I. C IM PHIN M EUKARYOTE Qu trnh phin m eukaryote cng tng t nh qu trnh phin m prokaryote.

te. - u s dng mt mch DNA c chiu 3 5 lm khun tng hp RNA. - Nguyn liu dng trng hp RNA l cc ribonucleotid triphosphat, gm: ATP, UTP, GTP, CTP. - ARN polymerase tham gia xc tc phn ng trng hp RNA. - Qu trnh phin m tri qua 3 giai on: khi s, ko di v kt thc. - Cc nhn t phin m tham gia gip cho ARN polymerase tng hp chnh xc hn. - RNA c tng hp theo chiu 5 3 v theo nguyn tc b sung. Tuy nhin, qu trnh phin m eukaryote phc tp hn qu trnh phin m prokaryote cc c im sau: - prokaryote ch c 1 loi enzym RNA pol tham gia phin m tng hp cc loi RNA. Nhng eukaryote c ba loi enzym ARN polymerase tham gia phin m tng hp cc loi phn t ARN khc nhau gm enzym ARN polymerase I, enzym ARN polymerase II v enzym ARN polymerase III. - pokaryote c yu t phin m l yu t sigma, eukaryote c nhiu yu t phin m v c chia thnh yu t phin m chung v yu t phin m c hiu. Cc yu t phin m chung cn cho vic phin m tt c cc gene v thng c k hiu l TFI, TFII, TFIII ng trc cc ch ci n. S I, II, III ch s tng ng vi cc loi RNA pol. Cc yu t phin m c hiu cn cho vic phin m cc gene nht nh trong hon cnh nht nh. Ngoi ra s ko di v kt thc phin m cng c cc yu t phin m tham gia. - Do cc gen eukaryote phn mnh nn mRNA va c phin m t DNA khng tham gia dch m ngay hnh thnh protein nh prokaryote. Cc tin mRNA c hnh thnh trong nhn phi chu mt s bin i v mt ha hc trc khi xut hin trong t bo cht di dng hot ng. Tng t, cc rRNA cng c hnh thnh trong nhn di dng mt phn t tin thn 45S. Phn t tin thn 45S sau s chu nhiu bin i ha hc, c phn ct thnh 3 RNA hot ng (18S, 28S, v 5,8S), ri mi c chuyn ra t bo cht.

- Do eukaryote l t bo c nhn nn qu trnh phin m to mRNA v qu trnh dch m hnh thnh protein xy ra khng ng thi nh prokaryote. II. ENZYM RNA POLYMERASE, CC YU T PHIN M V QU TRNH PHIN M EUKARYOTE 1. Enzym RNA polymerse Thnh phn cu trc enzym ARN polymerase ARN pol I RPA1 RPA2 RPC5 RPC9 RPB6 [+ 9 chui khc] ARN pol II RPB1 RPB2 RPB3 RPB11 RPB6 [+ 7 chui khc] ARN pol III RPC1 RPC2 RPC5 RPC9 RPB6 [+ 11 chui khc]

Bng 1. Thnh phn cc chui polypeptide ca cc ARN polymerase (ARN pol)

Hnh 1. M hnh cu trc cc loi enzyme ARN polymerase Hnh 1. M hnh cu trc cc loi enzyme ARN polymerase

Trong mi enzym ARN polymerase c vai tr v v tr khc nhau trong nhn t bo (Bng 2) 3 loi RNA polimerase RNA polimerase I RNA polimerase II V tr Hch nhn Dch nhn Dch nhn Sn phm rRNA18S, 28S, 5,8S - mRNA - snRNA U1, U2, U3, U4, U5 RNA polimerase III - tRNA, RNA 5S - snRNA U6, RNA 7S Bng 2. Ba loi enzyme RNA polymerase, v tr v sn phm 2. Cc nhn t phin m ca RNA pol II v qu trnh phin m cc mRNA do RNA pol II xc tc a. Cu trc gen m ha protein eukaryote. Gen m ha cho protein eukaryote bao gm 3vng: vng 5 kim sot biu hin gen

