Past exam of the mathematics course of the University of Cambridge 2006 iii Paper 84 4 a Solution Created 2026-10-03 Updated 2026-10-14
Fit a stimulus-conditioned linear-nonlinear-Poisson cascade model. Choose a causal stimulus-history window, subtract the stimulus mean, and estimate a temporal filter . A useful initial estimate is the spike-triggered average, . For white Gaussian stimulation and a one-filter response, this estimates the filter direction. With correlated Gaussian stimulation it estimates a direction proportional to , where is the history covariance matrix; use stimulus-correlated spike-triggered averaging to whiten or regularize this correction. An arbitrary stimulus or an even response nonlinearity does not guarantee that this average identifies the correct model.
A general fitting approach is to expand a causal filter in smooth basis functions, compute , and take a nonnegative rate . With , fit the coefficients by Poisson spike-train likelihood fitting:The first term rewards rate at observed spikes and the second penalizes overprediction elsewhere. Include a filter-size or smoothness penalty, and choose complexity on held-out data. Another option estimates by stimulus-projection histograms: divide the density of projection values at spikes by their occupancy density, multiplied by the mean firing rate. Avoid interpreting poorly sampled bins as reliable rates. Temporal train/test splits should avoid leakage through the filter-history window.
Given the fitted rate and the same stimulus, generate an Inhomogeneous Poisson process. For small bins of width , spike with probability , choosing the bins sufficiently short that multiple events are negligible. More exactly, advance integrated intensity until it increases by an independent unit-exponential random variable, then place the next event. Repeating this generates different synthetic spike trains with the same fitted stimulus-dependent rate, not copies of the original event times.
Compare held-out log-likelihood, stimulus-conditioned firing-rate predictions, count distributions, interspike-interval distributions and autocorrelation. With repeated identical stimulus presentations, compare trial-averaged time courses and trial-to-trial count variance, including the Fano factor. A single recorded train does not by itself supply a reliable repeated-trial average or variance; use held-out temporal data, model-based uncertainty and further recordings instead. Time-rescaled spike-train diagnostics test whether integrated-intensity intervals look independently exponential, rather than comparing raw intervals to an exponential law under a changing rate.
A successful model reproduces specified statistical features on new data; identical spike times are not the goal. The basic Poisson model can miss refractoriness, bursting, adaptation, history dependence or stimulus-unexplained trial variability. If needed, add a spike-history filter to the conditional rate, including an absolute dead time, and fit the resulting history-dependent model. A complex cell may also require multiple filters or quadratic stimulus features: its spike-triggered mean can vanish despite strong tuning. These failures identify model limitations rather than evidence that a random spike generator should match every observed interval.
Refractory limits on neuronal firing 2026-10-14
If every interspike interval is at least , a sustained spike train has mean firing rate at most . Across a window of duration , the event count satisfies ; dividing by and taking the long-time limit gives the bound. The extra one accounts for a possible first event. The bound permits much slower firing and says nothing about whether stronger drive can maintain the limiting rate in a real neuron.
For the chosen phenomenological paired-pulse recovery, the hold alone gives . Add the further assumption that every successful event resets precisely the same recovery curve, with no accumulation from earlier events. Then onset-timed pulses can sustain at most in that model. This repeatable reset is an extra assumption: a two-pulse result alone cannot establish it. A real neuron's sustained rate also depends on neural adaptation, input delivery and evolving ion-channel inactivation. The existing leaky integrate-and-fire model similarly adds integration time to its imposed hold.
A homogeneous Poisson process of rate has interspike intervals with an exponential distribution, soWith the purely illustrative choices and , this is . Thus about of intervals predicted by that model violate this chosen hold. Matching a mean spike train count does not make a stimulus-only Poisson process respect recent firing.
A minimal alternative is a renewal process with independent and identically distributed intervals , where has an exponential distribution of rate . Its hazard function is zero during the hold and afterwards. Its mean interval is and long-run firing rate isThe same rate after a hold gives , not . This separate stochastic example has no gradual relative recovery; it illustrates why an available-state rate differs from an observed mean firing rate. Smooth dependence on time since the last event can extend the hazard function, while longer histories require more than this simple renewal process. Time-rescaled spike-train diagnostics can test the resulting history-dependent predictions against recorded spike trains.