An effective hydrodynamic description of marching locusts.

A fundamental question in complex systems is how to relate interactions between individual components ('microscopic description') to the global properties of the system ('macroscopic description'). Furthermore, it is unclear whether such a macroscopic description exists and if such a description can capture large-scale properties. Here, we address the validity of a macroscopic description of a complex biological system using the collective motion of desert locusts as a canonical example. One of the world's most devastating insect plagues begins when flightless juvenile locusts form 'marching bands'. These bands display remarkable coordinated motion, moving through semiarid habitats in search of food. We investigated how well macroscopic physical models can describe the flow of locusts within a band. For this, we filmed locusts within marching bands during an outbreak in Kenya and automatically tracked all individuals passing through the camera frame. We first analyzed the spatial topology of nearest neighbors and found individuals to be isotropically distributed. Despite this apparent randomness, a local order was observed in regions of high density in the radial distribution function, akin to an ordered fluid. Furthermore, reconstructing individual locust trajectories revealed a highly aligned movement, consistent with the one-dimensional version of the Toner-Tu equations, a generalization of the Navier-Stokes equations for fluids, used to describe the equivalent macroscopic fluid properties of active particles. Using this effective Toner-Tu equation, which relates the gradient of the pressure to the acceleration, we show that the effective 'pressure' of locusts increases as a linear function of density in segments with the highest polarization (for which the one-dimensional approximation is most appropriate). Our study thus demonstrates an effective hydrodynamic description of flow dynamics in plague locust swarms.

Information integration for decision-making in desert locusts.

Locust swarms can extend over hundred kilometers, and starvation compels this ancient pest to devour everything in its path. Theory suggests that gregarious behavior benefits foraging efficiency, yet the role of social cohesion remains elusive. To this end, we collected high-resolution trajectories of individual and grouped gregarious desert locusts in a 2-choice assay with patches of either similar or different quality. Carefully maintaining animals' identities allowed us to monitor each individual's experience and estimate the leaky accumulation process of personally acquired and socially derived evidence. We fitted data to a Bayesian model to gain insight into the decision-making system for patch selection. By disentangling the relative contribution of each information class, our study suggests that locusts balance incongruent evidence but reinforce congruent ones. We provide insight into the collective foraging decisions of social (but non-eusocial) insects and present locusts as a powerful empirical system to study individual choices and consequent collective dynamics.

Ants prefer the option they are trained to first.

The temporal order in which experiences occur can have a profound influence on their salience. Humans and other vertebrates usually memorise the first and last items of a list most readily. Studies on serial position learning in insects, mainly in bees, showed preference for last encountered items. In bees, pheromone presence can also influence motivation, and thus learning. However, neither serial position learning nor the effect of recruitment pheromones on learning have been well investigated in ants. We trained Lasius niger ants to make multiple visits to sucrose on a runway which alternated between lemon or rosemary odour, and the presence or absence of trail pheromone, and then tested for preference between the odours on a Y-maze, in order to investigate the effect of pheromone presence on learning. Pheromone presence did not affect ant choice. However, unexpectedly, the ants strongly preferred the first odour encountered. This was explored by the addition of a familiarisation visit without pheromone or odour. The familiarisation visit disabled or reversed this preference for the first odour encountered, with ants now mostly taking their 'default' preference by choosing the left side of the maze. Our study found no effect of trail pheromone on learning, but a strong yet fragile preference for the first odour experienced. These different preferences could lead to spatial segregation of foraging activity depending on prior experience and might facilitate efficient resource exploitation by colonies.

Nitrogen fixation by diverse diazotrophic communities can support population growth of arboreal ants.

BACKGROUND: Symbiotic ant-plant associations, in which ants live on plants, feed on plant-provided food, and protect host trees against threats, are ubiquitous across the tropics, with the Azteca-Cecropia associations being amongst the most widespread interactions in the Neotropics. Upon colonization of Cecropia's hollow internodes, Azteca queens form small patches with plant parenchyma, which are then used as waste piles when the colony grows. Patches-found in many ant-plant mutualisms-are present throughout the colony life cycle and may supplement larval food. Despite their initial nitrogen (N)-poor substrate, patches in Cecropia accommodate fungi, nematodes, and bacteria. In this study, we investigated the atmospheric N2 fixation as an N source in patches of early and established ant colonies. RESULTS: Via 15N2 tracer assays, N2 fixation was frequently detected in all investigated patch types formed by three Azteca ant species. Quantified fixation rates were similar in early and established ant colonies and higher than in various tropical habitats. Based on amplicon sequencing, the identified microbial functional guild-the diazotrophs-harboring and transcribing the dinitrogenase reductase (nifH) gene was highly diverse and heterogeneous across Azteca colonies. The community composition differed between early and established ant colonies and partly between the ant species. CONCLUSIONS: Our data show that N2 fixation can result in reasonable amounts of N in ant colonies, which might not only enable bacterial, fungal, and nematode growth in the patch ecosystems but according to our calculations can even support the growth of ant populations. The diverse and heterogeneous diazotrophic community implies a functional redundancy, which could provide the ant-plant-patch system with a higher resilience towards changing environmental conditions. Hence, we propose that N2 fixation represents a previously unknown potential to overcome N limitations in arboreal ant colonies.

