Lecture 21 Notes, Tuesday 29 November 2005
Class notes on integrate-and-fire model neurons and thalamocortical neurons
Homework due: Tues 06 Dec 2005
In today's lecture we discuss the integrate-and-fire (IF) model neuron, where neuronal excitation is reduced to one state variable. This is a useful model because of ease of computation and simplicity. We will use the IF model as the basis for a thalamocortical (TC) neuron which is a very interesting type of cell that shows low threshold spikes (LTS) due to T-type calcium current. The TC cell will be modeled using the integrate-and-fire-and-burst (IFB) variation of the basic IF model.
# Integrate, fire, and burst model
#
init v=-64.99
init h=0.002
v'=(iapp-gl*(v-vl)-gt*minf(v)*h*(v-vt))/c
h'=if(v>vh)then(-h/tauhminus)else((1-h)/tauhplus)
global 1 v-vtheta {v=-50}
minf(v)=heav(v-vh)
iapp=step1+step2*heav(t-ton)*heav(toff-t)
par step1=0,step2=1
par ton=50, toff=350
par vtheta=-45
par vl=-65
par gl=0.035
par c=2
par vh=-70
par tauhminus=20
par tauhplus=100
par gt=0.2
par vt=120
@ dt=0.05,total=700,maxstor=10000000
@ xlo=0,ylo=-80,xhi=700,yhi=-40
done
Here are links to the files you can load and use in XPP:
Integrate-fire-and-burst neuron (IFB_model.ode)
tonic settings
post-inhibitory rebound settings
low-threshold calcium spike settings
Here are the three types of behavior in the IFB model, which can be run using the three settings files you can load into XPP for the model "IFB_model.ode" (above).
Homework (Due Tues 06 Dec 2005) answer key for Homework #10
"Square-wave" bursting behavior in the Morris-Lecar model is possible using the parameter set for type I dynamics, i.e., where oscillations emerge with zero frequency and then the parameter φ is raised to 0.23. These parameters for the model are listed in Figure 7.7 of the chapter. Rather than use Rinzel and Ermentrouts exact equation for the slow calcium-dependent potassium current I used:
dI/dt = ε (V0 - V),
where &epsilon = 0.05, and V0 = -26 mV.
So the full Morris-Lecar system is:
CV′ = Iapp - gCam∞(V)(V - ECa) - gKw(V - EK) - gL(V - EL)
w′ = φ ([m∞(V) - m]/Τm)
I′ = ε (V0 - V).
Here is the ODE file for the bursting system with three state variables ML_SqWav_3D.ode.
(1) Simulate bursting in this model for 4000 ms, plotting membrane potential as a function of time.
(2) Use XPP-AUT to calculate the bifurcation diagram for the model using the applied current as the bifurcation parameter. Make sure you edit the file so that there are just two state variables (V,w) to make the bifurcation diagram. You can use the original version of the model, which did not include the slowly changing current described here. Construct the bifurcation diagram in two stages: first calculate the equilibrium solutions. Next "grab" the Hopf bifurcation (Auto specifies this point with "HB") and then calculate the periodic solutions.
(3) Over what range of applied current does the model exhibit multistability?
(4) Does the slow current increase or decrease during the active (spiking) phase of the rhythmic bursts?
(5) Make two other time series to demonstrate that bursting oscillations can be made slower or faster by changing ε.
(6) Explain why the last several action potentials during the active phase of each burst have more space, i.e., time, between them compared to the first several spikes at the onset of each burst.
(7) In the diagram below:
Name the bifurcation that gives rise to oscillations as the steady-state destabilizes.
Label all the branches of fixed points, classifying their stability and whether they are nodes or spirals.
Identify any regions of bistability.
Identify the separatrix/separatrices between regions of bistability.
Label the branches of limit cycles and whether they are stable or unstable periodic orbits.
Identify the "saddle-node of limit cycles bifurcation" in the diagram.
Is there a homoclinic orbit in this bifurcation diagram? If so, where is it?
Sketch the phase planes (nullclines, fixed points, and periodic orbits) that correspond to the bifurcation diagram at Iapp = 90, 95, and 105 pA/μm2.
(8) In the diagram below:
Name the bifurcation that gives rise to oscillations as the steady-state destabilizes.
Label all the branches of fixed points, classifying their stability and whether they are nodes or spirals.
Identify any regions of bistability.
Identify the separatrix/separatrices between regions of bistability.
Label the brances of limit cycles and whether they are stable or unstable periodic orbits.
Identify the "saddle-node of limit cycles bifurcation" in the diagram.
Is there a homoclinic orbit in this bifurcation diagram? If so, where is it?
Sketch the phase plane at Iapp = 35 pA/μm2. Label all fixed points, draw heteroclinic orbits, and draw the stable and unstable manifolds of the saddle point (hint: some of these terms apply to the same structures).
At Iapp = 35 pA/μm2 the system is 'excitable', i.e., the membrane supports full amplitude single spikes for suprathreshold stimuli before returning to the stable point attractor or resting membrane potential. Why are the spikes called 'full amplitude' or amplitude preserving. Re-read the Rinzel and Ermentrout chapter if you need a refresher.
At Iapp = 35 pA/μm2 is there a separatrix present? If so, what is the separatrix and what does it separate in terms of solutions to the Morris-Lecar system?
(9) In the diagrams below (one is a zoomed-in version):
What features of this diagram are exactly the same as the one above (in #8)?
What features of this diagram are different than the one above (in #8)?
What parameter in the Morris-Lecar model will transform the diagram in #8 to the diagram here in #9? Explain how this parameter affects the solutions to the system.
Label all the branches of fixed points, classifying their stability and whether they are nodes or spirals.
Identify any regions of multistability.
Identify the separatrix/separatrices between regions of multistability.
Label the branches of limit cycles and whether they are stable or unstable periodic orbits.
Identify the "saddle-node of limit cycles bifurcation" in the diagram.
Is there a homoclinic orbit in this bifurcation diagram? If so, where is it? What is the special name for the homoclinic orbit in this case?
Sketch the phase plane at Iapp = 35 pA/μm2. Label all fixed points as well as stable and unstable periodic orbits. Draw the stable and unstable manifolds for the saddle point (use XPP with a small Dt to get the program to draw the manifolds for you, hint: use singular points 'Sing pts' and then 'draw invariant sets'.
Lecture 21 Top - Cellular Biophysics and Modeling -
Del Negro Homepage - Applied Science - The College of William and Mary