Hnh 2. Cu trc ca gen m ha cho protein eukaryote Vng 5 kim sot biu hin gen. Vng ny bao gm cc trnh t nucleotide iu ha biu hin gen v hot ha s phin m, bao gm: promoter v trnh t tng cng(enhancer) + Promoter: l nhng trnh t nh v u 5 khng dch m ca gen c chc nng xc nh v tr bt u phin m, kim sot s lng mRNA. Promoter ca phn ln gen m ha protein thng gm khong 200 bp (i khi di hn) nm ngc dng k t v tr bt u phin m (+1) v mang mt s trnh t (cn gi l yu t trnh t). Promoter c chia lm 2 vng chnh: vng li promoter v vng bin promoter. - Vng li promoter gm cc yu t trnh t tc ng gn, iu khin vic bt u phin m ng v tr ca RNA pol. Vng ny thng c kch thc khong 50 bp nm ngc dng v st im bt u phin m. Vng li promoter in hnh gm c 2 trnh t: trnh t ngn k hiu l Inr, nm ngc dng vng m ha ca gen v m rng ti v

tr bt u phin m; hp TATA thng nm v tr -30, c mt trnh t lin ng in hnh gm 7 nucleotid l [5-TATAAAA-3]. Cc yu t trnh t Inr v TATA c vai tr gip RNA nhn ra promoter v bt u phin m chnh xc (ti ng v tr +1). - Cc yu t vng bin promoter thng nm ngc dng hp TATA, trong khong t v tr -50 n -200 (k t v tr +1). Trong nhm ny c th c hp CAAT c trnh t lin ng [CAAT] thng v tr -75, v hp GC c trnh t lin ng [GGGCGG] thng v tr -90. C 2 hp CAAT v GC u c th biu hin chc nng theo c 2 chiu (ngha l d n cng chiu phin m hay ngc li nhng RNA pol u nhn ra). Ngoi ra cn c vng AP1 v Octamer l nhng trnh t trc gen (thuc vng bin) + Trnh t tng cng phin m (enhancer) cn thit gen c biu hin mc ti a. Cc enhancer c th nm ngc dng, xui dng thm ch nm trong trnh t m ha ca gen, nhng v tr ph bin l nm ngc dng gen. Cc enhancer thng mang mt chui cc on DNA ngn, trong c mt s ging vi cc yu t trnh t ca promoter. Trong qu trnh phin m, cc yu t phin m chung cho RNA pol II (TFII factors) bm vo vng promoter. Vi yu t phin m c hiu cng bm vo vng promoter, s khc bm vo vng tng cng.

Vng c phin m
Bao gm cc exon v intron nm xen k. y l mt c im phn bit vi gen ca prokaryote. Cc exon v intron u c phin m nhng ch c cc exon l c dch m. Cc intron c bt u bng GT v kt thc bng AG. Cc intron chim phn ln trong mi gen v chng s c loi b khi RNA mi c tng hp, cn cc exon c ni vi nhau to nn mRNA hon chnh (mature mRNA).

 hai u ca vng c phin m (coding region) cn c vng 5 khng dch m (5 untranslation region) v vng 3 khng dch m (3 untranslation region). Vng 5 khng dch m c tnh t v tr bt u phin m cho n codon khi u ATG. Vng 3 khng dch m bt u t codon kt thc n v tr gn ui poly(A).

Vng 3 khng dch m

Chc nng cha r, mt s gen vng ny mang cc trnh t iu ha chuyn bit. b. Cc nhn t phin m ca RNA pol II Cc nhn t khi u - TFIID (TBP v cc TFA) yu t v tr nhn bit v gn vo trnh t TATA. - TFIIA nhn t gn kt v n nh TFIID trn DNA. - TFIIB nhn t gn kt v n nh TFIID trn DNA. - TFIIF yu t lin kt vi enzym ARN polymerase II v hot ng helicase v do c vai tr vo s dui xon ca DNA cc Promoter l ra si mu. - TFIIH c mt s hot ng, bao gm mt ATPase, helicase, v kinase hot ng m c th phosphorylate v kch hot PoII RNA, n cng tham gia vo sa cha DNA b h hi. - TFIIE nhn t cm ng TFIIH vo phc hp khi u phin m V cc nhn t khc SRB-Mediator , SRB10-CDK, SWI/SNF COMPLEX v SAGA c nhim v php bt u phin m

