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HG8( ?  o2$ץt0xrv4s "$'ϓol5kBb$ s?)rF42$ݵ0D.*=2$ed7K&3qa?2$ANsiW] -A"$z?O\c=TxB"$Dw!W"J J2$bL'v.՜FUuCk2$I VV8 k( l2$ky %1azzn2$H_{}##\-p2$gY N߀0Cɮr2$bȷe rt2$, {dˀetu v"$߼(<n,\/x2$f)#":`^(;u "$e}Ip"2$Iֳ߁p)b$S1)`ҮYЄhSb$F']#%<f6b$Ԛ'~KۤDW$b$B<3\#h8=¬2$olrLI,}O, F2$v" rf{Pb$@I G>Pb$/w9Pͯd_>b$!KO)=Mb$Ru2 $3$b$ΑČ $ +ub$τo#  Lecture outline><Optimal Experimental Design: Where to find basic information==^-Theory of D-optimal designx43D-Optimal design example: Michaelis-Menten equation4452Realistic design for the Michaelis-Menten equation3367Theory of D-optimal design: The chicken-and-egg problem888<Special case: Time-course experiments with fixed time-points&= 098A typical kinetics experiment: Fixed mesh of time-points99:-Stopped flow experimentation: Fixed step size..;<Variation of the design problem: Optimize initial conditions==<=Theory of D-optimal design: Initial conditions in ODE systems>>=-Optimize initial conditions: DynaFit notation..?;Optimize initial conditions: Algorithm and DynaFit settings6<+ 71Case study: Kinetics of clathrin cage disassembly&2 'C)Clathrin structure: triskelions and cages**E%Clathrin biology: Role in endocytosis&&A<In vitro kinetics of clathrin disassembly: Experimental data(=5D=In vitro kinetics of clathrin disassembly: Theoretical models(>6F;In vitro kinetics of clathrin disassembly: DynaFit notation(<4B>In vitro kinetics of clathrin disassembly: Preferred mechanism(?7G3In vitro kinetics of clathrin disassembly: Raw data(4,H$How many assays are actually needed?%%I,Optimal Experimental Design: DynaFit results--J9Optimal Experimental Design: DynaFit results - discussion::@/Optimal Experimental Design in DynaFit: Summary00/hCby   0` ̙33` ` ff3333f` 333MMM` f` f` 3>?" dd@,|?" dd@   " @ ` n?" dd@   @@``PR    @ ` ` p>> <4(    6~ P  T Click to edit Master title style! !  0   RClick to edit Master text styles Second level Third level Fourth level Fifth level!     S  08s ``  X*  0x `   l*(    0 `   `* H  0޽h ? ̙33 Default Design 0 P (  4 4 NpF y y .   p* a00aa 4 N y y 3 .  r* a00aad 4 c $ ?  4 4 Nq y y 9 f  RClick to edit Master text styles Second level Third level Fourth level Fifth level!     S  4 T y y r   p* a00aa 4 TD y y r3   r* a00aaH 4 0ηo~ ? ̙3380___PPT10.mz$X \x(  \ \ N]e]e     p* AAA \ ND]e]e %    r* AAA  \ Tp]e]e    p* AAA \ T0]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 ZBinding and Kinetics for Experimental Biologists Lecture 8 Optimal design of experimentsN[0  9 % Z ?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 !(  x  c $`   XB  0D`X  0A?`m  T_- ?0 5  %The problem: How should we plan an experiment such we learn the most from it? The solution: Use the Optimal Design Theory of statistics An implementation: Software DynaFit An example: Kinetics of clathrin cage disassembly 3  ! 3   3    3&  @`H  0޽h ? ̙33y___PPT10Y+D=' = @B +  0   }(   x   c $``   XB   0D`X   0A?`m   <8`- iDOZENS OF BOOKS>  3 33X   0A?@ X   0A?`V h   <'- `   j Fedorov, V.V. (1972)  Theory of Optimal Experiments Fedorov, V.V. & Hackl, P. (1997)  Model-Oriented Design of Experiments Atkinson, A.C & Donev, A.N. (1992)  Optimum Experimental Designs Endrenyi, L., Ed. (1981)  Design and Analysis of Enzyme and Pharmacokinetics Experiments 6%!