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Y = ln X
2
f (y) ∼ e−(y−µ)
/2σ 2
f (x) f (ax) = , f (1) f (a)
a
F (ax) F (a)
F (x) =
=
f (ax) f (a)
= F (x)
f (x) f (1)
=
f (ax) f (a)
G(x) = log(F (x)) G(x) = α log(x)
F (ax) =
f (ax) f (1)
F (x) = xα
F (a) =
f (a) f (1)
G(ax) = G(x)+G(a) f (x) = xα · const
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new
new
'U
a)
b)
1
δE δ nn 1/s2
E ρE nn 1/s
10−1 P(>s)
10−2 10−3 10−4
c)
10−5 1
101 102 103 104 s
d) 1
101 102 103 104 δs
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6F< ,.@
' 8y),54 0!L8% '*4%,.- '*,5As8%(' (';254%';' L/0j8:4J/ >8%/ ,.BdN,5;ud@$:5J>N@$dZA2;)9$E q5; 4>N3>B@$EDq`9$HX;)£4q $g 4%>B2_`24%EDdZ4^4ZA]F6^44EA2,D4H ;uB44 q`;u2;)5G`AB;K9?E 6C406^9$G5:uqr:);K_?4FA29G5EDq`4%>BNdN,5;uPd@p(k):kJ4%@pA2G5>B4%7;KE 9pAB,54>T6C9?>BqDBdN,.-\J9$>!J>24%4j=O &@$2E54A!@$EDqF4§ÀH¨@p;)(: j( i59?>3A2,D@pA%e5:)4ZA8GDB9}Y`;KH¨@AB4\AB,54Fq`4%=$>B447q`;)B;)5G`A2;)9$E p @ N > N d . , γ = 2.0 3000 1.00 0.00 7 0.14 ¡\@pEDq`9?HX;K£%4%q 1.00 0.00 6 ± 1 − n¦@Y`;KH¨@$:@pEA2;K§À,D;K4%>B@$>BdN,.0.38 ± 0.00 0.00 10 ± 1 −1.24 e5no@pY.;)H¨@p:,5;)4>N@p>NdN,.- 3000 1.0 γ = 2.5 0.00 11 ± 1 0.44 ¡\@pEDq`9?HX;K£%4%q 0.88 ± 0.02 0.04 13 ± 1 − n¦@Y`;KH¨@$:@pEA2;K§À,D;K4%>B@$>BdN,.0.05 ± 0.0 0.00 14 ± 1 −1.51 e5no@pY.;)H¨@p:,5;)4>N@p>NdN,.- 3000 0.89 ± 0.05 0.00 15 ± 1 0.16 γ = 3.0 ¡\@pEDq`9?HX;K£%4%q 0.03 ± 0.03 0.00 80 ± 13 − n¦@Y`;KH¨@$:@pEA2;K§À,D;K4%>B@$>BdN,.0.004 ± 0 0.00 240 ± 20 −1.11 1#@$5:)4 ( mDa!13,D4FdZ9$HXLD@$>B@pA2;)g$4\AB@pD:K479pJ!,5;)4>N@p>NdN,-XL5>B9$L4>2A2;)4%vq`; 4>B4EA8>N@pEDq`9?H E54AM§ 6C9?>2_`B447q`;)B;)5G`A2;)9$Ek(¥©\4>B4 N ;u89pJkE59`q`4BdN,5;ud@$:&e68,5;):K4 S HX4@$24 9pJE59$E5§À,5;)4>N@p>NdN,5;ud@$:.MANdZG`AN2_e$@$EDq r ;uB9$D@$5;K:);KA]-A2,D@pA¤@:K;)E5_§À4%EDq 68;K:):c4 @AMAN@$dN,54qoAB9o@sE59`q`4¨9pJ3@rq`4=?>24%4,5;)=$,D4> A2,D@$E k ;u< Z ()' P (≥ k) ∝ k P (k )dk , 68,54>B4 P (k ) = A·k ;u24%4^q`;uB;K5G5A2;)9$E068;AB,Xª 4;)E5=S@TE59$>BH¨@p:);K£@AB;K9?E dZ9?ED2_e (#iD9$> A2,D4c;KEAB4=$>N@p:?q`;Kg?4>B=$4B4ZJ9$>B4¥JG5>2A2,54%>k6^4c68;):K:q`;)B447,5;)=$,54%>3A2,D@$E k ;u< J9?> γ < 2 N= ·k , J9?> γ > 2 N= , −γ
AD
AD
kmax
link−end
0
0
0
k
0
0−γ
max
max
2−γ A γ−2 A
max
γ−2
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pU *;AB,A2,5;u2H¨@p:);)£%@AB;K9?Ed9$ED8A2,D4F2
