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(                      &1:D>phs?"-(./H&D$        5    ,*&:#$+    D# (#'609 <e>02  9"7G55H#;  #&  (#  (0 '9  8 E9#* "#,1  $-wI  "&-h7+4 32#  4 ?,           (0  ?/#-I.#  ) *+P,*-O./)0F2E3'4%526)789#:+;<#=*>&@ABC(DEFGH b$k-YsGU r$?b$sJ먻' AfQ?b$s2S|t >3HKcb$3}2ULz9JBb$409}G=8l>@-Rb$zy+-_Nz:\ ?b$3Lj^c@4 %.Wb$':r5)|b$d/%vM㋃ordb$_2CB|b$;dᦴ4gb$xvuyN_m;EԾb$6YF9Xq/ ܶb$'H;֠;jU7Ab$>jǭ虾*iIVb$gIHCtb$ƋwI)ٰW$b$k`s`=Ϧb$Ԛ'~KۤDW$&ob$w^!UoAb@ =[b$X']X_E{09Y b$苔Ů|wxq~'GUb$NA j^%k" b$2s=(Ao|,b$Z3q~^şneqF%7b$Rotd~B"H+3'Lb$eU j2zX;bb$h$f!MUޑ&y8sb$i1l?b$Y 2>bfD?b$17-πy- b$K$w/X1%b$cSDm@ū檰 5<b$H~?jy ϪQ,:UXb$L XMK:eb}ˆHob$yd+=S.!!b$zV%ARQ3'b$7+]Nau, 0e0e     A@  A 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E|| 33f33@dg4tdtd x: 0*ppp@  <4!d!d@ 0t<4BdBd@ 0tg4dddd] x: 0Pp p#p ʚ;Xm8ʚ;<4dddd@U 0@r0___PPT10 2___PPT9/ 0?R, >DynaFit: Enzyme Progress CurvesO =">  }3Algebraic solution for time course of enzyme assays44FEnzyme kinetics in the  real world $$-Progress curvature at low initial [substrate]..1Problems with algebraic models in enzyme kinetics:2 /Numerical solution of ODE systems: Euler method00.Automatic derivation of differential equations//Software DYNAFIT (1996 - 2010)!DYNAFIT: What can you do with it?""1DYNAFIT applications: mostly biochemical kinetics22  /Traditional analysis of irreversible inhibition00TTraditional analysis   Take 2 : nonlinear++0Traditional analysis: Three assumptions (part 1)11 0Traditional analysis: Three assumptions (part 2)11 0Traditional analysis: Three assumptions (part 3)11.Simplifying assumptions: Requirements for data//:Actual experimental data (COURTESY OF Art Wittwer, Pfizer)(;# *  &Numerical model for Caliper assay data('&  \Caliper assay: Results of fit  optimized [E]0*/-  6Caliper assay violates assumptions of classic analysis*76 /Numerical model more informative than algebraic*0/   'Numerical modeling looks simple, but...*(' Residual plots* Direct plots of data: Example 1*  Residual plots: Example 1* "Residual plots: Runs-of-signs test*#" Residual plots: Example 2* +Relaxed inhibitor concentrations: Example 1*,+ 4Relaxed inhibitor concentrations: Example 1 (detail)*54 1Relaxed inhibitor concentrations: not all of themF2& \Initial estimates: the  false minimum problem*/. 3Large effect of slight changes in initial estimates*43 5Solution to initial estimate problem: systematic scan*65 'Model discrimination: Use your judgment*(' Summary and Conclusions*  Questions?*   /pC~   0` ̙33` ` ff3333f` 333MMM` f` f` 3>?" dd@,|?" dd@   " @ ` n?" dd@   @@``PR    @ ` ` p>> <4(    6lh  P   T Click to edit Master title style! !  0k     RClick to edit Master text styles Second level Third level Fourth level Fifth level!     S  0@  ``   X*  0  `    l*(    0l  `    `* H  0޽h ? ̙33 Default Design 0 P (  4 4 N* y y .    p* a00aa 4 N: y y 3 .   r* a00aad 4 c $ ?   4 4 Np= y y 9 f   RClick to edit Master text styles Second level Third level Fourth level Fifth level!     S  4 T$t y y r    p* a00aa 4 Tw y y r3    r* a00aaH 4 0ηo~ ? ̙3380___PPT10.R}X \x(  \ \ N/V]e]e    V p* AAA \ NX4V]e]e %   V r* AAA  \ TlV]e]e   V p* AAA \ T&}]e]e %  V r* AAAH \ 0ηo~ ? ̙3380___PPT10.R.IW 0 ~v0!(  X  0A?`mv  6l   RDynaFit in the Analysis of Enzyme Progress Curves Irreversible enzyme inhibitionS  1  <   APetr Kuzmic, Ph.D. BioKin, Ltd. WATERTOWN, MASSACHUSETTS, U.S.A.hB    XB  0D0  Td  ?< r PH@___PPT9" #TOPICS Numerical integration vs. algebraic models: DYNAFIT Case study: Caliper assay of irreversible inhibitors Devil is in the details: Problems to be aware ofN   @`XB  0D0H  0޽h ? ̙33y___PPT10Y+D=' }= @B +  0L0 5 - `DP  (  P r P S , `0   XB P 0D`X P 0A?`m P <X   @ONLY THE SIMPLEST REACTION MECHANISMS CAN BE TREATED IN THIS WAYLA  3 3/ 33~ 

