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<pory{|4zTt~,po}V~q4zTr~:V~S}qT}F~$}uFrt\p}rvru<}qTrv~uh~:uzKrtrtyT~S}Shrt4aptpshprt|4's}t}~:T~SurvVK}rtq4|~}0}~Sq4zK~,rtT}y,~SV|4rts|]u{haK|]~SpFqa~S' }
TThh~S~S{<MK~z]~S<vFprt}T~Ssh44o~ZF|Tv4hsh40 h}q4~xty r|4 '}@rtvu4T uryv}h rt&} qT~$~]po~rTrv~x}t~,]}rq]|}8}&}q4}q4~x~:}~,hy}qT|Ts|T~S~
Mf~sh]t{}q4~hx~Fvt~xTTv}~:of h4~'}sht'&v}q4~SByts4 |4y,]~SBrtuZps|]}~F
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i"SB9aN9JCFG89<CYg2SbM?CFCF=HC@BADxy9J>EH9<CF;J9W\9JQI"9<9JCQI"=B>=H\bMH\BAD][AQm@BADGeQ8>AD\FXYQA[=HCFG
ADG%ENADxy9<C\9 

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Information(Clusters,Response)/
Information(Stimulus,Response)

0.8

Fly 1

0.6

Fly 2

0.4
Fly 3

0.2

0 0
10

<|4~&4

>JZ bHF<P

10

10
# of Clusters

b<<PYZ?H@? AZ<B b

10

CEDYfYGF(PY

"i"SF9ADCBOP=H>K/MfQA[=HCQSFM?Q

QSF9%;<]DXBGeQ9J>G;<=HC2xy9JmdMH\=NXYQQSB9%CF9<XY> MH]F>9<G8=HCFG89CF=?>K/MH]DAD^<9_@a\nmWQSB9%A[CYO=?>K/M?QAD=NCaQSFM?Q QSF9kOXB][]


G89JQ=HO

8Z;<=NC2xy9 mBG M?\=NXBQ QSF9kGeQADKWXF]DXFG cNA[GG8SF=_I%CM?G"MLOXBCF;JQAD=HCa=HOQSF9kC2XBKd\9J>(=?O;<]DXFGeQ9J>G

QSbMfQ%QSF9M?][EH=H>AQSFK

XFG89JGG89J9Q9J2Q:OP=H>L@B9JQ M?AD][G h 9LSbM_xy9XFG89<@5HHNdQADK9=NADCnQG\FADCFG Ol>=NK

QSF9:9JY9J>ADK9<CnQMHGQSB9%A[CBAQAlM?]G89JQ"=?OIH

R KJyh(i"SF9%QADK9%=NADCnQGI"9J>9:; SF=NG89JC/> M?Cb@Y=NK]mdOl>=NK

QSF99 B9 >A[K9JC2Q_wF@BAML9J>9JC2Q:; SF=HA[;J9<GL=?OjQSF9QADK9=NADCnQGUEHADxy91G8A[KAD][M?>:>9<G8XB]DQGCF=?Q:G8SF=_I%C h%i"SB9


ADCBOP=H>K/M?QAD=HC;JXB>xy9kADG G8SF=_I%CO=H>N@BAML9J>9<CnQ `bAD9<G:GMHK9kQADK9k=NADCnQGXBG89_@dO=?>kM?][]BQSF9"`bAD9<G hPO]m
VM?Cb@SFMfxN9MWG8ADKAD][M?>kB>A[CBEW> MfQ9aMHCF@ADCBOP=H>K/MfQA[=HC> MfQ9<G cBI%SB9J>9_M?G"`Fm

