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" # 0 2 4 41 4 ;<}.*$ % :*&  ', :A?CB/b$,x[2ǸuWqb$AYӐ؂W:Yqb$"K)5 q2$Pܟ_FٞY]Ab$]hP.4ljm!Ւb$TzLrќD?.CKӴ"$L`ֻؘ31_="$o9l(_J^6}D$b$`WE \azKb$`kڅ>/u|ph+*!Cf"$ڃ{3|wFsn 6b$u!xUse a test statistic on all genes Rank them Select top k ?Z?#UProblem with current solutionDEach gene independently scored Top k ranking genes might be very similar and therefore no additional information gain Reason: genes in similar pathways probably all have very similar score What happens if several pathways involved in perturbation but one has main influence Possible to describe this pathway with fewer genesEZE8 Problem of redundancy y%OutlineoMotivating example Biological background Problem statement Current solution Proposed attack Results Future workDp  99 Proposed solutionSeveral possible approaches next neighbors correlation euclidean distance Approach: instead use clustering Advantages using clustering techniques natural embedding many different distance functions possible different shapes, models possible j.I_.I_7 ; 2Hard clustering  k-means MSoft - Fuzzy Clustering NFuzzy examples RFuzzy softness 1.3 +Fuzzy softness 1.25 *Fuzzy softness 1.2 (Fuzzy softness 1.15 )Fuzzy softness 1.05 r Selecting genes from clustersTwo way filter: exclude redundant genes, select informative genes Get as many pathways as possible Consider cluster size and quality as well as discriminative power VHow many genes per cluster?MConstraints: minimum one gene per cluster maximum as many as possible Take genes proportionally to cluster quality and size of cluster Take more genes from bad clusters Smaller quality value indicates tighter cluster Quality for k-means: sum of intra cluster distance Quality for fuzzy c-means: avg cluster membership probability 9 sb9bAb"B0bqBb)![Which genes to pick?Choices: Genes closest to center Genes farthest away Sample according to probability function Genes with best discriminative power& z zbComparison Evaluation ,Support Vector machinesFind separating hyperplane with maximal distance to closest training example Advantages: avoids overfitting can handle higher order interactions and noise using kernel functions and soft margin DbZiZ Zbi, O Wz&OutlineoMotivating example Biological background Problem statement Current solution Proposed attack Results Future workDp  I : Experimental setupRDatasets: Alons Colon (40 tumor and 22 normal colon adenocarcinoma tissue samples) Golubs Leukemia (47 ALL, 25 AML) Nottermans Carcinoma and Adenoma (18 adenocarcinoma, 4 adenomas and paired normal tissue) Experimental setup: calculate LOOCV using SVM on feature subsets do this for feature size 10-100 (in steps of 10) and 1-30 clusters ZZZpZ  #<"" "8""p"b $ >Results ?fuzzy c-means vs k-means  @Different test-statistics dComparing best resultseHow about randomly choosing?$t! Related work&Tusher, Tibshirani and Chu (2001): Significance analysis of microarrays applied to the ionizing radiation response, PNAS 2001 98: 5116-5121 Ben-Dor, A., L. Bruhn, N. Friedman, I. Nachman, M. Schummer, and Z. Yakhini (2000). Tissue classification with gene expression profiles. In Proceeding of the fourth annual international conference on computational molecular biology, pp. 54-64 Park, P.J., Pagano, M., Bonetti, M.: A nonparametric scoring algorithm for identifying informative genes from microarray data. Pac Symp Biocomput :52-63, 2001. Golub TR, Slonim DK, Tamayo P, Huard C, Gaasenbeek M, Mesirov JP, Coller H, Loh M, Downing JR, Caligiuri MA, Bloomfield CD, and Lander 18 ES. Molecular classification of cancer: class discovery and class prediction by gene expression monitoring. Science 286: 531-537, 1999. J. Weston, S. Mukherjee, O. Chapelle, M. Pontil, T. Poggio, and V. Vapnik. Feature selection for SVMs . In Sara A Solla, Todd K Leen, and Klaus-Robert Muller, editors, Advances in Neural Information Processing Systems 13. MIT Press, 2001. 