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TExample:  Good initial estimate  results++ >Example:  Bad initial estimate   RExample:  Bad initial estimate  results**\Example:  Good vs.  Bad results - comparison//\Example:  Good vs.  Bad results - comparison//VExample:  Good vs.  Bad results - summary,,Summary and conclusions/C     0` ̙33` ` ff3333f` 333MMM` f` f` 3>?" dd@,|?" dd@   " @ ` n?" dd@   @@``PR    @ ` ` p>> <4(    6L  P L  T Click to edit Master title style! !  0L   L  RClick to edit Master text styles Second level Third level Fourth level Fifth level!     S  0L  `` L  X*  0L  `  L  l*(    04L  `  L  `* H  0޽h ? ̙33 Default Design 0 P4 (  4 4 NH y y .    p* a00aa 4 NY y y 3 .   r* a00aad 4 c $ ?   4 4 N[ y y 9 f   RClick to edit Master text styles Second level Third level Fourth level Fifth level!     S  4 Th y y r    p* a00aa 4 Tm y y r3    r* a00aaH 4 0ηo~ ? ̙3380___PPT10.mz$X \x(  \ \ N\ ]e]e      p* AAA \ N ]e]e %     r* AAA  \ T ]e]e     p* AAA \ T` ]e]e %    r* AAAH \ 0ηo~ ? ̙3380___PPT10.nz01 0 0%(  X  0A?`m  <|/  (Petr Kuzmi , Ph.D. BioKin, Ltd. WATERTOWN, MASSACHUSETTS, U.S.A.hB    7p ! H A ? ` ~ " H A ?"` ); $ 6\| p kBinding and Kinetics for Experimental Biologists Lecture 1 Numerical Models for Biomolecular InteractionsNl0  /9 % ZxP| ?5U GI N N O V A T I O N L E C T U R E S (I N N O l E C)LH&    H  0޽h ? ̙33y___PPT10Y+D='  = @B +.  0L0 =5(  x  c $l8L `  L  XB  0D`X  0A?`mU  Th)| ?`   The problem: Traditional equations for fitting biomolecular binding data restrict the experimental design. Typically, at least one component must be present in very large excess. The solution: Abandon algebraic equations entirely. Use iterative numerical models, which can be derived automatically by the computer. An implementation: Software DynaFit. An example: Kinetics of forked DNA binding to the protein-protein complex formed by DNA-polymerase sliding clamp (gp45) and clamp loader (gp44/62). 3   '  L 3+     3     3   A      @`H  0޽h ? ̙33y___PPT10Y+D='  = @B +  0L0   6 @ (      <L "`  L  XB  0D`X  0A?`mp * H A/ ?   + T{| ? S c  TCold Spring Harbor Laboratory Press Cold Spring Harbor, NY, 2007 ISBN-10: 0879697369UU " , T| ?pD  `A typical  cookbook for experimental biologists"1 sl  N  0  N ,$D 02 -  T  >?"`  N rB .  B D?   /  T3| ?"`  Imolecular mechanism3Sl P @@  6  P@@ ,$D 0C@ P @@  5 P @@ 2 1  T  >?"`P @ rB 2 B D?P P  3  T&| ?"`  @  Ealgebraic model32 4  N ?"`@  H  0޽h ? ̙33___PPT10+hV'D~'  = @B D9' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*0 %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*6 %(+^  0L0    K (      <8|"`#  | XB  0D`X  0A?`mp  T| ?  ~EXAMPLE: Determine the association rate constant for A + B ABR@3.3p  H A ?   p  H A ?   ~  H A ?"` z l>P    0,$D  0x   H A ?l >P xB   H DjJ?` ` ` jB  B D?  jB  B D?P  P ~  H A ?"`Rd ~  H A ?"`  E  TL<| ? EGoodrich & Kugel (2006) Binding and Kinetics for Molecular Biologists>F- 6H  0޽h ? ̙333+___PPT10 +шUD'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-m6Bbox(in)*<3<* +-  0L0 WOP (  P P  <\|"`  | XB P 0D`X P 0A?