 Cc nhn t ko di - CTD (carboxy terminal domain) l nhn t lin kt vi tiu n v ln nht ca enzym ARN polymerase II, kch hot enzym ARN polymerase II chuyn dch qua promoter v phin m. CTD bao gm 52 ln lp i lp li ca chui axit amin Y-S-P-TS-P-S. Ser5 l v tr phosphoryl ha bi hot ng kinase ca TFIIH - DSIF l nhn t ko di phin m tm dng - NELF cng l mt nhn t khc ko di phin m tm dng, cho php cc enzym capping tham gia v sa i u 5 'ca chui ARN ang tng hp. - P-TEFb (mt kinase) mt yu t ko di th ba tham gia, c vai tr phosphoryl CTD v NELF, trung ha chng. P-TEFb phosphoryl CTD ti Ser2 - TFIIS nhn t phin m b sung, tham gia ko di phin m. Cc nhn t kt thc - Nhn t TTF1 lin kt v gp khc v tr kt thc u 3 ca chui sao chp. - Nhn t PTRF gii phng bn sao chp. c. Qu trnh phin m cc mRNA do RNA pol II xc tc. Khi s RNA pol II bt u phin m nh cc nhn t phin m c bn cht protein. Trc tin nhn t TFIID nhn bit v gn vo trnh t TATA. Tip theo l vic gn thm nhn t TFIIA. Lc , RNA pol II lin kt vi TFIIB, gn vo phc hp TFIID TFIIA. Mt phn t ATP c thy gii to nng lng dng tch 2 mch DNA, phc hp c m. Nhn t TFIIE lin kt vo phc hp khi ng phin m.

V cui cng cc nhn t SRB-Mediator , SRB10-CDK, SWI/SNF COMPLEX v SAGA lin kt vo phc hp v cho php bt u phin m.

Cc vng trc gene (vng bin) l cc v tr nhn bit cho cc yu t phin m c hiu. Nhng yu t ny thng tip xc vi b my phin m thng qua TFIID, TFIIB hay TFIIA, m khng trc tip chm vo RNA pol II. Ph bin nht l vic bm vo TFIID. Vic bm vo cc yu t phin m c hiu gip lp rp b my phin m v do tng tn sut khi u phin m.

Cc trnh t tng cng thc hin chc nng ng nh tn ca chng chng tng cng khi u phin m khi bm vo cc yu t phin m c th.

 Giai on ko di Vic gii phng RNA pol II khi promoter v ko di RNA cn c s h tr ca TFIIS. c bit TFIIH phi c phosphoryl ho phn ui ca RNA pol II trc khi n c th di chuyn. Phn ui, hay CTD (carboxy-terminal domain), gm mt trnh t 7 amino acid (Tyr Ser Pro Thr Ser Pro Ser) lp li khong 52 ln. Trnh t ny c th c phosphoril ho cc ui serine hay threonine. Tt c phc h TFII (tr TFIIH) c b li pha sau khi RNA pol II di chuyn i. Cc NTP b sung ton b gen to tin mRNA. RNA polymerase II ni cc ribonucleotide vo u 3ca si RNA ang c hnh thnh. Chui DNA vng phin m sau khi tho xon, tch hai mch, phi on DNA khun ra phin m, c ng xon tr li. - DSIF v NELF tm dng ny cho php cc enzym capping tham gia v sa i cc u 5 'ca chui ARN ang tng hp. - P-TEFb (mt kinase) mt yu t ko di th ba tham gia, c vai tr phosphoryl CTD v NELF, trung ha chng. P-TEFb phosphoryl CTD ti Ser2

