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!  0a  a RClick to edit Master text styles Second level Third level Fourth level Fifth level!     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"Hunches and intuitive impressions are essential for getting the work started, but it is only through the quality of the numbers at the end that the truth can be told. -Lewis Thomas L. Thomas (1977) "Biostatistics in Medicine", Science 198, 675j3.     F " Tg?"`p6,$D  0 4But how much confidence can you have in that number?05     l u  %  e ,$D 0x #  H A ?u  $  T_g ?   Gregor Mendel (1822-1884)4 l ` ` >  )  ` > ,$D  0x '  H A ?` ` >2  (  Tfg ?`   IGoogle - July 20, 2011l J  0 J| ,$D 0r .  TZ  ?0 P  /  Tig ?pJX  approximately 3:1 G / yV  3 $3  H  0޽h ? ̙33QI___PPT10)+~D' g= @B D' = @BA?%,( < +O%,( < +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?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<*0 %(D' =-o6Bdissolve*<3<*0 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 "(  x  c $4g`  g XB  0D`X  0A?`m:  Td ?` / PThe problem: How much (or how little) can we trust our rate and equilibrium constants? The solution: Always report at least some measure of parameter uncertainty: - formal standard error - confidence interval (a) by systematic search (profile-t method) (b) by stochastic simulations (Monte-Carlo method) An implementation: Software DynaFit. An example: The classic  Biological oxygen demand (BOD) problem 3K 3  {  > 3     35   @`H  0޽h ? ̙33y___PPT10Y+D=' = @B +E  0L0 TL(  X  0A?`m$  Hg?"0@NNN?NO t4Part 1: Confidence intervals by systematic searching$5.   Hd?"0@NNN?N#0  p$ Profile-t method0    H  0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0   0 dZ (  d d  B a"`  a XB d 0D`X d 0A?`m d <0d  t,A CLASSIC DATA SET IN STATISTICAL LITERATURE,-, 33p  d H A ?@ ? A  d Td ?t6~! 9BOD = measure of organic pollution in environmental waterN:    7A  d Td ?p O   eBates D. M. & Watts, D. G. (1988) Nonlinear Regression and its Applications Wiley, New York, p. 270<f# *  ^ d0 3 0e0e A     ? A@  A 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E||@ Q   d^l ` d`,$D 0~B d N D1?@0\ d Tod ?`- *BOD at t infinity ?p 3 3 3 3 3  3H d 0޽h ? ̙33___PPT10+:6D|' = @B D7' = @BA?%,( < +O%,( < +Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*d%(D' =+4 8?dCB0-#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*dD' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*d+E  0L0   p|E (  | |  B d"`y   d XB | 0D`X | 0A?`m | <0d  u-COMPARE ALGEBRAIC MODEL WITH DYNAFIT NOTATION,.- 33jB | B D?@  | T\3d ?p +  TALGEBRAIC MODEL:(  3  | Td8d ?p @ A  RDYNAFIT MODEL:( 3   | T>d ?  # g/[mechanism] Oxygen ---> Bacteria : k 00p  | S A ??#" `?   p  | H A ?0   | TEd?"`P `  @ time, days   | Z5d?"`@ Y BOD, mg/l   | H A ?0T,$D 0&l 0 |0,$D 0 | TBd ?0 BODmax = 19.1 mg/lB     2 | N ?"`H | 0޽h ? ̙33 ___PPT10+PD' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*|%(D' =-o6Bdissolve*<3<*|D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*|%(D' =-o6Bdissolve*<3<*|+i  0L0 7 / P"p (  p p  Bd"`  d XB p 0D`X p 0A?