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( xH  0޽h ? ̙33y___PPT10Y+D='  = @B + 0 (  vF J  V  HA  w?? w  HAI 4 x??J x  c $  l    rN EMP_ $`$H  0޽h ? ̙33y___PPT10Y+D='  = @B + 0 @8@P (  P< P 0l) @Dl  10a|v u\O(u ( SyEmbden -Meyerhof -Parnas_ {yEMP_)NY%3Bf ( 0RW\ P C 4Aglycolysis0005 P <<+ F X;mS P <x0   PyMO P <5 pNq \ 'lS P 0H:   g TxxS 2 P 6d@ 0~ a|o;mve_ 2 " 0  P 0C  ` V ]|_gv 2  P 0H  t  xxg|ov 2% 0  P 0DK    \g|Nxx)v 2 RB  P@ s *Dp pRB  P s *Dpp P 0O    tul-3-xx1"lv 2 $ P 0TT   0  Zxxulxov 2 P 0X @ p  X ulxSMOv 2 P 0h\    8pv 2 P 0`  pt V Nnxov 2H P 0޽h ? ̙33y___PPT10Y+D='  = @B + 0 j(  T  s ,A >f?`0   s *Dl  l3     EMP_yrp. '6"    s *xp   ~> a|RP[~lSb1 6 Nxxg|T (W)vvPS N ˆb$N*N NxSTirRP[ sSxxNNnT3-xxul0 3-xxulۏNek'lSub2RP[Nnx 1RP[a|SMb2RP[3-xxul v^m2RP[ATP02RP[3-xxul'lSub2RP[Nnx 2RP[NADH2T4RP[ATP0~ <26#!#@!)!  %       H  0޽h ? ̙33y___PPT10Y+D='  = @B + 0  (  b  C :A"glycolysis0000000H  0޽h ? ̙33y___PPT10Y+D='  = @B +6 0 v(    0  P` $S^ek10ek S^{_6k N6k 2NADH+H+ C6 2C3 2Nnx 2ATP 4ATP 2ATP ;`S^_ C6H12O6+2NAD++2ADP+2Pi 2CH3COCOOH+2NADH+2H++2ATP+2H2O yrpW,gN"_ NHesNO cOYy-NN"ir\O:NTbN"Se g'leNTCAsޏc e'leNnxSvQۏNekN"NirYN؏SbTySuNir NSu]N g[RsQ|0-0n<Z$0n<Z20n<ZZ@0n7ZU0n<Z     Rd @&, 8 F 0`  @ ZB  s *D@@@ZB  s *Dp@@ZB  s *D @@ZB  s *D @ @ZB  s *DP @ @ZB   s *D @@TB   c $D@TB   c $DZB   s *DZB   s *Dfr  6Z0` fr  6Z TB  c $DTB  c $D ZB  s *D`  0`P`  0``  0ZB  s *D  64Û 0 ` 1.EMP_6' ' H  0޽h ? 3333 0  &(    c $ϛ l `   bEMP_sQ.ek b(  s *H  1. a|xxS!1.6Nxxg|() 2. 1.6Nxxg|!2RP[3-xxul 3. 3-xxul!Nnx ;`S^_ a|+2NAD+2Pi+2ADP !2Nnx+2NADH2+2ATP CoA !Nnx1"lv YNpCoA ۏeQTCA  lP 2C& &($,5$&&$&'$&$&   <0H  0޽h ? ̙33 0 bZ0(    c $ě l<R  H  Za|o;mve_ &$   0 7 R}Y'l_uirǏMg++TATPv]|ov S'l_uirǏxxp_Nnx-xxlyv|~ (Wa|ۏeQ~ހesS[bNxxSnB '>''>#'#>   0H    X xxg|ov 2&$ r  0H  p` rEMP_vsQ.v (Wuir-N gdkv1\asT@wX[(WEMP_ ATPTMg++ (W;m~ހQPSvS^/f NSvS^6:  '30'FH  0޽h ? ̙33 0 jbPT(  TL T C $A u029 T 0#H 0  fNnxvS  2  f T 0z0P@ 0 0H T 0޽h ? ̙33 0  8(  X  c 0A  28 u028  0z0P@y 0 0H  0޽h ? ̙332 0 ``r(  `T ` C ,Aslide14uX ` 0  ` <-H   j20(  ` 0z0P@ 0 0H ` 0޽h ? ̙33 0 F>pl(  l^ l c 6A   img020|CQ l 02H  P h3-xxul 2 l 07H   P x(3-xxul1"lv)" 2  l < @  0H   1. a|~ǏQek'lSS^Nu8hn|-5-xxTCO2 2. 8hn|-5-xxSu TR_gSbh_gS R+RNu8h|-5-xxT(gn|-5-xx 3. NTy b|xx(We'lSNv`Q NSuxg͑c Nu]|xxTN|xx`p0Z'/R'%~$H  0޽h ? ̙33y___PPT10Y+D=' = @B + 0 @(    c $lH  l`  H  fHMP_sQ.ek &(`.  0H "  D1. a|!6-xxa|x 2. 6-xxa|x!5-xx8hn|!5-xx(gn| ! 5-xx8h|!SN8hxub 3. 5-xx8hn|!6-xxg|+3-xxul (ۏeQEMP_  lP'''g''''& H  0޽h ? ̙33l  0     (    0S  v R 6k C6 2C3 N6k 4 ATP 2ATP 2C3 2 Nnx 2NADH2 C6H12O6+2NAD++2ADP+2Pi 2CH3COCOOH+2NADH2+2H++2ATP+2H2O b0Z50Z70Z)0Z0Z3.  3  =  !)!)!)!)!) !)!)!)!6F5A[RB  s *DPpRB  s *DRB  s *D@ RB  s *D P` RB  s *D P RB  s *D @RB   s *Dp RB  @ s *Dp pRB   s *D ^R   6ԔP p *    <8S >  lS" ,l  S   N[hQHMP_v;`S^( %3!H  0޽h ? ̙33 0 F>(  J  0H 9  6 a|-6-xx+12NADP++6H2O 5 a|-6-xx+12NADPH+12H++6CO2+Pi S  2        X+  s *lH  lU  S  n[hQHMP_v;`S^ dB  <Do`~`H  0޽h ? ̙33 0 (    c $@H  l  H  jHMP_v͑aIN f  0@S m',$ 0 d1.O^TbSe:N8hxT8hxvuirTbcO b|-xx _-NvdӅ|0of)Y^n|I{S(uNe(lWxTb0xWTb0SY|Tb0 0 2.N؏SRNu'YϑNADPH2 Neb:Nx0VI{ir(vTbcO؏SR SebSǏ|T8TNu'Yϑvϑ0 3.ޏcEMP_NEMP_(Wg|-1 6-NxxTul-3-xxYޏc SNBR b|OsQ|0( '<'' '/)' ')'#+ (H  0޽h ? ̙33  ___PPT10~ +=D* ' S = @B D ' = @BA?%,( < +O%,( < +D' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*E%(D' =-6B'blinds(horizontal)*<3<*ED' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*E%(D' =-6B'blinds(horizontal)*<3<*ED' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-6B'blinds(horizontal)*<3<*+8+0+0 + 0  (  r  Z,ZS ??7g r4.ib'Yxn)R(uV_-NX[(W3-7xv| OwQ g__uirv@b)R(u)R(uvxn1f:Nf:N^l0 5.\O:NV[CO2v-NN/fIQ{Q_uirTS{Q_uirV[CO2v͑-NN0 Ǐ_SNuYy͑vSuNir0Y8hx0r^(lWx0vTsNx_WsNxSu I{0 HMP_(W;`vϑN"-N`SN[kO NN~ހN";mR[vQ-NNirvϑvsQ0 '(''/''+''['~ 8&H  0޽h ? 333380___PPT10.YH 0 (  f  0mS d LSy2-n-3-1'l-6-xxa|xKDPG ˆ_0 1952t^(WPseudomonas saccharophila-NSs TegfX[(WNYy~̃-NipQl4'`̃-NR^^ 0 ED_S NOVNEMPTHMP_ USrX[(W /f\pe:ON[teEMP_v_uirvNyfN_ *gSsX[(WNvQ[uir-N0r &&R& 3&  $    c $xS  l  S  n N ED_$H  0޽h ? ̙33y___PPT10Y+D=' S = @B +K 0 rj  (  ^   0S     ATP ADP NADP+ NADPH2 a| 6-xx-a| 6-xx-ax ~~ov NEMP_ޏc ~~'lSv (NHMP_ޏc) EMP_ 3-xx-ul ~~14lv 2-n-3-1'l-6-xx-a|x EMP_ Nnx ~~)v g'leNTCAsޏc e'leۏL~̃Su d  8   M    f>~au] H$  ,I kRB  s *D00f  0"` 0LB @ c $D 0 RB  s *D00pRB @ s *D  RB  @ s *DpRB   s *D` P`X   00 `RB   s *Dp RB  @ s *D `   s * S  lt  S  `ED_"$B  s *޽h ? ̙33y___PPT10Y+D=' S = @B + 0 0 f(      HA   ?V?    c $tS  l@  S  6ED_H   0޽h ? ̙33y___PPT10Y+D=' S = @B +  0 @`(  T  s ,A!a 3?wg  c $S  lZD  S  `ED_"$H  0޽h ? ̙33 0 |tP (    c $S  l,`   S  XED_vyrp  0pS 'w,$ 0 t41.N_:a|~lS:N2-n-3-1'l-6-xxa|xT ~1'ln|x)vPS ˆbNnxT3-xxul 3-xxulQ~EMP_lSb:NNnx0 2. ED_vyr_v/fKDPG)v. ED_vyr_S^/fsQ.-NN"ir2-n-3-1'l-6-xxa|xKDPG ˆ:NNnxT3-xxul0 3.S^ek{US NHesNO~g/f1RP[a|Nu2RP[Nnx 1RP[ATP0 4.dk_SNEMP_0HMP_TTCA_svޏc SNvOSNn_uir[ϑ0؏SRT N T-NN"irv0}Y'leNTCA_svޏ S'leۏLYNSu.0Z%>%$ %$$ $ $ $/$$$$O$       (H  0޽h ? ̙33___PPT10+=DB' S = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*R%(D' =-6B'blinds(horizontal)*<3<*RD' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*Rh%(D' =-6B'blinds(horizontal)*<3<*RhD' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*h%(D' =-6B'blinds(horizontal)*<3<*hD' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-6B'blinds(horizontal)*<3<*D' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-6B'blinds(horizontal)*<3<*+8+0+0 +  0   p (    0$ V   F ATP C6H12O6 ADP KDPG ATP 2ATP NADH2 NADPH2 2Nnx 6ATP 2YN ( g'le~Ǐ|T8T) e'leۏL~̃YNSu h  -     F v R= RB  s *D RB  s *D RB @ s *D RB @ s *Dp P ` RB  s *D@@ @ RB   s *D@ ` RB   s *D RB   s *D`u ``r   s *"` PRB  @ s *D ` RB  s *DP 0 0 RB @ s *D ` RB  s *DP 0 RB  s *D P 0 RB @ s *D 0   s *DV  lt   V  jED_v;`S^$ $H  0޽h ? ̙33y___PPT10Y+D=' S = @B + 0 $$ (      B1V ?? ' lsQ.S^2-n-3-1'l-6-xxa|xvˆ PSvv6-xx14lv KDPG)v vsQvSuuN~̃R|Su OpN"sؚ NirlSsؚ ̃SOub\ N"oRNir\ Su)n^ؚ N_[gO'l0 :ppH5 fg̃~̃[YNSRNO> B $ $ $ $) $ ~B (4   60V  P  ^$    B0JV ? ?   l~ ATP g'le~|T8T 6ATP e'le ۏLSu 2YNfPZP      @ 4H   BQV ? ?0 g  42ATP NADH+H+ NADPH+H+ 2NnxL (  ( ( 4    6mV  ?9 )  ATP C6H12O6 KDPGbx d F  @    * N  @p     @p VT    #  @xB   HD?  xB   HD?  xB   HD?rB   BD?  rB   BD?@p @rB   BD?p @p .T  pP 0   #  PT  p   # 0 pP 0 rB   BD? p xB   HD? p pxB   HD? xB   HD? xB   HD? rB   BD? rB   BD? JN        xB   HD? xB   HD?P xB   HD?PrB   BD?@P@rB   BD?P@Pr   BH/) ?@PxB   HD?z !  < ?"  z "  < ?" @z #  < ?" 0  $  c $V  l<  V  hED_v;`S^~  &$f H   0޽h ? ̙33y___PPT10Y+D=' S = @B +H  0   8H (  8 8 6V  jd v&a| NagM_(W N T_uir-NvR^ , 8 ZܜV ??mm ̃ T EMP% HMP% ED% Ruk 88 12  N gGPNuk 66~81 19~34  ppr ̃ 97 3  NĞR 77 23  'YFg̃ 72 28  ܔ~GPUSހ̃  29 71 U|GPUSހ̃   100 gIFg̃ 74 26  'lSa|Fg̃  100  w{QNx̃   100 ЏRSuUSހ̃   100 ĞkQSt̃ 70 30  D'+('(((('('&H 8 0޽h ? 333380___PPT10.gV8SN 0 ,(  ,V , 0V Pt   1uhS (W_uir~ހ-N gv TeX[(WYag_egMa| gvS gNy0(WgNwQSOagN N b gYag_vgy_uirvzz~UOy_N" [SuNirq_T_'Y0V 2V& VH , 0޽h ? ̙33T 0 <(  < <  <V   lS" ,1l  V  ^V xxn_   L < 0V 7,$D 0 X[(WNgN~̃Yf2Ns̃^\TsNFg̃^\-NvNN~̃-N0 ۏLxxn_v_uir:\)v @bN[ NY\xx]|ˆ:N2*N Nx|0 xxnv_ g$Ny xx b|n_PK _ xx]|n_HK _ ( 2#P;,H < 0޽h ? ̙33y 0 )!44@(  @L @ 0ظV P D ` a| 6-P-a| 6-P-a|x 5 -P-8hn| 5 -P-(gn|1 21R  @ 0пV  P` b  l 3 -P-ul Nnx 2  @ 0V p` P &YNpxx YNpCoA YN 2&RB @ s *D RB @ s *D0 RB @ s *DP RB @ s *D `LB  @ c $D  RB  @@ s *D@ ` RB  @ s *D 0 ^B  @ 6DoP 00^B  @ 6Do @ ^B @ 6Do0  RB @ s *D `Xb @ 0 P p @ 0V    ?ATP  2 @ 0TV P p 7ADP 2 @ 0(V p0 FNAD+& 2  @ 0V  INADH+H+& 2 Xb @ 0  @ 0V  ACO2" 2 @ 0 H @ 0޽h ? !@ ̙33 0 @8 D(  D D  <+X   lS"   X  fxx b|n_vyrp: >  D 0d/X   b`$R1RP[a|SNu1RP[ATP vS_NEMP_vNJS; a$QNNNuI{ϑvsNx0YNTCO2&2 20 t H D 0޽h ? ̙33 0 6.0 H(  H H  <;X 3  lS"    X  Xxx]|n_ >  H 0\X 0  2a| 2a|-6-xx 6-xxg| 6-xx-g|o 2ov& H 0LX @P `  :4-xx-dӅ| YNpxx 2$ H 0KX   @  p2(gn|-5-xx 2 $" H 0AX    R2ul -3-xx 2YNpxx* 2*6 H 0XXX @ R2sNx 2 H 0[X   R2YNx 2XB  H 0Do 0LB  H c $D  XB  H@ 0Do XB  H 0Do RB  H s *DoPPXB H@ 0DoPP@XB H 0DoPP@XB H@ 0Do  XB H 0Do@ ^B H 6Do  RB H s *Do  XB H@ 0Do 0 XB H 0Do  ^B H 6Do ^B H 6Do XB H 0Do`  H 0PbX  FYNx 2  H 0HfX `  Xxx]|nv 2  H 0jX  P  bxx]|nv b$ 2  H 0nX    f HMP_ 2XR H 0 XR H 000 H 0sX MqqG @ TEMP 2  H 0xX p D  RYNxov 2 H H 0޽h ? ̙33 0 @LD(  L  L  <`oX 3  lS" ,}   X  bxx]|n_vyrp &(>  L 0lX K >`$ g$N*NxxnvSRS^ a$(Wl g'lS\O(uT1"l\O(uvSN N 2RP[a|R:N3RP[YNxT2RP[3-xx-ul 3-xx-ul(W1"lvvSN NlS:NsNxYNpxxubYNxvS^RNADPubATPvS^vvPT b$kRP[a|Nu2.5RP[vATP c$Y_uirYSgkFg̃ v_WsNxSusSǑSdke_0V 20Z!% $ !H L 0޽h ? ̙33~ 0 t~(  t t 6X  Z ^N Nx_s "( t 0XX  'gg J SyTCA_s0Krebs_sbgjx_s0(W~'YYpe_{Q_uirv|T8TN"-NwsQ.\O(u0vQ-N'YYpev(Ww8huir-NX[(WN~|SOW(-N (W~̃-NX[(WN~ހ(-NS g%tsx1"lv/f~TN~ހb~|SO N0 ;NNir 4NADH+4H+ 12ATP FADH2 2ATP GTP^ir4ls^ ATP 3CO2 (Wir(N"-Nv0WMOg~MOn ]NSuNirgjx0gx0^"}x0%tsxT7(lx O   b ^% +  F @  t S 7 zB t <D?"@0 `0 zB t <D?" 0 0 zB  t <D?"0 0 tB  t 6D?"  tB  t 6D?"P 0 0 zB  t <D?"  zB  t <D?" zB t <D?"  zB t <D?"  t 6X  ?" (   C3 CH3CO~CoAb (2$,$,$ F  p  t   z t < ?" P p zB t <D?"`zB t <D?" zB t <D?"  z t < ?" z t < ?"@0 z t < ?"` P  t BDX ? ?"% S#E  t|T8T $    t BX ? ?"  t|T8T $   H t 0޽h ? 333380___PPT10.iVP  0 `X\(  \x \ c $X  lp P  X  H \ S A$a ?p \ 0X <p   Nnx(WۏeQ Nx_sKNHQ1ubYNpCoA YNpCoATIpYNx)TbgjxQۏeQ Nx_s0 _sv~g/fYNpCoA{_^'lSbCO2TH2O k'lS1RP[vYNpCoASNu12RP[vATP IpYNxSNS^ ,gv^ Nm0l{ <L!)!)*!                  H \ 0޽h ? ̙33b 0  `(  `j ` BA28 3Fig_4_117mH ` 0޽h ? ̙33T 0 d(  d\ d c 4A m HcitC_OVH d 0޽h ? ̙33 0 ` .(  b  c :A  HcitC_Step2` 0P  0X Z7 zLN4YxvvS^yr_PS14l 6qTSR4l N 9eSNRP[QOHTHvMOn ub_gjx0 v~g~b+Tv^_SIUˆ}v g4*NSP[04*Ne:gkxSP[S4*NJS(lxkxSP[~Tvkx-N_| SN^irvS4lTR4lvS^0F.BV!<M  0z0`P@ 0 0H  0޽h ? ̙33  0 80p (  ~  s *X  l?  X  b  c :A P citC_Step3 0`  0z` @ 0 0H  0޽h ? ̙33  0 80 (  ~  s * ]  l:  ]  b  c :A28 HcitC_step4T4S  0zp@ 0 0H  0޽h ? ̙33  0 80 (  ~  s *]  lZ  ]  b  c :Am HcitC_Step5<  0z@ 0 0H  0޽h ? ̙33  0   (  ~  s *]  l?8  ]  b  c :Am HcitC_Step6iH  0޽h ? ̙33  0 80 (  ~  s *&]  l@  ]  b  c :AP HcitC_Step8m  0z`@ 0 0H  0޽h ? ̙33< 0 l|(  l l s .] 0e0e3  lP  ]  ^ l 6Aa H3DX H l 0޽h ? ̙33N   0    (     0(3]   lS" L  ]     <3]   l"<  <\ ]  X2  0  RB  s *D  RB  s *D0 PRB  s *D` PRB @ s *D RB   s *D` RB  @ s *D`  ` RB   s *D` ` ` Xr   0`` RB   s *D RB  s *D RB  s *D RB  s *D RB  s *D PRB  s *D@ RB  s *D0 P RB  s *DP  RB  s *D` P@ RB  s *D`  RB  s *D ` RB @ s *D ` Xr  0  RB  s *Dp `p   0z0P@ 0 0H  0޽h ? ̙33V 0 p(  p^ p c 6A, RFig_4_18H p 0޽h ? ̙33 0 :(  Z  c 2AP tca037  0z0P@ 0 0H  0޽h ? ̙33 0 P N(     <(S] K  lt  ]  bTCA_sv͑yrp 3(  0`b]  10_sN!kv~g/fYNpCoAvYNpW'lS:N2RP[CO2,v^͑eub1RP[IpYNx 20'l NvcSNS^ FO/f g'lagNte*N_s gVek'lS؏SS^ vQ-N NekS^-N\NAD+؏S:NNADH+H+ SNek:NFAD؏S 30:N|00ˆ}v( N'Yir(lS-N_g~0 40_s-NvgN-NNir/fNN͑ir(uirTbvMRSO 50uirSOcOϑv;Nb__ kRP[NnxN15ATP 60:NN{|)R(uuirSuuN@bNTcO;NvN"_0YgjxSuGluSuI{0 2%%-;%- %-%4 ( H  0޽h ? ̙33P&  0 %%/]P%(  x  c $x]]  lU  ]  $  ] #"*kk  H Zp] ??S   L 2 @` F Z8] ??) S   J 1 @` D Z0] ?? S )  P  @` B Z] ??wS   K 2 @` @ Z] ??S w  aNADH+H+(J @` ; Z] ?? V  P  @` 9 Z] ??) V  J 1 @` 7 Z<] ?? ) V  K 12 @` 5 Z4] ??w  V  P  @` 3 Z] ?? wV  bNADPH+H+( J @` ) Z] ??S  S 2 2 @` ' Zl] ??) S  N 1    @` % Z] ?? ) S  P  @` # Z] ??w S  P 2    @`| ! Z] ??wS  V ATP ^ir4ls^ GTP6,$ @`  Zk ??  M 36-38 @`  Z k ??)  J 7 @`  Zk ?? )  K 35 @`  Zk ??w   J 8 @`  Z(k ?? w l QNATP @`  Z1k ??V   L 2 @`   Z:k ??) V   P  @`   ZBk ?? V )  P  @`   ZKk ??wV   P  @`   ZTk ??V w  _FADH2(J @`   Z0Ok ?? OEMP+TCA @`  Ztfk ??)  