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?܊r(7  DUexcdd``a 2 ĜL0##0KQ* Wä`.d3H1)fYifĒʂT/&`b]F"LL`0L !*'] ZZKy3@dY$ZR$fdD:Lv545.5X|E+-hL XYU,lm,^r:xvZJYm #zhI#27r&ml%Ǖ5}&.o&SR{(g3jQO[Whuΐ)s$78U}e\qk;-(%QǿMA}MaWSnu?Ze9\Ku%_fɹJ/E99bvUV!o`!O; &$8xڝ;KAgbaN#` B{!gRH*3bΈ tllD+ REbiea๳@icf۹a0И3BƔ}a&oF/=í` 6ܢK|b,(s>3LƞEbZ R?%y)2 Ci%7Nr镏aS .b2>ަHqΪuK) Z4r>dtNRΧ)SDe84& IV_(jH+%=GyMFg{x7G~Hҿ0nje.+{ڇ?vU (c\w/"<Wa*Hk2Eɨ2B{}Snx䙪(I6t ~b-(   & Equation Equation.30,Microsoft Equation 3.0 Equation Equation.30,Microsoft Equation 3.0Equation Equation.30,Microsoft Equation 3.0ZDrosophila Centrosome JORDAN RAFF, Cambridgedhttp://www.welc.cam.ac.uk/~rafflab/gfpfzrfrap.htmlhGFP-sialyl transferase in CEF cells Banting LAB, MRCVhttp://www.bch.bris.ac.uk/mrccif/Movie1.htmpFRAP demo Federica Brandizzi, University of Saskatchewandhttp://www.usask.ca/biology/brandizzi/movie7a.html BFilament FRAP Leube group, Mainzhttp://www.uni-mainz.de/FB/Medizin/Anatomie/Leube/publications/22/22_Movie06.avi"BVirus (SV40) FRAP Pelkmans et al.http://www.nature.com/ncb/journal/v3/n5/extref/ncb0501_473-s17.mov$ Equation Equation.30,Microsoft Equation 3.01-Equation Equation.30,Microsoft Equation 3.04/Equation Equation.30,Microsoft Equation 3.062Equation Equation.30,Microsoft Equation 3.0@4Equation Equation.30,Microsoft Equation 3.0A5Equation Equation.30,Microsoft Equation 3.0B6Equation Equation.30,Microsoft Equation 3.0C7Equation Equation.30,Microsoft Equation 3.0D8Equation Equation.30,Microsoft Equation 3.0E9Equation Equation.30,Microsoft Equation 3.0F:Equation Equation.30,Microsoft Equation 3.0G;Equation Equation.30,Microsoft Equation 3.0H<Equation Equation.30,Microsoft Equation 3.0,Reits & Neefjes (2001)http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=11389456DEquation Equation.30,Microsoft Equation 3.0EEquation Equation.30,Microsoft Equation 3.0FEquation Equation.30,Microsoft Equation 3.0GEquation Equation.30,Microsoft Equation 3.0,Phair & Misteli (2000)http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=11733769 *Axelrod et al. (1976)http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=786399*Ellenberg et al. 1997http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=92989764Presley et al. Nature 2002http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=12000962*Presley et al. (2002)http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=pubmed&dopt=Abstract&list_uids=12000962DAdriaan Houtsmuller Lab s web siterhttp://www.eur.nl/fgg/pathol/research/mcb/simulations.htmDhttp://cci.sahlgrenska.gu.se/ELMI/Dhttp://cci.sahlgrenska.gu.se/ELMI/\ EAMNET4http://www.embl.de/eamnet/J/ 0|DArialNew E(0(z[ 0 DMS PGothicE(0(z[ 0 " DSymbolhicE(0(z[ 0 0DWingdingsE(0(z[ 0 @DTimes New Roman(0(z[ 0 F .  @n?" dd@  @@`` x.  Tv6  XP M3 #$%&F'(*- 0 2#(R$gfPVY(wR$ άyF:wR$F´wЮU IOwTR$'%lW0Bc2$E=oco+/2$+ZPhIS1R$O@۷2$kAfhhG@2$X^,{g_9B2$?)4jh.3Q}6D2$Lj!)_!F2$+ʏGG2$?܊r(I2$w+Wn2t K2$O; M 0AA@=ʚ;ʚ;g4KdKd@z[ 0$Rppp@ <4ddddl 0E <4BdBdl< 00___PPT10 pp___PPT9 "?  %<=!@$              *  +  ,  H&    0  1    >"  d  ` 33` Sf3f` 33g` f` www3PP` ZXdbmo` \ғ3y`Ӣ` 3f3ff` 3f3FKf` hk]wwwfܹ` ff>>\`Y{ff` R>&- {p_/̴>?" dd@,|?" dd@   " @ ` n?" dd@   @@``PR    @ ` ` p>> &(    6yE " `} E X Click to edit Master title style!! (  0t|E " ` E RClick to edit Master text styles Second level Third level Fourth level Fifth level!    S   0 E "^ ` E d*b    0DE "^  E f*b    0ЋE "^ ` E f*b  H  0޽h ? 3380___PPT10.R!Ф Default Design   DE(  D8   D*  D 6X.E"`  What do FRAP curves tell us?4(2ff  Q D 63E   [Kota Miura EMBL Heidelberg, Germany Based on the EAMNET Practical Course (20, April, 2004 )X\(2 d?fff \ h D 0:E ,4,$ 0 *To proceed, press Page Down or click the mouse button. To go back, Press page Up. Clicking hyperlink (blue fonts) leads you to a website in the Internet.  2FB D s *޽h ? 33___PPT10z.чd%+n(D' = @B D' = @BA?%,( < +O%,( < +D' =%(%(D' =%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*D%(D' =-g6B fade*<3<*D+8+0+DE + A(  7  0 C=S ,$ 0 =What is FRAP? Quantitative Analysis of FRAP Modeling and FRAP > 2>f > 8X  0,K Y,$ 0 _In this presentation, I will give a short overview on what we can know about molecular dynamics by studying the Fluorescence Recovery After Photobleaching (FRAP). The presentation is intended for beginners of FRAP. The slides consists of the following topics. > 2FFfF  0F j Q,$ 0 ;Notes 2DB  s *޽h ? 33  ___PPT10 .f, U +>ΆD' = @B D' = @BA?