Emergent regulation of ant foraging frequency through a computationally inexpensive forager movement rule.

Ant colonies regulate foraging in response to their collective hunger, yet the mechanism behind this distributed regulation remains unclear. Previously, by imaging food flow within ant colonies we showed that the frequency of foraging events declines linearly with colony satiation (Greenwald et al., 2018). Our analysis implied that as a forager distributes food in the nest, two factors affect her decision to exit for another foraging trip: her current food load and its rate of change. Sensing these variables can be attributed to the forager's individual cognitive ability. Here, new analyses of the foragers' trajectories within the nest imply a different way to achieve the observed regulation. Instead of an explicit decision to exit, foragers merely tend toward the depth of the nest when their food load is high and toward the nest exit when it is low. Thus, the colony shapes the forager's trajectory by controlling her unloading rate, while she senses only her current food load. Using an agent-based model and mathematical analysis, we show that this simple mechanism robustly yields emergent regulation of foraging frequency. These findings demonstrate how the embedding of individuals in physical space can reduce their cognitive demands without compromising their computational role in the group.

Individual crop loads provide local control for collective food intake in ant colonies.

Nutritional regulation by ants emerges from a distributed process: food is collected by a small fraction of workers, stored within the crops of individuals, and spread via local ant-to-ant interactions. The precise individual-level underpinnings of this collective regulation have remained unclear mainly due to difficulties in measuring food within ants' crops. Here we image fluorescent liquid food in individually tagged Camponotus sanctus ants and track the real-time food flow from foragers to their gradually satiating colonies. We show how the feedback between colony satiation level and food inflow is mediated by individual crop loads; specifically, the crop loads of recipient ants control food flow rates, while those of foragers regulate the frequency of foraging-trips. Interestingly, these effects do not rise from pure physical limitations of crop capacity. Our findings suggest that the emergence of food intake regulation does not require individual foragers to assess the global state of the colony.