J9$> γ < 2 (VU J9$> γ > 2 13,54CLD>29?D@p5;):);A]-A2,D@pA!@\E59`q`4h9$JDq`4=?>24%4 k q`9.4%B;)68;KA2,0A29$L9$:)9$=?;)d%@p: Â?G@pEA2;KA2;)4%24FHX4@$24qJ9$>@ÁB_`C< p(v13,54>24B4L9$>2A24%q ;KEw1#@$5:K4 (Vx`(T13,54q`4%L4%EDq`4%EDdZ409pJ F (γ) J9?>>N@pEq`9$H 24%4FE54A]6C9?>2_`B4=?4E54%>B@pA2;)E5=0HX@pY`;KH¨@p:,5;)4>N@p>B,Dd;)@$:@$EDqHX@pY`;KH¨@p:@$E?AB;§Å,5;)4>N@p>NdN,5;ud@p:>B4ZJ§ 4>B4EDd48E54ZA]6^9$>B_`29`q`GDd4%2>B4:u@AB;K9?EL5>29$tD:)4%NdN,.-s68;):):4 @?B@$>BdN,.-q`;)BdN,5;udγ@$:$∼LD@p2A2,DB_68;KA2,0=$;)g$4%EF:K4%E5=pAB,D< 9pJ¥/MG5L LD@pA2,.Oje @pEDq/2q`968E¦LD@pA2,.Oje (Xi!;)=$G5>B4 (®
@L5>2424BEsJ9$>3AB,5l4S½PEA24>BE54A3;)EsA24%>2H¨B@pA2;)9 N (l , l ) ( H(l , l )= . up
down
up
down
up
down
Nrand (lup , ldown )
Up
l up
a)
b)
6 5 4 3 2 1
5
1.4
4 1
3 2
0.6
1 0 1 2 3 4 5 6
l down
0.2 0
1
2
3
4
l down
5
i!;K=?G5>24 (®
.a#©\;K4%>B@$>BdN,D;)d%@p:5LD@pA2,DNdN,5;)d%@p:3LD@pA2,dZ9?E?AB>2;)5G`AB4% 2A24%B@$>BdN,5;ud@$:5LD@pA2,(c13,54T@pDBdN,.@p>B4ED9$>BHX@$:K;)£4qÁ.-¨AB,54F3J>24Â?GD4EDd;K4B4%@$:½PEA24>BE54A%( 68,54>B4 N (l , l ) ;)89$J¥@$:K:,5;)4>N@p>NdN,5;ud@$:LD@pA2,D8=$;)g$4%E l @$EDq l ( ;u2>B4%¤J9?>AB,54s>B@$EDq`9$HX;)£4qµg$4%>BB(l_(#,13l,54>24%):)@pA2;)g$4%:K-0:K;)=$,A^>24%=$;)9$EDB4v;)EDq`;ud@pA24vAB,D@ACA2,5472A24B@$>M§ dN,5;ud@p:#LD@AB,DTHX9$>B4F9pJAB4Eo;)EoAB,54¤½PEAB4>BE54ZAA2,D@$Eo;KE ;ANN@pEq`9$HX;)£4%qrd9$G5EAB4>BLD@p>2A%( 13,54FA]6^9 H¨@p;)Er9$2g@AB;K9?ED24?aT' ^A2,D4:)4E5=$A2,r9$JA2,540MAB4%N@p>NdN,5;ud@p:kLD@AB,DB4%@$:`½PEA24>BE54Ah@$B4\HX9?>24J>24ÂG54EA%(¥½PEsi#;)=$GD>24 (®
}6C4T4ZY5@pHX;)E54T;KEÁA2,54S0AB,54H¨@pY.;)H¨@p:3,5;)4>N@p>NdN,.- ;u29?H¼A2,54s>24@p:^½PEA24>BE54A0.-µ;)EDB@pA2;)9 9pJcL@A2,wdZ9?G5EA4ZA]6^44EoA2,540H¨@Y`;KH¨@$:#,5;)4>N@p>NdN,.-@$EDqrAB,54¤>B4%@$:#½PEA24%>2ED4ZA%(T*w40B4%@p:½PEA24>BE54Ah;)hJ>29?H AB,54\H¨@Y`;)H¨@p:D,D;K4%>B@$>BdN,.-$e?@pEDq¨@$:)N@p>NdN,-X;uL@ @ 6F4:
M AB31 4 ,5>BLD4\@pA2>29?ABL04%>BA2@$,5>B4¦dN,.5-;K9?9pJk:K9?@p=$>2;uA2d;K@$t:^d;)ED@$4Z:DA]E56^49$A]>B6C_³9?>29p_`JTB@?dZdj9?ABG5;K9?E`E § [\Z4 A]6^9$>B_NdN,5;ud@p:J4@ABG5>24B4:u@A2;)g$47AB9 ;KABB@$EDq`9$H dZ9?G5E`§ AB4B> LD@p>2A%( M 4%v¡ B> @$@$ED>BdNq`,59?;uH d@$: E54Z A]6^9$>B'_`6829@$q¦BED4qÆ4XA2q`,54%4=$>BE5444XA]6Cq`;)9?2A2_`>B;)B;)49$Eµ:K4NL@p9?>N:KdN9?,5=$;u;udd@p@p:C:¥A2,5,D;K4 § v;uB447q`;u2;)5G`AB;K9?E 068,54%E Z( F
