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NI DUNG CHNH
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CU TRUC PROTEIN
* Cu truc: Amino acid la n vi c s (monomer) cu thanh protein. Co 20 amino acid u co cu truc 3 phn: + Gc NH2 + Gc COOH + Gc - R mach bn - Gc R (mach bn) cung la phn khac duy nht gia 20 loai amino acid, quy inh tinh cht cua tng loai. - Nhom amine (NH2) inh nguyn t C2. - Cac amino acid tn tai chu yu trong t nhin co nhom amine ng bn trai truc, c goi la amino acid dang L. Dang Damino acid chi tn tai ring bit, vi du trong thanh t bao vi khun.
CU TRUC PROTEIN
- Cu truc protein gm 4 bc: + Cu truc bc 1: La trinh t sp xp cac gc amino acid trong chui polypeptide. Cu truc nay c gi vng nh lin kt peptide (lin kt cng ha tri). + Cu truc bc 2: La tng tac khng gian gia cac gc amino acid gn nhau trong chui polypeptide. Cu truc c bn vng chu yu nh lin kt hydrogen hnh thanh gia cac lin kt peptide k gn nhau, cach nhau nhng khoang xac inh
CU TRUC PROTEIN
- Cu truc bc 3. La tng tac khng gian gia cac gc amino acid xa nhau trong chui polypeptide, la dang cun lai trong khng gian cua toan chui polypeptide. - Cu truc bc 4. L tng tac khng gian gia cac chui cua cc phn t protein gm hai hay nhiu chui polypeptide hnh cu. Mi chui polypeptide nay c goi la mt tiu n vi (subunit). S kt hp gia cac phn t nay long lo va chu yu la do lin kt hydrogen va ky nc. Bng cach nay hai phn t xac inh co th kt hp vi nhau tao thanh mt dimer
S PHIN MA
- Bc u trong s biu hin gen - Enzyme: ARN polymerase xc tc s tho xon, tch hai mach n cua ADN v b sung cc ARN nucleotides - Nguyn liu: NTP ATP, GTP, CTP, UTP trn DNA cua vi khun, vng trnh t m RNA polymerase c th gn vo c goi l vng khi ng (promoter); vng c cha tn hiu kt thc phin m c goi l vng kt thc (terminator) - oan ADN c phin m thnh mARN c goi l n vi phin m (transcription unit)
* S bin i hai u tn cng cua ARN H tr s chuyn mARN t trong nhn ra TBC Bao v mARN khoi tc dung cua cc enzyme thuy phn Gip ribosome gn c vo u 5 cua mARN khi giai m
S CT NI RNA
- Ban RNA mi tao ra co vung khng ma hoa (introns) va nhng vung ma hoa (exon) - Vung ma hoa exon giai ma thanh trinh t aa - Cac intron: bi ct va di i - Cac exon ni lai thanh mRNA co trinh t ma hoa lin tuc - S ct i c tin hanh bi: spliceosome (mt s loai protein va cac small nuclear ribonucleoproteinssnRNPs 4 loai)
Phin m
Qu trnh phin m sinh vt nhn thc khc g so vi nhn s?
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S GIAI MA
- Thng tin di truyn nm trong gen di dang m b ba: 3 nucleotide k tip nhau trn mach khun cua gen m ha cho mt acid amin trong phn t protein. - Cc b ba trong mARN c goi l cc codon, s c oc theo chiu 5 3
C IM MA DI TRUYN
- La ma b ba, khng phu ln nhau - La ma suy thoai: trong 64 b ba, c 61 b ba m ha cho 20 acid amin; 3 b ba kt thc khng m ha cho acid amin - Cc b ba khc nhau c th m ha cho cng mt acid amin. - M di truyn c tnh c hiu: mi b ba chi m ha cho mt acid amin. - M di truyn c tnh ph bin: cc loi khc nhau u c chung m di truyn
- Ribosomes.
- ARN vn chuyn (tARN).
Cc loai ARN
Cu trc
- Gm 1 mach polynucltit, mach thng. - u 5 c trnh t nuclit c hiu (khng uc dch m) gn codon m u ribxm nhn bit v gn vo.