P Tؼ  ? M &Kuzmic (2008) Anal. Biochem. 380, 5-12^'       @P TĜ  ?"`  |$D 0H@___PPT9" SIMPLIFYING ASSUMPTIONS: No substrate depletion No  tight binding>++ @` BP THȜ  ? d  yTASK: compute [P] over timeB 3 0l  30 DP  30,$D 0 ?P  H A ?"`@ 30 AP  N ?"` @ 2 CP  T  8c?"`@  H P 0޽h ? ̙33___PPT10+͹Dt' t = @B D/' = @BA?%,( < +O%,( < +D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*@P %(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*@P D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*@P D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*DP %(D' =-o6Bdissolve*<3<*DP +8+0+@P 0 +  0   > . (   x  c $Dߜ `   XB  0D`X ) 0A?`m * <P   /SUBSTRATE DEPLETION USUALLY CANNOT BE NEGLECTED>0 3 3 3~ 4 H A ?"`x 5 TT  ? M 6Sexton, Kuzmic, et al. (2009) Biochem. J. 422, 383-3927         l  8 ,$D 0 6  T  8c?"`W 7  T  8c?"`5 9 T  ?P=,$ 0 }8% systematic errorN 3 3  3 3l @  =  @,$D 0 ;  T  |?"`@  <  T  8c?"` P   > Tz  ?p  ,$ 0 ^residuals not random.  3  3H  0޽h ? ̙33  ___PPT10 +ۆ D '  = @B D ' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*8 %(DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*9 %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*= %(DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*> %(+p+0+9 0 ++0+> 0 +#  0 KC0" (   x  c $8 `   XB  0D`X  0A?`m3  <D 3 3SUBSTRATE DEPLETION IS MOST IMPORTANT AT [S]0 << KMx4 3 3 3 3 3 3Y  TH%  ?T M 7Kuzmic, Sexton, Martik (2010) Anal. Biochem., submittedN8     , ~  H A ?"`    H A ?"`p ? ,$D  0l pP0   Pp0 ,$D 0xB  H DjJ?p0    T8.  ?Pp U initial slope, 3 3l 0  "  0  ",$D 0xB  H DjJ?0  "   T`3  ?   S final slope,   `l     ,$D 02   T  |?"` F 2   T  |?"`F   TL9 ?"`   :1.343   T= ?"` :0.82  T@  ?t,$ 0 ]~40% decrease in rate,330l 0,  !  ,$D 0    TF  ?n0 ,   linear fit: 10% systematic errorX!   3  3 3    N  ?"`0@ " TM  ?v ~[S]0 = 10 M, KM = 90 MJ    H  0޽h ? ̙33___PPT10+Dd' V = @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<* %(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' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*! %(+8+0+ 0 +  0 Y Q  .  (   x  c $4 `   XB  0D`X % 0A?`m & <6   u/THERE ARE MANY SERIOUS PROBLEMS AND LIMITATIONS*0/ 33 + Tx=  ?d < Can be derived for only a limited number of simplest mechanisms Based on many restrictive assumptions: - no substrate depletion - weak inhibition only (no  tight binding ) Quite complicated when they do existl-33T 3 , 6H    ,$ 0 The solution: numerical modelsH B - 0D0 `0 ,$D 0 . T\N  ?V W:,$ 0 H Can be derived for an arbitrary mechanism No restrictions on the experiment (e.g., no excess of inhibitor over enzyme) No restrictions on the system itself ( tight binding ,  slow binding , etc.) Very simple to derivel333 H  0޽h ? ̙335 - ___PPT10 +D'  = @B D<' = @BA?%,( < +O%,( < +Ds' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*, %(D' =-o6Bdissolve*<3<*, D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*. %(D' =-o6Bdissolve*<3<*. D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*- %(D' =-o6Bdissolve*<3<*- +p+0+, 0 ++0+. 0 +[0  0 @+, (   R2  s *P R2  s *p x  c $8e `   XB  0D`C  <8h x d [A] / d t = - k [A]^B  6D0 0 ^B  6D 0     < v  @  8time   0 } 0  j[A]0J ^B   6D8c@0 ^B   6D ^B   6D ^B  6D0 pp pB  HDo ^B  6D3f8c p0 ^B  6D8c0 p``   <   ,$ 0 Mstraight line segment  <  =  _[A] t>   <|| P $ [D t<vB  NDjJ X  0 0[z x$   x$,$D 0o   <l Px$ D [A] / D t = - k [A]333Z2   s *p Zz pT    pT ,$D 0n   <L 0 T  D [A] = - k [A] D t 333Z2   s *pp z P `     P `  ,$D 0   0 @ `   %[A] t + D t - [A] t = - k [A] t D t f& 3333Z2   s *pP  %z ` ` T    ` ` T,$D 09 !  0Κ  ` T #[A] t + D t = [A] t - k [A] t D t $ 3333Z2 "  s *p`  X # 0 @@ $ <;   O[A]0X % 0A?`m & <4  @COMPLETE REACTION PROGRESS IS COMPUTED IN TINY LINEAR INCREMENTSLA/ 3 3  33" ' Tl ?"`p> 6 k mechanism: A B>7  ( <    [A] t+DtZ     ) H GH8H ?"``  Rdifferential rate equation pB * H D?) , T ?"` ,$D  0 c1practically useful methods are much more complex!22H  0޽h ?)  ̙33vn___PPT10N+D'  = @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<*  %(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<*, +p+0+ 0 ++0+, 0 +Q#  0 P"H S(   x  c $( `   XB  0D`X % 0A?`m8 & <  IT IS SO SIMPLE THAT EVEN A  DUMB MACHINE (THE COMPUTER) CAN DO IT*DC 33H , <X 0* :  D k1 E + S ---> ES k2 ES ---> E + S k3 ES ---> E + P |E    3  X - 0P` @ X2 . 0 / < Pps $ R Example input (plain text file):!!^B 0 6Dp 1 <\0 p  ( multiply [E] [S] <XB 2 0DP p  3 <,. F  = Rate terms: $ 4 <0 ` 4 k1 [E] [S]XB 5 0Dpp 6 0,@ p D  k2 [ES]n 33333 7 0G P  $  k3 [ES]n XB 8 0D@@  9 <O F& ARate equations:l ,  H , ,$D 0 :  <tT   d[E ]/dt = X  N ;  0d` @,  T- k1 [E] [S] + k2 [ES] + k3 [ES]v+33333bl qT F U8,$D 0fB ? B 6D` x@ qT D qTZ2 >  s *q5 @  <d< Tt R E disappears l Y  E Y ,$D 0Z2 <  s *mYZ2 =  s *j   A  <| p  P E is formed fB B 6D@0 fB C  6D0 @ p G T ?"`p 0O ,$D 0 *similarly for other species (S, ES, and P)d+H  0޽h ? ̙33. & ___PPT10 +D '  = @B Dm ' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*H %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*E %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*F %(D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*G %(D' =-o6Bdissolve*<3<*G +8+0+G 0 +  0 ` - I(   x  c $ `   XB  0D`X  0A?`m   <  nPRACTICAL IMPLEMENTATION OF  NUMERICAL ENZYME KINETICS *87 33! ' <l  `  ,$ 0 http://www.biokin.com/dynafit&3 ( <4  ] ,$ 0 <DOWNLOAD  H ) 3 A "` ~ * H A ?"``? + T  jJ?"``0pn , T@1  ? M *Kuzmic (2009) Meth. Enzymol., 467, 247-280p+       , - T4  ?0   :2009H  0޽h ? ̙33___PPT10t+}D'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*( %(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*' %(+p+0+' 0 ++0+( 0 +   0 ? 7  @  (  @ x @ c $h `   XB @ 0D`X @ 0A?`m @ <t  UANALYZE/SIMULATE MANY TYPES OF EXPERIMENTAL DATA ARISING IN BIOCHEMICAL LABORATORIES *VU 33H  @ TH  ?.  R Basic tasks: - simulate artificial data (assay design and optimization) - fit experimental data (determine inhibition constants) - design optimal experiments (in preparation) Experiment types: - time course of enzyme assays - initial rates in enzyme kinetics - equilibrium binding assays (pharmacology) Advanced features: - confidence intervals for kinetic constants * Monte-Carlo intervals * profile-t method (Bates & Watts) - goodness of fit - residual analysis (Runs-of-Signs Test) - model discrimination analysis (Akaike Information Criterion) - robust initial estimates (Differential Evolution) - robust regression estimates (Huber s Mini-Max)4 3536-  /A #H @ 0޽h ? ̙33y___PPT10Y+D='  = @B +  0  ( (  ( x ( c $* `   XB ( 0D`X ( 0A?`m ( <1 1 KBUT NOT NECESSARILY: ANY SYSTEM THAT CAN BE DESCRIBED BY A FIRST-ORDER ODEs*LK 33G~ ( H A ?"``D   ( T  ?   |~ 650 journal articles totalD33p ( H A  ?` H ( 0޽h ? ̙33y___PPT10Y+D='  = @B + 0 t J(  t X t 0A?`mv t 64   RDynaFit in the Analysis of Enzyme Progress Curves Irreversible enzyme inhibitionS  9 t <   APetr Kuzmic, Ph.D. BioKin, Ltd. WATERTOWN, MASSACHUSETTS, U.S.A.pB   XB t 0D0XB t 0D03 t Td  ?< r PH@___PPT9" WTOPICS Numerical integration vs. algebraic models: DYNAFIT Case study: Caliper assay of irreversible inhibitors Devil is in the details: Problems to be aware of 5630 @`H t 0޽h ? ̙33y___PPT10Y+D='  = @B +{/  0 &&=O| p&(  |  N| N ?"`0@x | c $^ `   XB | 0D`X | 0A?`m | <e   LBEFORE 1981 (IBM-PC) ALL LABORATORY DATA MUST BE CONVERTED TO STRAIGHT LINES>M  33 3 3  | Ti  ? F1962&00  | Th  ?o BKitz-Wilson plot l `   F| `  ,$D 0xB | H D?@P@ xB |  H D?@  |  TXs  ?P0   6time |  Tw  ?` 4 Olog (enzyme activity/control)rB |  B D?@p rB |  B D?@p rB |  B D?@ 0 rB |  B D?@ P 2 !|  N ?"`@P2 "|  N ?"`p2 #|  N ?"`p 2 $|  N ?"``0 2 &|  N ?"``@2 '|  N 3?"`P0 2 (|  N 3?"`0 p 2 )|  N 3?"` 0 2 *|  N 3?"` P 2 +|  N 3?"``X2 ,|  N 3f?"`0 P 2 -|  N 3f?"`  2 .|  N 3f?"`pP  2 /|  N 3f?"`` @ 2 0|  N 3f?"`p`2 1|  N ?"```  2 2|  N ?"` PP 2 3|  N ?"`  2 4|  N ?"`@ p 2 5|  N ?"`00 2 6|  N ?"`pp  7|  T̂  ?H P  Iincreasing [inhibitor]  l  `c  G| ` c ,$D 0xB 8| H D? P xB 9|  H D?   :|  T  ?P0 /  A1 / [inhibitor]) ;|  TT  ?` `c4 !1 / slope of straight line (kobs)>" rB <| B D?  2 =|  N ?"` 2 >|  N ?"`` 2 ?|  N ?"`@2 @|  N ?"`@`  xB A| B H D?  B|  T  jJ?"`  C|  T*  ?   L1/kinact(  r D|  Z  jJ?"` 0  E|  TȮ  ?  H1/Ki( D H| T8  ? RE + I EI X0*pB I| H D?4 4pB J| @ H D?d dpB K| H D?H H L| ZĻ  ?X n, dkinact(  M| Z\  ?X , `Ki( B O| T  ?0v ,$ 0 2Kitz & Wilson (1962) J. Biol. Chem. 237, 3245-3249<3$    H | 0޽h ? ̙33C;___PPT10+d%*D'  = @B D' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*F| %(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*G| %(DA' =%(D' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*O| %(+8+0+O| 0 +B6  0 (0 0J /(   x  c $4 `   XB  0D`X  0A?`mj  <,!  AFTER 1981 STRAIGHT LINES ARE NO LONGER NECESSARY ( NONLINEAR REGRESSION )NK  3) 3 3 3  T  ? F1981&003  T  ?8{ EIBM-PC (Intel 8086)1l `   ` ,$D 0~B 8 N D?@P@ ~B 9  N D?@  :  Z(0  ?P0   6time ;  Z8  ?`>4 Ienzyme activity/control2 @  T ?"`P 2 A  T ?"`p0 2 C  T ?"`` ` 2 D  T ?"`@`2 J  T 3f?"`` ` 2 K  T 3f?"` P 2 L  T 3f?"`P  2 M  T 3f?"`  2 N  T 3f?"`pp2 O  T ?"`p  2 P  T ?"` ` 2 Q  T ?"` Pp 2 R  T ?"`@ p 2 S  T ?"`@0 2 T  T ?"`pp  U  Z`J  ?H P  Iincreasing [inhibitor] 2 V  N ?"`pxB W  H D3f?@0pxB Y  H D3f?0p xB Z  H D3f? @ xB [  H D3f?@ 0 xB \  H D?@xB ]  H D?0 xB ^  H D?0  xB _  H D?  xB `  H D?  xB a  H D?@p xB b  H D?p  xB c  H D? 0 xB d  H D?0 @ xB e  H D? @ @p y f  TPT  ?"`p  A = A0 exp(-kobs t) 333   N ?"`0@D  T\`  ? RE + I EI X0*pB  H D?4 4pB  @ H D?d dpB  H D?H H  Z C  ?X n, dkinact(   ZL?  ?X , `Ki( l ^ `N  `^ N,$D 0~B h N D? P ~B i  N D?   j  ZZ  ?P0 J  = [inhibitor] 2 m  T ?"`P P 2 n  T ?"`p2 o  T ?"`p2 p  T ?"`@ `  s  Zn  ?` N4  dkinact( r t   `  jJ?"` 0  u  Z u  ? d  `Ki( 2 v  N ?"` 0 xB w H D? xB x B H D? 0 p xB y H D?p@0 xB z B H D?@@xB {  H D? xB |  H D?  xB }  H D?  ~  T  jJ?"`0    T0D  ?^ ` 4 pkobs6    T4  ?"`  kobs =kinact / (1 + Ki / [I])   6   T  ? Q  z 0.5 kinact2     H  0޽h ? ̙33___PPT10+JD~'  = @B D9' = @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<* %(+   0  N V(   x  c $p `   XB  0D`X  0A?`m  <|  z LINEAR OR  NONLINEAR ANALYSIS  THE SAME ASSUMPTIONS APPLY>> 3p 7 H A ? } ^ ~ 8 H A  ?"`bP  9 N  ?"`   : T  |?"`   ; N ?"` a  < T  ? -l  \vno  tight binding [I], Ki must not be comparable with [E]<3 3 3 3 3 3 3 3 3!* = Tl  ?PF $ ,1. Inhibitor binds only weakly to the enzyme^-  H  0޽h ? ̙33y___PPT10Y+D='  = @B +  0 0& Y(   x  c $ž `   XB  0D`X  0A?`m  <Ğ  z LINEAR OR  NONLINEAR ANALYSIS  THE SAME ASSUMPTIONS APPLY>> 3  T ?"` `p   Z͞  ? R2. Enzyme activity over time is measured  directly In a substrate assay, plot of product [P] vs. time must be a straight line at [I] = 0        3 3 3 3 3 3  3 J  Zޞ  ?d 8 8  RE + I EI X0*vB  N D? * vB  @ N D? * vB  N D?    `4  ?   dkinact(    `0  ?   `Ki( `  ZW  ? F  VE + S ES E + PB,3vB  N D? 4 vB  @ N D?< 4< vB  N D?     `  ?0 E   bkcat(    `  ?0   FKm(   N ?"`` D  TԱ  ?l @ RE + I EI X0*pB  H D?pB ! @ H D?pB " H D?`   # Z`  ?0  dkinact(  $ Zl  ?0% `Ki(  % Tæ  ? ` DASSUMED MECHANISM: & Tȿ  ?   QACTUAL MECHANISM IN MANY CASES: H  0޽h ? ̙33y___PPT10Y+D='  = @B +  0 @ $ z(   x  c $ `   XB  0D`X  0A?`m  <Ҧ  z LINEAR OR  NONLINEAR ANALYSIS  THE SAME ASSUMPTIONS APPLY>> 3  TЬ  ?0dw n3. Initial binding/dissociation is much faster than inactivation ( rapid equilibrium approximation )i   $ 3~  H A  ?"`P$ ~  H A ?"`> P^h p ! H A ? PXpB " H Dm?] 0 0 ] pB # H Dm?,p,pB $ H Dm?,p ,H  0޽h ? ̙33y___PPT10Y+D='  = @B +T  0 :2) (   x  c $ `   XB  0D`X  0A?`m"  <  LHOW MUST OUR DATA LOOK SO THAT WE CAN ANALYZE IT BY THE TRADITIONAL METHOD ?NM 3" 3  3  3~  H A  ?"`0 i  T  ?"``> .SIMULATED: [E] = 1 nM [S] = 10 mM Km = 1 mM |/  3  > B  Z DjJ?     T  ?d ?[I] = 03B  Z DjJ?   T!  ?p^D d [I] = 100 nM 3 B  Z DjJ?l l  T&  ?^ d [I] = 200 nM 3 B  Z DjJ?    TP+  ?p^D  d [I] = 400 nM 3 B ! Z DjJ?   " Tզ  ? ^  d [I] = 800 nM 3 l ` ` % ` `,$D 0 #  TѦ  ?