Chng 2

C IM CU TRC V HOT NG CA GEN

- S tit l thuyt: 2,0

- Mc tiu ca chng: (1) Phn bit c c im ca gen v cc vng iu khin
gen Prokaryot, Eukaryot. Phng php pht hin intron v exon; (2) Trnh by c
c im ca s ti bn v phin m ca gen t bo Prokaryot, Eukaryot; (3) c im
ca dch m v iu ho biu hin gen; (4) Mt s khi nim: Tnh tri, ln, c s phn
t ca tnh tri ln. Lai phn t, Southern blot, Northern blot, Western blot.
- Ni dung ging dy trn lp: (i) c im cu trc ca gen; (ii) C ch ti bn v sa
cha ADN; (iii) Phin m v kim sot hot ng phin m ca gen; (iv) Tnh tri, tnh
ln v c ch phn t ca s hnh thnh tnh tri ln.
- Ni dung sinh vin t nghin cu: Dch m v kim sot qu trnh dch m.
- Trang thit b dy hc: Bi ging, Sch gio trnh, sch tham kho.
- Phng php dy hc: Hng dn trn lp, t nghin cu liu.
- Ti liu hc tp:
1. David Clark (2005). Molecular Biology. Elsevier Inc.
2. Chu Hong Mu (2005). C s v phng php Sinh hc phn t. Nxb i hc
S phm.
3. Bi ging.

I. HNG DN SINH VIN HOC TRN LP

1. c im ca gen t bo Prokaryot v Eukaryot
1.1. c im ca gen t bo Prokaryot
- H gen ca t bo prokaryot gm cc gen thuc DNA vng nhn v DNA plasmid
- Phn t DNA dng vng, xon kp v khng lin kt vi histon. DNA plasmid c kh
nng nhn i c lp trong t bo ch.
- Gen c cu trc lin tc, bao gm nhng on m ha axit amin.
- Cc gen trn NST l a cistron. Mi cistron xc nh 1 chui polypeptit. a cistron
phin m tng hp phn t mRNA v sn phm dch m l nhiu polypeptit. Cc cistron
c ngn cch bi on m (spacer).

1.2. c im cu trc phn on gen sinh vt Eukaryot

- Gen Eukaryot c cu trc khng lin tc, gm cc on exon v intron xen k nhau.
-

Exon l trnh t nucleotit m ho thng tin di truyn. intron l trnh t nucleotit

khng m ho thng tin di truyn

Khi cc gen phn on phin m s to pre- mRNA c chiu di ln hn cc phn t

mRNA trng thnh. cc pre- mRNA phi tri qua giai on splicing, gn chp 7 mtG
v ui poliA to cc phn t mRNA trng thnh.

2. Phng php pht hin cc on intron v exon

Phng php pht hin cc on intron v exon gm 2 phng php
(1) So snh trnh t cDNA v trnh t gen
Bc 1. Gii trnh t cDNA
+ Tch mRNA trong t bo
+ S dng enzyme phin m ngc thc hin phn ng RT-PCR tng hp on
cDNA hai si t si n mRNA
+ To dng DNA ti t hp c cha on cDNA, nhn dng v tch dng gen
+ Gii trnh t on cDNA
Bc 2. Gii trnh t gen
+ Tch chit DNA h gen
+ Phn lp gen bng k thut PCR hay RE
+ To dng DNA ti t hp, nhn dng v tch dng gen
+ Gii trnh t gen
Bc 3. so snh trnh t DNA h gen vi trnh t cDNA xc nh on intron v exon.
(2) Lai phn t
+ Nguyn tc: dng cc phn t mRNA trng thnh c nh du phng x em
lai vi phn t DNA h gen, soi trn knh hin vi in t pht hin c on exon (c
s bt cp b sung mRNA- DNA) v on intron (khng c s bt cp b sung mRNA DNA).
+ Cc bc:
Tch chit DNA h gen v mRNA t bo cht
Phn lp gen
2

Lai phn t gia mRNA- DNA xc nh on exon v intron.