HED @`  Z ok ?? )  IHMP @`  Zxxk ??w  e EMP @`  Zk ??w dNb__  @`B  Zo ??B  T1 ??B  T1 ??  B  Zo ??B  Zo ??B  T1 ??wwB  T1 ??  B  T1 ??) ) B  T1 ??B  Zo ??B " T1 ??S S B 4 T1 ??V V B A T1 ??  r ,  ` ??2WH  0޽h ? 333380___PPT10.YD  0 x(  x$ x  < Vk   lS" `g  k  zN0"l0S"lTATPvNu'(3 x 0؛k @@` : 2  x 0w] tVY  @ &~ N1"l_ubvNADH0NADPH0FADI{؏SWvǏ|T8TI{e_ۏL"l g~NS"lSO'l0e:gb g:g'lSir ~T Nʑ>evQSf[\o02T S&'D1H x 0޽h ? ̙33y___PPT10Y+D=' X = @B + 0  (  n  0ئk mT  \&9hnc"lyr+R/fS"lǏ z-N"lSSO'`(v N T,b_uirϑN"R:N|T8T\O(uTSu\O(u$N'Y{|. Su\O(ul gNUOYcvg~5uP[SSOvuir'lS!j_ |T8T\O(u gYcvg~5uP[SSOvuir'lS!j_ &|T8T\O(uSSR:N$N{| g'l|T8T  g~5uP[SSO/fRP['lO2; e'l|T8T  g~5uP[SSO/fO2NYve:g'lSir Y NO3-0SO42-I{..Wx* %!%!%!!%%%#%% %-%% %-%--%--% Y   H  0޽h ? 3333___PPT10i.Y+D=' X = @B +   0  2(    s 4k 0e0e  l#" 0eI  k  F P  o N P  PT P # P  6k ?  C6H12O6 -[H] A -[H] [H] B -[H] C A0BbC AH2 BH2bCH2 -[H] SuNirYN0 CO2 sNxI{ 1"l "l S"l|(2$$,$,$,$ $$ $ $,$,$,$ $,$4  hX .   6k  ?`v <  B~|T8T `$|T8T a$e'l |T8T b$SuH (2((!$!R   6l  ?pP  R1/2O2 H2O NO3- SO42- CO2 NO2- SO32- CH4v(2( ((             &N H   HrB   BD?rB   BD?``rB  BD?` `rB  BD?0 ``rB  BD?x`H`rB  BD?rB  BD?pprB  BD?<< T @P  # @P N P   P l  < ?xB  HD?ccpxB  HD?ccxB  HD?c0 c xB  HD?c cP DN 0   0 l2  < ?r2  BL ?0 ~B  ND?@ @ ~B  ND? @ @@ N @    @ xB ! HD?r2 " B ?p0`xB # HD?0`0 2 $  LCYENHgQ ? `T0TY`T0TY`T0TY`T0 @ PN 04  % 04 xB & HD?r2 ' B ?0 xB ( HD?0  x2 ) HL ?t 04 B O  `D??m  B   `D?? B   `D??-B   `D?? vb  N??S vb  N??PB   fD?? CB @ ZD??B   `D??0 B @ ZD??6 B @ ZD?? H  0޽h ?o`"$') 3333___PPT10i.Y:H+D=' k = @B +Q  0 `(    0l  uPH@___PPT9"  Fermentation -- oxidation of an organic compound in the absence of external electron acceptor (no oxygen required). Uses SLP (substrate-level phosphorylation) Respiration -- oxidation of an organic compound where oxygen is the final electron acceptor. Uses ETS (electron transport system) as well as SLP Anaerobic respiration (unique to bacteria) -- oxidation of organic compounds where an external substrate other than oxygen serves as final electron acceptor. Exs: nitrate, sulfate, carbon dioxide l 2 2  43$ @`  c $L2l  l   l  \&Mechanisms of energy release: overview''H  0޽h ? ̙33 0 t(    6;l   PXi_:^ir1"lT ~|T8TT O g~YnRP['l\O:Ng~5uP[(b"l)SSOv'lSǏ z/fgnfM0g͑vuir'lSe_0 _EMP,TCA_s yrp_{cQ (W g'l|T8T\O(u-N ^irv'lS\O(u NN'lv؏S\O(uvcvPT /f^ir(W'lSǏ z-Nʑ>ev5uP[HQǏ5uP[ O1uTy5uP[ OSO YNAD,FAD,vQTTy~ހr }~b gTMb O0R'l0v3x%V'!$$V $$Vb$N* JT   f$Kl jJ  lc"$ `ZL   l  z 10 g'l|T8T('(`"  "  0Fl 9  D& 1udkS TCA_sN5uP[ O/f g'l|T8T-N$N*N;NvNs0># 2&V"&H  0޽h ? ̙33y___PPT10Y+D=' l = @B +" 0 IA(    A+|  H?E:\bacteria course\Procaryotic Metabolism.files\aeromodel.jpg  c $Vl  l  l  QFigure. Aerobic respiration`ZH  0޽h ? ̙33y___PPT10Y+D=' l = @B + 0  (       <8_l V  lS" ,  l  X5uP[ ON'lS|T8T   J   08bl ]T, [IN1uN|R'lS؏SRHTh^]v0rcRv"l OSO0R/fb"lb5uP[NNO'lS؏SRvSTirY~ O0Rؚ'lS؏S5uRvRP['lbvQN g:gir0e:gir0(W"lb5uP[v OǏ z-N ǏN'lSxxSS^SuvPT 1\SNuATPb__vϑ0 MOS8huirSu(W~ހ N w8huirSu(W~|SOQ N bXT5uP[ O/fNNAD0RO2 5uP[ O-Nv5uP[ OSO;NSbFMN 0CoQ0~ހr }b 0c 10c0 a 0aTNNkxˆ}v0ُN5uP[ OSO O5uP[vz^ cgq[Nv'lS؏S5uR'Y\cR 5uP[ O!k^Y N0n$Kq$$K$$K $,$$,$,1$j-'H   0޽h ? ̙33y___PPT10Y+D=' l = @B + 0 t(  tZ t c 2Ae 28 08-07F:H t 0޽h ? 333380___PPT10.fe0f[NGW&^5uP[[ wQ gؚvlyR [cR5uP[N؏SWvzaW N vly0RRP['l0 5uP[ O4OADPxxSbATPhQǏ z EeSy:N'lS|T8T0H8i 0n"*"*"*"*H"*"*"*"*P"*"*"*"*"*"*"*' ( (U $  CXB $ 0DjJXB $ 0DjJZ: XB $ 0DjJ 8XB $ 0DjJ0 `0XB $ 0DjJ XB  $ 0DjJRB  $ s *Djj XB  $ 0D1AXB  $ 0DjJ9B  $  `D??j~ 3  $ Tl  ??3  =ATP 2 $ Tl  ??ZA =ATP 2B $  `D??jv $ Tl  ??`  =ATP 2B $  `D?? CXB $ 0DjJS cS H $ 0޽h ? ̙33y___PPT10Y+D=' l = @B + 0 80D(  D` D 0l p NAD+T g[vvN^ir NyQN*N(P[T$N*N5uP[ b:N؏S`NDAH+H+0 FADTFMNĞ }ˆ}v'lS؏Sv vW0 kxˆ}vFe-S  O5uP[v'lS؏S}SOW :NRP[-Nv+Tkxv-N_R0X[(WN|T8T-NQyv YTSO-N SN Nv5uP[ O0(WV.l0Nkxx؏S0N]xx؏S0IQT\O(u0RP["lvo;mTʑ>eNSpv'lS\O(u-N_N g\O(u0(W|T8Tv 2Fe+2S -N_k!kN ON*N5uP[0 lvQ n'`"l}SO0^lX[(WNw8huir~|SOQTipQl4'`~̃v~ހ NipQl3'`~̃TgNipQl4'`~̃R+T2uW0(W|T8T-N{|v+TϑkvQN~RY10~15 P vQ\O(u/f6eƖeg|T8TTyvTW@bQv"lT5uP[ v^\[N O~~ހr }|~0 ~ހr }|~MON|T8TTz R/f O5uP[0jd $$!$,$$$$ $$f$!$$!$$$$|$$$$hKD( D  <l   l"<  I$  l  `_uir-N͑v|T8T~R $3 H D 0޽h ? ̙33y___PPT10Y+D=' l = @B +J 0 ((  (R ( 0l  s  |T8TvR N/f O5uP[N/f\5uP[ OǏ z-Nʑ>evϑTbATP  ُ1\/f5uP[ OxxS\O(uby'lSxxS\O(u 00B 2' K:# $H ( 0޽h ? ̙33 0  ..,(  ,oF 039 , uN 039 , 039KN 039 , 039$ , 6Pl ?03@R  bEMP 2NADH2 YNpCoA 2NADH2 TCA 6NADH2 TCA 2FADH2(2( (2( (2( (2(  (  (  (  ( L   N   ,   N |  , |  , <n  ?  F 'lS` ؏S` ؏S` 'lS` 'lS` ؏S` ؏S` 'lS` 'lS` ؏S` 1"lv|(( ((( ( ((( ( ((D  D   , 6n  ?| @  NAD FADH2 H2OD(2x  , 6n  ?| |  NADH2 FAD 1/2O2 +2H+Z(2 N @   , ,@, JN  @   ,  @ rR  , BG) ?@ rR ,B BG) ? @ JN  @  , @x rR , BG) ?@ rR ,B BG) ? @ JN  @  , @P rR , BG) ?@ rR ,B BG) ? @ JN  @  ,  @( rR , BG) ?@ rR ,B BG) ? @ JN  @  , t @ rR , BG) ?@ rR ,B BG) ? @ JN  @  , @@ rR , BG) ?@ rR ,B BG) ? @ JN  @  , @  rR , BG) ?@ rR  ,B BG) ? @ N  pp  !,  pp rB ", BD?   rB #, BD?   p rB $, BD?p p rR %, BG) ?@p rB &, BD?  ', <0n ??p9 bNO4ls^ ؚ'lS؏SR ( $ N  ` 0  (,  ` 0 rB ), BD? `  rB *, BD?  0 rB +, BD?0` 0  ,, 69n  ? .TFP Fe-S Cyt.b Cyt.c Cyt.a Cyt.a3 'lSvL!(2 (" HX  -,  0 ;n  lI0   n  V xQWv|T8T&(3  ., ZHn ?? T- | ؚ4ls^ NO'lS؏SR0 %% H , 0޽h ? ̙33y___PPT10Y+D=' n = @B + 0 @48(  4X 4 c 0A P fig17  4 0z0P@ 0 0H 4 0޽h ? ffD3fy___PPT10Y+D=' n = @B + 0 G?H(  H H A,p  ?E:\bacteria course\Procaryotic Metabolism.files\nad.jpgj\ H c $TVn  lw7}  n  [%Figure. The Structure of NAD and NADP&&`ZH H 0޽h ? ̙33s 0 4s(  4r 4 c JA`$ >20040413044515406/ 4 64^n  K [%Figure. The Structure of FMN and FAD&&`ZH 4 0޽h ? 333380___PPT10.fb 0 6.8(  8 8 s >Ab pic0722s  0H 8 0޽h ? 3333___PPT10i.f0+D=' n = @B + 0 T(  TZ T c 2Ah 28 08-05}cH T 0޽h ? 333380___PPT10.fα9r 0 0<r(  < < s 8Ac " R08-089g  0 < T(in  ??-  > 2  < T|kn  ??vV \ kxˆ}v" 2', H < 0޽h ? 333380___PPT10.foO$ 0 `D$(  DZ D c 2Ad 28 08-10Z  D s :An H5-30-1&    0H D 0޽h ? 333380___PPT10.f7 0 P8h(  8 8  Byn   l"<  ]TI<<$ 0 n   ~ހr }{|/f+TFe5uP[ OSO0SP[YN_SIUv~g-N_ gb@~ }0~ހr }N@~ }\O:NW0 ~|SOv5uP[ O\+T g5y N Tv~ހr } y:N~ހr }b c 0c 0a a3 0~ހr }b c c1 a a3teT(WNwX[(W0 ~ހr }a a3N YTirb__X[(W y:N~ހr }'lSv0~ހr }a a3+T g$N*N_vܔSP[01u؏SWa3\5uP[vc O~RP['l0 5uP[NCOQ O0Rb c c1 Fe-S.l}v, a a3 0^#+#+#+ #+#+#+#+#+#+#+ #+# :  8 0z0P@ 0 0 8  fn  ??Zw0  X ~ހr }{| '( H 8 0޽h ? ̙33___PPT10r.+ {D' n = @B D' = @BA?%,( < +O%,( < +D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*86%(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*86D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*86D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*86%(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*86D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*86D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*8%(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*8D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*8D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*8%(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*8D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*8+8+0+80 + 0 0(  (      s 8A( @ 408-117yz  0H   0޽h ? 3333___PPT10i.^q+D=' n = @B + 0 PB(  P P  <n K  l*b  n  ` ~ހr }|~&$   P 0n 2 RNl-NcS5uP[ v^\ TI{pevv(P[c0R~|SOb~ހYvnm-N0 R{|~|SOv5uP[ O\+T g5y N Tv~ހr } cvQ8T6eIQ1T'l؏5uMOv]+RR:Ncyt.a cyt.a3 cyt.b cyt.cTcyt.oI{0~ހr }b c c1 a a3teT(WNwX[(W0Cyta a3N YTirb__X[(W y:N~ހr }'lSv0~ހr }a a3+T g$N*N_vܔSP[01u؏SWa3\5uP[vc O~RP['l0 5uP[NCoQ O0Rb c c1 Fe-S.l}v, a a3 0 ~g~bN@~ }:NW ǏvQ_SIURP[-N_SP[vN5uwvSS O5uP[0 cyt.a3sS~ހr }'lSv/fY_uirv+gz'lSv PS4*N5uP[؏S'lvS^ o;mRP['l00Z%0ZW0Z$K#$$K3$,$,$,$,$,$, $, $,$$K'$,)$F6 'H P 0޽h ? ̙33y___PPT10Y+D=' n = @B +* 0   \*(  \Z \ c 2Aj 28 08-15,d   \  `(n  ??Y y z YTSOIb6RBR|n    Z \ c 2Ak 28 08-16, }$  \  f|n  ??Y@ y b }A[ YTSOIIIvb6R .H \ 0޽h ? 333380___PPT10.f@ 0 @ `H(  `Z ` c 2Al P 08-17  `  fn  ??,x L | YTSOIVb6RBR0lSir    Z ` c 2Am W R08-18ZM  2  `  fn  ?? p [ }['lDdSvb6R\O(u    0 H ` 0޽h ? 3333___PPT10i.fJ+D=' n = @B +  0 d\`<(  <~ < s *xn  lWt  n   <  BPn   l"<  f  n  RB < s *D U RB < s *D {  < 0z0P@ 0 0H < 0޽h ? ̙33`  0    T(  T T c $L l$   tATPv~gTub %`"   T 0d D  2. ATPvube_: F 2& ''"  & T 0   _uirϑN";mR-N@bmSv;N/fATPؚRP[ b__vSf[. ATP/fuirSOQϑv}SObAmb__.S__uir_ϑT,/fHQ\_vϑlbcbATP.S_ϑe,ATPRP[ Nvؚ.4l,͑eʑ>eQϑ.8r 2;"6#"       T 0"  { pIQTxxS 'lSxxS 2 'ff    T 0- Cc ^ir4ls^xxS 5uP[ OxxSh <2''''  T 04 .  h 1. ~g: 2&  T Bd9 # e D[B   T B=  D[B H T 0޽h ? ̙33 0 OGX(  X X A-p a H?E:\bacteria course\Procaryotic Metabolism.files\atp.jpg_ X s *D l9M   ]Figure. The structure of ATP $ `Z$H X 0޽h ? ̙33 0 L(  LZ L c 2Af  08-04L49H L 0޽h ? 333380___PPT10.fL 0 \(  \T \ 0Pz IQTxxS:)R(uIQTbATPvS^. IQTxxS\O(u\IQlSbSf[,N(uNNN'lSxTb~ހir(.;N/fIQT_uir0 IQT_uir:{|0݄~̃0IQT~̃Sb+}r~̃0~r~̃TUṽI{ 0 ~̃vIQT\O(uNؚI{ iir N Tv/f d݄~̃wQ gS~ }0ۏL4lvˆۏLN'lvIQT\O(uY vQN~̃l gS~ } S g̃~ }bvQNIQTr } Sˆe:girYH20H2SI{ b{US g:gir ۏL NN'lvIQT\O(u0 <2' '',''q'/'/'L      H \ 0޽h ? ̙33 0 `|(  `D ` 0D_ 'lSxxS)R(uSTir'lSǏ z-Nʑ>evϑubATPvS^0 'lSxxSubATPve_ g$Ny ^ir4ls^xxS  N'l 5uP[ OxxS  'l0K <2'$ff'$f'$'$'$'$'$'$'$'$L     ! H ` 0޽h ? ̙33y___PPT10Y+D=' n = @B + 0  l(  lV l Zo??@w ^ir4ls^xxS/f(WgySTir'lSǏ z-NSubNy+Tؚxx.vSTir ُ*NSTirǏv^vv\O(ubؚ.xx9hly~ADP OvQubATP0 ^ir4ls^xxSُy{|Wv'lSxxSe_(WuirN"Ǐ z-N:NnfM0PS^ir4ls^xxSvvX[(WN~ހ(Q0\}' A' '-'T  2 -H l 0޽h ? 333380___PPT10.Yp4J  0 dj(  d d  `A.a ?"  ,$D 0 d c $| lO<$D 0   l^ir4ls^xxS>NO '  T d 0 Y,$D 0  1uN1cN*N4lRP[ 2NxxulxvNOo.lS:N2NxxpNnx-Nvؚp.0ُyؚޏcvxxSNl~ADP NuATPRP[0(W_uirN";mR-N ͑vؚxxSTird NNNir(Y ؏ g1 3NNxxulxTYNpxxI{00x 2w%~"H d 0޽h ? ̙33___PPT10r.+E#mBD' n = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*dx%(D' =-6B'blinds(horizontal)*<3<*dx+8+0+d0 + 0 hh(  hV h 0dmgE  i_:(W5uP[ OxxS-N Ǐ|T8T O5uP[ \'lSǏ z-Nʑ>evϑTADPvxxSvPTweg b_bATP0 |T8T-Nv5uP[ OSO;N1uTyWTv~b g͑v5uP[ OSO/fl(tsSvQ T~ހr }|~0(W N Ty{|v_uir-N~ހr }vbXT/f N Tv0 ^6P' ~' 'j  '" h c $ lgS   f5uP[ OxxS*'(6 H h 0޽h ? ̙33y___PPT10Y+D=' n = @B +j 0 j(  2  Z8??M'-  Ǐ|T8TubvATPpeϑ;N/f9hnc|T8TbXTvY\ N T |T8Tv~bV_uiry{| _ Yuk̃Sub3*NATP ~̃'Y~Sub1*NATPxxS\O(u/f(W5uP[ꁛOSOTg~SSOv OǏ z-NSuv0 N'lS%{Qir(NuvN[5uP[b"lSP[Tg~5uP[SSOlye -N~ǏN|R5uP[ OSO k*N5uP[ OSOgbN*N'lS؏S|~ ُN|R5uP[ OSO(W N Tuir-N gvQ]N[vcR!k^ gbNag5uP[ O V y:N|T8T0AmRv5uP[Ǐ|T8Teekʑ>eQϑubATP0.'v* H  0޽h ? 333380___PPT10.[PFd 0  @p(  p@ p 0` |T8T(W O"lb5uP[vǏ z-N ǏN'lSxxS\O(uvvPT Nuuirv(un  ATP0 vMR_Ypef[cSv/fSf[nf[0 ;N‰p(W'lSxxSǏ z-N Ǐ|T8Tv|v\O(u \^irRP[ Nv(P[NvQO OYO N bN(P[(W$NOR^v NGWa sSb_bN(P[h^]Sy(P[RR0pHh^I{ 0ُ*Nh^]1\/fNuATPvϑegn V:N[SǏATPvvS^ b(P[NvYOQV0RQO ~gNebmdN(P[h^] Te1\TbNATP00Z9&&&&&&t&e )$ p  <  l"<  R   \'lSxxSN:g6R &$ H p 0޽h ? ̙33 0  x(  xZ x c 2Ai m 08-09t.f x  fƻ ??2 j~|SOTϑlbc * ' H x 0޽h ? 333380___PPT10.fRQu 0 Pt^(  tX t c 0A P fig20 t 6ɻ `'lSxxSN(P[h^] 2 &$ H t 0޽h ? ̙33 0  P(  PZ P c 2Ag P 08-12wJ  P  fԻ ??Pj rATP YTSO~g * ' H P 0޽h ? 333380___PPT10.fh   0 `xJ(  x x  `A/" ?"8  x 0F  B N*NNADRP[ Ǐ|T8TۏL'lS SNNu3*NATPRP[0[R+R(W N*NMOn TNuN*NATP0YV4 1@b:y ,{N*NATP'Y~(Wv1TĞ }ˆ}vKN,{N*NATP'Y~(W~ހr }bTclKN,{ N*NATP'Y~(W~ހr }cTaKN0$xx#   x c $ l    ^5uP[ OxxS>NO ' H x 0޽h ? ̙331 0 XP(    0P׻  ni_Ne:g'lSir-Nv'l\O:Ng~5uP[T"l SSOv'lS\O(u0 NNS'lT|Q'`S'l_uir(We'lagN NۏLe'l|T8T. e:g'lSirYNO3-0 NO2-0SO42-0S2O32-I{0 (We'l|T8TǏ z-N 5uP[OSOTSSOKN_N~ހr }I{-N5uP[SO v^4O gxxS\O(u ^irS{_^'lS SNuYϑ FO NY g'l|T8TNuvϑY0YN]xx:N5uP[SSOۏLe'l|T8Te Sʑ>eQ1796.14KJ1u0hv3x%!ai%!IIIIIIIIta A  J@  o     c $ l=   n 20e'l|T8T%`"  H  0޽h ? ̙33y___PPT10Y+D=' = @B +  0 0 D(  ~  s * l2      0 lS"   RB  s *D E RB  s *D  RB  s *D q Xr   0 ] Xr   0% :Z RB   s *D e RB   s *D ~ RB  s *D ~ H  0޽h ? ̙33y___PPT10Y+D=' = @B + 0 B:`d(  d d s @Ao 184244700#  0 d  f8 ?? le'l|T8Tv5uP[SSO * '  H d 0޽h ? 