11 'ZbBbbBbBb w c     (o{'OutlineoMotivating example Biological background Problem statement Current solution Proposed attack Results Future work<p  Q A Future workProblem how to find best parameters (model selection, model based clustering, BIC) Combine good solutions Incorporate overall cluster discriminative power into quality score Use of non integer error score ROC analysisu"Summary4Used clustering as a pre-filter for feature selection in order to get rid of redundant data Defined a quality measurement for clustering techniques Incorporated cluster quality, size and statistical property into feature selection Improved LOOCV error for almost all feature sizes and different related tests55fResult Notterman g Result Golub, Result Alon- Result Alon 2/35<=CDFL O P Q W XY\^cs ` ̙33` ` ff3333f` 333MMM` f` f` 3>?dd (? 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   c BCDEFf?%% #{*d2P98D$MW _gpx zsm"g0b?\PUdNoJB91( KL@h E   c BCDEFf?%% #|)d2Q99D%MW _gpx zsm#g0a>\PUdNoJB91' KL@g D   0%`p`v  : spot genes 2 Il    ,$D  0fH@ p Pf2  6(pf2  6P@f2  6@T    # @   BCDE,F4s?KK@_6X,Q$KD>8 2 -'"   "!%&*,-3296@:H>PCYHdNmTuY}_djpv{ys}lxdsZnPh@_@z <    BCDE(F0s?JJ@^5W,Q$KD>8 2 -'"  " %&*,-2296@:H>PCYHdNmSuY|_djou{ys}kxcsZmPh@^@     BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@     BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@d %    BCDEF A?FF  $y(s-l1c6Z<PA@K6R,X$_ekq w |  &,|3x9t@pHkPgYad[mVuP}JE?94.(" @ }    BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@ i    BCDEF A?FF !$y(s,k1c6Z;PA@K5Q,X$^djp v |  &,{2w9s@oHjPfYad[mUuO|JD>93-'!@5     BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@  T    # (   BCDE,F4s?KK@_6X,Q$KD>8 2 -'"   "!%&*,-3296@:H>PCYHdNmTuY}_djpv{ys}lxdsZnPh@_@z <    BCDE(F0s?JJ@^5W,Q$KD>8 2 -'"  " %&*,-2296@:H>PCYHdNmSuY|_djou{ys}kxcsZmPh@^@     BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@     BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@d %     BCDEF A?FF  $y(s-l1c6Z<PA@K6R,X$_ekq w |  &,|3x9t@pHkPgYad[mVuP}JE?94.(" @ }     BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@ i     BCDEF A?FF !$y(s,k1c6Z;PA@K5Q,X$^djp v |  &,{2w9s@oHjPfYad[mUuO|JD>93-'!@5      BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@  T     # X8   BCDE,F4s?KK@_6X,Q$KD>8 2 -'"   "!%&*,-3296@:H>PCYHdNmTuY}_djpv{ys}lxdsZnPh@_@z <    BCDE(F0s?JJ@^5W,Q$KD>8 2 -'"  " %&*,-2296@:H>PCYHdNmSuY|_djou{ys}kxcsZmPh@^@     BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@     BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@d %    BCDEF A?FF  $y(s-l1c6Z<PA@K6R,X$_ekq w |  &,|3x9t@pHkPgYad[mVuP}JE?94.(" @ }    BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@ i    BCDEF A?FF !$y(s,k1c6Z;PA@K5Q,X$^djp v |  &,{2w9s@oHjPfYad[mUuO|JD>93-'!@5     BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@    0% & ; cell tissue 2 al p  p ,$D  0"`@    }p   N     @     BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@     BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P   BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@     BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    #  @    BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@     BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P    BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   !  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BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P /  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   0  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    1# @ `  2  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   3  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P 4  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   5  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    6#  C  7  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   8  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P 9  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   :  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    ;#    <  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   =  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P >  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   ?  