Characterizing neural dependencies with copula models

The coding of information by neural populations depends critically on the statistical dependencies between neuronal responses. However, there is no simple model that combines the observations that (1) marginal distributions over single-neuron spike counts are often approximately Poisson; and (2) joint distributions over the responses of multiple neurons are often strongly dependent. Here, we show that both marginal and joint properties of neural responses can be captured using Poisson copula models. Copulas are joint distributions that allow random variables with arbitrary marginals to be combined while incorporating arbitrary dependencies between them. Different copulas capture different kinds of dependencies, allowing for a richer and more detailed description of dependencies than traditional summary statistics, such as correlation coefficients. We explore a variety of Poisson 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 and macaque motor cortex, and quantify the improvement in coding accuracy afforded by incorporating the dependency structure between pairs of neurons.