 Giai on kt thc Khi gp trnh t kt thc thng c RNA mi c tng hp hnh thnh cu trc chn vng hay cu trc kp tc. Nh nhn t TTF1 lin kt v gp khc v tr kt thc u 3 ca chui sao chp, RNA polymerase ngng li, kt thc qu trnh phin m. u 3 ca tin mRNA s kt thc bi mt trnh t (trailer), tng ng vi phn khng c dch m ca phin m ca exon 3. Nhn t PTRF gii phng tin mRNA. 3. Cc nhn t phin m ca RNA pol I v qu trnh phin m cc mRNA do RNA pol I xc tc a. Gen m ha cho rARN ln eukaryote Cc gene cho hai phn t rRNA ln c nhiu bn sao, sinh vt nhn chun chng to thnh cm cc trnh t lp lin tip. ngi, c cc cm gene rRNA trn 5 nhim sc th ring bit. rRNA 18S, 5.8S v 28S c phin m cng nhau thnh mt phn t RNA n (45S RNA) trc khi c ct ra thnh ba phn t rRNA. Gia cc n v phin m ny l vng m khng c phin m. cc t bo nhn chun, gene rRNA c RNA pol ring phin m ra chng, RNA pol I.

b. Cc nhn t phin m ca enzym ARN polymerase I Cc nhn t khi u - Nhn t UBF1 (upstream binding factor 1) l mt polypeptide n, c vai tr l nhn t kt ni ngc dng. - Nhn t SL1 chn lc, gm 4 polypeptide: + Mt trong s l TBA (TATA binding protein), TBA cng cn cho enzym ARN polymerase II v enzym ARN polymerase III.

+ Ba n v khc ca SL1 l TAF1 (TBP Associated Factors 1) c xem nh l nhn t kt hp viTBP. Cc nhn t ko di - Nhn t UBF cng c th phn hi thng tin, lm tng s ko di enzym ARN polymerase I qua khng cc cht c ch - Nhn t TFIC cng c th thc y tc qu trnh phin m v ngn chn tm dng ca enzym ARN polymerase I - Nhn t Topl cng l nhn t ko di qu trnh phin m Cc nhn t kt thc - Nhn t TTF1 lin kt v gp khc v tr kt thc u 3 ca chui sao chp. - Nhn t PTRF gii phng bn sao chp. c. Qu trnh phin m cc rARN do enzym ARN polymerase I xc tc Khi u - Cc trnh t khi u phin m (promoter) trn gene Gm 2 vng ring bit: + Vng trung tm (li) khi u phin m (core) cha c im bt u phin m c v tr t -31 n +6 + Vng iu khin ngc dng hay yu t kim sot trc gene UCE (upstream control element) c v tr t -180 n -107 v cch im bt u phin m khong 100 bp Hai vng ny ging nhau khong 80% - 90% v trnh t v u giu GC. Nhn chung cc v tr xung quanh im bt u phin m c xu hng giu AT AND tho xon d hn.

10

- Lp rp phc hp khi u phin m + Mt UBF1 lin kt vi vng UCE v mt UBF1 khc lin kt vi pha trc vng trung tm (core), sau hai UBE1 ny lin kt vi nhau lm DAN hnh thnh vng gia hai v tr c lin kt UBF1. + Nh t SL1 lin kt vi phn xui dng t do ca vng trung tm lm ne nh phc hp UBF-DNA. S kt hp ca SL1lm enzym ARN polymerase Ilin kt c vi phc hp v khi ng qu trnh phin m.

Ko di Tng t nh giai on ko di tng hp mARN ca enzym ARN polymerase II. Nhng khc bit y, l v cc nhn t phin m (khng ging vi on ko di tng hp mARN ca enzym ARN polymerase II) UBF, TIC v Topl. Kt thc Tng t nh giai on kt thc ca enzym ARN polymerase II. V c s tham gia ca hai nhn t kt thc: - Nhn t TTF1 lin kt v gp khc v tr kt thc u 3 ca chui sao chp. - Nhn t PTRF gii phng bn sao chp.

