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Fit a stimulus-conditioned linear-nonlinear-Poisson cascade model. Choose a causal stimulus-history window, subtract the stimulus mean, and estimate a temporal filter $k$. A useful initial estimate is the spike-triggered average, $\hat{a}(u)=N^{-1}{\sum }_i[s(t_i-u)-\bar{s}]$. For white Gaussian stimulation and a one-filter response, this estimates the filter direction. With correlated Gaussian stimulation it estimates a direction proportional to $Ck$, where $C$ 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 $a(t)=b+{\int }_0^{U}k(u)s(t-u)du$, and take a nonnegative rate $\lambda (t)=f(a(t))$. With $f(a)=e^a$, 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 $f$ 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 $\Delta t$, spike with probability $1-e^{-\lambda (t)\Delta t}$, 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.

A signed spatial filter model of a simple cell assigns a weight $K_{ij}$ to each sampled location and takes its dot product with the visual contrast image $C_{ij}$. Positive weights favor bright features; negative weights favor dark features. The signed driveis a linear function of the image, while the rectified linear unit output $R$ is nonlinear. $D$ and $R$ below are arbitrary model units. To obtain a predicted firing rate one would need a calibrated gain, background response and possibly a different output function. The model does not generate a spike train; a linear-nonlinear-Poisson cascade model adds a stochastic event-generation assumption.

For the first diagram, every one of five rows has weightsA bright central column, with $C_{i3}=1$ and zero elsewhere, gives $D=5/5=1$. A bright middle row gives $D=-1/20-1/20+1/5-1/20-1/20=0$; a uniformly bright patch also gives $D=0$, since every row sums to zero. Move the bright column one site right: $D=5(-1/20)=-1/4$, so $R=0$. Make that shifted column dark instead: $D=1/4$ and $R=1/4$. Thus an OFF-like weight can increase output for darkness. A negative weight is a descriptive sensitivity, not evidence for a particular inhibitory synapse.

For a smooth example use the real, even Gabor filterThe Gaussian function localizes the weights; the cosine supplies alternating polarity. Choose ${\sigma }_x=1/2$, ${\sigma }_y=1$ and $q=\pi$ in arbitrary spatial units. The stripes' spatial frequency is $q/(2\pi )=1/2$ cycles per unit. Their axis makes angle $\theta$ with horizontal, while brightness changes across the perpendicular axis. $\phi$ is their spatial phase. Normalize the full-plane integral of $K{C}_{\theta ,\phi }$ by its value at $\theta =\pi /2$, $\phi =0$. The Gaussian integral and the product identity for cosines givewhereAt $\phi =0$, horizontal stripes give $R\approx 0.004159$, $60^{\circ }$ stripes give $R\approx 0.286741$, and vertical stripes give $R=1$. At the same vertical orientation, $\phi =\pi /2$ gives $D=R=0$, while reversing the polarity with $\phi =\pi$ gives $D=-1$ and $R=0$. The preferred axis persists while the response changes dramatically with position. Shifting the vertical stripes by a quarter of their wavelength changes phase by $\pi /2$ up to sign, without rotating them.

These are two different toy filters, not discretizations of the same fitted receptive field. The discrete filter exactly rejects a uniform contrast patch; this Gabor filter need not reject uniform contrast because its weight integral need not vanish. Neither toy includes stimulus history, recurrent interactions, saturation or a measured response baseline. Their value is a transparent prediction: orientation selectivity can emerge from spatial summation, and it does not by itself imply tolerance to phase.