%,( < +O%,( < +D' =%(%(D@' =%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-g6B fade*<3<*D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-g6B fade*<3<*Dh' =A@BB BB0B%()))D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-g6B fade*<3<*++0+  ++0+  ++0+  +B4 Z"R"0KWH!(  H`2 H s *"`sVn`2 H s *"`n`2 H s *"`^A`2 H s *"`^/  A`2 H s *"` `2  H s *"``2  H s *"`( `2  H s *"`i`2  H s *"` `2  H s *"`a9D`2 H s *"`4`2 H s *"`4iL `2 H s *"`iL n`2 H s *"`)  `2 H s *"`w `2 H s *"`Y< `2 H s *"`& `2 H s *"`6  `2 H s *"`6 wZ `2 H s *"`6  `2 H s *"`6 <  `2 H s *"`t W `2 H s *"`  `2 H s *"`t jW `2 H s *"`t   W `2 H s *"`  `2 H s *"`aD`2 H s *"` (  `2  H s *"`Q 4`2 !H s *"`7  `2 "H s *"` n`2 #H s *"`2 n`2 $H s *"`^A`2 %H s *"`^fA`2 &H s *"`E(`2 'H s *"``2 (H s *"`| _ `2 )H s *"``2 *H s *"`4f`2 +H s *"`a pD`2 ,H s *"`4%`2 -H s *"`4`2 .H s *"`n`2 /H s *"`} ` `2 0H s *"`  `2 1H s *"` `2 2H s *"`& 9 `2 3H s *"`6  `2 4H s *"`6  `2 5H s *"`6 2  `2 6H s *"`6 s `2 7H s *"`t 9W `2 8H s *"` C& `2 9H s *"`t W `2 :H s *"`t sVW `2 ;H s *"`sV `2 H s *"``2 ?H s *"`7  `2 @H s *"`@ # 2 AH <D["`P ,$@ 0l  h PHP `  ,$@ 0h2 JH s *"`  h2 KH s *"` } hh2 LH s *"`m P h2 MH s *"`2 * h2 NH s *"` vh2 OH s *"` v Y SH 0v I,$ 0 t<We focus a strong laser beam to a spot (red dotted circle). = 2=F TH 0{ "`q H1. What is FRAP?(2D UH 0 z }1Fluorescence Recovery After Photobleaching (FRAP)2 22d 2 \ VH 0 Fw Let s think of fluorescence molecules dispersed in a field. White circles represent the molecules.c 2cF WH 0 I,$ 0 {CThen the strong irradiation BLEACHES the fluorescence at that spot.D 2DFB H s *޽h ? 33___PPT10h.&#@j+s]uD\' = @B D' = @BA?%,( < +O%,( < +D' =%(DZ' =%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*VHD' =1:Bhidden*o3>+B#style.visibility<*VH%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*AH%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*SH%(D' =-g6B fade*<3<*SHD' =%(DZ' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*PH%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*SHD' =1:Bhidden*o3>+B#style.visibility<*SH%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*WH%(D' =-g6B fade*<3<*WHD' =%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*AH%(++0+SH  ++0+SH  ++0+VH  ++0+WH  +V@   PHOL(  L`2 L s *"`sVn`2 L s *"`n`2 L s *"`^A`2 L s *"`^/  A`2 L s *"` `2 L s *"``2 L s *"`( `2  L s *"`i`2  L s *"`a9D`2  L s *"`4`2  L s *"`4iL `2 L s *"`iL n`2 L s *"`)  `2 L s *"`w `2 L s *"`Y< `2 L s *"`& `2 L s *"`6  `2 L s *"`6 wZ `2 L s *"`6  `2 L s *"`6 <  `2 L s *"`t W `2 L s *"`  `2 L s *"`t jW `2 L s *"`t   W `2 L s *"`  `2 L s *"`aD`2 L s *"` (  `2 L s *"`7  `2  L s *"` n`2 !L s *"`2 n`2 "L s *"`^A`2 #L s *"`^fA`2 $L s *"`E(`2 %L s *"``2 &L s *"`| _ `2 'L s *"``2 (L s *"`4f`2 )L s *"`a pD`2 *L s *"`4%`2 +L s *"`4`2 ,L s *"`n`2 .L s *"`  `2 /L s *"` `2 0L s *"`& 9 `2 1L s *"`6  `2 2L s *"`6  `2 3L s *"`6 2  `2 4L s *"`6 s `2 5L s *"`t 9W `2 6L s *"` C& `2 7L s *"`t W `2 8L s *"`t sVW `2 9L s *"`sV `2 ;L s *"` | _ `2 *B ppt_xB ppt_y=@0BBAApBBBn<*ALD' =4@BB1BB%()?)?Dq' =.97 BBBBBKM 0.00017 0.00232 L 0.03594 0.09121 *3>*B ppt_xB ppt_y=@0BBAApBB(}<Ba 6=<* LD!' =4@BB8BB%()?)?Du' =.=7 BBBBBOM 0.00434 -0.00301 L 0.03993 -0.06065 *3>*B ppt_xB ppt_y=@0BBAApBB<B{ <*LD+' =4@BB8BB%()?)?D' =.G7 BBBBBYM 3.05556E-6 -1.48148E-6 L 0.0434 -0.11018 *3>*B ppt_xB ppt_y=@0BBAApBBDZ<BŨa<*CLD!' =4@BB1BB%()?)?Du' =.=7 BBBBBOM -0.00364 -0.00532 L 0.03212 0.07338 *3>*B ppt_xB ppt_y=@0BBAApBB(}<Bh/!=<* LD' =4@BB1BB%(L)?)?Ds' =.;7 BBBBBMM 0.0007 -0.01065 L -0.05798 0.01551 *3>*B ppt_xB ppt_y=@0BBAApBB[BbT<<*&LD-' =4@BB1BB%()?)?D' =.I7 BBBBB[M -2.77778E-6 -7.40741E-7 L 0.02778 0.07894 *3>*B ppt_xB ppt_y=@0BBAApBB9c<Bh/!=<*DLD#' =4@BB1BB%()?)?Dw' =.?7 BBBBBQM -0.00364 -0.00532 L 0.03993 -0.07361 *3>*B ppt_xB ppt_y=@0BBAApBBDZ<BpS <*3LD!' =4@BB8BB%( )?)?Du' =.=7 BBBBBOM 0.00833 0.01041 L -0.05052 -0.00533 *3>*B ppt_xB ppt_y=@0BBAApBBB<*.LD ' =%(Db ' =%(D)' =4@BB BB%(D' =-g6B fade*<3<*JLD' =1:Bhidden*o3>+B#style.visibility<*JL%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*NLD' =1:Bhidden*o3>+B#style.visibility<*NL%(D[' =4@BB BB%()))D' =1:Bvisible*o3>+B#style.visibility<*KL%(D' =-g6B fade*<3<*KLD8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*OL%(D' =-g6B fade*<3<*OL+p+0+NL  ++0+OL  +   (  `  s *"`zF 2  < "`  ,$@ 02  < "`  ,$D 0  0   }1Fluorescence Recovery After Photobleaching (FRAP)2 22d 2   0 "`q H1. What is FRAP?