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TDNA~T ;bk^8O -EH w8hPola! TRNA Pola!~T F %~# ,  y uS p B220^240Kda N!jg~T NvwY ^8O gsQ v S_NS8h RNA Polv2 NWCz+T gW+gzR:S 'YNW B140^150Kda NDNA0^irTeuvRNA~T vS_NS8hRNA Pol  B44.5 vvޏc vS_NS8hRNAvNW RNA Pola! ABC 27KDa xxSˆ}v 1{| NyRNA PolqQ g Y ABC 25.5KDa NDNA~T gsQ ABC 23KDa B 12.6 \NW 2{| Pol a!yr g B 23 B 14.5 B 10 3{| (WgNagN NSdSvNW B 23 SNvvW,g~g B 16.5 ~ހhVvRNA Pol TlṼSOvRNAvPTA SNO1uPy~by:NwYP[initiator,Inr).(WS8hCATwY^R_N gُy`Q 0 N,1uPY2CAPY5gb MON-3^+5 cORNA pol a!Ƌ+R0 eTATA/f&TX[(W Inr[/TRP[v:_^TwYMOpv b/f͑v 0 s]~SNInr ~Tˆ}v0 ZZ%#'3 #'3#'3/3'3/3%#'3&##     zX 8h_CQN b TATAFhTSyHognessFh Goldberg-Hogness Fh O y:NёxGoldbrick vQN^R/fT85A97T93A85A63A83A50 8^(W-25]S vS_NS8hv-10^R0 vQ\O(u/f (1) bcknxvlU_wYMOp O|nxwY Ee_Ny:N bP[selector 0 (2) q_TlU_vs0 8#8?'#'#''#'3/3'3/3'3/3'3/3'3/3'3/3'3/3#"#'3##   A {Y. N8n/TRP[CQNUPE $   h12-4 TsNRirRNA Pol a! N8nlU_VP[~TvCQ CQN O[^R ~TDNAv^ ˆ}v(VP[ TATAbox TATAAAA ^10bp TBP CAAT box GGCCAATCT ^22bp CTF/NF1 GC box GGGCGG ^20bp SP1 Octamer ATTTGCAT ^20bp Oct-1 Octamer ATTTGCAT 23bp Oct-2 KB GGGACTTTCC ^10bp NFKB ATF GTGACGT ^20bp ATF B. Lewin: 0GENES 0e!.1997,Table 28.2 fH'#33  k g|Z X:_P[[/f(W1981t^1uBenerji,Rusconi\~TChambomI{Ssv Sy܏ N8n^R far upstream seguence 0 vQyrp/f `$ wQ g܏ݍyHe^0 a$ eeT'`0 b$ z_0 c$ eiryTWVvyr_'`0 d$ wQ g~~vyr_'`0 e$ gvMO'`0vQ\O(uTDNAvga gsQ0 f$ gvX:_P[SN[YOSNuS^0x/ZZZtZ)#'3##z  Y}[  ~\ FX:_P[:NNHNwQ g܏ݍy\O(ubT 1 b4gHe^b4gHe^􋤋:NX:_P[v \O(u/f[gr(~gSS O8h\ SONuDNaseIOea:S0 2 nR!jW 3 bs!jW0 Enhancer PromotorxZZP# ##"8B;_ * h12-5 N{|a!W/TRP[vlU_VP[ VP[ RP[ϑ R RNAPola! e"10K OV!jgTbRNA TFa!A 12,19,35K 3z[TFa!DTDNAv~T o;mTBPNW TFa!B 33K ~T!jg-10^+10 wYPola!~T TTFa!E/F vN\O(u TFa!D (TBP,30K) TBPNWƋ+RTATA \ZTv~eQ YTSO-N TAFsƋ+R yrk/TRP[ TFa!E 34K() ~T(WPola!vMR O YTSOvOb:S^8O0R N8n 57K() TFa!F 38, 74K 'YNWwQev;m'`RAP74 ,\NWTPola!