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S  0*< `` < X*  0< `  < l*(    0hE< `  < `* H  0޽h ? ̙33 Default Design 0 P4 (  4 4 N y y .   p* a00aa 4 N y y 3 .  r* a00aad 4 c $ ?  4 4 Nؗ 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 T  y y r3    r* a00aaH 4 0ηo~ ? ̙3380___PPT10.| >X \x(  \ \ N%^]e]e   ^ p* AAA \ Nl)^]e]e %  ^ r* AAA  \ Tt3^]e]e   ^ p* AAA \ T0>^]e]e %  ^ r* AAAH \ 0ηo~ ? ̙3380___PPT10.|  0 @ m(   X  0A?`m;  6`M p kBinding and Kinetics for Experimental Biologists Lecture 6 Enzyme kinetics: Analysis of reaction progressNl0  /  <0/  (Petr Kuzmi , Ph.D. BioKin, Ltd. WATERTOWN, MASSACHUSETTS, U.S.A.hB    7p  H A ? ` ~  H A] ?"` )H  0޽h ? ̙33y___PPT10Y+D=' a= @B +   0L0 4P 8(  P r P S hN`  N XB P 0D`X P 0A?`m 4P  `@^?"0@NNN?NT$  Goal: Demonstrate KM determination from progress curves using numerical methods. Use a  real-world problem arising from the instructor s consulting practice. Topics: The Michaelis constant: theory and practice. The illusion of linearity in enzyme assays. Global vs. local fit of multiple data sets: Mechanistic implications. The choice of proper time interval to measure the  initial rates. Assessing the randomness of residual plots. Enzyme deactivation mechanisms. Implementation: The DynaFit software package.3       F   3      d       3    @`H P 0޽h ? ̙33y___PPT10Y+D=' a= @B +  0L0  K H(   x  c $t@`   X  0A?`m J  `HP?"0@NNN?NP ' = 6or * K  `S?"0@NNN?Ns 0  b.A Day in the Life of an Enzymology Consultant // H  0޽h ? ̙33y___PPT10Y+D=' a= @B +`  0L0    < (  ~  s *U`   XB  0D`X  0A?`m 4 < 7FDA = UNITED STATES FOOD AND DRUG ADMINISTRATION AGENCY^8 3 3 3 33 5 TA1?"6@`NNN?N0 7 TA2?"6@`NNN?N l p q  ;p@A@ ,$D  0 8 TA3?"6@`NNN?Np q B 9 TDjJ?"0@NNN?N d ~d B : TDjJ?"0@NNN?N@  ( <  `?"6@`NNN?N ,$D 0 & The drug candidate is an enzyme. This is a small biotech company  no kineticist on staff. The solution: Hire a consultant to - design a biochemical assay; and - propose a method of data analysis.n  &    3P H  0޽h ? ̙33c[___PPT10;+5-D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*;%(D' =-m6Bbox(in)*<3<*;D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*<%(D' =-o6Bdissolve*<3<*<+8+0+<0 +  0L0    (  ~  s */`   XB  0D`X  0A?`m 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[ \]^_ `abP p *    `1?"0@NNN?Np  a vary [Substrate] from 3.5 to 56 M add [Enzyme] at 5 nM observe UV/Vis absorbance at 405 nMLb    / ,9$|z       ,$D  0  Z33?"6@`NNN?N p`` "    `<?"0@NNN?N  RKM ?2$ $$H  0޽h ? ̙333+___PPT10 +6D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+7  0L0 ['S'&&&(  x  c $L|`   XB  0D`X  0A?`m  <\G= * KM HAS TWO DIFFERENT MEANINGS, EVEN FOR THE SIMPLEST POSSIBLE ENZYME MECHANISMnQ 3 3 3 31 33   `4?"0@NNN?N4n, =E + SB  TD?"0@NNN?NPpB @ TD?"0@NNN?NPp    ` ?"0@NNN?N4 :ESB   TD?"0@NNN?N @    `L?"0@NNN?N4p0  =E + P     `l ?"0@NNN?NPHd Dk1&      `|?"0@NNN?NPH Dk2&     `(?"0@NNN?Nx Dk3& nz V 4)  p C,$D  0-   ` ?"0@NNN?NV C ]KM = <33B  ND?"0@NNN?N 88   `&?"0@NNN?Ni8`  Dk1& f   `HM?"0@NNN?N 4) k2 + k3 r   g   `LU?"0@NNN?N`   kcat = k3X 33    `?"0@NNN?Nlf  S!equilibrium dissociation constant"" ~  T?"0@NNN?N<f`  Kd = k2 / k1    J   ` ?"0@NNN?Nh@.   