(2D'  0 FWw mIn practice, the FRAP process looks like the following. Above is a microscope field filled with fluorophores.n 2nF`   0h X ,,$ 0 TBleach a spot&  2F  07 I,$  0 VGradual Fluorescence Recovery! 2FB  s *޽h ? 33YQ___PPT101.#7/+Q$,D%' = @B D' = @BA?%,( < +O%,( < +D' =%(DZ' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*D' =1:Bhidden*o3>+B#style.visibility<*%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-g6B fade*<3<*D] ' =%(D' =%(D' =4@BBBB%(D' =1:Bhidden*o3>+B#style.visibility<*%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*D' =1:Bhidden*o3>+B#style.visibility<*%(D' =%(D' =4@BBBB%())))?D' =1:Bvisible*o3>+B#style.visibility<*%(D' =%(D)' =4@BB BB%(D' =-g6B fade*<3<*D' =1:Bhidden*o3>+B#style.visibility<*%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* ++0+7  ++0+7  ++0+7  ++0+ 7  +4  K C `P(  P P 0$ 7   VExamples 2 f  m P <+7  U 0 Drosophila Centrosome JORDAN RAFF, Cambridge GFP-sialyl transferase in CEF cells Banting LAB, MRC a type 1 integral membrane protein which is localised to the trans face of the Golgi stack in control cells FRAP demo Federica Brandizzi, University of Saskatchewan Filament FRAP Leube group, Mainz A section of a PK18-5 cell producing HK18-YFP; filamentous fluorescence recovery occurs preferentially in the cell periphery. dt=2 min. Virus (SV40) FRAP Pelkmans et al. NATURE CELL BIOLOGY, vol 3 pp473- N-df"d$f|               -   8                     U 0-U 0/cU 0  U 0 ,"U 0 P 0&7 "`q H1. What is FRAP?(2D P 0,7 , X Many FRAP movies can be found on the web. Above is a list of examples. You could also find many others by searching keywords  FRAP and  movie . 2FB P s *޽h ? 33___PPT10i.)#`n+D=' = @B +.  wo T (  T` T s *"`]F f2 T 0 "`  2 T < "`  ,$D 0 T 0'7 fM }1Fluorescence Recovery After Photobleaching (FRAP)2 22d 2 dB T <DԔ #  T 087 ,3  ;Measure the temporal changes of the fluorescence Intensity!< 2<d <   T 0<7 "`qM   2. Quantitative Analysis of FRAP>!(2DdD   T 0I7 vqP ~ eTo gain information on molecular dynamics, time-course of the fluorescence recovery must be measured.f 2fFB T s *޽h ? 33_W___PPT107.#7/+ksD ' = @B D' = @BA?%,( < +O%,( < +D' =%(D' =%(D' =4@BBBB%())))?D' =1:Bvisible*o3>+B#style.visibility<*T%(D' =%(D)' =4@BB BB%(D' =-g6B fade*<3<*TD' =1:Bhidden*o3>+B#style.visibility<*T%(+  7/  (  T  C ,Afrapcurve &  0DN7  I  Reits & Neefjes (2001) 2d@     U 0  0dP7 w' X Frap Curve 2 f      0T7 "`qM   2. Quantitative Analysis of FRAP>!(2DdD 7   0LZ7 , This is an example of FRAP curve, a result of measuring the fluorescence intensity at the bleached spot. X-axis is time, Y-axis is the fluorescence intensity.  2FB  s *޽h ? 33___PPT10i.R!Ф+D=' = @B +6 \(  \V \ C .Aheiko_origB \ c $D"W",$D 0Rz    \   ,$@ 0`B \ 0D8c  `B \ 0D8c   \ 08a7 P  gTerms Used in FRAP analysis 2d  B  \ s *Dg,$@  0 l Jf y \f J y,$D  0 \ 6G7 JY y Half Life (t1/2)h 2 degme  rB  \ BD f ,l   \ ,$@  0~B  \ NDp" ~B  \ NDp"  \ 0DL"`qM   2. Quantitative Analysis of FRAP>!(2DdD  \ 00Lr  VThe actual measurement of fluorescence intensity tends to result in a graph like this.W 2WF \ 0Lr +,$ 0 GWe can extract several information out of this curve by drawing lines. H 2HF \ 0$ L ) ,$ 0 First is the  half-life : the time it takes for fl. Intensity to reach half the maximum of the plateau level.n 2nF \ 0LCt,$ 0 fPlateau 8 2DDB \ 0LL z f ,$  0 vThe recovery is not complete, plateau being lower than the pre-bleach intensity (in this graph, intensity during t<0).w 2wF \ 0TL  I ,$ 0 m5What could be the reason of this incomplete recovery?6 26FB \ s *޽h ? 33%%___PPT10y%.R!Ф+[<+1DU#' = @B D#' = @BA?%,( < +O%,( < +D ' =%(D4 ' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\D6' =A@BB BB0B%(D' =-g6B fade*<3<*\D' =1:Bhidden*o3>+B#style.visibility<*\%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\D ' =%(D' =%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*\D' =1:Bhidden*o3>+B#style.visibility<*\%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\D' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\D ' =%(D4 ' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* \%(D' =-g6B fade*<3<* \D6' =A@BB BB0B%(D' =-g6B fade*<3<*\D' =1:Bhidden*o3>+B#style.visibility<*\%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\D&' =%(D' =%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*\D' =1:Bhidden*o3>+B#style.visibility<*\%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-g6B