~T N[vQReQ YTSO TFa!H wQov;m'` SNxxSPola!CzvCTD OPola!8Q ^8O TFa!I 120K Ƌ+RInr wYTFa!F/D~T TFa!J (WTFa!FTReQ YTSO N9eSDNAv~Te_ TFa!S RNATb^8O D""I          ] (6lU_VP[ lU_ YTSO TBP TAFs TFIIA TFIIB TFIIF Pol II TFIIE RNA pol a!vlU_wYB9c<R9)^`( N) RNA pol`!/TRP[68h_/TRP[core promoter b8h_CQNcore element MON-450R+20 #lU_vwY0 N8nc6RCQNupstream,control element UCE N-180^8O0R-107 SXR8h_CQNvlU_wYvHes0Rn%36!%3@!@!a ` 170 160 150 140 130 120 110& & & 60 50 40 30 20 10 1 10 20 N8nc6R:S 8h_/TRP[ UBF1 UBF1~TNG C 0N[:S SL1 SL1NUBF1OS T~T Pol 1 ? 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S8huirlU_vwY^8O( V(N) /TRP[promoter v~gTlU_wY (1)~gxQW,+TƋ+RR),~TB)TwY I MOp; (2)^RO[; (3)MOnTݍykR`[ (4)vcTYZvv~T (5)8^Td~P[v (6)MONWVv52 z (7)Q[lU_v/TRTeT0 (8)yryrkvd~P[vR,B^R N T0R}%#'3#'3#'3#'3# '3#'3Y## #'3#'3#x  dlR E.coli-N N TvsVP[ SƋ+R N Tv/TRP[0*(((4 m0!xQW/TRP[v~g -35 -10 lU_wp TTGACA 16-19bp TATAAT 5-9bp ^c' *49,1. -10^R  N-10^R/f1uPribnowTSchaller1975 Ss Ee_Ny:NPribnowFhvPribnow box 0 O[^R:NT80A95T45A60A50T9 MON-10bp]S A.T0N[ fN0vQR/f: (1) RNA pol'}[~T; (2) b_b_>e/TR YTSO (3) ORNA pol[TlU_0b!%!% !%3!%3-3%3-3%3-3%3-3%3-3%3-3!%3N!--10^R[lU_vHesq_T  TATAAT!AATAAT lU_Hes NM,y:N NMzSdown mutation 0 TATGTT!TATATT lU_Hes NGS :N NGSzSup mutation 0 N NzS:NUOOq_TlU_Hes MRS1uNT-AvXƖ\NA-TvXy TS/f1uNXƖMNOT"l.vQ\ %3!%3!!!%3!%3 !!!%3-!Z /2. -35^R -35^RSy:NSextamavSextama box vQO[^R:NT82T84G78A65C54A45 vQR/f: (1) :NRNA polvƋ+RMOp0 NWƋ+R-35^R :NlU_ b!jg (2)-35T-10^Rvݍy/f3z[v dkNRNA polv~g gsQ0 n@L!%3!%3-3%3-3%3-3%3-3%3-3%3-3%3!%3!%30! (1"3. lU_wYMOpI 0 >lU__Ye!jg Nv,{N*NxW (WS8h-N8^:NAbG NMOnV[P !%3!%3!,2# lU_vwY(1.hQvN!jgvDNAc ub^N Nv, N3z[v YTir(W!jg NyR 2. wYƋ+RhQvN 35^R~T Nu\필vv-/TRP[NCQ YTir closed binary complex  3.hQv'}[0W~T(W-10^RY !jg DNA@\S'` b_b_>ev/TRP[N CQ YTSO 4. vyR0RI ,{N*NrNTPlU__Y, VP[ʑ>e,b_bv-/TRP[-rNTP NCQ Y TSOternary complex 0 ZZ;#'37# '3 #'3#? 