L strength of enzyme/substrate binding '' Kz pP $    p$l ,$D  0   `?"0@NNN?Np $p  Ospecial case: when dissociation (k2) is much faster than chemical reaction (k3)xP  ) B  TD1?"0@NNN?NP P B   `?"0@NNN?Np0   rk2 >> k3N   z @0     @ l ,$D  0  Z33?"6@`NNN?N@` 0    ft!?"0@NNN?N0   k2 + k3 k2   >z  0     l ,$D  0   Z33?"6@`NNN?N `   !  f0?"0@NNN?N@ 0   DKM = (k2 + k3) / k1 k2 / k1 = Kd#   +z  0 "  0,$D 0 #  `0E?"6@`NNN?N( 0 0 KM kcatj  $  `+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*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 C(  ~  s *Z`   XB  0D`X  0A?`m  <t\P $IF THE DOSE-RESPONSE CURVE IS HYPERBOLIC, THEN  KM HAS ONLY ONE MEANING rL 31 3 3  3  332 '  `8c?"6@`NNN?Ni (  `e?"0@NNN?ND  v = kcat [E] V33B ) ND?"0@NNN?Np *  ``n?"0@NNN?N  5[S]( +  `r?"0@NNN?N: `[S] + KM< 33 ,  `w?"0@NNN?N PMichaelis-Menten equation:  -  `dq?"0@NNN?NL i  Vmax = kcat [E] l33$4l  a  =  a ,$D  0 92 ZA ? ? t   8 ;  `?"0@NNN?Na hVmax.B < ZD>?"0@NNN?NEEl @   @@  ,$D  0B > TDjJ?"0@NNN?N@ 0 0 @ ?  ``?"0@NNN?N   p Vmax,dl & 0 P  C0 & P ,$D 0B A ZDjJ?"0@NNN?N 0  B  `?"0@NNN?N& P  NKM033H  0޽h ? ̙33  ___PPT10 + <D ' a= @B D~ ' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*=%(D' =-m6Bbox(in)*<3<*=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<*C%(D' =-o6Bdissolve*<3<*C+  0L0 rj`  (  x  c $,`   XB  0D`X  0A?`m  Z?"0@NNN?N2   `t?"0@NNN?N0D<4___PPT9 Perform a number of assays at constant [E] and varied [S] Fit each progress curve (or  initial portion ) to a straight line Consider the slope of each best-fit line as the  initial rate (v) Start up a computer program for nonlinear regression In the list of built-in nonlinear models, find the  rectangular hyperbola y = a x / (b + x) where x = [S] y = v a = Vmax b = KM Click the [OK] button. Paste KM and Vmax into your final Report. If kcat is also required, compute it as Vmax / [E] n 3 .  333333   ! Pv5! @`8  <H` tTHE SIMPLEST POSSIBLE APPROACH ( BIOCHEM-101 STUDENT LAB )<; 3 3 3  TA?"0@NNN?N0@ ,$D  0^l      ,$D  0  Z?"6@`NNN?N     `?"0@NNN?N 0  Bmodel parameters H  0޽h ? ̙33___PPT10+k~7D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+%  0L0 TL  (  ~  s *H``   XB  0D`X  0A?`m2  <Ts <WE EVOLVED TO SEE STRAIGHT LINES - EVEN WHEN THERE ARE NONE!n= 3 3 3 3 33  Z?"0@NNN?N 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[ \]^_ `ab0   l Z`   Z` ,$D  0    fjJ?"6@`NNN?NP`     `l?"6@`NNN?N Z.  zdoes this look like a  linear portion of the progress curve?0>   B   ND?"0@NNN?N      `?"0@NNN?NZ Vk,$  0 If this were a  linear portion , our problems would be over: We could get the initial rate by fitting a straight line to the first 100 seconds./ 3  H  0޽h ? ̙33aY___PPT109+reD' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-m6Bbox(in)*<3<* D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-m6Bbox(in)*<3<* +8+0+ 0 +&%  0L0 $\ 8(  \~ \ s *``   XB \ 0D`X \ 0A?