`mF p <Xd  *FIX Bmax AT AN ARBITRARY VALUE, OPTIMIZE kr+ 3 3 3! 3 33#v p N ?((~ p H A ?"`  p Td ? e@f +Bmax = 19.1 k = 0.53 sum of squares = 26.0, 33 3  (l pf !ppf,$D 0 p H A ?"`p  p T#d ? f *Bmax = 25.0 k = 0.28 sum of quares = 41.9+ 3 3  $l ` "p`,$D 0 p H A  ?"`` w p T4d ?  +Bmax = 30.0 k = 0.20 sum of squares = 57.6, 3  (  p Thd?"` B0 #optimized parameter fixed parameterF$  3     H p 0޽h ? ̙33___PPT10+^D~' = @B D9' = @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%(+   0L0  |  (     Bpd"`  d XB  0D`X  0A?`m  <d  y1THIS IS WHAT MOST PAPERS REPORT IN THE LITERATURE,21 33p  H A  ?P ` p  H A  ? `z 0  0  ,$D 0B  T d ?0 BODmax = (19.1 2.5) mg/lf    3 2  N ?"`2  T  V?"``   Td ? 0 p ,$ 0 zimplies the interval 19.1  2.5 = 16.6 19.1 + 2.5 = 21.6>>%    l  2  2,$D  0rB  B D3?    Td ? 2 )[settings] {Output} InferenceBands = y** H  0޽h ? ̙33, $ ___PPT10 + +0D ' d= @B Dk ' = @BA?%,( < +O%,( < +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<*%(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<*+8+0+0 +2  0L0    (     B8d"`^  d XB  0D`X  0A?`m  <dg ~6VERY RARELY REPORTED IN THE LITERATURE (UNFORTUNATELY),76 33z 0o  ! `,$D 0   TTd ?0o PBODmax = (19.1 2.5) [15.0  29.3] mg/l)       3 #2   N ?"`p   H A  ?0@<2  T  V?"`p 02  T  V?"`p 00p  H A  ?@ 0  Td?"`p0  B log [Oxygen]    ZLd?"`0 @  A mean square  H  0޽h ? ̙335-___PPT10 +5yD' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-o6Bdissolve*<3<*+a-  0L0 , (     B̨]"`t  ] XB  0D`X  0A?`m  <H]$ |4A SEQUENCE OF SEVERAL INDEPENDENT LEAST-SQUARES FITS,54 33  T] ?`  FINPUT:$  V  N8] ?` 2 j[mechanism] Oxygen ---> Bacteria : k [constants] k = 1 ? [concentrations] Oxygen = 10 ??RkD  3"  32  T  1?"`~ l S ,S,$D  0  T] ?^ J ALGORITHM:$   a@ qS $qS  T] ?q D<4___PPT9 Perform an initial fit with all parameters optimized Perform a series of follow-up fits focusing on a given parameter 2a.  Freeze the parameter at values progressively further away from optimal 2b. Optimize all remaining parameters 2c. Repeat (2a) and (2b) until sum of squares reaches a  critical value above minimumLB   4    @`  T@] ?   J REFERENCE:$   S  NԒ] ? PS uBates, D. M., and Watts, D. G. (1988) Nonlinear Regression Analysis and its Applications Wiley, New York, pp. 127-130<v& 3  l ` `,$D 0~  N A ?`   Z]?"` B log [Oxygen]    `_]?"`f3 A mean square Tl 0F %0F,$D  0B  < D?"0@NNN?N0--,  NLd]?"0@NNN?NF nSSQmin2  M # <i] ?"6@ NNN?Np},$  0 oSSQcrit2  l Wv +Wv,$D 0B ' ND8c?"0@NNN?N^B ( ND8c?"0@NNN?NW ) Hp]?"0@NNN?NWd =low 3 * Hs]?"0@NNN?Ncv >high 3H  0޽h ? ̙33___PPT10+6aDe' n]= @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' =%(DG' =4@BBB B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-6B'blinds(horizontal)*<3<*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?dCB1+#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*#D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*#D' =%(D' =%(D9' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*+%(D' =-u6Bdiamond(in)*<3<*++8+0+#0 +M  0L0 ),(     BL]"`:  ] XB  0D`X  0A?