Chc nng
mARN
tARN
rARN
RIBOSOME
Ribosomes h tr s bt cp c hiu gia anticodon cua tARN vi codon cua mARN trong s tng hp protein Hai bn n vi (ln v nho) cua ribosome u c cu thnh t cc protein v rRNA
Giai m
* C ch:
C CH GIAI MA
* Giai oan m u - S kt hp cua mARN, mt tARN c gn acid amin m u (Met) v hai bn n vi cua ribosome tao thnh phc h m u + Bn n vi nho cua ribosome gn vi mARN v Met tARN + Bn n vi nho di chuyn doc theo mARN n khi gp m m u (AUG) + Mt proteins goi l yu t m u (IF = initiation factor) mang bn n vi ln n gn vo phc h m u
C CH GIAI MA
Giai oan ko di - Cc acid amin gn vo acid amin trc o - Mi acid amin c gn thm vo nh c s h tr cua cc protein goi l yu t ko di (EF = elongation factor) - Giai oan ny gm 3 bc lp i lp lai: S nhn dang codon S thnh lp lin kt peptide S chuyn vi (translocation) nn c goi l chu k giai m
C CH GIAI MA
* Giai oan kt thc - Phin m s kt thc khi mt stop codon trong mARN i vo vi tr A cua ban n vi ln cua ribosome - Mt protein goi l yu t phng thch (RF = release factor) i vo vi tr A lm gn thm mt phn t nc thay v mt acid amin - Phan ng ny lm phng thch chui polypeptide v hai bn n vi cua ribosome tch nhau ra
POLYRIBOSOMES
- Mt mARN c gn cng mt lc vi nhiu ribosomes, tao thnh mt polyribosome (cn goi l polysome) - Polyribosome cho php mt t bo nhanh chng tao ra nhiu chui polypeptide ging nhau. - Chui polypeptid va c giai m cha th thc hin chc nng - Chui polypeptide cn phai c bin i: acid amin m u, mt s oan cua chui polypeptid bi ct bo, protein bin i cu hnh... - Protein hon chinh c chuyn n nhng vi tr cn thit thc hin chc nng.
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Dch m
Tnh trang
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Mt s amip biu hin s thay i hnh thi v sinh l c bit ap lai cc iu kin mi trng khc nhau. Khi cc amip c cho vo nc, chng chuyn t dang amip sang dang c lng bi. Khi mi trng thiu dinh dng, chng c th bin i thnh dang m biu b.
- Vi khun trong mi trng dinh dng ti thiu c kha nng tng hp amino acid. Nhng nu cho amino acid vo thng mi trng nui, vi khun ngng tng hp amino acid. n khi ngun amino acid ht, n mi t tng hp amino acid cho ban thn. - Cc bin i trn u thun nghich chng to s thay i chc nng y khng do bin di di truyn v s xut hin hay bin mt cc cu trc mi khng lm anh hung tim nng di truyn sn c. Cc hin tng trn do hin tng iu ha goi l iu ha thch nghi
Bit ha t bo
- Nhiu sinh vt bc cao nh con ngi cha nhiu ti t bo bt ngun t mt hp t do phn chia nguyn nhim. - T mt hp t khi trung thnh, c th ngi c khoang 200 loai t bo khc nhau (t bo gan, tim, thn, da, ). - Mi loai t bo chi biu hin mt phn thng tin. - Qu trnh chuyn mn ha chc nng cua t bo c goi l s bit ha t bo. - Tuy c s bit ha nhng t bo vn gi nguyn ven kha nng di truyn cua mnh.
C IM IU HA BIU HIN GEN S biu hin cua gen chiu s kim sot cua c ch iu ha. ap lai cc bin i cua mi trng bn trong v ngoi c th. Thc hin nhiu mc khc nhau v lin quan n tng giai oan pht trin. Gen iu ha gi vai tr quan trong trong vic ong m cc gen cu trc c biu hin tng hp protein ung lc, ung ni theo nhu cu cu th cua t bo
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m, sau dch m.
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CAC NGUYN LY IU HA
- Mc biu hin cua gene khc nhau gia cc t bo hoc khc nhau theo giai oan trong chu trnh t bo - Hoat tnh cua gene khc nhau theo chc nng t bo - Tt ca cc t bo u cha cng thng tin di truyn, nhng t bo khc nhau chi nhng gene hoat ng - Hoat tnh cua gene c iu ha mc phin m, ca qua nhng tn hiu bt u bn trong t bo v ca phan ng vi nhng iu kin bn ngoi. Tuy nhin thng tin di truyn c iu ha theo nhng cch khc nhau.
