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!j |"k l"pjlij! yli lij sj l!jkpliv ri sjs"!x llj i k" klj!k
(+ S'$$.+% ; & 7 3. ;7OO %.&&'$ P Q'[.+% )+/ D &-. %/ ; 0 7OO .Q ; S'$$.+% ; & 7 3. ;7OO 0/$ P (+ 7;
&'/$ '$ S-'0- &-. D /%& 7 0&'6. $.(+/$% %( RR +.%% &-. 7 0&'6'&'.% /) 7OO /&-.+ $.(+/$%
6' 7 7 $ '$-'V'&/+2 V 7 03 P +/($Q X%.. S/$ .+67 3%'4 >EE YT
= . O) ;= (%& 7 '$.Q ,-/( P -& 1+/0.%%.%
1
(0)
(1)
(2)
(3)
(4)
(5)
0.8
xi 0.6
0.4
0.2
0
(6)
(7)
(8)
(9)
(10)
(11)
0
250
(4,8,10)
(8,11)
500
(6,7,11)
1000
750
1
0.8
xi 0.6
(2,6)
0.4
(4,5,6,10) (4,8,10) (8,11)
(6,7,11)
(0,1,2)
(1,3,9)
(4,5,6,10)
0.2
0
0
50
100
150
200
250
300
350
N N ,-. 7 0&'6'&2 A X Y )/+ &-. &-/( P -& R +/0.%% X ?Y XA ? >@Y XA C >@Y XC >>Y X? >>Y X@ > Y X> B E Y XA ? >@Y )/+ &-. > ; %'&. $.&S/+3 '$ 8'P T
T ,/ R < /&&/ D Z 5%'$ P &-. R7 + 7D .&.+ %.&% X7 Y < XVY /) ,7 V O. >T /&. &-. Q'[.+.$& %0 7O'$ P
)/+ &-. +.% R .0&'6. &'D . ;7: '%
l!rl!j l l lij !jk"k "m |kri "m jsklr k"j yli k" l"k
!jl"k k |jlyjjk lijs v js"!x llj r"!!jz"kkp l" |""pr !jjtkl
"| jrl y i"yjtj! j i|l k pjkj! ij!!rir l!rl!j jjki|j! 4 "pp" &''' j j! v 3 jl ptj k j szj sjs"!x llj
jk"lkp n|"xq sx kt"tj p!"zkp "m r"!!jz"kkp l" mrj  i!l 
zkl  jp  !j  p!jjk k " "kv hi sjs"!x llj yj j€kj k
"! s"j yijkjtj! lij!j !j z"ltj ko + k |jlyjjk "m lijs v
hij!j k"y lij kjj m"! l"k km"!sl"k oj n!jq lij r""! "m
lij i!l "! "m lij l!"j! hil  lij!j lij kjj l" n|kq lij r""! !j
z!jmj!jklx l" "kj "m lij ly" zjrj "m r"lijv }l z"|j l" jkr"j li k~
lj!k km"!sl"k kl" lij sjs"!x llj n|"xq mrji!lzkljp!jp!jjkvv
|x zz!"z!lj s"l"k "m lij klj!k r"kkjrl"kv }k "!j! l" jkr"j m"!
klkrj lil !j lij r""! "m lij i!l "kj jl lij ko !j ~ i!l l" |j
sri l!"kpj! lik lij ko !j ~ zkl "! !j ~ jp v hi zj!mjrlx z"~
|j k k li yx lij |kkp "m !j l" i!l rijtjv hij l!rl!j
"m lij sjs"!x llj j€kj ij!j m"! lij lij!jm"!j uj |j jk"pi l"
"y m"! klj!k ij!!rir "| jrl !jz!jjkll"kv
" r"km"k !jp!kp lij r""! "m lij i!l k "m lij zkl !j k lij
|"tj j szj yijk t!|j ko~l!jkpli + !j jv "lj i"yjtj! lil
li z"|j "kx |jrj s k kjpltj ko + !j k"l "yj k "!
