Theoretical and Computational Neuroscience

(Lecture 7)
 What We Have Learned!

Membrane potential
changes with time

2
 What We Have Learned!

τm is so fast that m changes essentially instantaneously to follow m∞(V), i.e. that m(t)=m∞(V(t)). This eliminates the differential
equation for dm/dt=0. Note that the time scale of m is fast compared to the membrane time constant τ=RC of a passive
membrane, which characterizes the evolution of the membrane voltage when all channels are closed.
 What We Have Learned!

Hippocampus
RTM Model (excitatory neurons) WB Model ( inhibitory neurons)

Erisir Model (inhibitory neurons )

Somatosensory cortex
 Linear Integrate-and-Fire (LIF) Neurons

Nearly half a century before Hodgkin and Huxley, in 1907, Louis Edouard Lapicque proposed a mathematical
model of nerve cells. Lapicque died in 1952, the year when the famous series of papers by Hodgkin and Huxley
appeared in print. Lapicque’s model is nowadays known as the integrate-and-fire neuron. We will refer to it as the LIF
neuron. Most authors take the L in “LIF” to stand for “leaky,” for reasons that will become clear shortly. We take it to
stand for “linear,” to distinguish it from the quadratic integrate-and-fire (QIF) neuron discussed in Chapter 8.

The LIF neuron is useful because of its utter mathematical simplicity. It can lead to insight, but reduced models such as
the LIF neuron are also dangerous — they can lead to incorrect conclusions.

Chapter 7

The LIF model can be described as follows. We assume that the ionic conductances are constant as long as the neuron does not fire, and that a spike is
triggered if the membrane potential rises to a certain threshold voltage. We don’t model the process of spike generation at all. As long as no spike is
triggered, the equation governing the membrane potential v is assumed to be

Constant!
 Linear Integrate-and-Fire (LIF) Neurons

As long as no spike is triggered, the equation governing the membrane potential v is assumed to be:

This can be written in the form:

Resting potential

Where:

The membrane time constant.

( v Na =E Na ,v K =E K ,v L =E L )
 Linear Integrate-and-Fire (LIF) Neurons

At the rest: I k +I Na +I cl=0

 Linear Integrate-and-Fire (LIF) Neurons
As long as no spike is triggered, the equation governing the membrane potential v is assumed to be:

t is the time that spike is triggered

threshold voltage
The above equation is supplemented by a condition of the form:
reset voltage

Here “v(t − 0)” denotes the left-sided limit of v at t, and “v(t + 0)” denotes the right-sided limit. The underlying
assumption is that a very rapid voltage spike occurs when v reaches v thr , and v then “resets” to a low value.

One calls the LIF neuron leaky if:

and non-leaky if :
 Linear Integrate-and-Fire (LIF) Neurons

Hodgkin-Huxley-like model:

LIF model:
 Linear Integrate-and-Fire (LIF) Neurons
This might suggest that the classical Hodgkin-Huxley neuron is close to non-leaky below the threshold voltage.

Hodgkin-Huxley-like model:

LIF model:

The membrane time constant is in fact very small,

below 1 ms, along the entire limit cycle!
Linear Integrate-and-Fire (LIF) Neurons

The membrane time constant is in fact very small, below 1 ms,

along the entire limit cycle! The classical Hodgkin-Huxley
has nearly linear voltage traces below threshold not because
the sodium, potassium, and leak currents are small below
threshold, but because their sum happens to be close to constant
between spikes( see the figure.) Note that when the sum of the
ion currents is approximately constant, dv/dt is approximately
constant, and therefore v is approximately a linear function of t.

I Na IK IL
 Linear Integrate-and-Fire (LIF) Neurons

LIF model:

To reduce the number of parameters in the LIF model, we shift and scale the voltage:

The reset potential is now and the threshold voltage is

LIF model:

The time has not been scaled — τ m is the same membrane time constant, measured in milliseconds, as before.
 Linear Integrate-and-Fire (LIF) Neurons

LIF model:

We will usually drop units when discussing the normalized LIF model, but keep in mind that t and τ m are times (measured in
ms), and therefore I has to be reciprocal time (measured in ms−1 ).
 Linear Integrate-and-Fire (LIF) Neurons

LIF model:

Exercise 2.7 !

v(t) eventually reaches the threshold

voltage 1 if and only ifτ m I>1 , i.e.,
I>1 / τ m .We call 1/ τ m the threshold
drive.
 Linear Integrate-and-Fire (LIF) Neurons
time it takes for v(t)
to reach 1

τ m I>1
ms
T

Hz

x=I
 Linear Integrate-and-Fire (LIF) Neurons

I=0.500023
 Linear Integrate-and-Fire (LIF) Neurons
I=0.500023
Exercise 4.7 !

