr ht sc
quan trng trong qu trnh dch m
v cc protein ghi nhn m chp s
bm vo y v ribosome s bm
vo cc protein ny bt u cho
qu trnh dch m. Ngoi ra m
chp cn gip gi cho cu trc ca
mRNA c hng nh v nh
hng n qu trnh ct b cc
intron. (hnh 14)

GN UI POLYA

Ngay sau khi phin m xong,
cc tin mRNA (pre mRNA) s b
ct v tr nm pha sau mt trnh
t ng nht (consensus sequence)
AAUAAA khong 11 - 30
nucleotide. Sau enzyme polyA
polymerase v mt s yu t khc
s gip gn mt lng adenin v
u 3' ca mRNA, sau khi gn
khong 10 adenine, mt protein gi
l PABP (PolyA Binding Proein:
pretein lin kt vi polyA) s gn
vo ui poly A v gip pht trin
tip tc chui polynucleotide
adenin, cng nhiu PABP gn vo
s c cng nhiu adenine trong ui
poly A.S lng adenin trong ui
poly A khong t 50 - 250.

ui poly A gip duy tr s n
nh ca mRNA v ko di thi
gian hot ng dch m ca RNA
ny. S n nh ca mRNA ph
thuc vo cc protein gn vo ui
polyA. (hnh 15)










Hnh 14: Qu trnh hnh thnh m chp (cap) u
5' bng cch gn thm guanine v methyl ha.





41




Hnh 15: Gn ui poly A vo u 3' ca tin mRNA

CT NI (SPLICING)

Trong bc ny tin mRNA s ct b cc on intron khng mang m v ni cc
on exon mang m li vi nhau. C 3 trnh t ng vai tr quan trng trong qu trnh
ny l (hnh 16):


Hnh 16: Cc v tr quan trng trong vic ct ni tin mRNA

- Trnh t cho GU u 5' ca intron (mt s t l AU)

- im nhnh (branch point) l mt trnh t nucleotide vi mt adenine
nm pha trc v tr ct u 3' khang t 18 n 40 nucleotide. Trnh t
ny khng c ng nht cao nhng thng c dng YNYYRAY (Y:
nucleotide pyrimidine, N: bt c loi nucleotide no, R: nucleotide purine,
v A : adenine). t bin xy ra vng ny s lm cho qu trnh ct ni
khng xy ra c.

- Trnh t nhn AG u 3' (mt s t l AC)

Qu trnh ct ni c thc hin vi s tham gia ca mt phc hp ln c tn l
splicesome (th ct ni), c cu to t nhiu phn t RNA v protein. Cc phn
t RNA tham gia vo cu trc ca splicesome l cc RNA nhn nh (snRNAs),
nhng snRNA ny s kt hp vi cc protein to thnh cc snRNP (small
ribonucleoprotein particles: tiu phn ribonucleoprotein nh), mi snRNP cha
mt snRNA v nhiu protein. Splicesome gm 5 snRNP c gi tn theo loi snRNA
m n mang trong cu trc (U1, U2, U4, U5 v U6).



42
Trc khi qu trnh ct ni xy ra exon trc (exon 1) v exon sau (exon 2) c
tch ra bi 1 intron. Tin mRNA c ct ni theo hai bc sau (hnh 17):




Hnh 17: Qu trnh ct b cc on intron trong tin mRNA to thnh mRNA trng thnh

Bc 1
Tin mRNA b ct v tr ct 5' lm gii phng exon 1 ra khi intron v u 5'
ca intron s gn vo im nhnh bng cch un cong li to thnh mt cu trc hnh
"nt thng lng" (lariat). Nucleotide guanine u ct 5' ca intron s lin kt vi
nucleotid adenine ca im nhnh. Nhm OH trn carbon 2' ca adenine ti im
nhnh s ph v lin kt 5' phosphodiester ca guanine v tr ct 5', v to thnh mt
lin kt 5' - 2' phosphodiester mi gia guanine v adenine. Qu trnh ny c gi l
qu trnh chuyn ester ha (transesterification).

Bc 2
V tr ct 3' s b ct v ng thi u 3' ca exon 1 s gn vi u 5' ca exon 2
bng lin kt ha tr, phn ng ny cng c thc hin qua qu trnh chuyn ester
ha, trong nhm 3'-OH ca exon 1 s ph v lin kt phosphodiester v tr ct 3'
v to thnh lin kt phosphodiester mi gia u 3' ca exon 1 v u 5' ca exon 2.
Khi on intron c ct b di dng nt thng lng. nt thng lng intron ny s
tr thnh mch thng v nhanh chng b thoi ha di tc dng ca cc enzzyme ca
nhn. Gia cc exon v intron khc cng din ra qu trnh tng t to thnh
mRNA trng thnh (mature mRNA) ch gm cc on exxon. mRNA trng
thnh sau s vo bo tng tham gia qu trnh dch m.



CU HI N TP
1. Nu cc nguyn tc c bn ca qu trnh phin m ?

2. Trnh by cc giai on c bn trong qu trnh phin m prokaryote ?

3. Trnh by cc giai on c bn trong qu trnh phin m eukaryote ?


43
4. eukaryote, sau khi mRNA c tng hp, s tri qua nhng bin i nh th no
tr thnh mt mRNA trng thnh c th tham gia qu trnh dch m ?



