Adaptivity of Sensory Representations in the Adult Visual Cortex

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(1)Adaptivity of Sensory Representations in the Adult Visual Cortex. vorgelegt von Diplom-Informatiker. Lars Schwabe aus Berlin. von der Fakultät IV - Elektrotechnik und Informatik der Technischen Universität Berlin zur Erlangung des akademischen Grades. Doktor der Naturwissenschaften - Dr. rer. nat. genehmigte Dissertation. Promotionsausschuss: Vorsitzender: Prof. Dr.-Ing. Olaf Hellwich Berichter: Prof. Dr. rer. nat. Klaus Obermayer Berichter. Prof. Dr. Alessandra Angelucci Tag der wissenschaftlichen Aussprache: 16.12.2005. Berlin 2005 D 83.

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(180) =;<;KaV= ? =V5 ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ g~ >? bKqOaVb6b ? 3#= ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ C 6 89'(:;.=<?>@? <./#ABCD%'(#./%E0#1203F 6G CV~ EG=%;6? 4K3I? eIacq ; 3#=~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ C CV~ ) = BL

(181) 8tB13IeI798 ~`N ~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ CV~ C ;>79? qOag4K467O? =;=g7O; {\3%4 B 3IeI7O8~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ CV~ bKqOaVb6b 3#= ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ H+IJ./KMLON0BP? 1QBR%E$, "N;KS'(<>@ >@'-<$ GET c~ EG=%;64K3IeIacq ; 3#=~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ VU c~ s<rg7`h ? bKrg7O4 ? =IHJ3%4KBLM; ? 3%= 3s6 79q ; ? 2#7 HJag=Vq ; ? 3#= ~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ 7v c~ C ! A ; ? BL

(182) 8 qkrVL#=g5#7ub\3#HHJ3%ag4 bK7O8p79q ;679e B 79qkrVL#= ? bKBb ~ ~`~ ~`~`~ ~`~ ~`~ ~ 7 C c~ >? bKqOaVb6b ? 3#= ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~X7W C GZY[.=$\]^'(KS'-'-? J$#'-$#E#NA_'(K'($#'-./#$ ÈT g~ EG=%;64K3IeIacq ; 3#=~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~XWU g~ h ? 4 ? =g5 4kLM;67 B 3IeI798 L

(183) =Vef8p79L

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(185) Ht;6rg7 " 4 ? =V514kLM;67 B13IeI798~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~ba g~ >? bKqOaVb6b ? 3#= ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~baa I+cd+ e1E\? e]^cd:<'(N@.=f$N./Kg$'F>@'(< Te + ~ =V5#? 8p7 qO7O8p8 B 3IeI7O8 ~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~bU UU + ~~ C EGz 3#3 = ? bKq 7 q B ag463I4KeI7979=8`;6b ~`~~ ~`~`~~ ~`~`~~ ~`~`~`~`~~ ~`~`~~ ~`~`~~ ~`~`~~ ~`~`~`~`~~ ~`~`~~ ~`~`~~ ~`~`~`~`~~ ~`~`~~ ~`~`~~ ~b b ~ + ~ /h ? 4 ? =g5 4kLM;67@HJaV=Vq ; ? 3%=Vb ~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~bDU C + ~ =VL A bK7 B 3 eg7O8 ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~bUDC ?p?:.

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(188) b<4k3

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(190) 8=g? 7OA ; {\3%4 B 3IeI7O8 ~`~`~ ~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~ ~`~`~ ~`~ ~`~U U ~~ C L

(191) , 4k79L

(192) 4KB 79=%7O; ;K7O=4 ? x9agLM;;1? 3%~`= ~ 3#~`H ~b A ~`LM; ~ ? L#~`8 ~`bKq9~ L

(193) 8p~`79b~ L

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(196) ;kB b<793

(197) 4|H C; ;KrgL

(198) 7`3 h c? 3%bKrgag79=V4 e+? =g~`HJ3#~`46B ~ L

(199) ~`; ? ~ 3#= ~`~~ ~`~`~~ ~`~`~`~`~~ ~`~`~~ ~`~`~~ ~`~`~`~`~~ ~`~`~~ ~`~`~~ 7 g<%E1E./K"N T.