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K f2  6 <0  0t% p < extract cDNA 2 1ml  `   ` ,$D  0k@  c  ~ k  T    @#  C  A  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   B  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P C  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   D  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    E#    F  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   G  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P H  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   I  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    J# pX  K  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   L  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P M  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   N  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    O#   P  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   Q  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P R  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   S  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    T# c  U  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   V  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P W  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   X  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    Y#  0 C  Z  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   [  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P \  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   ]  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    ^#   h  _  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   `  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P a  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   b  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K  T    c# @  d  BCDE(F0?JJ\NfUp\xbhnuz|vpj{cw\sTnMjCe9_/Y'S NHC= 71+%      $(#,)0159:B?ME\N@   e  BCDE(F0?JJ\OfUo\wb~hntz|vpi{bw[sTnLjCe8^/Y'SMHB= 71,%     !$(#,)0058:B?LE\O@ P f  BCDEF ?FF #)}1x9tBnMi\_fXpQxKE?93.("  "|&v*p.j2c6\:T?MCCH9N/T'Y _ekp v|@   g  BCDEF ?FF #)}0y8tBnLi\_fXoQwK~E?93-("  "|&v*p.i2b6[:T?LDCH8N/T'Z_ekp v|@K T  ~> h#   xB i H1? j NԷ%xaxa? ~> 1*T  ~> k#  6xB l H1? m N4%xaxa? ~> 1*T  ~> n# XWvxB o H1? p N%xaxa? ~> 1*T  ~> q#  ?6xB r H1? s N%xaxa? ~> 1*T  ~> t#  p xB u H1? v NT%xaxa? ~> 1*T  ~> w# 0 0 N xB x H1? y N%xaxa? ~> 1*T  ~> z#   xB { H1? | N%xaxa? ~> 1*f2  6~    0Ժ%p`  : label cDNA 2 [l    ,$D  0Z@  w   w Y@ 0   T D` # ` `  0?DL  S BCMDEF????L_ @`  S BrCDEF?q @D`M  c BCDEF3? 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((hA~8/' w"g(U-K16:$AIPYajtuvut n id`"[.U@OUGhAQT@> n T  '#   xB ( H1?~2~} ) N4%xaxa? 1* - c BCDEF3?((QH9>%4, "   #0 @&Q,d3p7@HQYclvzuoje_|WdOQHQT@ s   . c BCDEF3?))QI9?%4, #  #0!>&P-d3o7AIRZdmw|vqkf`|XdPQIST@ p  / c BCDEFf?%% #{*d2P98D$MW _gpx zsm"g0b?\PUdNoJB91( KL@ v  0 c BCDEFf?%% #|)d2Q99D%MW _gpx zsm#g0a>\PUdNoJB91' KL@ u  7 c BCvDEF3? &&  ".@&U-h3}=FNU]dltutnic^XvRgMUFKC5<$3,$ MP@ ' u  8 c BCuDEF3? %% ".@&U-g3}=ENU\dkstmhb]WvRgLUFKB5;$3,$ KL@ B u  9 c BCDEF3? ((QH9>%4, "   #0 @&Q,d3p7@HQYclvzuoje_|WdOQHQT@ { +   : c BCDEF3? ))QI9?%4, #  #0!>&P-d3o7AIRZdmw|vqkf`|XdPQIST@ { *  ; c BCDEFf? %% #{*d2P98D$MW _gpx zsm"g0b?\PUdNoJB91( KL@ ( {  < c BCDEFf? %% #|)d2Q99D%MW _gpx zsm#g0a>\PUdNoJB91' KL@ ' {   = c BCxDEF? ))gA}8/'   v"g(U.K159$AIQYajtvwvto je`"\.V@OUHgAST@ u ,  > c BCwDEF? ((hA~8/' w"g(U-K16:$AIPYajtuvut n id`"[.U@OUGhAQT@ v  T e ?#   } < xB @ H1?II# A N%xaxa?e 1* C c BCvDEF3? &&  ".@&U-h3}=FNU]dltutnic^XvRgMUFKC5<$3,$ MP@ B <  D c BCuDEF3? %% ".@&U-g3}=ENU\dkstmhb]WvRgLUFKB5;$3,$ KL@ B V  E c BCDEF3? ((QH9>%4, "   #0 @&Q,d3p7@HQYclvzuoje_|WdOQHQT@ F H   F c BCDEF3? ))QI9?%4, #  #0!