Theoretical Neuroscience - Assignment 1

Gergo Bohner 11 October 2013

The Hodgkin-Huxley neuron

1.a 200nA/mm2 stimulus

The initial conditions for the simulation were equilibrium values of resting nonstimulated neuron, and are as follows: V (0) = 65mV n(0) = 0.3178 m(0) = 0.0530 h(0) = 0.5958
Membrane potential with 200 nA/mm stimulus
60 1 n m h 0.9 40 0.8 20

Gating variables with 200 nA/mm stimulus

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Pr(Ion channel open)

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1.b Equilibrium values

The expressions for the equilibrium values can be given in the form x (V ) = x (V ) x (V )+x (V ) for x = n, m, h .

Equilibrium values of gating variables

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1.c Firing rate

The ring rate simulations use the same resting state initial conditions as Section 1.a, and the ring rate is measured via calculating the number of spikes in the 200-1000 ms period, after the intial perturbations settle. An other possibility is to calculate r = 1/(tspikei+1 tspikei ) at large i values to get the ring rate, but due to the numerical errors it is safer to just count the spikes.
Firing rates
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1.d-e Conductance changes

In both cases, by decreasing gK or increasing gN a we only make the derivative of the voltage more positive as in most cases V EK > 0 and V EN a < 0. Therefore the ring starts already at lower stimulation and at the same stimulation the ring rate is higher.
Firing rates (varying g K)
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gK = 0.36 mS/mm2 gNa = 0.30 mS/mm2 gK = 0.24 mS/mm2

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gNa = 1.2 mS/mm gNa = 1.3 mS/mm gNa = 1.4 mS/mm

2 2 2

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The linear integrate and re neuron

dV dt

2.a Constant input current C = gL (V EL ) + I0

v = V EL , V0 = I0 /gL , = C/gL dv dt dv V0 v log (V0 v ) v V a =? V (t1 ) = Vt Vt t1 t1 r = = = = = V (0) = Vr t1 =? = = = = V0 + EL (Vr V0 EL )et/ Vt V0 EL log( ) Vr V0 EL C Vt I0 /gL EL log( ) gL Vr I0 /gL EL 1/t1 v + V0 1 dt t +a t V0 + ae t V0 + EL + ae a = Vr V0 EL

The logarithm function log(1+x) is linear for small x-es, so we expect that for high I0 -s the ring rate should grow linearly, and it does:

LIF neuron model, constant current

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2.b Sinusoid input current

This is a slightly more complicated case, but still can be computed analitically (or can be simulated by using some tricks to add the spikes). The same notation is used as in part a. The analitical solution uses the integrating factor (/ exp( P (x)dx)) method as well as integration by parts two times on trigonometric functions to return to the same integral form and compute the nal integral. Afterwards to nd the minimal value of V0 which creates repetitive spiking, one has to nd the maximum of the periodic V (t) function, and see if it reaches the Vt spike threshold: dv 1 V0 + v= sin(t) dt V0 d t/ (e v (t)) = sin(t)et/ dt V0 et/ v (t) = sin(t)et/ dt = X V0 1 1 1 X= cos(t)et/ cos(t) et/ = tau V 1 1 V0 1 0 = cos(t)et/ sin(t)et/ + 2 2 X tau tau 2 / exp( 1/ dt)

 1 X (1 + 2 2 ) v (t) = = V0 (sin(t) 2 2 +1 cos(t)) V0 1 V0 1 cos(t)et/ + sin(t)et/ tau tau 2

max(v (t)) d max((sin(t) cos(t))) dt tan1 (t) t = = =

max((sin(t) cos(t))) 0 1 (n tan1 (1/ ))/ n Z

max(v (t))

= >

V0 2 2 +

2 2 + 1 =

V0 2 2

+1

spike V0
20 Membrane potential V0 threshold 10 Spike threshold

v (t) > Vt EL = V V 2 2 + 1

LIF neuron model, increasing sinusoid current

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Nullclines

3.a Sketch nullclines

The stability of the equilibrium points is discussed later.

Simplified HH, Nullcline sketch

1 Vnullcline hnullcline (Vh=60) 0.9 hnullcline (Vh=50) hnullcline (Vh=40)

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3.b Condition on Vh
To nd a correct condition we need to move the green (Vh = 60mV) curve slightly up (i.e. increasing Vh ) so that it is tangent to the minima of the Vnullcline in the V > 50mVregion. Therefore we rst need to identify this minimum. If we model m(V ) as a step function, we get m(50 + dV ) = 1 which 15+dV leads to a minimum h value of h = 50 dV 0.3 to stay on the V-nullcline. Therefore we can safely nd the Vh value corresponding to the (50, 0.3) point in the set of h (50, Vh ) functions. From this, Vh = 10 log( 01 .3 1) 50 = 58.473mV.

3.c-d Trajectories
Simplified HH, Nullcline sketch
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Simplified HH, Nullcline sketch

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