`  V 1. control curve = straight line!!B $  Z D8c? `l , ) ,,$D 0  &  TD<  ?$ , I2. inhibitor concentrations must be much higher than enzyme concentrationJJ '  T  jJ?"` @`  (  T  jJ?"`@pH  0޽h ? ̙33___PPT10+*D~'  = @B D9' = @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<*) %(+  0 r j (  (   x  c $TI `   XB  0D`X  0A?`m  <LK  ?NEITHER OF THE TWO MAJOR SIMPLIFYING ASSUMPTION ARE SATISFIED !,@ 39 3~  H A  ?"`0 il ` # `,$D 0   ZW  ?` F1. control curveB    ` D8c?B  Z DjJ?     T@Z  ?d ?[I] = 03B  Z DjJ?  ! T^  ?peD v [I] = 2.5 nM, 33  " T d  ?`4 v [E] = 0.3 nM, 33 "l @, ( @,,$D 0 %  Zj  ?$ , t>2. inhibitor concentrations within the same order of magnitude?? &  Z  jJ?"` @`H  0޽h ? ̙33___PPT10+XD~'  = @B D9' = @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<*( %(+   0 RJ E (   x  c $y `   XB  0D`X  0A?`m  <{  }5NO ASSUMPTIONS ARE MADE ABOUT EXPERIMENTAL CONDITIONS,6 3' 3  T~  ?0 DDynaFit input:  N  ?"`P0  #s[mechanism] E + S ---> E + P : kdp E + I <===> EI : kai kdi EI ---> X : kxttt) l P E P,$D 0   T̊  ? p \&Automatically generated fitting model:'' D  H A ?"`0PH  0޽h ? ̙335-___PPT10 +UD'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*E %(D' =-o6Bdissolve*<3<*E +-  0 $$ -D $(   x  c $d `   XB  0D`X  0A?`m)  <D  CTHE ACTUAL ENZYME CONCENTRATION SEEMS HIGHER THAN THE NOMINAL VALUE^D  3 3 3 3 3~  H A ?"`  T ?"`  @B  ZH  ?\  0  JE + I EI X0&vB  N D? z vB  @ N D? z vB  N D? Z    `  ? *  `kinact$    `  ? M  ]kon$   ZX  ?<  ZE + S E + P&3vB  N D?      `,  ? 0  {kcat/Km>    Z4  ? S  ^koff$  + TX!  ?04  units: mM, minutes@     l ` @ ?  ` @,$D 0   T%  ?  ]kon$    T1  ?  ^koff$  !  T  ? &`  `kinact$  "  T  ? L   {kcat/Km>  rB $  B D?`  rB %  B D? @rB &  B D? @ '  T6  ?   b7.1 1044  (  T  ?   c M-1.sec-1>    ,  T8F  ?   I0.00007& -  TPO  ?   Esec-1$  .  T\W  ? t  I0.00014& 0  T[  ? `  Esec-1$  1  Tj  ? <  D[E]0$  4  TJ  ?` 4  b1.5 1054  5  Tp  ?L   c M-1.sec-1>    7  TPv  ?< E E1.0& 9  TX{  ?< 1 ^nM&rB :  B D?` @@rB ;  B D?` ` @rB <  B D? @l    A   ,$D 0B >  N ?"`  @  H<  ?  *Ki = koff/kon = 1 nM3336  B T  ?Z < r C Tp ?"`|,$D 0 Ki ~ [E]0  tight binding koff ~ kinact not  rapid equilibrium A6H  0޽h ? ̙33___PPT10+p3XDv'  = @B D1' = @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<*A %(D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*C %(D' =-o6Bdissolve*<3<*C +8+0+C 0 +f  0   P 1 i (   x  c $Ɋ `   XB  0D`X  0A?`m  <(̊  CALL THREE ASSUMPTIONS OF THE TRADITIONAL ANALYSIS WOULD BE VIOLATED,D 3. 3 . T\  ? D<4___PPT9 8Enzyme concentration is not much lower than [I]0 or Ki The dissociation of the EI complex is not much faster than inactivation The control progress curve ([I] = 0) is not a straight line (Substrate depletion is significant)       =    ,4 @`l P F 1 P F,$D 02 /  N ?"`P   0  T ?"` F 8bIf we used the traditional algebraic analysis, the results (Ki, kinact) would likely be incorrect.c<   ,<H  0޽h ? ̙335-___PPT10 +vD'  = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*1 %(D' =-o6Bdissolve*<3<*1 +t!  0 ,$`!( (   x  c $ `   XB  0D`X  0A?`m  <   -MORE INFORMATION EXTRACTED FROM THE SAME DATA>. 3 3  3  T,  ?   e'Traditional model (Kitz & Wilson, 1962)"(   TP  ?   IGeneral numerical modeljB  B D?@ @ jB  B D?jB  B D?@@<  TԊ  ?0  JE + I EI X0&pB  H D?HHpB  @ H D?xxpB  H D?\z*\  Z  ?lJ@ lkinact033  Zt  ?`@4 ikon033  Z)  ?d@8 jkoff033<  T  ?p4  JE + I EI X0&pB  H D?HZHpB  @ H D?xZxpB  H D?\j \  Z  ?lH @ lkinact033  Z   ?`u4 hKi033r  T  ?"`P@r  T  ?"`` @ ! T|  ? k  ? very fast  " T8  ? E  ? very slow  # T  ?X  Fno assumptions !l    '   ,$D  0rB $  B D?   %  T  ?f   TTwo model parameters$ &  T  ? `  VThree model parameters$ ( T! ?"` ,$D 0 vAdd another dimension ( la  residence time ) to the QSAR ?<<5H  0޽h ? ̙33___PPT10+i(#Dt' & = @B D/' = @BA?%,( < +O%,( < +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<*' D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*( %(D' =-o6Bdissolve*<3<*( +8+0+( 0 + 0 p 8(   X  0A?