3$$RNA8h_vThQv(WDNA NvR^: (,(1) T1/3vRNA pol~TbhQv b(W^yr_MOpv~gce YTSO-N b(W/TRP[-NvNCQ YTSO-N (2) vQ-NJSpev8h_vNNlU_ (3) YO Nv8h_v'YϑX[(WNT~gce YTSO-N (4)0Opeϑ_\vhQv/f8nyv0 Hsa8c^GaHcI5& N S8huirlU_v~bk  ~bkP[terminator,t :_~bkP[ Q~bkP[intrinsic terminators 0 1_~bkP[ VP[rho factor Sy:NOV'`~bkP[ Rho-dependent terminator /        7):_~bkP[v~g & NNAAGCGCCGNNNCCGGCGCTTTTTTNNN& & NNTTCGCGGCNNNGGCCGCGAAAAAANNN& DNA & NNAAGCGCCGNNNCCGGCGCUUUUUUNNN& RNA n  3  3"! *lJ :_~bkP[v~gyrp   (1) gVe~gX[(W 2 v:SW[Q+TG-C (3) :_~bkP[32 z N g6*NU N N~gyrp(W~bk-NwUO\O(u P133L mK N N N N N N G C C G C G G C C G G C C U & NNNNC U UUUU& -OH 3 :_~bkP[~gv!j_V D(P((((((,& nLpN , RNA PollU_DNA VP[D@w0RRNA Ƌ+RMOp N VP[ߍ(WRNAPol TlRNAyR RNA Pol(W~bkMO p\P N v^V P[ N (WlU_l-NVP[ ODNA-RNABgyS _ lU_~bk,ʑ>eQ RNA Pol,P[ T RNAnh./ Ztz43.6LU/717qO  ΑuW zSW 8h|SO~T0RmRNA N 8h|SO;x 8h|SO(Wz NVP[v SMOp㉻y D@wTyR VP[D@w VP[T8h 8h|SO;x 8h|SOc VP[yR lU_~~ lU_cMR ~bkT1ONVWV?  7UrP V RNAZTvTlṼSOlU_V{eu($ F\vlṼSO)R(u[;NvRNA polۏLlU_0 'YvlṼSO]xRNA Pol0 [;NvRNAlU_T7vRgrSO SbWV10 T7vWV1xRNAZTv0 vQ[WVvNirSb6R[;NRNAZTv0 T4lṼSOxNyˆ}vTE.colivRNA polvvN\O(u O[;NvZTvSlU_lṼSOvWV NlU_~̃vWV0 Bq##-#  $sQ& ,{V w8huirvlU_0" w8huirvlU_TS8hlU_v N Tp (1) S8hS gNyRNAZTv w8h~ ހ g NyZTv (2) /TRP[v~gyrp N T w8h g Ny N Tv/TRP[T gsQvCQN (3) w8hvlU_ g_Yˆ}v(VP[vNeQ0 $wx!< WtR N.w8hvRNAZTv$  h12-2 w8huirv NyRNAZTvvyrp RNA Pol MOn Nir v[;m'` [-EHvOe Pol `! 8hN 28s,18s,vRNA(W~ހ(-N dkhfNHN |S! %!%;!%3%z    1956t^E. VolkinT L.Astrachan (u TMO } QN*h hfT2lṼSOeTbvRNAvxWkTvDNAxW5.8s rRNAs 50^70% NOea Pol a! 8h( hnRNA,mRNA,gNSnRNA 20^40% ؚ^Oea Pol b! 8h( tRNA,5SrRNA,gNSnRNAs ^10% Grkyr_ -NI{Oea h12-3 lU_vb6RBR b6RBR vv b6R\O(u )Ry } ~̃hQv TNW~T b6RwY ñ } ~̃8h_v TNW~T b6RwY >e\~ }D w8hPol`! TDNA~T ;bk^8O -EH w8hPola! TRNA Pola!~T F %~# ,  y uS p B220^240Kda N!jg~T NvwY ^8O gsQ v S_NS8h RNA Polv2 NWCz+T gW+gzR:S 'YNW B140^150Kda NDNA0^irTeuvRNA~T vS_NS8hRNA Pol  B44.5 vvޏc vS_NS8hRNAvNW RNA Pola! ABC 27KDa xxSˆ}v 1{| NyRNA PolqQ g Y ABC 25.5KDa NDNA~T gsQ ABC 23KDa B 12.6 \NW 2{| Pol a!yr g B 23 B 14.5 B 10 3{| (WgNagN NSdSvNW B 23 SNvvW,g~g B 16.5 ~ހhVvRNA Pol TlṼSOvRNAvPTA SNO1uPy~by:NwYP[initiator,Inr).