CAC NGUYN LY IU HA
* Cc bc iu khin hoat ng gene bao gm: - Cu trc lai DNA, trong o nhng thay i biu hin gene phu thuc vo vi tr trnh t DNA trong genome. - iu ha phin m trong tng hp ban phin m RNA bng s iu khin s m u v s kt thc - Qu trnh ch bin RNA hoc iu ha qua qu trnh ct ni trn RNA (RNA splicing) - iu ha dich m qu trnh tng hp chui polypeptid
MC PHIN MA
- Prokaryote l nhng c th n bo sng t do, sinh trng v phn chia trong iu kin thch hp v c cung cp y u cht dinh dng. Do o hoat ng cua cc gene c iu ha do nhu cu cua t bo khi cn thit.
MC PHIN MA
- Eukaryote l nhng c th a bo. - Trong c th ang pht trin, mt t bo khng chi sinh trng v phn chia m t bo th h sau trai qua nhng thay i quan trong v hnh thi v ha sinh v duy tr trang thi bin i cua n. - Trong qu trnh pht trin phi, t bo eukaryote t chiu anh hng cua mi trng hn so vi vi khun. - C th trng thnh, s sinh trng v phn chia t bo cua hu ht cc loai t bo bi ngng, mi t bo chi phai duy tr cc tnh cht ring bit cua n
T bo c th kim sot s tng hp enzyme bng mi lin h ngc m tnh hoc bng s iu ha hoat ng gen.
IU HA M TINH
- Mt protein c ch c mt trong t bo, ngn can s phin m. - Trong h thng cam ng c iu ha m tnh, protein c ch hoat ng lm ngn can phin m. - Nhn t cam ng km hm cht c ch, cho php bt u phin m. - Trong h thng c ch, protein aporepressor gn vi co-repressor tao ra cht c ch c hoat tnh, lm ngn can phin m
IU HA DNG TINH
- S tng hp mRNA xay ra nu protein iu ha gn vo mt vng cua gene lm hoat ha phin m. - Nhng protein ny c xem l nhng nhn t hoat ha phin m. - iu ha m tnh v dng tnh khng loai tr ln nhau. - Mt vi h thng l ca iu ha m tnh v dng tnh, s dung ca 2 h thng iu ha phan ng vi cc iu kin khc nhau trong t bo. - iu ha m tnh l ph bin cho prokaryote, trong khi iu ha dng tnh lai ph bin cho eukaryote.
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CU TRUC OPERON
H thng s dung lactose gm 2 loai thnh phn: + Gene cu trc m ha protein cn thit cho s vn chuyn v chuyn ha lactose + Cc yu t iu ha: gene c ch lacI, promotor lac P v operator lacO.
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CU TRUC OPERON
- Gene cu trc c m ha bi mt phn t mRNA a gene (polycistronic). + Gene Z m ha cho enzyme - galactosidase (thuy phn ng lactose thnh galactose v glucose) + Gene Y m ha cho enzyme permease (cn cho vn chuyn lactose qua mng) + Gene A m ha cho enzyme transacetylase (vai tr chuyn ha lactose cha r).
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ngn phin m
Vng m ha
Vng iu ha
-Vng iu ha: +Vng khi ng (promoter): c trnh t nu c th gip enzim ARN polimeraza nhn bit mach m gc mARN v ni bt u phin m. +Vng vn hnh (operator): trnh t nu c bit protein iu ha bm vo ngn phin m.
ADN
P O
Vng khi ng
Opron l mt nhom gen co lin quan v chc nng v co chung mt c ch iu ho. V d: opron Lac E.Coli
Opron
Gen iu ho
Vng vn hnh
Cc gen cu trc (Z, Y, A) c lin quan v chc nng
ADN
P O
Vng khi ng
Opron Lac gm: + Vng khi ng (P): ARN polimeraza bm vo v bt u phin m. + Vng vn hnh (O): protein c ch lin kt ngn phin m + Nhm gen cu trc (Z, Y, A): tng hp cc enzim tham gia vo cc phan ng phn giai ng lactozo/mi trng nng lng 5
Opron
Gen iu ho
Vng vn hnh
Cc gen cu trc (Z, Y, A) c lin quan v chc nng
ADN
P O
Vng khi ng
ngn phin m protein c ch * Gen iu ha (R): khng nm trong operon nhng quan trng, tng hp protein c ch lin kt vng vn hnh ngn phin m
CU TRUC OPERON
- San phm cua gene lacI l cht c ch, n bm vo trnh t cc base cua DNA cu tao operator. - Cht c ch bm vo operator, ngn can s khi u phin m mRNA nh RNA polymerase. - Cht cam ng (lactose) kch thch s sinh tng hp mRNA bng cch kt hp v lm bt hoat cht c ch. - S c mt cua cht cam ng lm cht c ch khng gn vo operator, promotor cho php khi u tng hp mRNA.