γ = 2.0
M 13B4%4 >24\4Za Y`;)2E@A2;)g$4863@}-`2_¤A29¤dN,@pE5=?43A2,54,5;K4%>B@$>BdN,.-FHX4@§ F F ≈
γ=3
m
@! 6 @p>B-XA2,547AB9pAN@p:E.G5H4%>89pJ#,D;K4%>B@$>BdN,5;ud@$:LD@AB,D< AB,D@Av;uB- U Z( s13,5>B9$G5=?,Ág@p>B-.;KE5=0E.G5H¤4>\9pJ#ED9$E`§Å,5;)4>N@p>NdN,5;)d%@p:2ABdZG5AB< S j(
Bk +? Æ
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P½ H¨@p=$;)E54CA]6C9L4%>BN24%4%q`9?H A2,D@$E0@$9?G`Ac6^4:KJ~@p>B4^9$JDAB,54;)>h@$d%dZ9?H0LD:K;udZ4?(#13,54vL.)D0,6/8e!/#N, PL-100,S )/.0,\.6\!/#! 0,L !W. N(*'%)/0,!3. !Q&C&0 S$S>PQ0,. )X!/Sc#+I!) !/-#)5E5! )5-( .!W: /6$.'. . !BS,+T+D)$!/#!/8#)56[@!B0,P)) [!BP!/ ) '(70,&.CE)5..6 .10, [02!/!5! 5A5P3ba!/. )/!/# 6 02!/#!/EG /&CAX0,bMLAJP&%)5 J: `V #[.!7L5! ..& Q.'(02.! 6
$m 4%ÂG5;):K;)5>B;KG5H dZ9.9$L4>N@AB;K9?E¦:)4g$4%:¥;)2-A2;)HX40=?4ZAN 68;AB,@$E;KEDd>24@$89pJ¥:K9?E5=>N@pED=$4SdZ9?E5E54%dZA2;)9$ED2;uMAB;)d:K4 ! 13,54sÂGD@?B@pA2;)g$4%:K-¨:u@p>B=$4=?@p;)EJ9$>^A2,54Fq54ZJ4%dZA29?>BB=$4\J>B@?djAB;K9?ρE9$J#9$G5EDqD@p>B-¨4%q`=?4%)/*.@02;K!/... > E +I)5! B#:& -!/#)/$'/0,#'.)5.!/!BS/!4,+L-102!/#!/# !/[> #1 0,!8-! -(C;#10,-)\-(,e! N N(!/#$0,&a!,+L.+I)5&C02! .E! C-)5.[!8P[! #)806. > [> !8.*6.6 G :^WM B7)\W0,/ 0,c!/#%
Pi
Ii = − log2 (Pi )
Pi = 1/k
I = − log2 (1/k) = log2 (k)
'
t
P1(s → t)
P2(s → t)
!i ;K=?G5>24X5( m5a713,54%>24¨@p>B4A]6C9rq`4%=$4ED4>N@A24X2A242;)5G`AB;KE5=e!B9$D@$5;):K;KA]-rA29oJ9$:):K96 ;Ae P (s → t) 29?H A2,D;)NdN,ƽPE`J9?>2H¨@AB;K9?Ee S(s → t) e C6 9?G5:uqÁ4Sg$4%>2- N@$djAB4>B;)8NdZ4%< Ht =
X
S(s → t)/N
X
S(s → t)/N
BA 4:):)3@p:):AN@p>B=$4ZAN< t a s
As =
^6 4r9$`AN@p;)E@µÂ?G@pEA2;KA]-$e iLWYWXQh\Y\hkrhqVS_t UV^`PRqVk´AB,D@A AB4:):)< GDBq5;KE5=oA2,54%;K>¨ª8dd4%B4SE59`q`4B47@p>B4vE59S/];u29?LD@p=@A2;)9$EÁ9pJ2485( a!ªvED@$:K-`B9pAB4;)E¤;)EA24>N@$dZA2;)9$EE54ZA]6^9$>B_S;KEX-?4%@$B9pAB4;)E;KEA24%>B@?djAB;K9?ErE54ZA]6^9$>B_sd9$:)9$>\d9`q`4%qo@?dd9$>Nq`;KED=XA29 A e9$ E5
:)(wgÀ-µe139?;KE`,5ABAB44@$>N;KdZ@$ED9?dj4%>2ABq4;K9?EDq5.-w@B_.B9pAB4;)ED ,!/+T#))!/)/[[!BABP. ) 8c#,.+F)/!/0,#)/8 #a S70,& 0,)/ . #: ;+T)5&!/#*!/S
∼ log2 (N +1)−1−2/N +4log2 (N +1)/(N (N +1)) ≈ ∼ log2 (N/2)