>&P-d3o7AIRZdmw|vqkf`|XdPQIST@ C H  G c BCDEFf? %% #{*d2P98D$MW _gpx zsm"g0b?\PUdNoJB91( KL@ H I  H c BCDEFf? %% #|)d2Q99D%MW _gpx zsm#g0a>\PUdNoJB91' KL@ H H   I c BCxDEF? ))gA}8/'   v"g(U.K159$AIQYajtvwvto je`"\.V@OUHgAST@ S B  J c BCwDEF? ((hA~8/' w"g(U-K16:$AIPYajtuvut n id`"[.U@OUGhAQT@ 9 C T  K#  Z  / xB L H1?0t0 M N%xaxa? 1* O c BCvDEF3? &&  ".@&U-h3}=FNU]dltutnic^XvRgMUFKC5<$3,$ MP@s  X  P c BCuDEF3? %% ".@&U-g3}=ENU\dkstmhb]WvRgLUFKB5;$3,$ KL@s  r  Q c BCDEF3? 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TB B c $D0 `P TB  c $D0 `P T 0 `P  # @poTB B c $D0 `P TB  c $D0 `P f2  6P?f2  6@p f2  6p`r A 0` V 07p U%Randomly assign cluster to each point& 2&l  p g p,$D  0`r C 0` 0N@   U f 3 6 T 0 `P  7# `TB 8B c $D0 `P TB 9 c $D0 `P T 0 `P  :# @oTB ;B c $D0 `P TB < c $D0 `P f2 = 6 ? f2 ? 6pO  T 0 `P  Q# @0ooTB RB c $D0 `P TB S c $D0 `P `2 T 0P   X 0D8 p Q!Reassign points to nearest center" 2"l @ h@,$D  0r Z B8@ :  [ 0d9   IIterate until convergence 2|l @  f@ ,$D  0f % 6  T 0 `P  &#  O TB 'B c $D0 `P TB ( c $D0 `P T 0 `P  )# 0 o TB *B c $D0 `P TB + c $D0 `P T 0 `P  ,# @ o TB -B c $D0 `P TB . c $D0 `P f2 / 6 ? f2 0 6@ p0 f2 1 6p O `r B 0`@  W 0$:   `Find centroids 2 Z2 _ s *0  Z ` s * @  H  0޽h ? ̙33   aY04(  4x 4 c $: " *  %  4 0:  h8instead of hard assignment, probability for each cluster9 29X 4 C 0AD:\fuzz.pngp hX   4 0Dp@\  yVery similar to k-means but fuzzy softness factor m (between 1 and infinity) determines how hard the assignment has to bez 2zH 4 0޽h ? ̙33   g_@ <(  <x < c $ " *  % ? < 0 P`  Nottermans carcinoma dataset: 18 colon adenocarcinoma and 18 normal tissues data from 7457 genes and ESTs cluster all 36 tissues 2$ LH < 0޽h ? ̙33  ~PD(  Dx D c $d " *  % ^ D C 6A H:\cb\nc13.png=H D 0޽h ? ̙33  `(  ~  s *Ī " *  % `  C 8A H:\cb\nc125.png=H  0޽h ? ̙33  p(  ~  s *$ " *  z ^  C 6AH:\cb\nc12.png=H  0޽h ? ̙33  (  ~  s * " *   `  C 8A  H:\cb\nc115.png=H  0޽h ? ̙33  (  ~  s * " *   `  C 8A  H:\cb\nc105.png=H  0޽h ? ̙33  *(  x  c $D " *   r  S  P  H  0޽h ? ̙33J  \(  \x \ c $ĭ " *   x \ c $$ P  R \ s *H \ 0޽h ? ̙33  l0(  lx l c $ " *   x l c $ P  H l 0޽h ? ̙33q!  !!!H||0  (  |x | c $ " *   l p P s| p ,$D  0Z2 Q| s * `PZ2 R| s *0#`SZ2 S| s * #Z2 T| s *Z2 U| s * Z2 V| s *N~Z2 W| s *p0fZ2 X| s *c  Z2 Y| s *0`0ZB Z|B s *DfԔp PZ2 [| s *Z2 \| s *Z2 ]| s *Z2 ^| s *&Z2 _| s *@FZ2 `| s *0f` a| 0p Pl  s  x| s ,$D  0ZB  | s *D 0  "| <dg s  @extract features  j| XALarge grid   k| X3fALarge grid l p v|p,$D  0 ;| 0İ0  i9microarray data: n examples with m expression levels each: 2: b| XALarge grid 0@  c| X3fALarge gridp0 ` l| 0p0 l  p || p,$D  0t@  b  i| pZ2 | s *  6ZB |B s *DfԔ b ` | 0 b  $| 0 0  Hclassify held-out sample ZB n| s *D '  l `  w|` ,$D  0 !| 0`L g7Repeat for each of the n examples: leave out one sample8 8 <| 00P 9 test data 2  =| 0d@` : train data 2 ZB >|B s *DP@` e| XALarge grid   f| X3fALarge grid   g| X3fALarge grid  ZB r| s *D@ 0 `l   {| ,$D  0 #| <$   = train learner n@ }  h| 0 pZ2 @| s *@p#Z2 A| s *P  Z2 B| s *  Z2 C| s *@Z2 D| s *m  Z2 F| s *Z2 G| s *  Z2 H| s *PZB I|B s *DfԔ} Z2 J| s *-cZ2 K| s *MZ2 L| s *@vZ2 M| s *}Z2 N| s *0 ` Z2 O| s *0` P| 0} ZB t| s *D p  l 0 Y  z|0 Y ,$D 0 ,| 0p Y  o1apply same feature extraction to left out sample "1  22 m| X3fALarge grid  ZB o|B s *DP ` 0 ` ZB q|B s *DP` ` ZB y| s *D 9  H | 0޽h ? ̙33   ph(  x  c $ " *   x  c $ `Po  `  C 8A D:\image004.gif * `  C 8A D:\image008.gif `= H  0޽h ? ̙33  <(  ~  s * " *  " ~  s *t P  H  0޽h ? ̙33  0(  x  c $ " *   x  c $4 P  H  0޽h ? ̙33*   j(  x  c $ " *   R  s *bbX   C 0AH:\cb\a.png`P H  0޽h ? ̙33*    j(   x   c $t " *   R   s *bbX   C 0A"H:\cb\a.png`PH   0޽h ? ̙33*  0 j(  x  c $ " *   R  s *  X   C 0AH:\cb\a.png0H  0޽h ? ̙33  x@(  x  c $4 " *   X  C 0AH:\cb\a.pngF0LH  0޽h ? ̙33*  Pj(  x  c $ " *   R  s *99X  C 0AH:\cb\a.png0@H  0޽h ? ̙33  `*(  x  c $ " *   r  S T P  H  0޽h ? ̙33  p<(  ~  s * " *   ~  s * P  H  0޽h ? ̙33  0(  x  c $t " *   x  c $ P  H  0޽h ? ̙33  *(  x  c $T " *   r  S  P  H  0޽h ? ̙33j  (  x  c $4 " *   R  s *99R  s *99>  3 A?apH  0޽h ? ̙33  P(  x  c $ " *   R  s *99>  3 A?a@PH  0޽h ? ̙33  `X(  r  S  " *   >  3 A?a4@H  0޽h ? ̙3  .