`m \ Z?"0@NNN?N \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[ \]^_ `abP     \ TA?"0@NNN?N = l sW \,$D 0 \  `jJ?"6@`NNN?N @p \  `Ԕ?"6@`NNN?NRWB \ TDjJ?"0@NNN?Np \  `X?"0@NNN?Ns  $0rate D[Signal] / Dtime 3 3 3 3 3 3 3 3 3,l F  \F ,$D 0@ @F \@F \  `8c?"6@`NNN?N@`P \  `8c?"6@`NNN?NP \  `?"0@NNN?N6tF E relative rate 2 \ Z3f?"6@`NNN?N `p M \  `?"0@NNN?N ]  } Vr = 1.00> ]l    "\-  ,$D 0 \  f8c?"6@`NNN?N@ `  \  f8c?"6@`NNN?N  2  \  `?"6@`NNN?N  ` S !\  f?"0@NNN?NV  } Vr = 0.62>  #\  `+?"0@NNN?N @*& ZBy the time the assay reached 100 seconds, the reaction rate decreased by almost one half!J[ #3 $\ <1  o)CHECKING LINEARITY BY SIMPLE COMPUTATIONS**) 33H \ 0޽h ? ̙33___PPT10+*@D' a= @B Du' = @BA?%,( < +O%,( < +D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D4' =%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*"\%(+   0L0  'd(  d~ d s *`;`    d0  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 Q   XB d 0D`X d 0A?`m d Z?"0@NNN?N d  `=?"0@NNN?N & ZBy the time the assay reached 100 seconds, the reaction rate decreased by almost one half!J[ #3 d0  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[ \]^_ `abp Q    d  `&?"0@NNN?N4 <signal d  `A?"0@NNN?N-  4rate D [Signal] / D timep   d # ljJ?"6@`NNN?NP0`  #d  `o?"6@`NNN?N4 jl  P &dX ,$D 0B $d TD8c?"0@NNN?N * %d  `L?"6@`NNN?NP T constant rate   'd <Ę IF THERE WERE A  LINEAR PORTION , THE RATE PLOT SHOULD BE STRICTLY HORIZONTAL`N& 3 3 3 33H d 0޽h ? ̙335-___PPT10 +pD' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*&d%(D' =-o6Bdissolve*<3<*&d+  0L0 0 (  ~  s *m`    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[ \]^_ `ab   XB  0D`X  0A?`m  Z?"0@NNN?N  <pp vHOW BAD COULD IT BE, REALLY?> 3 33 l p $ p $,$D  0 2  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[ \]^_ `abp       `8c?"6@`NNN?Nn   ` w?"0@NNN?N`  $ |quite bad, actually: this plot should look completely  random $?  6 H  0޽h ? ̙333+___PPT10 + ND' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+k  0L0  { p  (  ~  s *LT`   XB  0D`X  0A?`m  Z?"0@NNN?N  <V <CAREFUL HERE: POLYNOMIALS ARE UNSUITABLE FOR EXTRAPOLATIONS!>= 3 33    f$]?"0@NNN?NR r  |quite bad, actually: this plot should look completely  random $?  6   0 6A  ?     0 6A !?p   !    `8c?"6@`NNN?N,t   `>?"6@`NNN?Nll {   `$d?"6@`NNN?N` M,$  0 y!Why would we need to extrapolate?<"    H  0޽h ? ̙33xp___PPT10P+:^D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+8+0+0 +!  0L0 ]U`9(  ~  s *`   XB  0D`X  0A?`m  Z?"0@NNN?Nl  ` `@|,$D  0X    `?"0@NNN?N ` @enzyme is UV/Vis transparent substrate concentration is the sameAA  @`2  Z33?"6@`NNN?Np2   `?"0@NNN?Nz b.absorbance at time zero should be identical...// l @$  $\O@ ,$D  02 " Z33?"6@`NNN?N@0 z #  `H?"0@NNN?N $  r... unless a lot of product was formed already at  t = 0 :: 3> & < IN MOST KINETIC EXPERIMENTS WE NEVER  SEE THE VERY INITIAL TIME POINT*GF 33 -0 ZA #? ?