Quantitative Methods in Neuroscience (neu 366M) Homework 6 Due: Tuesday October 21, by the beginning of class In this assignment, we’ll attempt to discover the unknown filter that a neuron applies to its inputs, from the known input stimulus and the measured neural response. 1) Recovering filters with the Wiener-Hopf equations. From the class website, download the Matlab script file generateWHdata.m. This file generates a white-noise input variable x, a filter h, and outputs xfilt and spks. The output xfilt is obtained by simply filtering x by h, whereas spks is a spike train generated from xfilt. You may think of xfilt as the sub-threshold voltage response of a cell to input x. a. From the Wiener-Hopf equations, we know that Cxy = C xx h where C xx is the autocorrelation, re-written as a Toeplitz matrix, and Cxy , h are vectors. For our previous work with the pseudoinverse, we know therefore that we can recover an estimate hinf of the filter h from the Wiener-Hopf equations by hinf = (C xx )−1 Cxy (1) where I’m using −1 to refer to the pseudoinverse. Let us use these insights to try to estimate the neural filter h from the input x and the (subthreshold) neural response y = xfilt. The filter is 200 time-steps long and causal, so M1 = 0 and M2 = 199. Recall that the correlation xcorr of a vector of length N with itself has length 2N − 1, and that the zero time-lag term is the N th term. Retain the appropriate terms of the autocorrelation, then use the command Toeplitz to convert these terms into a correlation matrix C xx of appropriate size. Compute the cross-correlation vector Cxy , also keeping only the appropriate terms. Normalize both correlation vectors by dividing by their lengths. Finally, use the \ command in Matlab (the backslash corresponds to the pseudoinverse operation in Matlab) to obtain hinf, through hinf = Cxx \Cxy . b. Evaluating the quality of the estimated filter hinf: Plot h (the ‘true’ filter) and hinf (the WH estimate of the filter) on the same plot: they should be in excellent agreement. Next, try to extract hinf by doing the same as in [a.], but with y = spks - mean(spks). That is, you are going to estimate the neural filter from its spiking output instead of its subthreshold response. The normalization of C xy should be by the number of spikes rather than the length of the vector. Compare the obtained filter against h. How good is the estimate? Why is it not as good as when y = xfilt? c. In this portion, we’ll evaluate when the spike-triggered average (STA) provides a good approximation to the filter estimate obtained from the WH equations. Recall that the spike-triggered average response of a neuron to a stimulus is simply the cross-correlation of the stimulus with the neural spike train: STA = Cxy Compare the mathematical expression for the STA with the WH filter estimate of Equation (1), to first infer under what conditions on C xx is the STA expected to produce a good approximation of the WH filter estimate? Next, compute the STA for the input and subthreshold response from [b.], and plot the STA together with the WH filter estimate hinf from the subthreshold response, as in part [b.]. They should be similar! Compute C xx in Matlab, to show that C xx does indeed look similar to what you’d expect, when the STA approximates the WH filter estimate. The input x used so far is called ‘white noise’: it is random from moment to moment, with no imposed continuity or correlation over time. d. Now set whitenoise=0 to generate a new stimulus x that is not ‘white’. It has temporal correlations in it (look at the autocorrelation, and contrast this autocorrelation with that from when x was white). Let y = xfilt (subthreshold response), and compute the WH estimate of the filter based on the non-white stimulus. Compare the result with the true filter h – it should still be an excellent fit. Finally, compute the STA estimate of the filter and the WH estimate of the filter based on the non-white stimulus. Are they similar? Using this observation, try to explain the role of the (C xx )−1 term in the Wiener-Hopf equation.
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