(  r  S T " *   |  C TA<H:\cb\alon_rand_fuzzy_err.png0@H  0޽h ? ̙3 0 y(  ^  S p   s  c $p F   IDNA, select genes, spot Cell, extract RNA, copy to cDNA, anneal, wash of, screen Oligonucleotide-based arrays 25mers spotted on a glass wafer, Affymetrix GeneChip arrays Custom spotted 50-80mers generated from known sequences. cDNA Inserts from cDNA libraries PCR products generated from gene specific or universal primers o x  \ o x \H  0i ? ̙33 0 D(  ^  S p     c $tp F   :&Selection process, filtering importantH  0i ? ̙33Z   0   0(  ^  S p      c $p F   genes = spots Feature selection currently T-test Probs. Assumption violations, repeated& P-value is the probability that we would get a more extreme t-statistic than the observed one strictly by chance Other test statistic suggestions: see related work Parametric Significance Analysis of Microarrays Tibshirani Fisher (x1-x2)2/(s12+s22) square of t-score not considering number of elements Golub x1-x2/(s1+s2) not considering number of elements different weighting for variance Non parametric TNoM (Threshold Number of Misclassification) Ben Dor number of misclassifications done with minimal error boundary Park, Mann Whitney, Wilcoxon rank sum score (rank elements, sum ranks for one group compare to what would be expected) Mututal information Zp B9 ( ( ( (( ((> ( ( ( (E 1 Oa,R Z H  0i ? ̙33  0 `(  ^  S p     c $p F   Vacute myeloid leukemia (AML), acute lymphoblastic leukemia (ALL) 18 colon adenocarcinoma as well as 4 adenomas and paired normal tissue (generally full-thickness colon) from the same patient obtained from the Cooperative Human Tissue Network  ,$ H  0i ? ̙33 0 )(  ^  S p     c $p F    Ask around H  0i ? ̙33< 0 (  ^  S p     c $4p F    make a call for cancer or normal DNA holds information for many genes (region on DNA for one protein) 35000 genes in humans Proteins are components of life Many of the genes are strongly regulated and only transcribed at certain times, environmental conditions and in certain cell types $! H  0i ? ̙33O 0  ((  (^ ( S p    ( c $Tp F   elots of noise in the microarray Given a series of microarrays experiments for a specific tissue under different conditions, find the genes most likely differentially expressed under these conditions. I.e. find the genes, which could be used to explain the effect of the conditions. This is also called feature selection. If the genes are the features describing the cell, the problem is to select the features, which have the biggest impact on describing the result and drop the features, which have little or no effect. Feature Selection is a filtering process, which reduces the total amount of features in a dataset to the ones, which are relevant. The learning task is: Given a set of n examples with m attributes each, the task is to find a subset of the attributes, which is sufficient to classify the n examples with high accuracy. Noisy or irrelevant attributes make the task for the learner more complicated as they can contain random correlation. As they do not provide any useful information it is in our interest to filter out these features. goal not just feature selection but also call for cancer or normalff H ( 0i ? ̙33 0 D<,(  ,^ , S p   6 , c $Tp F   Special cases/applications: more complex decision possible with non-linear kernel (e.g. RBF) if no perfect separation possible can use soft margin find linear decision boundary max margin separating hyperplane softmargin allows for noise complex decision surface can be learned by using a non linear kernel function An SVM expects a training data set with positive and negative examples as an input (i.e. a binary labeled training data set). It then creates a decision boundary (the maximal margin separating hyper plane) between the two groups and selects the most relevant