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Trng hp operon lactose: - Thng thng t bo vi khun s dung ng glucose nhng khi thiu glucose m c lactose, t bo s tao cc enzyme phn huy lactose. - T bo E.coli c 3 gen (G1, G2, G3) cu trc cua 3 enzyme (betagalactosidase, permease, transacetylase) xc tc cc phan ng chu trnh phn huy lactose.
- Gen iu ha R tng hp cht km hm l 1 protein. - Khi c ng lactose trong t bo, n s kt hp vi cht km hm lm cht ny mt hoat tnh. Nh o, vic phin m cc gen cu trc c tin hnh nh tng hp cc enzyme (E1, E2, E3) bin i ng lactose. - Khi ng lactose bin i ht, cht km hm c giai phng, n tr lai km hm gen chi huy lm qu trnh phin m bi inh tr, s tng hp cc enyme bi ngng.
Ribosome
RNA polymerase
b)
Ribosome
RNA polymerase
Design by HTV
* Khi mi trng khng c lactose: protein c ch c hoat tnh gn vo operator, lm s phin m cua tt ca cc gene cu trc cua operon lac bi dng.
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ADN
Khng hoat ng
Prtin c ch
Gen iu ha
Protein c ch
gn vao
Vng Operator
Ngn phin ma
ADN
Khng gn vo
Vng vn hnh (operator) ARN polimeraza Lin kt Vng khi ng
Phin m
mARN ca gen Z,Y,A
Dng phin m
ADN
ADN mARN
Prtin c ch b bt hoat
CU TRUC cAMP
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Khi lng glucose trong mi trng tng, CAP tch khoi lac operon, v tc phin m tr lai bnh thng CAP gip iu ha cc operon khc m ha cho cc enzymes c dng trong con ng di ha
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- Operon tryptophan (trp) cua E.coli cha cc gene cu trc m ha cho cc enzyme tng hp amino acid tryptophan - Operon ny c iu ha theo cch sau: khi tryptophan c mt y u trong mi trng sinh trng, s phin m cua operon bi c ch. Khi s cung cp tryptophan bi thiu, s phin m xay ra
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- S iu ha cua operon tryptophan tng t vi operon lactose, v s tng hp mRNA c iu ha m tnh nh cht c ch. - Khc vi iu ha operon lac, tryptophan hoat ng nh cht ng km hm, kch thch cht c ch gn vo operator ngng s tng hp. - Operon tryptophan hoat ng theo kiu c ch, iu ha m tnh.
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- Tryptophan c tng hp qua 5 giai oan, mi giai oan c s xc tc cua mt enzyme c bit. - Cc gene m ha cho cc enzyme ny nm k nhau trn nhim sc th cua E.coli.. - o l cc gene trpE, trpD, trpC, trpB, trpA. Cc enzyme c dich m t mt phn t mRNA a gene.
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- Vng m ha gene E c dich m trc tin. - Pha trc trpE v u 5' c promotor, operator v 2 vng xp ln lt l leader (trpL) v oan km hm phin m attenuator (trpa, khng phai l trpA). - Gene c ch trpR nm xa operon, tng hp protein aporepressor, l cht km hm m ring n khng c hoat tnh
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- Khi tryptophan d tha, n kt hp vi aporepressor tao cht km hm c hoat tnh, gn vo operator cua operon tryptophan lm dng phin m cc gene cu trc. - Khi nng tryptophan thp, n tch khoi phc hp km hm v aporepressor mt hoat tnh. Lc ny operator m ra, RNA polymerase dich m 5 gene cu trc tng hp 5 enzyme tao tryptophan
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IU HA M TINH OPERON TRYPTOPHAN Operon cam ng (inducible operon): Bnh thng khng hoat ng (ng). Khi c mt phn t goi l cht cam ng (inducer) lm bt hoat cht c ch gen phin m (m). Lac operon l mt operon cam ng.