`m  <]  {3HIGHER CONFIDENCE LEVEL = WIDER CONFIDENCE INTERVAL,43 33 0  0e0eA     ?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[ \]^_ `ab0P   ]  H]?"0@NNN?N @  990%   H]?"0@NNN?NJ 995%    HD]?"0@NNN?N@ 999% B ! C xD8c?"0@NNN?N  B " # lD8c?"0@NNN?N  B #  fD8c?"0@NNN?N 5  $ H ]?"0@NNN?NL  F ;? & H]?"0@NNN?N@  ,$  0 GUpper limit for BODmax could not be determined at 99% confidence level.DH     2l pP  )Pp ,$D 0 ' 0A ?"6@ NNN?NpP B ( ND?"0@NNN?N@@H  0޽h ? ̙33A9___PPT10+WD' ]= @B D' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*)%(D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*&%(D' =+4 8?dCB1+#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<*&D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<*&+8+0+&0 +{  0L0   0 ! (     B #]"`  ] XB  0D`X  0A?`m:  <%] RUPPER LIMITS FOR BIMOLECULAR ASSOCIATION RATE CONSTANTS OFTEN CANNOT BE DETERMINED`S 3  3 3* 33  0A ?"6@ NNN?N  0A ?"6@ NNN?N }  <4.] ?"6@ NNN?NS 7Moss, Kuzmic, et al. (1996) Biochemistry 35, 3457-3464.8           HD7]?"0@NNN?Nt! ? MECHANISM     H;]?"0@NNN?Ns  oCONFIDENCE INTERVAL FOR k48    2  H8c?"6@`NNN?Np02  B8c?"6@`NNN?N b R   H4A]?"0@NNN?N ,$ 0 <k4 = (5 200) [3  ] M-1s-13H  0޽h ? ̙33zr___PPT10R+V aD' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-o6Bdissolve*<3<*+8+0+0 +  0L0 NFp5(     B|R]"`+  ] XB  0D`X  0A?`m  <T] :AN OCCASIONAL SIDE-BENEFIT OF CONFIDENCE INTERVAL SEARCHES*;: 33 #  0e0e    BCDE@F  A5% 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E||hp(@P@8@L0 @     "0e@     @ABC DEEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `abN E5%  N E5%  N F   5%    !"?N@ABC DEFFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `ab/ B $  fDjJ?"0@NNN?N B % ZDjJ?"0@NNN?N  2 & H?"6@`NNN?N@  2 ' H?"6@`NNN?N0 B (@ TD?"0@NNN?N1  ) HX]?"0@NNN?N_ Hinitial estimate  * Hp]?"0@NNN?N }  V false minimum   + H]?"0@NNN?N }  = PARAMETER   , ND]?"0@NNN?N BSUM OF SQUARES l P3  5P3 ,$D 0B - ND?"0@NNN?N@  0 B .B ND?"0@NNN?NP @ 2 1 H3?"6@`NNN?NP R  2 H]?"0@NNN?N 3 ] A CI search   3 H]?"0@NNN?NP +  Fglobal minimum 3B 4B ZD?"0@NNN?N P ` H  0޽h ? ̙335-___PPT10 +D' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*5%(D' =-o6Bdissolve*<3<*5+  0L0   Pe (     B,"`   XB  0D`X  0A?`m8  H8E?"0@NNN?N`#  ( Confidence intervals are asymmetrical for all nonlinear parameters Frequently much wider (more realistic) than formal standard errors Sometimes half-open intervals:  better than nothing , e.g. for bimolecular association Can have mechanistic implications (reversible / irreversible steps) Sometimes CI search helps in falling out of false minima In DynaFit scripts, CIs are requested by the  ?? syntax Should always be reported with their corresponding confidence levels (%) CIs are wider at higher confidence levels Frequently used confidence levels: 90%, 95%, or 99% Computation can be time consuming (many repeated least-squares fits)      <  O  ^  0  =  W  G  ,{pl  {   {  ,$D  0G  H@'E?"6@`NNN?N{   /[settings] {Marquardt} ConfidenceLevel = 9000 B  ND?"0@NNN?N  H  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<*+  0L0 *"(  X  0A?