Regulation of harvester ant foraging as a closed-loop excitable system.

Ant colonies regulate activity in response to changing conditions without using centralized control. Desert harvester ant colonies forage for seeds, and regulate foraging to manage a tradeoff between spending and obtaining water. Foragers lose water while outside in the dry air, but ants obtain water by metabolizing the fats in the seeds they eat. Previous work shows that the rate at which an outgoing forager leaves the nest depends on its recent rate of brief antennal contacts with incoming foragers carrying food. We examine how this process can yield foraging rates that are robust to uncertainty and responsive to temperature and humidity across minute-to-hour timescales. To explore possible mechanisms, we develop a low-dimensional analytical model with a small number of parameters that captures observed foraging behavior. The model uses excitability dynamics to represent response to interactions inside the nest and a random delay distribution to represent foraging time outside the nest. We show how feedback from outgoing foragers returning to the nest stabilizes the incoming and outgoing foraging rates to a common value determined by the volatility of available foragers. The model exhibits a critical volatility above which there is sustained foraging at a constant rate and below which foraging stops. To explain how foraging rates adjust to temperature and humidity, we propose that foragers modify their volatility after they leave the nest and become exposed to the environment. Our study highlights the importance of feedback in the regulation of foraging activity and shows how modulation of volatility can explain how foraging activity responds to conditions and varies across colonies. Our model elucidates the role of feedback across many timescales in collective behavior, and may be generalized to other systems driven by excitable dynamics, such as neuronal networks.

Fast and slow processes underlie the selection of both step frequency and walking speed.

People prefer gaits that minimize their energetic cost. Research focused on step frequency selection suggests that a fast predictive process and a slower optimization process underlie this energy optimization. Our purpose in this study was to test whether the mechanisms controlling step frequency selection are used more generally to select one of the most relevant characteristics of walking - preferred speed. To accomplish this, we contrasted the dynamic adjustments in speed following perturbations to step frequency against the dynamic adjustments in step frequency following perturbations to speed. Despite the use of different perturbations and contexts, we found that the responses were very similar. In both experiments, subjects responded to perturbations by first rapidly changing their speed or step frequency towards their preferred pattern, and then slowly adjusting their gait to converge onto their preferred pattern. We measured similar response times for both the fast processes (1.4±0.3 versus 2.7±0.6 s) and the slow processes (74.2±25.4 versus 79.7±20.2 s). We also found that the fast process, although quite variable in amplitude, dominated the adjustments in both speed and step frequency. These distinct but complementary experiments demonstrate that people appear to rely heavily on prediction to rapidly select the most relevant aspects of their preferred gait and then gradually fine-tune that selection, perhaps using direct optimization of energetic cost.