We introduce a model for ant trail formation, building upon previous work on biologically feasible local algorithms that plausibly describe how ants maintain trail networks. The model is a variant of a reinforced random walk on a directed graph, where ants lay pheromone on edges as they traverse them and the next edge to traverse is chosen based on the level of pheromone; this pheromone decays with time. There is a bidirectional flow of ants in the network: the forward flow proceeds along forward edges from source (e.g. the nest) to sink (e.g. a food source), and the backward flow in the opposite direction. Some fraction of ants are lost as they pass through each node (modeling the loss of ants due to exploration observed in the field). We initiate a theoretical study of this model. We note that ant navigation has inspired the field of ant colony optimization, heuristics that have been applied to several combinatorial optimization problems; however the algorithms developed there are considerably more complex and not constrained to being biologically feasible. We first consider the linear decision rule, where the flow divides itself among the next set of edges in proportion to their pheromone level. Here, we show that the process converges to the path with minimum leakage when the forward and backward flows do not change over time. On the other hand, when the forward and backward flows increase over time (caused by positive reinforcement from the discovery of a food source, for example), we show that the process converges to the shortest path. These results are for graphs consisting of two parallel paths (a case that has been investigated before in experiments). Through simulations, we show that these results hold for more general graphs drawn from various random graph models; proving this convergence in the general case is an interesting open problem. Further, to understand the behaviour of other decision rules beyond the linear rule, we consider a general family of decision rules. For this family, we show that there is no advantage of using a non-linear decision rule, if the goal is to find the shortest or the minimum leakage path. We also show that bidirectional flow is necessary for convergence to such paths. Our results provide a plausible explanation for field observations, and open up new avenues for further theoretical and experimental investigation.

Creating a routing backbone is a fundamental problem in both biology and engineering. The routing backbone of the trail networks of arboreal turtle ants (Cephalotes goniodontus) connects many nests and food sources using trail pheromone deposited by ants as they walk. Unlike species that forage on the ground, the trail networks of arboreal ants are constrained by the vegetation. We examined what objectives the trail networks meet by comparing the observed ant trail networks with networks of random, hypothetical trail networks in the same surrounding vegetation and with trails optimized for four objectives: minimizing path length, minimizing average edge length, minimizing number of nodes, and minimizing opportunities to get lost. The ants' trails minimized path length by minimizing the number of nodes traversed rather than choosing short edges. In addition, the ants' trails reduced the opportunity for ants to get lost at each node, favoring nodes with 3D configurations most likely to be reinforced by pheromone. Thus, rather than finding the shortest edges, turtle ant trail networks take advantage of natural variation in the environment to favor coherence, keeping the ants together on the trails.

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.

Many dynamical networks, such as the ones that produce the collective behavior of social insects, operate without any central control, instead arising from local interactions among individuals. A well-studied example is the formation of recruitment trails in ant colonies, but many ant species do not use pheromone trails. We present a model of the regulation of foraging by harvester ant (Pogonomyrmex barbatus) colonies. This species forages for scattered seeds that one ant can retrieve on its own, so there is no need for spatial information such as pheromone trails that lead ants to specific locations. Previous work shows that colony foraging activity, the rate at which ants go out to search individually for seeds, is regulated in response to current food availability throughout the colony's foraging area. Ants use the rate of brief antennal contacts inside the nest between foragers returning with food and outgoing foragers available to leave the nest on the next foraging trip. Here we present a feedback-based algorithm that captures the main features of data from field experiments in which the rate of returning foragers was manipulated. The algorithm draws on our finding that the distribution of intervals between successive ants returning to the nest is a Poisson process. We fitted the parameter that estimates the effect of each returning forager on the rate at which outgoing foragers leave the nest. We found that correlations between observed rates of returning foragers and simulated rates of outgoing foragers, using our model, were similar to those in the data. Our simple stochastic model shows how the regulation of ant colony foraging can operate without spatial information, describing a process at the level of individual ants that predicts the overall foraging activity of the colony.