333380___PPT10.f;v 0 >6h(  h h FʼnApO 185038105Zd N  0H h 0޽h ? 333380___PPT10.f@{i^ 0  (  f  >A3a H1?'H  0޽h ? ̙33 0 P(  .  0?@[,$ 0 @&i_(Wuir'lS-NSu/fce'lagN N ^ir1"lT@bNuv؏SR N~Ǐ|T8T O vcN~NQn'lS'`-NN"NirvN{|NOHeNS^0 (WSu]N N Su/fcNUO)R(uS'lb}Y'l_uireguN g(uN"NirvN{|uNe_0 &Su_a|(WS'lagN NRa|vN_;N gEMP0HMP0EDTPK_0D v0 v- v&&"&B"&f#"&&'&L  c $M lp   v 30Su\O(u$!6"  H  0޽h ? ̙33  ___PPT10~ +3D* ' = @B D ' = @BA?%,( < +O%,( < +D' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*D%(D' =-6B'blinds(horizontal)*<3<*DD' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*Dt%(D' =-6B'blinds(horizontal)*<3<*DtD' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<*t%(D' =-6B'blinds(horizontal)*<3<*t+8+0+0 +P 0 pP(  pb p c :Aq pic_86000  p TX ??C > 2  p T[??m ` Su( 2!! H p 0޽h ? 333380___PPT10.fЇWM+  0 0 +(    Ta ?? X^ir H 2B   `D??Cw C  Tg ?? T^ir 2  T??"`"3   T k ??0  TADP+Pi 2   Te ??  7ATP 2   Tq ??} d jeO2. 2K   N??"` 6    T\v ??c   R -NN"ir 2     Tz ??c   $-NN"ir H (SuNir) 2$H  0޽h ? 333380___PPT10.Wa 0 vn@(  L  ZT??G8 $&SuQm(W N_-NGW g؏SW"lOSO  NADH+H+TNADPH+H+Nu FONuvϑv^ NY Y NSeO[N'lSQu |vR㉧N\O-Ne ُ7h_uir1\Na|RǏ z-Nb_bvTy-NNir:N"l5uP[ SSOegcSNADH+H+TNADPH+H+v"l5uP[ N/fNuNTyT7hvSuNir0(%%'o%'oF%'o%'o%R Fz  Z̓??pG 8^vSuy{| 1.1uEMP_-NvNnxQSvSu YNSu TWsNxSu NxSu 2,3-NNSu mTxSu NxWSu 2.ǏHMP_vSu _WsNxSu 3.ǏED_ۏLvSu ~̃vR|Su_WR|Su 0 %% )H  0޽h ? 333380___PPT10.Y`k 0 #(    0j +Fermentations: A. In the absence of oxygen, NADH is not usually oxidized by the electron transport chain because no external electron acceptor is available B. However, NADH must still be oxidized to replenish the supply of NAD+ for use in glycolysis C. Fermentations are reactions that regenerate NAD+ from NADH in the absence of oxygen 1. Fermentations involve pyruvate or pyruvate derivatives as electron acceptors 2. Fermentations may or may not produce additional ATP for the cell D. Alcoholic fermentations produce ethanol and CO2 E. Lactic acid fermentations produce lactic acid (lactate) 1. Homolactic fermenters reduce almost all pyruvate to lactate 2. Heterolactic fermenters form substantial amounts of products other than lactate F. Formic acid fermentation produces either mixed acids or butanediol ,0F2$ V" * I" * " * "  q    t H  0޽h ? ̙33E 0 ld(  T  s ,A) ?n>  0 {  \`$ukWR|Su a$ TWsNxSu b$NxSu c$mTxSu d$2,3 NNSu e$NxSu4/ <2.* "   f8??Z $1.1uEMP_-NvNnxQSvSu ' $ H  0޽h ? ̙33y___PPT10Y+D=' = @B +  0 ` >(     0 lS"   RB  s *D {RB  s *D   0z0P@ 0 0   6 ,< RN YNSu( B    `D??  B    `D??}}B   `D??} }B   `D??V  V H  0޽h ? ̙33y___PPT10Y+D=' = @B +8 0 _ W   (  g  040Pf  3 C6H12O6 2CH3COCOOH 2CH3CHO 2CH3CH2OH4 2 XB  0DjJP XB  0DjJ XB  0DjJ Xb  0 p   0t   ;NAD 2  0p @ WNADH26 2Xb   0 p  RB   s *DjJP RB   s *DjJ RB   s *DjJ  RB   s *DjJp pRB  s *DjJp RB  s *DjJ    0LP $ G-2CO2& 2   0$P  ;EMP 2RB @ s *D@P @  0P 82ATP 2  0X   T YN1"lv 2    0, lS" ,   j`$uk̃vYNSu( & 6   0t p i_ ̃y _ yrp SuagN" 2   0  <; YNSuǏ zS(WpH3.5~4.5NSS'lvagN NSu0 2$  H  0޽h ? ̙33y___PPT10Y+D=' = @B +  0    v (    0\q p&S_SumY(Wx'`agN N ukvYNSuO9e:NulSu0 SVagN NNuvYN N\O:Nck8^S"lSO ~g2RP[YNSugkSS^ ub1RP[YNT1RP[YNxlQ 2%f!%f!%1!@5  0|$T4  2CH3CHO+H2O+NAD+ CH3COOH+NADH+H+ CH3CHO+NADH+H+ CH3CH2OH+ NAD+ dke_N1uxxNNnbNS"lSOcS3-xxul1 Nv"l ub a-xxul T~a-xxulovPS ubul0 2   ! H_ LB  c $DpM pLB  c $Dz LB  c $DC sLB  c $D6 " f LB   c $D6  6 LB   c $D  LB   c $D[ LB   c $D m D  0T=Ii J2a| 2ul+YN+YNx+2CO2&& 2$ (PXB  0DjJH  0޽h ? ̙33y___PPT10Y+D=' = @B + 0 0 F(    0Ks ~v0___PPT106___PPT9 ^i_ g'lagN N Su\O(uSb6Rvsab'l[Suvb6Rsa 0 aINTt)R(un :gt%B0 2&&&. 0./  0PM3 m XΘ[ukN"vq_T 2  h q@   #"=/@q   <(T?a_  pؚ Y _1_m1Y    @`  <8e?_a  tNOcя \ ev    @`   <o?q_  (R|ubϑ |ϑ/USMOe ~ހvA~k   @`  <x?a@_ j :'lSu   @`   <tz?@a_ nΘ g'l|T8T     @`   <?q@_ V  @``B   0o ?q@@ZB   s *1 ?q__`B   0o ?q  `B  0o ?q@q ZB  s *1 ?@ ZB  s *1 ?a@a ZB  s *1 ?@ D   <`  lS" w'p   4]e_He^(The Pasteur effect )0("   <}     0pMm Jsa 2 H  0޽h ? ̙33y___PPT10Y+D=' = @B +< 0 P d(    0   ` a| 6-xx-a| 6-xx-g| 1,6-Nxxg| xxp_Nnx Nnx YNpCoAU 2 Lv'd2  <jJ p 8  0@ @` Jgjx 2 B  0DjJ  ,$D 0RB  s *D ` @RB  s *D` p   0DI `P,$D 0 JIpYNx 2 B   s *Dpp0,$D 0RB   s *D ppRB   s *DpppRB   s *DppPRB   s *D  RB  s *D@pp  0$pp0 hxxg|ov(PFK) 2   0X `  T Nnxov 2   0H ` R]|ov 2f   0@@ & -------ATP0gjx> 2 ,   0 $ % --------ADP,AMPP 2  J  0`  0   6-xx-a|   xxp_NnxP 2  f  fF   0 %   1,6-N-xx-g|, 2@   <  lS" w    \]e_He^~  ( H  0޽h ? ̙33 0 f^` ""(     <  lS" ,    Ta$~̃vYNSu     0 P  La| 2    BL` Z $2-n-3-1'l-6-xx-a|x"H  00P B3-xxul Nnx" 2"    00 0 P  LNnx 2    08 0   FYN YN$ 2$  $  6DfjJ , T2YN 2 ^B   6DjJRB   s *DjJP@XB  @ 0DjJ@ `XB   0DjJ@XB   0DjJ  ` XB  0DjJ XB  0DjJP RB  s *DjJ XB  0DjJ RB  s *DjJ   6fjJ   B2CO2" 2XB @ 0DjJRB  s *D   0 p 42H 2RB  s *DjJXB @ 0DjJLB  c $DXB @ 0DjJ   0,@ 42H 2RB @ s *DjJRB  s *DjJXB  0DjJ   0(p 6+ATP 2   0D  62ATP 2XB !@ 0DjJp0 p  " 0]q  "̃yЏRSuUSހ̃I{ _EDT 2 $ $ $ H  0޽h ? ̙33  0   p  (    0T(_ (&uk̃(WpH3.5-4.5e vYNSu ~1v ~1"lv Nnx YN YN ǏEMP_NuYN ;`S^_:N C6H12O6+2ADP+2Pi 2C2H5OH+2CO2+2ATP &~̃(Zymomonas mobilis)vYNSu ǏED_NuYN ;`S^Y N a|+ADP+Pi 2YN+2CO2+ATP &~̃(Leuconostoc mesenteroides)vYNSu ǏHMP_NuYN0sNxI{ ;`S^Y N a|+ADP+Pi sNx+YN+CO2+ATP TWYNSuNir-NN gYNNy g:girRP[vR|Su _WYNSud;NNirYNY ؏X[(W gvQ[ g:girRP[vSu40vZ0vZ0vZA0vZ$0ZC0vZ90Z$*$#&;$%-%-%-%-%-%-%$$"$$2$,$$$$ $$/$$$,$$$$$P ?  (       9XB  0DjJzzXB  0DjJz0 ` zXB  0DjJmZ mXB  0DjJpj pXB  0DjJs =M s H  0޽h ? ̙33y___PPT10Y+D=' = @B +u 0  ,(    s *^ l9'   $)R(uZ.mobilisI{~̃uNR|#    0d`}\ FOpN"sؚNirlSsؚ̃SOub\N"oRNir\Su)n^ؚ0$ 2'!# $  0e /O  &:ppH5fg̃YNRukNO( 2H  0޽h ? ̙33y___PPT10Y+D=' = @B +b 0   (     0p  lZj   \N sNxSu    0 u : 2   0w7 &sNx~̃)R(ua|SvQNv^vSSuv|NusNx y:NsNxSu0 1uÑy N T N"_ N T ubvNir g@b N T \sNxSuSR:N TWsNxSu0_WsNxSuTSgkFg̃Su0 TWsNxSu~EMP_ _WsNxSu~HMP_ SgkFg̃Su: ~HK_ xx]|nv_  <2X%% %% %%%W             H  0޽h ? ̙33 0  T(    0    La| 2    0P V  \3-xxul 2   0p p  R xxNNn 2    0  2( 1,3-N-xxulx  2 $0  00@ `2sNx 2Nnx(1 2$V. $+   <  lS" A   R`$ TWsNxSu  RB  s *DjJP P XB   0DjJ0 P XB   0DjJP @ XB   0DjJ@0 @   0$  \2NAD+ 2NADH2 2  X   0 ` @XB @ 0DjJ p XB  0DjJX  0  0 Xb  0  0@ 84ATP 2  0į`4 84ADP 2  0 P  : 2   6@0  < 2ATP 2ADP X  0 `P XB  0DjJ` ppp p  0 ) nLactococcus lactissNxsNt̃ Lactobacillus plantarum iirsNFg̃<8 2B    0V Y "i_ ̃y _ EMP yrp" 2 H  0޽h ? ̙33 0 ##D(  8   <4  lS" <d   &a$ǏHMP_vSu-_WsNxSu.  0dP p  Ja| 2   0|   Z6-xxa| 2  0   \6-xxa|x 2   0t p p  Z5-xx(gn| 2  0$  Z3-xxul 2  0` @  LsNx 2V    0 `   LYNpxx 2    0`  Y NAD+ NADH2 2    04`  Y NAD+ NADH2 2 XB   0DjJ p XB   0DjJ P XB  0DjJ  XB  0DjJ ` XB  0DjJ@ 0 XB  0DjJ pp X  0   X  0 P p  X  0 p   0`  <ATP ADP 2 X  0 X  0P p   00`P LYN YN YNpCoA$' 2V%#XB  0DjJP0 X  0 0@  0H@p A 2ADP 2ATP 2  0 ` 4  7-2H 2RB  s *DjJ ` RB  s *DjJ  RB  s *D     0  fi_ ̃y _ yrp" 2   ! 0@   F-CO2& 2 XB "@ 0D1` XB #@ 0DjJ@H  0޽h ? ̙33z  0 *"';(    6 <z  P b$SgkSu    0] R ~ a| 6-P-g| 6-P-g| dӅ|-4-P YNp-PX^ 2 H  0*` Z >3 -P-ul of)Y^n|-3 -P2 20  Z/ ?? (  (gn|- 5  P 8h|-5  P 8hn|-5  P (gn|- 5  P`n  H%%B @  `D??# MB   `D??MXb  0)B  ZD??  0<j)I> 82ADP 2  0 @ZV. @2ATP  2B @  `D??&cB   `D?? B   ZD?? B ! ZD??@B " ZD?? }B # ZD?? }B $ ZD??} B %@  `D?? `B &  `D?? `B ' ZD??C B ( ZD??C B ) ZD?? S B *@  `D??S B +  `D??S  B , ZD??c B - ZD?? C B . ZD??V C B / ZD?? CC B 0@  `D?? C B 2  `D?? Cc 4 0dL  42ul-3  P 2YNp-P4 2,B 5  `D?? B 6  `D??  7 0Q0  >2sNx 2YNx& 2$B 8  `D?? 9  fHY ??~g] JYNx 2 B : ZD?? ; 0\"TS  .i_ ̃y SgkFg̃ _ HMP yrp " 2H  0޽h ? ̙33C 0 56[(  L  0$dP D ` a| 6-P-a| 6-P-a|x 5 -P-8hn| 5 -P-(gn|1 21R   0i P` b  l 3 -P-ul Nnx 2   0op` P &YNpxx YNpCoA YN 2&RB  s *D RB  s *D0 RB  s *DP RB  s *D `LB   c $D  RB  @ s *D@ ` RB   s *D 0 ^B   6DoP 00^B   6Do @ ^B  6Do0  RB  s *D `Xb  0 P p  0|w   ?ATP  2  0|P p 7ADP 2  0p0 FNAD+& 2   0؄ INADH+H+& 2 Xb  0   0\ ACO2" 2  0P& FsNx 2   0P0 0V FYN 2 LB @ c $D RB  s *D^B  6DoP `   <P P ` P _gS\O(u ^b  6HKP   08 NNAD+. 2   0@ QNADH+H+. 2    0 P 0 Vxx b|nv 23 ^b ! 6G0*H`I P   " 0tP   QCoA 2 # 0`   6Pi 2Xb $ 0    % 0̱  @  ;2ADP+Pi 2 & 0@   82ATP 2 ' 0l p  7-2H 2RB ( s *Df`@@RB )@ s *Df @RB *@ s *Df @ RB + s *Df@@RB , s *Df`P@` - 0 @  7-2H 2 . 0p  7-2H 2Xr / 0   0 0 @@  FNAD+& 2  1 00 @p  INADH+H+& 2 LB 2 c $Df  RB 3 s *D 0  4  <3  lS" Mp   Xxx b|n_ >  6 6 8 `_WsNxSu_ 3H  0޽h ? ! ̙33_ 0  "*(    64 <g l TWsNxSuN_WsNxSuvk p " * #""7 y3"\  <?~ " F Lactobacillus brevis wsNFg̃ <$, @`  <<? ~ " U 2ATP @`  <?[ " 1sNx 1YNx 1CO2  , @`  <? [" R HMP @`   < ? 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TRvm0  ,$D 0B  s *޽h ? 33̙3 0 @8(    BA< jJ?)YQ(lxTbp,$D  0B  s *޽h ? f3f3ff.%  0 $$TT\$(  x  c $Ц< lP  <   c $< lP < * <RB  s *D~N\F @ PP   P@ P ZB  s *Ds`B  0D@  lb  <GP ZB   s *D6  f2   6 P`H F 0,   N    z   <1"`z2  <1"`( `B  0D1 `B  0D1  `B  0D1 TN p  pz  <1"`@pz  <1"` z2  <1"`  ZB  s *D10@0ZB  s *D1  `B  0D1  `B  0D1 ZB  s *D1 z2  <L1"`0tN 0,  0,`B  0D1  p`B  0D1  p`B  0D1  pt   61"`p0 `B ! 0D1 ` `B "B 0D1phN `   # `  `B $ 0D1` 8 8z2 % <1"`,fB & 6D1000 `B ' 0D100 0`B ( 0D100zb ) <1"`0 `B * 0D1@  hN  ,H  +  ,H `B , 0D1  `B - 0D1P P z2 . <1"` `B / 0D1`B 0 0D1@0 H zb 1 <1"`,0 fB 2B 6D1p  0N 0Pp 3 0Ppz2 4 <1"`0Pz2 5 <1"`H8`B 6 0D10zR 7 <1"`0 `B 8 0D1 zR 9 <1"` 0zR : <1"`` P`B ; 0D1 00`B < 0D1P`B = 0D1P` @` `B > 0D1 `fB ? 6D1pfB @ 6D1PfB A 6D10` ` z B <1"` z2 C <1"`h0 `B D 0D1  `B E 0D1 `B F 0D1 `B G 0D1 zR H <1"`@ zR I <1"`l P<zR J <1"`84h`B K 0D1P`B L 0D1P<<`B M 0D1 fB N 6D1@` ` fB O 6D1p@ @fB P 6D10 @ @`B Q 0D1 @@z2 R <1"` |z2 S <1"`P @ B T BD1?"pB  s *޽h ? 33̙3}  0 (  D  c $< l  < P2 OS TSb6R  concerted feedback inhibitionB) '$'#" d  6<?"qJ 2[INR/eN"_-NQ*N+gzNir TeǏϑeMbb6RqQ T_-Nv,{N*NvvNySe_0 >NO7(lxhFg̃Corynebacterium glutamicum Y|b[Fg̃Bacillus polymyxa )YQ(lxe(lWxTb-N)YQ(lxovSV(lxTς(lxvOS TSb6RT;O06 $$d8  +  >A=m Ho?OS TSb6RB ,$D 0B  s *޽h ? f3f3ff1)___PPT10 +:SD' <= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D/' =4@BBB B%(E' =4 B`BPB`B?<*%(/%,( < +D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-6B+checkerboard(across)*<3<*+!  0 !|!LL (    c $x< lP < * <x  c $<< l,<  < F LP  LPZB  s *D @@N LP  LPZB  s *DP<<ZB  s *D t2   6"` |6N LP   LPt2   6"`,ZB   s *D0 `0 ZB   s *D 00ZB  s *DPt  6"` t2  6"`h0 ZB  s *D  ZB  s *D ZB  s *D ZB  s *D tR  6"`@ tR  6"`l P<tR  6"`84hZB  s *DP`B  0D@` ` `B  0Dp@ @`B  0D0 @ @t2  6"`P @ ZB B s *D 0 TB  c $D nN LP  LPt   6"`t2 ! 6"`( ZB " s *D ZB # s *D  ZB $ s *D N p % pt & 6"`@pt ' 6"` t2 ( 6"`  TB ) c $D0@0TB * c $D  ZB + s *D  ZB , s *D TB - c $D t2 . 6L"`0ZB / s *D  pn 0 0"`p0 ZB 1 s *D ` ZB 2B s *DpZB 3 s *D` 8 8`B 4 0D000 ZB 5 s *D00 0ZB 6 s *D00tb 7 6"`00 ZB 8 s *D  t2 9 6"` `ZB : s *D0tb ; 6"``0 `B <B 0Dp  0t2 = 6"`H8tR > 6"` 0tR ? 6"`` PZB @ s *DP` @` ZB A s *D ``B B 0DP`B C 0D0` ` t2 D 6"` L~N  P E  PzR F <o"` P2 G CTENGJTQo `Tr/`TT`Tr/`TTT/`TTT"`>ZB H s *Do `B I 0Do0   ZB J s *Dop``2 K CTENGJTQo `Tr/`TT`Tr/`TTT/`TTT"`tVZB L s *Do  B  s *޽h ??0.GK 33̙3 0 L(  4  c $| E l  E T3 T\OSb6R  cooperative feedback inhibition.+ &$3 &3""L  6E?" [IN$Ny+gzNir TeX[(We qQ TvSb6R\O(u'YNNUSr\O(uKNT0 >NO(W VdT8hxTb-N xx8h|&qxxvSAMPTGMP TIMP vT\OSb6R NqQ TX[(We SN[hQb6Rvv;m'`0 NUSrǏϑe R+Rb6RvQ;m'`v70%T10%0L~ $!$$W$n:-  HA>,   ??m   B  s *޽h ? f3f3ff $ 0 (  >  c $h"E lq  E R4 y/}Sb6R  cumulative feedback inhibition:* & 333&333"!p  6(E?"/1 [INkNR/e_+gzNir cN[~vRkUSrb6RqQ T_-NMRbvv @bNS_Qy+gzNirqQ TX[(We[Nvb6R\O(u/fy/}v T+gzNirKNeeOS THe^ Nebb\O(u0&Q P$P  >A?, o?