`mE P Tp| ? EGoodrich & Kugel (2007) Binding and Kinetics for Molecular Biologists>F- 6 P H A ?"`xF,$@ 0 P T I  ?w07,$ 0 ;p. 79 P H A ?"`G 1 ,$@ 0  P Tp I  ?7  ,$ 0 <p. 120~  P H A ?"`,  P T_I  ?  ;p. 34  P H A  ?"`G  ,$@  0  P TaI  ?  ,$  0 <p. 126pB P H D>?'@'B P H D>?p ,$D 0B P H D>?g ` pg ,$@ 0B P H D>?U ` U ,$D  0B P H D>?  ,$D 0 P TLhI  ?P $,$   0 6etc. pB P H D>? H P 0޽h ? ̙332*___PPT10 +^ID'  I = @B D' = @BA?%,( < +O%,( < +Df' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-o6Bdissolve*<3<*PD@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-o6Bdissolve*<3<*PD3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-o6Bdissolve*<3<*PD ' =%(DI ' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-o6Bdissolve*<3<*PD@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* P%(D' =-o6Bdissolve*<3<* PD3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-o6Bdissolve*<3<*PD3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-o6Bdissolve*<3<*PDf' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* P%(D' =-o6Bdissolve*<3<* PD@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* P%(D' =-o6Bdissolve*<3<* PD3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-o6Bdissolve*<3<*PD{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*PD' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*P++0+P0 ++0+ P0 ++0+ P0 ++0+P0 +  0L0 @ ^(      <I "`r  I  XB  0D`X  0A?`m  <4DI  SFROM A VARIETY OF ALGEBRAIC EQUATIONS TO A UNIFORM SYSTEM OF DIFFERENTIAL EQUATIONS<T= 3 33  T4I  ? 'EEXAMPLE: Determine the rate constant k1 and k-1 for A + B ABF33333333pB   H DjJ?pjB   B D?0    TqI  ?$  IALGEBRAIC EQUATIONS   TI  ?0e  LDIFFERENTIAL EQUATIONSpB  H D? @0 pB  @ H D?<@0<  TI  ? 80 Lk1.  H|I  ?l@`@ Mk-1.~  H A ?"`^4 ~  H A ?"`` ' M  H0I  ?6  d[A]/dt = -k1[A][B] + k-1[AB]   M  HI  ?   d[B]/dt = -k1[A][B] + k-1[AB]   l  HI  ? 6   <d[AB]/dt = +k1[A][B]  k-1[AB]   l p@    @ p ,$D  0rB   B D?p@ @    TI  ?  ^Applies only when [B] >> [A]&   ThxI  ?4    ^Applies under all conditions&  T  V?"` ,$D 0H  0޽h ? ̙33og___PPT10G+G(#D+'  = @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<* %(+  0L0  z `  (      <$I "`  I  XB  0D`X  0A?`m  <h'I   l&THERE IS NO SUCH THING AS A FREE LUNCH*'& 33pB  H DjJ?jB  B D?`   T-I  ?`P  gALGEBRAIC MODEL<  pB  H DjJ?`   T3I  ?D` X  `can be derived for any molecular mechanism can be derived automatically by computer can be applied under any experimental conditions can be evaluated without specialized software requires very little computation time does not require an initial estimate is resistant to truncation and round-off errors has a long tradition: many papers publishedaa  T9I  ?<`P  \- - - + + + + +* 3  T{I  ?`   A ADVANTAGE 3l `(T   `(T ,$D  0   T(@I  ?`( jDIFFERENTIAL MODEL<  $   TI  ?@T  + + + - - - - -f333  Z  8c?"`` @,$D 0H  0޽h ? ̙33___PPT10+!w D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-m6Bbox(in)*<3<* D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* +  0L0 : 2 (  (  (  (  <(pI "`  I  XB ( 0D`X ( 0A?