~  # l    7  ,$D 0B .B TD>?"0@NNN?N b B /B TD>?"0@NNN?N b B 0B TD>?"0@NNN?N* b B 1B TD>?"0@NNN?N_ B 3B TD>?"0@NNN?NX b B 4B TD>?"0@NNN?N  e B 5 ZD?"0@NNN?N\  'l    9  ,$D 0 6  `?"6@`NNN?N v  ;-40  8  `?"6@`NNN?N   Vactual start of the assay:  time = -40 sec J,     H  0޽h ? ̙33  ___PPT10 +zڒD ' a= @B Di ' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*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<*7%(D' =-o6Bdissolve*<3<*7D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*9%(D' =-o6Bdissolve*<3<*9+  0L0   (j (  (x ( c $`   XB ( 0D`X ( 0A?`m ( Z?"0@NNN?N$ ( <\! NA FEW INNOCENT-SOUNDING CHALLENGES IN ANALYZING THE PROGRESS OF ENZYME ASSAYS:NO4 3 3 33 (  `l2(?"0@NNN?NP$ D<4___PPT9 The time-course of enzyme reaction is nonlinear. It can be considered  almost linear only in special cases. There is a finite mixing delay requiring extrapolation. In our experiments we always  miss the initial rate. Extrapolation is dangerous - unless we use a mechanistic model. Generic mathematical models such as polynomials are  out . Y& 3;  3 3(   / 33:  @` (  `?"6@`NNN?N ` ,$D  0 All these problems would disappear if/when the progress of enzyme assays can be treated as linear. This is almost never the case.dI       H ( 0޽h ? ̙33xp___PPT10P+ j D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*(%(D' =-m6Bbox(in)*<3<*(+8+0+(0 +   0L0  w(  x  c $X0   X  0A?`m  Z?"0@NNN?Nl f@   @f ,$D 0   TA?"0@NNN?N0@ I    `/(?"0@NNN?Nfy   y#... so why not do it the right way?:$ 3     `D(?"0@NNN?N@M Dlinear, polynomial H  0޽h ? ̙335-___PPT10 +D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* +  0L0 Nn(  x  c $+B#style.visibility<*M%(D' =-m6Bbox(in)*<3<*MD' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*N%(D' =-o6Bdissolve*<3<*N+8+0+N0 +W  0L0   D (     <+("`  ( XB  0D`X  0A?`m<  <'  hMORE THAN 600 PAPERS PUBLISHED WITH IT (1996  2009)L5  3  3  337z  0 0   0 0 ,$D  0  <' 0 0  http://www.biokin.com/dynafit&3  <' 2   <DOWNLOAD  H   3 A "` ~   H A ?"``?   T  jJ?"``0p   Hp' ?0 pv  oY1. Kuzmic (1996) Anal. Biochem. 237, 260-273 2. Kuzmic (2009) Meth. Enzymol. 467, 247-280Z >    T ' ?0   :2009   `'?"0@NNN?N4 p  > REFERENCES  H  0޽h ? ̙333+___PPT10 +4`D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+  0L0   (     <'"`  ' XB  0D`X  0A?`m  <'  f JUST AS GOOD AS MICHAELIS-MENTEN*!  33   `'?"0@NNN?Nn,a =E + SB  TD?"0@NNN?NPp   `'?"0@NNN?Na :ESB  TD?"0@NNN?N @   `'?"0@NNN?Np0 a =E + P    `'?"0@NNN?NPH Dk1&     `'?"0@NNN?N Dk3& +   f0 '?"0@NNN?Np =  ])Mathematical model for reaction progress:** 9   `'?"0@NNN?Npm qVan Slyke-Cullen mechanism:   TA?"0@NNN?N  yl & 0 0 : ,$D  0  Z'?"6@`NNN?N` 0 w [task] task = fit data = progress [mechanism] E + S ---> E.S : k1 E.S ---> E + P : k3 ... Lw 28:3,F   `2'?"0@NNN?N& O BDynaFit input:    Z5'?"0@NNN?N P "Application of the Van Slyke Cullen irreversible mechanism in the analysis of enzymatic progress curves Kuzmic (2009) Anal. Biochem. 394, 287-289nh       ,`H  0޽h ? ̙333+___PPT10 +H8D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+f  0L0 *"0 (     BE'"`sm  ' XB  0D`X  0A?`m  TA?"6@`NNN?N~   `G'?"0@NNN?Np 0  Meth. Enzymol. 210, 37-54 (1992)N!