Implications of hierarchical Markov models of behavior: on irreversibility, predictability, and dimensionality

The maturation of quantitative tools for studying the high-level structure of animal behavior, and especially tools which represent spontaneous behavior as a sequence of stereotyped and neurally well-defined 'syllables', demands that the field revisit a fundamental theoretical question: if the coarse structure of behavior can be accurately described by Markov models, what do these models really tell us about behavior? In this work, we explore the theoretical implications of these models and discuss how they allow us to quantitatively formulate questions about the sequence-like nature and effective dimensionality of behavior. One important insight is that the eigenvalues and eigenvectors of various model-associated matrices furnish interpretable time scales and modifications of behavior that occur on those time scales. We illustrate our points using both toy examples and Markov models fit to real data. By analyzing the consequences of Markov representations, we clarify the theoretical meaning of progress in quantifying behavior.