`mv  6   RDynaFit in the Analysis of Enzyme Progress Curves Irreversible enzyme inhibitionS  9  <@   APetr Kuzmic, Ph.D. BioKin, Ltd. WATERTOWN, MASSACHUSETTS, U.S.A.pB   XB  0D0XB  0D0!  TZ  ?< r PH@___PPT9" ETOPICS Numerical integration vs. algebraic models: DYNAFIT Case study: Caliper assay of irreversible inhibitors Devil is in the details: Problems to be aware ofp k033 @`H  0޽h ? ̙33y___PPT10Y+D='  = @B +  0 %" (   x  c $\P `   XB  0D`X  0A?`m  <I   /A RANDOM SELECTION OF A FEW TRAPS AND PITFALLS^0 3 3 3 3 3  " Tb  ?R   Residual plots we must always look at them Adjustable concentrations we must always  float some concentrations in a global fit Initial estimates: the  false minimum problem nonlinear regression requires us to guess the solution beforehand Model discrimination: Use your judgment the theory of model discrimination is far from perfect   =  2$   +7 H  0޽h ? ̙33y___PPT10Y+D='  = @B +  0 )* (   x  c $ `   XB  0D`X  0A?`m  < ] &RESIDUAL PLOTS SHOWS THE DIFFERENCE BETWEEN THE DATA AND THE  BEST-FIT MODELpN 3  3  3 3 3 3pB  H D? pB  H D? @   Tg  ? "  6time  T4  ?P; 8signal2  NL  8c?"` 2  N ?"`@p2  N ?"`0 P 2  N ?"``2  N ?"` 2  N ?"`  |B  T D?p |B  T D?  ` |B  T D?|B  T D?   T  jJ?"` 0   T  ?p = =model   T$  ?@M <data 3  T  ?@ @residual  9 l     *   ,$D  0rB   B D? p ~B   N DjJ?  2   N ?"`P P B   T D?  2   N ?"` 0 B   T D? 2   N ?"`PB   T D?  2 !  N ?"`@@B "  T D?p2 #  N ?"``p $  T  ? T <residual  %  T<  ?p   6time &  T ?"` p  1+ '  TL ?"`  2 - (  T  jJ?"``  )  TxÞ ?"` @   4 0H  0޽h ?  ̙33___PPT10+╊D|' = @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<** +  0    A (   x  c $Ξ `   XB  0D`X  0A?`m2  <Ϟ   nTHESE TWO PLOTS LOOK  INDISTINGUISHABLE , DO THEY NOT ?<8 3 3 3p  H A ? P p  H A ?   <  T֞  ?  JE + I EI X0&pB  H D?ZpB  @ H D?8Z8pB  H D?:  Z  ?,  lkinact033  Z|  ? E  ikon033  Z  ?$P jkoff033  T\6  ? @\ FE + I XpB  H D? P   ZX;  ?   lkinact033  Tȇ  ?U SOne-step inhibition$   TH  ?  STwo-step inhibition$ H  0޽h ? ̙33y___PPT10Y+D='  = @B +~  0 d\ " (   x  c $ `   XB  0D`X  0A?`m,  <  hTHESE TWO PLOTS LOOK  VERY DIFFERENT , DO THEY NOT ?<5 3 3 3<  T  ?  JE + I EI X0&pB  H D?ZpB  @ H D?8Z8pB  H D?:  Z  ?,  lkinact033  Z  ? E  ikon033  Z  ?$P jkoff033  T#  ? @\ FE + I XpB  H D? P   Zx  ?   lkinact033  T3  ?U SOne-step inhibition$   T8  ?  STwo-step inhibition$ p  H A ? P p  H A ?   bl p`   p` ,$D 0   Z  8c?"`P `    T  8c?"` `p0    T\/  ?# L horseshoe     T|,  ?0 @  log Gl  `  "  ` ,$D 0   TlA  ? @  <+2.5@  `  !  ` B  # l D8c?p B  # l D8c?0 0    TG  ? :+5   T,-  ?p 0  ;-10   THM  ? `  :-5H  0޽h ? ̙33___PPT10+JD~'  = @B D9' = @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<*" %(+C  0 5 - )  (   x  c $xV `   XB  0D`X  0A?`m  <X  pWE DON T HAVE TO RELY ON VISUALS ( LOG VS.  HORSESHOE )99 3  T]  ?U SOne-step inhibition$   Tb  ? . STwo-step inhibition$ ~  H A ?"` 8x ~  H A ?"`  x p ! H A ? ? p # H A ?  Yl 0 (  (  0( ,$D 0 $  T  8c?"`  `  %  T  8c?"`` P  &  Ti  ?0T U (  Dprobability 0%3 '  Tn  ?L   Eprobability 31%3 ) Tr  ?  ,$ 0 `passes p > 0.05 test0 H  0޽h ? ̙33___PPT10+}5D:'  = @B D' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*( %(D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*) %(D' =-o6Bdissolve*<3<*) +8+0+) 0 +  0  % (   x  c $ `   XB  0D`X  0A?`m:  <D n tSOMETIMES IT S O.K. TO HAVE  OUTLIERS  USE YOUR JUDGMENT>;) 3 3 3~ ! H A ?"`0 i " T  8c?"`P pB # @ H D1? P " $ T  ?` h j something happened with the first three time-points666 % T'  ?v It s not always easy to judge  just how good the residuals are:AA H  0޽h ? ̙33y___PPT10Y+D='  = @B +?  0 f^  (   x  c $ܸ`   XB  0D`X  0A?`m  <  DWE ALWAYS HAVE  TITRATION ERROR !N# 3 3 3 3p  H A  ?   T ?n qResidual plots: fixed [I]<pB  H D>? p  H A! ?    Tll ?  sResidual plots: relaxed [I]<H  0޽h ? ̙33y___PPT10Y+D='  = @B +D   0 k c    (   x  c $w`   XB  0D`X  0A?