s"j ojx {j!jkrj l" lij s"l r"ss"kx j kj!~kjly"!o s"jv }m
yjo ki|l"!x ko y" |j z!jjkl lij |"k!x "m sjs"!x llj r" k"l
|j j€kj z!jrjxv hij!j y" |j k" lltj {j!jkrj k |jlyjjk s
kjpltj k s z"ltj xkzkp l!jkpliv !lij!s"!j lij l|lx
r"kl"k % v ( y" |!jo "ykv
C
O7 (Q'(% E+/%
[t=27]
[t=297]
(6)
(5)
(7)
(4)
(8)
(2)
(10)
(1)
(7)
(4)
(9)
(3)
(9)
(3)
(2)
(11)
(10)
(1)
(7)
(9)
(3)
(2)
(10)
(1)
(2)
(7)
(4)
(9)
(3)
(10)
(11)
(8)
(9)
(3)
(2)
(10)
(1)
(11)
(0)
(11)
(0)
[t=1099]
(5)
(6)
(5)
(8)
(1)
(8)
(0)
(4)
(7)
(4)
(11)
[t=821]
(6)
(5)
(6)
(5)
(8)
(0)
[t=629]
[t=539]
(6)
(5)
(0)
(6)
(7)
(4)
(8)
(9)
(3)
(2)
(10)
(1)
(11)
(0)
N N ,-. &-/( P -& R +/0.%% X ?Y XA ? >@Y XA C >@Y XC >>Y X? >>Y X@ > Y /) 7 > ; %'&. $.&S/+3 S'&- ; 0.$&.+4 B ; 0.$&.+ 7 $Q /$. A ; 0.$&.+ D . D /+2 %& 7 &.T
,-. $/$ ; \.+/ . : 0'& 7 &/+2 O'$3% ; ? @ Q'[.+ )+/ D &-. ($')/+ D O.6. O ; + 7 $Q/ DO2 V2 7 &
D /%& T / DR7 +. 8'P T B )/+ &-.> &'D . ; .6/ O (&'/$ /) &-. 6 7 +' 7 V O.%
` gf ce a bcf }k p v k pv yj z!jjkl k l"k"s" li"pil z!"rj ylik & ~rjklj!
kjly"!o yli &) l|j sjs"!x llj l!lj k pv v wj itj ri"jk s kjly"!o ij!j l" r lij z!"zj!lj "m lij xksr li"pil z!"rj k
jlv hij s"j i"yjtj! r"szjljx r|j k yj itj zj!m"!sj s~
l"k "m kjly"!o r"klkkp jtj! li"k "m lj yli"l kx z!"|js "k
lk! r"szlj!v hij ri"rj "m xkri!"k" "! xkri!"k" zlkp z!"~
rj!j !|l!!x ij!j j l" lij r"klk"~lsj m"!sl"kv " z!lr!
r!j kjj l" |j lojk m"! lij kljp!l"k "m lij {j!jkl jl"k & k lij xksr z!"rj i !j l"k z!"zj!lj m"! lij i"!l~lsj xksr
"m |"li lij rltlj *+ yj m"! lij !jj!t"! jtj 6 + v hij s"j
ksj!rx !"|lv hij xksr ksj!r l|j " m"! lij "kp~lsj
jt"l"k "m lij rltlj *+ yiri !tjk |x lij !jzjrltj !jj!t"! jtj
t lij !jj!t"!~mkrl"k 6 k 6 v
hij sl"k z!jjklj k pv yj!j zj!m"!sj kp lij ly" lkrl
z!sjlj! jl lj k h|j & v "! lij z!sjlj! jl yj "|j!tj tj!x !z
!j l"k l"y! "kj "m lij sjs"!x~llj jkr"j k lij ko~sl!rj "m
lij kjly"!o i"yk k pv v hij !jj!t"! jtj !j jzjlj tj!x "yx
k lij sjs"!x~l|j |jr"sj kl|j k {j!jkl sjs"!x~llj loj
"tj! "kx mlj! |lkl lsj i zjv "sz!kp lij li"pil z!"rj
i"yk k pv yli lij kjly"!o "m j rll"!x ko "m lij i"yk k pv
"kj rk k"lrj lil } kx ly" |jjkl sjs"!x llj !j r"kkjrlj |x
"kj "! s"!j ko k lil }} j rll"!x ko itj k mrl mkrl"klx '
= . O) ;= (%& 7 '$.Q ,-/( P -& 1+/0.%%.%
ri
E
0
-0.5
50
100
200
150
1
(0)
(1)
(2)
(3)
(4)
(5)
0.8
xi 0.6
0.4
0.2
0
50
(4,8,10)
(8,11)
(6)
(7)
(8)
(9)
(10)
(11)
(6,7,11)
100
(0,1,2)
200
150
N N ,-. P +/S&- ; + 7 &.% X Y X&/ R Y 7 $Q &-. 7 0&'6'&2 A X Y XV/&&/ D Y )/+ &-. &-/( P -&
+/0.%%
XA ? >@Y XA C >@Y XC >>Y X? >>Y X@ > Y )/+ &-. > ; %'&. $.&S/+3
R
%-/S$ '$ 8'P T 4 (%'$ P &-. R7 + 7D .&.+ %.& XVY /) ,7 V O. >
h" l|‚j l!kjkl sjs"!x llj k l" "rlj "kj sjs"!x llj yli
|jjkl "kjv hij s"j "j lij!jm"!j !j‚j lij ly" z"llj m"! jm~
!jplj "rltj li"pil z!"rjj jl m"!li k lij kl!"rl"kv
""prx zjokp l nylj "m lsjq m kt sjs"!x llj !jsk
rltj m"! k j rjjkpx "kp klj!tv hij jzjl"k~!lj
1 2 ' m"! lij
!jj!t"!~jtj tj!x s m"! lij z!sjlj! jl lj k h|j & v "! lij
z!sjlj! jl | yj itj ri"jk |lklx !pj! jzjl"k~!lj 1 2 l"pjlij!