11

12

5. Cc nhn t phin m ca enzym ARN polymerase III v qu trnh phin m cc mRNA do enzym ARN polymerase III xc tc a. Cc nhn t phin m ca enzym ARN polymerase III Cc nhn t khi u - TFIIIA l nhn t lp rp ni vo hp A. Nhn t ny l cn thit ch dnh cho cc gen phin m cc rRNA 5S - TFIIIB nhn t ny c ba tiu n v, trong c TBP (TATA box-binding protein), B v BRF. L mt nhn t v tr . - TFIIIC bao gm 6 tiu n v. N cng c chc nng nh mt nhn t lp rp Cc nhn t ko di (khng cn nhn t ko di) Cc nhn t kt thc (c th ging vi t phin m ca enzym ARN polymerase I v t phin m ca enzym ARN polymerase II) b. Qu trnh phin m cc rARN do enzym ARN polymerase III xc tc Gm 3 loi: Loi 1: Phin m to rRNA 5S - Cc trnh t khi u phin m (promoter) trn gene Vng khi ng ca rRNA 5S gm hp C cch im khi u phin m 81 99 bp v hp A (5-TGGCNNAGTGG- 3) cch im bt u phin m khong 50 65 bp.

- Lp rp phc hp khi u phin m + TFIIIA lin kt vi hp C. + TFIIIC lin kt vi hp A v bao trm c yu t TFIIIA. + Tip n TFIIIB tin vo v lin kt vi AND trc v tr bt u phin m. + Sau enzym ARN polymerase III lin kt vo v tr bt u phin .

13

 loi 2: phin m to tRNA - Cc trnh t khi u phin m (promoter) trn gene Vng khi ng phin m tARN nm sau v tr bt u phin m l A (5TGGCNNAGTGG- 3) v hp B (5 GGTTCGANNCC 3), m ha cho vng D v TC.

14

- Lp rp phc hp khi u phin m - TFIIIC lin kt vi c hp A v hp B trn vng khi ng, l yu t xc nh v tr bt u lin kt TFIIIB. - TFIIIB lin kt vi 50 bp trc hp A, cho php enzyme RNA polymerase III lin kt v dch m.

c. Dng 3: phin m to snRNA - Cc trnh t khi u phin m (promoter) trn gene + Vng khi ng pha trc v tr khi s phin m. + TATA l trnh t ch yu cn cho s phin m, + TBP v cc protein ni vi n xc nh v tr chnh xc cho enzyme RNA polymerase III im khi s phin m. Oct v PSE lm tng hiu qu phin m

15

- Lp rp phc hp khi u phin m + SNAPc (snRNA Activating Protein complex) (cng gi l PBP v PTF) lin kt vi cc PSE ( Proximal Sequence Element) cch v tr bt u phin m khong 55bp v pha trc, v kch thch cc yu t phin m Pol II Oct1 v STAF lin kt vi yu t kim sot thng ngun DSE ( Distal Sequence Element) t nht 200 bp pha trc v tr bt u phin m. Nhng yu t ny v cc promoter c s dng cho enzym ARN polymerase II v enzym ARN polymerase III phin m gen snRNA. + SNAPc lp rp TFIIIB ti hp TATA cch v tr pha trc bt u phin m 26. S hin din ca mt hp TATA cho php cc gene c phin m snRNA bi enzym ARN polymerase III hn l enzym ARN polymerase II. + Cc TFIIIB phin m U6 snRNA cha mt paralogue nh Brf1, Brf2. + TFIIIB l yu t lin kt enzym ARN polymerase III ti v tr bt u phin m.

Ko di Tng t nh giai on ko di ca enzym ARN polymerase I v enzym ARN polymerase II, khng c yu t ko di. Kt thc C th tng t nh giai on kt thc ca enzym ARN polymerase I v enzym ARN polymerase II.