/}ySb6R/Z ,$D 0B  s *޽h ? f3f3ff-%___PPT10+B8UD' E= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D+' =4@BBB B%(E' =4 B`BPB`B?<*%(/%,( < +D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-6B'blinds(horizontal)*<3<*+ 0 0((  Z  66E?"0  xTry 16% CTP 14% (l2upxx 13% AMP 41% & & (= 2<&"  FA@P Ro?7(lxTbvS/,$D  0  c $9E l ):  Ev0___PPT106___PPT9 0y/}Sb6R  E.coli7(lpTbvv'& B  s *޽h ? f3f3ffP% 0  (    c .AA o?z^S@ZE,$D  0<  c $DE l,/  E V5 z^Sb6R  sequential feedback inhibition4, #$ "$"#  6JE?"mq  Ny~Nirvy/},[MRN-NNirvy/} ǏTSb6RTb_sQ.vv;m'` OTb~bk0 >NOgIb[Fg̃e(lWxTbv67( K$ $:B  s *޽h ? f3f3ffV   0  @T(  x  c $UE l p  E   c $@_E lp E    F P5  ZB  s *D``ZB  s *D``ZB  s *D`` ZB  s *D` `` fB   6D>PP 0ZB   s *D0 `2   0 @ `2  B 0`  `B  B 0D>0xR  H+HG0*>ZB  s *D` @ l2  <> 0 @ N % 5  % 5rR  BPsG0*> % 52 B CRENG\JRQ> r`T2`TRr`T2`TRR`TRRB  s *޽h ?O@   33̙3   0   PT (  ~  s *$gE l  E    00qE lS" z  E *JGR X2  0jJRB  s *D32d2LB  c $D2 2LB  c $D2z * 2LB  c $D2U2LB   c $D LB   c $DF  LB   c $D6  LB   c $D  LB   c $D LB  c $D LB  c $D #  LB  c $D  LB  c $D  LB  c $D  LB  c $D  LB  c $D ; XB  0D>2  XB  0Df> CRB  s *Df>,u,RB  s *Df> >RB  s *Df>SS RB  s *Df> S B  s *޽h ? 33̙3& 0 h``(    <tE?""/  p (WR/eTb_-N R/epTv$NyvzN TNy^ir YAMPNGMP ThrNLys0Met 1uN$Nyv[^irvKm$ ~v0___PPT106___PPT9 Sgvt Sgv:NNySgˆ}v vRP[zzgavSS q_Tv;m0vQ NwQ g$N*NN NzSONN'` N TvcSMO N*N/f;m'`-N_ SN*N/f-N_0 K 2K'J  6(E ?"  ] ~v0___PPT106___PPT9 ";m'`MOp:N^ir~T SgMOp:J('f''f$ ,  6E ?"  8___PPT10N___PPT90( TNb6RBR~T,gaSS, NN^ir~T No;mBR~T, gaSS,OۏN^ir~T @+%%%Zr  0ԷEf3" ` ,$D 0 >3f B  s *޽h ? f3f3ff___PPT10h+E4D' E= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(DV' =A@BBB*B0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-6B)barn(outHorizontal)*<3<*+8+0+0 + ( 0 <(  x  c $lE l   E   HAB >  ?3f?0y   B  s *޽h ? 33̙3y___PPT10Y+D=' E= @B +H) 0 (     <`E!  ldD  E ZN0vTbv &(3 p  6E?"H 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&, 0 0l(  f  0 ?"   ACt P ?http://gened.emc.maricopa.edu/bio/bio181/BIOBK/operon.gif/  c $AK l,7W  K ~Structure of a typical operon "$0B  s *޽h ? f3f3ff  0 H @ P (       <JK3  l $  K b NN͑/g(&$3$    6*  { , 6K?"_   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>[ 2<$ & & B , s *޽h ? f3f3ff / 0 0,(  0 0 c $P lwqW  Kv0___PPT106___PPT9 2vTbv;OEnzyme Repression X'(3'$'(&03 0 6TP?"<|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$3j) 0 c 4AGo?r(lxTbwV,$D  0j 0 BAH28 Regfig5" 4 0 6$P ?",i& JFigure 5. Genetic organization of the Trp operon and its control elements &K 2I`Z,&B 0 s *޽h ? f3f3ff___PPT10+:D' P= @B D' = @BA?%,( < +O%,( < +D' =%(Dm' =%(D' =4@BBBB%(E' =4 B`BPB`B?<*%(/%,( < +D' =1:Bvisible*o3>+B#style.visibility<*0%(D' =-m6Bbox(in)*<3<*0+  0 \ T 8(  8 8 6\%P ?"   FTable 2. The Trp operon and its control elements R = Regulatory gene that encodes for the trp Repressor protein that is concerned with regulating the synthesis of the 5 gene products. An active repressor binds to a specific nucleotide sequence in the operator region and thereby blocks binding of RNAp to the promoter to initiate transcription. O = Operator specific nucleotide sequence on DNA to which an active Repressor binds. P = Promoter specific nucleotide sequence on DNA to which RNA polymerase binds to initiate transcription. If the repressor protein binds to the operator, RNAp is prevented from binding with the promoter and initiating transcription. Therefore, none of the enzymes concerned with tryptophan biosynthesis are synthesized. A = Attenuator DNA sequence which lies between the operator and the structural genes for trp biosynthesis. The attenuator is a barrier that RNA polymerase must traverse if it is to transcribe the genes for tryptophan biosynthesis. In the presence of trp, most RNAp molecules fall off the DNA before transcribing the trp genes. In the absence of trp, RNAp is able to traverse the attenuator region to successfully transcribe the trp genes. Trp A, B, C, D, E = Trp Genes structural genes for enzymes involved in tryptophan biosynthesis. Trp = tryptophan end product of the tryptophan biosynthetic pathway. When combined with the repressor protein the Repressor is active. Trp is called a corepressor.  203333 3J3 353333D333 Dy yr "4J 0   Y B 8 s *޽h ? f3f3ff 0 <#(  <  < 609P ?"`@ 'Figure 6. Repression of the trp operon "( 2&`Z, < c 4AGo?r(lxTb ,$D  0j < BAIm Regfig6)& < c $  l  P ` ~Nirv;O&&(3 B < s *޽h ? f3f3ff0 0  @W(  @f @ 0 ?"    @ AJb m 3?E:\wuw-t-2\Other Regulated Systems.files\trp.gif[ @ c $UP lP  P {The Tryptophan Operon, B @ s *޽h ? f3f3ff1 0 ME@H(  H H c $^P l  Pv0___PPT106___PPT9 @2.M㉧Nirv;OCatabolite Repression F! ' 3''$3,   H 6LfP?"P~v0___PPT106___PPT9 1 N!kusaSvQ:g6R Ra|vv  ~bvV gv RsN|vv  [v Sa|RN"Nirv c0 W '3' 3'3 %3' 3 %3' 3%3' 3%3' 3 %3>!j H BA7 Regfig9 x o H 6wP ?" 5 sFigure 9. The Diauxic Growth Curve of E. coli grown in limiting concentrations of a mixture of glucose and lactose Jt 2&33E33_B H s *޽h ? f3f3ff 0 |`P(  Pl P DAK Regfig10<DD P 6tP?"0z  x1 cAMPNCAP~T0 2 S_cAMPNCAP YTir~T(W/TRP[ N,RNApolase MbN/TRP[~T lU_ۏL0 3 S_:\cAMPTCAPe, RNApolase NN/TRP[~T0Q  2333/333333    z P 6P ?"'J Figure 10. Catabolite repression is positive control of the lac operon. The effect is an increase in the rate of transcription. $ 2`Z> ';B P s *޽h ? f3f3ff|2 0 ,$pT(  T T 6 P?"" ~v0___PPT106___PPT9 $ ga|e   cAMPvNuSa|b6R0 a|Mirb6RzxsSvv;m'` cAMPSm^ NM cAMP NNCAP~T RNAYZv N~T lU_ NۏL0 ea|e   cAMPSm^ NGS cAMPNCAP(Mirvo;mˆ}v)~T,N/TRWV~T lU_ۏL.j %3' 3A%3' 36%3  B T s *޽h ? f3f3ff! 0 V!N!EE\ (  \ \ 0\P  d1 X[(W[ire mRNA_0RlU_ WV c6RR ~gWV P P R t O z y a t -JSsN|v nv lYNpWv [ir ;Oir 1Y;m;Oir RNAYZv cAMP CRP n, @ZG ]&&(',&j$-? < XJ O1]emu \ c $P lO  P dsN|d~P[ 1 F @ \  fr \ 6Zp <fr \ 6Z  <N n  \ @N  ^ p \ n TB  \ c $D00TB  \ c $D0TB  \ c $DP0PTB  \ c $DZB  \ s *D ZB \ s *D PPTB \ c $DppTB \ c $DTB \ c $D  TB \ c $DTB \ c $DTB \ c $DTB \ c $DppZ \ s *pPZ \ s *p0PZB \ s *D0 ZB \ s *D0P PZ2 \ s *`Pfr \ 6Z^Z2 \ s *pZ2 \ s *t \ DB CDElF[[@ $0Hpppp0pPppppp0pPpppp0Ppppp0pPppppp0pPpppp0`0Ppppp 0Xp ppp 0 X p   P p p p  8 P `p p x p?@@                              `@ \ BC0DE4F 0 @`000 @`0 @    @ZB  \ s *D@Z !\ s * 0Z2 "\ s *PPZB #\B s *D0`ZB $\ s *D0PZ2 %\ s * ` &\ 0BtdZR '\ s *DXN @ (\ 0ZR )\ s *D@lR *\ <G%@lR +\ <G&0TB ,\B c $D0TB -\ c $DZR .\ s *@p&N v @  /\ `*p`r 0\B 0G @ `r 1\ 0Gv  ZB 2\ s *D @ 0ZB 3\ s *D@PZB 4\ s *D@0ZB 5\ s *D0ZB 6\ s *DZB 7\ s *D`` 8\ 0  Z2 9\ s * p Z :\ s *p 0 &N 0 @  ;\ p : `r <\B 0G @ `r =\ 0G0 @ f" >\ 6G$ p  ` ?\ 00 Z2 @\ s * L  A\ 6hP  z sN|d~P[&(LB B\ c $DppLB C\ c $DdR D\ <G%p  E\ 0tPp`j  vDNA RNA ˆ}v(( 2 & B \ s *޽h ? f3f3ff  0 bZ((`(  `~ ` s *P lP   P  ` <P  l  P L8XJ O1]emubF .  ` P@@ TB ` c $DPTB ` c $D@@PT N .  ` . TB ` c $D@@P N .   ` . TB  ` c $DPZB  ` s *DpZB  ` s *D p Z  ` s *@ ZB ` s *DZB ` s *D   N .  ` . TB ` c $DpTB ` c $Dp  TB ` c $DPTB ` c $D  PTB ` c $DPTB ` c $DPTB ` c $DPZ ` s *@ Z2 ` s *p0` fr ` 6Z.pZ2 ` s *pP`@Z2 ` s *p` ` BC0DE4F 0 @`000 @`0 @    ZB ` s *DPPZ ` s * ` Z2  ` s *   ZB !`B s *D0  ZB "` s *D  Z2 #` s *p  ` $` 0D 4 ZR %` s *&N 0 @  &` p :` `r '`B 0G @ `r (` 0G0 @ B ` s *޽h ? 33̙3   0 | t d (  d~ d s *\ W l   W ( d 0W 0  3 NX[(W[ire mRNAellU_ WV c6RR ~gWV P P R t O z y a t `-0Z$Z0ZP&'&4 $68XJ O1]emuLB d c $D00LB d c $D 0 LB d c $D0LB d c $DRB d s *D   RB  d s *D LB  d c $DppLB  d c $DLB  d c $D LB  d c $DLB d c $DLB d c $DLB d c $DppR d s * pR d s * p0RB d s *D 0  RB d s *D0 R2 d s *^r d 6ZR2 d s *R2 d s * d BC0DE4F 0 @`000 @`0 @    @@R d s *RR d s *D@jR d BG @ B d s *޽h ? 33̙3 0 hB(  hn h FALP lacoperonF h 6.W ?" 5 This assortment of genes and their regulatory regions is called the Lac operon. The Lac operon has been examined in detail. Here is a diagram of it: 2 2D I4H 8B h s *޽h ? f3f3ff  0 ++lR(  lL  l# BN  l N   l   l B8W ?"  GElement$t l 6 ?" N   l   l B=W ?"  Gpurpose$t  l 6 ?" N  @  l  @  l BPBW ?" @ cOperator (LacO) t  l 6 ?" @N  @  l  @ l BGW ?" @ Lbinding site for repressort l 6 ?" @N @` l @` l B,LW ?"@` cPromoter (LacP) t l 6 ?"@`N @` l @` l BpPW ?"@` Qbinding site for RNA polymerase t l 6 ?"@`N ` l ` l BTUW ?"` dRepressor (LacI) t l 6 ?"`N ` l ` l BXZW ?"` w#gene encoding lac repressor protein$$t l 6 ?"`N  l  l B_W ?" {IBinds to DNA at operator and blocks binding of RNA polymerase at promoterJJt l 6 ?"N  l   l BdW ?" 4Pit !l 6 ?"N  "l  #l B|iW ?" epromoter for LacI t $l 6 ?"N  %l  &l BdnW ?" 5CAPt 'l 6 ?"N  (l  )l BsW ?" u!binding site for cAMP/CAP complex"" t *l 6 ?"t +l 6 h+?"B l s *޽h ? f3f3ff3 0 pY(  pZ p c 2AM m laconl`P p  0xyW l  W GWhen lactose is present(H p 0޽h ? f3f3ff4 0 x](  x\ x c 4AN" lacoff`Pe x c $pW l`  W U%When the inducer (lactose) is removed&& H x 0޽h ? f3f3ff%5 0  e(  R  C *AOlacgluc^   0W lP   W [+When levels of glucose in the cell are high,,H  0޽h ? f3f3ff 0 @##7(  8  NW@AB`TC`T S" p 8___PPT10N___PPT90( ( (N)uirTb_vN"c: 10~Nir;Oend product repression) ~Nir;O_@b gvvTb0 20Sfeedback)bSgb6R(allosteric inhibition) ~Nir[Tb_,{NvTbvb6R& '%'%'% '''%'% !    0 %  c $@AB`TC`T= },$D 0  <Wf3" U0  o)A B C D** 2)$ NB  S " p NB  S " @   <hWf3" p t  HE1* 2$   <Wf3" p   HE2* 2$ NB   S "0 @    <Wf3" p  HE3* 2$ 4    @AB CDEFf3"""`"1RmIEB >19V*|**9>ZSIA<z*-+4Be4hn, o r u | |@ t HB   C "00HB   C "HB  C "0HB  C "0 00 HB  C " ` HB  C "`  <Wf3">  KGene1* 2$   <Wf3"0   KGene2* 2$   <Wf3" ~  KGene3* 2$ NB  S "@NB  S "P P P NB  S "P @P NB  S "P P P NB  S "P @P NB  S " P NB B S "PNB  S "  HB  C "  HB  C "   <Wf3"   f~NirSb6R* 2$     < 0   p (  L P p # = ]T  c $f3" P pNB  S "NB  S "@` @NB  S "  NB  S "p \  ND]@AB`TC`T ,$ 0 1 [Nv~_N"_ %{Q:w'`zS*hSy/}-NN"Nir A a B b C c D d E a"%        "$;6   Z ]@AB`TC`T! S" G  1.1%{Q:wW̃*hv^(u,$NN&$NN    6]o9 q^ : +gzNirE[uCN/f_v @bN ^(WW{QW-NPϑO~E OKNN~c̃*hu FOS NN bS;Obb6R ُ7hMb g)RÑ*hy/}-NNirC 0,S & 3N$F -F  `     , zB   <Do?" P zB   <Do?"@ B  BDo?"p zB  <Do?" @ B  BDfo?" ` B B BDfo?"0` ` B  HDfo?"0 0` zB  <Df>?"p p zB  <Df>?"     <<]!  lrT  ] h1.W Of[el$$B  s *޽h ? ̙33  0 L D   (  h  N ]@AB`TC`TS" * v(2) R/eN"_`Q YBg S)R(u%{Q:w'`KQ gOS Tb/}RSb6Ry/}+gzNir NS)R(uS͑:wSuuN-NNir<<" 8:  p  # ,$D 0h  s *f3""`Ppn  0f3""`0 p   B$]f3""`Pv ZA B C&$   B0*]f3""`   CD&$   BP.]f3""`   CE&$   <3]f3""` &  EF( 2$   <p8]f3""`` e F  EG( 2$ TB  c $"@TB  c $"0TB  c $" TB  c $"p 0 TB  c $" ` PTB  c $" 0 n   0f3""``  B  s *޽h ? ̙337 0 F(  h  B@]@AB`TC`T : R/e_  `$ V(lxSu:7(lxhFg̃vHom X '$'$ &$3.$3@  BAP28 o?)YQ(lxN"/,$D 0B  s *޽h ? ̙33/'___PPT10+hD' ]= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D-' =4@BBBB%(E' =4 B`BPB`B?<*%(/%,( < +D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-6B)randombar(vertical)*<3<*+ 0 xp(  T  BXK]@AB`TC`Td L R/e_  a$xSu IMPTb_vN"c D' '$ &$'$.     