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S phin m - VOER

S phin m

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S phin m (Transcription)
Qu trnh tng hp phn t RNA t khun DNA c gi l qu trnh phin m (transcription).
Nhng thng tin di truyn c trong phn t DNA c s dng tng hp protein. Tuy nhin, phn t DNA
khng phi l cht lm khun tng hp protein, bi v, cc phn t DNA tp trung ch yu trong nhim sc
th,phn b trong nhn t bo, cn qu trnh tng hp protein li din ra trong t bo cht. Nh vy, phi c
mt cht trung gian lm nhim v chuyn thng tin di truyn t DNA trong nhn ra t bo cht v lm khun cho
qu trnh tng hp protein.
Nm 1957, sau khi khm ph ra cu trc phn t DNA, Francis Crick nu tng v mt cht trung gian
truyn thng tin di truyn cho qu trnh tng hp protein, v d on "ngi phin dch" trung gian l RNA.
D on ny c khng nh bng cc nghin cu chng minh RNA c cu to ha hc tng t nh DNA
v c tng hp trn khun ca DNA.
Nm 1960, c 3 nhm nghin cu c lp M chit tch c enzyme RNA-polymerase, enzyme ny c
kh nng tng hp phn t RNA t khun DNA.
Benjamin Hall v Sol Spiegelman (1961) tch v tinh sch RNA v DNA t thc khun th T2 . Ly DNA ca
T2 lm bin tnh tch thnh 2 si n. Khi cho cc si n DNA bt cp vi si n RNA th chng to thnh
dng phn t lai DNA/RNA. Tuy nhin, khi h ly DNA ca cc c th khc trn vi RNA ca T2 th chng khng
bt cp, ngha l khng to phn t lai. iu chng t rng, RNA ca T2 c tng hp trn khun DNA
ca n theo c ch b sung nn chng c th bt cp v to thnh phn t lai.
Nhng bng chng khc nh RNA tp trung ch yu t bo cht nhng chng cng c mt trong nhn t
bo. nhng loi t bo m s tng hp protein xy ra mnh m th hm lng RNA tp trung nhiu hn so
vi nhng t bo t tng hp protein.
Nh vy, RNA chnh l cht trung gian truyn thng tin di truyn tng hp protein. Mi quan h gia DNA,
RNA v protein c th hin nh sau

Mi quan h gia DNA, RNA v protein

V sau, ngi ta cn pht hin qu trnh phin m ngc, ngha l, phn t DNA c tng hp trn khun ca
phn t RNA. Qu trnh phin m ngc c enzyme phin m ngc (reverse transcriptase) xc tc. Enzyme
ny c pht hin ln u tin khi nghin cu s sao chp b gen ca virus gy ung th (retrovirus). u tin,

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enzyme phin m ngc s tng hp si DNA n (cDNA) trn khun ca phn t RNA, sau , enzyme DNApolymerase s tng hp mch b sung to thnh si DNA kp (cDNA kp). Qu trnh c th biu din qua s
sau:

Qu trnh phin m ngc

c im chung ca qu trnh phin m

S phin m c thc hin theo mt s nguyn tc chung sau
- Sn phm phin m l mt si polyribonucleotide mch n (RNA).
- thc hin phin m, phn t DNA phi dn xon cc b. i vi mi gen, ch mt trong hai si DNA c
chn lm khun tng hp phn t RNA. Enzyme RNA-polymerase s quyt nh vic chn si lm khun v
xc tc qu trnh tng hp phn t RNA.
- Phn t RNA mi c tng hp theo chiu t u 5'phosphat n u 3'OH.
- S phin m c thc hin khi c mt ca cc enzyme v 4 loi nucleotide l GTP, CTP, UTP v ATP.
- Phn t RNA c tng hp theo c ch b sung tng t nh s sao m DNA nhng i din vi A trn
si DNA s l U trn RNA.
- Khng c c ch sa sai i km vi qu trnh phin m, do vy, nhng sai st s dn n s sai khc trong
biu hin gen. Tuy nhin, s sai khc trong qu trnh phin m khng nh hng g n vic truyn t thng
tin di truyn cho th h sau.