`m< ( <I   hMORE THAN 600 PAPERS PUBLISHED WITH IT (1996  2009)L5  3  3  33)l  0 0  ( 0 0 ,$D  0 (  <pI  0 0  http://www.biokin.com/dynafit&3 (  <I  2   <DOWNLOAD  H ( 3 A "` ~ ( H A  ?"``? ( T  jJ?"``0pn ( T8tI  ?_ P *Kuzmic (2009) Meth. Enzymol., 467, 247-280p+ , ( T,n ?0   :2009 ( Tn ? Nx ,$  0 [FREE for academic users(H ( 0޽h ? ̙33___PPT10+=Dr' I = @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' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*( D' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*( +8+0+( 0 +  0L0 ?(  XB  0D`X  0A?`m  <n }7APPROXIMATELY 850 JOURNAL ARTICLES PUBLISHED SINCE 1998*87 33   Bn `  n p  H A ?0 @  Tn ?   Kuzmic, P. (1996) "Program DYNAFIT for the analysis of enzyme kinetic data: Application to HIV proteinase" Anal. Biochem. 237, 260-273. Kuzmic, P. (2009) "DynaFit - A software package for enzymology Meth. Enzymol. 467, 247-280.~k    L     P_ QH  0޽h ? ̙33y___PPT10Y+D='  = @B +!  0L0    (   x  c $DRn`  n XB  0D`X  0A?`m  <4n}  n(RATE IS PROPORTIONAL TO CONCENTRATION(S)*)( 33z      ,$D 0Z   s *A   <n  OMONOMOLECULAR REACTIONS`   0    <n  krate is proportional to [A]4Iz  -T     -T ,$D 0Z   s *Ad  `   04 T    <0n   MBIMOLECULAR REACTIONS,   < n b -6  Brate is proportional to [A] [B]Z"z      ,$D  0U   <Hn  - d [A] / d t = k [A] fffffffff`2   0  fB  6DP     <HIn  $monomolecular rate constant 1 / time@%3z  J @   J @,$D  0   <n J ~  X- d [A] / d t = - d [B] / d t = k [A] [B]-fffffffffffffff`2   0J m fB  6D=)  V   <n @ hbimolecular rate constant 1 / (concentration time)^53  HnGHy ` b* rate &  derivative H  0޽h ?  ̙33___PPT10+wYDj' = @B D%' = @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?dCB1+#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<* D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<* 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?dCB1+#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<* D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<* +%  0L0 @ (   x  c $h\n`  n XB  0D`X  0A?`m  <t^n u=PRODUCTS ARE FORMED WITH THE SAME RATE AS REACTANTS DISAPPEAR>>X  0z z zL N   zL N,$D 0   <4nzL N - d [A] / d t =t3333`2   03f+:R  s *A@E  <pn  9EXAMPLE-z ` 4    ` 4 ,$D 0   <+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<* %(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<* 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<* +6  0 ` .(   x  c $dn`  n XB  0D`F fp   fpZ   s *APp   <nf: IEXAMPLE MECHANISM3`   0pz fa   fa,$D 0   <Lmnv fa: FRATE EQUATIONS3   0G  P $ v d[P] / d t =N 3f2   63jJ z    ,$D 0   0M   xd[EAB] / d t =Nf2   6jJ 0f2   6jJ~G 7 f2   6jJ  <v } 5Q,$D   0 PSimilarly for other species...X  0A?