Backtracking behaviour in lost ants: an additional strategy in their navigational toolkit

Ants use multiple sources of information to navigate, but do not integrate all this information into a unified representation of the world. Rather, the available information appears to serve three distinct main navigational systems: path integration, systematic search and the use of learnt information—mainly via vision. Here, we report on an additional behaviour that suggests a supplemental system in the ant''s navigational toolkit: ‘backtracking’. Homing ants, having almost reached their nest but, suddenly displaced to unfamiliar areas, did not show the characteristic undirected headings of systematic searches. Instead, these ants backtracked in the compass direction opposite to the path that they had just travelled. The ecological function of this behaviour is clear as we show it increases the chances of returning to familiar terrain. Importantly, the mechanistic implications of this behaviour stress an extra level of cognitive complexity in ant navigation. Our results imply: (i) the presence of a type of ‘memory of the current trip’ allowing lost ants to take into account the familiar view recently experienced, and (ii) direct sharing of information across different navigational systems. We propose a revised architecture of the ant''s navigational toolkit illustrating how the different systems may interact to produce adaptive behaviours.     

蚂蚁捷径返巢及其朝向机制的研究

定量研究了铺道蚁Tetramorium caespitum的载物返巢行为。实验证明这种蚂蚁在获得食物后,并不重复搜寻食物期间所走过的迂回路线,而是沿着近似直线的新路径迅速回巢。太阳及天空偏振光模式能用作它们返巢的提示,非偏振光源也能指引蚂蚁成功地捷径返巢。这些结果表明有高级社会性结构的铺道蚁通过脑以光导航。     

Optimal cue integration in ants

In situations with redundant or competing sensory information, humans have been shown to perform cue integration, weighting different cues according to their certainty in a quantifiably optimal manner. Ants have been shown to merge the directional information available from their path integration (PI) and visual memory, but as yet it is not clear that they do so in a way that reflects the relative certainty of the cues. In this study, we manipulate the variance of the PI home vector by allowing ants (Cataglyphis velox) to run different distances and testing their directional choice when the PI vector direction is put in competition with visual memory. Ants show progressively stronger weighting of their PI direction as PI length increases. The weighting is quantitatively predicted by modelling the expected directional variance of home vectors of different lengths and assuming optimal cue integration. However, a subsequent experiment suggests ants may not actually compute an internal estimate of the PI certainty, but are using the PI home vector length as a proxy.     

A new navigational mechanism mediated by ant ocelli

Many animals rely on path integration for navigation and desert ants are the champions. On leaving the nest, ants continuously integrate their distance and direction of travel so that they always know their current distance and direction from the nest and can take a direct path to home. Distance information originates from a step-counter and directional information is based on a celestial compass. So far, it has been assumed that the directional information obtained from ocelli contribute to a single global path integrator, together with directional information from the dorsal rim area (DRA) of the compound eyes and distance information from the step-counter. Here, we show that ocelli mediate a distinct compass from that mediated by the compound eyes. After travelling a two-leg outbound route, untreated foragers headed towards the nest direction, showing that both legs of the route had been integrated. In contrast, foragers with covered compound eyes but uncovered ocelli steered in the direction opposite to the last leg of the outbound route. Our findings suggest that, unlike the DRA, ocelli cannot by themselves mediate path integration. Instead, ocelli mediate a distinct directional system, which buffers the most recent leg of a journey.     

Behavioral ecology of odometric memories in desert ants: acquisition, retention, and integration

Assuming that the acquisition and retention of memories havecosts, properties of memories should fit the functional requirementsfor the system of memory. Based on a functional analysis ofwhat path integration is meant to do, we predicted that odometricmemories in desert ants should show (1) little improvement withrepeated training: performance should be as good after one trainingtrial as after six training trials, (2) decay of memory after24 h, and (3) performance based solely on the most recent outboundtrip, with no integration over multiple memories. Desert ants(Cataglyphis fortis) traveled in narrow straight plastic channelsto forage for cookie crumbs in a feeder at 6- or 12-m distance.Each ant was tested once by being taken from the feeder andreleased 2 m from the end of a 32-m channel to run home. Thedistance at which the ant first turned back (first turn) constitutedthe data. In acquisition, groups trained one or six times beforebeing tested had unsystematic scatter that did not differ significantly.In retention, ants tested after a 24-h delay showed larger unsystematicscatter than control animals tested after no delay. In integration,ants were trained five times at 6 or 12 m and then tested at12 or 6 m, respectively. No evidence of integration of multipleodometric memories was found. The results show that the propertiesof odometric memories are indeed tailored to what the memorysystem is used for.     

Long-term memory of individual identity in ant queens

Remembering individual identities is part of our own everyday social life. Surprisingly, this ability has recently been shown in two social insects. While paper wasps recognize each other individually through their facial markings, the ant, Pachycondyla villosa, uses chemical cues. In both species, individual recognition is adaptive since it facilitates the maintenance of stable dominance hierarchies among individuals, and thus reduces the cost of conflict within these small societies. Here, we investigated individual recognition in Pachycondyla ants by quantifying the level of aggression between pairs of familiar or unfamiliar queens over time. We show that unrelated founding queens of P. villosa and Pachycondyla inversa store information on the individual identity of other queens and can retrieve it from memory after 24h of separation. Thus, we have documented for the first time that long-term memory of individual identity is present and functional in ants. This novel finding represents an advance in our understanding of the mechanism determining the evolution of cooperation among unrelated individuals.