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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. LCG95L@_=H  0޽h ? ̙33 D 0 y q  (  ^  S T   bk  c $0bT @  b 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> jk 0pxH  0޽h ? ̙33E 0 $((  $^ $ S T   b $ c $bT @  b  H $ 0޽h ? ̙33H 0 4((  4^ 4 S T   b 4 c $bT @  b  H 4 0޽h ? ̙33K 0 0D((  D^ D S T   d D c $dT @  d  H D 0޽h ? ̙33L 0 xPL(  L^ L S T   dr L c $'PT @  d  Catabolite Repression Enzyme Induction is still considered a form of negative control because the effect of the regulatory molecule (the active repressor) is to decrease or downregulate transcription. Catabolite repression is a type of positive control of transcription, since a regulatory protein affects an increase (upregulation) in the rate of transcription of an operon. The process was discovered in E. coli and was originally referred to as the glucose effect because it was found that glucose repressed the synthesis of certain inducible enzymes, even though the inducer of the pathway was present in the environment. The discovery was made during study of the regulation of lac operon in E. coli. Since glucose is degraded by constitutive enzymes and lactose is initially degraded by inducible enzymes, what would happen if the bacterium was grown in limiting amounts of glucose and lactose? A plot of the bacterial growth rate resulted in a diauxic growth curve which showed two distinct phases of active growth (Figure 9). During the first phase of exponential growth, the bacteria utilize glucose as a source of energy until all the glucose is exhausted. Then, after a secondary lag phase, the lactose is utilized during a second stage of exponential growth. Figure 9. The Diauxic Growth Curve of E. coli grown in limiting concentrations of a mixture of glucose and lactose During the period of glucose utilization, lactose is not utilized because the cells are unable to transport and cleave the disaccharide lactose. Glucose is always metabolized first in preference to other sugars. Only after glucose is completely utilized is lactose degraded. The lactose operon is repressed even though lactose (the inducer) is present. The ecological rationale is that glucose is a better source of energy than lactose since its utilization requires two less enzymes. Only after glucose is exhausted are the enzymes for lactose utilization synthesized. The secondary lag during diauxic growth represents the time required for the complete induction of the lac operon and synthesis of the enzymes necessary for lactose utilization (lactose permease and beta-galactosidase). Only then does bacterial growth occur at the expense of lactose. Since the availability of glucose represses the enzymes for lactose utilization, this type of repression became known as catabolite repression or the glucose effect. Glucose is known to repress a large number of inducible enzymes in many different bacteria. Glucose represses the induction of inducible operons by inhibiting the synthesis of cyclic AMP (cAMP), a nucleotide that is required for the initiation of transcription of a large number of inducible enzyme systems including the lac operon. The role of cyclic a cAMP is complicated. cAMP is required to activate an allosteric protein called CAP (catabolite activator protein) which binds to the promoter CAP site and stimulates the binding of RNAp polymerase to the promoter for the initiation of transcription. Thus to efficiently promote gene transcription of the lac operon, not only must lactose be present to inactivate the lac repressor, but cAMP must be available to bind to CAP which binds to DNA to facilitate transcription. In the presence of glucose, adenylate cyclase (AC) activity is blocked. AC is required to synthesize cAMP from ATP. Therefore, if cAMP levels are low, CAP is inactive and transcription does not occur. In the absence of glucose, cAMP levels are high, CAP is activated by cAMP, and transcription occurs (in the presence of lactose). Many positively controlled promoters, such as the lac promoter, are not fully functional in the presence of RNAp alone and require activation by CAP. CAP is encoded by a separate Regulatory gene, and is present in constitutive levels. CAP is active only in the presence of cAMP. The binding of cAMP to CAP causes a conformational change in the protein allowing it to bind to the promoter near the RNAp binding site. CAP can apparently interact with RNAp to increase operon transcription about 50-fold. Positive control of the lac operon is illustrated in Figure10. Figure 10. Catabolite repression is positive control of the lac operon. The effect is an increase in the rate of transcription. As a form of catabolite repression, the glucose effect serves a useful function in bacteria: it requires the cells to use the best available source of energy. For many bacteria, glucose is the most common and readily utilizable substrate for growth. Thus, it inhibits indirectly the synthesis of enzymes that metabolize poorer sources of energy.  !(Y'=0&`Z`ZE`Zt5*"g6Y`Z^<   l $"h Je&F H   ,   Vk 4m7]%k6b/ 5 + & G EH L 0޽h ? ̙33lN 0 ,$X(  X^ X S T   d X c $3dT @  d faIN O_uirHQ)R(usX-N[f)R(uvn dk:g6RO_uir(WNUOAQsX-NNg'Y^A~k0 42  4H X 0޽h ? ̙33T 0 RJt(  t^ t S T   dD t c $^dT @  d lSo let's look at the lac operon in action. When lactose is present, it acts as an inducer of the operon. It enters the cell and binds to the Lac repressor, inducing a conformational change that allows the repressor to fall off the DNA. Now the RNA polymerase is free to move along the DNA and RNA can be made from the three genes. Lactose can now be metabolized. P 2 kH t 0޽h ? ̙33U 0 |'(  |^ | S T   d | c $$kdT @  d =When the inducer (lactose) is removed, the repressor returns to its original conformation and binds to the DNA, so that RNA polymerase can no longer get past the promoter. No RNA and no protein is made. Note that RNA polymerase can still bind to the promoter though it is unable to move past it. That means that when the cell is ready to use the operon, RNA polymerase is already there and waiting to begin transcription; the promoter doesn't have to wait for the holoenzyme to bind. We could say that the operon is primed for transcription upon the addition of lactose. $>3k S v B =H | 0޽h ? ̙33V 0 0((  ^  S T   d  c $|dT @  d  H  0޽h ? ̙33^ 0 ((  ^  S T   d  c $dT @  d  H  0޽h ? ̙33a 0 :(  ^  S T   d  c $dT @`  dH@___PPT9" 4. 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YCoQSMKSN |T8T-Nv'l؏}SOvpeϑ(W N Tvy N TvsXagN NSXSQ gR/e|T8TvX[(W hs(Weg N Tv^irv؏SRۏeQ|T8Te g N TvR/e N Tv_uir~ހr }|~ g+R YE.Coli(W:'le |T8TR:N$N/e sSCyt.b556 Cyt.c; Cyt.b558 Cyt.dZv###+#+ #nXWXYZ[\]^_`co6e'l|T8Tanaerobic respiration 2'$'Ji_N{||T8T+gzv"lSSO:NYne:g'lSir*N+R:N g:g'lSir vuir'lS /fNye'lagN NۏLvNHesNOvyrk|T8T0 e'l|T8Tv{|W ]xxv|T8TNO3- NO2-, NO, N2 e:gv|T8T kxxv|T8TSO42- SO32-,S3O62-,S2O32-,H2S kx |T 8T S0 S-2 xxv|T8T CO2,HCO3- CH3COOH CO2, HCO3- CH4 ^"}x|T8T^"}x %tsxFv`v&;"&$                  + -      :    Rh? 7 l-.  , [IN1u]xxv؏SbNO2 v^ۏNek؏SbN2vǏ z^IN 0rINvS]xS\O(uNc1uN]xx؏SbN2vǏ z0 agNS'lm4lvWXb{k4lXX-N ̃y\pe_{QTS{Q̃0YBacillus lichenoformis0Wcb[Fg̃ 0Paracoccus denitrificans1.loRt̃ 0Pseudomonas aeruginosaܔ~GPUSހ̃ 0Ps. stutzerielGPUSހ̃ 0Thiobacillus denitrificans1.lkxFg̃ NSSpirillum̃^\ TMoraxellabl̃^\ I{0 aINWX-N.lCQ }Am1Yv͑SVKNN04l;z0u-Ne(uSf[.l gHe)R(usS g25%]S0SYSN)R(u4lu'`S]xS~̃Sdal4l-Nv]xxv0U' #++#+#+#'#'L## # # ## # ## # #'A# '    .fpq uk̃(WpH3.5-4.5e vYNSu 1v 1"lv Nnx YN YN ~̃(Zymomonas mobilis)vYNSu ǏED_NuYN ;`S^Y N a|+ADP+Pi 2YN+2CO2+ATP ~̃(Leuconostoc mesenteroides)vYNSu ǏHMP_NuYN0sNxI{ ;`S^Y N a|+ADP+Pi sNx+YN+CO2+ATP TWR|SuNir-NN gYNNy g:girRP[vR|Su _WsNxSud;NNirYNY ؏X[(W gvQ[ g:girRP[vSuvZJvZvZAvZ#vZCvZ9ffZ!##)##B#+## #'4#+#'#'## Q      8 0 0%&'(rs  tv.(4)1u(lWxSuN SticklandS^#. w)*xN0{Q_uirNATPT؏SR  yz "e:g^ir1"lT5uP[ۏeQ|T8TvMO& ( H2 NH4+ S2- SO32- S2O3- Fe2+ NO2- NAD FP Q ,Cyt.b Cyt.cc1 Cyt.a1.aa3 O2(NO3) ATP ATP ATP lckT OSNuATP, T OmATP v^Nu؏SR[H]             .      mhn~ +{ | } 2.^_sIQTxxS #(  ~  3.Uṽ+}vIQN[ATPTb$$  , ,{N RN"TTbN"vT|& #   ghijk,{ N _uirryrTbN">NO$N;NQ[ N0{Q_uirvCO2V[ N0uirV.leu N0Z|uirTbnZ"Z$ $ld$$cN0{Q_uirvCO2V[4  h`N Calvin_s Z dBN S'lYNpCOA_  N TTCA_s V WNx_""# 6  joe N0uirV.l"(   !lifN V.lvuS:g6R "$ 1.uirV.lv6 } 1 ATPvO^ 2 ؏SR[H]SvQ O}SO 3 V.lv 4 ؏S^ir N2 5 AyP[ 6 %N Z|USSOvTb  "$ Z|USSOvTb  ~̃b $$ ,{ N6k(W~ހYvTb"$   !"#z {$%&'()*+,-.&1 TRv  isoenzyme'$3" &[INPSv TvuSS^ vRP[~g g]+RvN~v0 aIN(WN*NR/eN"_-N Yg(WR/epNMRvN*NevS^/f1uQ*N TRvPSe RR/eN"vQ*Ng~Nir__R+R[ُQ*N TRvSub6R\O(u0   gNNirǏϑNb6Rv^v;m [vQNNirlq_T0 >NO'YFg̃v)YQ(lxe(lWxTbvZ&&&G&&&&&$c-/0 )YQ(lxe&$f` )YQ(lx E,R III I E,R 4-xx)YQ(lx E )YQ(lxJS E,R II I E N"l!TvUNx TWN(lx TWN(lxxx E,R R O-%tsp TWN(lx ς(lx E,R mQ"l!TvUNx kx 2-nNx R N(lW^Nx TWJS(lx R V(lx 2ukx(lx _N(lxvEZ0Z("Z00Z(Zg )g    12$ )YQ(lxe 7(lxhFg̃ "( )YQ(lx E,R 4-xx)YQ(lx E )YQ(lxJS E,R E N"l!TvUNx TWN(lx TWN(lxxx E,R R O-%tsp TWN(lx ς(lx E,R mQ"l!TvUNx kx 2-nNx R N(lW^Nx TWJS(lx R V(lx 2ukx(lx _N(lxj5Z 0Z("Z00Z(Z   345676 s^aTb"(" (WĞrwFg̃Brevibacterium flavus -N)YQ(lxvTb PEP Py IpYNx YNpCoA Asp gjx ATP CO2.nlz  87 N"N&("6hb[hQ NvsQv$Nag_KNv0ُy\O(uN,(WؚSm^ NMb>f:y N:NRb6R0 Asp Asp- p Asa DAP Lys N"l!TvUN xTbv Hse Thr Ile Py _NW Leu gx"g X*+9:;Sgv&( <=/"2.;Orepression @&$ &&$"  0123456789:?@ABCDEFGHIJKLMN  OPQRSTUVWXYZ[\]^ N c6R~ހn'`  $  _`a,gz\~  1.uir'lS@bv1"l0"lTS"lv N*N6k0 2.9hnc"lSSOv N T _uirNv Nye_|T8T0e'l|T8TTSu0 3.STIQ{Q̃Nve_0 4.RN"NTb N"vT|0 5._uirryrvTbN"_0 6._uirN"v;Ne_{Z{!l  b Y`N(  c N0T{   d>"2.;Orepression @&$ &&$"  ?@ABCDEFGHIJKMNOPsN|d~P[ 2#(2 [irT;OirX[(We WV c6RR ~gWV P P R t O z y a t 2#Z40ZW'"'.QsN|d~P[ 3#( RSTUVWX/<  !(NOVn !"#$%&'()*+,-./02345;8<9=:e;f<g=h>i?j@kAlBmCnDoEP   0 2* (     <P51ff  l"<  w7  |   6`1 ?"` @ : 2   6Eff?"  p /f;xN"Ǐ z-NSbsQ.v(WQvN|RvvTbvsa N f{_^0Wc6RTQ\+gzNirvTb0 &;O\O(uv{|W `$+gzNir;Oend-product repression 1uN~NirvǏϑy/} [uirTb_-NvTbv;Ovsa 8^8^Su(W(lWx0 VdTT'VvUI{ُN͑~gCQNuirTbveP0 OYǏϑv|(lx;ONSNTb|(lxvYvvTb0  & &&&&!&&&!q&H6 YB  s *޽h ? f3f3ffy___PPT10Y+D=' = @B ++ 0 B: (    Z 1ff??',f a$RN"ir;Ocatabolite repression  S__uir(W+T g$NyYR㉕^irvW{QW-Nue )R(u_vyR㉕^irO;O)R(ubav^irv gsQvvTbvsa0geSsN'YFg̃u(W+Ta|TsN|vW{QWe,EeSya|He^0RN"ir;O[Qs N!kudiauxic growth  . vc\O(u/fOHQ)R(uvxnv-NN"ir  [(/fVN "S^-NgN-NN"irb+gzN"irvǏϑy/} ;ON"-NNNvvTbvsa00g!mgg! gg g+gx   S/H  0޽h ? fff___PPT10i.@=+D=' = @B +J8 0 qi (  "   B1ff  l"<    1v0___PPT106___PPT9 @2.M㉧Nirv;OCatabolite Repression F! ' 3''$3,    6 ff?"p~v0___PPT106___PPT9 1 N!kusaSvQ:g6R Ra|vv  ~bvV gv RsN|vv  [v Sa|RN"Nirv c0 P ## # !# !# !# !# !>!j  BA7 Regfig9 x o  6   ?" 5 sFigure 9. The Diauxic Growth Curve of E. coli grown in limiting concentrations of a mixture of glucose and lactose Jt 2&33E33_B  s *޽h ? f3f3ffy___PPT10Y+D=' |= @B + 9 0 4, (  .   <ff  l"<  dR    f3.vTbv:g6R &$3 \  6(ff?"D d~P[f[i 10d~P[operon /fWVhTc6RvN*N[teUSCQ vQ-NSb~gWV WV d\OP[T/TRP[0 `$~gWV(structural genes)/fQ[gNYvDNA !jg S9hncvQ NvxWz^lU_Qv^vmRNA 6qTQSǏ8h|SOlыQv^vv;xˆ}v(vDNA^R a$/TRP[(promoter)OVNDNAvRNAZTv@bƋ+RvxWz^ /fRNAZTvv~TMOTlU_wp(WY`Q N؏SbOۏُNǏ zvˆ}v~TMOp0 b$d~P[operator MON/TRWVT~gWVKNvNkxWz^ /f;Oˆ}vv~TMOp ǏN;Oirv~TegQ[~gWVvlU_/f&TۏL c$WVregulator gene (uNx~bWˆ}vvWV N,܏yd~P[,FO(WS8huir-N,SNMONd~P[e,xˆ}v0 2 &5&&[5&& & & &&[G&&[>&&[&"&3 '  *< B  s *޽h ? f3f3ffy___PPT10Y+D=' |= @B +: 0  r(  f  0 ?"   ACt P ?http://gened.emc.maricopa.edu/bio/bio181/BIOBK/operon.gif/  s *  l-   ~Structure of a typical operon "$0B  s *޽h ? f3f3ffy___PPT10Y+D=' |= @B + 0 !p(  *   <\ff  l"<  w79    b NN͑/g(&$3$   6pff?": 20[irinducer N;Oircorepressor   [ir  /fwYv[Tbvir( YsN|I{Nˆ}v~T b6RvQNd~WV v~TOۏlU_ۏL  ;Oir  /f;OvNuvir( Y(lWxT8hxI{Nˆ}v~T OۏvQNd~WVv~Tb6RlU_ۏL  [N/f\RP[OSir( 8^;`y:NHe^ireffecor SNˆ}vv~TNOTSuSg\O(u v^ۏNekcؚbMNONd~WVv~TR00Z ' ' ' ' '3' ' ' "'3'' ' *'  8O)B  s *޽h ? f3f3ffy___PPT10Y+D=' |= @B + 0 *" !(    Zff?? 30;Oirrepressor N;Oirˆ}vaporepresseor   N/f1uWVxNuyr_'`ˆ}v(regulatory potein) /fN{|NORP[ϑSgˆ}v g$N*N~TMOp N*NNd~WV~T SNMOpSNHe^ir~TS_ˆ}vNHe^ir~TT 1\SuSg\O(u SgTNd~WVv~TRScؚb NM0 g;m'`  SNO~Te;m'`  NNO~T ;Oir:(Wl g[ireNd~WV~Tvˆ}v ;Oirˆ}v:S(W g;OirX[(WeMbNd~WV~T00s0s%ggg!ggIgg3gggg   ZH  0޽h ? fff___PPT10i.