(WS8hCATwY^R_N gُy`Q 0 N,1uPY2CAPY5gb MON-3^+5 cORNA pol a!Ƌ+R0 eTATA/f&TX[(W Inr[/TRP[v:_^TwYMOpv b/f͑v 0 s]~SNInr ~Tˆ}v0 ZZ%#'3 #'3#'3/3'3/3%#'3&##     zX 8h_CQN b TATAFhTSyHognessFh Goldberg-Hogness Fh O y:NёxGoldbrick vQN^R/fT85A97T93A85A63A83A50 8^(W-25]S vS_NS8hv-10^R0 vQ\O(u/f (1) bcknxvlU_wYMOp O|nxwY Ee_Ny:N bP[selector 0 (2) q_TlU_vs0 8#8?'#'#''#'3/3'3/3'3/3'3/3'3/3'3/3'3/3#"#'3##   A {Y. N8n/TRP[CQNUPE $   h12-4 TsNRirRNA Pol a! N8nlU_VP[~TvCQ CQN O[^R ~TDNAv^ ˆ}v(VP[ TATAbox TATAAAA ^10bp TBP CAAT box GGCCAATCT ^22bp CTF/NF1 GC box GGGCGG ^20bp SP1 Octamer ATTTGCAT ^20bp Oct-1 Octamer ATTTGCAT 23bp Oct-2 KB GGGACTTTCC ^10bp NFKB ATF GTGACGT ^20bp ATF B. Lewin: 0GENES 0e!.1997,Table 28.2 fH'#33  k g|Z X:_P[[/f(W1981t^1uBenerji,Rusconi\~TChambomI{Ssv Sy܏ N8n^R far upstream seguence 0 vQyrp/f `$ wQ g܏ݍyHe^0 a$ eeT'`0 b$ z_0 c$ eiryTWVvyr_'`0 d$ wQ g~~vyr_'`0 e$ gvMO'`0vQ\O(uTDNAvga gsQ0 f$ gvX:_P[SN[YOSNuS^0x/ZZZtZ)#'3##z  Y}[  ~\ FX:_P[:NNHNwQ g܏ݍy\O(ubT 1 b4gHe^b4gHe^􋤋:NX:_P[v \O(u/f[gr(~gSS O8h\ SONuDNaseIOea:S0 2 nR!jW 3 bs!jW0 Enhancer PromotorxZZP# ##"8B;_ * h12-5 N{|a!W/TRP[vlU_VP[ VP[ RP[ϑ R RNAPola! e"10K OV!jgTbRNA TFa!A 12,19,35K 3z[TFa!DTDNAv~T o;mTBPNW TFa!B 33K ~T!jg-10^+10 wYPola!~T TTFa!E/F vN\O(u TFa!D (TBP,30K) TBPNWƋ+RTATA \ZTv~eQ YTSO-N TAFsƋ+R yrk/TRP[ TFa!E 34K() ~T(WPola!vMR O YTSOvOb:S^8O0R N8n 57K() TFa!F 38, 74K 'YNWwQev;m'`RAP74 ,\NWTPola!~T N[vQReQ YTSO TFa!H wQov;m'` SNxxSPola!CzvCTD OPola!8Q ^8O TFa!I 120K Ƌ+RInr wYTFa!F/D~T TFa!J (WTFa!FTReQ YTSO N9eSDNAv~Te_ TFa!S RNATb^8O D""I          ] (6lU_VP[ lU_ YTSO TBP TAFs TFIIA TFIIB TFIIF Pol II TFIIE RNA pol a!vlU_wYB9c<R9)^`( N) RNA pol`!/TRP[68h_/TRP[core promoter b8h_CQNcore element MON-450R+20 #lU_vwY0 N8nc6RCQNupstream,control element UCE N-180^8O0R-107 SXR8h_CQNvlU_wYvHes0Rn%36!%3@!