Phin m tng hp mRNA procaryote

Enzyme RNA-polymerase procaryote
cc sinh vt procaryote ch c mt loi enzyme RNA-polymerase, n chu trch nhim xc tc qu trnh tng
hp c ba loi RNA l mRNA, tRNA v rRNA. RNA-polymerase ca vi khun E. Coli c nghin cu k nht.
Enzyme ny c cu trc bc 4 rt phc tp, bao gm hai hp phn chnh l enzyme li v nhn t . Nhn t
c th tch ra khi enzyme li, c trng lng phn t l 70.000dalton (1dalton=1,66x10-24g=1/12 khi lng
nguyn t carbon). Enzyme li c cu to t 5 chui polypeptide: hai chui , mt chui , mt chui ' v
mt chui . Ngoi ra, enzyme li cn c nhn t phin m l v nus A. Cc nhn t phin m ny d dng
tch ra khi enzyme li, nn trong qu trnh tch v tinh sch enzyme thng khng tm thy.
Cc tiu n v RNA-polymerase ca E.Coli
C c tiu n v RNA-polymerase ca E.C oli
Tiu n v

S lng

Trng lng phn t(dalton)

C hc nng

' (beta)

160.000

- lin kt vi DNA khun

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(beta)

150.000

- ko di mch RNA

(alpha)

42.000

- cha r

(omega)

11.000

- cha r

(sigma)

70.000

- nhn bit promoter

(rho)

46.000

- kt thc phin m

Nus A

70.000

- ko di v kt thc

T liu da theo S.C. Rastogi

Gia cc si polypeptide trong enzyme li khng c lin kt ng ha tr bn vng m ch hnh thnh cc lin
kt th cp. Trng lng phn t ca cc chui polypeptide c biu din bng trn.
Chc nng ca chui ' l lin kt vi si DNA khun, cn chui c tc dng xc tc hnh thnh lin kt
phosphodiester. Vai tr ca hai chui hin cha bit r.
Nhn t ng vai tr quan trng trong vic tng hp RNA, n gip cho enzyme li c th khi u qu trnh
phin m ti mt im c th.
Nhiu th nghim (in vitro) cho thy, khi thiu nhn t , enzyme li s khi u phin m mt cch ty tin,
khng ng v tr quy nh. E. Coli, ngi ta tm thy 3 loi nhn t , mi loi c trng lng phn t
ring. RNA-polymerase pht hin ra promoter, sau gn ln phn t DNA, m xon cc b v thc hin qu
trnh tng hp phn t RNA. S c mt ca Mg+2 l cn thit cho hot ng xc tc ca enzyme RNApolymerase.

Cu trc RNA-polym erase procaryote

Phin m tng hp mRNA procaryote

c im ca mRNA procaryote l mi phn t mang d liu thng tin cho s tng hp nhiu si polypeptide
ng thi (polycistronic mRNA).
Phin m tng hp mRNA procaryote c nghin cu kh k vi khun E. Coli. Qu trnh ny c th chia
lm 3 giai on: khi ng, ko di v kt thc.

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Giai on khi ng
Tiu n v ca enzyme RNA-polymerase nhn bit v gn enzyme vo promoter khi ng qu trnh
phin m. Promoter l on phn t DNA c cu trc c bit, gip enzyme RNA- polymerase nhn bit v tr bt
u ca s phin m.
Enzyme RNA-polymerase gn vo trnh t 35 ca ptomoter, trt dc theo phn t DNA n trnh t 10 th
m xon, lm l si lm khun, on m xon c di khong 1217 nucleotide, qu trnh phin m bt u
ti v tr xc nh.

Khi u phin m prom oter UV5 operon lactose

Sau khi tng hp c on RNA ngn khong 810 nucleotide th nhn t tch khi enzyme li v si khun
DNA, kt thc giai on m u. Sau , nhn t c th kt hp vi mt enzyme li khc khi u mt s
phin m mi.

Giai on ko di
Sau khi nhn t tch khi phc hp, mt nhn t ko di l protein Nus A gn vo, enzyme DNA-polymerase
tip tc chuyn dch dc theo gen, lm gin xon si DNA v thc hin qu trnh polymer ha, ko di si RNA
mi. Si mRNA c tng hp theo nguyn tc b sung: A i din vi U, T i din vi A, G i din vi C v
C i din vi G, nh sau

Giai on ko di

Khi enzyme RNA-polymerase trt dc theo gen n u th s tho xon v tng hp mi c thc hin n
, cn on gen sau khi tho xon, phn t RNA-polymerase trt qua ri th c xon tr li cu trc
ban u.