`m$  <{ P $,$D 0  + k+5 [EAB]\  #  <83  @ T ,$D  0  - k+5 [EAB]\  d  <g .,$D  0 + k+2 [EA][B] #  < @ ,$D  0  - k-2 [EAB]\  d  < @t ,$D  0 + k+4 [EB][A] #  <  @ d ,$D  0  - k-4 [EAB]\  H  0޽h ? ̙33$$___PPT10}$+瀒pD"' 9 = @B D"' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D{' =%(D#' =%(D' =A@BBBB0B%(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<* D{' =%(D#' =%(D' =A@BBBB0B%(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<* D{' =%(D#' =%(D' =A@BBBB0B%(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<* D{' =%(D#' =%(D' =A@BBBB0B%(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<* D{' =%(D#' =%(D' =A@BBBB0B%(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<* D{' =%(D#' =%(D' =A@BBBB0B%(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<* ++0+ 0 ++0+ 0 ++0+ 0 ++0+ 0 ++0+ 0 ++0+ 0 ++0+ 0 +H  0  p'' (   x  c $p ``   XB  0D`[z j    j ,$D 0;   0ʅ J* T  G E + S ---> ES : k1 ES ---> E + S : k2 ES ---> E + P : k3 pH   3   `   0`    <0 pjf> JInput (plain text file):`B  0D C  <R ,$D 0  d[E ] / dt = X n  0\ @,$D  0  - k1 [E] [S]X  0A?`m  < ą   v0HOW DYNAFIT PROCESSES YOUR BIOCHEMICAL EQUATIONS*10 33R  s *A` JX  00` z      ,$D 0,   <p߅    k1 [E] [S]`   0 z       ,$D 0   0@  z  k2 [ES]n 33333`   0  z l \    l  \ ,$D 0   0\  v J  k3 [ES]n `   0l \  z  `@@   ` @@,$D 0`B   0D j @   <<Յ  ` 4 = Rate terms: `B   0D @z @`@   `@@,$D 0`B   0D@@@   <ȃ &`4 ARate equations:`B   0D@R  <t 0t ,$D   0 + k2 [ES] 33333N ! <` L < ,$D   0 + k3 [ES]| D " <x  p ,$D  0 d[ES ] / dt = X  $z ,   #   , ,$D   0B $  0$a <    + k1 [E] [S]& %  <, ,\ 0  - k2 [ES] 33333" &  <c8   - k3 [ES]|  ' <c p ,$D   0 PSimilarly for other species...H  0޽h ? ̙33++___PPT10*+vD)' c= @B D=)' = @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<* %(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<* %(DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D{' =%(D#' =%(D' =A@BBBB0B%(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<* D{' =%(D#' =%(D' =A@BBBB0B%(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<* D{' =%(D#' =%(D' =A@BBBB0B%(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<*! DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*" %(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<*# D{' =%(D#' =%(D' =A@BBBB0B%(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<*' +P+0+ 0 ++0+ 0 ++0+ 0 ++0+! 0 ++0+" 0 ++0+' 0 +H  0 og// (   x  c $c``  c XB  0D`[z j    j ,$ 0;   0cJ* T  G E + S ---> ES : k1 ES ---> E + S : k2 ES ---> E + P : k3 pH   3   `   0`    < cpjf> JInput (plain text file):`B  0D X  0A?`m  <pc  v0HOW DYNAFIT PROCESSES YOUR BIOCHEMICAL EQUATIONS*10 33R  s *A` JX  00` z      ,$ 0,   <c   k1 [E] [S]`   0 z       ,$ 0   0\"c z  k2 [ES]n 33333`   0  z l \    l  \ ,$ 0   0)c v J  k3 [ES]n `   0l \  z  `@@   ` @@,$ 0`B   0D j @   <@2c ` 4 = Rate terms: `B   0D @z @`@   `@@,$ 0`B   0D@@@   <|7c&`4 ARate equations:`B   0D@R  C *Alego-001`  <   0 N A 8c?"`@ l ,$D  0 0 Tuc ? @ ,$  0 [#Typical email from a Ph.D. student:$$ 3H 0 0޽h ? ̙33  ___PPT10 +AMpDO ' eL = @B D ' = @BA?%,( < +O%,( < +DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*0 %(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*0 D' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*0 D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*0 %(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*0 D' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*0 +8+0+0 0 +  0L0    8 n (  8  8  <c"`  c XB 8 0D`X 8 0A?