The coding of information by neural populations depends critically on the statistical dependencies between neuronal responses. However, there is no simple model that can simultaneously account for (1) marginal distributions over single-neuron spike counts that are discrete and non-negative; and (2) joint distributions over the responses of multiple neurons that are often strongly dependent. Here, we show that both marginal and joint properties of neural responses can be captured using copula models. Copulas are joint distributions that allow random variables with arbitrary marginals to be combined while incorporating arbitrary dependencies be- tween them. Different copulas capture different kinds of dependencies, allowing for a richer and more detailed description of dependencies than traditional sum- mary statistics, such as correlation coefficients. We explore a variety of copula models for joint neural response distributions, and derive an efficient maximum likelihood procedure for estimating them. We apply these models to neuronal data collected in macaque pre-motor cortex, and quantify the improvement in cod- ing accuracy afforded by incorporating the dependency structure between pairs of neurons. We find that more than one third of neuron pairs shows dependency concentrated in the lower or upper tails for their firing rate distribution.

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