3. Cu trc ca promoter trong h gen ca Prokaryot. ng dng ca vic nghin
cu promoter trong k thut sinh hc phn t
3.1. Cu trc ca promoter ca sinh vt Prokaryot (E.coli)
-

Cu trc ca promoter

-----TTGACA-------------TATAAT--------- ----------- mRNA

-35 bp

-10 bp

+1

+Base u tin c phin m ra mRNA c nh s +1. V tr pha trc +1

gm 6bp xung quanh -10bp l hp TA (pribnow box) (cc nucleotit khng c phin
m). y l v tr c i lc vi RNA polymerase.
+ Xung quanh -35bp c trnh t nht tr. Trnh t nht tr lm tng tc qu
trnh phin m.
-

Chc nng ca promoter:

+ L vng cha tn hiu nhn bit ca enzym phin m v c i lc vi enzym

phin m, nh hng s di chuyn ca enzym phin m
+ iu khin tc phin m ca gen nh trnh t nht tr v v tr CAP nm pha
trc hp TA
ng dng nghin cu promoter
- Thit k vector biu hin gen: S dng cu trc promoter E. coli (promoter lac.,
promoter tryp.) thit k vector biu hin. l nhng promoter mnh v iu khin
c.
- Cc promoter c kh nng biu hin gen cao c nhng c im chung sau: (1) promoter
mnh c kh nng to protein ngoi lai ln hn 10 n 30% protein tng s ca t bo; (2)
Promoter ch hot ng phin m gen mc c s (phin m khi khng cm ng) vi
mc nh ti a: (3) Promoter c cm ng theo cch n gin nhng hiu qu cao.
- Tm hiu c s phn t ca cc c tnh, tnh trng c lin quan n trnh t promoter.
3.2. Cu trc v chc nng ca promoter, operator sinh vt Eukaryot
- Promoter ca sinh vt Eukaryot
------GGGCGGG-------CCAAT--------TATA----- ------mRNA
-110 bp

-40 bp

-30 bp

+1

+ Hp TA pha trc im khi u phin m khong 30bp (ng vt c v),

60- 120bp (nm men) c vai tr lm tng tc phin m do c i lc vi loi protein
hot ha chuyn ha (v tr CAP).
+ Trc hp TA c hai trnh t CCAAT (-40bp) v trnh t giu GGGCGGG (110bp)
- Chc nng ca promoter:
+ L vng cha tn hiu nhn bit ca enzym phin m v c i lc vi enzym
phin m, nh hng s di chuyn ca enzym phin m
+ iu khin tc phin m ca gen nh trnh t nht tr v v tr CAP nm pha
trc hp TA.
c im v cu trc v chc nng ca operator
+ Cu trc: l trnh t nucleotit nm trc im khi u phin m v sau trnh t
promoter
+ Chc nng: l vng DNA c i lc vi protein c ch, khi protein c ch bm
vo s ngn cn s di chuyn ca enzym phin m
4. Ti bn DNA theo c ch na gin on ca Okazaki E.coli
-

H enzym tham gia ti bn DNA E.coli

+ Enzyme DNA polymerase: DNA polymerase I (c v sa cha), DNA
polymerase II (xc nh s bt u v kt thc tng hp), DNA polymerase II (xc
tc tng hp chui polynucleotit mi)
+ DNA gyrase: ct v lm tho xon 2 pha ngc nhau bt u t im khi u
sao chp.
+ DNA helicase: b gy lin kt hidro, gii phng cc chui n to chc ti bn.
+ DNA ligase: ni cc on okazaki to thnh si DNA di lin tc
+ RNA polymerase: tng hp on mi to u 3OH t do.

Cc yu t khc tham gia: DNA, cc nucleotit t do, protein B (nhn ra im khi

u ti bn), protein SSB (gi cho cc si n tch ring khi cha ti bn)

C ch ti bn DNA theo Okazaki l c ch ti bn na gin on gm cc khu

sau:
+ Hin tng dui xon:
+ Khi u ti bn bng RNA mi (primer):
+ Ko di, loi b mi v hnh thnh cc phn on Okazaki:
4

+ Cc phn on Okazaki c ni li nh enzyme ni ligase to thnh mch

DNA mi:
5. c im ca s phin m ca gen t bo Prokaryot, Eukaryot
5.1. c im ca s phin m ca gen t bo Prokaryot
-

Gen a cistron phin m to 1 phn t mRNA

Cc giai on phin m:
Li RNA-polymerase cng nhn t nhn bit v bm vo trnh t khi ng trn
promoter (hp TA). Nhn t tch khi li enzyme, RNA polymerase trt dc gen
theo chiu 3- 5, xc tc bin tnh cc b 2 si DNA l ra si khun tng hp RNA.
Qu trnh tng hp RNA bt u, s lp rp cc ribonucleotit t do thnh chui
polyribonucleotit theo nguyn tc b sung A- U, G- C v si RNA c sinh ra c
chiu 5- 3.
Kt thc: on kt thc ra tn hiu cho RNA-polymerase dng phin m. on
kt thc gm 2 phn: phn c t l GC cao v mt loi base T trn si antitemplate.

