Active contributions to computation Dendrites as computational elements Examples Dendritic computation r V m I m R m Current flows uniformly out through the cell I m I ID: 915996
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Slide1
Dendritic computation
Slide2Passive contributions to computation
Active contributions to computation
Dendrites as computational elements:
Examples
Dendritic
computation
Slide3r
V
m
=
I
m
Rm
Current flows uniformly out through the cell:
I
m
=
I
0
/4pr2
Input resistance is defined as RN = Vm(t∞)/I0 = Rm/4pr2
Injecting current I
0
Geometry matters: the
isopotential cell
Slide4r
m
and
r
i
are the membrane and axial resistances, i.e.the resistances of a thin slice of the cylinder
Linear cable theory
Slide5r
i
r
m
c
m
For a length L of membrane cable:
r
i
r
i
L
r
m
r
m
/ L
c
m
c
m
L
Axial and membrane resistance
Slide6(1)
(2)
(1)
or
where
Time constant
Space constant
The cable equation
Slide70
Decay of voltage in space for current injection at
x
=0, T
∞
+
I
ext
(x,t
)
Slide8 Electrotonic length
Properties of passive cables
Slide9Johnson and Wu
Electrotonic
length
Slide10Properties of passive cables
Electrotonic
length
Current can escape through additional pathways: speeds up decay
Slide11Johnson and Wu
Voltage rise time
Current can escape through additional pathways: speeds up decay
Slide12Koch
Imp
ulse
response
Slide13General solution as a filter
Slide14Step response
Slide15 Electrotonic length
Current can escape through additional pathways: speeds up decay
Cable diameter affects input resistance
Properties of passive cables
Slide16 Electrotonic length
Current can escape through additional pathways: speeds up decay
Cable diameter affects input resistance
Cable diameter affects transmission velocity
Properties of passive cables
Slide17Step response
Slide18Step response
Slide19Other factors
Finite cables
Active channels
Slide20Rall
model
Impedance matching:
If a
3/2
= d
1
3/2 + d23/2
can collapse to an equivalent
c
ylinder with length given
b
y
electrotonic
length
Slide21Active
conductances
New cable equation for each
dendritic
compartment
Slide22Who’ll be my
Rall
model, now that my
Rall
model is gone, gone
Genesis, NEURON
Slide23Passive computations
London and Hausser, 2005
Slide24Linear filtering:
Inputs from dendrites are broadened and delayed
Alters summation properties..
coincidence detection to temporal integration
Segregation of inputs
Nonlinear interactions within a dendrite
--
sublinear
summation
-- shunting inhibition
Delay lines Dendritic inputs “labelled”Passive computations
Slide25Spain;
Scholarpedia
Delay lines: the sound localization circuit
Slide26Passive computations
London and Hausser, 2005
Slide27Mechanisms to deal with the distance dependence of PSP size
Subthreshold
boosting: inward currents with reversal near rest
Eg
persistent Na+
Synaptic scaling
Dendritic spikes
Na
+
, Ca
2+
and NMDA
Dendritic
branches as
mini computational units backpropagation: feedback circuit Hebbian
learning throughsupralinear interaction of backprop
spikes with inputs
Active dendrites
Slide28Segregation and amplification
Slide29Segregation and amplification
Slide30Segregation and amplification
The single neuron as a neural network
Slide31Currents
Potential
Distal: integration
Proximal: coincidence
Magee, 2000
Synaptic scaling
Slide32Synaptic potentials
Somatic action potentials
Magee, 2000
Expected distance dependence
Slide33CA1 pyramidal neurons
Slide34Passive properties
Slide35Passive properties
Slide36Active properties: voltage-gated channels
Na
+
, Ca
2+ or NDMA receptor block eliminates supralinearity
For short intervals (0-5ms), summation is linear or slightly supralinear
For longer intervals (5-100ms), summation is sublinearIh
and K
+
block eliminates
sublinear
temporal summation
Slide37Active properties: voltage-gated channels
Major player in synaptic scaling:
hyperpolarization
activated K current, I
h
Increases in density down the dendrite
Shortens
EPSP duration, reduces local summation
Slide38Synaptic properties
While active properties contribute to summation, don’t explain normalized amplitude
Shape of EPSC determines how it is filtered .. Adjust ratio of AMPA/NMDA receptors
Slide39Rall
; fig London and
H
ausser
Direction selectivity
Slide40Back-propagating action potentials
Slide41Johnson and Wu, Foundations of Cellular Physiology
, Chap 4
Koch,
Biophysics of Computation
Magee, Dendritic
integration of excitatory synaptic input, Nature Reviews Neuroscience, 2000London and Hausser,
Dendritic Computation, Annual Reviews in Neuroscience, 2005
References