The figure shows that during a large fraction of the period, v is very close to the threshold voltage, 1. As a result, the LIF
neuron will be highly sensitive to noisily fluctuating input: While v is nearly at threshold, any small input fluctuation can
cause a spike. This is true whenever T / τ m ≫1 .

When using the LIF model, τ m should therefore not be chosen much smaller than the largest period T we are
interested in to avoid excessive noise-sensitivity.

Since the most typical firing periods are on the order of tens of milliseconds, τ m should certainly not be much smaller
than 10 ms.
 Linear Integrate-and-Fire (LIF) Neurons

LIF model: H-H model:

Hodgkin-Huxley-like model neurons often do have very small membrane time constants, yet they are not extremely sensitive to noisy input!

In the LIF model, the subthreshold dynamics of a neuron are replaced by

one-dimensional linear dynamics. The LIF neuron is therefore not a good
model of neurons with more complex subthreshold dynamics. For example,
subthreshold voltage traces often have inflection points. A one-
dimensional linear equation describing subthreshold dynamics cannot
reproduce this feature. However, inflection points can be obtained by
making the subthreshold dynamics quadratic; this is the main topic of
Chapter 8.
 Course Presentation

Izhikevich neuron:

[80] E. M. Izhikevich, Resonate-and-fire neurons, Neural Networks, 14 (2001), pp. 883–894.

[81] E. M. Izhikevich, Simple model of spiking neurons, IEEE Transactions on Neural Networks, 14 (2003),
pp. 1569–1572.
 Quadratic Integrate-and-Fire (QIF) and Theta Neurons

v is a non-dimensionalized
Chapter 8 membrane potential

Exercise 8.1 !

The right-hand side of the above equation is positive for all v if and only if I > 1 . Thus 1
is the threshold drive here.
4 τm 4 τm
 Quadratic Integrate-and-Fire (QIF) and Theta Neurons

Exercise 8.2 !

v (0)= v 0
 Quadratic Integrate-and-Fire (QIF) and Theta Neurons

Changing coordinates:
 Quadratic Integrate-and-Fire (QIF) and Theta Neurons

Theta neuron (proposed by Ermentrout and Kopell):

The reset plays no role for subsequent calculations, since the right-hand side of the equation is periodic with period 2π; it can
therefore be omitted without consequence. Thus our model is now a differential equation, without any discontinuous reset.

1
I> the right-hand side of thee first equation is positive for all θ, and the point therefore moves around the
4 τm
circle indefinitely in the counter-clockwise direction

1
I< the right-hand side of the equation is zero at are fixed points
4 τm
 Quadratic Integrate-and-Fire (QIF) and Theta Neurons

Stable fixed point

1 fixed points
I<
4 τm
Halfstable (semistable) fixed point
Unstable fixed point

Unstable fixed point

< >

Stable fixed point

 Quadratic Integrate-and-Fire (QIF) and Theta Neurons

1− cos θ has no biological interpretation, but it has narrow

“spikes,” reaching its maxima when θ = π modulo 2π

Exercise 8.5 !
The firing period:
 Simple Pendulum

Dynamical Systems: Many engineering and natural systems are dynamical systems. For example a pendulum is a dynamical system.

State: The state of the dynamical system specifies it conditions. For a pendulum in the absence of external excitation shown in the figure, the
angle θ and the angular velocity uniquely define the state of the dynamical system.

The dynamical system is linear if the governing equation is linear. For the pendulum the dynamical system is nonlinear. This is the second order
nonlinear ODE describing the motion of the pendulum. The nonlinear pendulum equation can be reformulated as two first-order equations:

.
 Simple Pendulum

Phase Space: Plots of the state variables against one another are referred to as the phase space representation. Every point in
the phase space identifies a unique state of the system. For the pendulum, a plot of θ versus is the phase space representation.

θ
 A Geometry Way of Thinking

We will introduce one of the most basic techniques of dynamics: interpreting a differential equation as a vector field. Consider
the following nonlinear differential equation::

To evaluate the constant C, suppose that x=xo at t=O. Then :

A Geometry Way of Thinking
A Geometry Way of Thinking
A Geometry Way of Thinking
A Geometry Way of Thinking
 A Geometry Way of Thinking


Find fixed point Exercise !

●
Plot phase portrait

●
Plot qualitatively Q versus time
 Summary


Linear integrate-and-fire (LIF) neurons

●
Quadratic integrate-and-fire (QIF) neurons

●
Theta neurons

●
Introduction to dynamical systems

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 Next Lecture

In the next lecture we will cover:

●
Spike frequency adaptation (chapter 9)

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