#,2 B._H   0޽h ? ̙33y___PPT10Y+D=' = @B +(  0L0 " -\(  \x \ c $`A-`  - XB \ 0D`X \ 0A?`m\ \ <b- vMAXIMIZE THE DETERMINANT ( D ) OF FISHER INFORMATION MATRIX^<  3  3  3 3 3p \ S A 0??#" `?ox 0 \ T -?"0@NNN?Nu f algebraic fitting function xi independent variable, ith data point (i = 1, 2, ..., N) yi dependent variable, ith data point p vector of M model parameters                      >6+l @`S #\`@S,$D 0x \ S A 1??#" `?@` 1  \ Tl-?"0@NNN?NvSQ -Csi,j sensitivity of f with respect to jth parameter, ith data pointD            6"  ,l @9  *\9@ ,$D 0~ !\ c &A  2??#" `?@V  2 $\ ZF-?"0@NNN?Na  6Fi,j (i,j)th element of the Fisher information matrix\7       $+~ %\ c &A  3??#" `?`9  3Sl S  -\S ,$D  0 (\ 2A  4?jJ?#" `?@0 G 4 +\ T4P-?"0@NNN?NS d  WD-Optimal Design:*  3  " ,\ Tr-?"0@NNN?NF  ^Choose the independent variable x1, ..., xN (e.g., total or initial concentrations of reagents) such that the determinant of F is maximized.  S)dH \ 0޽h ? ̙33  ___PPT10n +DR ' = @B D ' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*#\%(D' =-o6Bdissolve*<3<*#\D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<**\%(D' =-o6Bdissolve*<3<**\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<*-\+*2  0L0 !! G Y!(  ~  s *-`  - XB  0D`X  0A?`m  <-. xRONALD DUGGLEBY  UNIVERSITY OF QUEENSLAND, AUSTRALIA (1979)^= 3 3( 3 3 3.  T$[?"0@NNN?Nu "vi initial rate of enzyme reaction, ith data point Si substrate concentration, ith data point (i = 1, 2, ..., N) V, K vector of model parameters K ... Michaelis constant, V ... maximum rate   "                   ,$(mp  S A <??#" `? <l `  ) E,$D 0 # Z[?"0@NNN?N m Gsensitivity functions x % S A  ???#" `?` ?x ' S A @??#" `?`  @'l v   -vt ,$D 0+ * T[?"0@NNN?Nv   gBox-Lucas two-point design:0    x + S A A??#" `? L  A.l F  j  1 ` ,$D 0 . N?"6@`NNN?N p  / T[?"0@NNN?NF   S1  33333333b 0 Hl[?"0@NNN?Np j  S2 = K  3333333y l  @ < @`,$D 0 72  0e0eA D    ?A A5% 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E|| "0e@     @ABC DEEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `abN E5%  N E5%  N F   5%    !"?N@ABC DEFFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `ab @ D [1 ; Tl[?"0@NNN?NF T Dt  mK = 0.5 V = 1.0B    l @ 9 F@ ',$D 0B ? BD?"0@NNN?NB @ BD?"0@NNN?NB AB ND?"0@NNN?N9B BB ND?"0@NNN?N>2 = T?"0@NNN?N G 2 > T?"0@NNN?NG@ C Z0 [?"0@NNN?N  LS1.33 D ZD[?"0@NNN?N@ S  LS2.33Z G T[?"0@NNN?NM "J. Theor. Biol. 81, 671-684 (1979)>#   H  0޽h ? ̙33___PPT10+D' = @B DX' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*)%(D' =-o6Bdissolve*<3<*)D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*-%(D' =-o6Bdissolve*<3<*-D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*1%(D' =-o6Bdissolve*<3<*1D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*<%(D' =-o6Bdissolve*<3<*<D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*F%(D' =-o6Bdissolve*<3<*F+,  0L0 7 / F@(  ~  s *2[`  [ XB  0D`X  0A?`mx  <9[=  INFINITE SUBSTRATE CONCENTRATION (TO GET Vmax) IS IMPOSSIBLE TO ACHIEVE<J, 3 3 3+  Z?[?"0@NNN?N`  ?Box-Lucas two-point design with one point (Smax) already given:X@        +]  NM[?"0@NNN?N2   S2 = 3333333p + S A J??