yli tj!x s""li !jj!t"!~mkrl"k 6 k 6 k €klj tj m"!
+ k "!j! l" t" !k"s !ml m"! rjklj!
rltj "tj! z!""kpj zj!"
k mx jzjlj !jj!t"! jtjv
hij ly" li"pil z!"rjj i"yk k pv !j m"! jklr ko~sl!rj
"kx lij z!sjlj! jklj!kp % v &
k {j!v wj k"lj lil lij jjkrj
"m sjs"!x~llj jklr m"! |"li z!sjlj! jl lij xksr l|j "tj!
yj !kpj "m z!sjlj!~tjv hij il"!x "m sjs"!x llj p"j li!"pi
rxrj l jtjkl m"! lij sl"k kp lij z!sjlj! jl |  krj
lij zij~zrj €kljv hij & ~lj rlj! j k li sl"k r"klk &)
{j!jkl sjs"!x llj k lij rxrj !k "tj! ( lkrl llj |lkl
m!rl"k "m lij l"lv "! !pj! kjly"!o yli lij! tj!x ipi ks|j! "m l"!j
sjs"!x~llj jj iskk to" riljk &'5' lij rxrj jkpli y
|j k pjkj! tj!x "kp m"! kx z!rlr z!z"jv wj itj i"yjtj! k"l xjl
zj!m"!sj xljslr lx "m lij rxrj~jkpli "k lij xljs z!"zj!lj "kp
lij sj kj x "kj m"! !k"s |""jk kjly"!o rilj! & v
wj k"lj lil |k!x rxrj " k"l "rr! m"! lij z!sjlj! j ij!jv wj
k lij !jj!t"! jtj lij!jm"!j loj yij l" €~
itj ri"jk
z pk v rltj sjs"!x llj rk lij!jm"!j k"l !jrltlj lij! z!jjrj"!
yiri itj kjrj!x "y !jj!t"! jtj k s tj m"! 6 v hij sj
!"
>@
O7 (Q'(% E+/%
1
(0)
(1)
(2)
(3)
(4)
(5)
0.8
xi 0.6
0.4
0.2
0
0
(6)
(7)
(8)
(9)
(10)
(11)
(1,3,6,9)
250
500
(5,7,8,11)
(1,2,4,11)
750
1000
1
0.8
xi 0.6
(6,8,9,11)
0.4
(0,3,4,5,6,7,10)
(0,1,4,5,6,7)
(2,6,7,8)
0.2
0
0
100
50
200
150
N N ,-. 7 0&'6'&2 A X Y )/+ &S/ > ; 0.$&.+ 0 O (%&.+% S'&- 7 -' P - Q.$%'&2 /) O '$3% (%'$ P
&-. R7 + 7D .&.+ %.&% X7 Y 7 $Q XVY /) ,7 V O. > X&/ R < V/&&/ D YT
,/ R Z ,-. &-/( P -& R +/0.%% '% X> A ? >>Y X@ ? C >@Y X> B ? E Y X C >>Y X C >>Y X> A >>YT
/&&/ D Z ,-. &-/( P -& R +/0.%% '% X@ B A ? >@Y XB ? C >@ >>Y X? C E >>Y X@ B A ? >@Y X@ > A ? Y X ? CY
ljsz"! xssjl!x r" |j rijtj |x ri""kp + 1 + v wj " k"l !j~
"l lij j "m xssjl!r ko~sl!rj m"! lij  |l l k sz"!lkl
z!"zj!lx "m lij l" |j |j l" jl|i lsj !jrl"k l"k"s"xv
hij lxzj "m sjs"!x~llj l"!|j k lij y" "lij!yj |j slj
l" xssjl!r lljv
a g c a ` gf }k pv ) yj z!jjkl |"yz "m lij li"pil z!"rj z!jjklj k pv m"! lij
z!sjlj! jl |  l"pjlij! yli lij lsj~jzjkjkrj "m lij p!"yli !lj + v
wj rk rj!x "|j!tj i"y r"szjll"k s"kp lij zx r!r !"jv
"! lij €!l l!kl"k "rr!!kp l
5 lj & k 5 r"szjlj yli jri
"lij!v "li itj ly" j rll"!x ko yli