@!a p 170 160 150 140 130 120 110& & & 60 50 40 30 20 10 1 10 20 N8nc6R:S 8h_/TRP[ UBF1 UBF1~TNG C 0N[:S SL1 SL1NUBF1OS T~T TFa!B Pol 1 ? b# +>[wb  c  1 WQ/TRP[ 2 WQ/TRP[ lU_wYp lU_wYp boxA boxC boxA boxB TFIIIA TFIII C TFIII C TFIII B TBP A B TFIII C TFIII B Plo III Pol III cxi didnUe h12-6 RNA pol b! /TRP[vlU_VP[v~gTR VP[ ~g R TFb!A 38Kda, g9*N c ~TN1WQ/TRP[(5sRNAWV)vCFh Ob!C~T(WCFh N8n Rb! 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RNATbvW,gyrp 1960t^Weiss,S,BI{SsRNAZTvRNA Pol 0 vQyrp/f 1 N8h|8h NxxrNTR :N^ir 2 NDNA:N!jg 3 c52 -32 eTTb 4 e_irvX[(WUSrwYvTb 5 ,{N*N_eQvrNTP/fN Nxxb__X[(W 6 (WSOQDNAS-NNNag\O:N!jg 7 RNAv^RT!jg/fNev0 XZ(ZZZ# #   2 # nx[ gIN\1963 J.MarmurTDoty Spiegelnan:SR gIN0Ǒ(ugIFg̃vSP8lṼSO:NPge SP8 DNA S g { 0 ͑ ]_f>f0 s(W\\O:NlU_!jgvDNAUSy:N!jgbSINantisen strand ^!jgy:N gINsense strand bx0 (WSOYDNAv$NagS\O:NRNATbv!jg0 !%3>!%3!%3!%3!%3!%3!     %h Gene& J`7h(u[[mRNAvTb;`/f^@w5-32 eTۏLv E.coli(W 0Ce13yMbR NN*N8hx FO(W37Cky1\SR N40*N8hx; )R(uُ*N]+RN14CeghU,(W0CW{QE.coli 0 cSُyck(W8OvmRNARP[ Ss14ChHQQs(W8Ov32 z VdkSNfTb/f^@w52 -32 eTۏLv0 /;!!!!!)!!!!)'! ''$ RNATbTDNA Y6Rv:S+R,(1 lU_eS gNagDNA:N!jg Y6R e$NagS\O:N!jg 2 DNA-RNABgTS N3z[ RNATbTʑ>e DNA Y6RSb_bTNvSb_ eTkbP[ 3 RNATb N_ir DNA Y6R_ir 4) lU_v^ir/frNTP Y6Rv^ir/fdNTP 5 ZTv| N T0 (ZZ!  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S8huirlU_vwY^8O( V(N) /TRP[promoter v~gTlU_wY (1)~gxQW,+TƋ+RR),~TB)TwY I MOp; (2)^RO[; (3)MOnTݍykR`[ (4)vcTYZvv~T (5)8^Td~P[v (6)MONWVv52 z (7)Q[lU_v/TRTeT0 (8)yryrkvd~P[vR,B^R N T0R}%#'3#'3#'3#'3# '3#'3Y## #'3#'3#x  dlR E.coli-N N TvsVP[ SƋ+R N Tv/TRP[0*(((4 m0!xQW/TRP[v~g -35 -10 lU_wp TTGACA 16-19bp TATAAT 5-9bp ^c' *49,1. -10^R  N-10^R/f1uPribnowTSchaller1975 Ss Ee_Ny:NPribnowFhvPribnow box 0 O[^R:NT80A95T45A60A50T9 MON-10bp]S A.T0N[ fN0vQR/f: (1) RNA pol'}[~T; (2) b_b_>e/TR YTSO (3) ORNA pol[TlU_0b!%!% !%3!%3-3%3-3%3-3%3-3%3-3%3-3!%3N!--10^R[lU_vHesq_T  TATAAT!AATAAT lU_Hes NM,y:N NMzSdown mutation 0 TATGTT!TATATT lU_Hes NGS :N NGSzSup mutation 0 N NzS:NUOOq_TlU_Hes MRS1uNT-AvXƖ\NA-TvXy TS/f1uNXƖMNOT"l.vQ\ %3!%3!!!%3!%3 !!!