#" `?d Jj - T(X[?"6@`NNN?N`  assume K = 0.5, Smax = 2.0P       8   ;  72  0e0eA L    ?A A5% 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E|| "0e@     @ABC DEEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `abN E5%  N E5%  N F   5%    !"?N@ABC DEFFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `ab  L [2 9 N?"0@NNN?N` : Te[?"0@NNN?N LS1.33l @  ?@ ,$D 0  T?"6@`NNN?N  Z,i[?"0@NNN?N,z  S1 = Smax  333333333B < BD?"0@NNN?N  f = Zm[?"0@NNN?Ni z  Smax K`3333j > ZC[?"0@NNN?N< @   Smax + 2 K` 3333l  = ] E= ],$D 0B @ ZD3fjJ?"0@NNN?N y B A ZD>?"0@NNN?Nk k ^ 2 B N?"0@NNN?N5 e  C T|[?"6@`NNN?N = $  LS2.33 D T([?"0@NNN?N ] >0.3333 3J F Tؐ[?"6@`NNN?N  ,$D  0 TRECIPE In the determination of KM, always include a substrate concentration that corresponds to a reaction rate approximately one half of maximum achievable rate.       ! 3    H  0޽h ? ̙33  ___PPT10 +QV.D_ ' [= @B D ' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*?%(D' =-o6Bdissolve*<3<*?D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*E%(D' =-o6Bdissolve*<3<*ED{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*F%(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*FD' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*F+8+0+F0 +   0L0   @ 7 ( (  x  c $̮[`  [ XB  0D`X  0A?`m  <x[ GPROBLEM: TO DESIGN AN EXPERIMENT WE MUST FIRST GUESS THE FINAL ANSWER! ,H 3A 3 2  Z[ ?  'optimal design for very few data points>( 2  T4B[ ?`  'perform a smallest possible experiment @(  2  Z)[  ? @ 2guess model parameters (literature, hunches, ...)@3   p  H ?@ v  N ?  2   Z[ ?` p \refine parameter estimates&p " H ? 0 ` v # NK ?  % T[ ? P  Voptimal design theory use here  & T([ ? x <repeat3 6 TxL?"0@NNN?NPX [SEQUENTIAL OPTIMAL DESIGN&  7 T[ ?p p@  D START HERE   3H  0޽h ? ̙33y___PPT10Y+D=' = @B +q  0L0 x (  x  c $4L`p  L X  0A?`mH  0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0 )!  ' (  ~  s *dL  L XB  0D`X  0A?`m  <pL GPROBLEM: WE DO NOT ALWAYS HAVE A CHOICE OF INDEPENDENT VARIABLE VALUES ,H 3A 3  HA?"0@NNN?N ` #0  0e0eA f    ?A A5% 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E|| "0e@     @ABC DEEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `abN E5%  N E5%  N F   5%    !"?N@ABC DEFFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `ab  f L % T8L?"0@NNN?N O  F[Hsc70] = 1 M  ' TL?"0@NNN?NM FRothnie, Kuzmic, et al. (2011) Proc. Natl. Acad. Sci. USA 108, 6927-35pG       $*H  0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0   y(  ~  s * L`Pp  L  0  0e0eA i    ?A A5% 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E|| "0e@     @ABC DEEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `abN E5%  N E5%  N F   5%    !"?N@ABC DEFFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `ab> 0  i LXB  0D`X  0A?`m  <Lo wAAT BEST, WE CAN CHANGE THE (FIXED) STEP SIZE FOR A GIVEN INTERVALBB 3   TL?"0@NNN?N NRothnie, Kuzmic, et al. (2011) Proc. Natl. Acad. Sci. USA 108, 6927-35, Fig. 4O        $* 0  0e0eA k    ?A A5% 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E|| "0e@     @ABC DEEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `abN E5%  N E5%  N F   5%    !"