Solutions for the suggested exercises to accompany the Neuron Tutorial QUESTION 1: In 20 words or less, what is the relationship between what you see in window #4 and window #3? Why do you see action potentials in window #3 (Dendrite #1 Vm)? Ans: #4 Shows the Soma membrane potential and the action potentials arising in the soma. In #3 (Dendrite #1 Vm) we are seeing passive properties. The dendrite compartment is coupled through an axial resistance to the soma, and we see a smaller amplitude version of the soma Vm. QUESTION 2: In 10 words or less, what principle does the overlayed plot in window #2 demonstrate? (There is a two-word phrase for this.) Ans: Temporal summation. (The plots of the conductance of the Dendrite #1 excitatory channel show that for spike intervals of less than 3 milliseconds, the conductance is able to build up to a point where the Vm in the soma has increased to a value which will trigger action potentials.) QUESTION 3: Make a plot of the input-output transfer function. That is, plot output rate vs input spikes rate. Ans: There is a fairly sharp threshold for the generation of multiple action potentials at an input rate of about 300 spikes/sec. The curve tends to flatten out or saturate at input rates greater than 1000 spike/sec. This is somewhat similar to the sigmoid curve used to represent the input/output relation in "connectionist" artificial neural networks. QUESTION 4: Using only the "synaptic weight for dendrite #1 inhibitory input" and the three "Source B" parameters (delay, width and interval, inhibit (suppress) the middle of the three action potentials produced by "Source A" input. You may not modify any "Source A" parameters, and both the first and last action potentials must remain. Answer the question by stating the parameter values you had to use and by submitting hard copy of window #2 if your system allows you to make prints of the screen. Ans: The initial set of parameters, (Source A: delay=10, width=50, interval=2, Dend #1 Exc. Wt.=12) and (Source B: delay=20, width=50, interval=10, Dend #1 Inh. Wt.=12), had little effect upon the generation of action potentials. One solution of the problem is to reduce the source B burst width to 20 msec and to reduce the spike interval to 2 msec. QUESTION 5: Toggle "Plot Soma" to "Plot Dendrite 2". Move the inhibitory input ("Source B") to Dendrite #2. For the above configuration, is the inhibitory synapse more or less effective at suppressing the middle action potential? Defend your answer with numbers by varying the synaptic weights used in this configuration and the previous one. Give reasons for the results which you see. Ans: Many of the solutions for QUESTION 4 will have the same effect for this new configuration. However, to properly judge the relative effectiveness of the two configurations, we need to compare the minimum inhibitory synaptic weights or spike rates needed to suppress the middle spike. With a Source B spike rate of 2 msec, as in the solution above, a minimum weight of 8.5 will be needed to suppress the middle spike. In this new configuration, a weight of at least 9 is required, because the stimulus is applied further from the soma. Thus, the decrease in soma membrane potential will be less, due to the increased axial resistance and the shunting conductance in dentrite section #1. QUESTION 6: Reverse the inputs. That is, place the excitatory input on dendrite #2 and the inhibitory input on dendrite #1. For the above configuration, is the inhibitory synapse more or less effective at suppressing the middle action potential? Defend your answer in the same manner as in QUESTION 5, and explain the differences between this situation and the previous one. Ans: With a Source B spike interval of 2 msec as before, a synaptic weight of only 8 is sufficient to prevent the middle action potential from occuring. For the reasons given in QUESTION 5, the effect of the inhibition will be greater when it is applied closer to the soma. QUESTION 7: Using various numbers of inserted segments, with a sub-threshold epsp at dendrite section #2, estimate the length constant, "lambda", for this dendrite. Express your answer in units of "number of segments" and compare your result with the predictions of theory. Ans: Present an excitatory input to dendrite #2 from Source A with a weight of 10, using the default timing parameters (a spike interval of 10 msec). Compare the heights of the peaks of the membrane potential in the two dendrite sections by toggling the "Plot Soma/Plot Dend2" button to show the dendrite #2 membrane potential. Use the "scale" buttons on the graphs to pick an appropriate vertical scale (-75 to -65 mV). If we call the peak height V1 for dendrite #1 and V2 for dendrite #2, we should expect an attenuation with distance x given by V1/V2 = exp(-x/lambda). If we measure x in terms of the number of cable segments, N, we have x = N+1, since we have to count dendrite #2 as well. By progressively adding cable sections and plotting V1/V2 vs. x, we can find the value x=lambda at which V1/V2=0.37. Depending upon the care with which the peak heights are measured, this produces values of the attenuation constant roughly in the range of 9 to 11. Theory predicts that lambda is given by the square root of Rmem/Raxial. For the values given for the dendrites in the "Cell Parameters" menu, this predicts lambda=10.0.