%3-!Z /2. -35^R -35^RSy:NSextamavSextama box vQO[^R:NT82T84G78A65C54A45 vQR/f: (1) :NRNA polvƋ+RMOp0 NWƋ+R-35^R :NlU_ b!jg (2)-35T-10^Rvݍy/f3z[v dkNRNA polv~g gsQ0 n@L!%3!%3-3%3-3%3-3%3-3%3-3%3-3%3!%3!%30! (1"3. lU_wYMOpI 0 >lU__Ye!jg Nv,{N*NxW (WS8h-N8^:NAbG NMOnV[P !%3!%3!,2# lU_vwY(1.hQvN!jgvDNAc ub^N Nv, N3z[v YTir(W!jg NyR 2. wYƋ+RhQvN 35^R~T Nu\필vv-/TRP[NCQ YTir closed binary complex  3.hQv'}[0W~T(W-10^RY !jg DNA@\S'` b_b_>ev/TRP[N CQ YTSO 4. vyR0RI ,{N*NrNTPlU__Y, VP[ʑ>e,b_bv-/TRP[-rNTP NCQ Y TSOternary complex 0 ZZ;#'37# '3 #'3#? 3$$RNA8h_vThQv(WDNA NvR^: (,(1) T1/3vRNA pol~TbhQv b(W^yr_MOpv~gce YTSO-N b(W/TRP[-NvNCQ YTSO-N (2) vQ-NJSpev8h_vNNlU_ (3) YO Nv8h_v'YϑX[(WNT~gce YTSO-N (4)0Opeϑ_\vhQv/f8nyv0 Hsa8c^GaHcI5& N S8huirlU_v~bk  ~bkP[terminator,t :_~bkP[ Q~bkP[intrinsic terminators 0 1_~bkP[ VP[rho factor Sy:NOV'`~bkP[ Rho-dependent terminator /        7):_~bkP[v~g & NNAAGCGCCGNNNCCGGCGCTTTTTTNNN& & NNTTCGCGGCNNNGGCCGCGAAAAAANNN& DNA & NNAAGCGCCGNNNCCGGCGCUUUUUUNNN& RNA n  3  3"! *lJ :_~bkP[v~gyrp   (1) gVe~gX[(W 2 v:SW[Q+TG-C (3) :_~bkP[32 z N g6*NU N N~gyrp(W~bk-NwUO\O(u P133L mK N N N N N N G C C G C G G C C G G C C U & NNNNC U UUUU& -OH 3 :_~bkP[~gv!j_V D(P((((((,& nLpN , RNA PollU_DNA VP[D@w0RRNA Ƌ+RMOp N VP[ߍ(WRNAPol TlRNAyR RNA Pol(W~bkMO p\P N v^V P[ N (WlU_l-NVP[ ODNA-RNABgyS _ lU_~bk,ʑ>eQ RNA Pol,P[ T RNAnh./ Ztz43.6LU/717qO  ΑuW zSW 8h|SO~T0RmRNA N 8h|SO;x 8h|SO(Wz NVP[v SMOp㉻y D@wTyR VP[D@w VP[T8h 8h|SO;x 8h|SOc VP[yR lU_~~ lU_cMR ~bkT1ONVWV?  7UrP V RNAZTvTlṼSOlU_V{eu($ F\vlṼSO)R(u[;NvRNA polۏLlU_0 'YvlṼSO]xRNA Pol0 [;NvRNAlU_T7vRgrSO SbWV10 T7vWV1xRNAZTv0 vQ[WVvNirSb6R[;NRNAZTv0 T4lṼSOxNyˆ}vTE.colivRNA polvvN\O(u O[;NvZTvSlU_lṼSOvWV NlU_~̃vWV0 Bq##-#  $sQ& ,{V w8huirvlU_0" w8huirvlU_TS8hlU_v N Tp (1) S8hS gNyRNAZTv w8h~ ހ g NyZTv (2) /TRP[v~gyrp N T w8h g Ny N Tv/TRP[T gsQvCQN (3) w8hvlU_ g_Yˆ}v(VP[vNeQ0 $wx!< WtR N.w8hvRNAZTv$  h12-2 w8huirv NyRNAZTvvyrp RNA Pol MOn Nir v[;m'` [-EHvOe Pol `! 