CU HI TRC NGHIM
1. M chp G trn mRNA cho php cc ...... (P: protein; A: amino acid; N:nucleotide)
ghi nhn m chp bm vo y v .......(R: ribosome; T: tRNA; E:enzyme) s bm
vo cc protein ny bt u cho qu trnh dch m v gip gi cho cu trc ca
mRNA c ........(H: hng nh; L: linh hot)
A. P, T, L B. A, R, H C. N, E, H
D. A, R, L E. P, R, H


2. Trong qu trnh phin m ca Eukaryote, ui poly A c nhim v:
A. Gip duy tr s n nh ca mRNA

B. Ko di thi gian hot ng dch m ca RNA
C. Ni cc on exon li vi nhau
D. A, B ng

E. A, B, C u ng

3. Trn mRNA, b ba m ........ (A: AUG; B: UAA; C: UAG) m ho cho methionine
(Met) s bt u cho qu trnh dch m. Mc d ch c 1 codon m ho cho Met
nhng c...... ( 1. mt; 2. hai; 3. ba) loi tRNA mang Met n ribosome.
A. A, 2 B. A, 1 C. B,2 D. C, 3 E. B, 1

4. Sau khi qu trnh tng hp mRNA bt u, u 5' ca mRNA ang c tng hp s
c gn thm:
A. Enzyme RNA polymerase II
B. Mt chui poly A
C. Mt on base gm 100 n 200 Uracyl

D. Mt nucleotide Adenine c methyl ha

E. Mt nucleotide Guanine c methyl ha

5. to thnh u "5' cap" th u 5' ca phn t mRNA ang c tng hp s
c "bt" li bng cch gn thm mt nucleotide c methyl ha loi:
A. Adenine B. Cytosine C.Uracyl
D. Thymine E. Guanine




P N

1. E 2. D 3. A 4. E 5. E






44
6. CU TRC V CHC NNG CA PROTEIN


Mc tiu
Trnh by c:

- Cu trc chung ca cc amino acid

- Cu trc chung ca protein

- Cc chc nng c bn ca protein



CU TRC CA PROTEIN
Protein c cu trc a phn t vi n phn l amino acid. C tt c 20 loi
amino acid khc nhau, tt c u c cu trc chung nh sau. (hnh 1)

- Mt nhm amine (-NH2)

- Mt nhm carboxyl (- COOH)

- Mt gc R c trng cho tng loi
aminoacid, quy nh tnh cht ca
tng loi amino acid. Da vo in
tch ca gc ny m ngi ta chia cc
amino acid thnh 5 nhm: (1) Nhm
mang in tch dng; (2) Nhm
mang in tch m; (3) Nhm gc R
c nhn thm; (4) Nhm phn cc v
(5) Nhm khng phn cc (hnh 2, 3,
4).

Cc phn t amino acid ni vi nhau bi
cc lin kt peptide c hnh thnh gia

Hnh 1: Cu t ruc chung cua
mt amino acid
nhm carboxyl ca amino acid trc vi nhm amine ca amino acid k tip.
Phn ng s gii phng mt phn t nc. (hnh 5)

Nu mt chui c s amino acid di 30 s c gi l chui peptid, nu s
amino acid ln hn s c gi l polypeptide. Mi polypeptide c hai u tn cng,
mt u mang nhm amin t do (N - teminus), u kia mang nhm carboxyl t do
(C-terminus). T protein dng ch mt n v chc nng c cu trc khng gian
phc tp. Mt protein c th c cu to t 1 hoc nhiu chui polypeptide.

Protein c bn bc cu trc:

CU TRC BC 1

L s lng cc chui polypeptide tham gia vo cu trc, s lng v trnh t ca
cc amino acid trong mi chui polypeptide. Ty thuc vo s lng, thnh phn v
trt t sp xp ca cc amino acid m s c v s loi protein khc nhau.

Cu trc bc mt c mt vai tr quyt nh trong vic xc nh cu trc khng
gian cc bc tip theo ca phn t protein.


45




Hnh 2: Amino acid mang in tch dng v in tch m



Hnh 3: Amino acid phn cc v khng phn cc




Hnh 4: Amino acid cha nhn thm Hnh 5: Hnh thnh lin kt peptide

CU TRC BC 2 (HNH 6, 8)

S tng tc gia cc peptide trong protein s to nn cu trc bc 2 ca protein.
C 2 loi cu trc bc 2 l xon alpha v xon beta.





46
Xon alpha
Hnh thnh thng
qu cc lin kt hydro
gia cc nhm amin
ca amino acid vi
oxygen ca amino acid
th ba tnh t amino
acid to nn cu trc
xon u quanh mt trc
hnh tr tng tng.
Cu trc ny c thy
mt s protein dng si.

Xon beta
Cc chui polypep-
tide nm cnh nhau ni
vi nhau bng cc lin
kt hydro to nn cc
tm xp (plated sheet).
Vi cch sp xp ny




























Hnh 6: Cturuc bc 1 va bc2 acuphnt protein
phn t protein do v chc hn nh cc protein ca mng nhn, lng v, t la v.v...

CU TRC BC BA (hnh 7)





Hnh 7: Cu t ruc bc 3 va bc4 acuphnt protein

Chui polypeptide cun
xon to thnh khi cu thng
qua lin kt gia cc gc R. Cc
protein ny gm cc enzyme,
cc hormone protein, cc khng
th v.v.... Trong cu trc bc ba
thng c s an xen gia on
xon alpha v on xon beta.