x 9pJ@pEr@$qDq`;AB;K9?ED@p:kG5EDq`4%>2:)-.;KE5=¨BGDdjABG5>B4~N@pL5,D;)d%@p:Àe4ABd$( ¨A2,D@pAvL5>29g.;uq`4%BHX@pA2;)9$Ee$GD29`d4%B;AB,5H¨85;K= N@A24?e ρ → ∞ e ek68,54%>24 ;)MAN@pEDd4¨9pJ3A2,54ÁE54;)=$,.9$>Ne?;)EÁJ9?>84ZY5@pHXL5:)4SB_.NdN,Ád@$E4FBHX@pA2;)9$EE544q`4%q A29µNdN,Æ;KJ79$E54r@p:)>B4%@$q5-µ_ED96vB@$:K:^AB>B@ ¨d¾D96S( ½PE A2,5;u< d%@$B4:u@A2;)g$4 6C4%;K=?,A\9pJc4Y.;KA:);)E5_`B9$H @ E59`q`4 ;)!@$:K:.;)Eg?4%B_`N@p=?4=?@p;)EDBdN,FN@AB4=?-$(k©\96C4%g$4>e%A2,54^dZ9$EAB>2;)5G`AB;K9?EF;) l ≤ 3 @pEDqFA2,.GD2ED< 9$G`AvAB9 4F;KED4 dZ;)4EAvAB9¨J9$:):K96 A2,D4S¾D96É68,54EAB,54SAN@p>B=$4ZAv;uB.-aC.- J9$:):)968;KE5=XAB,54 ¾D96-$9?G 68;K:):E54%@$>2:)-X@p:)6^@}-`B:K9.9?_B9?@$qDCED4;)=$,.9?>2,D99`q(#½PE AB4>BHX2EB9$H -?9$G5>h4EDq0AN@p>B=$4ZA(!13,D4E¨;KAhED@ABG5>N@p:):K-LD@}-`NqÁAB,540HX@$;KEw24@pH(13,D40q`;uB4 4dZ9$HX424J9$>B40q54ZtDE54N@$djAB4>B;)B;K£%9$Ee l S (l) − S(l) e@A 68,5;)dN,9?E54S2405( 5a\ª\ED@p:)-`B:)qoE54ZA]6^9$>B_`B_ 9pJkno@$:KHeVx$xHgÀ(¥½PEA24%>2E54AC;uB_9$Jk@pG`AB9$E59?H09?GDB9?24< S(l) 68;KA2,wA2,54 BdN, S (l) HX4%@$B4%q ;)E @ >B@$EDq`9?H0;)£4qµg$4%>BB_( E54s9$2g?4%2_`B4 68;KA2, A2,54XNdN,r9$5AB@p;)E54qo68,54%E¦9$ED4GD2_(#13,D4v=$:)9$D@$:`A2>N@ d8;)E`J9?>2H¨@AB;K9?EX,54:)LD
i
i
ki
i
ij
j=1
2
ij
S = 10.5 T = 0.14 R = 0.245
S = 7.9 T = 0.55 R = 0.72
S = 6.8 T = 0.81 R = 0.82
a
b
c 5
S = 14.5 T = 0.16 R = 0.20
S = 7.9 T = 0.55 R = 0.72
S = 6.6 T = 0.73 R = 0.84
d
e
f 0
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3.5 T (ER) 3
T (SF)
2.5 2 T 1.5 1 0.5 0
γ
2.0 8.8
2.1 6.4
2.2 5.0
2.3 3.9
2.4 3.2
2.5 2.9
2.6 2.6
2.7 2.4
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B
Bacteriophage 186 (30.6 kbp)
Bacteriophage
λ
(48.5 kbp)
E. coli proteins Regulatory proteins Early lytic proteins Late lytic proteins Lysogenic proteins N
B
RecA
A
ClpXP Q
RecA
O P
Tum CI
CI
LexA
LexA Lon
Apl
Int
Cro
CP69 Xis
CII
Int
RexA
RexB
CIII HflB
CII
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Signaling Distance: DS (A, B) = ∑i, j |ASij − BSij | λ
186
DS = 43 186r
λr
DE = 32
Random, P(k) preserved DS = 108 ± 34 λrr
DE = 32 ± 1
186rr Random, P(k) – Poisson
DS = 1050 ± 90 DE = 1060 ± 200
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1
2
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σ12 N1
σ22 N2
1
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iE 36 17 2
λ 186
iS 17 0.7 11
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