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PHIN MA D (ATTENUATION) - Kiu iu ha th hai c pht hin operon tryptophan c goi l attenuation. - N dng s dich m iu khin s phin m. - Khi c mt tryptophan ni bo, ngay ca vi nng thp, s dich m mt phn vng leader cua mRNA ngay khi va c tng hp, kt qua lm dng s phin m trc khi gene cu trc u tin cua operon c sao chp
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PHIN MA D (ATTENUATION)
- Attenuation l kt qua s tng tc gia cc trnh t DNA trong vng leader cua ban phin m trp. - Khi c mt mt lng nho tryptophan, hu ht phn t mRNA kt thc vng 28 base c bit trong trnh t leader. - Kt qua cua s kt thc sm ny tao phn t mRNA cha 140 nucleotide chm dt mt oan ngn cua cc gene m ha cho cc enzyme trp. - Vng 28 base xay ra s kt thc c goi attenuator. Trnh t base cua vng ny thng c cc tnh cht im kt thc, gm dang oan v vng (stem-loop) trn mRNA theo sau l trnh t cua 8 uridine. 95
PHIN MA D (ATTENUATION)
* Trnh t leader c cc c im: - Mt vng c codon AUG v pha sau l codon kt thc UGA, m ha cho mt polypeptide cha 14 amino acid c goi l leader polypeptide. - Hai codon tryptophan vi tr 10 v 11 trn mRNA cua leader polypeptide. Trnh t lp lai ngn ny c ngha trong iu ha. - Bn oan cua RNA leader l vng 1, 2, 3 v 4 tao thnh do kha nng kt cp cua cc base vi nhau. Cc base vng 1 kt cp vi vng 2, vng 3 kt cp vi 96 vng 4
A. S phin m ca leader trp B. Chi tit cu trc ca 2 codon trp vng 1-2
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PHIN MA D
- Mt kiu kt cp bin i c th xy ra, trong cc base vng 2 kt cp vi vng 3 nh cc cp base 2 vng ny gn nh b sung nhau. - Qua m hnh kt cp base bin i ny (3-4 hoc 23), s t chc trnh t mRNA c th iu ha phin m qua dch m ca leader polypeptide. - Khi vng leader c phin m, s dch m leader polypeptid cng bt u. V c 2 codon trptophan trong trnh t m ha, nn s dch m nhy cm vi s lng tRNAtrp a vo
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PHIN MA D
- Nu mi trng cung cp y u tryptophan, ribosome trt qua codon tryptophan v i vo vng 2. - S c mt cua ribosome loai bo kha nng kt cp cua vng khoang 10 base mi pha cua codon ang dich m. - S c mt cua ribosome vng 2 ngn can n kt cp vi vng 3. - Vng 3 kt cp vi vng 4, tao ra im kt thc phin m. - S phin m kt thc khi qua cc uridine nm pha sau vng 4.
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A. mRNA t do co s kt cp base gia 1-2 v 3-4 B. nng cao ca tryptophan, ribosome tin n vng 2 v s kt cp 3-4 lm kt thuc phin m C. nng tryptophan thp, ribosome vng codon trp cho php kt cp 2-3 v phin m khng b kt thuc sau khi qua
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PHIN MA D
- Khi s lng tRNAtrp khng u, s dich m leader polypeptide bi dng lai t ngt cc codon tryptophan. - S dng lai ny ngn can ribosome tin vo vng 2, v vy vng 2 c t do s kt cp vi vng 3 lm can tr s hnh thnh cu trc kt thc. - V vy phn t trp mRNA hon chinh c tao thnh, cha ca trnh t m ha cho gene cu trc
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PHIN MA D
- Attenuation l c ch iu ha tinh t trn c s iu ha m tnh: Khi tRNAtrp n u cung cp cho s dich m leader polypeptide, s phin m bi dng, cc trp enzyme khng c tng hp. Khi nng tRNAtrp qu thp, s phin m xay ra cho n ht, cc trp enzyme c tao nn.
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PHIN MA D
- iu ha kiu attenuation khng th xay ra eukaryote v eukaryote s phin m v dich m khng xay ra ng thi. S phin m xay ra trong nhn, cn s dich m xay ra t bo cht. - iu ha operon lac v operon trp l v du v mt trong s cc c ch quan trong iu ha hoat ng gene mc phin m cua prokaryote.