CU TRC BC BN (HNH 7)

Khi c hai hoc nhiu
chui polypeptide c lp, ging
nhau hoc khc nhau gn li vi
nhau thng qua cc lin kt yu
s to thnh cu trc bc bn
nh trng hp cu trc ca
phn t hemoblobine ca ngi.



CHC NNG SINH HC
Protein thc hin hu ht cc chc nng c bn ca cht sng, di y lit k
mt s chc nng c bn ca protein:




47




Hnh 8:
(a) (b)

(a) Xon alpha: Cc lin kt hydro gia cc nhm amin ca amino acid vi oxygen ca
amino acid th ba tnh t amino acid to thnh cu trc alpha

(b) Xon beta: Cc chui polypepotide nm cnh nhau ni vi nhau bng cc lin kt
hydro to nn cc tm xp (pleated sheet)

Enzyme
ng vai tr xc tc cho cc phn ng sinh hc. Cc enzyme tham gia qu trnh
t nhn i ca DNA, qu trnh sao m ... cho thy tm quan trng ca loi protein
ny.

Protein cu trc
C mt trong hu ht cc cu trc ca t bo t mng t bo, mng nhn v cc
bo quan, bo tng v.v... cho n cu trc ca nhim sc th.

Protein vn chuyn
Cc protein ny nm trong cu trc ca cc h thng mng ca t bo hoc c
mt trong t bo nh hemoglobine trong hng cu ngi lm nhim v vn chuyn
oxygen hoc trong huyt tng v.v...

Protein vn ng
Mt s protein c kh nng co dui, to nn kh nng vn ng mc t bo
v mc c th. Cc protein myosine v actine ca t chc c vi kh nng co dui
to nn kh nng vn ng l mt v d minh ha.







48
Protein bo v
Khng th l nhng protein do h thng min dch ca c th sn xut, c chc
nng bo v c th chng li cc tc nhn c hi. Ngoi ra cng c th k n cc
protein c tnh cht bo v nh nh fibrinogen, thrombin tham gia vo qu trnh cm
mu ....

Hormone
Nhiu hormone c bn cht l protein ng vai tr ht sc quan trng trong qu
trnh iu ha, tng trng, trao i cht.



CU HI N TP
1. M t cu trc chung ca mt amino acid, chui polypeptide ?

2. Gii thiu cu trc khng gian cc bc ca phn t protein ?

3. Nu cc chc nng c bn ca protein ?



CU HI TRC NGHIM
1. Protein c cu trc a phn t vi n phn l amino acid. Cc amin acid c cu
trc chung:

A. Mt nhm amine, mt nhm phosphate, mt gc R c trng cho tng loi
aminoacid

B. Mt nhm amine, mt nhm carboxyl, mt gc R c trng cho tng loi
aminoacid

C. Mt base nitric, mt nhm carboxyl, mt gc R c trng cho tng loi
aminoacid

D. Mt base nitrid, mt ng pentose, mt gc R c trng cho tng loi
aminoacid

E. Mt nhm amine, mt base nitric, mt gc R c trng cho tng loi
aminoacid

2. Trong chui polypeptide, cc phn t amino acid ni vi nhau bi cc lin kt:
A. Lin kt hydro B. Lin kt peptide C. Lin kt phosphodiester
D. Lin kt ion E. Tt c u sai


3. Chc nng no di y khng phi l chc nng ca protein:
A. Cc enzyme v hormone

B. Protein cu trc hoc vn chuyn

C. T nhn i

D. Protein vn ng

E. Protein bo v



49
4. Trong chui polypeptide cc phn t amino acid ni vi nhau bi cc lin kt .........
(P: peptide, E: phosphodiester; H: hydro) c hnh thnh gia nhm ...........(A:
amine; C: carboxyl) ca amino acid ny vi nhm .......(A: amine; C: carboxyl) ca
amino acid k tip, phn ng s gii phng mt phn t nc.
A. P, A, C B. P, C, A C. H, A, C
D. H, C, C E. E, A, C




P N

1. B 2. B 3. C 4. A























































50
7. QU TRNH DCH M V IU HO SINH
TNG HP PROTEIN


Mc tiu
Trnh by c:

- Qu trnh gn amino acid v tRNA

- Cc giai on c bn trong qu trnh dch m

- Qu trnh iu ho sinh tng hp protein prokaryote v eukaryote



QU TRNH DCH M
GN AMINO ACID VO tRNA

Trong bo tng enzyme amino acyl-tRNA synthetase c hiu cho tng loi amino
acid s gn amino acid vo tRNA qua lin kt ester giu nng lng ti v tr
nucleotide adenine u 3' ca tRNA. Trong qu trnh dch m, b ba i m
(anticodon) ca tRNA s b sung vi b ba m sao (codon) trn mRNA trn c s
nguyn tc b sung m bo cc amino acid c lp t chnh xc vo chui
polypeptide. (hnh 1)





Hnh 1: Qu trnh gn amino acid c hiu (Phenylalanine) vo tRNA qua tc ng ca
enzyme amino acyl-tRNA synthetase v hin tng b sung gia b ba m sao (codon)
trn mRNA vi b ba i m (anticodon) trn tRNA.