- Sn phm phin m l si n RNA c chiu 5- 3, cc mRNA c tng hp xong

mang thng tin di truyn lin tc v tham gia ngay vo qu trnh dch m.
5.2. c im ca s phin m sinh vt Eukaryot
-

Enzym phin m gm:

+ RNA-polymerase I, enzyme khng b c ch bi - amanitin (cht c ch tng

hp RNA), chuyn trch vic phin m cc gen tng hp rRNA (28S, 58S v 18S).
+ RNA-polymerase II: enzyme b c ch bi - amanitin, n cn cho tng hp
mRNA
+ RNA-polymerase III: enzyme b c ch bi - amanitin nng cao, phin
m tng hp cc tRNA 4S, rRNA 5S.
-

Cc vng kim sot phin m

+ Hp TATA nm cch im khi u phin m 25- 30 bp v pha trc, c chc

nng gip RNA-polymerase nhn ra v bm ng im khi u phin m.
+ on kt thc nm u 5 ca gen
+ Operator l trnh t nucleotit ngn c i lc vi protein c ch qu trnh phin
m.
-

Cc giai on phin m:
5

+ Giai on khi ng: chu s kim sot ca hp TA. RNA-polymerase hot ng

phin m nh nhiu nhn t TF c bn cht l protein.
+ Giai on ko di: phn t RNA c tng hp t mch khun DNA nh nhn t
TFIIS.
+ Giai on kt thc: Kt thc phin m c lin quan n cu trc kp tc tip sau
trnh t giu GC.
-

Qu trnh hon thin cc pre-mRNA to mRNA trng thnh

+ Gn thm chp 7-methylguanosine u 5P ca mRNA
+ Gn thm cn ui polyadenine di 100- 200 A cui phn t mRNA

+ Ct b intron, ni cc exon li vi nhau nh phc SnRNP ca nhn to phn

t RNA trng thnh.
7. Lai phn t
7.1. Khi nim v lai phn t
nh ngha
Cc kiu lai phn t : Southern, Northern, Western
7.2. Phng php Southern blot
- S dng mt trnh t DNA bit trc lai vi on DNA cn tm.
- Cc bc tin hnh c th thc hin theo cc bc sau:
+ Ct DNA bi enzyme gii hn thnh cc on DNA nh c th dch chuyn trn
gel agasore.
+ Bin tnh cc si DNA kp trn gel agarose bng phng php x l vi kim.
+ Chuyn DNA bin tnh sang mng lai.
+ Lai vi trnh t DNA c nh du bng phng x hoc hunh quang.
+ Pht hin phn t lai bng k thut phng x t ghi hoc knh hin vi in t.
- ng dng :
+ Phng php Southern blot cho php xc nh c trnh t on DNA cn tm,
+ C th pht hin c t bin mt on hoc thm on khi so snh c th t
bin vi c th bnh thng.

7.3. Khi nim, quy trnh v ng dng ca phng php Northern blot
- Phng php Northern blot l k thut lai gia mRNA vi trnh t mu d c nh du
phng x.
- Cc bc
+ Tch chit RNA tng s v bin tnh RNA khng tr v cu trc bc 2 ca
n.
+ in di RNA trn gel agarose.
+ RNA c chuyn sang mng v lai vi trnh t DNA c hiu (mu d c
hiu)
+ V tr v s lng cc on ca RNA lai vi DNA c pht hin trn phim
bng phng phng x t ghi.
- Cc ng dng
+ Xc nh kch thc v hm lng ca 1 mRNA c trng trong 1 hn hp
RNA.
+ Xc nh v tr chnh xc v s phn b ca mt mRNA trong mt nhm t bo
chuyn bit ca t chc, t xc nh vai tr ca mRNA trong t chc .
+ Nghin cu cu trc RNA, pht hin t bin.
7.4. Phng php Western blot
- K thut ny s dng tnh gn bm c hiu ca khng th vi khng nguyn tng
ng; v s xc tc c hiu ca enzyme (c gn vo khng th th hai)
- Bc 1: Tch protein bng in di trn gel polyacrylamide
- Bc 2: Chuyn protein t gel sang mng lai nh in trng
- Bc 3: Nhum mng bng khng th c hiu
- Bc 4: Nhum mng bng cng hp (khng th khng khng th gn enzyme)
- Bc 5: B sung c cht cho enzyme chuyn ha c th nhn thy vt protein phn
ng vi khng th c hiu trn mng.
- Enzyme s chuyn ha c cht thnh hp cht c mu c th nhn thy trn mng.
- Nh vy ch vt protein no phn ng vi khng th th mi hin th trn mng.
II. HNG DN THO LUN NHO TRN LP
1. im ging v khc nhau gia phin m ca sinh vt Prokaryot v Eukaryot
1. Nhng im ging nhau
7