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S b iu din qu trnh tng hp RNA

Giai on kt thc
Trn si DNA khun c cha on m kt thc, c chc nng lm dng qu trnh phin m im qui nh. Ti
im kt thc, enzyme RNA- polymerase dng qu trnh ni cc nucleotide, gii phng phn t RNA mi tng
hp ra khi phc h enzyme v DNA khun, ng thi, enzyme cng tch khi si DNA c th n vi mt
trnh t khi ng mi.

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C hai kiu kt thc phin m

- Th nht: Enzyme RNA-polymerase tip nhn tc nhn kt thc .
- Th hai: Kt thc xy ra khng cn s c mt ca tc nhn .
Khi nghin cu v cch kt thc phin m theo kiu th hai vi khun v phage, ngi ta nhn thy c hai
im c trng v cch sp xp cc baz nit on cui trn si lm khun nh sau
- C 2 trnh t i xng b sung pha trc im kt thc.
- Mt on poly A nm ngay trc im kt thc
Nh cu trc c bit nh vy nn sau khi hai trnh t i xng b sung c hnh thnh trn si RNA, chng
c th bt cp vi nhau, to thnh cu trc c dng "kp tc", ngn cn RNA-polymerase phin m tip tc,
ng thi, c tc dng ko si RNA mi c tng hp ra khi phc h enzyme v si khun. Qu trnh tng
hp s dng li on poly-U.

S b iu din cch hnh thnh cu trc dng "kp tc"

Ti cc v tr kt thc cn s c mt ca protein kt thc phin m , cu trc ca on DNA khun thng thy

thiu on poly-A trc im kt thc v khng phi lc no cng c cc trnh t i xng b sung c th
hnh thnh dng "kp tc" nu trn.
Hin nay, chng ta cn cha bit r, bng cch no m nhn t tc ng ln qu trnh kt thc phin m. C
th n s dng nng lng t phn t ATP thc hin s tc ng ny.
Mt c im ni bt t bo sinh vt procaryote l cc phn t DNA nm trong t bo cht, nn qu trnh
phin m v dch m c tin hnh ng thi.

Phin m tng hp mRNA eucaryote

S phin m eucaryote xy ra phc tp hn nhiu so vi procaryote. c im ca mRNA eucaryote l mi
phn t mang d liu thng tin cho s tng hp mt polypeptide. Qu trnh phin m c thc hin trong nhn
t bo, cn s dch m xy ra t bo cht, hai qu trnh ny xy ra hai v tr khc nhau.

Enzyme RNA-polymerase eucaryote

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t bo eucaryote c 3 loi enzyme RNA-polymerase, mi loi m nhn mt chc nng ring bit, RNApolymerase I chu trch nhim tng hp rRNA; RNA-polymerase II chu trch nhim tng hp mRNA v RNApolymerase III chu trch nhim tng hp cc loi RNA c kch thc nh nh tRNA, RNA 5S ribosome v cc
phn t RNA nh khc.
Cu trc ca cc RNA-polymerase sinh vt eucaryote rt phc tp, c trng lng phn t ln v cng c
hnh thnh t nhiu tiu n v nh enzyme RNA-polymerase sinh vt procaryote. S lng tiu n v cu
to thng khong 10.
Mi loi RNA-polymerase chu trch nhim chnh tng hp mt loi RNA, do vy, cc RNA-polymerase khc
nhau s c kh nng nhn bit promoter ca mnh khi u phin m ng v tr. Nhiu kt qu nghin cu
cho thy, eucaryote, trnh t on DNA promoter di hn so vi t bo procaryote, c bit l i khi cn c
s hin din ca nhm trnh t khuych i (enhancer), c tc dng lm tng biu hin gen.
Ngoi 3 loi RNA-polymerase ch yu k trn, trong t bo sinh vt eucaryote cn c cc RNA-polymerase ca
ty th v ca lp th. Cc enzyme ny chu trch nhim phin m cc gen nm trn DNA ty th v lp th.