(200)

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(202) 8:7u=]eI5#3#7 465%HJL

(203) 3#= 41bK8pB.L

(204) ;K79q 4D3%=%7; A = 8:? ag3 7? ;kLM;63; ? 3#7= C s< Ur U 7bw H L#? bB1LD3%aV4kb<LM;6L

(205) rg=V7Oe 4r{\? 795%8:rIC C8p7O=V2%3M7O{8 =A 463 V"8:!$79#&B %')(+46*,7O#)8p-/7O2M.01L

(206) =;<!$#)HJ3#*2(34 ')aV(+*,=V#)eI-7O(44k%b|'6;6L

(207) 57=c8#:e 9 ? =V5DN`;6rgL

(208) 7O7 8 g 8 7OrV=VL5R2 7 ? ; 3#4L#83

(209) ~ H ? 3%8:3%5 ? qOL

(210) 8w3#465%L#= ? b|Bb L#=Ve!HJ3%4 ga ? 8 e ? =g5 ? =;K798: ? 5%7O=;1L#4|; ?%" q ? L

(211) 8<L

(212) 5#79=;6b9~EFH@L#= L#5#7O=; =V3#;K7Org2#797944rV7L

(213) A=c8:e 3% 4K;K7u3 b 31? ;6BRb@aV7O=qkrf2 ? 7463# =gA B8p3#794kLM=%;; ? 3%? =; { { ? ? 8p8p8 8L

(214) =g8 79bK2#3R79=g43#8p;79L

(215) A 46L =" bK3-3#, B 7 c;67ur qO? =VL

(216) ac5 b|L7 ? 3#= aI;K; r ?? ;ub ~rq9@!L#bK= 7 ;6;6rgrg77 LrV#5#3M79{\=%79;w2#b|7O794 8p eIrV3%L#B b 7L 8:A 3M8:{<3 ?CF;k8:b792#;679rg8 7@qOL%3#q ag1 =a ;K?794K4 7ue A L

(217) N 4K=g; 3M {{8p79r eg? qk5#r 7<?;K="3R;63 agI46;k= L ? =? b 4Kr 79?{w5%L#rg468 egb94K~79s<8:79r2 L#? b =A;@4KHJ33%g4 8p7OaVB =IC egeg7O7O4k2 b ? ;kqOL

(218) 79=Vb e 3

(219) ? H =g5? =;6 7O? 8p3%8 8:? 3%5#579=%? qO;`L#8L

(220) 4KbK; 7O?#=V" bKq 3#? L#4 8 L#b 5# 7Ob =;67O;kBbO~ b s<L

(221) rg=V7e bK3

(222) 3#CGHoqOqL#3%8:pag7946e;bK7 " !$'=HJ3%<>(44 *,? g*2a(4@&? 8.3e <>(3? =g'75A *2!B;6*2rg(+@C7 (4b|%79'6=V57b 8? =g#:9 5 ?=y ;K4K7934K=VbKb L# 8 7Ob|46795 =V bK3#U(4 W 7 4KT7 46A 7 4KHJ7u79b|4679b=;K;63DLM; ;K? rg13%7= A 3#46H3 ;6gA rg8p7`7OB 3#aI3#;kHtb ? ? eIeI797 q ?{we 3#? =g468p51e{~ rg7O= L

(223) =Ve rg3M{X;K31L#egL A ;oA ;6rg7 e7gg aVL

(224) q B@!? g=A8p7"8p7 ;Kb| MrV79;67f=Vrgb|7 3%3%=geI4 7 79q rV46? b7L

(225) ? A=c3#46e= 79 wbK 7ObK7 =a

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(227) 3I; q ?A 7u3#7ObK=Rb4KHJL#=V3#=V465]eDB b|7? ;6=V L#5A 5#8p793;6b1rg? ;6746LM?7O;48 ? 3%q = 3%3#~{B =X@! A b =DaI;k;6L;KL

(228) grV; 8p7w?7 3#3#=VL

(229) ;Kb1=Vrge798 ?:4N 46rV7 7 L

(230) =cHJ463%e3

(231) 4C bK7O=cb|3%4 467 A 4679bK7O=;6L

(232) ; ? 3#=Vb=g7O7ue;K3 7 A 8 L#b|; ? q ? = 3%46eg7O4;63D79=VbKag4K7 L r ? 5#r " eI798 ? ; 4K7 A C. 6HJ4 3%794KbK797OrV=;kL

(233) LM=c; e? 3#~T=1EG= 3

(234) H3#bK;K7Org=V79bK43#{\4 3%46b|eg; ? b BRb|ag798 =V? {b|3%rg4 3b|467 7 A 7O4KrV7ub|L 792 =%? ;k3#LM4k;L

(235) ? 8 3%46=V7Obo8p7O=V2M7OL

(236) 79=Ve qO7<;K3 B L 7`L#=geg3

(237) L ; A ; 7? 2#N 7%=g~ 3MEG{=ceI= 7Oc7ue7OC L3%%="egL BRA ;6aVLM8i;; ??:3%A = 8p7 ? ; b? B L]7O{CFb6?qOeIL#798:7ub AbO~4679EGL#= e ;6rgA 7Drg79qO=g3#3#=B ;K77O=V; 3#3

