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"Mutation, genetic diversity"%"Mutation, genetic diversity"&"Mutation, genetic diversity"'"Mutation, genetic diversity"E=JDE  undocumented settings in DynaFit&&  "Selection"  0Basic Differential Evolution Algorithm - Summary11+1+Example: DNA + clamp / clamp loader complex,,/22Example: DynaFit script for Differential Evolution33'.Example: Initial population(/!Example: The evolutionary process"",0Example: Final population13bExample: The  fittest member of final population22452Example: Comparison of DE and regular data fitting33?;)Significant disadvantage of DE: very slow**C<,Example of Differential Evolution in DynaFit--762Example: Systematic scan of many initial estimates3397DExample: Systematic scan  Phase 1##;8DExample: Systematic scan  Phase 2##=9DExample: Systematic scan  Phase 3##@:Summary and conclusions/~!"$)*-.0268:<>ABDF   0` ̙33` ` ff3333f` 333MMM` f` f` 3>?" dd@,|?" dd@   " @ ` n?" dd@   @@``PR    @ ` ` p>> <4(    6#L  P L  T Click to edit Master title style! !  0UL   L  RClick to edit Master text styles Second level Third level Fourth level Fifth level!     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This initial estimate must be close enough to the  true values. The solution: Use a data-fitting method that does not depend on initial estimates. An implementation: The Differential Evolution algorithm (Price et al., 2005). An example: Kinetics of forked DNA binding to the protein-protein complex formed by DNA-polymerase sliding clamp (gp45) and clamp loader (gp44/62). 3+  4   3   3      3   A      @`H P 0޽h ? ̙33y___PPT10Y+D='  = @B +g  0L0 vn`') (   x  c $#I `  I  XB  0D`X  0A?`m  <%I   +SELECT AMONG POSSIBLE MOLECULAR MECHANISMS <, 3 33dB  <D8cdB  <D8c-  <,I  A concentration  <0I } Fsignal$R2  s *MR2  s *}R2  s *A=R2  s *-1R2  s *1}R2  s *a mX  0f{   6H7I 5( P  <computer  R2  s *I R2  s *N ^B  6Dp& M & ^B  6DV V^B  6DpS ! ^B  6DjJS XdB  <Df)A  dB  <Df)D ^B  6Df)0p p0^B  6Df)D1 DdB  <Df)P   0(I    OSelect most plausible model  < I a W competitive ?. X  0f_ R ! s *f@R " s *fR # s *A]M $ <]I ${ U mechanism B.  % <hI @]  C mechanism C  & <TI  U mechanism A.  ( <LlI 0 P  YEXPERIMENTAL DATA, ff ) 0I    h A VARIETY OF POSSIBLE MECHANISMS,!  ffH  0޽h ? ̙33y___PPT10Y+D='  = @B +   0   "@  ( ?j, @ r @ S 0I ``  I  XB @ 0D`X @ 0A?`m @ <X s *b!R2 ?X s *"R2 @X s *qR2 AX s *qR2 BX s *X CX 0f^R DX <0np]  YEXPERIMENTAL DATA, ffb EX Tp(n?"` ,$D  0 *The Initial Estimate Problem: Estimated parameters most be  close enough . How can we guess them? 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Enzymol., 467, 247-280p+ ,  Tn ?0   :2009  T@q ?"` Y ,$D  0 aDynaFit implements the Differential Evolution algorithm for global sum-of-squares minimization.@b   4 H  0޽h ? ̙33aY___PPT109+F:D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-m6Bbox(in)*<3<* D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-m6Bbox(in)*<3<* +8+0+ 0 +  0 :2 (   r  S I `  I  XB  0D`X  0A?