?N@ABC DEFFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `ab   k LQ  TL?"0@NNN?N ~ #t = 0 to 5 sec: step size 20 msec4$S  T`bL?"0@NNN?N #0  %t = 5 to 10 sec: step size 100 msec4&3!H  0޽h ? ̙33y___PPT10Y+D=' = @B +   0L0   P  (  x  c $`L`  L XB  0D`X  0A?`m  <lL IF WE CAN T CHOOSE OBSERVATION TIME, AT LEAST WE CAN CHOOSE INITIAL CONCENTRATIONS ^T< 3 3 3 3 3  T,L?"0@NNN?N4 DBASIC PRINCIPLE:   TL?"0@NNN?NV!8   In kinetic studies - the independent variable is time; - initial concentrations are considered  parameters of the model. D-Optimal design theory is concerned with optimal choice of independent variable - in this case the observation time. Unfortunately, in the real-world we cannot choose particular observation times: - usual instruments are offering us only a fixed mesh of output points. But we can turn things around and - treat initial concentrations as  independent variables . - Then we can optimize the choice of initial concentrations, using the usual formalism of the D-Optimal Design theory.o          N   7 }  H  0޽h ? ̙33y___PPT10Y+D=' = @B +(  0L0 p >0 (  x  c $D7L`  L XB  0D`X  0A?`m\  <xL vMAXIMIZE THE DETERMINANT ( D ) OF FISHER INFORMATION MATRIX^<  3  3  3 3 3p  S A  t??#" `?@f t  TjJ?"6@`NNN?NP0  TL?"0@NNN?N Finitial value problem (first-order ordinary differential equations) c ... vector of concentrations k ... vector of rate constants c0 ... concentrations at time zero 0        " p  S A u??#" `?  uwl m  .m ,$D 0x  S A v??#" `?[ v{  TteL?"0@NNN?Nm  yi experimental signal at ith data point (time ti) c concentrations at time ti r vector of molar responses and/or offset on signal axis                8 H: # c &A  x??#" `?* [ 8 x$D 0 + N?"0@NNN?N - N?"0@NNN?Nul    /  ,$D 0y ! ZoL?"0@NNN?N   usi,j sensitivity of f with respect to jth parameter, ith data point p ... model parameters: vectors k and r combinedv                  6" >x , S A |??#" `?   | : N?"0@NNN?N5l @` = ;` @=,$D  0! 7 ZlL?"0@NNN?N@  WD-Optimal Design:*  3   9 8A ?8c?#" `? ` = l U 9 >U 9,$D  02 < N8c?"6@`NNN?NU2 = N8c?"6@`NNN?N 9H  0޽h ? ̙33___PPT10+lD' = @B Dn' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*.%(D' =-o6Bdissolve*<3<*.D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*/%(D' =-o6Bdissolve*<3<*/D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*#%(D' =-o6Bdissolve*<3<*#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<*>+   0L0   (  x  c $H`   XB  0D`X  0A?`m  <$  ]'THE SOFTWARE TAKES CARE OF ALL THE MATH(( 3  T)?"0@NNN?NzY  ,v[task] data = progress task = design [mechanism] ... [data] set ... concentration X = 1 ?? (0.01 .. 100)w3933Nl  0 {  0 { ,$D 0   T>?"6@`NNN?N^0  B   TD?"0@NNN?N  D   Z|&?"0@NNN?N  {  z<this value is ignored (present for syntactical reasons only)"=' Nl    ,$D 0   T>?"6@`NNN?N0 p B  TD?"0@NNN?N0 D  Z$?"0@NNN?N z.syntax otherwise used for confidence intervals0/,l 0   0  ,$D 0  T>?"6@`NNN?N0 0 B  ND?"0@NNN?N (  T8?"0@NNN?Nc   d$lower and upper bounds must be given$%H  0޽h ? ̙33  ___PPT10 +FD ' = @B D ' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-o6Bdissolve*<3<*D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-o6Bdissolve*<3<*D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-o6Bdissolve*<3<*+|  0L0  $"(  $x $ c $C`   XB $ 0D`X $ 0A?