@_+D=' |= @B +W 0 ~v0!(    <`ff?" ~v0___PPT106___PPT9 RcklU_Ǐ zOVNˆ}vvX[(W0 lU_Ǐ z NOVNˆ}vvX[(W0\* 2''3''3 *B  s *޽h ? f3f3ffy___PPT10Y+D=' |= @B +F 0 me@! (  j  BAsI *gh-5 b+v  Nv2 @  C Induction G$  Nv @ C  ?InducerE  Nlv @| ?  ?InducerE  6ff" JPositive controlGf  6ffR6  JNegative controlGf  <ff ; } X c[|~    <ff X ckc[|~ H  0޽h ? ̙333y___PPT10Y+D=' |= @B +{ 0 P! 2(   j  BAz *gh-6 by  6  JNegative controlGf  6P ff<  JPositive controlGf  N<v2 @n  D Repression G   Nv @   Y Corepressor A   Nh'v @z   Y Corepressor A   <1ff;}  X c;O|~   <;ff;I  X ckc;O|~ H  0޽h ? ̙333y___PPT10Y+D=' |= @B + 0 `! `(     BA{a 3lacop05,$D 0xB  B Ԕ?" Q  HI ?" < |PlacB0n<ZGOH  0޽h ? f3f3y___PPT10Y+D=' |= @B +Q 0 22p!DD !2(   5 z  Z   @ ,$D 0`   0o Z    `tX1?G  ;,$D 0 7i    Bb1 h ;,$D 0 Ep(      Bl1h H ;,$D  0 Co&    Hw1H ; Cz&    H؁1 ; Ey(       BX1 k; Ca& 6    3 E e T,$D 0T y   # y2    S ~vd @?y2    S ~vd @?yT y  # @2   S ~vd @?y2   S ~vd @?ylB   <DogM   0 |T ]Very low level of lac mRNA&( 2   0 8p TAbsence of lactose&( 2   0   <Active 2   `D1? 6 ,$D 0 Ci&   B1 6 ,$D 0 Ep(     BL1 6 ,$D  0 Co&   H16  Cz&   Hx1*6  Ey(     B1*y6  Ca& dB  <D3> M dB  <D3o B dB  @ <D3o *,< dB  <D3o B dB  <D3o yyB     ZD ?  b-Galactosidase<( 2  ! ZH  ?K h NPermease* ( 2 " Z ?  nTransacetylase*( 22 # S ~vd @?j  2 $ S ~fvd @?j v 2 % S ~vd @? ^ 2 & S ~vd @? m 2 ' S ~vd @?  ? 2 ( S ~vd @? E 2 ) S ~vd @? E? 2 * S ~fvd @? P 2 + S ~fvd @? 1 2 , S ~fvd @? ? 2 - S ~fvd @? V 2 . S ~fvd @? v 2 / S ~ffvd @? "? 2 0 S ~ffvd @? ` 2 1 S ~ffvd @? A 2 2 S ~ffvd @? ; 2 3 S ~ffvd @?j  2 4 S ~ffvd @? j .  5 3  5 L ,$D 0T yOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOP 6 # y2 7  S ~vd @?y2 8  S ~vd @?yT y 9 # @2 :  S ~vd @?y2 ;  S ~vd @?y < 0  ~ UPresence of lactose&( 2R = B+jJ o  >   > 0$5  JInactive& 2 dB ? <Do~   @ <?ff?" ,$D 0 `Lack of inducer: the lac repressor block all but a very low level of trans-cription of lacZYA .Za0n<Z23IfffI A 6?" p,$D 0 B <(Mff?"D |,$D 0 Lactose is present, the low basal level of permease allows its uptake, and-galactosidase catalyzes the conversion of some lactose to allolactose. Allolactose acts as an inducer, binding to the lac repressor and inactivate it. 0n<Z2 3#f3f 3;f  3f 35fP+; ) C B ?"  @ ,$D 0 D 6?"&z v,$D 0H  0޽h ? ff̙___PPT10+ѕyD ' h= @B D' = @BA?%,( < +O%,( < +Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<* D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<* Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*A %(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*A D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*A D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*@ %(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*@ D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*@ Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*C %(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*C D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*C Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*D %(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*D D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*D D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*5 %(D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*B %(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*B D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*B +p+0+@ 0 ++0+B 0 + 0 ! (   \  S 4A|"Image2831H  0޽h ? fff___PPT10i.C`4+D='  = @B +; 0 ! m(   j  BAD Hlactose9   6<}  ?"`  l6The enzymes that are required for lactose degradation:7 27  6\ ff?"]  gbeta-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 ? f3f3ffy___PPT10Y+D=' = @B +< 0 "! (      <H ff  l"<  7    v0___PPT106___PPT9 0vTbv[Enzyme Induction L' 3' $3j  BAE Regfig7  0  6P  ?"@ 2Figure 7. The Lac operon and its control elements "3 21`ZB  s *޽h ? f3f3ffy___PPT10Y+D=' = @B +T 0 {s! (   j  BAFa HRegfig8   6 ff?"%E KFigure 8. Enzyme Induction. Induction (or derepression) of the lac operon is a form of negative control because the effect of the regulatory protein (Active repressor) is to stop transcription. & 2`Z>*  {  6 ff?"<4___PPT10 B___PPT9$  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.[ 2$ U%B  s *޽h ? f3f3ffy___PPT10Y+D=' = @B ++= 0 RJ!$ (  $  $  < ff  l"<  w   v0___PPT106___PPT9 2vTbv;OEnzyme Repression X'(3'$'(&03 $ 6 ff?" x1.~Nirv;O(end product repression): S;O) sS(WTbN"-N ~Nir;O_ @b gvvTb0:NWVhvc6R 0 Yr(lx(Try)Tbvc(ck)&c  " Z j)j $ BAH28 Regfig5 4 $ 6/  ?",i& JFigure 5. Genetic organization of the Trp operon and its control elements &K 2I`Z,&B $ s *޽h ? f3f3ffy___PPT10Y+D=' = @B + 0 80", (  ,  , c 4AGo?r(lxTb<dY,$D  0H , 0޽h ? fff'___PPT10.P/+0D' = @B D' = @BA?%,( < +O%,( < +D' =%(Dm' =%(D' =4@BBBB%(E' =4 B`BPB`B?<*%(/%,( < +D' =1:Bvisible*o3>+B#style.visibility<*, %(D' =-m6Bbox(in)*<3<*, +* 0 QI"0 (  0   0 6| ?"`@ 'Figure 6. Repression of the trp operon "( 2&`Z,j 0 BAIm Regfig6<d_( 0  <<|ff  l"<  w   | ` ~Nirv;O&&(3 B 0 s *޽h ? f3f3ffy___PPT10Y+D=' = @B +> 0  "4 W(  4 f 4 0 ?"    4 AJb m 3?E:\wuw-t-2\Other Regulated Systems.files\trp.gifI' 4 c $0| lP  | {The Tryptophan Operon, B 4 s *޽h ? f3f3ffy___PPT10Y+D=' = @B +? 0 @"< E(  < " <  B,|ff  l"<    |v0___PPT106___PPT9 @2.M㉧Nirv;OCatabolite Repression F! ' 3''$3,   < 6|ff?"V~v0___PPT106___PPT9 1 N!kusaSvQ:g6R Ra|vv  ~bvV gv RsN|vv  [v Sa|RN"Nirv c0 W '3' 3'3 %3' 3 %3' 3%3' 3%3' 3 %3>!j < BA7 Regfig9 x o < 6dz ?" 5 sFigure 9. The Diauxic Growth Curve of E. coli grown in limiting concentrations of a mixture of glucose and lactose Jt 2&33E33_B < s *޽h ? f3f3ffy___PPT10Y+D='  = @B + 0 `"D (  D  D ZA} ??  D BlGM a'CRP-binding site is an inverted repeat.((GfH D 0޽h ? f3f3y___PPT10Y+D='  = @B + 0 +#p"H (  H  H C *A{lacop05| ,$D 0B H H3?"  KC.8n<ZGB H Hk3?"   KA.8n<ZGB H HR3?" - KB.8n<ZG H H ';B L s *޽h ? f3f3ffy___PPT10Y+D='  = @B +@ 0 ,$"P (  P  P 6\ff?"Tt&~v0___PPT106___PPT9  ga|e   cAMPvNuSa|b6R0 a|Mirb6RzxsSvv;m'` cAMPSm^ NM cAMP NNCAP~T RNAYZv N~T lU_ NۏL0 ea|e   cAMPSm^ NGS cAMPNCAP(Mirvo;mˆ}v)~T,N/TRWV~T lU_ۏL0p '3'$3>'3'$36'3! B P s *޽h ? f3f3ffy___PPT10Y+D=' = @B + 0 "X (  X V X C .A~Image2836ZWZH X 0޽h ? fff___PPT10i.I`;+D=' = @B + 0 &"\ (  \ ~ \  `Dff??t' lacd~P[vckc:g6R/fcAMP-CAP YTirN DNA~T9eSNُN:Sk DNA!k~~g OۏNRNAZTv~T:Sv0ُS/fcAMP-CAPǏNRNAZTv~T QNDNA~T V OۏNRNAZTvN/TRP[v~T N X:_NlU_(V9-12B)0cAMP-CAP YTirvb_bSQN~ހQcAMPvSm^ S_Na|:Nne 1uNvQb6RzxsSvv;m'` ATP NlS:NcAMP ~ހQcAMPSm^MNO b_ NbCAP-cAMP YTir V sN|~gWV NlU_0~ހe ga|\O:Nn _N1\l g[sN|)R(uvQyvv0S(WsN|d~P[-N1uCAP-cAMPo;mvُyckc/fNyASRTtvc|~ wQ g^aIN0ck0cqQ TMT SOlU_g gHe0WۏL0(WcAMPckc-N_N g[irvX[(W NO;Oir1Y;m lU_MbۏL0 cAMP-CAPvc/fgvQ^lv dNgN|{|vN"v ~̃-NYvQNvR_Nhs:N[a|He^vOea Y Nx_sT|T8Tv|~-Nv'YYpev0RTyxn^irvv0MgN(lWxvv0YNx_vv0bu }TbvNS#kb_bvv|~I{I{000a#    82tH \ 0޽h ? fff___PPT10i.IQ+D=' = @B +  0 "` v(  ` ~ `  BAW Y??, Y ` 0Ÿ0 X 2H ` 0޽h ? ̙33y___PPT10Y+D=' = @B + 0 "d B(  d n d FALP lacoperonF d 6$ ?" 5 This assortment of genes and their regulatory regions is called the Lac operon. The Lac operon has been examined in detail. Here is a diagram of it: 2 2D I4H 8B d s *޽h ? f3f3ffy___PPT10Y+D=' = @B + 0 "++h R(  h L  h # BN  h  N   h    h  B ?"  GElement$t h  6 ?" N   h    h  B0  ?"  Gpurpose$t  h  6 ?" N  @  h   @  h  B>  ?" @ cOperator (LacO) t  h  6 ?" @N  @  h   @ h  BC  ?" @ Lbinding site for repressort h  6 ?" @N @` h  @` h  BM  ?"@` cPromoter (LacP) t h  6 ?"@`N @` h  @` h  BW  ?"@` Qbinding site for RNA polymerase t h  6 ?"@`N ` h  ` h  B,b  ?"` dRepressor (LacI) t h  6 ?"`N ` h  ` h  Bm  ?"` w#gene encoding lac repressor protein$$t h  6 ?"`N  h   h  Bw  ?" {IBinds to DNA at operator and blocks binding of RNA polymerase at promoterJJt h  6 ?"N  h    h  B  ?" 4Pit !h  6 ?"N  "h   #h  B  ?" epromoter for LacI t $h  6 ?"N  %h   &h  B  ?" 5CAPt 'h  6 ?"N  (h   )h  B  ?" u!binding site for cAMP/CAP complex"" t *h  6 ?"t +h  6 h+?"B h s *޽h ? f3f3ffy___PPT10Y+D=' = @B +A 0 #l Y(  l Z l c 2AM m laconK l  0ܮ  l   GWhen lactose is present(H l 0޽h ? f3f3ffy___PPT10Y+D=' = @B +B 0  #t ](  t \ t c 4AN" lacoff`Pe t c $  l`   U%When the inducer (lactose) is removed&& H t 0޽h ? f3f3ffy___PPT10Y+D=' = @B +C 0 @#| e(  | R | C *AOlacglucZC |  0P  lP    [+When levels of glucose in the cell are high,,H | 0޽h ? f3f3ffy___PPT10Y+D=' = @B +~ 0 `### ;(   b   Z @AB`TC`Tff "<  p 8___PPT10N___PPT90( ( (N)uirTb_vN"c: 10~Nir;Oend product repression) ~Nir;O_@b gvvTb0 20Sfeedback)bSgb6R(allosteric inhibition) ~Nir[Tb_,{NvTbvb6R& '%'%'% '''%'% !    0 +&  c $@AB`TC`T6 |,$D 0   <$|f3""` [6  g)A B C D"* 2) TB   c $" p TB   c $" @    <p|f3""` p z  HE1* 2$    <f3""` p   @E2" 2 TB   c $"0 @    <@ f3""` p  >E3  2 Z   S @AB CDEFf3"""`"1RmIEB >19V*|**9>ZSIA<z*-+4Be4hn, o r u | |@ "`t NB   S "00NB   S "NB   S "0NB   S "0 00 NB   S " ` NB   S "`   <f3""`D & KGene1* 2$    <#f3""`0  & CGene2" 2    <-f3""` & CGene3" 2 TB  c $"@TB  c $"P P P TB  c $"P @P TB  c $"P P P TB  c $"P @P TB  c $" P TB  B c $"PTB   c $"  NB   S "  NB   S "    <9f3""`   f~NirSb6R* 2$     <Df3""`@+f rmRNATb;O* 2 $  !  <TNf3""`0K ADNA" 2  "  <8Xf3""`0 F BmRNA" 2  #  <bff  l"<  7   v0___PPT106___PPT9 ^;`~,&(&( B  s *޽h ? ̙33y___PPT10Y+D=' |= @B +- 0 TL# (      <  l!`   r( N0N"vN]c6RSvQ(WSu]N-Nv^(u& 3   6 ?"T @]NSuvvv'Yϑy/}NN@bv_uirN"Nir0 N"vN]c6RN:N0WSb4x_uirvN"c6RSO| ON"g@wNN ^gveTۏL0 N]c6RN"vKbk 9eS_uirW Oyr'`(W Of[el  c6RSuagNuirSf[el  9eS~ހ'`l 2$!$$ $ $y$$qTB  s *޽h ? f3f3ffy___PPT10Y+D=' |= @B + 0   #  (   L P p  # = ]T   c $f3" P pNB   S "NB   S "@` @NB   S "  NB   S "p \   N(@AB`TC`T ,$ 0 1 [Nv~_N"_ %{Q:w'`zS*hSy/}-NN"Nir A a B b C c D d E a"%        "$;6   Z-@AB`TC`T! S" G  1.1%{Q:wW̃*hv^(u,$NN&$NN   6TAo9 q^ : +gzNirE[uCN/f_v @bN ^(WW{QW-NPϑO~E OKNN~c̃*hu FOS NN bS;Obb6R ُ7hMb g)RÑ*hy/}-NNirC 0,S & 3N$F -F  `     , zB   <Do?" P zB   <Do?"@ B   BDo?"p zB   <Do?" @ B   BDfo?" ` B  B BDfo?"0` ` B  HDfo?"0 0` zB   <Df>?"p p zB   <Df>?"     <lP!  lrT   h1.W Of[el$$B  s *޽h ? ̙33y___PPT10Y+D=' |= @B +%  0 L D #  (   h   NHY@AB`TC`TS" * v(2) R/eN"_`Q YBg S)R(u%{Q:w'`KQ gOS Tb/}RSb6Ry/}+gzNir NS)R(uS͑:wSuuN-NNir<<" 8:  p   # ,$D 0h   s *f3""`Ppn   0f3""`0 p    B_f3""`Pv ZA B C&$    Bjf3""`   CD&$    Btf3""`   CE&$    <~f3""` &  EF( 2$    <f3""`` e F  EG( 2$ TB   c $"@TB   c $"0TB  c $" TB   c $"p 0 TB  c $" ` PTB   c $" 0 n   0f3""``  B  s *޽h ? ̙33y___PPT10Y+D=' |= @B +7 0 # F(   h   BL@AB`TC`T : R/e_  `$ V(lxSu:7(lxhFg̃vHom X '$'$ &$3.$3@  BAP28 o?)YQ(lxN"/,$D 0B  s *޽h ? ̙33/'___PPT10+hD' |= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D-' =4@BBBB%(E' =4 B`BPB`B?<*%(/%,( < +D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-6B)randombar(vertical)*<3<* +Q 0 xp# (   T   Bt@AB`TC`Td L R/e_  a$xSu IMPTb_vN"c D' '$ &$'$.     Bf3"4 jctv[^(u* &   :AQ " o?Sua,$D 0B  s *޽h ? ̙33y___PPT10Y+D=' |= @B +D 0 # P(      <T3  ld,   x1.2 bSc6RzS*hv^(u$$   6?"'  &bSc6RzS*h  /fc[Sb6R NOeab[;O gb'` b$N|Q gKNṽ*h0 &bSc6RzS*hSNN~Nir~g{|P?P@PAPBPCPDPEPFPGPHPIPJPKPLPMPNPOPPPQPRPSPTPUPVPWPXPYPZP[P\P]P^P_P`PaPbPcPdPePfPgPhPiPjPkPlPmPnPoPpPqPrPsPtPuPvPwPxPyPzP|P}P~PPP(ff&  6/ ?" ,$D 0 ~& YgWVSuzS NNueHev;Oir NTd~WV~T bd~WVzS N b~gWV NSc6RvlU_ v vub\ NQ[BRb NQ+gzNirbRN"ir;O ُ7hvzS*hy:N ~bWzS*h0\pe`Q N ~bWzS*hSNu'Yϑv0kN,gؚvYvv ُyzS*hy:NNzS*h0"0x'B  s *޽h ? f3f3ff___PPT10j+mI(D' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(DX' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-6B+checkerboard(across)*<3<* +8+0+ 0 +%E 0 LD$ (   k  6 ?"0p   D A B C E F( 2=w   <!  