examples involved in the decision process (the so called support vectors). This is always possible as long as the data is linearly separable. If this is not the case SVMs can use  kernels which provide a non-linear mapping in a higher dimensional feature space. If a separating hyper plane in this feature space is found this can correspond to a non-linear decision boundary in the input space. If there is noise or inconsistent data SVM still tries to find a perfectly separating hyper plane and this might lead to over fitting of the training data. Soft-margin SVMs solve this problem by tolerating some training errors. They try to exclude a minimal set of training examples so that they can separate the rest of the examples with maximal margin. SVM classification is a supervised learning technique as it expects a user to have knowledge about all the examples in the training set. It is also only able to make a binary decision (whether the example to be tested is negative or positive). It is possible to embed a multiple group distinction process into several binary steps. But the number of binary trees with n leaves is given by Catalan numbers and therefore the number of possible embeddings is exponentially growing. The main advantages of SVMs are that they are robust to outliers, converge pretty fast and find the optimal decision boundary if the data is separable. Another advantage is that the input space can be mapped into an arbitrary high dimensional working space and the linear decision boundary can be drawn here. This allows for higher order interactions between the examples and can also find correlations between examples. Finally SVMs are very flexible as they allow for a big variety of kernel functions. tw-w !.H , 0i ? ̙33q  0 1)p0(  0^ 0 S p   # 0 c $p F   5k-means clustering (see (Forgy, 1965 MacQueen, 1967)) Cluster analysis is a technique for automatically grouping and finding structures in a dataset. Clustering methods partition the data set into clusters or classes, where similar data are assigned to the same cluster whereas dissimilar data should belong to different clusters. k-means clustering is a technique which tries to minimize the within cluster distance. Obviously three design decisions of this algorithms are: how to set the initial cluster centers, what is the distance function to measure distance between cluster center and element, how is a cluster center assigned (is it the most representative element (then it is called k-median) or is it the mean of all elements (then it is called k-means)). Another big factor is the number of cluster centers. 26$.$ ,  H 0 0i ? ̙33 0 ~8(  8^ 8 S p   x 8 c $p F   Bezdek 1973 Fuzzy clustering deals with the problem, that there is often no sharp boundary between clusters in real applications. Instead of making a hard assignment to one specific cluster for an element we assign a probability (between zero and one) to be in each cluster. In doing so an element can be a member of several clusters. The fuzzy c-means (FCM) algorithm was introduced by J. C. Bezdek 2QA $H 8 0i ? ̙33^ 0 @(  @^ @ S p    @ c $p F   @FCMeans Clustering MATLAB Toolbox V2-0 Regular n-sided polygon .A&  :H @ 0i ? ̙33i  0 )!PH(  H^ H S p    H c $4p F   ]Adenomas are considered the pathological and genetic precursor to adenocarcinoma of the colon^^ AH H 0i ? ̙33 0 jb@T(  T^ T S p   \ T c $Ժp F   pathway: e.g. gene A controls gene B which controls gene C If we pick up gene A with a good test score it is highly likely that we get a good score for gene B as well. This is not a big problem if we would have no limit on the genes we want to extract. But usually biologists are interested in a small set of genes (for financial, personal workload or experimental reasons), which describe the perturbation as well as possible. . If there are several pathways involved in the perturbation but one pathway has the main influence we will probably pull out all genes from this pathway. If we then have a limit for the number of genes we might end up with genes only from this pathway. If many genes are highly correlated we could describe this pathway with fewer genes and reach the same precision. Additionally we could substitute correlated genes from this pathway by genes from other pathways and possibly increase the prediction accuracy.