fade*<3<*\++0+\L ++0+\L ++0+\L ++0+\L ++0+\L ++0+\L ++0+\L ++0+\L ++0+\L +$% o g FLX (  X`2 X s *"`sVn`2 X s *"`n`2 X s *"`^A`2 X s *"`^/  A`2 X s *"` `2 X s *"``2 X s *"`( `2  X s *"`i`2  X s *"`a9D`2  X s *"`4`2  X s *"`0  `2  X s *"`iL n`2 X s *"`)  `2 X s *"`w `2 X s *"`Y< `2 X s *"`& `2 X s *"` s `2 X s *"`6 wZ `2 X s *"`6  `2 X s *"`6 <  `2 X s *"`t W `2 X s *"`  `2 X s *"`t jW `2 X s *"`t   W `2 X s *"`  `2 X s *"`aD`2 X s *"` (  `2 X s *"`7  `2 X s *"`} ``2 X s *"`2 n`2  X s *"`^A`2 !X s *"`^fA`2 "X s *"`E(`2 #X s *"``2 $X s *"`@ # `2 %X s *"``2 &X s *"`4f`2 'X s *"`a pD`2 (X s *"`4%`2 )X s *"`4`2 *X s *"`n`2 +X s *"`  `2 ,X s *"` `2 -X s *"`& 9 `2 .X s *"`6  `2 /X s *"`~ a `2 0X s *"`6 2  `2 1X s *"`6 s `2 2X s *"`t 9W `2 3X s *"` C& `2 4X s *"`t W `2 5X s *"`t sVW `2 6X s *"`sV `2 7X s *"` | _ `2 8X s *"``2 9X s *"`7  `2 :X s *"``jM C`2 ;X s *"`6 z ]  `2 X s *"` `2 ?X s *"`2P 3 f2 DX 0D["`" c  FX 0HU q%Mobile Fraction and Immobile Fraction& 2&d &   HX 0wU"`qM   2. Quantitative Analysis of FRAP>!(2DdD  IX 0DU GY D The incomplete recovery is due to some fraction of the molecules that are immobilized at the BLEACHED spot. We call these molecules the  immobile fraction . Rest of the molecules are contributing to the fluorescence recovery. We call them the  mobile fraction . 2Fl  < LX< ,$D 0`B JXB 0DԔ V KX 0U< JImmobile molecule! 2FB X s *޽h ? 33]U___PPT105.&#@j++݂D ' = @B D' = @BA?%,( < +O%,( < +D' =%(%(D' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*LX%(D' =-g6B fade*<3<*LX+   ( (  (V  ( C .Aheiko_origLB ( c $D"W"F    (   `B ( 0D8c  `B ( 0D8c    ( 0 w   }1The time constant and mobile / immobile fractions2 22d 2 RB  ( s *DgvB  ( NDp" vB ( NDp" l dz  (zd ,$@ 0 ( 0XU  [Mobile Fraction 2d   ( 0dzg ]Immobile Fraction 2d  F Jf y ( f J y ( 0JY y Half Life (t1/2)h 2 degme  lB ( <D f ,  ( 0"`qM   2. Quantitative Analysis of FRAP>!(2DdD / ( 0  ,$ 0 cIn the FRAP curve, the immobile & mobile fraction can be measured by determining the plateau level.d 2dFB ( s *޽h ? 33___PPT10.R!Ф+K*DI' = @B D' = @BA?%,( < +O%,( < +D;' =%(%(D' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*(%(D' =-g6B fade*<3<*(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*(%(D' =-g6B fade*<3<*(+8+0+( +]  @!5 0 (   T  C ,Aheiko_fitV  C .Aheiko_origB  c $DW,$@  08l       ,$@ 0ZB   s *D  ZB   s *D    0  Y Curve Fitting 2d    s *"`u h,$@ 0   S A  ??"? :8  $@ 0  64I ,$  0 QA  2f  B  c $DW,$@  0l VV  ! VV ,$@  0B   TDpV"V B   TDpVV  6l4,$ 0 S1-A  2f  z Jf y "  f J y,$@ 0 #  084JY y Half Life (t1/2)h 2 degme  lB $ <D f ,l q d7  )  qd7 ,$D 0B '  TDpd d7  (  0$b4q 6  RA  2f   * s *"`S  ,$@ 0 +  S A  ??"?S  8  $@ 0  , 0O "`qM   2. Quantitative Analysis of FRAP>!(2DdD X - 0    A more automatic way of obtaining Half life and mobile/immobile fractions is by  Curve Fitting .a 2aFJ . 0   ,$ 0 ~In curve fitting, parameters in an equation are optimized by computer. Here, we use a simple exponential equation for fitting. 2F / 0  ,$ 0 j2This is the fitted curve&  2F 0 0 S  ,$  0 zBOptimized parameters can be directly related to the FRAP kinetics.C 2CF 1 0   ,$ 0 j2A is the mobile fraction&  2F 3 0P   ,$ 0 Y!and 1-A is the immobile fraction." 2"F& 4 0  q,$ 0 HHalf Life is the time when the recovery is the half of A, by definition..I 2 F@F 5 0 m ,$ 0 ZSubstitution of f(t) by A/2 gives the formula below. Half Life can be calculated from t. [ 2FFFF;FGFHB  s *޽h ? 33=D5D___PPT10D.R!Ф+ܩD@' = @B D\@' = @BA?%,( < +O%,( < +DN' =%(D' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*. %(D' =-g6B fade*<3<*. D ' =%(Dm ' =%(DY' =4@BB BB%()))D' =-g6B fade*<3<* D' =1:Bhidden*o3>+B#style.visibility<* %(D6' =A@BB BB0B%(D' =-g6B fade*<3<*- D' =1:Bhidden*o3>+B#style.visibility<*- %(D6' =A@BB BB0B%(D' =-g6B fade*<3<*. D' =1:Bhidden*o3>+B#style.visibility<*. %(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*/ %(D' =-g6B fade*<3<*/ D ' =%(D) ' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*! %(D' =-g6B fade*<3<*! D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*0 %(D' =-g6B fade*<3<*0 DX' =%(D' =%(Dh' =A@BB BB0B%()))D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*1 %(D' =-g6B fade*<3<*1 DX' =%(D' =%(Dh' =A@BB BB0B%()))D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*3 %(D' =-g6B fade*<3<*3 D' =%(D' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*" %(D' =-g6B fade*<3<*" D6' =A@BB BB0B%(D' =-g6B