8hN 28s,18s,5.8s rRNAs 50^70% NOea Pol a! 8h( hnRNA,mRNA,gNSnRNA 20^40% ؚ^Oea Pol b! 8h( tRNA,5SrRNA,gNSnRNAs ^10% Grkyr_ -NI{Oea h12-3 lU_vb6RBR b6RBR vv b6R\O(u )Ry } ~̃hQv TNW~T b6RwY ñ } ~̃8h_v TNW~T b6RwY >e\~ }D w8hPol`! TDNA~T ;bk^8O -EH w8hPola! TRNA Pola!~T F %~# ,  y uS p B220^240Kda N!jg~T NvwY ^8O gsQ v S_NS8h RNA Polv2 NWCz+T gW+gzR:S 'YNW B140^150Kda NDNA0^irTeuvRNA~T vS_NS8hRNA Pol  B44.5 vvޏc vS_NS8hRNAvNW RNA Pola! ABC 27KDa xxSˆ}v 1{| NyRNA PolqQ g Y ABC 25.5KDa NDNA~T gsQ ABC 23KDa B 12.6 \NW 2{| Pol a!yr g B 23 B 14.5 B 10 3{| (WgNagN NSdSvNW B 23 SNvvW,g~g B 16.5 ~ހhVvRNA Pol TlṼSOvRNAvPTA SNO1uPy~by:NwYP[initiator,Inr).(WS8hCATwY^R_N gُy`Q 0 N,1uPY2CAPY5gb MON-3^+5 cORNA pol a!Ƌ+R0 eTATA/f&TX[(W Inr[/TRP[v:_^TwYMOpv b/f͑v 0 s]~SNInr ~Tˆ}v0 ZZ%#'3 #'3#'3/3'3/3%#'3&##     zX 8h_CQN b TATAFhTSyHognessFh Goldberg-Hogness Fh O y:NёxGoldbrick vQN^R/fT85A97T93A85A63A83A50 8^(W-25]S vS_NS8hv-10^R0 vQ\O(u/f (1) bcknxvlU_wYMOp O|nxwY Ee_Ny:N bP[selector 0 (2) q_TlU_vs0 8#8?'#'#''#'3/3'3/3'3/3'3/3'3/3'3/3'3/3#"#'3##   A {Y. N8n/TRP[CQNUPE $   h12-4 TsNRirRNA Pol a! N8nlU_VP[~TvCQ CQN O[^R ~TDNAv^ ˆ}v(VP[ TATAbox TATAAAA ^10bp TBP CAAT box GGCCAATCT ^22bp CTF/NF1 GC box GGGCGG ^20bp SP1 Octamer ATTTGCAT ^20bp Oct-1 Octamer ATTTGCAT 23bp Oct-2 KB GGGACTTTCC ^10bp NFKB ATF GTGACGT ^20bp ATF B. Lewin: 0GENES 0e!.1997,Table 28.2 fH'#33  k g|Z X:_P[[/f(W1981t^1uBenerji,Rusconi\~TChambomI{Ssv Sy܏ N8n^R far upstream seguence 0 vQyrp/f `$ wQ g܏ݍyHe^0 a$ eeT'`0 b$ z_0 c$ eiryTWVvyr_'`0 d$ wQ g~~vyr_'`0 e$ gvMO'`0vQ\O(uTDNAvga gsQ0 f$ gvX:_P[SN[YOSNuS^0x/ZZZtZ)#'3##z  Y}[  ~\ FX:_P[:NNHNwQ g܏ݍy\O(ubT 1 b4gHe^b4gHe^􋤋:NX:_P[v \O(u/f[gr(~gSS O8h\ SONuDNaseIOea:S0 2 nR!jW 3 bs!jW0 Enhancer PromotorxZZP# ##"8B;_ * h12-5 N{|a!W/TRP[vlU_VP[ VP[ RP[ϑ R RNAPola! e"10K OV!jgTbRNA TFa!A 12,19,35K 3z[TFa!DTDNAv~T o;mTBPNW TFa!B 33K ~T!jg-10^+10 wYPola!~T TTFa!E/F vN\O(u TFa!D (TBP,30K) TBPNWƋ+RTATA \ZTv~eQ YTSO-N TAFsƋ+R yrk/TRP[ TFa!E 34K() ~T(WPola!vMR O YTSOvOb:S^8O0R N8n 57K() TFa!F 38, 74K 'YNWwQev;m'`RAP74 ,\NWTPola!