lij rltj rlj! ) (  &  jj pv
k pv 5 v hi r"szjll"k !j"tj ij!j |x s !k"s {j!jkrj k
|jlyjjk lij j rll"!x ko + j ij!jv
&' k |jlyjjk lij ly" krl sjs"!x
hij l!kl"k "rr!!kp l
llj (   &&
k &  loj |lkl s"kl "m lsj l" r"szjlj l
p"j li!"pi lij klj!sjlj llj  v hij sjs"!x llj  rlx t sjs"!x llj |x ljm |l k"l l|‚j ij!j lij ""j "l
k lij r"szjll"k yli &
k  r"sz!j pv ) v
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k"l "rr!v hij klj!sjlj llj 5 &&
l|‚j |jrj lij !jj!t"! jtj
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>>
1
0.8
xi 0.6
0.4
0.2
0
(10)
(64)
(12)
(83)
(21)
(85)
(26)
(86)
(39)
(93)
(54)
(94)
(58)
(98)
0
50
100
150
200
0
50
100
150
200
1
0.8
ϕi
0.6
0.4
0.2
0
N N ,-. 7 0&'6'&2 A X Y 7 $Q &-. +.%.+6/'+ X Y )/+ 7 EE ; 0.$&.+ 0 O (%&.+% S'&- >? < >AB < ?
%& 7 V O. D . D /+2 %& 7 &.% 0/$& 7 '$'$ P < B < A 0.$&.+%4
(%'$ P &-. R7 + 7D .&.+ %.& XVY /) ,7 V O.
> ,-. &-/( P -& R +/0.%% '% X ? ?A E BY X C ?A E BY XB E C CBY XB E CB E CY X>@ B E E CY X A C C? E AY X > A C? E AY X> > AY
"m ( i xjl k"l |jjk !j€j r"szjljx j l" z!jrjjkl rltl"kv }k li
rj ( k ""j "l k r"szjll"k yli 5 v
}k pv ( yj i"y lij !jl "m ly" sl"k "k {j!jkl & ~lj rlj!
yli tj!x jkj ko~sl!rj + yiri r"klk sjs"!x~llj yli z l" jtjk
rjklj!v sl"k m"! |"li jl "m z!sjlj! !j i"yk k yj "|j!tj lil
lij xksr y"!o zj!mjrlx €kjv w li lij zz!"z!lj ri"rj % v ( m"! lij
ko~l!jkpli jtjk kjly"!o yli |lklx !pj! sjs"!x llj "y m"!
ksj!r l|j sl"kv
kx yj z!jjkl k pv lij rltlj k !jj!t"!~jtj m"! '' ~lj
kjly"!o kp lij z!sjlj! jl | v 4 "llj k pv !j "kx lij rltlj
k lij !jj!t"!~jtj "m lij lj rltj !kp lij klj!t "m "|j!tl"kv wj
rk krjx "|j!tj lij jzjl"k "m lij !jj!t"!~jtj m"! lij rltj rjklj! k
lij "sjyil "yj! !jr"tj!x "krj lij rltlx m pk v s! !jl !j
rijtj " |x sl"k "m tj!x |p kjly"!ov
g fe gc`
wj itj ij!j ktjlplj lij l"k"s" xksr "m lij k kjpjrlj
kx klj!rl"k yli lij "lj y"!v jk"!x kzl y" l" lij p!"yli
!lj |x zz!"z!lj lsj~jzjkjkl s"l"k "m lij !jzjrltj | + 
jj % v v "! kx jk"!x kzl lij | "m lij kt"tj rltlx rjklj! y"
r!j €klj z"ltj tj !kp lij klj!t "m lij jk"!x lsl"kv
hokp ""o l lij p!"yli !lj z"llj k p v ) l |jr"sj ssjljx
rj! lil lij jk"!x kzl kjj k pjkj! rj!lk r!lr l!jkpli k "!j!