$  <P' ^A VERY IMPORTANT CONCEPT* 33B  ND3?"0@NNN?NP0PP  Z?"6@`NNN?N` 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 H $$  0 '   `4X'?"0@NNN?NDf v8 ,$  0 Lglobal analysis: analyze all five curves together as a single data setZM3    `_'?"0@NNN?N @ p ,$  0 Plocal analysis: analyze each curve individually as five separate data setsZQ3  H  0޽h ? ̙33  ___PPT10 +AD ' a= @B D ' = @BA?%,( < +O%,( < +D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*++(+ 0+'+0+0 ++0+0 +f   0L0 ` S(     Bq'"`sm  ' XB  0D`X  0A?`m  <s' }7WE NEED INITIAL RATES FOR THE MICHAELIS-MENTEN EQUATION*87 33  TA?"6@`NNN?N` l    ,$D  0  TA"?"0@NNN?N P  TA$?"0@NNN?N0  H  0޽h ? ̙333+___PPT10 +7Z{HD' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+_  0L0   HJ (  H H  BT'"`sm  ' XB H 0D`X H 0A?`m8 H <`'S  INITIAL RATE IS COMPUTED AT A SPECIFIC REACTION TIME (E.G. 1 SEC)*DC 33 H TA?"6@`NNN?N  H  `jJ?"6@`NNN?N <h F P&  H PV  H TA?"6@`NNN?NP&  H  `jJ?"6@`NNN?Nz p $  H  @dh ,$D 0  H TA?"6@`NNN?Np $v H  `jJ?"6@`NNN?N  B H ND?"0@NNN?NP PH H 0޽h ? ̙335-___PPT10 +D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* H%(D' =-o6Bdissolve*<3<* H+e  0L0    (     B؎'"`sm  ' XB  0D`X  0A?`m  <' ^ALWAYS EXTREMELY HELPFUL* 33  TA?"6@`NNN?N    `jJ?"6@`NNN?NP `  l p v p v,$D 0   ZA?"6@`NNN?Np v   fjJ?"6@`NNN?N     TA?"0@NNN?N`p   `'?"0@NNN?NpP  9E    `8'?"0@NNN?Np =  UE.S 33   `̠'?"0@NNN?N`  ,$  0 t steady state : when E.S stays constant (at least briefly)P;   33# #H  0޽h ? ̙33c[___PPT10;+mBD' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-o6Bdissolve*<3<*D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+8+0+0 +  0L0 (     B'"`sm  ' XB  0D`X  0A?`m  <0' DWHEN A BIOCHEMICAL ASSAY REACHES THE STEADY-STATE IS SYSTEM-SPECIFICLE% 3  3 33  TA?"0@NNN?NP0   TA?"6@`NNN?NP     `jJ?"6@`NNN?Np     `('?"0@NNN?N0  9E    `x'?"0@NNN?N   UE.S 33l p  Z   p Z ,$D 0   `<'?"6@`NNN?Np Z  <0.01 3   `t| ?"6@`NNN?N$ Z  91 3   ` ?"6@`NNN?N4  Z  @100 sec  3   `xd ?"6@`NNN?N Z  ;0.1 3   `Ħ ?"6@`NNN?N Z  :10 3pl p 0  p 0 ,$D  0   `jJ?"6@`NNN?N 0    `.?"0@NNN?Np  N50 ms  1 s  l  P P ,$D  0  TA?"6@`NNN?N Pr   `.?"6@`NNN?N  tell DynaFit at what time we want to read-off the  initial reaction rate$J2  H  0޽h ? ̙33  ___PPT10 +5D ' a= @B D~ ' = @BA?%,( < +O%,( < +D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-o6Bdissolve*<3<*D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+/!  0L0 $H(     B."`sm  . XB  0D`X  0A?`m  <̒.  yFINALLY READY TO COMPUTE THE KM>  3 33 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[ \]^_ `abP p & .|z       ,$D  0  Z33?"6@`NNN?N p`` "   `K?"0@NNN?N  RKM ?2$ $$ l u k  !u k ,$D  0   `K?"6@`NNN?N 0k  :1.0981    `8K?"6@`NNN?N k }  :0.9022    `(K?"6@`NNN?N k  :0.7456    `XK?"6@`NNN?Nu  `  :0.4530    `8K?"6@`NNN?N k  :0.2674     ` L ?"6@`NNN?N "  A initial rates Vl   $ ,$D 0B " ND?"0@NNN?N pn p #  `DW?"0@NNN?NP  Labsorbance units / sec H  0޽h ? ̙33  ___PPT10 +``D ' a= @B D~ ' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*!%(D' =-m6Bbox(in)*<3<*!D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*$%(D' =-o6Bdissolve*<3<*$D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*+9  0L0 H@  (       B< "`sm   XB   0D`X   0A?