`m  <y  DWE ALWAYS HAVE  TITRATION ERROR !N# 3 3 3 3  Tl ?n qResidual plots: fixed [I]<  T ?  sResidual plots: relaxed [I]<p  H A ?  p  H A" ?  @   T  8c?"`    ZO  8c?"`` 0   T ?( * 00 #nD = 100, nP = 55 nR = 44 p = 0.08$   3 36  TH ?( 0 (nD = 100, nP = 44 nR = 18 p < 0.0000001)   336H  0޽h ? ̙33y___PPT10Y+D='  = @B +   0    (   x  c $ `   XB  0D`X  0A?`m0  <( } HONE (USUALLY ANY ONE) OF THE INHIBITOR CONCENTRATIONS MUST BE KEPT FIXED`I 3  3 38 3 3  N ?"`  ... [data] directory ./users/COM/.../100514/C1/data extension txt file 0nM | offset auto ? file 2p5nM | offset auto ? | conc I = 0.0025 ? file 5nM | offset auto ? | conc I = 0.0050 ? file 10nM | offset auto ? | conc I = 0.0100 file 20nM | offset auto ? | conc I = 0.0200 ? file 40nM | offset auto ? | conc I = 0.0400 ? ...   @Y3353l3633b4424  T) ?0 EDynaFit script:"  N ?"` `   T(.?"`` 4  9FIXEDH  0޽h ? ̙33y___PPT10Y+D='  = @B +  0 0' (   x  c $5`   XB  0D`X  0A?`m  <7 ANONLINEAR REGRESSION REQUIRES US TO GUESS THE SOLUTION BEFOREHANDNB$ 3 3 3 3~  H A ?"`o   NX? ? @ xl[constants] kdp = 10 ? kai = 10 ? kdi = 1 ? kx = 0.01 ? [concentrations] S = 0.85 ?m P   ,  TN ?` !  Hinitial estimate   NR?"`P  <LE + S ---> E + P : kdp E + I <===> EI : kai kdi EI ---> X : kxXM t l     ` ,$D 0   N ?"`  p   T ? D  jiterative refimenement l   $  ,$D  0   N A# ?"`   !  T ?  sM kdp = 72 kai = 4.5 kdi = 1.8 kx = 0.048 [S] = 0.22N   P    #  Ta ? =  L best fit   l    '   ,$D  0 %  H A$ ?"`   &  TD. ' = @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<*$ 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<*' +"  0 p Y(   x  c $? `   XB  0D`X  0A?`m@  <Ԋ  HIN UNFAVORABLE CASES EVEN ONE ORDER OF MAGNITUDE DIFFERENCE IS IMPORTANTpI 3  3  3 3 3 3  N䓰  ? @ xl[constants] kdp = 10 ? kai = 0.1 ? kdi = 0.01 ? kx = 0.1 ? [concentrations] S = 0.85 ?m   P   ,  N% ?"`P  <LE + S ---> E + P : kdp E + I <===> EI : kai kdi EI ---> X : kxXM t z      ` ,$D 0   N ?"`  p   T02  ? D  jiterative refimenement ~  H A% ?"`o   T7  ?0   Hinitial estimate l  J   J,$D  0   T;  ?  sM kdp = 96 kai = 0.36 kdi =0.0058 kx = ~ 0 [S] = 0.27N   P      H A& ?"` J    TN  ?0   @best fit  l  J    J ,$D  0   H A' ?"` J    TLS  ?P`   A residuals  l  p   p,$D 0   T  8c?"` Pp{   NpW ?"` Z 5E + S ---> E + P : kdp E + I <===> EI : kai kdi>6P H  0޽h ? ̙33]U___PPT105+mD' = @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<* 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' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* +  0 . &    (   x  c $$`   XB  0D`X  0A?`m"  < LDYNAFIT-4 ALLOWS HUNDREDS, OR EVEN THOUSANDS, OF DIFFERENT INITIAL ESTIMATESNM 3 3 3 3  Nt?"``P &  [mechanism] E + S ---> E + P : kdp E + I <===> EI : kai kdi EI ---> X : kx [constants] kdp = { 10, 1, 0.1, 0.01 } ? kai = { 10, 1, 0.1, 0.01 } ? kdi = { 10, 1, 0.1, 0.01 } ? kx = { 10, 1, 0.1, 0.01 } ?*) $  T ?   j Try all possible combinations of initial estimates. 66  T ? Q  8MEANS:   T`< ? ,$  0  4 rate constants 4 estimates for each rate constant 4 4 4 4 = 44 = 256 initial estimates d:33333H  0޽h ? ̙33___PPT10+. D' = @B DD' = @BA?%,( < +O%,( < +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 +V  0 ` (   x  c $E`   XB  0D`X  0A?`m   <G 4DYNAFIT IMPLEMENTS TWO MODEL-DISCRIMINATION CRITERIAN5 3 3 3  3T  TI ?" d L1. Fischer s F-ratio for nested modelsh' <  T ? d  .2. Akaike Information Criterion for all modelsL/&jB  B D?00~  H A ?"`0~  H A ?"` 0$  Tx ?@0" gOne-step model:<   T  ?0g  gTwo-step model:< ~  H A f?"` l |z  t     t,$D 0   T  8c?"`P 0 @   Th ? t Jprobability (0 .. 1)3~  H A f?"`Pn l 0$  0$,$D  0   H A( ?"`0   H A) ?"`0 $sl 02 @  20 @,$D 0   T  8c?"`0 @rB  B D?@`    T?"`V 2 @ Irelative sum of squaresH  0޽h ? ̙33  ___PPT10 ++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' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* +>  0   @  A (   x  c $%`   XB  0D`X  0A?`m   <' JNUMERICAL MODELS ENABLE US TO DO MORE USEFUL EXPERIMENTS IN THE LABORATORYNK! 