Abstract
Recent developments in automated tracking allow uninterrupted, high-resolution recording of animal trajectories, sometimes coupled with the identification of stereotyped changes of body pose or other behaviors of interest. Analysis and interpretation of such data represents a challenge: the timing of animal behaviors may be stochastic and modulated by kinematic variables, by the interaction with the environment or with the conspecifics within the animal group, and dependent on internal cognitive or behavioral state of the individual. Existing models for collective motion typically fail to incorporate the discrete, stochastic, and internal-state-dependent aspects of behavior, while models focusing on individual animal behavior typically ignore the spatial aspects of the problem. Here we propose a probabilistic modeling framework to address this gap. Each animal can switch stochastically between different behavioral states, with each state resulting in a possibly different law of motion through space. Switching rates for behavioral transitions can depend in a very general way, which we seek to identify from data, on the effects of the environment as well as the interaction between the animals. We represent the switching dynamics as a Generalized Linear Model and show that: (i) forward simulation of multiple interacting animals is possible using a variant of the Gillespie’s Stochastic Simulation Algorithm; (ii) formulated properly, the maximum likelihood inference of switching rate functions is tractably solvable by gradient descent; (iii) model selection can be used to identify factors that modulate behavioral state switching and to appropriately adjust model complexity to data. To illustrate our framework, we apply it to two synthetic models of animal motion and to real zebrafish tracking data.
Highlights
The three behavioral states that we introduce for this purpose are: (i) a run state, where the bacteria move at a constant speed and direction; (ii+iii) left/right tumble states, where the bacteria are stationary but rotate to the left or right at a constant angular velocity
We introduced a novel approach for understanding the behavior of individual animals or groups of interacting animals
The probabilistic model of animal behavior combines deterministic dynamics, which describe the motion of animals at each of the possible behavioral states, and stochastic switching between these states
Summary
The combination of stochastic state transitions with deterministic laws of motion through space distinguishes our model from the classical models where behavioral rules are assumed fixed in time Such rules are inferred by fitting to all available data, resulting in simulated trajectories which tend to be much smoother than real data. [23], [24], and [25] distinguish behavioral states in ant foraging due to either multiple food sources or due to recruiting of the resting ants While these works describe different “behavioral states”, they do so in terms of population averaged quantities, while our goal is to incorporate a set of different behaviors at the individual level and fit that to individual trajectory data. The second half of the paper focuses on three examples: two synthetic toy models of animal behavior (ant motion, bacterial chemotaxis) to showcase and validate the inference as well as illustrate an interesting extension to coarse-grained behavioral states, and one real data example (tracked zebrafish data) to illustrate inference without prior knowledge of laws of motion, as well as model selection to identify kinematic variables that affect behavioral state switching
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