Qu trnh phin m eucaryote

DNA ca t bo eucaryote tp trung ch yu trong cc nhim sc th trong nhn, c phn t lng ln v cha
nhiu gen. Phn ln cc gen, phn mang m tng hp protein, c s an xen gia cc on mang m
(exon) v cc on khng mang m (intron). Do vy, qu trnh phin m xy ra phc tp hn v thng bao
gm hai qu trnh chnh
- Qu trnh phin m to nn phn t mRNA u tin, gi l phn t tin mRNA.
- Qu trnh bin i phn t tin mRNA thnh phn t mRNA trng thnh.
Qu trnh phin m to nn phn t tin mRNA cng c thc hin theo cc giai on tng t nh qu trmh
phin m t bo procaryote: Tc l, enzyme RNA-polymerase cng vi tc nhn khi ng phin m nhn
bit promoter v khi ng phin m v tr chnh xc, tip theo l qu trnh ko di mch mRNA v kt thc
phin m ti v tr xc nh. Enzyme RNA- polymerase II s chu trch nhim phin m tng hp phn t mRNA.
Phn t tin mRNA tri qua mt s bin i mi tr thnh phn t mRNA trng thnh. Qu trnh bin i bao
gm: gn m, hnh thnh ui polyA v loi b cc intron, ni cc exon.

Gn m (capping)
Phn t mRNA c tng hp theo hng t u 5P n u 3OH. Sau khi phn t tin mRNA hnh thnh
c mt on th u 5P c gn thm mt nucleotide l 7methylguanosine (Hnh 2-11). Vi chiu
ngc li to lin kt phosphodiester gia hai cacbon th 5 ca hai nucleotide. To mch lin kt 55 thng
c gi l m. Sau khi c gn m th u 5 ca phn t mRNA cng s c nhm 3OH t do gc
ng ging nh u 3. Tuy vy, nh nhm metyl (CH3) v tr nit th 7 ca baz guanosine c th phn
bit c u 5. Nh vy, sau khi gn m, u 5 khng cn nhm phosphat t do na. M gip cho
ribosome nhn bit v gn vo u 5 ca mRNA, khi u dch m ng v tr qui nh, ng thi, m bo v
mRNA khi b cc enzyme nuclease phn hu.

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Cu to "m " chp u 5' ca m RNA

Hnh thnh ui poly-A u 3OH

Ngi ta pht hin rng, trong phn ln cc phn t mRNA ca nm men v t bo eucaryote, u 3 c cha
mt trnh t di khong 200 nucleotide vi cc baz adenine (A) gi l ui poly-A. ui poly-A l im c
trng ca cc phn t mRNA eucaryote. cc loi RNA khc nh rRNA, tRNA khng c ui poly-A. ui
poly-A c gn vo sau khi phn t tin mRNA c tng hp xong. Enzyme chu trch nhim gn ui
poly-A vo tin mRNA l poly-A-polymerase.
Qu trnh hnh thnh ui poly-A c tin hnh theo m t Hnh 8. u 3'OH ca phn t tin mRNA c
cha mt trnh t nhn bit l AAUAAA. Mt enzyme endonuclease c hiu nhn bit v ct si tin mRNA
mt v tr khong t 11 n 30 nucleotide sau trnh t nhn bit v pha u 3'OH, tip sau , enzyme poly-Apolymerase s xc tc gn cc nucleotide A vo u 3'OH to thnh ui poly-A. di ui poly-A ca cc
mRNA thay i tu theo loi v giai on pht trin ca t bo.
Khi mRNA di chuyn t nhn ra t bo cht cng nh trong qu trnh tn ti, ui poly-A b thoi ha ngn dn.
ui poly-A cng di, thi gian tn ti ca mRNA trong t bo cng lu. Trong mt s t trng hp, mRNA
khng c ui poly-A nh cc mRNA phin m t gen m ha histon khng c ui poly-A.