`m $ <E >THE DIFFERENTIAL EVOLUTION ALGORITHM REQUIRES SPECIAL SETTINGS<? 3 3% 3o $ TK?"0@NNN?NV  O[task] data = progress task = design [mechanism] ... [settings] {DifferentialEvolution} PopulationSizeFixed = 300 MaximumEvolutions = 1 MinimumEvolutions = 1 TestParameterRange = n TestParameterRangeAll = n TestParameterRangeFull = n StopParameterRange = 0.1 TestCostFunctionRange = y StopCostFunctionRange = 0.01 TestCostFunctionChange = y StopCostFunctionChange = 0.00001 TestCostFunctionChangeCount = 5 NP  3  J  $ N>?"6@`NNN?NpP `  B $ ND?"0@NNN?N`  $ T`?"0@NNN?NpF   Xpopulation not too large$    $ N>?"6@`NNN?N0 P ` B $ ND?"0@NNN?N `  $ T`f?"0@NNN?NC F   b"perform the optimization only once$#   $ N>?"6@`NNN?N P ` B $ ND?"0@NNN?N `    $ TTl?"0@NNN?N F `  d$relatively weak convergence criteria$%  i $ Tn?"6@`NNN?NH ,$D 0 sAcopy these settings from one of the distributed example problemsBBH $ 0޽h ? ̙33zr___PPT10R+)oiD' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*$%(D' =-o6Bdissolve*<3<*$+8+0+$0 +q  0L0 x` (  x  c $<|p`   X  0A?`mH  0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0 0  @i(  @x @ c $X`   XB @ 0D`X @ 0A?`m# @ <d MCLATHRIN CAGES ARE LARGE ENOUGH TO VISIBLE IN MICROSCOPY AND LIGHT SCATTERINGNN. 3  3 3 3 @ HA#?"0@NNN?NS  @ TX?"0@NNN?N# < j  A clathrin cage   @ T?"0@NNN?N# =  Uclathrin triskelion&   H @ 0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0 |p  P(  Px P c $ܚ`   XB P 0D`X P 0A?`m P <  0CLATHRIN IS INVOLVED IN INTRACELLULAR TRAFICKING,10 3 3&   P HA?"0@NNN?N0r  P T\?"0@NNN?N a *Eisenberg et al. (2007) Traffic 8, 640-646p+        dl P  PP ,$D  0  P T`?"0@NNN?N @   Mclathrin-coated vesicle3  P Tܬ?"0@NNN?N 0   MATP-dependent uncoating2 P H?"6@`NNN?NP H P 0޽h ? ̙33___PPT10+LAED|' = @B D7' = @BA?%,( < +O%,( < +Dn' =%(D' =%(D' =4@BBBB%(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+   0L0    0} (  0x 0 c $$`   XB 0 0D`X 0 0A?`m 0 <T6B GWATCHING CLATHRIN CAGES TO FALL APART BY PERPENDICULAR LIGHT SCATTERING,H) 3 3 0 TL?"0@NNN?NM NRothnie, Kuzmic, et al. (2011) Proc. Natl. Acad. Sci. USA 108, 6927-35, Fig. 4O        $* 0 HA?"0@NNN?N 2 x 0 ZXQ6 ?"0@NNN?N  0  Clathrin cages (0.09 M triskelia) premixed with 0.1 M auxilin were mixed with Hsc70 (concentrations in M shown on graph) and 500 M ATP, and perpendicular light scattering was measured using stopped-flow...         3  J P3N 0 T[6 ?"0@NNN?N`Gp :M Hsc70 H 0 0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0   P  H3 (  Hx H c $0c6 `  6  XB H 0D`X H 0A?`m& H <e6  @MODEL SELECTION USING THE AKAIKE INFORMATION CRITERION (DYNAFIT)<A 3& 3 3! H TDm6 ?"0@NNN?NM FRothnie, Kuzmic, et al. (2011) Proc. Natl. Acad. Sci. USA 108, 6927-35pG       $* H HA?"6@`NNN?NO H HA ?"6@`NNN?N \    H Tv6 ?"0@NNN?N`  NTHREE-STEP SEQUENTIAL: 3  H T{6 ?"0@NNN?Ns|  LTWO-STEP SEQUENTIAL: 3  H HA!?"6@`NNN?NL P"   H T\6 ?"0@NNN?N tXM  MTHREE-STEP CONCERTED: 3v  H T̄6 ?"6@`NNN?N`,$D  0 4Etc. In total five different models were evaluated.05"H H 0޽h ? ̙33___PPT10+f D' 6 = @B DD' = @BA?%,( < +O%,( < +D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* H%(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<* HD' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<* H+8+0+ H0 +U  0L0    X@ (  Xx X c $6 `  6  XB X 0D`X X 0A?