l'   \N uirSf[el &03   6l ?" 81. mRMRSO~ǏSc6RpNOgyN"Nir'YϑNu2(2%%HF p    p B   BDjJ?"0B   BDjJ?" B  BDjJ?"  B   BDjJ?"  ` B   BDjJ?"` ` B  BD1?"ppp B  B HD1?"0 pppB   BD1?"p p B  B BD1?" p B  HD1?"p B  BD1?"@@0B  B BD1?"@B   HD1?"  6 ?"0 ;(-) 23  6< ?" `z  C(-)$ 23  6du ?"ppj  C(-)$ 23  6  ?"@  22. mR[BRNcؚ[vTbϑeg g}Yv[BR__ N/fvv^ir /f^irvMuir 3. SuNRyǏ z&T 4. c6RSuvW{QWbRBL 2 %&%%6,  B  s *޽h ? f3f3ffy___PPT10Y+D=' = @B +R#  0 0$ V(      B֤!  l      6 ?" ,$D 0 OހQvN"NirŏnoQS,d+gzNirvSb6R0 1. (uutf[Kbk  vcb6RvTbbOS:_c Y: (WGluSu-Nbuir }Sm^c6R(WNϑS'YϑRlGlu c6Ruir }v+TϑS9eS~ހvbR ۏ 9eS'` S_W{Qm-Nuir }+TϑؚeǑ(uϑmRR }vel QYN(lwFg̃v8hxSu-Nc6RV }/fMn2+ Mn2+v\O(uNuir }v+B#style.visibility<* %(D' =-o6Bdissolve*<3<* D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* 9%(D' =-o6Bdissolve*<3<* 9D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* 9Y%(D' =-o6Bdissolve*<3<* 9YD' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* Yt%(D' =-o6Bdissolve*<3<* YtD' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* t%(D' =-o6Bdissolve*<3<* tD' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* T%(D' =-o6Bdissolve*<3<* T+8+0+ 0 +x 0 nf@$ (      < K  lS" wt    N gjxSu    0( *9,$D 0 2BN0̃yNugjxṽy_Y FON ̃:N;N vQ-NSNўf NugjxvR:_ v^)R(uYyxn Ee8^/fuN NO(uṽy0 N0Su:gt~ހQ g Nx_sTYNx_sgjxTbv;mRؚ LN4Yxvb_gjx1"lvSgNV } Yё^\yP[v:ON S0Rb6R ُ g)RNgjxvy/}0 N0]zAm z`$SumvpH+B#style.visibility<* @%(D' =-6B'blinds(horizontal)*<3<* @D' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<* @%(D' =-6B'blinds(horizontal)*<3<* @D' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-6B'blinds(horizontal)*<3<* D' =%(D' =%(DT' =A@BBB B0B%(D' =1:Bvisible*o3>+B#style.visibility<* "%(D' =-6B'blinds(horizontal)*<3<* "+8+0+ 0 + 0 P$ L(      < K  ld    N 7(lxSu    0  : 2   0 8rN07(lxSũy Corynebacterium pekinense Corynebacterium glutamicum Brevibacterium flavu N0Su:gt `$7(lxNa-n bNx:NxgS_N|(:NSuSee Tb_SbEMP,HMP,TCA_s,YNx_sI{ a$7(lxNũva-n bNx'lSv;mR_1_b:\ 7(lx1"lvv;mR_ؚ b$uir }/f7(lxNũ_vNy~u } (W7(lxuirTb-Nw@w͑\O(u :ONbϑ*YؚOO7(lxTbS;0 uir }Ǐq_T~ހv'` q_T7(lxSu0 : 2! %L!%!  .!% D% / K    H  0޽h ? ̙33y___PPT10Y+D=' = @B +  0 `$ b(   x  c $ l       08 lS"  )2  H  0޽h ? 3333___PPT10i.wde_+D=' = @B +  0 p$ (     B@o   l D     TH??m y  &N0 T͋ʑ eHN"uir'lS^ir4ls^xxS0'lSxxS|T8T0e'l|T8T0SuSticklandS^+}IQTxxSuirV.l{|̃SORuN"ir!kuN"ir~bvSd 2d# / ,H  0޽h ? 3333___PPT10i.yd+D=' = @B +  0 4,$ (      TXЕ??/ 1.S_{Q_uiruir'lSǏ z-N W(1"lv;N gTVy_ ՋNNTꁄvyrp0 2.ՋRhk|T8T0e'l|T8TTSuv_ Tp0 3.NHNSticklandS^ fvQNSt0 4.ՋUṽ+}IQTxxSvW,gSt0 5.uirV.lv6y }/fNHN T gNHNRՋuirV.lvǏ z0 6.NHNS!kuN"ir?[NRuN" gNHNT| 7._uirN"(WSu]N-N gUO͑'`Ջ>NOf0  2%(   x  c $ԕ lZW3 u   H  0޽h ? 3333___PPT10i.xd0ҏ+D=' = @B + 0 p# j(      Z&&[&"&3 '  *< B  s *޽h ? f3f3ffy___PPT10Y+D=' = @B +!  0 H @ P (       <6 3  l $   b NN͑/g(&$3$    6 ?"@@ D20[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$$$!&3&$ $+$&$$$!$$I$&3$$$$$  :P*   Y  B   s *޽h ? f3f3ffy___PPT10Y+D=' = @B +) 0 pP(    c $  l  v0___PPT106___PPT9 &E.coli sN|d~P[f[( '$3.  c 4AGo?r(lxTb,$D  0B  s *޽h ? f3f3ff___PPT10+H&D' = @B D' = @BA?%,( < +O%,( < +D' =%(Dm' =%(D' =4@BBBB%(E' =4 B`BPB`B?<*%(/%,( < +D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+/ 0 0(  0 0 c $O  lwqW  v0___PPT106___PPT9 2vTbv;OEnzyme Repression X'(3'$'(&03 0 6W ?"<|{ 1.~Nirv;O(end product repression): S;O) sS(WTbN"-N ~Nir;O_ @b gvvTb0:NWVhvc6R 0 Yr(lx(Try)Tbvc(ck).c  & 3Z$3j)j 0 BAH28 Regfig5" 4 0 6X[  ?",i& JFigure 5. Genetic organization of the Trp operon and its control elements &K 2I`Z,&B 0 s *޽h ? f3f3ffy___PPT10Y+D=' = @B +f 0 <#(  <  < 6  ?"`@ 'Figure 6. Repression of the trp operon "( 2&`Z, < c 4AGo?r(lxTbkI,$D  0j < BAIm Regfig6)& < c $  l<,   ` ~Nirv;O&&(3 B < s *޽h ? f3f3ffy___PPT10Y+D='  = @B + 0 @##a(  b  Z @AB`TC`T "<  z 8___PPT10N___PPT90( ( (N)uirTb_vN"c: 10~Nir;Oend product repression) ~Nir;O_@b gvvTb0 20Sfeedback)bSgb6R(allosteric inhibition) ~Nir[Tb_,{NvTbvb6R& '%'%'% '''%'% !    0 %  c $@AB`TC`T= },$D 0  <@ f3" U0  o)A B C D** 2)$ NB  S " p NB  S " @   <t f3" p t  HE1* 2$   <D f3" p   HE2* 2$ NB   S "0 @    < f3" p  HE3* 2$ 4    @AB CDEFf3"""`"1RmIEB >19V*|**9>ZSIA<z*-+4Be4hn, o r u | |@ t HB   C "00HB   C "HB  C "0HB  C "0 00 HB  C " ` HB  C "`  <( f3">  KGene1* 2$   <h f3"0   KGene2* 2$   <h f3" ~  KGene3* 2$ NB  S "@NB  S "P P P NB  S "P @P NB  S "P P P NB  S "P @P NB  S " P NB B S "PNB  S "  HB  C "  HB  C "   < f3"   f~NirSb6R* 2$     < f3"@%` rmRNATb;O* 2 $  ! <L% f3"0E IDNA* 2$  " <) f3"0 @ JmRNA* 2$  # c $'  l    V;`~$$  B  s *޽h ? ̙33y___PPT10Y+D=' Q = @B + 0 x (  ^  S T    r  c $8T @     Catabolite Repression Enzyme Induction is still considered a form of negative control because the effect of the regulatory molecule (the active repressor) is to decrease or downregulate transcription. Catabolite repression is a type of positive control of transcription, since a regulatory protein affects an increase (upregulation) in the rate of transcription of an operon. The process was discovered in E. coli and was originally referred to as the glucose effect because it was found that glucose repressed the synthesis of certain inducible enzymes, even though the inducer of the pathway was present in the environment. The discovery was made during study of the regulation of lac operon in E. coli. Since glucose is degraded by constitutive enzymes and lactose is initially degraded by inducible enzymes, what would happen if the bacterium was grown in limiting amounts of glucose and lactose? A plot of the bacterial growth rate resulted in a diauxic growth curve which showed two distinct phases of active growth (Figure 9). During the first phase of exponential growth, the bacteria utilize glucose as a source of energy until all the glucose is exhausted. Then, after a secondary lag phase, the lactose is utilized during a second stage of exponential growth. Figure 9. The Diauxic Growth Curve of E. coli grown in limiting concentrations of a mixture of glucose and lactose During the period of glucose utilization, lactose is not utilized because the cells are unable to transport and cleave the disaccharide lactose. Glucose is always metabolized first in preference to other sugars. Only after glucose is completely utilized is lactose degraded. The lactose operon is repressed even though lactose (the inducer) is present. The ecological rationale is that glucose is a better source of energy than lactose since its utilization requires two less enzymes. Only after glucose is exhausted are the enzymes for lactose utilization synthesized. The secondary lag during diauxic growth represents the time required for the complete induction of the lac operon and synthesis of the enzymes necessary for lactose utilization (lactose permease and beta-galactosidase). Only then does bacterial growth occur at the expense of lactose. Since the availability of glucose represses the enzymes for lactose utilization, this type of repression became known as catabolite repression or the glucose effect. Glucose is known to repress a large number of inducible enzymes in many different bacteria. Glucose represses the induction of inducible operons by inhibiting the synthesis of cyclic AMP (cAMP), a nucleotide that is required for the initiation of transcription of a large number of inducible enzyme systems including the lac operon. The role of cyclic a cAMP is complicated. cAMP is required to activate an allosteric protein called CAP (catabolite activator protein) which binds to the promoter CAP site and stimulates the binding of RNAp polymerase to the promoter for the initiation of transcription. Thus to efficiently promote gene transcription of the lac operon, not only must lactose be present to inactivate the lac repressor, but cAMP must be available to bind to CAP which binds to DNA to facilitate transcription. In the presence of glucose, adenylate cyclase (AC) activity is blocked. AC is required to synthesize cAMP from ATP. Therefore, if cAMP levels are low, CAP is inactive and transcription does not occur. In the absence of glucose, cAMP levels are high, CAP is activated by cAMP, and transcription occurs (in the presence of lactose). Many positively controlled promoters, such as the lac promoter, are not fully functional in the presence of RNAp alone and require activation by CAP. CAP is encoded by a separate Regulatory gene, and is present in constitutive levels. CAP is active only in the presence of cAMP. The binding of cAMP to CAP causes a conformational change in the protein allowing it to bind to the promoter near the RNAp binding site. CAP can apparently interact with RNAp to increase operon transcription about 50-fold. Positive control of the lac operon is illustrated in Figure10. Figure 10. Catabolite repression is positive control of the lac operon. The effect is an increase in the rate of transcription. As a form of catabolite repression, the glucose effect serves a useful function in bacteria: it requires the cells to use the best available source of energy. For many bacteria, glucose is the most common and readily utilizable substrate for growth. Thus, it inhibits indirectly the synthesis of enzymes that metabolize poorer sources of energy.  !(Y'=0&`Z`ZE`Zt5*"g6Y`Z^<   l $"h Je&F H   ,   Vk 4m7]%k6b/ 5 + & G EH  0޽h ? ̙33. 0  ~(  ^  S T     c $?T @   tR{|[WT;OW 0  & H  0޽h ? ̙33  0 !h(  ^  S T     c $PT @   ^*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. LCG95L@_=H  0޽h ? ̙33  0 y q !  (   ^  S T    k  c $ T @    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> jkX 0pxH  0޽h ? ̙33 0 ! ((   ^  S T      c $  T @     H  0޽h ? ̙33 0 !( ((  ( ^ ( S T    ( c $\|T @    H ( 0޽h ? ̙33 0 0"8 ((  8 ^ 8 S T   | 8 c $p|T @  |  H 8 0޽h ? ̙33 0 xP"@ (  @ ^ @ S T   |r @ c $T @  |  Catabolite Repression Enzyme Induction is still considered a form of negative control because the effect of the regulatory molecule (the active repressor) is to decrease or downregulate transcription. Catabolite repression is a type of positive control of transcription, since a regulatory protein affects an increase (upregulation) in the rate of transcription of an operon. The process was discovered in E. coli and was originally referred to as the glucose effect because it was found that glucose repressed the synthesis of certain inducible enzymes, even though the inducer of the pathway was present in the environment. The discovery was made during study of the regulation of lac operon in E. coli. Since glucose is degraded by constitutive enzymes and lactose is initially degraded by inducible enzymes, what would happen if the bacterium was grown in limiting amounts of glucose and lactose? A plot of the bacterial growth rate resulted in a diauxic growth curve which showed two distinct phases of active growth (Figure 9). During the first phase of exponential growth, the bacteria utilize glucose as a source of energy until all the glucose is exhausted. Then, after a secondary lag phase, the lactose is utilized during a second stage of exponential growth. Figure 9. The Diauxic Growth Curve of E. coli grown in limiting concentrations of a mixture of glucose and lactose During the period of glucose utilization, lactose is not utilized because the cells are unable to transport and cleave the disaccharide lactose. Glucose is always metabolized first in preference to other sugars. Only after glucose is