NWSFMHG">=HXFENSB]mQzI%AD;<9

MHG%SBA[EHS|F>ADCBEd> M?Q9M?Cb@ADCYO=H>K/MfQAD=NC|> MfQ9<G h

p|4tzt}|T~S}~:hm4y{&r}qT}@v}rtu}qT}q4~~ |44yurtKy{~xvrv}umhrt&}h}yqT|4}@sh|T}fq4u~x~:u|4}sh|4'~:z4ht}q]~}}t]m~q4]rposhr4<~SvBKhrtB|4}8~S}shvqT}~xht~S}shy|4ysh|Tv sh2


z4rt4 r~S}qT~~o~$4tv~$y{zKv~Sz4 psh)~t]p}r ]posho|T }~rvFu}q4~$qTrthpFhq^thTvsZh~:urt|Typ}}r"r-uMp}zTrt~:rv~Su~SrtF~o~4Fp}rvu&vysh|4v~tT4~}zK|T~T~ 4
44fK4! V <4 "  4 ! ! T
}rtqTh~0trthTv|4y{}~0~pspo|Tush|Trt}y~S}~hrt4x4qT~pot|Tr~TT}h~y|Tshh'psvh}4Brm~SmzKv}shrtTrv4uB~|4}}q4hrt~m}}Thh}4y{q4|~S~S}rv}Tshq4rt~T\u8}t~q4sFT~v}q4~0}~:y,~|44}s|]hh4}|}}qTTv|r}&}ahrtKT~~SshBrtpoT~:\hv]sh}~:r
}rtqTT~,~4rtru@h}q4~~&y{Trh~SF~Ss<4rtR uf~t}q4}~qT~$4~S|4t~rFvm&~@@ht|T~ \rtu<q4~:rtpFq-}rvq4u@~}q4p~$~'p}s|Trt}4~S0Srv uf}q4~{\posh|T}~FDurt
}qT~~%~\T~v~Fs}vrshrvSut4`rtF}%q4qT~v4}-hz4}t~~T|}M}rtq4hrt~%Txp rtry{R] y{~\rt}^}hty{u~S~:~v}$~S|4*~Sq4\thpFrv~Sq*utt}tq4~~~`}h}yhy|4|44sh|Tsh~S|T%-}r~SBt}y{q4~\~ ~'p}vt\y{shpz4~|4s~Sppos}|T~:~ }}h~S4qTS~
Vtq4~S~:t~Vt~ tr~Stt~t~}q4}~`hy|4}hshy |4sh|T~ourt~yy ~S'p}s{|]z4}~~S@p}~Sh4t~~S}q4~a}~hyFv~:{~ }]q]ur}\@po~:sht|TpFq,}~Srvu~:}"qT~V}rtps|]~o}qT}~~F:
q4r urtV]`vT }q4~$p~rvuposh|T}~FV-}q4~s~uD}zTvT~smrtu<<|4~]
<~Sz]|4~:~0}h4x}~q4~Vprty{ty{zTt~xv~:]v}shq4 ~0!psu|]rt@}}~q4~4rv}q4~S~f|4s}TBhrv~Su!<}q4q4~r'~BTh~SSurTyrtvB}}hrtq4~ppS|4t~~rvu8@q4psr|]}$~hF
u rt|4<y~SKpF~qrtrvuKu4}qTq4~xvT~S&~:}y{~B~:M~'T}pu~$}pq4rt~T~:h~SzK<rtVT4}pF4qT0~Sposh}|Tq4~}~Ft@y{m~xry{4r~S @~q4<pFVq$q4qT~pt~~B'}}q4rt~x$shtrt~SF }
(

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k!

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z.

f

H

b
1

0
1

0
0.2

0
0.2

Probability

0
1

0
0.2

0
0.2

0
0.2
0
0

0
0.4

<|4~$4

20
40
60
80
100
Binary Words (ordered by # of spikes)
UB2fWVb

XW 

0
0

120

20
40
60
80
100
Binary Words (ordered by # of spikes)

b|ZYZ[?N.A<<2?F\bZYZ]FjPY a 1

xy9J> M?EN9

120

R OP=H>

9_M?; S=?OjQSF9;J][XBGeQ9J>G:=HOj`FmVHcbADG%G8SF=_I%CO=?>:QSB9;_MHG891=?OW;<]DXFGeQ9J>G<h =?> @BG:M?>9=?> @B9J>9<@|\nmQSB9