Trail Pheromone Does Not Modulate Subjective Reward Evaluation in Lasius niger Ants.

Comparing the value of options is at the heart of economic decision-making. While an option may have an absolute quality (e.g. a food source has a fixed energy content), the perceived value of the option may be malleable. The factors affecting the perceived value of an option may thus strongly influence which option is ultimately chosen. Expectations have been shown to be a strong driver of perceived value in both humans and social insects, causing an undervaluation of a given option if a better option was expected, and an overvaluation if a poorer one was expected. In humans, perceived value can be strongly affected by social information. Value perception in some insects has also been shown to be affected by social information, showing conformism as in humans and other animals. Here, over a series of experiments, we tested whether pheromone trail presence, a social information source, influenced the perceived value of a food source in the ant Lasius niger. We found that the presence of pheromone trails leading to a sucrose solution does not influence food acceptance, pheromone deposition when returning from a food source, drinking time, or frequency of U-turns on return from the food. Two further assays for measuring changes in food acceptance, designed to increase sensitivity by avoiding ceiling effects, also showed no effect of pheromone presence on food acceptance. In a separate study, L. niger have also been found to show no preference for, or avoidance of, odors associated with foods found in the presence of pheromone. We are thus confident that trail pheromone presence does not affect the perceived value of a food source in these ants.

Meat ants cut more trail shortcuts when facing long detours.

Engineered paths increase efficiency and safety but also incur construction and maintenance costs, leading to a trade-off between investment and gain. Such a trade-off is faced by Australian meat ants, which create and maintain vegetation-free trails between nests and food sources, and thus their trails are expected to be constructed selectively. To test this, we placed an artificial obstacle consisting of 300 paper grass blades between a sucrose feeder and the colony, flanked by walls either 10 cm or 80 cm long. To exploit the feeder, ants could detour around the walls or take a direct route by traversing through the obstacle. We found that, when confronted with a long alternative detour, 76% of colonies removed more grass blades and ants were also 60% more likely to traverse the obstacle instead of detouring, with clearing activity favouring higher ant flow or vice versa. An analysis of cut patterns revealed that ants did not cut randomly, but instead concentrated on creating a trail to the food source. Meat ants were thus able to collectively deploy their trail-clearing efforts in a directed manner when detour costs were high, and rapidly established cleared trails to the food source by focusing on completing a central, vertically aligned trail which was then followed by the ants.

Transmission of fungal partners to incipient Cecropia-tree ant colonies.

Ascomycete fungi in the nests of ants inhabiting plants (= myrmecophytes) are very often cultivated by the ants in small patches and used as food source. Where these fungi come from is not known yet. Two scenarios of fungus recruitment are possible: (1) random infection through spores or hyphal fragments from the environment, or (2) transmission from mother to daughter colonies by the foundress queen. It is also not known at which stage of the colony life cycle fungiculture is initiated, and whether the- symbiont fungi serve as food for the ant queen. To clarify these questions, we investigated four Azteca ant species inhabiting three different Cecropia species (C. insignis, C. obtusifolia, and C. peltata). We analysed an rRNA gene fragment from 52 fungal patches produced by founding queens and compared them with those from established Azteca colonies (n = 54). The infrabuccal pockets of winged queens were dissected to investigate whether young queens carry fungi from their mother colony. Additionally, 15N labelling experiments were done to verify whether the queen feeds on the patches until she is nourished by her first worker offspring. We infer from the results that the fungi cultivated in hollow plant structures are transferred from the parental colony of the young queen. First, fungal genotypes/OTU diversity was not significantly different between foundress queen patches and established colonies, and second, hyphal parts were discovered in the infrabuccal pockets of female alates. We could show that fungiculture already starts before queens lay their eggs, and that the queens do not feed on fungal patch material but feed it to the larvae. Our findings suggest that fungiculture may be crucial for successful colony founding of arboreal ants in the tropics.