~T N[vQReQ YTSO TFa!H wQov;m'` SNxxSPola!CzvCTD OPola!8Q ^8O TFa!I 120K Ƌ+RInr wYTFa!F/D~T TFa!J (WTFa!FTReQ YTSO N9eSDNAv~Te_ TFa!S RNATb^8O D""I          ] (6lU_VP[ lU_ YTSO TBP TAFs TFIIA TFIIB TFIIF Pol II TFIIE RNA pol a!vlU_wYB9c<R9)^`( N) RNA pol`!/TRP[68h_/TRP[core promoter b8h_CQNcore element MON-450R+20 #lU_vwY0 N8nc6RCQNupstream,control element UCE N-180^8O0R-107 SXR8h_CQNvlU_wYvHes0Rn%36!%3@!@!a p 170 160 150 140 130 120 110& & & 60 50 40 30 20 10 1 10 20 N8nc6R:S 8h_/TRP[ UBF1 UBF1~TNG C 0N[:S SL1 SL1NUBF1OS T~T TFa!B Pol 1 ? b# +>[wb  c  1 WQ/TRP[ 2 WQ/TRP[ lU_wYp lU_wYp boxA boxC boxA boxB TFIIIA TFIII C TFIII C TFIII B TBP A B TFIII C TFIII B Plo III Pol III cxi didnUe h12-6 RNA pol b! /TRP[vlU_VP[v~gTR VP[ ~g R TFb!A 38Kda, g9*N c ~TN1WQ/TRP[(5sRNAWV)vCFh Ob!C~T(WCFh N8n Rb! B[MO~T TFb!B +TTBPTSYNyˆ}v [MOVP[ OPol~T(WwYMOp N TFb!C +TATB, g5*NNW B~Ta!WQ/TRP[(tRNAWV)vBFh wX:_P[v\O(u0 A~TAFh w/TRP[ v\O(uRb! B[MO~T TBP /fb!B,a!D,SL1vNW Tyr_DNA^RSRNA Pol~T OPol~T (WcknxvMOp N TFa!D +TTBPNW ~TTATAFh nx[ bPolb! PBP !kяz~Tˆ}v STa!DNSRb! B[MO~TvSN _ N p, "  ),) *d  TBP Pol III /T R P[ TFIIIB TFIIIC TBP RNA Pol III Pol I /T R P[ SL1 RNA Pol I TBP UBF1 Pol II /T R P[ TFIID TATA lU_wYp RNA Pol II Z   %"2 "g J "3'" "3"F &dS=/ ALRn  lU_wYp OSE PSE TATA OCT-1 PBP TFa!D OSE (OCT sequuence element) TFb!B PSE(prixima sequuence element) RNA pol b! p@\#-##?/#   fgR  (,:(  2 + Zu! @ K|2  Ti @F@E|2  T @t IM|2  Tt @o /|2  Tt @ yR  s *s *R  s *& R  s *}Zr  S \@ %  XB  0DoP4tPXB  0DoY2 Z^B   6D3o}}XB   0D3o}m2    BCE$G>bHkLI`TQo FH᧎gFH᧎g`T`TFHg`T`T2iXB   0DoRXB   0Do YYiXB @ 0DoNZNXB  0DoLiP|2  T. @`2   B"JCRENGIHyJRQ3o `T~d"J)`T~d"J)Rd"J)R |2  Ti @ |2  T @k @ |2  Tt @ K |2  Tt @| U\ R  s *s  R  s *  R  s *} 6w XB   0Do P XB ! 0Do   ^B " 6D3o= = XB # 0D3o= - 2 $  BCE$G>bHkLI`TQo FH᧎gFH᧎g`T`TFHg`T`TwE XB % 0Do  XB & 0Do 55) XB '@ 0Do 6 XB ( 0Do E |2 ) T. @ e  2 *  CYENGH.I`TJnLQ3o Π Y0Π Y0`TnL Y0`TnL . 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