fade*<3<*/ D' =1:Bhidden*o3>+B#style.visibility<*/ %(D6' =A@BB BB0B%(D' =-g6B fade*<3<*0 D' =1:Bhidden*o3>+B#style.visibility<*0 %(D6' =A@BB BB0B%(D' =-g6B fade*<3<*1 D' =1:Bhidden*o3>+B#style.visibility<*1 %(D6' =A@BB BB0B%(D' =-g6B fade*<3<*3 D' =1:Bhidden*o3>+B#style.visibility<*3 %(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*4 %(D' =-g6B fade*<3<*4 D' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*) %(D' =-g6B fade*<3<*) DN' =%(D' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<** %(D' =-g6B fade*<3<** D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*+ %(D' =-g6B fade*<3<*+ D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*5 %(D' =-g6B fade*<3<*5 +H+0+   ++0+  ++0+-  ++0+.  ++0+.  ++0+/  ++0+/  ++0+0  ++0+0  ++0+1  ++0+1  ++0+3  ++0+3  ++0+4  ++0+5  +    ` (  ` ` C *AoriginalQD,$@ 0 ` 0S JThe FRAP curve equation we will use& .& 2&f &  ` 6T|% 3 W+  cAcquisition Bleaching 2f  >L  }) `# "ph ` s *"` })r ` S A $??"?: $XB ` 0Dp   ` 0 Z"`qM   2. Quantitative Analysis of FRAP>!(2DdD m ` 0a  In this course, we will use same type of exponential equation as it was shown in the previous slide. This equation however is corrected for the bleaching due to the image acquisition bleaching (underlined term).  2FN ` 0T"vI,$ 0 In this graph, acquisition bleaching curve is plotted in blue color. Since we illuminate the sample for taking images, the fluorescence intensity decays slightly. This decay is considered in the above fitting equation. The kinetics of acquisition bleaching is actually unknown. It is also possible to directly correct the Frap curve by the acquisition bleaching at each time point.  2FB ` s *޽h ? 33___PPT10.R!Ф+=oDy' = @B D4' = @BA?%,( < +O%,( < +Dk' =%(D' =%(D<' =A@BB@BB0B%()?)?D' =.K7 BBBBB]M -2.77778E-6 -3.7037E-6 L -0.00382 -0.05439 *3>*B ppt_xB ppt_y=@0BBAApBBOBR߼<*`D'' =4@BB8BB%()?)?D{' =.C7 BBBBBUM -2.5E-6 -2.96296E-6 L 0.21268 -0.14166 *3>*B ppt_xB ppt_y=@0BBAApBB=B<*`D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*`%(D' =-g6B fade*<3<*`Df' =A@BB BB0B%()))D' =-g6B fade*<3<*`D' =1:Bhidden*o3>+B#style.visibility<*`%(D)' =4@BB BB%(D' =-g6B fade*<3<*`D' =1:Bhidden*o3>+B#style.visibility<*`%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*`D' =1:Bhidden*o3>+B#style.visibility<*`%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*`%(D' =-g6B fade*<3<*`++0+` ++0+` ++0+` ++0+` +    <+ (  <l W < W,$@ 0h < s *"`d Wr < S A ??"? ) \ < C ,Aheiko_fit  < <X  TModeling f   < 0P`  $ 0NF___PPT9(  (Specific Interactions Diffusion Advanced& Analytical Approach Numerical ApproachB* 2' 2*f'f Q 8X < 0x"`qw 3. Modeling & FRAP>(2DdD   < 0ô ,,$ 0 &What does the fitting and obtained parameters actually mean, in terms of molecular dynamics? This question leads us to the world of  Modeling . We will consider following three molecular dynamics in next slides. ` 2IFF+FFHF  < 0Z- Y,$ 0 Specific interaction is also called chemical interaction. Specific interaction and diffusion are two basic models. In actual research, there are more various and advanced models.  2FB < s *޽h ? 33___PPT10.#7/+?D ' = @B DR ' = @BA?%,( < +O%,( < +D ' =%(D' =%(D6' =A@BB@BB0B%()?)?D}' =.E7 BBBBBWM 5.55556E-7 1.11111E-6 L 0.00104 -0.3169 *3>*B ppt_xB ppt_y=@0BBAApBB:B^"<*<D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*<%(D' =-g6B fade*<3<*<D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* <%(D' =-g6B fade*<3<* <D' =%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*<%(D' =-g6B fade*<3<*<D' =%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* <%(D' =-g6B fade*<3<* <++0+<- ++0+<- ++0+ <- ++0+ <- +K0 Pu:(    0$b-ZM z fA. Specific Interactions 2f  `2 $ s *"`}``2 % s *"` `2 ) s *"`pjS `2 * s *"`#J-`2 + s *"`s V `2 , s *"`5 3  `2 - s *"`c F `2 7 s *"`}& ``2 = s *"`pS 8  /  o 3 h2 & s *"` / h2 ' s *"`/ ^ Ah2 ( s *"`  h2 . s *"` m Ph2 2 s *"` n h2 3 s *"` @ #h2 4 s *"` /~ h2 8 s *"`#  h2 < s *"`0  }h2 @ s *"`@ # h2 F s *"`  }h2 [ s *"`m P XB p 0Dp%  / b XB q@ 0Dp z  r 0E-F   dPre-Bleach&  2 f   s 0H-"`qw 3. Modeling & FRAP>(2DdD r t 0M-dY Let s consider a cluster of molecules, and the same molecular species unbound to the cluster. There are continuous exchange between the bound and the un-bound molecules. The system is in a steady state. The cluster size is not changing.  2FB  s *޽h ? 33___PPT10i.R!