`m   TA?"6@`NNN?NJ T   TA?"6@`NNN?N p$    TA?"0@NNN?N `    Z33?"6@`NNN?N `     `T ?"0@NNN?NP n N File .. Try      `N?"0@NNN?Nvq Finitial estimate H   0޽h ? ̙33y___PPT10Y+D=' a= @B +r  0L0 ((  ( (  BN"`sm  N XB ( 0D`X ( 0A?`m  ( TA?"0@NNN?N0   ( TA ?"0@NNN?N6 @6 (  `\N?"0@NNN?N0  n"KM = (13.1 1.6) [6.5 .. 26.8] M0# !l n  (n ,$D 0@ H  (H  (  `8c?"6@`NNN?N  (  `jJ?"6@`NNN?NH (  `N?"0@NNN?N@n0 Dstandard error Rl ^ B  (B^  ,$D 0 (  `jJ?"6@`NNN?NB (  `8c?"6@`NNN?N^  * (  `tN?"0@NNN?N @ Z$confidence interval (99% probability%% l    (  ,$D  0 ( TA!?"6@`NNN?N0^ B ( ND?"0@NNN?N  , (  `N?"0@NNN?N6 +  \log KM:3 B ( ND?"0@NNN?N   (  `ԿN?"0@NNN?N c  Bsum of squares 1 ( <N IREMEMBER: THE STANDARD ERROR ( VALUE FREQUENTLY REPORTED) IS MEANINGLESS`J 3 3 3  33H ( 0޽h ? ̙33  ___PPT10 +W,D ' a= @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' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*(%(D' =-m6Bbox(in)*<3<*(+  0L0   0&(  0 0  BN"`sm    XB 0 0D`X 0 0A?`m 00  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[ \]^_ `abP p ( NF    0    0 Z33?"6@`NNN?N p`` d  0  `N?"0@NNN?N   KM = 13.1 Ml $ $3333B 0  `N?"0@NNN?N f ,$ 0 Fare we done yet?  0 <N pALMOST DONE .... MAYBE> 3 33H 0 0޽h ? ̙33zr___PPT10R+laD' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*0%(D' =-o6Bdissolve*<3<*0+8+0+00 +7  0L0 F>@ 8(  8x 8 c $N0  N X 8 0A?`m 8 Z?"0@NNN?N. 8  flN?"0@NNN?Ny fz  `*what else we can learn from the same data?++  8 TA#?"6@`NNN?NpV H 8 0޽h ? ̙33y___PPT10Y+D=' a= @B +.  0L0 + # `@ (  @ @  BN"`sm  N XB @ 0D`X @ 0A?`m  @ TA%?"6@`NNN?NB   @ TA'?"6@`NNN?N ,  @  `>?"6@`NNN?N @p @  `>?"6@`NNN?N % @ <N" =VAN-SLYKE CULLEN MECHANISM (SAME AS BEFORE, IN THE LOCAL FIT)>> 3- 335l F 00  @0F 0 ,$D  0 @ TA&?"0@NNN?N 00  @ Z33?"6@`NNN?N P0  @  ``<?"0@NNN?NF z  ? File .. Run  \ @  `c<?"0@NNN?N@ @ ,$ 0 `.looks  reasonably good ? @  `b<?"0@NNN?NP Ld,$ 0 C or does it ?!3H @ 0޽h ? ̙33  ___PPT10k +6mD ' a= @B D ' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*@%(D' =-m6Bbox(in)*<3<*@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](  P P  Bp<"`sm  < XB P 0D`X P 0A?`m  P <r< aDOES NOT LOOK GOOD AT ALL, 33 P TA(?"0@NNN?N 0  P  `hx<?"6@`NNN?N (,$D  0 `If you see a residual plot as bad as this one, completely ignore any other result from DynaFit.Ba31` P  `<?"6@`NNN?NL  ,$D  0 ^Always look at residual plots.$H P 0޽h ? ̙33___PPT10~+_D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-m6Bbox(in)*<3<*PD' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-m6Bbox(in)*<3<*P+p+0+P0 ++0+P0 +;   0L0 J B  X (  X X  B\<"`sm  < XB X 0D`X X 0A?`m  X TA)?"6@`NNN?NK  X  `<?"0@NNN?Ng6; =GOOD:  X TA*?"6@`NNN?N   X  `H<?"0@NNN?N <BAD:3k  X  `<?"0@NNN?N@K 9Draper & Smith (1981) Applied Regression Analysis, p. 146N:$ X <<7 NASSESSING RESIDUAL PLOTS IS ONE OF THE MOST IMPORTANT SKILLS OF A DATA ANALYSTNO' 3 3 33n X  `إ<?"0@NNN?N` Nrandom sequence no  trend or  pattern <(   X  `X<?"0@NNN?N 1g dnon-random sequence shows a  trend or a  pattern <3 3 3 3H X 0޽h ? ̙33y___PPT10Y+D=' a= @B +-  0L0 me`(  ` `  B<"`sm  < XB ` 0D`X ` 0A?