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S v s hnh thnh ui poly-A

Ct b intron v ni cc exon (splicing)

Cc gen sinh vt eucaryote thng rt di. Phn ln c cha cc on khng mang m cho protein (intron)
xen ln vi on mang m (exon). Nhiu gen c tng di ca cc intron ln hn tng di ca cc exon. V
d nh gen m ha cho -globulin c 2 intron: Mt intron nh c di 116 cp baz v mt intron ln c
di l 646 cp baz, thm vo , u 5 c 52 cp baz v u 3 c 110 cp baz khng mang m cho
protein. Tng di ca cc on khng mang m ln hn tng di ca cc on mang m
Tng t nh vy, gen m ha cho ovalbumin cng c tng di ca cc on intron ln hn tng di
ca cc on exon cng li.

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So snh di ca m RNA ca -glob ulin ca chut vi di ca gen m ha

i vi cc gen c cha intron th sau khi hnh thnh bn phin m u tin phi tri qua giai on ct b intron
v ni cc exon to thnh phn t RNA trng thnh. Qu trnh ny xy ra theo hai bc chnh: Cc
endonuclease thc hin vt ct ranh gii gia intron v exon, tip theo l s ni cc exon li v loi b
intron.
Trong qu trnh loi b intron v ni cc exon c s tham gia ca cc phn t ghp ni (spliceosome) l
phc hp gia RNA c phn t lng nh giu uracil c mt trong nhn t bo vi mt s protein chuyn bit
trong nhn.
Sau khi cc intron c loi b, cc exon c ni li, phn t mRNA trng thnh lc ny c m chp u
5, ui poly-A u 3 v cc trnh t mang m cho protein. mRNA trng thnh s i qua l mng nhn vo t
bo cht kt hp vi ribosome thc hin qu trnh tng hp protein.
Tuy nhin, khng phi tt c cc gen trong t bo sinh vt eucaryote u c cha intron. C mt s gen khng
c intron, v d nh phn ln cc gen m ha cho cc protein histon. Trong trng hp ny, qu trnh phin m
to mRNA s khng c giai on splicing.

Phin m tng hp tRNA

Trong b gen ca t bo procaryote cng nh eucaryote c cc gen m ha cho tRNA. vi khun, thng
thng mt gen c th m ha cho mt s tRNA khc nhau. V d E. Coli, mt gen c th m ha cho 7 loi
tRNA khc nhau. Tuy vy, i lc ngi ta cng tm thy c nhng gen ch m ha cho mt tRNA.
Qu trnh phin m tng hp tRNA vi khun do enzyme RNA- polymerase xc tc, cn nm men v cc sinh
vt a bo do enzyme RNA- polymerase III xc tc. S phin m tng hp tRNA sinh vt ni chung thng
din ra theo 2 giai on
- Phin m tng hp phn t tin tRNA chung,
- Ct b cc intron to phn t tRNA hon thin.
Trong trng hp mt gen m ha cho nhiu tRNA th bn phin m u tin c on dn u u 5, on
cui u 3 v cc on m nm gia cc tRNA cn c loi b to cc phn t tRNA hon thin. Phn
t tin tRNA s c mt endonuclease l ribonuclease P (RNase P) ct ti cc v tr xc nh u 5, to u
5phosphat v mt exonuclease khc l ribonuclease D ct b on khng mang m u 3, to u 3OH.

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Tuy nhin, u 3 ca tRNA lun c b ba CCA cui cng, l ni axit amin gn vo. Nu ribonuclease D ct sai
v tr u 3 th enzyme c hiu CCA- nucleotidyltranferase s sa sai u 3 lun c b ba CCA. Mi loi
tRNA c hiu cho mt axit amin, do vy, c t nht l 20 loi tRNA.
t bo eucaryote, s lng gen m ha cho tRNA nhiu hn t bo vi khun. t bo nm men S.
Cerevisiae, ngi ta xc nh c 46 loi tRNA khc nhau, nhng c n khong 360 gen m ha cho cc
tRNA ny. Nh vy, tnh trung bnh, mi tRNA c khong 8 gen m ha ring bit. Tuy nhin, con s ny khng
th chia u nh vy m thng thng, nhng loi axit amin no c mt trong protein vi s lng ln th s
lng gen m ha cho tRNA vn chuyn axit amin s nhiu hn. Cc gen m ha cho mt loi tRNA thng
phn b trn cc nhim sc th khc nhau. Do , cc promoter ca cc gen ng vai tr quan trng trong s
kim sot qu trnh phin m.