`m^B  @ 6DԔ@   <py  Jgene*  <} pL9 ;BIOLOGY  <y p  <COMPUTER XB  0D`  < $a O...AAGTCG...GTAACCGG...3  <  yp  Z& four-letter alphabet variable length''XB  0D@PXB  0D@PdB  <D  <H | ? "keratin" fl p   p ,$D 0   <譅  k' sequence of bits representing a number(('   < 0 0  X ...01110011000001101101110011...!!3   < 0  p  f2 two letter alphabet fixed length (16 or 32 bits)33`B  0D @ P`B  0D @ PlB   <D     <В @   \"KM"<fff`B  0D@@@PlB   <D @   <p P "kcat"<fffH  0޽h ? ̙33~___PPT10^+gDB'  = @B D' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(+  0 ## T(   r  S  `   XB  0D`X  0A?`m^B  @ 6DԔ@   < v Ngenotype*   <df pL9 ;BIOLOGY  <p# p  <COMPUTER XB  0D`^B  6D`@@   <F Sa U...AAGTCGGTTCGGAAGTCGGTTTA... 3^  6jJt`  <K  P  Okeratin,^B  @ 6D`pp ^  6jJD `  <4    m oncoprotein,    <  p  A phenotype z      ,$D 0   <  s/ particular combination of all model parameters(0/z   6A  ?? 0    z \ `0    \ `0 ,$D 0fB   6Dl ^l @   < \ `_ \$011010110110011110011010001111101101%% 3fB  B 6D ` 0 fB   6D ` f   6jJ.`f   6jJ `f   6jJz  `X2  0 P z 0 pT     0 pT ,$D 0   <X | 0 N  KM=4.56d   <5  @q    Vmax=1.23b  !  <( } Kis=78.9d `2 "  00 p  #  <  T  Lfull set of parametersH  0޽h ? ̙33___PPT10+}yD'  = @B Du' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(+0  0 RJ ,, (   r  S  `   XB  0D`X  0A?`m^B  @ 6DԔ@   < v Rgenotype.   <`S pL9 ;BIOLOGY  <x p  <COMPUTER XB  0D`^B  6D`@@   <k Sa U...AAGTCGGTTCGGAAGTCGGTTTA... ^  6jJt`  <P:  P  ?keratin^B  @ 6D`pp ^  6jJD `  <ԅ    ] oncoprotein  ^2  6jJ P R2  s *3f  ^B  @ 6DԔ `& ^B  6DԔ PP ^B  6DԔP  ^B  6DԔ P ^B  6DԔP P ^B  6DԔ `P ^B  6DԔ ` ^B  6DԔ ` R2  s *  ^B  6DjJP     BCPDE4FjJ P H@ 0hH0 x @    @8    BCPDE4FjJ P H@ 0hH0 x @    ^p R2  s *FR2  s *F@v^B ! 6D8c ` ^B " 6D8c ` ^B # 6D8c <z ~ $  ~,$D 0 %  <…  6 FITNESS: agreement between the data and the modelJ7. &  <A  ??` p~  f2 '  6jJP  fB (  6D\|.fB ) B 6Dp@`B *  0Dvv`B +  0D>pp , <`W  %  * FITNESS: "agreement" with the environment<+#H  0޽h ? ̙33~___PPT10^+놢mDB'  = @B D' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*$ %(+}:  0 c4[4@ggL 3( , L r L S Ĕc`  c XB L 0D`X L 0A?`m^B L @ 6DԔ@  L <TcpL9 ;BIOLOGY L <̅ p  <COMPUTER XB L 0D`z  `  L   `,$D 0`2  L  03 P Z2  L  s *33@  fB  L B 6D3ԔP  fB  L  6D3Ԕ *0fB L  6D3Ԕ*0`fB L 6D3Ԕ*0fB L 6D3Ԕ*00fB L  6D3Ԕ @ fB L 6D3Ԕ @ fB L 6D3Ԕ @ Z2 L  s *30 ` fB L B 6DjJz ~ fB L 6D38cP @ fB L B 6D38c  @ fB L B 6D38c @P  L  <0c3P   S low fitness,  3  3fB L  6D fB L  6D Z2 L  s * J Z Z2 L  s *p * Z2 L  s * : D Z2 L  s * Z2  L  s *P 0 j Z2 !L  s *@ z: Z2 "L  s *  Z2 #L  s * @ zfB $L  6D38c @ %L  <$х  v  jVmax033 &L  <ةc   NKM033 'L  0c   iKis033z P`8  (L  P`8 ,$D 0`2 )L  0pP Z2 *L  s *  fB +L B 6DԔp  fB ,L  6DԔ` `T fB -L  6DԔp 8 fB .L 6DԔp  fB /L 6DԔp  Z2 0L  s * P fB 1L 6DjJ   2L   BCPDE4FjJ P H@ 0hH0 x @    8   3L   BCPDE4FjJ P H@ 0hH0 x @    p fB 4L 6D8cP