Cc yu t tham gia: ribonucleotit t do, RNA polymerase, ATP, promoter, operator,

si khun DNA c chiu 3- 5...

Vng DNA cha gen c phin m phi c s m xon cc b lm l ra si khun

(si c ngha - strand sense).

S khi u v kt thc phin m ph thuc vo tn hiu khi u v kt thc, l

nhng trnh t nucleotit c th nm trc v sau gen.

RNA c tng hp theo chiu 5 3, enzym RNA-polymerase di chuyn theo chiu

3 5 trn si strand sense.

Sn phm l 1 chui poliribonucleotit c chiu 5- 3 l bn sao ca gen.

2. Nhng im khc nhau

Phin m ca sinh vt Prokaryot

Phin m ca sinh vt Eukaryot

- Xy ra trong t bo cht ng thi - Xy ra trong nhn t bo, trc qu trnh dch

vi qu trnh dch m
m
- C 1 loi enzyme RNA polymerase - C 3 loi enzyme RNA polymerase ph trch
ph trch tng hp 3 loi RNA khc tng hp 3 loi RNA khc nhau
nhau
- Si RNA c hnh thnh l pre- mRNA, cc
- Si n RNA c tng hp tham phn t ny tri qua bc hon thin mi tr
gia ngay vo qu trnh dch m
thnh phn t RNA trng thnh tham gia dch
m

2. Khi nim tnh tri, tnh ln. C s phn t ca tnh tri v tnh ln
- Khi nim tnh trng tri: l tnh trng c biu hin con lai F1 khi cho lai gia hai
c th b m thun chng khc nhau v cp tnh trng tng phn
- Khi nim tnh trng ln: l tnh trng khng c biu hin con lai F1 khi cho lai
gia hai c th b m thun chng khc nhau v cp tnh trng tng phn v c tnh
trng ging 1 bn (b hoc m)
- Khi nim tnh trng tri hon ton: l nhng tnh trng do gen tri hon ton quy nh.
- Khi nim tnh trng tri khng hon ton: l tnh trng trung gian c biu hin
con lai F1 khi cho lai gia hai c th b m thun chng khc nhau v cp tnh trng
tng phn.
8

- Tnh trng trung gian do gen tri khng hon ton quy nh, vai tr ca gen tri v gen
ln l ngang nhau trong vic hnh thnh tnh trng.
- C s phn t ca tnh trng tri, ln c gii thch:
- S tc ng qua li gia cc sn phm ca cc gen trong s hnh thnh tnh trng gm:
Tng tc gia sn phm ca cc gen alen v tng tc gia cc sn phm ca cc gen
khng alen. Bn cht ca hin tng tng tc gen l: S tng tc ca cc sn phm
biu hin gen
- S hnh thnh tnh trng l do hot ng chuyn ho cc cht trong chui phn ng ho
sinh, vi s tham gia ca cc enzyme.
- S khc nhau v hot ng chuyn ha c cht ca enzym do gen tri v do gen ln
tng hp.
- S khc nhau v hm lng enzyme, protein do gen tri v do gen ln tng hp.
3. Ti bn DNA trong t bo sng cn mi (primers), primer l trnh t RNA. im
khc ca primer trong ti bn in vivo v primer trong phn ng PCR. Hin tng cc
phn on okazaki ni vi nhau trong qu trnh ti bn DNA
- Mi l mt trnh t RNA ngn, c nhm OH 3 (3-OH).
- Qu trnh tng hp mch DNA mi thc hin theo chiu t 5 n 3. Mi c tng
hp to 3-OH l c s hnh thnh lin kt phosphodiester ko di chui
polynucleoptit.
- Mi l trnh t RNA m khng phi l DNA l v lin quan n s khc nhau v chc
nng ca enzyme RNA-polymerase v DNA-polymerase.
+ RNA-polymerase c kh nng xc tc gn kt cc ribonucleotit t do to thnh
chui polyribonucleotit m khng cn primer c u 3-OH.
+ DNA-polymerase ch xc tc tng hp chui polynucleotit khi c mt ca
primer. Do vy trong ti bn DNA cn c RNA-polymerase xc tc tng hp 1 on
primer ngn, v primer l trnh t RNA.
- Primer trong ti bn l trnh t RNA ngn do RNA-polymerase xc tc tng hp, cn
trong phn ng PCR l trnh t DNA ngn do con ngi thit k da trn trnh t gen
bit.
Hin tng cc phn on okazaki ni vi nhau trong qu trnh ti bn DNA
- Cc phn on okazaki ni vi nhau theo c ch:
+ Trnh t mi c ct ta tng ribonucleotit ca mch b sung nh nuclease, ct
n u tng hp nucleotit b sung vi mch gc n
9