RIBOSOME

Trong tt c t bo, ribosome tn ti di dng 2 tiu phn: tiu phn ln v tiu
phn b. Mi tiu phn cha cc rRNA vi chiu di khc nhau v cc phn t protein
khc nhau. Hnh 2 gii thiu thnh phn rRNA v protein tham gia vo cu trc ca
cc tiu phn ln v ca ribosome prokaryote v eukaryote. Lu l eukaryote,
rRNA 5,8S bt cp vi rRNA 28S.



51



Hnh 2: Cc thnh phn tham gia vo cu trc ca ribosome prokaryote v eukaryote.

CC GIAI ON CA QU TRNH DCH M

Qu trnh dch m xy ra ti ribosome qua ba giai on: giai on m u
(initiation), giai on ko di (elongation) v giai on kt thc (termination). Khng
c s khc bit ln trong qu trnh dch m gia prokaryote v eukaryote.

Giai on m u
Giai on ny c thc hin vi s tham gia ca mRNA, cc ribosome v cc
yu t khi ng (IF: Initiation Factor), eukaryote cc yu t t ny c k hiu l
eIF.

Trn mRNA, b ba m AUG, m ho cho methionine (Met) s bt u cho qu
trnh dch m. Tuy nhin cn lu l mc d ch c 1 codon m ho cho Met nhng
c ti hai loi tRNA mang Met n ribosome. tRNA
Met
kt hp vi codon nm
gia phn t mRNA v tRNA
iMet
kt hp vi codon v tr khi ng bt u cho
vic dch m, gn Met u tin vo chui polypeptide.

Vic dch m bt u khi tiu n v nh ca ribosome gn vi Met-tRNAiMet
bm vo mRNA mt v tr gn vi v tr ca codon AUG khi ng u 5' ca
mRNA vi s tham gia ca cc IF v nng lng do GTP cung cp. Hnh 3 gii thiu
cc bc trong giai on khi u eukaryote.

Giai on ko di
Giai on ny khng phc tp nh giai on m u. Sau khi amino acid u tin
(Met) c t vo ng v tr trn c s s bt cp theo nguyn tc b sung gia
anticodon trn tRNA vi codon tng ng trn mRNA, qu trnh ko di chui
polypeptide bt u.





52
Phc hp aminoacyl-tRNA k tip
s n xp vo ng v tr trn ribosome
nh cc EF (elongation factor: yu t
ko di). S tip xc gia peptidyl-tRNA
v aminoacyl-tRNA s lm hnh thnh
lin kt peptide gn amino acid mi
vo chui polypeptide ang tng hp
di s xc tc ca enzyme peptidyl
transferase. Qu trnh ny c lp i
lp li theo chiu t 5' n 3' trn
mRNA cho n khi tip xc vi du
hin kt thc dch m. Hnh 4 gii thiu
cc bc trong giai on ko di chui
polypeotide.













Hnh 3: Giai on m u. Cc tiu
n v ln v b ca ribosome ln lt gn
vi eIF6 v eIF3 to thnh phc hp bt
u dch m. (1) Phc hp 40S, eIF3 gn
vi eIF1 v phc hp eIF2, GTP v
tRNA
iMet
to thnh phc h tin khi u
(preinitiation complex). (2) mRNA gn vi
eIF4 v tr m (cap) v cho php mRNA
kt hp vi phc hp khi u to thnh
phc hp khi u (initiation complex). (3)
Phc h tin khi u cng vi eIF4 trt
trn mRNA n v tr codon khi u,
qu trnh i hi nng lng do ATP cung
cp, sau cc eIF c gii phng. (4)
Phc h tiu phn 60S v eIF6 ca
ribosome n gn vo tiu phn b vi s
h tr ca eIF5 v GTP














53
Hnh 4: Giai on ko di: (1)
Khi ribosome 80S v Met-tRNAiMet
nm v tr P (v tr ko di chui
polypeptide) ca ribosome, th phc hp
aa-tRNA mang amino acid th 2(aa2)
n codon tng ng trn mRNA ang
nm v tr A (v tr tip nhn amino
acid) ca ribosome. (2) S thu phn
ca GTP trong phc hp EF1-GTP
thnh EF1-GTP, qu trnh ny lm
thay i cu trc ca ribosome. (3)
Phn t rRNA ln xc tc cho qu trnh
to thnh lin kt peptide gia aa2 v
Met. (4) S thu phn ca GTP trong
phc hp EF2-GTP s gy ra tip s
thay i cu trc ca ribosome lm cho
ribosome i tip n codon tip theo
trn mRNA v tRNA
Met
tch Met tin
vo v tr E (v tr chun b gii phng
tRNA) v tRNA vi cc amino acid gn
vi nhau qua lin kt peptide s nm
v tr P ca ribosome. Qu trnh lp cc
phc hp tRNA mang aa3, aa4 ... tip
tc din ra theo cch trn.






