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1. iu ha v biu hin bng thay i cu trc NST hay cu trc phn t DNA 2. iu ha mc phin m 3. Ct bo intron, ni exon (Splicing) 4. mARN gn vi mt s protein c hiu xuyn qua l nhn ra bo tng 5. Hu cc mARN khng c dng dich m 6. Dich m, tng hp Protein 7. Bin i protein 8. iu ha bng c ch phn huy Protein
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1. iu ha v biu hin gen bng thay i cu trc NST hay cu trc phn t DNA Chu yu l iu ha cu trc cua cht nhim sc (chromatin) Trong vng di nhim sc (heterochromatin) l ni cht nhim sc ong xon cht, cc gen thng khng biu hin S bin i ha hoc cua cc histone v DNA cua chromatin c anh hng n cu trc cua chromatin v s biu hin cua gen
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Nhm acetyl c gn vo lysine tch in dng trong ui histone chromatin tho xon phin m.
b. S methyl ha DNA
Nhm methyl c gn vo chromatin. S gn thm nhm phosphate gn cc acid amin bi methyl ha s lm cho chromatin tho xon.
Trong cc sinh vt a bo, metyl ho ADN l mt du hiu biu hin gen lin quan ti s bit ho cua m t bo. Trnh t nhn bit cho s metyl ho rt ngn, ng vt thng l CG cn thc vt l CNG.
2. iu ha hoat ng gen bng kim sot phin m a/iu ha biu hin bng cc trnh t tng cng (Enhancer) phin m:
- Mt s yu t trnh t iu khin, goi l cc yu t iu khin gn, nm ngay gn promoter, cn cc yu t iu khin xa nm cch promoter mt oan xa hn v chng tp hp thnh mt nhm c goi l cc trnh t tng cng (enhancer)
Cc yu t hoat ho sinh vt bc cao nh l cc yu t h tr RNA polymerase di chuyn ln pha trc k t vi tr promoter trong qu trnh phin m. Cc phc protein c kha nng gn vi phn pha trn cua RNA polymerase II do RNA polymerase c cc vi tr c hiu gn cho cc yu t ny c bit i vi cc yu t c kha nng lin kt vi cc enhancer.
c.Protein hot ha (activator protein): Protein hoat ha kt hp vi nhau, gn vi trnh t enhancer v cc nhn t phin m, enzyme RNA polymerase II kch hoat qu trnh phin m. Cc protein hoat ha gn cng nhiu trnh t enhancer, phin m cng manh, c th cc ai.
3. S ct ni cc intron v exon
Qa trnh ny nh spliceosome (gm snRNP v cc protein) Hai Nu u tin GU u 5 v AG u 3 cua mi intron l vi tr nhn bit ct intron ra khoi phn t tin mARN
Ba bc ca c ch ct-ni (splicing) pre-mRNA sinh vt bc cao (1) lp m 5 (xy ra cng lc phin m); (2) tch b v gn ui poly(A); v (3) splicing (xy ra trong nhn trc khi mRNA i ra t bo cht)
5 - Hy mARN (nu mARN khng c dch m) Nu mARN khng c dich m, di tc dung cua ribonuclease s phn huy phn t mARN ny. S thoi ha mRNA l mt c ch kim sot chu yu trong biu hin gen sinh vt bc cao, thi gian tn tai cua mRNA phu thuc vo di trnh t AU vng 3 khng dich m v s kh ui poly A.
7 Bin i Protein
Sau khi c tng hp, chui polypeptide cn phai c bin i: acid amin m u, mt s oan cua chui polypeptid bi ct bo, protein bin i cu hnh, gn thm mt s chc ha hoc nh acetat, photphat, lp cu ni disulfide hoc ct bo 1 oan peptid gia protein thc hin chc nng cua n.
Cc protein iu ha thng c hoat ha hoc bt hoat mt cch ph bin tng ng bng vic c gn thm nhm phosphate (phosphoryl ha) hoc loai bt i nhm phosphate (loai phosphoryl ha), trong khi o cc protein c chuyn n b mt t bo ng vt thng c gn thm cc gc ng. Cc protein b mt t bo v nhiu protein khc phai c vn chuyn n ich trong t bo l ni chng c th biu hin chc nng. S biu hin cua gen c th xut hin trong mi bc lin quan n qu trnh hon thin v vn chuyn protein trc y.
THANK YOU
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