Phin m tng hp rRNA

Cc phn t rRNA cng c tng hp t cc gen m ha cho chng trn DNA. Trong t bo procaryote c 3
loi rRNA tham gia vo qu trnh tng hp protein l 16S, 23S v 5S; cn t bo eucaryote, cc rRNA c mt
trong ribosome l 18S, 28S v 5,8S.
Qu trnh phin m tng hp rRNA cng xy ra theo hai bc nh tRNA. Phn t tin rRNA c tng hp u
tin da trn khun DNA. t bo eucaryote, qu trnh c xc tc nh enzyme RNA-polymerase I. Phn t
tin rRNA tri qua giai on ct loi intron to cc phn t rRNA hon thin.

Nhiu rRNA c m t trong cng m t phn t tin RNA E. Coli

vi khun E. Coli, mt gen thng m ha cho mt s phn t rRNA. V d 3 loi rRNA l 16S, 23S v 5S c
m ha trong mt gen. Khi phin m s to mt phn t tin rRNA chung cho c 3 loi rRNA trn, sau , cc
enzyme ribonuclease s thc hin qu trnh ct loi cc intron to 3 phn t rRNA hon thin. Cng c
trng hp, trong mt operon m ha cho rRNA c c phn m ha cho c tRNA .

Thch

Chia s

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Vit bnh lun

Bnh lun bng...

Quan Nguyen i Hc Y H Ni
Thanks.
Tr li Thch 8 Thng 8 2014 lc 2:34
Plugin x hi ca Facebook

(/user/reuse/m/a72951e7/1)

T I V

T I S DNG

Bi Ch Bu (/profile/229)
0 GIO TRNH (/PROFILE/229?TYPES=2)

18 TI LIU (/PROFILE/229?TYPES=1)

(/profile/229)
NH GI:
0 da trn 0 nh gi

Cng tc gi
S sao m (/m/su-sao-ma/70596231)
Lc s pht trin ca di truyn hc v k thut di truyn (/m/luoc-su-phat-trien-cua-di-truyen-hoc-va-ky-thuat-ditruyen/20e2f7d7)
Phng php gii trnh t ca axit nucleic (/m/phuong-phap-giai-trinh-tu-cua-axit-nucleic/fe6f8717)
C c phng php lai phn t (/m/cac-phuong-phap-lai-phan-tu/7b1d9e0f)
Dch m(Translation) (/m/dich-matranslation/2e55cf1e)
Bn cht ca vt cht di truyn (/m/ban-chat-cua-vat-chat-di-truyen/1f802bed)
C c Enzyme s dng trong cng ngh di truyn (/m/cac-enzyme-su-dung-trong-cong-nghe-di-truyen/0ee91ddc)
Phng php nhn bn (PC R) (/m/phuong-phap-nhan-ban-pcr/02bcb4b4)
Vt cht di truyn ca mt s nhm sinh vt (/m/vat-chat-di-truyen-cua-mot-so-nhom-sinh-vat/5003ab76)
Vector chuyn gen (/m/vector-chuyen-gen/7d602ed9)
TRC
TIP

Ni dung tng t
Qu trnh nhn ln (/m/qua-trinh-nhan-len/a31cc96e)
Qu trnh nhn ln ca virus (/m/qua-trinh-nhan-len-cua-virus/8f5aa0bf)
Gen l n v chc nng ca b my di truyn (/m/gen-la-don-vi-chuc-nang-cua-bo-may-di-truyen/1a0af523)

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C c Enzyme s dng trong cng ngh di truyn (/m/cac-enzyme-su-dung-trong-cong-nghe-di-truyen/0ee91ddc)

C c phng php tch chit axit nucleic (/m/cac-phuong-phap-tach-chiet-axit-nucleic/ffa54dfb)
S sao m (/m/su-sao-ma/70596231)
Phng php nhn bn (PC R) (/m/phuong-phap-nhan-ban-pcr/02bcb4b4)
Qa trnh nhn i DNA (/m/qua-trinh-nhan-doi-dna/ff99e894)
Polymerase C hain Reaction(PC R ) (/m/polymerase-chain-reactionpcr/07a59598)
Qu trnh nhn ln (/m/qua-trinh-nhan-len/77358052)
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