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R*  p  0 ?    <= ? @  8USQdkYkHre,g7h_ ,{N~ ,{ N~ ,{V~ ,{N~    BI ?`P   P*    BLM ?`   R*  H  0޽h ? ̙3380___PPT10. Vv &@(     6 f!    XN0vTbv (3   6 ?"H 2 ǏvvTbϑۏ N"sv:g6R /fWV4ls^ Nv ^\N|>ev c ba0 N vTbv{|W 1.[(induction)/fvOR㉕^irbNirO_uir~ހTbRN"_-N gsQvvǏ z0_uirǏ[\O(u Nuvvy:N[v:N^Yeg^irbvQ~g{|NOE.coli(W+TsN|vW{QW-NTb-JSsN|vTJSsN|nvI{0 [ir(inducer)^irb~g{|$$[$"&3 '  * < B  s *޽h ? f3f3ff pT(  f  0 ?"   s At ?http://gened.emc.maricopa.edu/bio/bio181/BIOBK/operon.gif/  S Ѝ   xStructure of a typical operon $"((B  s *޽h ? f3f3ffn    (     6Y3    ^ NN͑/g$$3   6] ?"@: $20[irinducer N;Oircorepressor   [ir  /fwYv[Tbvir( YsN|I{Nˆ}v~T b6RvQNd~WV v~TOۏlU_ۏL  ;Oir  /f;OvNuvir( Y(lWxT8hxI{ˆ}v~T OۏvQNd~WVv~Tb6RlU_ۏL  [N/f\RP[OSir( 8^;`y:NHe^ireffecor SNˆ}vv~TNOTSuSg\O(u v^ۏNekcؚbMNONd~WV v~TR0 30;Oirrepressor NT;Oirˆ}vaporepresseor   N/f1uWVxNuyr_'`ˆ}v(regulatory potein) /fN{|NORP[ϑSgˆ}v g$N*N~TMOp N*NNd~WV~T SNMOpSNHe^ir~TS_ˆ}vNHe^ir~TT 1\SuSg\O(u SgTNd~WVv~TRScؚb NM0 g;m'`  SNO~Te;m'`  NNO~T ;Oir: (Wl g[ireNd~WV~Tvˆ}v ;Oirˆ}v:SQ g;OirX[(WeMbNd~WV~T00Z0 Z $ $$$&3$$$!&3&$ $+$&$$$!$$I$&3$$$$$`  :P*   Y  B  s *޽h ? f3f3ff@ (  N  <R ?" RcklU_Ǐ zOVNˆ}vvX[(W0 lU_Ǐ z NOVNˆ}vvX[(W0T* 2&&3&&3 *B  s *޽h ? f3f3ff J(    S 3    &E.coli sN|d~P[f[( ,& 3 36B  s *޽h ? f3f3ff  O(  ^  c 6AlactoseY  6L ?"`  l6The enzymes that are required for lactose degradation:7 27  6 ?"0 p Ubeta-galactosidase: This enzyme hydrolyzes the bond between the two sugars, glucose and galactose. It is coded for by the gene LacZ. Lactose Permease: This enzyme spans the cell membrane and brings lactose into the cell from the outside environment. The membrane is otherwise essentially impermeable to lactose. It is coded for by the gene LacY. Thiogalactoside transacetylase: The function of this enzyme is not known. It is coded for by the gene LacA. 0ir0i0i0i!0iM0irM   G  IB  s *޽h ? f3f3fft $(    S  6    0vTbv[Enzyme Induction B& 33^  c 6ARegfig7    6; ?"@ 2Figure 7. The Lac operon and its control elements "3 21`ZB  s *޽h ? f3f3ff>     (  !  6 ?"C Lac R = Regulatory gene (also known as Lac I) gene that encodes for the lac Repressor protein that is concerned with regulating the synthesis of the structural genes in the operon. Lac I is adjacent to the Promoter site of the operon. An active repressor binds to a specific nucleotide sequence in the operator region and thereby blocks binding of RNAp to the promoter to initiate transcription. The lac repressor is inactivated by lactose, and is active in the absence of lactose. O = Operator specific nucleotide sequence on DNA to which an active Repressor binds. P = Promoter specific nucleotide sequence on DNA to which RNA polymerase binds to initiate transcription. (The promoter site of the lac operon is further divided into two regions, an upstream region called the CAP site, and a downstream region consisting of the RNAp interaction site. The CAP site is involved in catabolite repression of the lac operon.). If the Repressor protein binds to the operator, RNAp is prevented from binding with the promoter and initiating transcription. Under these conditions the enzymes concerned with lactose utilization are not synthesized. Lac Z, Y and A = Structural Genes in the lac operon. Lac Z encodes for Beta-galactosidase; Lac Y encodes the lactose permease; Lac A encodes a transacetylase whose function is not known. lac = lactose the inducer molecule. When lactose binds to the Repressor protein, the Repressor is inactivated; the operon is derepressed; the transcription of the genes for lactose utilization occurs. (23pHb0s0)x/ 4 p C#  S    ATable 3. Functional and regulatory components of the lac operon ,BA&33,5B  s *޽h ? f3f3ff ([(  (^ ( c 6A Regfig86#  ( 6Ą ?"  GFigure 8. Enzyme Induction. Induction (or derepression) of the lac operon is a form of negative control because the effect of the regulatory protein (Active repressor) is to stop transcription. " 2`Z>*  { ( 6ʄ̙?"_   gsN|esN|\O:NNyHe^irN;Oir~T ,OvQSg, NNO~T,N lU_ۏL,TbsN|v,RsN|0 esN|e~ހQNuv;Oir~T(Wd~WV N,lU_ NۏL*U 26$3&3B ( s *޽h ? f3f3ffs   0(  0$ 0 c $؎   2vTbv;OEnzyme Repression P&(3&(&(3(3 0 6 ?"<|u 1.~Nirv;O(end product repression): S;O) sS(WTbN"-N ~Nir;O_ @b gvvTb0:NWVhvc6R 0 Yr(lx(Try)Tbvc(ck).c  & 3Z$3r) 0 c 4Ao?r(lxTbPw,$D 0^ 0 c 6A Regfig5" 4 0 6ڄ ?",i& JFigure 5. Genetic organization of the Trp operon and its control elements &K 2I`Z,&B 0 s *޽h ? f3f3ff___PPT10+RD' b= @B D' = @BA?