`m ` <<  i!DOES NOT LOOK GOOD AT ALL, EITHER,"! 33  ` TA+?"0@NNN?NZ   `  `>?"6@`NNN?N `   `  `<?"0@NNN?N  KM = kdp/kas = 18.2 M333  3    ` TA,?"0@NNN?N L  `  `<?"0@NNN?N ~h,$ 0 P13.1 M from initial ratesl P  `P ,$D 0 `  `8c?"6@`NNN?NP `  `8c?"6@`NNN?N>P `  `8c?"6@`NNN?N|P  `  `8c?"6@`NNN?N Pb  `  `8c?"6@`NNN?N P jl   ` ,$D 00 `  ``<?"0@NNN?N  `These  best fit concentrations are highly improbable: It has been established that the actual titration error is much smaller than this.0`   B ` ND?"0@NNN?N` ` H ` 0޽h ? ̙33P H ___PPT10( +HwD ' a= @B D ' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(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<*`+8+0+ `0 +7   0L0 F >  h (  h h  B<"`sm  < XB h 0D`X h 0A?`m h <<X  ZENZYMES DO  DIE IN THE COURSE OF MANY ASSAYS,.- 33 h Z<?"0@NNN?N`  [mechanism] E + S ---> E.S : kas E.S ---> E + P : kdp E ---> E* : kie E.S ---> E.S* : kies.L:3 h  `jJ?"6@`NNN?N  h  `<?"0@NNN?Nm A denatured  3H h Z<?"0@NNN?N`   Mechanical Effects on the Kinetics of the HIV Proteinase Deactivation Kuzmic et al. (1996) Biochem. Biophys. Res. Commun. 221, 313-317F         P. # h  ``?"0@NNN?N  nSadana (1991) Biocatalysis - Fundamentals of Enzyme Deactivation Kinetics Prentice-Hall, Englewood Cliffs, NJ.<o < % $ U h TA-?"0@NNN?N @  h Z33?"6@`NNN?NPP H h 0޽h ? ̙33y___PPT10Y+D=' a= @B +  0L0 @x(  x x  Bk"`sm   XB x 0D`X x 0A?`m x <m ~6IT LOOKS GOOD EVEN UP CLOSE (RESIDUALS AND PARAMETERS),76 33  x TA.?"0@NNN?N`   x  `r?"0@NNN?N =  A residuals   x TA/?"0@NNN?N`   x  `>?"6@`NNN?N,0Lf x  `Dw?"0@NNN?N`0   KM = 13.6 Mv 3333 W x  `?"0@NNN?NPp,$ 0 [13.1 M initial rates*  x  `jJ?"6@`NNN?N> x  `jJ?"6@`NNN?N| x  `jJ?"6@`NNN?N0   x  `jJ?"6@`NNN?N   x  `jJ?"6@`NNN?N Z B x ND?"0@NNN?N`   x  `\?"0@NNN?N v vq NThese concentration look OK. H x 0޽h ? ̙33zr___PPT10R+9D' a= @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D@' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*x%(D' =-o6Bdissolve*<3<*x+8+0+x0 +  0L0 JB`  (     B"`sm   XB  0D`X  0A?`m  TA.?"0@NNN?N`   TA0?"0@NNN?N`@  "   `<?"0@NNN?Nd  $  Zwith enzyme deactivation&%   `?"0@NNN?N` ` *  ]without enzyme deactivation&B   `DԔ?"0@NNN?NPpp` ,$D  0l pP `  Pp ` ,$D  0B   `DԔ?"0@NNN?N  B  ND?"0@NNN?NpP B  ND?"0@NNN?Np ` $  <7 NASSESSING RESIDUAL PLOTS IS ONE OF THE MOST IMPORTANT SKILLS OF A DATA ANALYSTNO' 3 3 33kl c q  c q,$D 0M   f?"0@NNN?Nd  wno recognizable pattern: random<  3 3 3   f?"0@NNN?N` c q Ha clear  zig-zag pattern non-random>% 3  3 3 H  0޽h ? ̙33  ___PPT10 +E_lD ' a= @B D~ ' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-o6Bdissolve*<3<* +8  0L0 G ?  (     Bغ"`sm   XB  0D`X  0A?`m  TA4?"0@NNN?NPP B  TD?"0@NNN?N B  TD?"0@NNN?N B  TD?"0@NNN?N B  TD?"0@NNN?N@ @    `<?"0@NNN?N V =  5E    `?"0@NNN?N  QE.S    `p?"0@NNN?N`  :E* 3   ` ?"0@NNN?N  @  VE.S* 3"   `?"0@NNN?ND Zexample assay: [S] = 14 M$  K   `?"0@NNN?N`  Cdenatured enzyme is the main molecular form at the end of the assay$D 4 B  TD?"0@NNN?N H  0޽h ? ̙33y___PPT10Y+D=' a= @B +   0L0    (     B"`sm   XB  0D`X  0A?