16

III. QU TRNH CH BIN CC TIN mARN 3.1. Ch bin tin mARN * gm 3 bc: Lp m Gn ui poliA Ct b cc intron v ni cc exon

Qu trnh bin i tin mARN

exo n
M7

premRNA mRN A

M7

cap G

intro n

exo n RNA splicing

AAAAAAA200
poly(A) tail

AAAAAAA200

nh n T bo cht ribosom es
a. Lp m

S vn chuyn
M7

AAAAAAA200

protei n

Bc ny c thc hin sau khi enzym ARNpolimerase II tng hp c mt on tin mARN di khong 20-30 Gn thm mt nucleotit (b ci bin) l 7-methylguanosin (7-mG) vo u 5 ca chui ARN.

- M c gn nh: + Enzym guanyltransferase: ni GTP vi nucleotit u tin ca mARN bng lin kt triphotphat 5-5. + Enzym methyl trasferase gn nhm CH3 vo nit s 7 ca vng guanin, ng thi gn thm c vo nhm 2-OH ca ng ribose ca 2 nucleotit k tip. * Cc chc nng c bn ca m

17

 Bo v u 5 ca mARN khi b phn hy bi exonuclease trong t bo cht. Lm tn hiu cho ribosom nhn bit im khi u ca phn t mARN. Tng cng kh nng dch m ca mARN. Gp phn vn chuyn mARN ra khi t bo cht. Tng hiu qu ct ni mARN.

b. Gn ui poliA - u 3 ca phn t tin mARN c sa i bng cch gn vo mt trnh t poly A c th di t 50- 250 base adenin. - Cn c mt tn hiu trn phn t tin mARN: 5-AAUAAA- 3 nm gn u 3 ca phn t tin mARN; khong 11-20 base tip theo c trnh t YA (y=pyrimidin); tip n l on trnh t giu GU nm xui dng.

18

- C s tham gia ca cc protein: CPSF (c v tr lin kt vo mARN l trnh t 5AAUAAA- 3 ), CF1 (c v tr lin kt vo mARN l on giu GU), C F2 (yu t ct mARN), enzym poli A polimerase (PAP), nhn t kch thch ct (CStF) v protein lin kt vi trnh t poliA (PABP) cn gi l PAB II. - Khi cc protein v enzym trn hnh thnh mt phc h poliadenin ho hon chnh lin kt vi mARN, phn t mARN c ct ti v tr 5-CA-3 khi mARN c u 3 t do c gn ui poliA bi hot ng ca PAP.

* Chc nng ca cc ui poliA Tng thi gian tn ti, tnh n nh ca mARN. Tng kh nng dch m ca mARN. Bo v u 3 ca mARN khi exonuclease.

c. Ct b cc intron v ni cc exon

19

* Gm 2 giai on: - Giai on 1: nhm 2OH ca nucleotit A trnh t im phn nhnh tng tc vi lin kt photphodieste u 5 ca G thuc trnh t GU (im ct u 5). Trc tin lin kt photphodieste b ph v, gii phng u 5- G ca intron v u ny c gn vi trnh t im phn nhnh. Lc ny intron hnh thnh mt cu trc vng. - Giai on 2: u 3 ca ca intron b ct sau G ca trnh t AG (im ct u 3), intron c gii phng ra v 2 exon c ni vi nhau. * Thnh phn tham gia: - Qa trnh ct b nh phc h xn intron (spliceosome) gm tin mARN kt hp vi cc ht ribonucleoprotein ( c k hiu l snRNP) - snRNP c to thnh t s lin kt snARN v protein, c 5 loi snARN ph bin nht l U1, U2, U4, U5 v U6 * Qu trnh ct intron tri qua mt s bc nh sau b1) - U1 snRNP gn vo v tr ct u 5 ca intron. Vic gn ny da trn nguyn tc b tr ca U1 snARN c trong snRNP vi trnh t on ni vi exon gn u 5 ca intron. b2) - U2 snRNP gn vo mt trnh t gi l im phn nhnh nm ngc dng so vi on ni vi exon v pha u 3 ca intron. im phn nhnh l v tr c th ca cc intron, ti cha mt adenyl l v tr gn vo ca u 5 t do ca intron trong qu trnh ct b intron. b3) - Phc h U4 / U6 snRNP tng tc vi U5 snRNP ri gn vo cc phc h U1 v U2 snRNP lm hai u 5 v 3 ca intron tin li gn nhau, to thnh cu trc thng lng. b4) - U4 snRNP tch ra khi phc h, lc ny spliceosome chuyn thnh dng c hot tnh ct (exonuclease). b5) - snRNP ct intron u 5 to ra mt u 5 t do. u ny s lin kt vi nucleotit A ti im phn nhnh vo v tr nhm 2- OH (lin kt phosphodieste 5-2). Nhm 3- OH ca adnyl ny vn lin kt bnh thng vi nucleotit khc trong chui. b6) - Intron c ct pha u 5 (intron vn dng thng lng) v cc exon lin k hai u 5 v 3 ca intron lin kt vi nhau. Lc ny phc h snRNP ri khi phn t ARN. V qu trnh ct intron nh vy c lp i lp li.