Giai on kt thc
Khi b ba m kt thc (UAA, UAG hoc UGA) c nhn bit bi cc RF
(Release Factor: yu t phng thch), phc hp peptidyl-tRNA lp tc tch thnh phn
t tRNA t do v chui polypeptide hon chnh c u l -NH2 v ui l


54
-COOH. Ribosome s tch tr li thnh hai tiu
n v bt u cho chu k dch m mi. (hnh
5)

















Hnh 5: Giai on kt thc: khi robosome di chuyn
n b ba m kt thc (UAA, UAG hoc UGA),
eRF1 s i vo phc hp ribosome v tr A hoc
gn cng vi eRF3-GTP. S thu phn guanine
eRF3-GTP s ct chui peptide khi tRNA v tr P,
gii phng cc tRNA v cc tiu phn ribosome.





POLYSOMES




Hnh 6: C ch gia tng tc dch m trn mRNA qua polysomes v
to nn cu trc dng vng ca mRNA.





55
tng cng tc tng hp protein (trung bnh eukaryote, mi protein c
tng hp trong vng 30 - 60 giy), nhiu ribosome s cng tham gia dch m trn
1mRNA gi l polyribosomes hay polysomes v tng cng tc quay vng ca cc
tiu n v ribosome sau khi chng hon tt qu trnh dch m. Nhiu nghin cu cho
thy cc protein gn vo ui poly A (PABPI) va c th tc ng trn ui polyA
u 3' va c th tc ng ln tiu n v 4G v 4E ca eIF4 u 5' lm cho mRNA
tr nn c dng vng to iu kin d dng cho cc tiu n v ca ribosome sau khi
kt thc dch m u 3' c th bt u ngay qu trnh dch m mi u 5' (hnh 6).

IU HO S BIU HIN CA GENE
IU HO S BIU HIN CA GENE PROKARYOTE

prokaryote mc ch ca vic iu ho s biu hin ca gene nhm iu chnh
h enzyme cho ph hp phc v cho qu trnh tng trng v sinh sn ca t bo.
Tu thuc vo iu kin mi trng m cc gene m ho cho cc enzyme c th c
biu hin hoc ngng biu hin bt c mt thi im no trong chu k t bo.

Qu trnh iu ho ch yu xy ra qu trnh phin m thng qua operon. Mi
operon bao gm:

- Mt s gen cu trc m ho cho cc polypeoptide nm gn nhau gi l
cistron.

- Cc trnh t DNA khc tham gia vo hot ng iu ho vi nhng vai tr
khc nhau.

Cn c vo cch thc iu ho ngi ta chia lm hai loi operon chnh: operon
cm ng v operon km hm.

Operon cm ng (inducible operon)
Mt v d in hnh cho kiu operon cm ng ny l operon lactose.

ng lactose ng vai tr ca cht cm ng. Khi khng c lactose cc gene m
ho cho enzyme chuyn ho lactose s khng hot ng phin m. Khi lactose l
ngun carbohydrat duy nht c mt trong mi trng thi vi khun s sn xut cc
enzyme cn thit cho qu trnh chuyn ho lactose qua hot ng phin m cc gene
m ho cho cc enzyme ny. Hnh 7 m t qu trnh iu ho hot ng phin m ca
operon lac E.coli.

(a) Khi mi trng khng c lactose, mRNA c tng hp rt t do mt protein
gi l lac repressor (yu t km hm) gn vo operator, ngn cn hot ng phin m
ca RNA polymerase-70 (hnh 7a).

(b) Khi mi trng c c glucose v lactose, mt phn lac repressor s gn vi
lactose v gii phng operator cho php RNA polymerase-70 thc hin phin m
cng thp (hnh 7b).

(c) Khi mi trng ch c lactose v khng c glucose, qu trnh phin m s
din ra mc ti a do tnh trng khng c glucose, cAMP s to thnh phc hp
CAP-cAMP, gn vo v tr CAP v qua tc ng ln RNA polymerase thc y
qu trnh phin m (hnh 7c).




56




Hnh 7: iu ho qu trnh phin m kiu cm ng ca operon lac E. coli: Vng kim
sot phin m di khong 100 cp base gm 3 vng gn protein: (1) V tr CAP gn protein
hot ho qu trnh chuyn ho (catabolite activator protein); (2) lac promotor (vng khi
ng lac) gn vi phc h RNA polymerase-(70; (3) lac operator (vng vn hnh lac ). Gene
lacZ l mt trong 3 gene ca operon.

(a) Khi mi trng khng c lactose.

(b) Khi mi trng c c glucose v lactose.

(c) Khi mi trng ch c lactose v khng c glucose

Operon km hm (repressible operon)
Mt v d in hnh cho kiu operon cm ng ny l operon tryptophan E. coli.
Amino acid tryptophan ng vai tr ca cht cm ng (inducer) gn vi
repressor. Triptophan s gn vi trp repressor lm thay i cu trc ca trp repressor
lm cho n c kh nng gn vi trp operon c ch khng cho operon ny phin m.
Khi nng tryptophan trong mi trng v trong bo tng cao, t bo s khng
tng hp cc enzyme c m ho trn trp operon xc tc cho qu trnh tng
hp tryptophan.






57
Khi nng tryptophan trong mi trng v bo tng thp, trp repressor s
khng c kh nng c ch trp operon v cho php thc hin phin m tng hp cac
enzyme cn thit cho vic tng hp triptophan. (hnh 8)



Hnh 8: iu ho qu trnh phin m kiu km hm ca trp operon E. coli. Khi nng
tryptophan trong mi trng v bo tng thp, trp repressor s khng c kh nng c ch
trp operon v cho php thc hin phin m tng hp cac enzyme cn thit cho vic tng
hp tryptophan. Khi nng tryptophan trong mi trng v trong bo tng cao, t bo s
khng tng hp cc enzyme c m ho trn trp operon xc tc cho qu trnh tng hp
tryptophan.

