0 20 cell number 0 cell number 20rubin/ls04bump.pdf · bumps in neuroscience models with mexican...
TRANSCRIPT
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Activity Patterns in Purely Excitatory Networks
Jonathan Rubin
Department of Mathematics, University of Pittsburgh
SIAM Life Sciences Meeting - July 11, 2004
MAIN POINTS:
• spatially localized, temporally sustained activity (bumps) canoccur in purely excitatory networks
• the go curve provides a useful construct for understanding theunderlying mechanism and properties
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Bumps: what are they and why do we care?
cell population/stimulus feature
firing rate/firing probability/averaged voltage
• spatially localized, temporally sustained activity
•which cells are active depends on some external feature
• activity persists without persistent stimulus
• seen in visual system, head direction system, and pre-frontal cortex (working memory):
stimulus shown, subject must later recall position;localized group of cells fire until recall
stimulus memory recall
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Recipes for bumps
1. E
I
e.g. Wilson/Cowan/Amari: ut(x, t) = h− σu(x, t) +∫ ∞−∞w(x− y)f(u(y, t))dy
−4 −2 0 2 4
−0.2
0
0.2
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0.6
x
w
w(x) > 0 on (−x, x)
w(−x) = w(x) = 0
w(x) < 0 on (−∞,−x)∪ (x,∞)
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Bumps in neuroscience models with Mexican hat
• Guo and Chow, 2003
• Coombes, Lord, and Owen, 2003
• Renart, Song, and Wang, 2003
• Laing and Troy, 2002 & 2003
• Laing, Troy, Gutkin, and Ermentrout, 2002
• Gutkin, Laing, Colby, Chow, and Ermentrout, 2001
• Laing and Chow, 2001
• Pinto and Ermentrout, 2001
•Werner and Richter, 2001
• Compte, Brunel, Goldman-Rakic, and Wang, 2000
• Taylor, 1999
• Camperi and Wang, 1998
• Hansel and Sompolinsky, 1998
• Amit and Brunel, 1997
• Kishimoto and Amari, 1979
• Amari, 1977
•Wilson and Cowan, 1972 & 1973
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2. Rubin, Terman, and Chow, JCNS, 2001: no E-E; PIR
E
I
TC cells
time
7500
9000
10 40
-80 10
mV
RE cells
time
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9000
10 40
-80 10
mV
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3. Rubin and Troy, SIAP, to appear:
E
I
⇓
w(x)
x
off-center coupling
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4. One-layer network with purely excitatory coupling[Drover/Ermentrout, SIAP, 2003; Rubin/Bose, 2004]:
Equations
v′i = f(vi,wi)− gsyn[vi−Esyn]cosi + Σ
j=3j=1cj[si−j + si+j]
w′i = [w∞(vi)−wi]/τw(vi)
s′i = α[1− si]H(vi− vθ)− βsi (on a ring)
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v
w
. o
SNIC
(movie)
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Geometry - θ model:
θ′i = 1− cosθi + (1 + cosθi)(b+ gisyn)
gisyn = gsync0si + Σ
j=3j=1cj(si−j + si+j)
s′j = α[1− sj]e−γ(1+cos θj)− βsj, j = i− 3, . . . , i+ 3
with α,γ large
synaptic decaydynamics:
θ′ = f(θ, gisyn),
g′isyn = −βgisyn
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θ
gisyn
−b
(θS,0) (θ
U,0)
P
go curve
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Geometry - θ model - (2):
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θ
gisyn
−b
(θS,0) (θ
U,0)
P
go curve
synaptic decaydynamics:
θ′ = f(θ, gisyn),
g′isyn = −βgisyn
main ingredients:
• threshold of synaptic decay dynamics determinesresult of stimulation
• delay to firing depends on proximity to threshold
• variable delays desynchronize and stop spread
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Geometry - Morris-Lecar with w′ = 0:
v′ = f(v,w)− gisyn(v−Esyn)
w′ = 0
s′i = α[1− si]H(vi− vθ)− βsi
decay dynamics: replace si-equations with g′isyn = −βgisyn
l 2
(v , w )l
w=w
w
v
g=gg=g
1g=0
g
v
g=g
g=g2
1P
(v , 0) (v , 0)l mc
g=0
l
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V
gisyn
go curve
P
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Geometry - Morris-Lecar full system:
v′ = f(v,w)− gisyn(v−Esyn)
w′ = g(v,w)
s′i = α[1− si]H(vi− vθ)− βsi
decay again: replace si-equations with g′isyn = −βgisyn
w
v
w=w
m
go curvefor w=w
(v , w , 0)
RK
RK
m
gisyn
v’=0,isyn
v’=0, g =0
g >0
isyn w’=0, g =0isyn
W s
note: can still project into (v, gisyn) phase plane!
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Analysis: who jumps?
w
v
w=w
m
go curvefor w=w
(v , w , 0)
RK
RK
m
gisyn
v’=0,isyn
v’=0, g =0
g >0
isyn w’=0, g =0isyn
W s
• suppose that cell i gets a synaptic input from cell j
• check the projected (v, gisyn) phase plane for cell i atthe moment cell j falls down through vthresh
• position of cell i relative to go curve determinesrecruitment
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Implications - identify recruitment
−0.32 −0.3 −0.28 −0.26 −0.24 −0.22 −0.20.005
0.01
0.015
0.02
0.025
0.03
0.035
V
g isyn
−0.24 −0.22
0.026
0.028
0.03
0.032
V
g isyn
first
second
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0.015
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0.025
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0.035
V
g isyn
shut off
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Implications - recruitment depends on more than |gisyn|
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0.01
0.015
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0.025
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0.035
V
g isyn
0 50 100 150 2000
0.005
0.01
0.015
0.02
0.025
0.03
time
gisyn
• also, faster synaptic decay hurts in two ways...
• ...and bump size is non-unique and nonrobust
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Bump formation:
• recruitment as above; initial synchrony helps
−− start near same rest state
−− common input overrules synaptic coupling
• localization via desynchronization after shock (delayedescape from go surface)
• sustainment via self-coupling (not sufficient!) andasynchrony
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Synaptic depression:
Bose et al., SIAP, 2001:
s′ = α[d− s]H(v− vthresh)− βsd′ = ([1− d]/τγ)H(vthresh− v)− (d/τη)H(v− vthresh)
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0
0.5
v
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0.5
1
time
sd
Results: filtering and toggling
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Filtering: strong input moderate input
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Toggling:
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Discussion
• bumps can exist in purely excitatory networks -inhibition is not necessary
– initiated by transient input
– go curve forms recruitment threshold
– localized by desychronization (Type I dynamics)
– sustained by self-coupling (or e.g. ICaL) anddesynchronization
• bump details are sensitive to parameters
• synaptic depression ⇒ band-pass filter, toggling
•OPEN: rigorous analysis, role of short-term spikefrequency changes, bump mechanism for Type II (D/E)
• IDEA: go curve/surface is useful for predicting theimpact of transient inputs
• IDEA: intrinsic dynamics can be important, andexcitatory networks can do interesting things
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• IDEA: go curve/surface is useful for predicting theimpact of transient inputs
• IDEA: intrinsic dynamics can be important, andexcitatory networks can do interesting things
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