+ Cui cng nh enzym ni DNA ligase ni vi nucleotit ca phn on okazaki

trc n.
III. HNG DN T HOC
Sinh vin t nghin cu theo cc ni dung sau:
1. Dch m
- Cc yu t tham gia: cc loi RNA, ribosome, aa t do, ATP, aminoacyl synthetase, IP...
- Cc giai on ca qu trnh dch m.
1) Khi ng
- Giai on c s tham gia ca nhiu nhn t protein nhn t khi ng (IF Initiation
Factor); Prokaryot c 3 nhn t IF, cn Eukaryot c 6 nhn t IF.
- Cc bc ca giai on khi ng:
+ Aminoacyl tRNA synthetase chuyn bit (methionyl tRNA synthetase)
gn 1 phn t met vo mt u ca tRNAmet to thnh Met tRNAimet .
+ Hnh thnh tiu n v nh ribosome met tRNAi met mRNA vi s tham gia
ca nhn t khi ng (IF2 Prokaryot ; IF4 Eukaryot).
2) Ko di
- Khi a.a met t vo v tr P, chui polypeptit bt u c tng hp. Aminoacyl tRNA
k tip n n xp vo ng v tr A trn ribosome nh mt nhn t ko di (EF). Lin
kt peptit gia Met v aa th nht hnh thnh.
- Qu trnh c lp li cho n khi xut hin du hiu kt thc dch m. Mt phn t RNA
c th c nhiu ribosome cng trt gi l chui polixom
3) Kt thc
Khi nhn t (TF) kt thc nhn bit c du hiu kt thc (cc codon kt thc: UAG,
UAA, UGA), phc hp peptidyl tRNA lp tc tch thnh tRNA t do v chui
polypepetit. Ribosome tch ra thnh 2 tiu n v v mRNA c gii phng. aa m u
b loi khi chui.
- im khc bit trong phin m Prokaryot v Eukaryot:
+ Prokaryot : qu trnh phin m v dch m din ra ng thi trong t bo cht
+ Eukaryot : qu trnh phin m v dch m din ra khng ng thi, phin m
xy ra trc trong nhn cn dch m xy ra sau trong t bo cht