IU HO S BIU HIN CA GENE EUKARYOTE

M hnh iu ho biu hin ca gene eukaryote v prokaryote c mt s im
ging nhau nh vai tr ca cc protein gn vi DNA nh hng n hot ng ca
enzyme RNA polymerase trong vic bt u qu trnh phin m. Tuy nhin trong vic
iu ho s biu hin ca gene ca eukaryote vn c nhng im khc bit: (1) Cc
gene ca eukaryote khng to thnh cc operon v him khi cc gene c phin m
vi nhau vo cng mt mRNA thay vo mi gene c mt promotor ring v c
phin m mt cch ring r. (2) Cu trc ca chromatin nh hng n s biu hin
ca gene eukaryote, c th phin m DNA phi tho xon khi cc protein
histone. (3) Mc d cc cht hot ho v c ch u c mt trong qu trnh iu ho
biu hin ca gene eukaryote v prokaryote, nhng eukaryote cc cht hot ho c
vai tr ph bin hn. (4) C ch iu ho biu hin ca gene eukaryote lin quan n
nhiu c ch khc nhau tc ng n nhiu khu khc nhau trong qu trnh truyn
thng tin t DNA n protein. Di y gii thiu vn tt mt s yu t lin quan n
vic iu ho s biu hin ca cc gene eukaryote.

Mi lin quan gia cu trc ca chromatin v s biu hin ca gene
Cu trc chromatin c ch s biu hin ca gene, gene c th phin m, cc
yu t phin m, cc cht hot ho, RNA polymerase phi gn vo DNA do trc
khi phin m, cu trc chromatin phi thay i. S thay i ny lin quan n mt s
hin tng sau:



58
V tr nhy cm vi enzyme DNase I
Trn DNA c nhng v tr nhy cm vi enzyme DNase I, mt enzyme ging ho
DNA ch tc ng ln DNA khi c s lng lo trong cu trc ca cromatin. V tr ny
nm trc v tr bt u phin m khong 1000 bp. S c mt ca nhng v tr ny gi
chromatin vng ny tr nn lng lo hn khi
DNA vng ny thc hin phin m. V tr ny l
ni gn ca cc protein iu ho iu khin
hot ng phin m.

Acetyl ho protein histone
Vic acetyl ho histone s lm thay i cu
trc ca nucleosome v to iu kin thun li cho
qu trnh phin m. (hnh 9)

Qu trnh demethyl ha DNA
S methyl ho s thc y cc enzyme
deacetylase tch cc nhm acetyl ra khi cc
protein histone do s lm cho cu trc ca
chromatin tr nn hng nh v qua c ch qu
trnh phin m. Hin tng demethyl ho
(demethylation) ngc li s lm cn tr qu trnh
trn do to thun li cho qu trnh phin m.

KIM SOT QU TRNH PHIN M

Cc protein iu ho qu trnh phin m c
th nh hng ti vic bt u phin m thng qua
vic lm hng nh hoc tp trung cc thnh phn

Hnh 9: Acetyl hoa histone lam
thay i cu truc acuchromatin
cho phep cac yut phin ma gn vao
DNA.
trong h thng phin m. Mt s cc protein ny l cc cht hot ho (activator) kch
thch hot ng phin m v mt s protein khc ng vai tr ca cht c ch
(repressor).

Vng tng cng v vng c lp
Mt s cht hot ho gn vo vng tng cng (enhancer) nm cch xa gene
qua kch thch qu trnh phin m. Bn cnh cc vng c lp (insulator) c tc
dng c lp hoc cch ly vng tng cng lm c ch qu trnh phin m.(hnh 10)













Hnh 10: Insulator (vng c lp) nh ch tc ng ca enhancer (vng tng cng) ln
promotor (vng khi ng) khi nm gia enhancer v promoter.





59
Cc yu t a p ng (multiple response elements)
Mt gene c th c hot ho bi nhiu yu t p ng khc nhau, nhng yu
t ny c mt c vng tng cng v vng khi ng (promotor). Cc yu t a p
ng cho php cng mt gene c th p ng vi nhiu tc nhn kch thch khc nhau.
Trong cng mt thi im, s c mt ca cng mt yu t p ng nhng gene khc
nhau cho php mt kch thch c th tc ng cng mt lc ln nhiu gene khc nhau.
Bng cch ny nhng yu t p ng s cho php cc gene c lin quan vi nhau trong
cc hot ng chuyn ho phc tp c th phi hp vi nhau (hnh 11).




Hnh 11: Cc yu t a p ng (MRE) nm pha trc gene m ha cho metallothionein.
Cc protein hot ha gn vi MRE kch thch hot ng phin m.

KIM SOT S BIU HIN CA GENE THNG QUA QU TRNH HON THIN
mRNA

Qu trnh hon thin mt mRNA sau khi phin m c th din ra theo cc cch
kiu khc nhau. S thay i v tr ct trn tin mRNA s lm xut hin cc mRNA
trng thnh khc nhau v tng hp nn cc loi protein khc nhau.(hnh 12).