3 3 3 32  H ?Pp   No constraints on experimental conditions EXAMPLE: large excess of [I] over [E] no longer required No constraints on the theoretical model EXAMPLE: dissociation rate can be comparable with deactivation rate Theoretical model is automatically derived by the computer No more algebraic rate equations Learn more from the same data EXAMPLE: Determine kON and kOFF, not just equilibrium constant Ki = kOFF/kON+  # *      <"                 Pk B  T, ?0c zADVANTAGES of  Numerical Enzyme Kinetics (the new approach):(> 34l  p `  p `,$D 0A   H ?t `  Change in standard operating procedures Is it better stick with invalid but established methods ? (Continuity problem) Training / Education required Where to find time for continuing education ? (Short-term vs. long-term view)  "   !       #    T ? M  RDISADVANTAGES:( 3 rB   B D? p p 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<* +R  0 yqP   (   x  c $ޥ`   XB  0D`X  0A?`m  Tĵ ?vZ  OMORE INFORMATION AND CONTACT:{  T ?Z   cBioKin Ltd. Software Development Consulting Employee Training Continuing Education since 1991x K 3L    Tp ?pZ D DPetr Kuzmic, Ph.D.  T ? f   mhttp://www.biokin.comjB  B D?  H  0޽h ? ̙33y___PPT10Y+D='  = @B + 0 @0 <( @Y@ 0 X  C 4     S  49 f   >@  H 0 0ηo~ ? ̙33  0 T p(  T ^ T S 4    T # l԰ss49 f    H T 0ηo~ ? ̙33 0 x T(  x d x c $4   V x s *`V49 f  V >@  H x 0ηo~ ? ̙33 0  T(   d  c $4     s *TV49 f   >@  H  0ηo~ ? ̙33r@IP>VCr} c]trUʇC,!2]:R/v*iB)ѩ-;\d;xl>КV/sYj> -Gz1Oh+'0T hp  Algebraic Model..497Microsoft PowerPoint@ҥ0@E1(@s}*@Iv?GSg  )'    """)))UUUMMMBBB999|PP3f333f3333f3ffffff3f̙3ff333f333333333f33333333f33f3ff3f3f3f3333f33̙33333f333333f3333f3ffffff3f33ff3f3f3f3fff3ffffffffff3ffff̙fff3fffff3fff333f3f3ff3ff33f̙̙3̙ff̙̙̙3f̙3f333f3333f3ffffff3f̙3f3f3f333f3333f3ffffff3f̙3f3ffffffffff!___www14'A x(xKʦ """)))UUUMMMBBB999|PP3f3333f333ff3fffff3f3f̙f3333f3333333333f3333333f3f33ff3f3f3f3333f3333333f3̙33333f333ff3ffffff3f33f3ff3f3f3ffff3fffffffff3fffffff3f̙ffff3ff333f3ff33fff33f3ff̙3f3f3333f333ff3fffff̙̙3̙f̙̙̙3f̙3f3f3333f333ff3fffff3f3f̙3ffffffffff!___www¼rrIrrrrrnnrr ՜.+,0D    On-screen Show.] & ,Times New RomanVerdanaArialSymbol Courier NewDefault DesignSlide 14Algebraic solution for time course of enzyme assays$Enzyme kinetics in the real world.Progress curvature at low initial [substrate]2Problems with algebraic models in enzyme kinetics0Numerical solution of ODE systems: Euler method/Automatic derivation of differential equationsSoftware DYNAFIT (1996 - 2010)"DYNAFIT: What can you do with it?2DYNAFIT applications: mostly biochemical kinetics Slide 110Traditional analysis of irreversible inhibition+Traditional analysis Take 2: nonlinear1Traditional analysis: Three assumptions (part 1)1Traditional analysis: Three assumptions (part 2)1Traditional analysis: Three assumptions (part 3)/Simplifying assumptions: Requirements for data;Actual experimental data (COURTESY OF Art Wittwer, Pfizer)'Numerical model for Caliper assay data/Caliper assay: Results of fit optimized [E]07Caliper assay violates assumptions of classic analysis0Numerical model more informative than algebraic Slide 23(Numerical modeling looks simple, but...Residual plots Direct plots of data: Example 1Residual plots: Example 1#Residual plots: Runs-of-signs testResidual plots: Example 2,Relaxed inhibitor concentrations: Example 15Relaxed inhibitor concentrations: Example 1 (detail)2Relaxed inhibitor concentrations: not all of them/Initial estimates: the false minimum problem4Large effect of slight changes in initial estimates6Solution to initial estimate problem: systematic scan(Model discrimination: Use your judgmentSummary and Conclusions Questions?  Fonts UsedDesign Template Slide Titles&_+{0..  !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ\]^_`abdefghijwRoot EntrydO)PictureswCurrent UsercSummaryInformation(0TPowerPoint Document(rO{DocumentSummaryInformation8[