P  5L  <c   Vmedium fitness,  Z2 6L  s *T  fB 7L  6D`P`p fB 8L  6D`p p B2 9L   4cBuENG)RH5ISQ8c ` #Ju` #JuS`T#JuS`Ty`   > Z2 :L  s * * Z2 ;L  s *  Z Z2 L  s * :: Z2 ?L  s ** Z2 @L  s *j  Z2 AL  s *n J  BL  <\c V  jVmax0 CL  < c   NKM0 DL  0Xc" `  iKis0z 0  EL  0 ,$ 0`2 FL  0D|Z2 GL  s *3fDpfB HL B 6DԔdfB IL  6DԔT fB JL  6DԔ 0 fB KL 6DԔ fB LL 6DԔ  fB ML  6DԔdfB NL 6DԔdfB OL 6DԔdZ2 PL  s *4dfB QL 6DjJ"0` RL   BCPDE4FjJ P H@ 0hH0 x @     |p SL   BCPDE4FjJ P H@ 0hH0 x @    4>fB TL 6D8cpP`fB UL 6D8cP`fB VL 6D8cpP`Z2 WL  s *@ Z2 XL  s * P YL  <cDR T high fitness,    fB ZL  6D  fB [L  6D B2 \L   !cBENG)RH ISQ8c ` #J"` #J"S`T#J"S`T # > Z2 ]L  s * * ZZ2 ^L  s *P  *Z2 _L  s *  DZ2 `L  s *p  Z2 aL  s *0 0j jZ2 bL  s * Z :Z2 cL  s *  Z2 dL  s * @ z eL  <&c  p jVmax0 fL  <H+c  p NKM0 gL  0/c .p iKis0H L 0޽h ?/ 9L \L  ̙33___PPT10+Nz'D~'  = @B D9' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*(L %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* L %(+  0 ` w A(   ~ w H A# ?"`"] T r  S D4c`  c XB  0D`X  0A?`=XB n 0Do_XB o 0D p <?c+p` :number of population members per optimized model parameter<;  q <c+  3number of population members per order of magnitude<4 ^ s 6jJ  ^ t 6jJ @XB u 0D_ _XB v 0D H  0޽h ? ̙33y___PPT10Y+D='  = @B +H  0 h@`@` 0 ?(   r  S c`  c XB  0D`X  0A?`m^B  @ 6DԔ@   <DcpL9 ;BIOLOGY  <,Icp  <COMPUTER XB  0D`N F 0    Pp`2   0D|Z2   s *3fDpfB  B 6DԔdfB   6DԔT fB   6DԔ 0 fB  6DԔ fB  6DԔ  fB   6DԔdfB  6DԔdfB  6DԔdZ2   s *4dfB  6DjJ"0`    BCPDE4FjJ P H@ 0hH0 x @     |p    BCPDE4FjJ P H@ 0hH0 x @    4>fB  6D8cpP`fB  6D8cP`fB  6D8cpP`Z2   s *@ Z2   s * PN F `     `2   0   Z2   s *  p0 fB  B 6DԔ@ :fB !  6DԔ0 `pfB "  6DԔ`pfB # 6DԔ`@pfB $ 6DԔ`ppfB %  6DԔ@4dfB & 6DԔ@4fB ' 6DԔ@44Z2 (  s *@ @p fB ) 6DjJ p  *   BCPDE4FjJ P H@ 0hH0 x @     X  +   BCPDE4FjJ P H@ 0hH0 x @      fB , 6D8c , fB - 6D8c , fB . 6D8c , Z2 /  s *`  Z2 0  s *v  X 1 0  8 `  `f2 6  6#`2 7  0aGlB 8 B <DԔXwlB 9  <DԔlB :  <DԔlB ; <DԔlB < <DԔlB =  <DԔXlB > <DԔXlB ? <DԔX`2 @  0BQ lB A <DjJ<[) B  # BCPDE4FjJ P H@ 0hH0 x @    oY C  # BCPDE4FjJ P H@ 0hH0 x @    BylB D <D8c)lB E <D8c)lB F <D8c)`2 G  06`Fo`2 H  0corN F 0  I  0 p `2 J  0D|Z2 K  s *3fDpfB L B 6DԔdfB M  6DԔT fB N  6DԔ 0 fB O 6DԔ fB P 6DԔ  fB Q  6DԔdfB R 6DԔdfB S 6DԔdZ2 T  s *4dfB U 6DjJ"0` V   BCPDE4FjJ P H@ 0hH0 x @     |p W   BCPDE4FjJ P H@ 0hH0 x @    4>fB X 6D8cpP`fB Y 6D8cP`fB Z 6D8cpP`Z2 [  s *@ Z2 \  s * P 8 p`   p` f2 ^  6s `2 _  0pTlB ` B <DԔiG lB a  <DԔ T lB b  <DԔT ` lB c <DԔH T lB d <DԔT T lB e  <DԔiE Q lB f <DԔi9 E lB g <DԔiE E `2 h  09FlB i <DjJ9T j  # BCPDE4FjJ P H@ 0hH0 x @    Mp k  # BCPDE4FjJ P H@ 0hH0 x @    9$pxlB l <D8cplB m <D8c|lB n <D8cp`2 o  0.<`2 p  0Vd( l X Dv   DX v ,$D 0 2  <hjc HV [#01101011011001111001101 00011111011$$ 3  <4oc D [#01101011011001111001101 11100011011$$f`B 4  0D@@ q  <scX (  :mother  r  <xc[ c  :father lB t  <D00  u  Buc` n  #"sexual mating" probability pcross8$  v  Bc $ v  p&01101011011001111001101 11100011011.' f z  Bc` p   7child  |  <Pc6D ^random crossover point,el  B `   B ` ,$D  0f }  6> B f   6> B  f   6>B `    <ĺc  -  pVmax6       <c   TKM6       < }c   oKis6    H  0޽h ? ̙33___PPT10+.