Ф+D=' = @B + @|W(  |`2 | s *"`}``2 | s *"` `2 | s *"`pjS `2 | s *"`#J-`2 | s *"`s V `2 | s *"`5 3  `2  | s *"`c F `2  | s *"`}& ``2  | s *"`pS F  /   |  3 h2  | s *"` / h2 | s *"`/ ^ Ah2 | s *"`  h2 | s *"` m Ph2 | s *"` n h2 | s *"` @ #h2 | s *"` /~ h2 | s *"`#  h2 | s *"`0  }h2 | s *"`@ # h2 | s *"`  }h2 | s *"`m P XB | 0Dp%  / b XB |@ 0Dp z  | 07-F   UBleach! 2f   | 0|-"`qw 3. Modeling & FRAP>(2DdD  | 0-ZM z fA. Specific Interactions 2f   | 0-F* 0& then we bleach only the cluster. Immediately after the bleaching, all molecules consisting the cluster loses fluorescence. } 2}FB | s *޽h ? 33___PPT10i.R!Ф+D=' = @B +K0 P#%N(  `2  s *"`}``2  s *"` `2  s *"`pjS `2  s *"`#J-`2  s *"`F 0  ) `2  s *"`5 3  `2  s *"`}& ``2  s *"`pS F  /     3 h2   s *"` / h2  s *"`/ ^ Ah2  s *"`  h2  s *"` m Ph2  s *"` n h2  s *"` @ #h2  s *"` /~ h2  s *"`#  h2  s *"`0  }h2  s *"`@ # h2  s *"`  }h2  s *"`m P XB  0Dp%  / b XB @ 0Dp z `2  s *"` `2  s *"`  m`2  s *"` s   0|-V Yv  `Recovery&  2 f  $l   ! ,$@ 0  0`-]  hk16 2fn     0l-  hk26 2fn   " 0$-ZM z fA. Specific Interactions 2f   # 0,x-"`qw 3. Modeling & FRAP>(2DdD  $ 0- W9 aSince there is a continuous exchange of the molecules, the cluster regains fluorescence by time. b 2bF % 0-VW,$ 0 GWe define the exchange rate k1 and k2 to see this event in more detail.H 2FFNFFN#FB  s *޽h ? 33  ___PPT10z .R!Ф+zD ' = @B D ' = @BA?%,( < +O%,( < +D ' =%(D' =%(D' =4@BB4BB%(D' =1:Bvisible*o3>+B#style.visibility<*!%(D' =0l9 BBzCzCBB*<3<*!%())?D' =.7 BBBBBM -0.46736 0.92887 C -0.37517 0.88508 -0.02552 0.75279 0.0908 0.66613 C 0.20747 0.57948 0.21649 0.50394 0.23177 0.40825 C 0.24705 0.31256 0.22118 0.15964 0.18264 0.09152 C 0.1441 0.02341 0.03802 0.0 0.0 0.0 *3>*B ppt_xB ppt_y=B0B<*!%())?D' =-g6B fade*<3<*!D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%%(D' =-g6B fade*<3<*%+8+0+%- +    0xv (  xhF Z x 7 &N  6 x   x 0- P3  l[F]180( 2fd   x 0-#j P(0( 2f   x 08,-a }& Z!(0( 2f   x 0-a &< Z!(0( 2f   x 0|- 6&} rk1@0( 2fn    x 0- `& rk2@0( 2fn    x 0$-P l[F]280( 2fd  n  x 08c"`Z# n  x 08c"`#   x 0D-"`qw 3. Modeling & FRAP>(2DdD  x 0-w3  fA. Specific Interactions 2f   x 0h G, We consider a simplified model. The exchange of molecules between the cluster and the surrounding can be thought as molecule exchange between two chambers as above. The exchange is limited by a wall in between. The transfer rates in two directions are k1 and k2 respectively. We define the number of molecules in the cluster as [F]1. The number of molecules in surrounding environment as [F]2. 2FFNFFNDFFN8FFNFB x s *޽h ? 33___PPT10i.R!Ф+D=' = @B +  !"[(  hF Z  ^ O&N  6     0Ŀ P3  l[F]180( 2fd    0`ċ#j P(0( 2f    0ȋa }& Z!(0( 2f    0ŋa &< Z!(0( 2f    0l΋ 6&} rk1@0( 2fn     0Ӌ `& rk2@0( 2fn     0ԋP l[F]280( 2fd  n   08c"`Z# n   08c"`#  F  ,<    ,w@N   P Iv<h  s *"`r  S A ??"?M @N  ,    ,`h  s *"` , r  S A ??"? C  N 6v   6v h  s *"`6v r  S A ??"?% 1  r  S A ??"? I  lB  <DԔ   0ߋ"`qw 3. Modeling & FRAP>(2DdD   0w3  fA. Specific Interactions 2f  k  0T] m Using this model, we can compose a differential equation. See equation (1). In this differential equation, the speed of the changes in the number of molecules in the chamber 1, which is the cluster, is defined.  2F  B(I   Kchamber 1 (cluster)F  BxI ;  Ochamber 2 (surrounding)F  0, ;(1) 2F  0 ,  <(2)  2F ! 0  <(1a) 2F " 0$j t,$ 0 C Solving the equation (1) and substituting the rate constants as shown in (1a), we get the equation (2). This is the same equation as we have used for the fitting in the previous slides. We now know the meaning of the parameters we obtained by fitting the FRAP curve.   2 FB  s *޽h ? 33___PPT10.R!Ф+3D4' = @B D' = @BA?%,( < +O%,( < +D&' =%(D' =%(D6' =A@BB BB0B%(D' =-g6B fade*<3<*D' =1:Bhidden*o3>+B#style.visibility<*%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*"%(D' =-g6B fade*<3<*"+p+0+ ++0+" + ; `GS(  `2  s *"`sVn`2  s *"`n`2  s *"`^A`2  s *"`^/  A`2  s *"` `2  s *"``2  s *"`( `2  s *"`i`2  s *"`a9D`2  s *"`4`2  s *"`4iL `2  s *"`iL n`2  s *"`)  `2  s *"`w `2  s *"`Y< `2  s *"`& `2  s *"`6  `2  s *"`6 wZ `2   s *"`6  `2 ! s *"`6 <  `2 " s *"`t W `2 # s *"`  `2 $ s *"`t jW `2 % s *"`t   W `2 & s *"`  `2 ' s *"`aD`2 ( s *"` (  `2 ) s *"`7  `2 * s *"` n`2 + s *"`2 n`2 , s *"`^A`2 - s *"`^fA`2 . s *"`E(`2 / s *"``2 0 s *"`| _ `2 1 s *"``2 2 s *"`4f`2 3 s *"`a pD`2 4 s *"`4%`2 5 s *"`4`2 6 s *"`n`2 7 s *"`  `2 8 s *"` `2 9 s *"`& 9 `2 : s *"`6  `2 ; s *"`6  `2 < s *"`6 2  `2 = s *"`6 s `2 > s *"`t 9W `2 ? s *"` C& `2 @ s *"`t W `2 A s *"`t sVW `2 B s *"`sV `2 C s *"` | _ `2 D s *"``2 E s *"`7  `2 F s *"` `2 G s *"`%  `2 H s *"`P 3`2 I s *"`} ` `2 J s *"`3P 3 `2 K s *"`@ # F   L  fB M 6D ] CfB N 6D  S fB O 6D @ Q }2 P <D["`" c ,$D 0 Q 0tF"`qw 3. Modeling & FRAP>(2DdD  R 0$FT \ B. Diffusion (2 f  / S 0uFI FRAP could also occur without the specific interactions. Simply the diffusion drives the exchange of molecules between bleached and non-bleached area.  2FB  s *޽h ? 