completely utilized is lactose degraded. The lactose operon is repressed even though lactose (the inducer) is present. The ecological rationale is that glucose is a better source of energy than lactose since its utilization requires two less enzymes. Only after glucose is exhausted are the enzymes for lactose utilization synthesized. The secondary lag during diauxic growth represents the time required for the complete induction of the lac operon and synthesis of the enzymes necessary for lactose utilization (lactose permease and beta-galactosidase). Only then does bacterial growth occur at the expense of lactose. Since the availability of glucose represses the enzymes for lactose utilization, this type of repression became known as catabolite repression or the glucose effect. Glucose is known to repress a large number of inducible enzymes in many different bacteria. Glucose represses the induction of inducible operons by inhibiting the synthesis of cyclic AMP (cAMP), a nucleotide that is required for the initiation of transcription of a large number of inducible enzyme systems including the lac operon. The role of cyclic a cAMP is complicated. cAMP is required to activate an allosteric protein called CAP (catabolite activator protein) which binds to the promoter CAP site and stimulates the binding of RNAp polymerase to the promoter for the initiation of transcription. Thus to efficiently promote gene transcription of the lac operon, not only must lactose be present to inactivate the lac repressor, but cAMP must be available to bind to CAP which binds to DNA to facilitate transcription. In the presence of glucose, adenylate cyclase (AC) activity is blocked. AC is required to synthesize cAMP from ATP. Therefore, if cAMP levels are low, CAP is inactive and transcription does not occur. In the absence of glucose, cAMP levels are high, CAP is activated by cAMP, and transcription occurs (in the presence of lactose). Many positively controlled promoters, such as the lac promoter, are not fully functional in the presence of RNAp alone and require activation by CAP. CAP is encoded by a separate Regulatory gene, and is present in constitutive levels. CAP is active only in the presence of cAMP. The binding of cAMP to CAP causes a conformational change in the protein allowing it to bind to the promoter near the RNAp binding site. CAP can apparently interact with RNAp to increase operon transcription about 50-fold. Positive control of the lac operon is illustrated in Figure10. Figure 10. Catabolite repression is positive control of the lac operon. The effect is an increase in the rate of transcription. As a form of catabolite repression, the glucose effect serves a useful function in bacteria: it requires the cells to use the best available source of energy. For many bacteria, glucose is the most common and readily utilizable substrate for growth. Thus, it inhibits indirectly the synthesis of enzymes that metabolize poorer sources of energy.  !(Y'=0&`Z`ZE`Zt5*"g6Y`Z^<   l $"h Je&F H   ,   Vk 4m7]%k6b/ 5 + & G EH @ 0޽h ? ̙33l 0 ,$"T (  T ^ T S T    T c $<TPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPPQ @   faIN O_uirHQ)R(usX-N[f)R(uvn dk:g6RO_uir(WNUOAQsX-NNg'Y^A~k0 42  4H T 0޽h ? ̙33 0 RJ#p (  p ^ p S T   D p c $l T @   lSo let's look at the lac operon in action. When lactose is present, it acts as an inducer of the operon. It enters the cell and binds to the Lac repressor, inducing a conformational change that allows the repressor to fall off the DNA. Now the RNA polymerase is free to move along the DNA and RNA can be made from the three genes. Lactose can now be metabolized. P 2 kH p 0޽h ? ̙33 0 0#x '(  x ^ x S T    x c $` T @   =When the inducer (lactose) is removed, the repressor returns to its original conformation and binds to the DNA, so that RNA polymerase can no longer get past the promoter. No RNA and no protein is made. Note that RNA polymerase can still bind to the promoter though it is unable to move past it. That means that when the cell is ready to use the operon, RNA polymerase is already there and waiting to begin transcription; the promoter doesn't have to wait for the holoenzyme to bind. We could say that the operon is primed for transcription upon the addition of lactose. $>3k S v B =H x 0޽h ? ̙33 0 P# ((   ^  S T     c $ T @    H  0޽h ? ̙33 0 # ((   ^  S T     c $$jT @    H  0޽h ? ̙33 0  $ <(   ^  S T      c $H*A T @b   JB___PPT9$ 4. 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"SystemX9-1_GB2312-. >2 A ΢Ĵл .---$ % nn% %--'1_GB2312-. 2 , ³´л.-1_GB2312-. 12 ,*ڻϸеĸַֽл.-1_GB2312-.  2 2.-@Times New Roman-. 2 2 catabolism.-1_GB2312-.  2 2,.-1_GB2312-. 2 22 ͺϳɴл.-@Times New Roman-. 2 2S anabolismm.-1_GB2312-.  2 2i.-1_GB2312-. 2 2oܺ͡.-1_GB2312-. 2 ;3³´л.-@Times New Roman-.  2 ;L= .-1_GB2312-. 2 ;Oֽл.-@Times New Roman-.  2 ;g+ .-1_GB2312-. 2 ;jϳɴл.-1_GB2312-. 2 E ֽл.-1_GB2312-. @2 E*&ָӵлֽͨлøϵĴ.-1_GB2312-. 42 L򵥷ӡᣨ.-@Times New Roman-.  2 L`ATP.-1_GB2312-.  2 Li.-1_GB2312-. 2 Lo ʽ.-1_GB2312-. 2 Tԭá.-1_GB2312-. 2 ] ϳɴл.-1_GB2312-. =2 ]*$ָںϳɴлøϵĴ߻£ɼС.-1_GB2312-.  2 dӡ.-@Times New Roman-.  2 dATP.-1_GB2312-. C2 d$(ʽͻԭһϳɸӵĴӵ.-1_GB2312-. 2 m̡.-՜.+,D՜.+,H((     l'在屏幕上显示天津商学院9=y/9 CTimes New Roman宋体Tahoma Wingdings楷体_GB2312 Wingdings 2Arial隶书仿宋_GB2312Symbol CG Times 华文楷体 华文行楷Georgia Arial BlackWP Greek CenturyBlends 位图图像 BMP 图象第五章 微生物的代谢 幻灯片 2第一节 代谢概论FFigure 1. The relationship between catabolism and anabolism in a cell 幻灯片 5 幻灯片 6 幻灯片 7#第二节 微生物的能量代谢微生物氧化的形式生物氧化的功能:生物氧化的过程(一、底物脱氢的四条主要途径.一、化能异养微的生物氧化和产能(一)EMP途径 幻灯片 15EMP途径特点: 幻灯片 17 幻灯片 18EMP途径关键步骤葡萄糖激活的方式 幻灯片 21 幻灯片 22 幻灯片 23 幻灯片 24EMP途径的生理功能O(二) HMP途径 (戊糖磷酸途径) (Hexose Monophophate Pathway) 幻灯片 27 幻灯片 28(HMP途径降解葡萄糖的三个阶段HMP途径关键步骤:不完全HMP途径的总反应完全HMP途径的总反应HMP途径的重要意义 幻灯片 34(三)ED途径 ED途径 ED途径 ED途径ED途径的特点ED途径的总反应ED途径的总反应(续) 幻灯片 42 幻灯片 43(四)磷酸酮解途径磷酸戊糖酮解途径#磷酸戊糖酮解途径的特点:磷酸己糖解酮途径%磷酸己糖酮解途径的特点: 幻灯片 49 幻灯片 50 幻灯片 51 幻灯片 52 幻灯片 53 反应特点:这是TCA中第一次氧化作用,被异柠檬酸脱氢酶催化。 该酶是TCA中第二个调节酶,当细胞在高能状态时(即ATP/ADP,NADH/NAD+比值高),酶活性被抑制;在低能状态时被激活。这是TCA循环中第二个氧化脱羧反应,由α-酮戊二酸脱氢酶系所催化。 该酶与丙酮酸脱氢酶系的结构组成相似。酶的催化活性受其产物NADH、琥珀酰CoA和Ca2+抑制;细胞高能荷时,ATP、GTP也可反馈抑制酶的活性。 琥珀酰CoA合成酶催化琥珀酰CoA的硫酯键水解,使GDP磷酸化为GTP,(GTP的作用:[1]在二磷酸核苷激酶作用下,推动ADP生成ATP;[2]用于蛋白质的合成。) 该步反应是TCA中唯一底物水平磷酸化产能的反应。 琥珀酸脱氢酶是第三个氧化还原反应。 该酶构成:分子量100000(二亚基),酶的辅酶是FAD;酶直接与呼吸链联系,将FADH2交给酶的铁硫中心进入呼吸链。 该步反应是第四次氧化还原反应,由L-苹果酸脱氢酶催化,NAD是氢的受体。 在生理情况下,该反应是向右进行的。三羧酸循环的枢纽位置.TCA循环在微生物代谢中的枢纽地位 幻灯片 61 幻灯片 62TCA循环的重要特点1葡萄糖经不同脱氢途径后的产能效率%二、递氢、受氢和ATP的产生 幻灯片 66(呼吸、无氧呼吸和发酵示意图'Mechanisms of energy release: overview1、有氧呼吸Figure. Aerobic respiration电子传递与氧化呼吸链 幻灯片 72 幻灯片 73%微生物中重要的呼吸链组分 幻灯片 75典型的呼吸链 幻灯片 77&Figure. The Structure of NAD and NADP 幻灯片 79 幻灯片 80 幻灯片 81 幻灯片 82 幻灯片 83 幻灯片 84 幻灯片 85细胞色素系统 幻灯片 87 幻灯片 88原核生物呼吸链的特点ATP的结构和生成Figure. The structure of ATP 幻灯片 92 幻灯片 93 幻灯片 94 幻灯片 95底物水平磷酸化举例:电子传递磷酸化 幻灯片 98氧化磷酸化产能机制幻灯片 100幻灯片 101幻灯片 102电子传递磷酸化举例2、无氧呼吸(无氧呼吸(anaerobic respiration)幻灯片 106幻灯片 107幻灯片 108幻灯片 109幻灯片 110幻灯片 1113、发酵作用幻灯片 113幻灯片 114幻灯片 115幻灯片 116幻灯片 117幻灯片 118①酵母菌的乙醇发酵:幻灯片 120%巴斯德效应(The Pasteur effect )巴斯德效应(续)②细菌的乙醇发酵%利用Z.mobilis等细菌生产酒精(二)2,3-丁二醇发酵 鉴别肠道细菌的V.P.试验=鉴别肠道细菌的产酸产气、甲基红(M.R)试验IMViC试验:(三)乳酸发酵 ①同型乳酸发酵-EMP途径2②通过HMP途径的发酵-异型乳酸发酵:幻灯片 132磷酸戊糖酮解途径幻灯片 134(三)混合酸发酵.(4)由氨基酸发酵产能—Stickland反应幻灯片 137幻灯片 138底物水平磷酸化举例:(二、自养微生物产ATP和还原力幻灯片 1414无机底物脱氢后电子进入呼吸链的部位幻灯片 143幻灯片 144幻灯片 145幻灯片 146幻灯片 1472.非循环光合磷酸化'3.嗜盐菌紫膜的光介导ATP合成幻灯片 150幻灯片 1515 第二节 分解代谢和合成代谢的联系二、代谢物回补途径+CO2固定作用补充TCA环的中间产物乙醛酸循环乙醛酸循环幻灯片 157,第三节 微生物独特合成代谢举例"一、自养微生物的CO2固定T(二)厌氧乙酰COA途径 (三)逆向TCA循环 (四)羟基丙酸途径幻灯片 161幻灯片 162幻灯片 163二、生物固氮幻灯片 165幻灯片 166幻灯片 167幻灯片 168幻灯片 169(二)固氮的生化机制幻灯片 171幻灯片 1723.固氮的生化途径幻灯片 1744.固氮酶的产氢反应幻灯片 176幻灯片 177幻灯片 178幻灯片 179幻灯片 180幻灯片 181幻灯片 182幻灯片 183幻灯片 184幻灯片 1857三、微生物结构大分子——肽聚糖的合成幻灯片 187幻灯片 188(第一阶段:在细胞质中的合成8由葡萄糖合成N-乙酰葡糖胺和N-乙酰胞壁酸“Park”核苷酸的合成(第二阶段:在细胞膜中的合成肽聚糖单体的合成+肽聚糖单体的合成——细菌萜醇(第三阶段:在细胞膜外的合成幻灯片 196幻灯片 197幻灯片 198:肽聚糖的生物合成与某些抗生素的作用机制幻灯片 200幻灯片 2012第四节 微生物的代谢调控与发酵生产.一、微生物代谢过程中的自我调节(☆微生物自我调节代谢的方式幻灯片 205幻灯片 206幻灯片 207K☆Possible points for Regulation of Various Metabolic Control Mechanisms 一、酶活性的调节Feedback Inhibition(二)反馈抑制的类型&(1)同功酶调节——isoenzyme幻灯片 213天冬氨酸族=(2)协同反馈抑制——concerted feedback inhibition- 天冬氨酸族 (谷氨酸棒杆菌)?(3)合作反馈抑制——cooperative feedback inhibition>(4)积累反馈抑制——cumulative feedback inhibition=积累反馈抑制——E.coli谷氨酰胺合成酶的调节@(5)顺序反馈抑制——sequential feedback inhibition(6)平衡合成(7)代谢互锁(8)优先合成(三)酶活力调节的机制 变构酶二、酶合成的调节$(一)酶合成调节的类型-22.阻遏(repression):幻灯片 22912.降解产物的阻遏Catabolite Repression :3.酶合成调节的机制Structure of a typical operon 一些重要术语幻灯片 234幻灯片 235幻灯片 236幻灯片 237幻灯片 238幻灯片 239幻灯片 240幻灯片 241'酶合成的诱导Enzyme Induction: 幻灯片 243(酶合成的阻遏Enzyme Repression: 幻灯片 245终产物的阻遏The Tryptophan Operon12.降解产物的阻遏Catabolite Repression :幻灯片 249幻灯片 250幻灯片 251幻灯片 252幻灯片 253幻灯片 254幻灯片 255幻灯片 256幻灯片 257When lactose is present&When the inducer (lactose) is removed,When levels of glucose in the cell are high总结=三、代谢的人工控制及其在发酵工业中的应用1.遗传学方法幻灯片 264幻灯片 265幻灯片 266&1.2 抗反馈控制突变株的应用幻灯片 268.1.3选育组成型突变株和超产突变株(二)生物化学方法"(三)控制细胞膜渗透性柠檬酸发酵谷氨酸发酵 本章小结 复习题二、问答题$(二)分解途径的代谢调控:2.阻遏(repression):"(二)酶合成调节的机制Structure of a typical operon 一些重要术语幻灯片 282*E.coli 乳糖操纵子学说(负调节)幻灯片 284'酶合成的诱导Enzyme Induction: CTable 3. Functional and regulatory components of the lac operon  幻灯片 287(酶合成的阻遏Enzyme Repression: 幻灯片 289终产物的阻遏The Tryptophan Operon幻灯片 292幻灯片 293乳糖操纵子—1乳糖操纵子—2乳糖操纵子—3幻灯片 297幻灯片 298When lactose is present&When the inducer (lactose) is removed,When levels of glucose in the cell are high总结$(二)分解途径的代谢调控: 已用的字体演示文稿设计模板嵌入 OLE 服务器幻灯片标题/ 8@ _PID_HLINKSA@334,5,a|o;mve_ 335,6,xxg|ov 284,7,{^opGr 7..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[333,18,a|o;mve_..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[297,17,ED_v;`S^323,44,_WsNx(YN)Su_..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[..\ߘT_uirf[,gy .ppt -1,1,ߘT_uirf[/http://www.bjkp.gov.cn/bjkpzc/kpxx/53341.shtmlJhttp://courseware.ecnudec.com/zsb/zsw/zsw04/zsw065/zsw06503/zsw065030.htm5http://gene.tmmu.com.cn/biosoft/mobio/2.6/J02_fm.htmKhttp://www.labmedicine.cn/lab_cn/_resources/images/pics/08/Charpter_08.htm7http://202.116.33.235/wswx/info_Show.asp?ArticleID=147347,8,Z|USSOvTb9http://esg-www.mit.edu:8001/cgi-bin/biosearch.pl?lactose?http://www.foodmate.net/ddimg/uploadimg/20050612/121317254.jpgQhttp://tbxt.educast.cn/GZ03-04-1/03/XK26_NJ12/ZY20030819095413390/TPJJ_ZOOM.html)http://www.0-100.com/200406/ca455629.htm+http://202.116.64.20/wjf/7/course/c741.htmKhttp://www.labmedicine.cn/lab_cn/_resources/images/pics/08/Charpter_08.htmKhttp://www.labmedicine.cn/lab_cn/_resources/images/pics/08/Large/08-18.jpgGhttp://etc.lyac.edu.cn/courseware/03_04shengwuhuaxue/ch05/section2.htm.http://www.foodmate.net/4images/3/1/6276.html.http://www.foodmate.net/4images/3/1/6285.htmlDhttp:/_8=0wuwu  $"a|o;mve_ % xxg|ov( EMP_h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[rxMOVVP Paint.Picture0MOVVPzw BMP Va Paint.Picture0 BMP Vazx BMP Va Paint.Picture0 BMP Vaz BMP Va Paint.Picture0 BMP Va%hy ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[za BMP Va Paint.Picture0 BMP Vazb BMP Va Paint.Picture0 BMP VaEDWDYNh ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[h\ ߘT_uirf["..\ߘT_uirf[,gy .ppt0-1,1,ߘT_uirf[b\http://www.bjkp.gov.cn/bjkpzc/kpxx/53341.shtml\http://www.bjkp.gov.cn/bjkpzc/kpxx/53341.shtml@n999:::::G:;X;<y<=y==|>>y??{@@}AA}BBxCC~DElEE[FF}GG}HH~IIhJJ~KLYLLNMMyNN|OP{PP|QQzRRS  !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~                           ! 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