C2XFKW\9J>%=HO G8FADy9JGwFxy9J>8QAD;_MH]Z\B>=NN9<C]DADCF9<GUK/M?>QSF91\=NXBCb@BM?>8m\9JQzI9<9<CI"=?> @BG%I%AQS@BAML9J>9<CnQ
C2XFKW\9J>%=HOG8FADy9<GG8SF=_I%CADCQSF9Q=N5FMHCB9<]l hd

xy9J> M?EN9

R OP=H>L`BmN2hW91CF=HQ9QSFM?Q

QSF9">9JG8XF]QADCFE@BADGeQ8>AD\FXYQA[=HCFGM?>9%CF=?Q xy9J>8mWG89<CFG8AQADxy9kQ=1@BAML9J>9<CnQ(> MHCF@B=HKA[^J9_@d; SF=NAD;<9:=HOQSB9:HHN


QADK9=NADCnQG<h

q4r}}qah]4z]|r}rTxhqpqT}0zTht4v~t4sh,~S~$s8hhzTy{`<s~vx~o|4}tpt~S~ 4oT~\F4v|4qTsK}r}|TKpo~S}}qT|4~h-~,posh}2|Tq4$~}s~v~ot}}F~vhps st|]~Srt}u~:~-}q4rv~:um~$~4Frvz4~S}h rv4Tu|4T}}hqT rt~,vT4,{rt}T4q4hrt~S~S}


uh rtB}q4~$prt~}~:vhzKsrt T44posh|T}~FSv~T~Spo}h4&hy{svf~4~Sts }|Tpo}|4~t4|4}sht~!~~Tp~
Z
por~hqT4~Vrvu&urtpo|4rTyzT}sqT~v}Su rtTv}y{q4~p]}y,|4s4~h|T{rq4mhttqT}sq4"~0p4rthpoy{}hz4rtT~Sth4}sh~tqT
v}wxrt~:p~:h]zK~:rt]}q4T~0t4sht~B|Tt s
y}h|]y}&|4sh]|T~&~z]sht~Spopo~}ht~$urr~Sr t~y:Kvsh<q4 sh|4~,y,}]q4~S~{&4rv~Sux|4}thsfypo|4rsh|T~S0u~Srtvux}|4K~S~S~Sha}}q4T~{h~S&q4yq%hy{K~~ST$hrtportT ts
h~St4sha}q4~h&T4F}~Sv]`hurty}r%F}~SSm}qT~porty{z4~S4hs }$vshy{r}
^



20

stimulus velocity (/sec)

Probability

10
0

10
20
30
40

<|4~T

50
100

80

60

40

20

time (ms)
OF=?>"9_M?; S/=HOQSB9;<]DXFGeQ9 >G"ADCUOA[EYheNNMYc
a`WbA<<2Zc<?bVVb2fY<_]AbAdB2fWVbY e

I"9/;<=HKFXYQ9dQSF9/M_xy9J> M?EN9/GeQADKdXB][XBGIkM_xy9JOP=H>KI%SFAD; SB>9<;J9_@B9_@MH]D]QSF9aQADK9<GQSFM?Q;<]DXFGeQ9J>9<@


Q=NEH9JQSF9 >]D9JOlQ1bM?CF9<]l h

f=HKbM?>ADG8=HC=HOQSF9dI"MfxN9JO=?>KG=HO:/;<=H>8>9<G8=HCb@YA[CBE;<]DXFGeQ9 >G=?O"QSF9 N

`bAD9<G1>ADENSnQ:bM?CF9<]l hjMfxN9JO=?>KG"I"9J>91]D=_I%bMHG8G%b]Q9J>9<@O=?>LY>9<G89<CnQ M?QAD=HC;<][M?>AQzmnh