KIM SOT S BIU HIN CA GENE THNG QUA S HNG NH CA RNA

S hng nh ca mRNA s nh hng n s biu hin ca gene thng qua thi
gian tn ti ca mRNA v qua nh hng n hot ng dch m.

S hng nh ca mRNA ph thuc v u 5' (5' cap), ui poly A, vng khng
dch m (UTR: untranslated region) 5' v 3' v c vng mang m.

QU TRNH LM IM LNG RNA (RNA silencing)

Qu trnh lm im lng RNA c bt u khi mt phn t RNA chui kp
(double stranded RNA) c ct v hon thin bi enzyme Dicer to thnh nhng
on RNA xen nh (siRNA: small interfering RNA). (hnh 13)

siRNA s n b sung vi mt trnh t trn mRNA trn c s thu ht mt
phc hp RNA - protein n ct mRNA on gia siRNA, sau khi b ct mRNA s
tip tc thoi ho.

siRNA cng c th vo trong nhn, b sung vi mt on DNA v kch thch qu
trnh methyl ho on DNA ny v qua nh hng n hot ng phin m.




60
Hnh 12: Vic ct nhng v tr
ct khc nhau trn tin mRNA s
dn n s to thnh cc mRNA
m ha cho cc protein khc nhau






















































KIM SOT QU TRNH DCH M V SAU DCH M

Vic khi ng qu trnh dch m c th b nh ch do cc protein c hiu gn
vo u 5' ca mRNA ngn cn s tip xc ca cc ribosome. Cc ribosome, tRNA,
cc yu t khi u v ko di chui polypeptide cng nh hng n qu trnh dch
m.




61
Rt nhiu protein ca
Eukaryota tip tc bin i
sau khi phin m bng cch
ct mt cch chn lc, ta b
cc amino acid cc u tn
cng, acetyl ho, b sung
cc nhm phosphate,
carbohydrate, nhm methyl
vo phn t protein. Nhng
thay i ny s nh hng
n vic vn chuyn, chc
nng v hot ng ca cc
protein ny.



Hnh 13: Qua trnh RNA
silencing dn nsthoai aho
cua mRNA hoc methyl
DNA





CU HI N TP
1. M t qu trnh gn amino acid vo tRNA ?

2. Hy nu mt s c im trong cc bc ca qu trnh gii m ?

3. Polysomes l g ?

4. Qu trnh iu ho s biu hin ca prokaryote din ra nh th no ?

5. Qu trnh iu ho s biu hin ca eukaryote din ra nh th no ?
hoa



CU HI TRC NGHIM
1. tng cng tc tng hp protein, nhiu ribosome s cng tham gia dch m
trn 1mRNA gi l:
A. Lysosome B. Polysomes C. Chromosome
D. Replicon E. Nucleosome


2. t bo nhn ribosome c hnh thnh t 2 tiu phn:
A. Tiu phn ln gm 50 phn t protein v 3 phn t rRNA; Tiu phn b gm
33 phn t protein v 2 phn t rRNA

B. Tiu phn ln gm 45 phn t protein v 1 phn t rRNA; Tiu phn b gm
33 phn t protein v 3 phn t rRNA





62
C. Tiu phn ln gm 50 phn t protein v 3 phn t rRNA; Tiu phn b gm
33 phn t protein v 1 phn t rRNA

D. Tiu phn ln gm 34 phn t protein v 2 phn t rRNA; Tiu phn b gm
21 phn t protein v 1 phn t rRNA

E. Tiu phn ln gm 45 phn t protein v 2 phn t rRNA; Tiu phn b gm
33 phn t protein v 1 phn t rRNA

3. Hai tiu phn ln v b kt hp li vi nhau hnh thnh nn ribosome khi:
A. Cc tiu phn ra khi nhn vo bo tng

B. Cc rRNA c tng hp xong ti hch nhn

C. Hon tt vic tng hp protein

D. giai on m u ca qu trnh tng hp protein

E. ARN mang amino acid n mRNA

4. Trong qu trnh gii m, amino acid t do:

A. trc tip n ribosome phc v cho qu trnh gii m

B. n ribosome di dng c ATP hot ha

C. c hot ha thnh dng hot ng nh ATP, sau lin kt vi tRNA c
hiu to nn phc h amino acid - tRNA, qu trnh ny din ra di tc dng ca cc
enzyme c hiu