33___PPT10.R!Ф+D[' = @B D' = @BA?%,( < +O%,( < +D+' =%(%(D' =%(D[' =4@BB BB%()))D' =1:Bvisible*o3>+B#style.visibility<*P%(D' =-g6B fade*<3<*PD9' =%(D' =%(D/' =4@BB8BB%()?)?D' =.K7 BBBBB]M -2.77778E-7 1.11111E-6 L -0.07083 -0.02894 *3>*B ppt_xB ppt_y=@0BBAApBBBn<*FD' =4@BB1BB%()?)?Dq' =.97 BBBBBKM 0.00017 0.00232 L 0.03594 0.09121 *3>*B ppt_xB ppt_y=@0BBAApBB(}<Ba 6=<*D!' =4@BB8BB%()?)?Du' =.=7 BBBBBOM 0.00434 -0.00301 L 0.03993 -0.06065 *3>*B ppt_xB ppt_y=@0BBAApBB<B{ <*&D+' =4@BB8BB%()?)?D' =.G7 BBBBBYM 3.05556E-6 -1.48148E-6 L 0.0434 -0.11018 *3>*B ppt_xB ppt_y=@0BBAApBBDZ<BŨa<*HD!' =4@BB1BB%()?)?Du' =.=7 BBBBBOM -0.00364 -0.00532 L 0.03212 0.07338 *3>*B ppt_xB ppt_y=@0BBAApBB(}<Bh/!=<**D' =4@BB1BB%(L)?)?Ds' =.;7 BBBBBMM 0.0007 -0.01065 L -0.05798 0.01551 *3>*B ppt_xB ppt_y=@0BBAApBB[BbT<<*0D-' =4@BB1BB%()?)?D' =.I7 BBBBB[M -2.77778E-6 -7.40741E-7 L 0.02778 0.07894 *3>*B ppt_xB ppt_y=@0BBAApBB9c<Bh/!=<*ID#' =4@BB1BB%()?)?Dw' =.?7 BBBBBQM -0.00364 -0.00532 L 0.03993 -0.07361 *3>*B ppt_xB ppt_y=@0BBAApBBDZ<BpS <*<D!' =4@BB8BB%( )?)?Du' =.=7 BBBBBOM 0.00833 0.01041 L -0.05052 -0.00533 *3>*B ppt_xB ppt_y=@0BBAApBBB<*7D' =%(D' =%(D)' =4@BB BB%(D' =-g6B fade*<3<*LD' =1:Bhidden*o3>+B#style.visibility<*L%(+- D<p  (    0Fw \ B. Diffusion (2 f    0LFD[ `Theoretical approach 2d  "8  ra h  s *"`j  C A 1?"a 1(  0lF,wP  Phair & Misteli (2000) 2f@     U 06L  `  # E ? h   s *"` ` j   C A 4?"   4  6XF w7 ECu(r) : unbleached fluorophore concentration r: radial distance K: bleach constant I(r) is the Gaussian intensity profile of the laser w: half width at e-2 intensity. D 2fn f4 P  S   0F`w  JC0: plateau fluorescence tD: the characteristic diffusion time. tD= w2/4D K 2fnfgn%fgnfnf@  %    l2  68c"`g^B @ 6DԔ   0r7 "`qw 3. Modeling & FRAP>(2DdD   07  R  WFRAP kinetics based on diffusion was examined theoretically many years a go by Axelrod et al. (1976). The formula they proposed are still in used by many researchers with small modifications. The equation shown in the left needs a parameter before fitting. Bleaching constant K is obtained by measuring the laser intensity distribution Cu(r).X 2WFFFF;FFNFF4OU 0OdB  s *޽h ? 33___PPT10i.R!Ф+D=' = @B +  <(     0<7 !   Ellenberg et al. 1997 2d&     U 0  0܏7 '   ^Empirical approach 2d  *8 *   9h   s *"`* r   S A 6??"? 6  087 "`qw 3. Modeling & FRAP>(2DdD   07 T \ B. Diffusion (2 f  /  07    There are also empirical formulas for fitting the FRAP curves. Above equation is an example. w is the width of bleached spot. Ifinal is the plateau level after recovery. The diffusion coefficient D can be directly obtained by fitting this equation.  2]FF FFN@FF7F~xB  s *޽h ? 33___PPT10i.R!Ф+D=' = @B +! yqp(     07 -n JC. More Advanced& Analytical Approach0&ff & Z  C 2A PresleyModelW)B   0p7  W  Presley et al. (2002)> 2ddd  U 0.l &i /- ,$@ 0h  s *"`&ir  S A @??"?jCK @r   S A A??"?jCJK  Ar   S A B??"?. V6  Br   S A C??"?j6 R>  Cr   S A D??"?jF :<  Dr   S A E??"?j) 1  Er  S A F??"?j9 ,A Fr  S A G??"?jIQ Gr  S A H??"?jYC H  0H7 "`qw 3. Modeling & FRAP>(2DdD   C 2A PresleyModelw,$@ 0S  0d7  C,$ 0 Presley et al. (2002)> 2ddd  U 0T  0D7 7  Advanced techniques for the modeling deal with complex molecular interactions such as the one shown above. This is a diagram of molecular interactions involved in the vesicle trafficking.  2F  07 S<,$ 0 For all these interactions, it is possible to design differential equations for each molecular species involved in the system. These equations can be solved and fitted to the FRAP curves from the experiment to study their dynamics. 2FB  s *޽h ? 33 ___PPT10.R!Ф+p>D' = @B D' = @BA?