Ku~S~ov|T}}4|4~~~S~ F4}tqT~osm}~&ut|4u~Ss]Sp }}|4hrt~SVMrv<u@hT}q4~$t4!<~T~'t}shtT!h~SS~$4T|]}}}qTq]}}}}q4q4~&~T~h~S&rporTK~Sh~S-t}4q4sh~$~$rzKt~S~SpsvKz p


4R|4 ~t}|4~tTrt  q4r |4vshT{u|Tprt}y{q4zT~v~o~}z4q4shrt~~~:}|4qTs~}fFvurt4B~rv!u8~}~h'y}B|4sh}|Ty,xzT|4vsh]Fvv]y{,~}~~SFzKtrtTT{~~:~StzKy{rt]~'~:}:
}qTus|T~srT8}r]y,|4rt|4$}s}v|]sh~]$TvV~Ssh pF\t&~oy{urrv~uB}yq4}rq4{-~\}$v}h}yq4rt~S|4|4Vshs|T%}&qTttush~:\r}}|4q4]~$~\~:vrvuVtqTsh'rtrt}|4y{~Ss~`~:Fh}&T}p}sra|T~4v~z4~Fz4tshrhartmK}~rt}rq4T`z4}q4rt~Ty{s~S~Sy t}qTrv~ u
}vqT`4~S~~~y{y{z4~Sh|4q'ps}}ts<ry{'|4~S~:poq'}}}}r4qTmrt}}]~~-}K z4~shpv~Svts!~,hrt4}u@qT~t'v~S}}0T}r~~S|440Ft4}z4~S~|4z4rrtr"t~:pFqhzK^~S}prvht}sq4h~, tpop~Srrty{}}~hz4pS4vF~Ss!}po~trh4h~:tsT}"VrvqTq4uf~T}qTph~~
4hpor~S}|4hr4tas@|]ppor ~S4p }~$r%Thsv8vp %rt-'}qTth4}zK|~}~y~T!}rvpoTrtv]Tx~$~~SFq4vhr4Tz],sa~}q4}rqT~:|4v~}''}|4qT|4~:~~:}po}rrryTT$zT|v}Kvrt}|4hrt}T}tq4s8~\y{~o}}q4|Trpo}4|4~~$rvuVqT}qT~~
8
o$h~!~tfD \B4 tsvTmv ,fh:Bt ~ ]a|4 '}B~~:vSTwn v}y~T~4hTtpF~{!:t vj hts~SK !to
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}SttTuo~S 0h%q4r}zT~~S'|4~STpo~: :: ZS: vKtv ! A v ']
$artTtST ~!m Mo<! F )n ~Z:4vr !':Rv]&4 K]TS~shv 4hs }-rvufz4ht~$}hyh4h4~Sr prt}ptsZ4~|
v$rtVu8v}h:qT4~]~S qT &4r{pFqmy!tw0pStK'8|4~S~&y{y{z]rrt4tsm~z4~Spor- rtu z4hKt~$}FRMvhThZa~oS } Ftv}']}~Sprtt}o~o
$t4~ShshpFq}tTn!h }y{ph}4rt-z4] ,~:po h&rt4rtuBq'}4~S|4 rThvs:z4v <~$&} tT0h`vz4 sv`rTh Z~Sz]rT T~, vMh
ZTtt TtK:t t
tn'T}\r4hTu'rn!yh} r& r]h~S4sh~t~'|Ttsm'z4~ht~|T}}t~TSt\n'}o~St~TT pF!'oR Zh:t s~S4Kv\B:t}rz
:$VBrq po ~St~S]h$4h'K]rt~Su ~STF44SDh )]TThtvs ~S0K]D]vVq4~,vSourtKyFV}rFrt},}sh~]TM~SpFv 2Mav4~om}FqTr TmR]vh
vK\Fv !2D T4ht~F}rtu<shh4rhSto
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$0D]v|T~:m {f Vq4~:-~srs4R|4Ts*v poKrtyv zTs~v"T'rvu0Bs}~Sq4'~-|TyTB:ht v 4 F KRMK:M D
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