( !H T 0i ? ̙33  0 B:`X(  X^ X S p   4 X c $p F   Find similar genes automatically group similar genes together and dissimilar in different clusters One way to pull out correlated genes would be to calculate the correlation between all genes. Instead, we decided to use clustering algorithms for this task for several reasons. First, clustering is more versatile as it can use different distance functions. Correlation could be one way to calculate a similarity distance, other possible options are Lk metric (where L2 is the Euclidean distance), Mahalanobis distance, etc. Second, clusters can have different underlying models, shapes and densities, which are not captured by just correlation. Third, it is not clear where to draw the boundaries for correlation. Should we say genes are in the same pathway when they have a correlation of 0.85 or is 0.4 enough? Clustering methods naturally put genes in clusters and generate this grouping for us. K-means does a fixed assignment of one gene into one cluster. But a gene may be involved in several pathways. Fuzzy clustering methods solve this problem as it can assign a membership probability for a cluster. If a gene has a similar expression profiles to genes in one pathway and in another it would get a membership probability for both clusters. y^! ( (7 y!\ ,- H X 0i ? ̙33 0 `^(  `^ ` S p    ` c $4p F   TILP NP complete, so used a builtin non linear zero finding algo why at least one? don t wanna miss small clusters with good discriminative power pseudocounts, get at least some information about a cluster if cluster has very low quality score, I.e. very tight we might loose it drawback, might include totally unrelated pathways HS-5-M!P3 H ` 0i ? ̙33W 0 p(  p^ p S p     p c $p F   WNow we know how many representatives to choose from a cluster the remaining question is which representatives from each cluster to choose. There are several options. We could choose genes closest to the center or genes farthest from the center or genes with the best discriminative power (best test-statistic) or do a sampling of the elements in the cluster according to the probability function for that cluster or choose some from the center, some from the border, some from the fuzzy border. We decided to select the genes with the best test-statistic as these yielded good preliminary results. *VV!H p 0i ? ̙33M 0  (  ^  S p     c $p F   SWant to have a way to compare different gene subsets. LOOCV classifiers learn on n-1 samples and tests on the remaining one. I.e. we take n-1 samples, extract the most revealing genes for samples and use a machine learner to build a classifier for these samples. 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&՜.+,D՜.+,8    oOn-screen ShowUWs$!A*j 5Times New RomanVerdana StarBatsArialComic Sans MSArial Unicode MS WingdingsCUTEMicrosoft Excel WorksheetMicrosoft Equation 3.0!Microsoft Photo Editor 3.0 PhotoNo Slide Title MotivationMajority voteOutlineBiological task MicroarraysOutlineFinding informative genesOutlineCurrent solutionProblem with current solutionProblem of redundancyOutlineProposed solutionHard clustering k-meansSoft - Fuzzy ClusteringFuzzy examplesFuzzy softness 1.3Fuzzy softness 1.25Fuzzy softness 1.2Fuzzy softness 1.15Fuzzy softness 1.05Selecting genes from clustersHow many genes per cluster?Which genes to pick?Comparison EvaluationSupport Vector machinesOutlineExperimental setupResultsfuzzy c-means vs k-meansDifferent test-statisticsComparing best resultsHow about randomly choosing? 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