l" kujkrj lij "kp"kp li"pil z!"rjv hij l"k"s" li"pil z!"rj k
>
O7 (Q'(% E+/%
(6)
(5)
(6)
(7)
(5)
(8)
(4)
(2)
(9)
(3)
(2)
(10)
(1)
(8)
(4)
(9)
(3)
(7)
(11)
(10)
(1)
(0)
(11)
(0)
N N 2$ 7D '0 7O 7 &&.$&'/$ ; )/0(%'$ P '$ &-. > ; %'&. $.&S/+3 S'&- &-. &-/( P -& R +/0.%%
'OO(%&+ 7 &.Q '$ 8' P T 4 )/+ &S/ Q'[.+.$& %& 7 V O . D . D /+2 %& 7 &.%T ,-. Q 7 %-.Q < Q 7 %-.Q ; Q/&&.Q
02 7 $ 7 $Q R '$3 O'$3% < 0'+0 O .% Q.$/&. 0.$&.+% O'$3.Q S. 7 3 O2 < %&+/$ PO2 XV2 /$. < &S/ . : 0'& 7;
&/+2 O '$3%Y &/ &-. 7 0&'6. D . D /+2 %& 7 &.
lij jk"!x kzl r"szjlj yli jri "lij!v hi ok "m r"szjll"k "j k"l
"rr! k szj ll!rl"! kj! kjly"!o rilj! &  m"! yiri kx k"k~
‚j!" kzl krj lij kjly"!o l" u"y kl" lij kj!jl rrj|j ll!rl"!v "li
m"! kzl !jjs|kp r"jx z!jt"x l"!j zllj!k yj m"! !k"s
k k"kjkr kzlv
hij r"kj!j ij!j y "k lij "lij! ik !jr"pk‚j j lj!k zl~
lj!k "kx yijk lij kzl ri lil l yk lij r"szjll"k yli lij "kp"kp
li"pil z!"rj jkp l" lij rltl"k "m lij r"!!jz"kkp sjs"!x lljv
hij l!jkpli "m lij jk"!x kzl kjrj!x m"! li !jr"pkl"k z!"rj l" |j
r"szjlj rrjmx jzjk r!rx "k lij ks|j! "m ko |jlyjjk lij r!~
!jkl rltj sjs"!x llj k lij lslj |x lij jk"!x kzl "kj
rk jj k pv ) k pv 5 v hij lj &
k 5 itj ly" ko l" lij sjs"!x
llj ) ( &  "lij! rjklj! itj jlij! ‚j!" "! l "kjv kx jk"!x kzl
!!tkp "k lj &
"! 5 y" |j !jr"pk‚j jtjk m"! s pk~l!jkpli
yijk ) (  & rltj jk"!x kzl !!tkp l lj y" kjj l" |j
"k lij "lij! ik tj!x !pjv hi z!"zj!lx "m lij lijk jtjkl l"
lij rz|lx "m m"r lljkl"k l"k"s"x k sz"!lkl z!jr"kl"k m"!
"| jrl !jr"pkl"k |x r"pkltj xljs jj jxk" js"kj &''' v % t~
j!x rltj sjs"!x llj r!!j yli l k n"rl"k r"q k kx j lj!k
kzl !!tkp ylik li !j "m koj !jrlx l" lij rltj sjs"!x llj
y jk "x z!jmj!jkl l!jlsjkl |x lij v
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'
l €!l pil lij s"j % v & z"jj k k|""pr mjl!jv jpjrlkp
lij 6 k 6 m"! s"sjkl lij l"l j{jrltj ko l!jkpli +
+ r"klk" % lij! l!"kpx kjpltj + 1 . +
 "! yjox z"ltj
+
. + 1  l!lj k pv ' v hi z!"zj!lx "m lij ko~
sl!rj |jlyjjk lij r"klljkl rltlx rjklj! r!r m"! lij yi"j s"jv
= . O) ;= (%& 7 '$.Q ,-/( P -& 1+/0.%%.%
>B
allowed values for
link−strengths
(1)
(2)
inhibitory interneuron
w
excitatory
0
−|z|
inhibitory
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