{VD' = @B Ds' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<* D' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<* +R  0 JJ J(   r  S `c`  c XB  0D`X  0A?`m^B  @ 6DԔ@   <cpL9 ;BIOLOGY  <cp  <COMPUTER XB  0D`X2  0$BR2  s *~ig^B  @ 6DԔ>F ^B  6DԔBJ^B  6DԔJEp^B  6DԔ%EJ^B  6DԔJEJ^B  6DԔkA^B  6DԔk^B  6DԔkR2  s *C^B  6DjJZ:  # BCPDE4FjJ P H@ 0hH0 x @    vU~  # BCPDE4FjJ P H@ 0hH0 x @    ^B  6D8c~-:^B  6D8c-:^B  6D8c~-:R2  s *P(vR2  s *vlX2  0 # R2  s * a\ ^B  @ 6DԔ B ^B  6DԔ  ^B  6DԔ = ^B ! 6DԔ = ^B " 6DԔ = ^B # 6DԔ f ^B $ 6DԔp f ^B % 6DԔ f R2 & s ** 8R ^B ' 6DjJy \  ( # BCPDE4FjJ P H@ 0hH0 x @    J  ) # BCPDE4FjJ P H@ 0hH0 x @     ^B * 6D8c , ^B + 6D8c , ^B , 6D8c , R2 - s *R2 . s *aX / 0 N F 0  D  0 p `2 E  0D|Z2 F  s *3fDpfB G B 6DԔdfB H  6DԔT fB I  6DԔ 0 fB J 6DԔ fB K 6DԔ  fB L  6DԔdfB M 6DԔdfB N 6DԔdZ2 O  s *4dfB P 6DjJ"0` Q   BCPDE4FjJ P H@ 0hH0 x @     |p R   BCPDE4FjJ P H@ 0hH0 x @    4>fB S 6D8cpP`fB T 6D8cP`fB U 6D8cpP`Z2 V  s *@ Z2 W  s * P| z 0  v   ` ,$D 0Z2 w  s *D|Z2 x  s *DpfB y B 6DԔdfB z  6DԔT fB {  6DԔ 0 fB | 6DԔ fB } 6DԔ  fB ~  6DԔdfB  6DԔdfB  6DԔdZ2   s *4dfB  6DjJ"0`    BCPDE4FjJ P H@ 0hH0 x @     |p    BCPDE4FjJ P H@ 0hH0 x @    4>fB  6D8cpP`fB  6D8cP`fB  6D8cpP`Z2   s *@ Z2   s * P l 0 p   p0  ,$D 0   B0  z & 01101011011 001111001101 11100011011@'f%f   B 0 p  8father QT  B `   #  !Tf   6> B f   6> B  f   6>B `    <8   -  pVmax6       <    VKM8        <T    qKis8        B(  A   & 11100111011 001011010101 11001011001@'f%3f   B% 8 L a  ? mutant father jT  B `   #  e ^ f   6> B f   6> B  f   6>B `    <    Vmax(*)F         <  ?  YKM(*)8        <  S  tKis(*)8     lB  B <D  L    <  \ 0  <mutation Z F `   ``2   0#Z2   s *aGfB  B 6DԔXwfB   6DԔfB   6DԔfB  6DԔfB  6DԔfB   6DԔXfB  6DԔXfB  6DԔXZ2   s *BQ fB  6DjJ<[)   # BCPDE4FjJ P H@ 0hH0 x @    oY   # BCPDE4FjJ P H@ 0hH0 x @    ByfB  6D8c)fB  6D8c)fB  6D8c)Z2   s *6`FoZ2   s *corZ F p`    p` `2   0s Z2   s *pTfB  B 6DԔiG fB   6DԔ T fB   6DԔT ` fB  6DԔH T fB  6DԔT T fB   6DԔiE Q fB  6DԔi9 E fB  6DԔiE E Z2   s *9FfB  6DjJ9T   # BCPDE4FjJ P H@ 0hH0 x @    Mp   # BCPDE4FjJ P H@ 0hH0 x @    9$pxfB  6D8cpfB  6D8c|fB  6D8cpZ2   s *.<Z2   s *Vd(l   0    0 ,$D 02   T  jJ?"`  2   T  jJ?"`  0 H  0޽h ? ̙33___PPT10+% D'  = @B Du' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*v %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(+'  0 #6d ( _  d r d S ) `   XB d 0D`X d 0A?`m d <'   ^& 01101011011 001111001101 11100011011''f d <(: - 9aunt #1 b  B `  d # 1,$D  0f d  6> B f  d  6> B  f  d  6>B `   d  <,?    iVmax(1)F        d  <E  ?  YKM(1)8       d  <J  S  ZKis(1)8      d < O  g  & 11100111011 001011010101 11001011001@'f%ff d <T  9aunt #2 b  B `  d # )P ,$D  0f d  6> B f d  6> B  f d  6>B `  d  <Y    iVmax(2)F       d  <_  ?  