`m&  HPT^?"0@NNN?N v6Part 2: Confidence intervals by stochastic simulations$70  H# ?"0@NNN?N#R  DMonte-Carlo method H  0޽h ? ̙33y___PPT10Y+D=' = @B +   0L0 a(     BL^"`#  ^ XB  0D`X  0A?`m5  ZTa ?P@`"D<4___PPT9 _yPerform an initial fit as usual Perform a large series (> 1000) of follow-up fits 2a. Simulate an artificial data set with random errors superimposed in ideal data 2b. Perform a fit of the artificial data 2c. Compile a histogram of distribution for model parameters from many repeated fits 2d. Determine the range of plausible values for model parameters from the histogramsz           T    @`  Z ? "=  J REFERENCE:$     T5e ? `  zStraume, M., and Johnson, M. L. (1992)  Monte-Carlo method for determining complete confidence probability distributions of estimated model parameters Methods Enzymol. 210, 117 129.^      $H  0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0  0 (     B#"`.  e XB  0D`X  0A?`m  <'&  {)A SINGLE LINE ADDED TO THE DYNAFIT SCRIPT6* 3  3   BAe?"6@`NNN?N3  ?[task] task = fit data = progress confidence = monte-carlo [mechanism] Oxygen ---> Bacteria : k [constants] k = 1 ?? [concentrations] Oxygen = 10 ?? ...@43q D s   BIe?"6@`NNN?N3 `,$D  0 p[settings] {MonteCarlo} Runs = 1000 ... ConcentrationErrorPercent = 0qq , 5 H   HPe?"0@NNN?N @ ,$ 0 d2plus a number of other advanced control parameters33 H  0޽h ? ̙335 - ___PPT10 +"D' [e= @B D<' = @BA?%,( < +O%,( < +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@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* +p+0+ 0 ++0+ 0 +#  0L0 P$ t(     Bce"`  e XB  0D`X  0A?`m  <fe  1HISTOGRAMS OF DISTRIBUTION PLUS CORRELATION PLOTS^2 3 3 3  3 3   HA?"0@NNN?N4G    HA?"0@NNN?N@ c    Toe?"6@`NNN?Nw@ e  >[Oxygen]     Nse?"6@`NNN?N `=  7k    TPwe?"6@`NNN?N   >[Oxygen]  l  ]  #c 0 ,$D 0B  TD?"0@NNN?N @ B  TD?"0@NNN?N @ B  S ~D>?"0@NNN?N     N}e?"0@NNN?N ]  Kconfidence interval 3l     S ,$D 0B B HD?"0@NNN?N  B  HD?"0@NNN?N P  B  S ~D>?"0@NNN?N    Ne?"0@NNN?Nc   ^conf. interval for k. 3 l  P  S #,$D 0B  BD?"0@NNN?N PPB  BD?"0@NNN?N B  3 rD?"0@NNN?N@@P  Ne?"0@NNN?Np# @[Oxygen]  E ! HVe?"0@NNN?Nt0! ,$ 0 ajoint confidence interval,  0 $ H$Ue?"0@NNN?N$} NDistribution of best-fit values from 1000 least-squares fits of simulated dataOOH  0޽h ? ̙33; 3 ___PPT10 +qD ' = @B Dz ' = @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<* D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*!%(D' =-o6Bdissolve*<3<*!+8+0+!0 +A  0L0   p&A (     BDe"`e  e XB  0D`X  0A?`m  <e MCONVEX HULL = SHORTEST PATH COMPLETELY ENCLOSING A GROUP OF POINTS IN A PLANE,N  3C 3  HA?"0@NNN?N0  He?"0@NNN?N ' m+EPS (PostScript) file generated by DynaFit:&, ) nl  p % p,$D 0B B TD?"0@NNN?N pp.   He?"0@NNN?N` vsolid line: convex hull plot>  3  3 3Il P s  &P s ,$D 0B " TD?"0@NNN?N B # TD?"0@NNN?N P E $ He?"0@NNN?N s  3intensity of squares ~ frequency of best-fit values>4    H  0޽h ? ̙33 ___PPT10+˜HoD' = @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<*&+  0L0 '(     BLe"`  e XB  0D`X  0A?