(238) = HT? ;K=XrV7R=g79794KL#2%4K3#8 aVb 2 ? b b|acb L

(239) ;687OBb b b w;67OL

(240) B =Ve ;KrV? ;7DL#rVq L ; AV? 2A ? 79; =VbC e8gL#7 b|A ;6bw7O=V{weI7O797 =N ; b 4K7 A " 79=g46qO7O7B A 79;6=%aV; L

(241) 83#8pHw79L

(242) qO463#= 4K;? =g? qO5RL

(243) 8T3Iq9Bq agL A4kb\b egrVagL 4 Ag? =gA 517O=crgb`3#eIaVag46bw4 ? L

(244) =g=V5 e L egqL 4 ? b; ?qO8 L

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(246) ; ? 3#= ? =;Krg7 A 4 ? B? L#4 2 ? bKaVL

(247) A 8T?q 3%4|;6" 7 Jl A A {\3%4 ? N b 3%3%=1= ;K;Krgrg77 3%3#464keIeI797O4T4`3

(248) 3#HH<LLfHJ79HJ{ 79{ bK79;6qO7O3#=V=VbReVb3##4 L#r =VageR=c;KeIrg4K77ue]H L%bB ;? e 8p8 ? b|=V7uL#q B3%=Vq9egbTb`3

(249) BH4KL 7uq 7 L#8pbK; 3 2#7c7 7O2 8 e? 7O{w46793 e L%7O4Kb ; L

(250) 79= b L%egL sA ;3#? 5%2#7 7 ;6rgA 7O4643 qO{ 79b6? ;KbOr ~ ;KrV7L AgA L#4K79=; 5%7O=g7946L#8TL AgA 8 ? q9L ? 8 ? ; 3#H\;6rg7 ? eI79Lf3

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(252) =5%;wag7O3

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(255) H ? =fe ;Krg=V7L

(256) B 2 ?? b|q ac46L

(257) 798 q q7 3%A 4|; ;6? 2#7 7 " L%7Ob8 eb|3%A B 4K73 A b|793%4|4|; ;? 7u3

(258) boH3%L#4\eV79L 2#A 7O;k=fLM; LM? 3#;K=;K7O =; ? 3%8:7u=VL

(259) L

(260) 468c8 ;K3gAagCF=Vegeg3M7O{4kb=;kL

(261) B =V3 e eg? =gaI5 C LaV%eg8pL

(262) L ; A ? 3#;6=LM; ?H 3%L#=8: b 3%;6=g3 8 3 ? bK= rg3#;Krg4K;97 ~ =Vs<L#rg4K467O3M46{7 HJ3%b|4K797 =VVb|HJ7 463#3

(263) BH LR=g=g3M79{ ag463#3#==VL#{\8c7DH L

(264) aV; b|? 5#7Rag;K7 rgLM7RH};6;K7O7944KB A 463#& L#8peg3#=gL A5%;79?e2 ? b|; ; ? B13

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(267) 4679;{ bK7O2? eg=L#7O;64 4DL

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(275) ag;78:; {\L#bR7 > L"bK4k7OL

(276) 8b|5#eIaV3 3%4 ? B L

(277) 4 =cb|b|79e77 ;KgagrgaI4 7R;D {\ ;Krg 3%7 <4K8C e2 ~;6b|rgacs<7L

(278) r 8ob6? b@L

(279) b B B1b 7`;67u7OL

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(282) H ;6rg7 bKL#B172 ? bKaVL

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(284) hV= ;3#;K4 -3 rV,77bI? b|L#eI7uB 7 q 3%A A =V8:3%7eDb ? ; ; ? B13%7 ?=f7=g 5 L# 8i;6;K797O3%q94k8pL

(285) ebwaV;KbK;6rg7rV7`LM{w;{<7 ;6L L#rg q 7;6rgab6q7 463#79q4 e13%? =g4K=g546797O{b3

(286) A H3%L =V=ge3M5#{?3%=gL#8:5 C8 ?? A A ? 46?b6=7q ;K79A 7O=g464 797 A bK467Ob7 =;k;kLM=LM; ;;K? ?7O3#3#4 =t= ~t463#7hgHT;63%7924 e? 7 b|~ gacL

(287) $ L

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(291) r]718%eb|qka

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(294) q ; ;K? ac3#L

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(300) L

(301) 46B = A? =g8p795 b A rV{L#r bK?7#qk~ r eI7OB 3#=cb ;646L

(302) ;K7;6rVLM;1;KrV7 2L#=V? b|e aVL# K8\L#eVqOL3#A4K;K; 7? 2%!7 q 3%{<=gL =g 7uq ;K;6rVbDL#=fL

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(306) A ?

(307) i

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(309) 79;6b|rg; ? 7O5%4L

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(314) =g5%79e ? =? ;Krg7 " 4kb ; A 8pL%q 7A L%b q 3%B A L

(315) ? 4679e ;K3 ? L

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(318) ;6L

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(401) 3

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