10

2. Mc iu ha biu hin gen

1. Mc cht nhim sc
- c thc hin bi hot ng ca mt s enzym nh DNAaza I (lm tho xon NST
cc gen biu hin) hoc DNA Z (dng cu trc siu xon lin quan n ng m gen).
- S methyl ho cc base. V d: s methyl ho C v tr s 5. S methyl ho lm gen
ngng hot ng. V d: S bt hot ca NST X ngi do siu methyl ho.
- Do s thay i cu hnh NST.
2. Mc phin m
- iu ho mc phin m nh hng trc tip n ng m gen, kiu ny thng gp
iu ho trao i cht hoc s bit ho t bo.
- S tc ng ca trnh t cis (enhancer) lm tng s sao m, cc nhn t trans, chn la
promoter thch hp, attenuatinon (s gim bt tnh tit)
3. Mc sau phin m
Khi hnh thnh mRNA trng thnh c nhiu yu t nh hng v c th kim tra
gin tip biu hin ca gen tng ng: Splicing khc nhau, im polyadenin ho khc
nhau, t bin trn mRNA, bn chu k phn hu ca mRNA, s bo tn cc RNA trong
t bo.
4. Mc dch m
S bin i ca nhn t khi u IF tc ng n biu hin gen.
5. Mc sau dch m
Sau khi chui polypeptit c tng hp, cc protein c th b bin i th cp
trc khi biu hin hot tnh hoc c th chu nhng bin i lp th nh s kt hp cc
enzym vi mt s sn phm c bit c th lm thay i cu trc khng gian ca chng
dn n mt hot tnh: glycosylation, phosphorylation, gn thm cc tn hiu petit
3. iu ha biu hin gen Prokaryot
- Cu trc ca operon
+ S iu ho biu hin gen Prokaryot phn ln xy ra giai on phin m. C
ch iu ho thng qua operon. Mi operon bao gm: nhm gen a cistron cnh nhau v cc
trnh t DNA khc tham gia iu ho.
+ Operon gm promoter, operator, k tip l nhm a cistron. Prokaryot thng
to ra mRNA a gen (polycistronic), nhng mRNA ca Eukaryot ch c 1 gen.
- Cu trc ca promoter
11

+ Thc cht ca khi u phin m l quan h tng tc gia RNA -polymerase

vi promoter.
+ Phn ln promoter E. coli v cn bn c cng mt cu trc: hp TA v trnh t
nht tr. Hai trnh t phi hp vi nhau cho php RNA- polymerase gn vo v qu trnh
phin m v dch m bt u.
-----TTGACA-----------------TATAAT---------- --------- mRNA
-35 bp

-10 bp

+1

- Operator: l trnh t nucleotide ngn nm trc gen cu trc c i lc vi protein c

ch, ngn cn qu trnh phin m
- Protein iu ho (TF) tng tc vi trnh t DNA c hiu kim sot phin m ca gen
cu trc. Gen iu khin tng hp protein iu ho gi l gen iu ho. TF c th bm
vo promoter hoc operator lm tng cng hoc c ch phin m.
-

Cc hnh thc kim sot phin m

+ Kim sot m: S gn ca repressor protein vo operator lm cn tr s phin m

ca tt c cc gen trong cng mt operon.
+ Kim sot dng: Cc protein hot ha biu hin gen (activator) gn vi cc im
khi ng ca operon hay gn vi enhancer site s kch thch s phin m ca cc gen
cu trc.
+ Cht gip cho gen phin m c gi l cht cm ng, cht c tc dng ngc li
gi l cht km hm. Cc gen cm ng thng thng tham gia vo cc phn ng thoi
dng (kim sot m, cm ng); cn cc gen c ch thng tham gia vo cc phn ng
bin dng.
+ Kh nng c th xy ra 4 kiu t hp kim sot s phin m: m, cm ng; m,
c ch; dng, cm ng; dng, c ch
IV. CU HOI V BI TP
1. Cc khi nim: exon, intron, on m, yu t di truyn vn ng, on xen v gen nhy.
Nu c ch xen ca gen nhy.
2. c im s phin m ca gen a cistron t bo Prokaryot. Ti sao trong qu trnh phin
m li khng cn c s tham gia ca mi?
3. c im ca s phin m t bo Eukaryot.
4. c im ca s ti bn DNA theo c ch na gin on ca Okazaki E.coli. Ti sao ti
bn DNA trong t bo sng li cn c mi (primers), mi li l trnh t RNA m khng phi
trnh t DNA? C ch ni cc phn on Okazaki thnh mt chui polynucleotide.

12

5.

Phn bit khi nim tnh tri, tnh ln v tnh trng trung gian. C s phn t ca tnh tri,
tnh ln v tnh trng trung gian.

6.

c im v cu trc v chc nng ca promoter, operator sinh vt Prokaryot v

Eukaryot. ng dng promoter trong k thut sinh hc phn t.

7.

Hin tng bin tnh, hi tnh phn t DNA l g ? Khi nim v nhit chy DNA
(Tm). ng dng ca hai hin tng ny trong sinh hc phn t.

8.

Khi nim lai phn t v nu cc k thut lai phn t v ng dng.

9.

Nguyn tc, k thut v ng dng ca lai Southern blot.

10. Nguyn tc, k thut v ng dng ca lai Northern blot.

11. Nguyn tc, k thut v ng dng ca lai Western blot.

13