20

21

d. Kh nng t ct b intron khi mARN

Ngoi phn ng ct b intron trn, th mt s phn t tin mARN ty th hoc lc lp c kh nng t loi cc intron v ni cc exon m khng cn snRNP hoc protein. Gm c 2 nhm khc nhau tu thuc vo cch thc xy ra phn ng l intron nhm 1 v intron nhm 2. * Intron nhm 1 - iu kin: khi phn t mARN v nuceotit G c nhm OH v tr 3. - C ch ct- ni: S c mt ca G dng ny dn n vic ct ti bin gii 5 exonintron, ng thi G c gn vo u intron va c gii phng (bng vic chuyn lin kt photphat t ng ny sang ng kia m khng nh hng n lin kt photphodieste). u t do 3OH ca exon s tng tc vi ni tip gip 3 intron-exon. Tip theo 2 exon c ni vi nhau v intron tn ti dng cu trc vng, cu trc ny sau c chuyn sang dng thng. * Intron nhm 2 - iu kin: Cc intron cha GU u 5. Cc on nucleotit trong intron c th tng tc to cp vi nhau. S thay i cu trc ny lm cho phn t mARN c kh nng t xc tc cho phn ng ct b mt on nucleotit ca chnh n - C ch t ct: xy ra tng t phn ng i hi xc tc ca snRNP. Nhng trong phn ng ct ny khng h c s tham gia ca bt k yu t no ngoi bn thn tin mARN.

22

3.2. Ch bin cc tin rARN - eukaryote, c 4 loi rARN l: 5S; 5,8S; 18S; v 28S - 5S c tng hp t 1 locut c lp - 3 loi rARN cn li c hnh thnh t 1 tin rARN - u tin, cc enzym ribonuclease ct cc tin rARN ti cc v tr c hiu. Sau , cc enzym exonuclease ct ta t hai u 5 v 3 hnh thnh cc rARN hon thin

3.3. Ch bin tin tARN Tin tRNA to cu trc bc hai c endonuclease nhn bit v ct b cc on u mt 5 v 3 tRNA nucleotidyl transferase gn trnh t 5-CCA-3 vo u 3. Sau , chng c loi b intron v ghp thm acceptor vi nhau.

23

TI LIU THAM KHO 1. C s di truyn hc phn t v t bo, inh on Long, NXB HQG HN 2009 2. Di truyn hc, Phm Thnh H, NXB GD 2008 3. Sinh hc phn t, Nguyn Hong Lc,NXB H Hu Nm 2007 4. Sinh hc phn t, H Hunh Thy Dng NXB GD 2008 5. Bn dch t Molecular biology-understanding the genetic revolution ca David D. Clark bi ng Xun Nghim 2010 6. Cc trang web http://thuviensinhhoc.com http://violet.vn/main http://vi.wikipedia.org

http://www.mun.ca/biochem/courses/3107/Topics/euk_transcription.html

24