YKM(2)8      d  <d  S  ZKis(2)8     7 *d <Hj  ?THE "DIFFERENTIAL" IN DIFFERENTIAL EVOLUTION ALGORITHM - STEP 1p@ 3  3 3& 3 33" +d < r 7 Compute difference between two randomly chosen  auntie phenotypesCCdB ,d <D P  -d <@w   <subtract  .d <z    & 11100111011 001011010101 11001011001@'f%f /d < C   aaunt #2 minus aunt #10     B {  0d #  ,$D  0f 1d  6> B f 2d  6> B  f 3d  6>B `  4d  <   &  Vmax(2)-Vmax(1)f           5d  <  h   KM(2)-KM(1)f            6d  <  {   Kis(2)-Kis(1)f          H d 0޽h ? ̙33___PPT10+hDh' = @B D#' = @BA?%,( < +O%,( < +Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*d %(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*d D' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*d Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*d %(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*d D' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*d Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*0d %(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*0d D' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*0d +  0 #*l f( Aډ  l r l S  `   XB l 0D`X l 0A?`m l <   p& 01101011011 001111001101 11100011011.'f& l <X ;- 8father }  B `  l # 1,$ 0f l  6> B f  l  6> B  f  l  6>B `   l  <   -  pVmax6      l  <    VKM8       l  <    qKis8      l <l f ,  & 11100111011 001011010101 11001011001@'f%3f l <`  S  ? mutant father 7 l <@  ?THE "DIFFERENTIAL" IN DIFFERENTIAL EVOLUTION ALGORITHM - STEP 2p@ 3  3 3& 3 33< l <  Add weighted difference between two  uncle phenotypes to  father 0C 6dB l <D P  l <<   aadd a fraction of4 l < T  & 11100111011 001011010101 11001011001@'f%f l <P  0  aaunt #2 minus aunt #10     B {  l # = ,$ 0f l  6> B f l  6> B  f  l  6>B `  !l  <h   &  Vmax(2)-Vmax(2)f           "l  <  h   KM(2)-KM(1)f            #l  <5 {   Kis(2)-Kis(1)f            B `  $l #  ",$ 0f %l  6> B f &l  6> B  f 'l  6>B `  (l  < 5  [  Vmax*F       )l  <5   eKM*F       *l  <D5   Kis*F      H l 0޽h ? ̙33y___PPT10Y+D='  = @B +   0   3t U ( _  t r t S L'5`  5 XB t 0D`X t 0A?`m t <D)5 6THE "DIFFERENTIAL" IN DIFFERENTIAL EVOLUTION ALGORITHM^7 3  3 3# 33 $t <35  >KM* = KM + F (KM(1) - KM(2)) &    3     %t <\@56  T"EXAMPLE: Michaelis-Menten equation##x &t  <AX  ?? +  ^B )t @ 6D f ^B *t 6DV V ^B +t 6D ^B ,t 6D -t <L5!  N"father"*   .t <Q5!  V aunt 1"*   /t <V5!  V aunt 2"*   0t <p[5Bk  U"mutant father"* ^B 1t 6D   h  2t <`5 l `  Gweight (fraction)3 3t <Pe50 p y  ? mutation rate H t 0޽h ? ̙33y___PPT10Y+D='  = @B +  0 B : `   (   ~  H A% ?"`ofx  c $l5`  5 XB  0D`X  0A?`=XB  0D |  <n5 0r  probability that  child inherits  father's genes, not  mother's genesdI    XB  0D `  ~  TK5 ?