`m 8 <c  .ALL COMPONENTS PRESENT AT EQUAL CONCENTRATIONS@/ 3 33 8 Tc ? D<4___PPT9 pre-mix sliding clamp (C) + clamp loader (L) to form C.L complex; add DNA solution; observe the formation of C.L.DNA ternary complex over time final concentrations: 100 nM clamp 100 nM loader 100 nM DNA 3b557   @`Z  8 TTc ?2` HCourtesy of Senthil Perumal, Penn State University (Steven Benkovic lab),IH>    8 Tx}c ?PX  X... gp45 labeled with Cy5 acceptor dye ... gp44/62 ... primer labeled with Cy3 donor dyeYYz  8 Tc ?"`@  ,$D  0 C.L complex has estimated Kd = 1 nM, so C.L C+L dissociation upon adding DNA should be negligiblepd 8Z3H 8 0޽h ? ̙33xp___PPT10P+шUD'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* 8 %(D' =-m6Bbox(in)*<3<* 8 +8+0+ 8 0 +c   0L0 r j  @  (  @  @  <0c"`  c XB @ 0D`X @ 0A?`m% @ <  MJUST BECAUSE THE DATA FIT TO A MODEL DOES NOT MEAN THAT THE MODEL IS CORRECT!PN* 3 3  33Z @ Tx  ?2` HCourtesy of Senthil Perumal, Penn State University (Steven Benkovic lab),IH>  ~  @ H A ?"`   @ T} ?"` d  :time  @ N ?"`  B fluorescence y  @ H  ?F  1raw fluorescence F fit to F = A0 + A1 exp (-k t)2   $ @ T ?"`P   Fexponential model fits the data well but it is theoretically invalid!0G>H @ 0޽h ? ̙33y___PPT10Y+D='  = @B +  0L0  %P @(  P  P  <l "`   XB P 0D`X P 0A?`mp P <x  DYNAFIT SOFTWARE IS DRIVEN BY TEXT  SCRIPTS - MINIATURE  COMPUTER PROGRAMS PM 3 3 33~  P H A ?"`0`l   P  ,$D 0 P  T  8c?"`xB P  H D8c?@ P  TD  ? p >sections 3rl P  P P ,$D 0 P  Z  8c?"`P~B P  N D8c?@ P  Z  ?  >keywords 3l @P  $P @P ,$D 0 P  Z  8c?"`@ ~B P  N D8c? P  P  Z$  ? ~  Earbitrary names3 P  T  8c?"` P xB P B H D8c?` P {l @ ` 0  %P @ ` 0 ,$D 0 !P  Z  8c?"`@ p0 ~B "P  N D8c?p   #P  Z0  ?@ `   Ginitial estimates3H P 0޽h ? ̙33: 2 ___PPT10 +D '  = @B D ' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*P %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*P %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*$P %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%P %(+   0L0   @X  (  X  X  <@‡ "`   XB X 0D`X X 0A?`m X <(LJ 0  r(YOUR FIRST DYNAFIT DATA-ANALYSIS SESSION.)( 33 X H  ?. D<4___PPT9 Start DynaFit Select menu  File ... Open or press Ctrl+O Navigate to file ./courses/bkeb/lec-1/a+b/fit-001.txt Select menu  File ... Try or press Ctrl+T This is the initial estimate Select menu  File ... Run or press Ctrl+U Wait several seconds to finish the analysis Select menu  View ... Results in External Browser Navigate in the output filesh3 $ 3  3 - (3 >4^D @`~ X H A ?"``  X T ?"`03 :TUTORIAL H X 0޽h ? ̙33y___PPT10Y+D='  = @B +^  0L0 me` ` (  `  `  <8i "`   XB ` 0D`X ` 0A?`m ` <Dk  [A BIT OF THEORY0 33  ` Hp  ?pp D<4___PPT9 Reaction order Units and dimensions (scaling) The DynaFit model for biomolecular kinetics Initial estimates of model parameters @`H ` 0޽h ? ̙33y___PPT10Y+D='  = @B +  0L0 jb%)h (  h  h  << "`   XB h 0D`X h 0A?`mc h <8ɇ  GIN PRACTICE WE ENCOUNTER ONLY ZERO-, FIRST-, AND SECOND-ORDER REACTIONSH 3 3 3 3 3  3  33 h T  ? ;zero- h Tه  ? Pconstant-rate influx or effluxjB  h @ B D? pB  h H D8c?0u0  h T  ?uz 9ORDER  h HP  ?uu DPHYSICAL MEANING  h T  ?0   <first- h T  ?   =second-jB h @ B D?5 5 jB h @ B D?EE  h T<  ?0   _-isomerization or dissociation of one molecule.. h T($  ?   X&binding (association) of two molecules'' h T%  ? DDYNAFIT NOTATION h T 2  ?E IX --> : v3 h T3  ?0   LA --> B : k1 C --> A + B : k1 ''3 h T5  ?   JA + B --> C : k23 h T(=  ?  C A B  h H\@  ?% &  <NOTATION pB h H D?` `  h T(I  ?p D  Lk1. h TD,  ?L  HA + B CpB h H D?  h TlG ?  Lk2. h T,nG ?  iA B + C> pB  h H D?r r  !h ThG ? V  Pk 1.jB "h B D?pupjB #h B D?@ u@ l 8Nf=  )h N= ,$D 0 $h  TG ? 8  muni-molecular monomolecular 3 &h  TG ? =  C bimolecular  32 (h  N  ?"`8NfH h 0޽h ? ̙335-___PPT10 +6D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*)h %(D' =-o6Bdissolve*<3<*)h +&  0L0 h`*Mx (  x  x  <G"`  G XB x 0D`X x 0A?`m x <G MDYNAFIT CAN HANDLE AN ARBITRARY NUMBER OF ELEMENTARY REACTIONS IN A MECHANISM.NM 33 &x TG ?SI' hA + B C>  'x T8G ?;2 Lk1.pB (x H D?;+pB )x @ H D?;+ *x H6G ?;[ Lk2.pB +x H D8c?0PjB ,x @ B D?&  -x T:G ?p AF DDYNAFIT NOTATION /x TtG ?6p   OA + B <==> C : k1 k23 9x T ?G ?t ! EREVERSIBLE REACTION > 8x ZX|G ? p @,$ 0 HA + B <==> AB : k1 k2 A + C <==> AC : k3 k4 AC ---> X : k5II3l 0P Mx 0P,$D  0 0x  ZЁG ? ,  GA AB 1x  ZG ?%P $  Lk1.~B 2x N D?e\ \ ~B 3x B N D?e   4x  NG ? t  Lk2.~B 5x  N D8c?0 P xB 6x B H D? @  7x  ZXG ?p A`  DDYNAFIT NOTATION :x  ZG ? M  FMULTI-STEP MECHANISM  ;x   `0G ? J  :+B\T    >x C#   xB +B#style.visibility<*Kx %(D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Mx %(D' =-m6Bbox(in)*<3<*Mx DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*8x %(+8+0+8x 0 +-  0L0 )] (      <0G"`  G XB  0D`X  0A?`m  <,G @CAREFUL ABOUT DIMENSIONS OF RATE CONSTANTS! DIMENSIONAL ANALYSIS@A, 3 33 6 TTG ? iA + B AB>  7 T,G ?$ Lk1.pB 8 H D?pB 9 @ H D?RR : HZG ?\0 Lk2.  ; HTRG ?,Z  v = k1 [A] [B]Z   < H0G ?|Z P v = k2 [AB]Z   = TxG ?|P  c#forward and reverse reaction rates:$$  jB B B D?0: C TcG ?`  quantity v [X] k1, k2   jB D B D?P PJ F T  ?`l 2dimension concentration / time concentration ?x3  3v G N  ?0 l xt  W xt ,$D  0< >  T  ?h x<  Bdimensional analysis of k1 (bimolecular association rate constant)RC  ?  H  ? t  v = k1 [A] [B]Z   H  TD<  ?lT  9EXAMPLE(l `  Y  ` ,$D  0 @  H[  ? ` |  Sk1 = 2  A  T ?"`` L xB L  H D?    M   ``  ?`  w  Fv& N   `p  ?7  9[A] [B]`l @    Z  @  ,$D  0 I  Tt  ?`    B{conc.} / {time}rB J B D? P  O  Z\y  ?` M   n"{conc.} {conc.}0 P  N ?"`@ K tl  P  [  P ,$D 0xB Q B H D8c? 0 P xB R B H D8c?  l P   \  P ,$D 0 S  T͖  ?P 3  5= T  TK  ?   51rB U  B D?@   V  Z0Ж  ?M   r {conc.} {time}6H  0޽h ? ̙33,$___PPT10+*D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*W %(D' =-m6Bbox(in)*<3<*W D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Y %(D' =-m6Bbox(in)*<3<*Y D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*Z %(D' =-m6Bbox(in)*<3<*Z D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*[ %(D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*\ %(D' =-o6Bdissolve*<3<*\ +  0L0  @ (      < "`   XB  0D`X  0A?`m  <Ԗ  SSUMMARY0 33 * Tܺ  ? iA + B AB>  + TȖ  ?