D. c gn vi tRNA nh enzyme c hiu to thnh phc hp amino acid
tRNA phc v cho qu trnh gii m ribosome

E. kt hp vi tiu phn b ca ribosome tham gia vo qu trnh gii m

5. Trong qu trnh gii m di truyn, ribosome s:
A. Tr li dng rRNA sau khi hon thnh vic tng hp protein

B. Bt u tip xc vi mRNA t b ba m UAG
C. Trt t u 5' n 3' trn mRNA
D. Gi nguyn cu trc sau khi hon thnh gii m

E. Ch tham gia gii m cho 1 loi mRNA

6. Trong qu trnh gii m, amino acid n sau s c gn vo chui polypeptide
ang c hnh thnh:

A. Khi tiu phn ln v b ca ribosome tch nhau

B. Trc khi tRNA ca amino acid trc tch khi ribosome di dng t do

C. Khi ribosome i khi b ba m khi u

D. Khi ribosome di chuyn n b 3 m tip theo

E. Gia nhm carboxyl ca amino acid mi kt hp vi nhm amin ca amino
acid trc hnh thnh mt lin kt peptide

7. Qu trnh gii m kt thc khi:

63
A. Ribosome ri khi mRNA v tr li dng t do vi 2 tiu phn ln v b

B. Ribosome di chuyn n m b 3 AUG

C. Ribosome gn amino acid mtinin vo v tr cui cng ca chui polypeptide

D. Ribosome tip xc vi 1 trong cc m b 3 UAU, UAX, UXG

E. Ribosome tip xc vi 1 trong cc m b 3 UAA, UAG. UGA

8. Chui polypeptide sau khi c tng hp xong bi qu trnh gii m s:

A. hon thin cu trc khng gian bc 2, 3 hoc bc 4 tr thnh nhng protein
c hot tnh sinh hc

B. c hon thin cu trc khng gian trong li ni sinh cht hoc bo tng

C. C th kt hp vi cc protein khc to nn nhng phn t protein c cu
trc bc 4

D. A v C ng

E. A, B v C u ng

9. Trong qu trnh gii m, khi ribosome tin ti m b 3 tip theo th hin tng no
sau y s xy ra:
A. Lin kt peptide gia 2 amino acid c v mi s c hnh thnh

B. tRNA mang amino acid n ribosome tip xc vi b 3 m sao qua b 3 i
m

C. Chui polypeptide hon thin cu trc khng gian bc 2, 3

D. Chui polypeptide mi c tng hp s tch khi ribosome

E. Phn t amino acid tng ng vi b 3 m sao m ribosome mi trt n s
n ribosome v gn vo chui polypeptide ang c tng hp

10. S hon thin cu trc khng gian ca chui polypeptide t bo c nhn xy ra
:

A. Ty th v li ni sinh cht

B. T bo cht v li ni sinh cht

C. B Golgi v li ni sinh cht

D. Lysosome v th vi

E. Trung th v thoi v sc

11. vi khun, trong c ch iu ha sinh tng hp protein bng operon cm ng,
cht cm ng c vai tr:
A. Hot ha enzyme RNA polymerase

B. c ch vng vn hnh (operator), ngn cn qu trnh tng hp cc yu t km
hm


C. Hot ha vng promotor (vng khi ng)

D. Gii c ch v kch thch hot ng phin m ca cc gen cu trc

64
A. Ribosome ri khi mRNA v tr li dng t do vi 2 tiu phn ln v b

B. Ribosome di chuyn n m b 3 AUG

C. Ribosome gn amino acid mtinin vo v tr cui cng ca chui polypeptide

D. Ribosome tip xc vi 1 trong cc m b 3 UAU, UAX, UXG

E. Ribosome tip xc vi 1 trong cc m b 3 UAA, UAG. UGA

8. Chui polypeptide sau khi c tng hp xong bi qu trnh gii m s:

A. hon thin cu trc khng gian bc 2, 3 hoc bc 4 tr thnh nhng protein
c hot tnh sinh hc

B. c hon thin cu trc khng gian trong li ni sinh cht hoc bo tng

C. C th kt hp vi cc protein khc to nn nhng phn t protein c cu
trc bc 4

D. A v C ng

E. A, B v C u ng

9. Trong qu trnh gii m, khi ribosome tin ti m b 3 tip theo th hin tng no
sau y s xy ra:
A. Lin kt peptide gia 2 amino acid c v mi s c hnh thnh

B. tRNA mang amino acid n ribosome tip xc vi b 3 m sao qua b 3 i
m

C. Chui polypeptide hon thin cu trc khng gian bc 2, 3

D. Chui polypeptide mi c tng hp s tch khi ribosome

E. Phn t amino acid tng ng vi b 3 m sao m ribosome mi trt n s
n ribosome v gn vo chui polypeptide ang c tng hp

10. S hon thin cu trc khng gian ca chui polypeptide t bo c nhn xy ra
:

A. Ty th v li ni sinh cht

B. T bo cht v li ni sinh cht

C. B Golgi v li ni sinh cht

D. Lysosome v th vi

E. Trung th v thoi v sc

11. vi khun, trong c ch iu ha sinh tng hp protein bng operon cm ng,
cht cm ng c vai tr:
A. Hot ha enzyme RNA polymerase

B. c ch vng vn hnh (operator), ngn cn qu trnh tng hp cc yu t km
hm


C. Hot ha vng promotor (vng khi ng)

D. Gii c ch v kch thch hot ng phin m ca cc gen cu trc

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E. V hiu ha yu t km hm (repressor)

12. vi khun E. Coli, cht c ch trong c ch iu ha sinh tng hp qua operon
km hm, cht cm ng hot ng theo cch:
A. Gn vo vng khi ng (promotor)v lm mt tc dng ca vng ny

B. Gn vo vng vn hnh (operator) cn tr s hot ng ca enzyme ARN
polymerase

C. Kt hp vi enzyme polymerase lm mt kh nng xc tc cho qu trnh phin
m ca gen ny

D. Gn vi cht c ch (repressor), lm cht ny c kh nng c ch vng vn
hnh (operator) khng cho operon thc hin phin m

E. A v C ng



P N

1. B 2. C 3. D 4. C 5. C 6. E 7. E 8. E 9. B 10. B

11. E 12. D











































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