} `,$ 0 These DE tuning constants are  undocumented in the DynaFit distribution.JJ   <H50 Y'fractional difference used in mutations((XB  0D`   <v50 >KM* = KM + F (KM(1) - KM(2)) &    3    ^  6jJr4 ` ^  6jJ4 ` PXB  0D@XB  0D@`@^  6jJ: f ZXB  0D0` 0XB  0D 0XB  0D `   T5 ?00v S!six different mutation strategies""H  0޽h ? ̙33zr___PPT10R+E D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* +8+0+ 0 +,  0 &&@GH o&(   r  S  5`  5 XB  0D`X  0A?`m^B  @ 6DԔ@   <5pL9 ;BIOLOGY  <Į5p  <COMPUTER XB  0D` F 0    0 `2   0D|Z2   s *3fDpfB  B 6DԔdfB   6DԔT fB   6DԔ 0 fB  6DԔ fB  6DԔ  fB   6DԔdfB  6DԔdfB  6DԔdZ2   s *4dfB  6DjJ"0`    BCPDE4FjJ P H@ 0hH0 x @     |p    BCPDE4FjJ P H@ 0hH0 x @    4>fB  6D8cpP`fB  6D8cP`fB  6D8cpP`Z2   s *@ Z2   s * P   <@5DR T high fitness,       <L5^ Tmore likely to breed$Yl  p G  p,$D 0  p    p,$D 0   <`5 ~/ T"0110101101100111100110100011111011## f !  6jJ  0f "  6jJ 0f #  6jJN0 $  <̪5 V  jVmax0 %  <5  NKM0 &  0(5bp iKis0 '  <5 p p0more likely to be carried to the next generation$1- E  <5  Hlow sum of squaresl P f` H P f`,$D 00  f` (   f`,$D 0}  ` )   `,$ 0`2 *  03 P Z2 +  s *33@  fB , B 6D3ԔP  fB -  6D3Ԕ *0fB .  6D3Ԕ*0`fB / 6D3Ԕ*0fB 0 6D3Ԕ*00fB 1  6D3Ԕ @ fB 2 6D3Ԕ @ fB 3 6D3Ԕ @ Z2 4  s *30 ` fB 5 B 6DjJz ~ fB 6 6D38cP @ fB 7 B 6D38c  @ fB 8 B 6D38c @P  9  <L53P   S low fitness,  3  3 :  <5   Tless likely to breed$  @ f. ;   @ f.,$ 0 <  < 5 @ N  T"0000000000111111111111100000000000## f =  63jJ ^ f >  63jJ ^ T f ?  63jJl^   @  <5 &  jVmax033 A  <p5P ^  NKM033 B  0H52 p  iKis033 C  <5 f. p0less likely to be carried to the next generation$1- F  <,ȇ  P S$  Yhigh sum of squares*33H  0޽h ? ̙33___PPT10+aD~'  = @B D9' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*G %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*H %(+  0   ` < (   r  S ч `   XB  0D`X  0A?`m  <} &X 31 Randomly create the initial population (size N) ^4%fl d   d ,$D 0fB  B 6Dd d fB  6Dd lB   <D  0H Z,$D 0 Repeat until almost all population members have very high fitness: 2 Evaluate fitness: sum of squares for all population membersnB3,Cz      p ,$D 0K   <0 L   E5 Natural selection: keep child in gene pool if more fit than mothervFfB   6D` xz 2      ,$D 0   <    @4 Sexual reproduction: random crossover with probability PcrossAff:fB   6D2  iz  P     ,$D 0q   < | P  ?3 Mutation: random gene modification (mutate father, weight F)@  f fB   6D  H  0޽h ? ̙33  ___PPT10 +U GD? '  = @B D ' = @BA?%,( < +O%,( < +DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(+8+0+ 0 +\  0L0 kc`  (      <\ "`   XB  0D`X  0A?`m  <T  DETERMINE ASSOCIATION AND DISSOCIATION RATE CONSTANT IN AN A + B AB SYSTEMdM; 3 3 3 33p  H A ?T Z  T8  ?2` HCourtesy of Senthil Perumal, Penn State University (Steven Benkovic lab),IH>     Tc ?"`` f4  esee Lecture 1 for details0  H  0޽h ? ̙33y___PPT10Y+D='  = @B +   0L0  (      <l "`   XB  0D`X  0A?`m  <n $  r*INSERT A SINGLE LINE IN THE [TASK] SECTION,+* 33p  H A ?0ym   T  jJ?"`  l     y,$D 0x   H A ?    T  jJ?"`p `    T  jJ?"