`mV  <ei LMONTE-CARLO INTERVALS ARE ALMOST ALWAYS NARROWER THAN PROFILE-t AT 90% LEVELM  3 3 3 3 3  3 3  HA?"0@NNN?N    Te?"6@`NNN?N >[Oxygen]    Ne?"6@`NNN?N }  @ time, days  E  He?"0@NNN?N& f 3 k BmaxP       B  BD?"0@NNN?N B  ND?"0@NNN?N\0 \  H(e?"0@NNN?N v- <low high  )  He?"0@NNN?N  yMONTE-CARLO METHOD (n = 1000)@      He?"0@NNN?N y3 X0.24 1.20 16.0 27.5(   33  He?"0@NNN?N }  'PROFILE-t METHOD (90% confidence level)@(   B  ND?"0@NNN?N0 B  ND?"0@NNN?N0 E   HVe?"0@NNN?N 0 p 0  k BmaxP       B ! BD?"0@NNN?N} } B " ND?"0@NNN?NY : Y  # He?"0@NNN?N} *  <low high   $ HXe?"0@NNN?N 0  X0.20 1.27 15.0 29.3(   3B % ND?"0@NNN?N} : } B & ND?"0@NNN?N} : } H ' N?"6@`NNN?NP,$D  0 X&good agreement between the two methods'' H  0޽h ? ̙33___PPT10+GYD' e= @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 +a   0L0 p h (  (       B4("`  e XB   0D`X   0A?`m    <*% FREAGENT CONCENTRATIONS ARE ALWAYS AFFECTED BY RANDOM TITRATION ERRORS!>G5 3 3 3o   B81?"6@`NNN?NG` # [mechanism] E + S <===> ES : k ks ES ----> E + P : kr [constants] k = 100 ks = 1000 ? kr = 1 ? ... [settings] {MonteCarlo} ConcentrationErrorPercent = 10 [end]R   3"3 t2    !  HA?"0@NNN?NG   "  H<?"0@NNN?N L  Y%Enzyme kinetics: Substrate conversion&&  #  HA?"0@NNN?N 4  OMechanism: Michaelis-Menten 2 $  B?"6@`NNN?NP2 %  B?"6@`NNN?NE2 &  <?"6@`NNN?N  '  N,F?"6@`NNN?N   :time  (  TxJ?"6@`NNN?N@|  ? [product]  H   0޽h ? ̙33y___PPT10Y+D=' = @B +E  0L0 TL  (     BQ"`n  e XB  0D`X  0A?`m  <xT$  ,JOINT CONFIDENCE INTERVAL (AS A CONVEX HULL)>- 3' 3 3 0  0e0eA     ? 5% 8c8c     ?1 d0u0@Ty2 NP'p<'pA)BCD|E|| "0e@     @ABC DEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `abN 5%  N 5%  N    5%    !"?N@ABC DEFGHIJK5%LMNOPQRSTUWYZ[ \]^_ `ab C  B  ND?"0@NNN?N@ @  N [?"6@`NNN?Nj K10% titration error 3B   ND?"0@NNN?Nr "    HP`?"0@NNN?N`  Qerror-free concentrations H  0޽h ? ̙33y___PPT10Y+D=' = @B +K  0L0   p  <6 (  < <  Bi"`   XB < 0D`X < 0A?`mb < H,l?"0@NNN?Ne  Method makes no assumptions about the statistical distribution of model parameter errors Often uncovers  strange effects such as half-open confidence intervals - mechanistic implications (reversible / irreversible steps) Reveals special patterns in the statistical correlation between model parameters Does not require an arbitrary choice of confidence levels (%) x    F  5  6  D    < H@o?"0@NNN?NG1 9PROS: 7l  @  < @,$D 0  < H?"0@NNN?N @  Method makes heavy assumptions about the statistical distribution of experimental errors - could be overcome by the  shuffling and  shifting methods in DynaFit Can take a very long time to compute (multiple hours) Does not help in discovering false minima         L    $  !    < H?"0@NNN?N Le  9CONS: 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 %0 ,(  ,X , 0A?`m , H?"0@NNN?N  m-Side comment: The issue of significant digits$. !H , 0޽h ? ̙33y___PPT10Y+D=' = @B +  0L0   %= (     BX"`   XB  0D`X  0A?`m  <  c-RESULTS FROM A SEMESTER-LONG RESEARCH PROJECT.. 