`m X <6   e/THE MOST PLAUSIBLE MODEL: THREE STEP SEQUENTIAL00 3 X T$6 ?"0@NNN?NM FRothnie, Kuzmic, et al. (2011) Proc. Natl. Acad. Sci. USA 108, 6927-35pG       $* X N6 ?"6@`NNN?N@  H[task] task = fit data = progress model = AHAHAH ? [mechanism] CA + T  > CAT : ka CAT  > CAD + Pi : kr CAD + T  > CADT : ka CADT  > CADD + Pi : kr CADD + T  > CADDT : ka CADDT  > CADDD + Pi : kr CADDD  > Prods : kd ... [task] task = fit data = progress model = AHAH ? ... Zd(  3  3 3J 8 X Th6 ?"0@NNN?N4 FDYNAFIT INPUT: 3l  3 &  X3 & ,$D 0 X HA"?"0@NNN?Np @ &  X T6 ?"0@NNN?Nj 3? [#AUTOMATICALLY GENERATED MATH MODEL:$$ 3 X N?"6@`NNN?N @ H X 0޽h ? ̙335-___PPT10 +WD' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*X%(D' =-o6Bdissolve*<3<*X+^  0L0 me 8(  8x 8 c $6 `  6  XB 8 0D`X 8 0A?`m 8 <6  OCONCLUSIONS: THREE ATP MOLECULES MUST BE HYDROLYZED BEFORE THE CAGE FALLS APART<P  3 30 3 8 T6 ?"0@NNN?NM FRothnie, Kuzmic, et al. (2011) Proc. Natl. Acad. Sci. USA 108, 6927-35pG       $* 8 HA?"0@NNN?N@H 8 0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0 SK `(  `x ` c $p6 `  6  XB ` 0D`X ` 0A?`m ` <|6   /THIS WAS A VERY EXPENSIVE EXPERIMENT TO PERFORMN0  3 3 3  3 ` Tx6 ?"0@NNN?NM FRothnie, Kuzmic, et al. (2011) Proc. Natl. Acad. Sci. USA 108, 6927-35pG       $*M  ` T$6 ?"0@NNN?Npa ,$ 0 ]M six assays / experiment 90 nM clathrin 100 nM auxilin up to 4 M Hsc70N  3 33,p  ` T6 ?"6@`NNN?N  ,$D  0 z<a lot of material expensive and/or time consuming to obtain"=* X  ` 0A&?[ P   ` < 26  c m 90  ` <$06 pN  <0.25  ` <0>6 pK j   ;0.5  ` <B6 ` C  91  ` <E6  k#  92  ` <I6  K]  94 LB ` c $D;  ` T46 ?"0@NNN?N 0 ;[Hsc70]   ` T@6 ?"0@NNN?N6*  MACTUAL EXPERIMENTAL DATA: H ` 0޽h ? ̙335 - ___PPT10 +˒6D' D6 = @B D<' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* `%(D' =-o6Bdissolve*<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<* `+p+0+ `0 ++0+ `0 +m  0L0 G?  h(  hx h c $x{`  { XB h 0D`X h 0A?`m h < ^ QD-OPTIMAL DESIGN IN DYNAFIT 3   h 6- >[task] task = design data = progress [mechanism] CA + T  > CAT : ka CAT  > CAD + Pi : kr CAD + T  > CADT : ka CADT  > CADD + Pi : kr CADD + T  > CADDT : ka CADDT  > CADDD + Pi : kr CADDD  > Prods : kd [data] file run01 | concentration CA = 0.1, T = 1 ?? (0.001 .. 100) file run02 | concentration CA = 0.1, T = 1 ?? (0.001 .. 100) file run03 | concentration CA = 0.1, T = 1 ?? (0.001 .. 100) file run04 | concentration CA = 0.1, T = 1 ?? (0.001 .. 100) file run05 | concentration CA = 0.1, T = 1 ?? (0.001 .. 100) file run06 | concentration CA = 0.1, T = 1 ?? (0.001 .. 100) file run07 | concentration CA = 0.1, T = 1 ?? (0.001 .. 100) file run08 | concentration CA = 0.1, T = 1 ?? (0.001 .. 100)Z  )())))))u  j  h < -p b 8[constants] ka = 0.69 ? kr = 6.51 ? kd = 0.38 ?v9    ,    h 6jJ"` ,$D 0v  h <- ns ,$ 0  Choose eight initial concentration of T such that the rate constants ka, kr, kd are determined most precisely. s'        ! ,K"H h 0޽h ? ̙33___PPT10+vD:' = @B D' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* h%(D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* h%(D' =-o6Bdissolve*<3<* h+8+0+ h0 +  0 u m p (  px p c $-``  - XB p 0D`X p 0A?