` [rB   B D? p rB  B D? p   TXv  ?p `  C constraints !H  0޽h ? ̙335-___PPT10 +h D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* +0  0 WO (   x  c $ `   XB  0D`X  0A?`mp  H A ?`(   <f  3BOTH RATE CONSTANTS SPAN TWELVE ORDERS OF MAGNITUDE<4 3 33H  0޽h ? ̙33y___PPT10Y+D='  = @B +  0     (   x  c $P `   XB  0D`X  0A?`m~  H A ?"`   <  RSNAPSHOTS OF k1 / k2 CORRELATION DIAGRAM - SPACED BY 10  GENERATIONS F  3 3 3 3 3  3 3  33~  H A ?"`J ~  H A ?"` ~  H A ?"`0~  H A  ?"`s  ~  H A  ?"`s J ~  H A  ?"`s  ~  H A  ?"`s 0   T N  ?p 70   T\6  ?p 810   T@-  ?b P p 820   TX<  ?@.p 830   T0@  ? @ 840   TD  ? @ 850   TG  ? ` N @ 860   T,>  ? @ 870 H  0޽h ? ̙33y___PPT10Y+D='  = @B +   0 um@ (   x  c $ `   XB  0D`X  0A?`m  <  EBOTH RATE CONSTANTS SPAN AT MOST 30% RANGE RELATIVE TO NOMINAL VALUELF 3 3 33p  H A ?` l P  P,$D  0x   H A ? xB  H D1?PP xB   H D1?P* 0* H  0޽h ? ̙33___PPT10+6D|' = @B D7' = @BA?%,( < +O%,( < +Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =+4 8?dCB1+#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<* D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<* +U  0 w o   (   x  c $! `   XB  0D`X  0A?`m   <#  9THIS IS (PRESUMABLY) THE GLOBAL MINIMUM OF SUM-OF-SQUARESL: 3 3 33p  H A ?0  p  H A ?0@ p  H A ?0 @   T  ?Z ] :data jB  B D?PP P   T*  ?P  ;model   T%  ?m ? residuals  Hl  p   p,$D 0x   H A ?     TZ  ?t0 p  Vcompare with  good estimate from Lecture 1,,2   T  >?"`. 2   T  >?"`XH  0޽h ? ̙33~___PPT10^+Y DB'  = @B D' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(+  0 +# 1$ (  $ x $ c $LG`  G XB $ 0D`X $ 0A?`mr $ <XG~ DIFFERENTIAL EVOLUTION (DE) FOUND THE SAME FIT AS THE  GOOD ESTIMATE`F 3 3  3 33pB $ H D8c?0@0jB $ @ B D?4 ]  $ TG ? $ @sum of squares jB $ B D? - 0 ]  $ T0.002308  $ TXG ?4   >0.002354   $ T8G ?xJ L :1.00 !$ TضG ?4J   :1.02 "$ TdG ?T !k1 = 2.2 0.5 k2 = 0.030 0.015^" #$ T{G ?  k1 = 0.2 3.4 k2 = 0.2 0.6^  $$ ZdG ?1@ B  good  %$ ZG ?@  @  bad 3jB *$ B D? @  ,$ TG ? FK  Bk1 = 10-6  10+6 k2 = 10-6  10+6"       -$ TTG ?   J0.002308&  .$ TЪG ? V  61.00  /$ TG ?   !k1 = 2.2 0.5 k2 = 0.030 0.015J"   0$ TG?"` p  K1000 random estimates  1$ ZDG?"`@j  ? lecture 1 H $ 0޽h ? ̙33y___PPT10Y+D='  = @B +  0   0p (  p x p c $G`  G XB p 0D`X p 0A?`m p <G 7DYNAFIT CAN TAKE MULTIPLE DAYS TO RUN A COMPLEX PROBLEM`8 3 3  3 33 !p T`G ?`8  = algorithm  "p TPG ?P  Dcomputation timejB #p B D? @ jB $p @ B D?  M %p TG ?`X D  iLevenberg-Marquardt with two restarts Differential Evolution with four restarts (population size: 1000)Dj+ &p TtaG ?p @ :0.88 sec  'p T G ? F T  ? 12 min 31 sec (p TG ?` 4 j4DynaFit 4.065 on DNA / clamp / clamp loader example:55jB )p B D?``@  *p TG ?p+@ 31 +p TXG ? KP  ;8533 ,p TOG ?Q ? relative timel @ p 0p @ p,$D 0 -p  TxG?"`h p P1 second 1 minute 10 minutes .p  N ?"`  /p  T^G?"` @ H N15 minutes 15 hours 6 daysH p 0޽h ? ̙335-___PPT10 + D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*0p %(D' =-o6Bdissolve*<3<*0p +  0 %0  x (  x x x c $JG`  G XB x 0D`X x 0A?