3  <A?"0@NNN?N F    bW=  H?"0@NNN?Nx  y = Bmax >  B  BD?"0@NNN?N@p p  H,?"0@NNN?NL @ 3x&  H?"0@NNN?N `  nKd + x2    B?"6@`NNN?N  ! <A?"0@NNN?NU @B " ZD?"0@NNN?N  # BV?"6@`NNN?N>; $ H?"0@NNN?N  ,$ 0 Wwhat is wrong with this result?  3 % H,?"0@NNN?N B@ p$ 0<4___PPT9 w)no measure of uncertainty too many digits**  @`H  0޽h ? ̙33___PPT10+(aD' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*$%(D' =-o6Bdissolve*<3<*$D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%%(D' =-o6Bdissolve*<3<*%+p+0+$0 ++0+%0 +/(  0L0 )!P 4(  4 4  BHҺ"`   XB 4 0D`X 4 0A?`m 4 <Ժ\  ZOUTPUT GENERATED BY SOFTWARE PACKAGE  ORIGIN ,. 3 3 4 <A?"0@NNN?N F   4 2 r = 4 Hں?"0@NNN?Nx  y = Bmax >  B  4 BD?"0@NNN?N@p p  4 H?"0@NNN?NL @ 3x&  4 H?"0@NNN?N `  nKd + x2   4 B?"6@`NNN?N z P   4 P ,$D  0 4 HA?"0@NNN?NP B 4B TD?"0@NNN?NB 4B TD?"0@NNN?N22" 4 HP?"0@NNN?N2 jBmax0  4 H ?"0@NNN?N  hKd0k 4 H<?"0@NNN?NP  Kd = (442.3346 67.39583) nMV    $_ 4 H_^?"0@NNN?NS  ,$ 0 {what is wrong with this result?@  3 3  3 4 HDe^?"0@NNN?N 9 NDIRECT OUTPUT FROM SOFTWARE: l     4  ,$D 0 4 Hi^?"0@NNN?N    LSENSIBLE WAY TO REPORT IT: h 4 Hn^?"0@NNN?N &  Kd = (440 70) nMV     $A 4 Nu^?"6@`NNN?N |$D  0H@___PPT9" uRECIPE: Round standard error to a single significant digit Round best-fit value to the same number of decimal pointsZ m       @`H 4 0޽h ? ̙33___PPT10n+krD' ^= @B D' = @BA?%,( < +O%,( < +Dn' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* 4%(D' =+4 8?dCB1+#ppt_w/2BCB#ppt_xB*Y3>B ppt_x<* 4D' =+4 8?\CB#ppt_yBCB#ppt_yB*Y3>B ppt_y<* 4D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*4%(D' =-o6Bdissolve*<3<*4D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*4%(D' =-o6Bdissolve*<3<*4D{' =%(D#' =%(D' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*4%(D' =+4 8?\CB#ppt_xBCB#ppt_xB*Y3>B ppt_x<*4D' =+4 8?dCB1+#ppt_h/2BCB#ppt_yB*Y3>B ppt_y<*4+p+0+40 ++0+40 +)   0L0 8 0 D(  D D  <^"`  ^ XB D 0D`X D 0A?`m D H܏^ ?pbD<4___PPT9 ZVAlways report at least some measure of statistical uncertainty for all nonlinear model parameters (rate and equilibrium constants). At the very least report the formal standard errors. Confidence intervals are more informative than standard errors. DynaFit offers two different methods for confidence intervals: a. Systematic search (profile-t method) b. Stochastic simulation (Monte-Carlo method) The two methods have their own merits and drawbacks When in doubt, use both. DynaFit is not a  silver bullet : You must still use your brain a lot. *?D  L7 4 33  @` D <^P PANY NUMERICAL RESULT REPORTED WITHOUT SOME MEASURE OF UNCERTAINTY IS MEANINGLESS,QE 3  3H D 0޽h ? ̙33y___PPT10Y+D=' = @B + 0 @<( @Y@ X  C 4   e  S Sg49 f  e >@  H  0ηo~ ? ̙33, 0 P |(   d  c $4   ^  3 r^ss49 f  ^  H  0ηo~ ? ̙33, 0  |(   d   c $4   ]   3 r^ss49 f  ]  H   0ηo~ ? ̙33, 0 @h|(  hd h c $4   ^ h 3 r+ ss49 f  ^  H h 0ηo~ ? ̙33, 0 `t|(  td t c $4   .  t 3 r. ss49 f  .   H t 0ηo~ ? ̙33, 0 |(  d  c $4   .   3 rX . ss49 f  .   H  0ηo~ ? ̙33, 0 |(  d  c $4   .   3 r(^ss49 f  .   H  0ηo~ ? ̙33, 0 |(  d  c $4   .   3 r`. ss49 f  .   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