`4 Lk1.pB , H D?PPpB - @ H D? . H<Ӗ  ?l Lk2. / T4  ? ec  NA A   0 Tז  ? ri  Lk3.pB 1 H D? rb pB 2 @ H D? rb  3 H8  ?( r  Lk4.2 4 Td  ?p  zdissociation rate constant association rate constant dissociation equilibrium constant association equilibrium constant{{pB 5 H DjJ?   6 Hh}  ? w| Lk2. 7 Hd  ? w Lk1.? 8 H  ? p  Kd = k2 /k1  % 9 H  ? p  Ka = k1 /k2  : T  ?fOt P1/sec 1/(M sec) M 1/M3 ; T  ? x = 0 rate constant rate constant equilibrium constant equilibrium constanttQ < H#  ?   Lk3. = H  ?   Lk4.. > HÖ  ?` 4  K = k3 /k4 . ? H3  ?P $ K = k4 /k3  @ TP  ?,  L1/sec 1/sec -- --3H  0޽h ? ̙33y___PPT10Y+D='  = @B +I  0L0  v(      <|D "`   XB  0D`X  0A?`m#  <(F  KALL UNITS ARE ARBITRARY BUT MUST BE IDENTICAL THROUGHOUT THE ENTIRE SCRIPT!PL  3  3 33;  T,)  ?& [mechanism] DNA + Clamp.Loader <==> Complex : kon koff [constants] kon = 1 ? koff = 1 ? [concentrations] DNA = 0.1 Clamp.Loader = 0.1 [responses] Complex = 1 ? [data] file ./.../d1-edit.txt offset auto ? t  2 az @     @,$D  0O   Tޖ  ?"`0  _time, s signal, eV 0.52 0.3526 0.56 0.3484 0.60 0.3485 0.64 0.3454 0.68 0.3499 0.72 0.3502 ...`` N   N  ?"`@n B   T D? 0 l  U    U ,$D 02   Z  V?"`r U ~B  N D?    Z  ?   s!time units of rate constants: sec6"l 6w   6w ,$D 02   T  8c?"`6 xB  H D?pp    T  ?pw {)concentration units of rate constants: M6*    Tw  ?`  ^100 nM3#l  P   P,$D 02   T  V?"`PxB  H D?`   T\}  ?  = 1 M-1s-1 = 106 M-1.s-1H  0޽h ? ̙33  ___PPT10 +aF/D '  = @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<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(+&  0L0 G ? @  (      < "`   XB  0D`X  0A?`m~  <t  MOLAR  RESPONSE = PROPORTIONALITY FACTOR LINKING CONCENTRATIONS TO SIGNALbK2 3 3 3 33b  Tؗ  ?& [mechanism] DNA + Clamp.Loader <==> Complex : kon koff [constants] kon = 1 ? koff = 1 ? [concentrations] DNA = 0.1 Clamp.Loader = 0.1 [responses] Complex = 1.00 ? [data] file ./.../d1-edit.txt offset auto ? @?t  2 dz @     @,$D  0O   T  ?"`0  _time, s signal, eV 0.52 0.3526 0.56 0.3484 0.60 0.3485 0.64 0.3454 0.68 0.3499 0.72 0.3502 ...`` N   N  ?"`@n B   T D? 0 Hl l   l ,$D 02   T  8c?"`p p 2   T  8c?"`     Zȓ  >?"` l  |one concentration unit (in this case 1 M) of Complex will produce an increase in the signal equal to 1.00 instrument unitsl}." rB  B D?   rB   B D?  H  0޽h ? ̙33og___PPT10G+Dr D+'  = @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<* %(+  0L0 *" (      < "`   XB  0D`X  0A?`m'  </  ONONLINEAR REGRESSION ANALYSIS ALWAYS REQUIRES INITIAL ESTIMATES OF THE SOLUTIONPP. 3 3 33  T<  ?& K[mechanism] DNA + Clamp.Loader <==> Complex : kon koff [constants] kon = 1 ? koff = 1 ? [concentrations] DNA = 0.1 Clamp.Loader = 0.1 [responses] Complex = 1 ? [data] file ./.../d1-edit.txt offset auto ? Z [7t  3 c2  T  1?"`;2  T  1?"`;2  T  1?"` `j 2  T  1?"` `  pB  @ H D?` pB  @ H D?` pB  @ H D? ``  jB  B D?` ` pB  @ H D? `   TP  ? U Poptimized model parameters3  TX  ?  ,$  0 A VERY DIFFICULT PROBLEM: HOW TO GUESS  GOOD ENOUGH INITIAL ESTIMATES OF RATE CONSTANTS?(\A  T  8c?"` jB  B D?  Tt  ?   ^&the initial estimate of rate constants''H  0޽h ? ̙33xp___PPT10P+MMD'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-m6Bbox(in)*<3<* +8+0+ 0 +0  0L0 ?7 (     <d "`   XB  0D`X  0A?`m~  H A" ?"``np  H A# ?`   N ?"``p  N ?"`@ pj  C BA*data-model-t001-g001`0    T  V?"`0P  Th  ?J :data   TTm  ? ;model   T  V?"`N H  0޽h ? ̙33y___PPT10Y+D='  = @B +  0L0  8(     <l "`   XB  0D`X  0A?`m~  H A" ?"``n  N ?"``p  N ?"`@ p   T  V?"`0Pp   H A ?T   T  V?"` > p  H A$ ?`0  p  H A% ?x p  H A& ?x H  0޽h ? ̙33y___PPT10Y+D='  = @B +   0L0 /'  (       <xē "`   XB   0D`X   0A?`mp   H A# ?`    N ?"``p   N ?"`@ p   T  V?"`N p   H A' ?`\   T  V?"`0@P   T  ?q  l a hundred times smaller / larger0!   p   H A( ?`0     T̟  ?@0z :data    T  ?0 ;model H   0޽h ? ̙33y___PPT10Y+D='  = @B +  0L0 @ (*(  ( (  <] "`   XB ( 0D`X ( 0A?`m ( N ?"``p ( N ?"`@ pp  ( H A ?T   ( T  V?"` > p ( H A' ?`\ ( T  V?"`0@Pp ( H A) ?`0  p ( H A* ? jp ( H A+ ? @6H ( 0޽h ? ̙33y___PPT10Y+D='  = @B +  0L0 RJ`40 (  0 0  <0+ "`   XB 0 0D`X 0 0A?`mpB 0 H D8c?jB 0 B D?00G 0 T$J  ?& @sum of squares jB 0 B D?] ` G 0 T,'  ?S  Erelative sum of sq. jB 0@ B D? G 0 HM  ? @ Z( best-fit constants   0 T  ? Binitial estimate jB "0 B D? #0 T`  ?;  s k1 = 1 k2 = 1J   $0 T+B#style.visibility<*40%(D' =-m6Bbox(in)*<3<*40+   0L0 -% $8(  8 8  < "`   XB 8 0D`X 8 0A?`mp  8 H A- ?`0  p !8 H A- ?`   "8 T  ?T f (  8From  good initial estimate2 #8 T@  ?` 4  6From  bad initial estimate23  $8 Tޞ ?"`ps,$D 0 Gnot very encouraging!H 8 0޽h ? ̙33zr___PPT10R+^D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*$8%(D' =-o6Bdissolve*<3<*$8+8+0+$80 +  0L0 F >  @ (  @ @  < "`   XB @ 0D`X @ 0A?`m  @ T  ?*^ D<4___PPT9 Initial estimates  off by a factor of 100 can produce misleading results. The data/model overlay may  look good , but the results may be invalid. The same applies to the residual sum of squares (only 2% difference). The only indication that something went wrong might be: a. huge standard errors of model parameters; and b. various warnings from the least-squares fitter The simplest possible safeguard: Use several different initial estimates? Disadvantage: how do we know which multiple estimates?333W ' @`|l  w  @ w,$D 0  @ T| ?"` w LOOKING AHEAD DynaFit offers more reliable and convenient safeguards Global minimization Systematic combinatorial scan Confidence interval searchPGP )P\  @ C ,A.hand-icon < H @ 0޽h ? ̙335-___PPT10 +D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* @%(D' =-o6Bdissolve*<3<* @+   0L0    H4 (  H H  <,ɞ "`   XB H 0D`X H 0A?`mT  H Tʞ  ?prD<4___PPT9 Numerical models are applicable to all experimental conditions. No more  large excess of this over that . Numerical models apply uniformly to all types of experiments: a. reaction progress (kinetics); b. equilibrium composition (binding); c. enzyme catalysis. Numerical models can be automatically derived by computer. No more looking up algebraic equations  if they exist at all. Main disadvantage: requirement for specialized software. But DynaFit is free to academic users. Not a  silver bullet ! Example: the initial estimate problem. 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