`m p <- 9SURPRISE: WE DID TOO MUCH WORK FOR THE INFORMATION GAINED*:9 33X p 0A$?P LB p c $D` `  p <`-b 6  ] = 0.70 M, 0.73 MR LB  p c $D `   p <T->   #[T] = 2.4 M, 2.5 M, 2.5 M p$ LB  p c $D~ `   p <- )  [T] = 76 M, 81 M, 90 Mp   p <- v i!D-Optimal initial concentrations:,"  p <t-"`P  ,$D  0 xJust three experiments would be sufficient for follow-up! One half of the material compared to the original experiment.Xy1 & @z 0   p  f ,$D 0t p 6jJ"`0  : p <d   ~ maximum feasible concentration upswing phase no longer seen,@!  8 p T?"0@NNN?N0g  |HSIMULATED DATA  OPTIMAL EXPERIMENT:%% H p 0޽h ? ̙33A9___PPT10+9WD' -= @B D' = @BA?%,( < +O%,( < +D{' =%(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<*pD4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*p%(+8+0+p0 +   0    t (  tx t c $``   XB t 0D`X t 0A?`mF t <% rEACH OF THE THREE UNIQUE ASSAYS TELLS A DIFFERENT  STORY L:  3 3( 33X t 0A$?BP B t BD?"0@NNN?N  t H?"0@NNN?N P  :mostly disassembly (kd) association ( upswing ) is no longer visiblerF     .  0 t N?"0@NNN?NP  [mechanism] CA + T  > CAT : ka CAT  > CAD + Pi : kr CAD + T  > CADT : ka CADT  > CADD + Pi : kr CADD + T  > CADDT : ka CADDT  > CADDD + Pi : kr CADDD  > Prods : kdN(G B t@ BD?"0@NNN?N@? t H?"0@NNN?N `  mostly ATP association (ka)X     H t 0޽h ? ̙33y___PPT10Y+D=' = @B +&   0 M E ,(  ,x , c $$``   XB , 0D`X , 0A?`m , <,  mNOT A SILVER BULLET !< 3 33 , <dP|  \ Useful for follow-up (verification) experiments only - Mechanistic model must be known already - Parameter estimates must also be known Takes a very long time to compute - Constrained global optimization:  Differential Evolution algorithm - Clathrin design took 30-90 minutes - Many design problems take multiple hours of computation Critically depends on assumptions about variance - Usually we assume constant variance ( noise ) of the signal - Must verify this by plotting residuals against signal (not the usual way) / **  F%9 H , 0޽h ? ̙33y___PPT10Y+D=' = @B + 0 @<( @Y@ X  C 4     S x49 f   >@  H  0ηo~ ? ̙33, 0  |(   d   c $4      3 rass49 f  a  H   0ηo~ ? ̙33,- 0  `|(  `d ` c $4    ` 3 rgss49 f    H ` 0ηo~ ? ̙33,4 0  |(  d  c $4     3 r6ss49 f    H  0ηo~ ? ̙33,5 0 0 |(  d  c $4     3 r;ss49 f    H  0ηo~ ? ̙33,6 0 P |(  d  c $4     3 r?ss49 f    H  0ηo~ ? ̙33,7 0 p |(  d  c $4     3 rEss49 f    H  0ηo~ ? ̙33,8 0  |(  d  c $4     3 r4Kss49 f    H  0ηo~ ? ̙33,9 0  |(  d  c $4     3 rPss49 f    H  0ηo~ ? ̙33,: 0  |(  d  c $4     3 rVss49 f    H  0ηo~ ? ̙33,; 0 `  |(   d   c $4      3 r@\ss49 f    H   0ηo~ ? ̙33,< 0  |(  d  c $4     3 r`ss49 f    H  0ηo~ ? ̙33,= 0  |(  d  c $4     3 rfss49 f    H  0ηo~ ? ̙33,? 0  (|(  (d ( c $4    ( 3 r8lss49 f    H ( 0ηo~ ? ̙33,A 0  4|(  4d 4 c $4    4 3 rpss49 f    H 4 0ηo~ ? ̙33,B 0  <|(  <d < c $4    < 3 rvss49 f    H < 0ηo~ ? ̙33,C 0 @ D|(  Dd D c $4    D 3 rL|ss49 f    H D 0ηo~ ? ̙33,D 0 ` L|(  Ld L c $4    L 3 rss49 f    H L 0ηo~ ? ̙33,E 0  T|(  Td T c $4    T 3 rss49 f    H T 0ηo~ ? ̙33,F 0  \|(  \d \ c $4    \ 3 rDss49 f    H \ 0ηo~ ? ̙33,G 0  d|(  dd d c $4    d 3 r쐻ss49 f    H d 0ηo~ ? ̙33,H 0  l|(  ld l c $4    l 3 rss49 f    H l 0ηo~ ? ̙33xp^RЀ3ÿ lHb@ 000A$R )Đ% y@=)b3 1@Ppzl`)TD\ L!V~gs Hb"~! 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