`m   `|?"0@NNN?NYD<4___PPT9 The progress curves must be treated as nonlinear There is no  linear portion, even if it seems that there is one. Michaelis constant, KM, was determined by two independent methods A. local fit of progress curves fit of initial rates (KM = 13.1 M) B. global fit of progress curves (KM = 13.6 M) Both methods gave the same results. Global fit revealed that the enzyme undergoes deactivation. Deactivation (denaturation) is almost complete after 15 minutes. Deactivation does not interfere with KM determination. If we use the initial rate method; or if we take deactivation into account in the global fit. Overall: This substrate is very suitable for a KM assay. If we use an appropriate (nonlinear) method of data analysis. ' D   "    0   4 .  @  %  < &33333A  @`H  0޽h ? ̙33y___PPT10Y+D=' a= @B +!0  0L0 (<(     B"`sm   XB  0D`X  0A?`m  <a(  .THE FDA WANTS US TO DO TWO THINGS: KM AND kcat/ 3 3 3 3 3 3 3 33*z p q   `0,$D  0  TA3?"6@`NNN?Np q B  TDjJ?"0@NNN?N d ~d B  TDjJ?"0@NNN?N@  }l    ,$D  0l   `(?"0@NNN?Nh   v = Vmax T B  ND?"0@NNN?N     `(?"0@NNN?N&f 5[S]0   `Dp(?"0@NNN?N 9  `[S] + KM< 33   `0(?"0@NNN?N'l @  Vmax = kcat [E] l33$   `u(?"0@NNN?N NMichaelis-Menten equation:   Z?"6@`NNN?N l   ,$D  0   ` r(?"6@`NNN?N !YEven with the differential-equation model, we can (essentially) measure only KM and Vmax.Z@        3 3 T   `T(?"0@NNN?N Q @ The problem:   "  fȴ(?"0@NNN?NP ,$  0 kcat = Vmax / [E]|33$(z   #   ,$D  0 $  `X(?"6@`NNN?N  cAt some point we must determine the concentration of active enzyme [E] (active site concentration).0d5    %  `t(?"0@NNN?N Q A The solution:  z    & 0  ,$D  0\ '  `(?"6@`NNN?N  D<4___PPT9 :PDetermine  apparent kcat , from [E] as total protein, for a reference sample of enzyme. Relate every other  apparent kcat value to the reference sample in the future.      #  +   / ,^/ @` (  `P(?"0@NNN?N#Q OThe only other possibility: H  0޽h ? ̙33___PPT10+l9.D' a= @B DD' = @BA?%,( < +O%,( < +D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-m6Bbox(in)*<3<*D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-m6Bbox(in)*<3<* D' =%(D' =%(D>' =A@BBBB0B%(D' =1:Bvisible*o3>+B#style.visibility<*"%(D' =-m6Bbox(in)*<3<*"D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*#%(D' =-m6Bbox(in)*<3<*#D' =%(D' =%(D1' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*&%(D' =-m6Bbox(in)*<3<*&+8+0+"0 +  0 b(  x  c $(`  ( XB  0D`X  0A?`m  T( ? D<4___PPT9 Even a  simple task such as KM determination can be quite tricky if it is to be done correctly and rigorously. DynaFit can help in dealing with the nonlinearity of reaction progress. Local fit of individual progress curves is not sufficiently powerful to reveal the true kinetic mechanism. Global fit of multiple curve is much preferred for mechanistic studies. Residual plots are crucially important for model validation. 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