%,( < +O%,( < +D' b=%(Dm' =%(D' =4@BBBB%(E' =4 B`BPB`B?<*%(/%,( < +D' =1:Bvisible*o3>+B#style.visibility<*0%(D' =-m6Bbox(in)*<3<*0+  \ T <(  < < 6 ?"   FTable 2. The Trp operon and its control elements R = Regulatory gene that encodes for the trp Repressor protein that is concerned with regulating the synthesis of the 5 gene products. An active repressor binds to a specific nucleotide sequence in the operator region and thereby blocks binding of RNAp to the promoter to initiate transcription. O = Operator specific nucleotide sequence on DNA to which an active Repressor binds. P = Promoter specific nucleotide sequence on DNA to which RNA polymerase binds to initiate transcription. If the repressor protein binds to the operator, RNAp is prevented from binding with the promoter and initiating transcription. Therefore, none of the enzymes concerned with tryptophan biosynthesis are synthesized. A = Attenuator DNA sequence which lies between the operator and the structural genes for trp biosynthesis. The attenuator is a barrier that RNA polymerase must traverse if it is to transcribe the genes for tryptophan biosynthesis. In the presence of trp, most RNAp molecules fall off the DNA before transcribing the trp genes. In the absence of trp, RNAp is able to traverse the attenuator region to successfully transcribe the trp genes. Trp A, B, C, D, E = Trp Genes structural genes for enzymes involved in tryptophan biosynthesis. Trp = tryptophan end product of the tryptophan biosynthetic pathway. When combined with the repressor protein the Repressor is active. Trp is called a corepressor.  203333 3J3 353333D333 Dy yr "4J 0   Y B < s *޽h ? f3f3ff sk8 (  8 8 6X ?"`@ 'Figure 6. Repression of the trp operon "( 2&`Z$ 8 c 4Ao?r(lxTb ,$D 0^ 8 c 6ARegfig6)& 8 S    ` ~Nirv;O&& 3 B 8 s *޽h ? f3f3ff' 0E(  f  0 ?"     s Ab 3?E:\wuw-t-2\Other Regulated Systems.files\trp.gif[  S ZP   {The Tryptophan Operon , B  s *޽h ? f3f3ff0  >(  X  C      S  @   @ "  H  0޽h ? ̙33 0 m e p(  X  C    e  S ȍ @   Jacob and Monod were the first scientists to elucidate a transcriptionally regulated system. They worked on the lactose metabolism system in E. Coli. When the bacterium is in an environment that contains lactose: It should turn on the enzymes that are required for lactose degradation. These enzymes are: beta-galactosidase: This enzyme hydrolyzes the bond between the two sugars, glucose and galactose. It is coded for by the gene LacZ. Lactose Permease: This enzyme spans the cell membrane and brings lactose into the cell from the outside environment. The membrane is otherwise essentially impermeable to lactose. It is coded for by the gene LacY. Thiogalactoside transacetylase: The function of this enzyme is not known. It is coded for by the gene LacA. These three enzymes appear adjacent to each other on the E. Coli genome. They are preceded by a region which is responsible for the regulation of the lactose metabolic genes. Note that there is more to regulation than the obvious. It would seem that the cell would want to turn these genes on when there is lactose around and off when lactose is absent. But the story is more complicated than that. A bacterium's prime source of food is glucose, since it does not have to be modified to enter the repiratory pathway. So if both glucose and lactose are around, the bacterium wants to turn off lactose metabolism in favour of glucose metabolism. There are sites upstream of the Lac genes that respond to glucose concentration. Hr!MrM9 )& F   H> jkn!0pxH  0޽h ? ̙33 0 `\(  X  C      S e @   ^*Bacterial genes are organized in by gene systems known as operons. %The operon model of prokaryotic gene regulation was proposed by Fancois Jacob and Jacques Monod. Groups of genes coding for related proteins are arranged in units known as operons. An operon consists of an operator, promoter, regulator, and structural genes. The regulator gene codes for a repressor protein that binds to the operator, obstructing the promoter (thus, transcription) of the structural genes. The regulator does not have to be adjacent to other genes in the operon. If the repressor protein is removed, transcription may occur. Operons are either inducible or repressible according to the control mechanism. Seventy-five different operons controlling 250 structural genes have been identified for E. coli. 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