%,( < +O%,( < +D' =%(Ds' =%(DY' =4@BB BB%()))D' =-g6B fade*<3<*D' =1:Bhidden*o3>+B#style.visibility<*%(Df' =A@BB BB0B%()))D' =-g6B fade*<3<*D' =1:Bhidden*o3>+B#style.visibility<*%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-g6B fade*<3<*D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-g6B fade*<3<*Dh' =A@BB BB0B%()))D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-g6B fade*<3<*D6' =A@BB BB0B%(D' =-g6B fade*<3<*D' =1:Bhidden*o3>+B#style.visibility<*%(D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =-g6B fade*<3<*++0+7  ++0+7  ++0+7  ++0+7  +[  rj(    07 "`qw 3. Modeling & FRAP>(2DdD    07 -T LC. More Advanced& Numerical Approach0'ff '   07   VThe example shown in the previous slide was a modeling by analytical approach  which is to compose differential equations and solve them for the fitting. There is another approach for an advanced modeling of biological system. That is to derive the estimated FRAP curve by simulation and compare it with the FRAP curve from the experiments. Simulation has potential in including many more parameters than the analytical approach. This approach is powerful but not yet widely used and still in progress. Simulations in this sense can be viewed on the web, such as Adriaan Houtsmuller Lab s web site. Z 2ZF45 U 05WB  s *޽h ? 33___PPT10i.h,PHm+D=' = @B +$) 0 L(   z '    C' ,$@ 0   07    |"B([E], [S], [ES], [P], t, x, y, z)*#0( 2#f # &   07 ' M  :Topo-biochemistry (in vivo)0:0( 2ff&    8 D   D _ @ D"   D" *   6`BLDF  .Biochemistrysolution 0:0( 2ff0        6SL v"  sB([E], [S], [ES], [P], t)*0( 2f  T jI}  # M    0UL  _[E]+[S](0( 2f     0TL  \[ES](0( 2f     04^L  I Z!(0( 2f     0YL z  Z!(0( 2f     0XeL#   \k1*0( 2f     0|cL#  , \k2*0( 2f  h   s *"`j}   0DlL7z Z!(0( 2f     0oLFI [[P](0( 2f     0sLd \k3*0( 2f  V   C &Abeaker? ~  C "A cell ,$@ 0z ",   ",,$@ 0z2   s *3C"?J}z2   s *3C"?"}z2   s *3C"?9  z2   s *3C"? ,  0xL"`g WSummary (2f    0\|LV 7, $What is the significance of FRAP technology? Let s compare it with traditional biochemistry. Biochemistry studies enzymes in the solution. Within beakers such as the one shown in above, enzyme  substrate kinetics is measured and characterized.  2F  0Lx i,$ 0 ^Now, we can measure the molecular interaction directly inside cells through microscope. This enables us to consider the spatial information that was disregarded in biochemistry. Maybe it would be more suited to call this technology as  topo-biochemistry . Since the site of interaction does matter within cell, these technology are now inevitable in understanding the cell as a system. 2FB  s *޽h ? 33 ___PPT10.o+ @oDF' = @B D' = @BA?%,( < +O%,( < +D8' =%(D' =%(D' =4@BB8BB%()?)?Dm' =.57 BBBBBGM -4.44444E-6 0.0 L 0.004 -0.2625 *3>*B ppt_xB ppt_y=@0BBAApBBO:Bff<* D6' =A@BB BB0B%(D' =-g6B fade*<3<* D' =1:Bhidden*o3>+B#style.visibility<* %(D) ' =%(D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D+' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* D8' =A@BB BB0B%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =-g6B fade*<3<* +p+0+ L ++0+ L +# ^ V  8 (  8RB 8 s *DԔdd! 8 0L,' ~Thanks to: Heiko Runz (EMBL Heidelberg) Joel Beaudouin (EMBL Heidelberg) Adriaan Houtsmuller (JNI EMC, Rotterdam) EAMNET 2d                U 0x~ 8 0hL ?,$ 0 QFor Suggestions, Comments and Questions, please e-mail Kota Miura (miura@embl.de)R 2RFRB 8 s *DԔIIRB 8@ s *DԔ'RB 8@ s *DԔ-6  8 0LzZIP,$  0 ~Thanks to: Heiko Runz (EMBL Heidelberg) Joel Beaudouin (EMBL Heidelberg) Adriaan Houtsmuller (JNI EMC, Rotterdam) EAMNET.~fF               U 0x~  8 0L! Gf ,$  0 aThe European Advanced Light Microscopy Network (EAMNET) is a EU funded network of eight European laboratories and two industrial partners working in the field of light microscopy. The aim of EAMNET is to assist scientists in exploiting the power of imaging by organizing practical teaching courses, creating online teaching modules and offering software packages for microscopy. All EAMNET partners are also members of the European Light Microscopy Initiative (ELMI). 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(1976) Biophys J. 16(9):1055-69 "Mobility measurement by analysis of fluorescence photobleaching recovery kinetics. Ellenberg et al. (1997) J Cell Biol.;22;138(6):1193-206. "Nuclear membrane dynamics and reassembly in living cells: targeting of an inner nuclear membrane protein in interphase and mitosis." Phair RD & Misteli T. (2001) Nat Rev Mol Cell Biol. 2(12):898-907. "Kinetic modelling approaches to in vivo imaging." Presley JF et al. (2002) Nature.9;417(6885):187-93. "Dissection of COPI and Arf1 dynamics in vivo and role in Golgi membrane transport." 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