`mp x H A! ?P0  x T  ?  V J. Biol. Chem. 283, 11677 (2007)!!; x T ?"``@  ,$D 0 uCThis took one week of computing on the Linux cluster in Heidelberg.DD~ x H A$ ?"` ppB x H DV?   pB x H DV? @ pB x H DV?pP0p  x H A" ?"` w,$D  0H x 0޽h ? ̙33c[___PPT10;+PՅD'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* x %(D' =-m6Bbox(in)*<3<* x D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*x %(D' =-o6Bdissolve*<3<*x +8+0+x 0 +}  0   *, F (  , x , c $9 `   XB , 0D`X , 0A?`m , <;  MCAREFUL! THIS IS FASTER THAN DIFFERENTIAL EVOLUTION BUT DOES NOT ALWAYS WORK,NM 33p ", H A ?`r  #, T  8c?"` `` $, T  8c?"``  l  0  ',  0 ,$D  0N %,  T(  ?"` 0 D<4___PPT9 hgenerate all possible combinations of rate constants compute initial sum of squares for each combination rank combinations by initial sum of squares select the best N combinations perform a full fit for those N rank results againL  ,    @` &,  T  ?  = ALGORITHM l  CT *,  CT,$D 0rB (,  B D? P  ),  TK  ? CT N7 7 = 49 combinations of kon and kofff(,H , 0޽h ? ̙33og___PPT10G+vD+'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*', %(D' =-m6Bbox(in)*<3<*', D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<**, %(+  0 @< Z(  < x < c $W `   XB < 0D`X < 0A?`m\ < <Do 7 PAFTER EVALUATING THE INITIAL SUM OF SQUARES FOR ALL 49 COMBINATIONS OF k1 and k2Q 3 3, 3 3 3 33~ < H A  ?"`0fH < 0޽h ? ̙33y___PPT10Y+D='  = @B +o  0 P &(  P x P c ${ `   XB P 0D`X P 0A?`m( P <˖  PAFTER RANKING THE INITIAL ESTIMATES AND SELECTING 20 BEST ONES BY SUM OF SQUARESPQ2 3  3 33~ P H A ?"``@H P 0޽h ? ̙33y___PPT10Y+D='  = @B +w  0   X z (  X x X c $` `   XB X 0D`X X 0A?`m X <,ږ c FAFTER PERFORMING FULL REFINEMENT FOR 20 BEST ESTIMATES OUT OF 49 TRIEDPG% 3 3 33~ X H A ?"`g pB X H DԔ?0 pB  X H DԔ? pB  X H DԔ?` pB  X H DԔ? ` pB  X H DԔ?P0pB  X H DԔ?`00PpB X H DԔ?p0`pB X H DԔ? ppB X H DԔ? ` pB X H DԔ?0 0   pB X H DԔ? 0 0 jB X @ B D?` 0 `  X T4  ?   N success zone l   X  ` ;,$D 02 X  T  8c?"` rB X  B D?   R X  T9 ?"` R  DThe best initial estimates do not produce the best refined solution!XE H X 0޽h ? ̙335-___PPT10 +csD'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*X %(D' =-o6Bdissolve*<3<*X +s   0   h * (  h x h c $ `   XB h 0D`X h 0A?`m h Tpc  ?PD D<4___PPT9 Finding good-enough initial estimates is a very difficult problem. One should use system-specific information as much as possible. This includes using the literature and/or general principles for  intelligent guesses. Always use the  Try method in DynaFit to display the initial fit. Make sure that the initial estimate is at least approximately correct. The Differential Evolution algorithm almost always helps. However, it can be excruciatingly slow (running typically for multiple hours). The systematic scan (task = estimate) sometimes helps. However, the  best initial estimates almost never produce the desired solution! 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