Unbalanced visual cues do not affect search precision at the nest in desert ants (Cataglyphis nodus).

Desert ant foragers are well known for their visual navigation abilities, relying on visual cues in the environment to find their way along routes back to the nest. If the inconspicuous nest entrance is missed, ants engage in a highly structured systematic search until it is discovered. Searching ants continue to be guided by visual cues surrounding the nest, from which they derive a location estimate. The precision level of this estimate depends on the information content of the nest panorama. This study examines whether search precision is also affected by the directional distribution of visual information. The systematic searching behavior of ants is examined under laboratory settings. Two different visual scenarios are compared - a balanced one where visual information is evenly distributed, and an unbalanced one where all visual information is located on one side of an experimental arena. The identity and number of visual objects is similar over both conditions. The ants search with comparable precision in both conditions. Even in the visually unbalanced condition, searches are characterized by balanced precision on both sides of the arena. This finding lends support to the idea that ants memorize the visual scenery at the nest as panoramic views from different locations. A searching ant is thus able to estimate its location with equal precision in all directions, leading to symmetrical search paths.

The abundance, biomass, and distribution of ants on Earth.

Knowledge on the distribution and abundance of organisms is fundamental to understanding their roles within ecosystems and their ecological importance for other taxa. Such knowledge is currently lacking for insects, which have long been regarded as the "little things that run the world". Even for ubiquitous insects, such as ants, which are of tremendous ecological significance, there is currently neither a reliable estimate of their total number on Earth nor of their abundance in particular biomes or habitats. We compile data on ground-dwelling and arboreal ants to obtain an empirical estimate of global ant abundance. Our analysis is based on 489 studies, spanning all continents, major biomes, and habitats. We conservatively estimate total abundance of ground-dwelling ants at over 3 × 1015 and estimate the number of all ants on Earth to be almost 20 × 1015 individuals. The latter corresponds to a biomass of ∼12 megatons of dry carbon. This exceeds the combined biomass of wild birds and mammals and is equivalent to ∼20% of human biomass. Abundances of ground-dwelling ants are strongly concentrated in tropical and subtropical regions but vary substantially across habitats. The density of leaf-litter ants is highest in forests, while the numbers of actively ground-foraging ants are highest in arid regions. This study highlights the central role ants play in terrestrial ecosystems but also major ecological and geographic gaps in our current knowledge. Our results provide a crucial baseline for exploring environmental drivers of ant-abundance patterns and for tracking the responses of insects to environmental change.

Transfer of Visual Learning Between a Virtual and a Real Environment in Honey Bees: The Role of Active Vision.

To study visual learning in honey bees, we developed a virtual reality (VR) system in which the movements of a tethered bee walking stationary on a spherical treadmill update the visual panorama presented in front of it (closed-loop conditions), thus creating an experience of immersion within a virtual environment. In parallel, we developed a small Y-maze with interchangeable end-boxes, which allowed replacing repeatedly a freely walking bee into the starting point of the maze for repeated decision recording. Using conditioning and transfer experiments between the VR setup and the Y-maze, we studied the extent to which movement freedom and active vision are crucial for learning a simple color discrimination. Approximately 57% of the bees learned the visual discrimination in both conditions. Transfer from VR to the maze improved significantly the bees' performances: 75% of bees having chosen the CS+ continued doing so and 100% of bees having chosen the CS- reverted their choice in favor of the CS+. In contrast, no improvement was seen for these two groups of bees during the reciprocal transfer from the Y-maze to VR. In this case, bees exhibited inconsistent choices in the VR setup. The asymmetric transfer between contexts indicates that the information learned in each environment may be different despite the similar learning success. Moreover, it shows that reducing the possibility of active vision and movement freedom in the passage from the maze to the VR impairs the expression of visual learning while increasing them in the reciprocal transfer improves it. Our results underline the active nature of visual processing in bees and allow discussing the developments required for immersive VR experiences in insects.

How to Navigate in Different Environments and Situations: Lessons From Ants.

Ants are a globally distributed insect family whose members have adapted to live in a wide range of different environments and ecological niches. Foraging ants everywhere face the recurring challenge of navigating to find food and to bring it back to the nest. More than a century of research has led to the identification of some key navigational strategies, such as compass navigation, path integration, and route following. Ants have been shown to rely on visual, olfactory, and idiothetic cues for navigational guidance. Here, we summarize recent behavioral work, focusing on how these cues are learned and stored as well as how different navigational cues are integrated, often between strategies and even across sensory modalities. Information can also be communicated between different navigational routines. In this way, a shared toolkit of fundamental navigational strategies can lead to substantial flexibility in behavioral outcomes. This allows individual ants to tune their behavioral repertoire to different tasks (e.g., foraging and homing), lifestyles (e.g., diurnal and nocturnal), or environments, depending on the availability and reliability of different guidance cues. We also review recent anatomical and physiological studies in ants and other insects that have started to reveal neural correlates for specific navigational strategies, and which may provide the beginnings of a truly mechanistic understanding of navigation behavior.

Using virtual reality to study visual performances of honeybees.

Virtual reality (VR) offers an appealing experimental framework for studying visual performances of insects under highly controlled conditions. In the case of the honeybee Apis mellifera, this possibility may fill the gap between behavioural analyses in free-flight and cellular analyses in the laboratory. Using automated, computer-controlled systems, it is possible to generate virtual stimuli or even entire environments that can be modified to test hypotheses on bee visual behaviour. The bee itself can remain tethered in place, making it possible to record neural activity while the bees is performing behavioural tasks. Recent studies have examined visual navigation and attentional processes in VR on flying or walking tethered bees, but experimental paradigms for examining visual learning and memory are only just emerging. Behavioural performances of bees under current experimental conditions are often lower in VR than in natural environments, but further improvements on current experimental protocols seem possible. Here we discuss current developments and conclude that it is essential to tailor the specifications of the VR simulation to the visual processing of honeybees to improve the success of this research endeavour.

Associative visual learning by tethered bees in a controlled visual environment.

Free-flying honeybees exhibit remarkable cognitive capacities but the neural underpinnings of these capacities cannot be studied in flying insects. Conversely, immobilized bees are accessible to neurobiological investigation but display poor visual learning. To overcome this limitation, we aimed at establishing a controlled visual environment in which tethered bees walking on a spherical treadmill learn to discriminate visual stimuli video projected in front of them. Freely flying bees trained to walk into a miniature Y-maze displaying these stimuli in a dark environment learned the visual discrimination efficiently when one of them (CS+) was paired with sucrose and the other with quinine solution (CS-). Adapting this discrimination to the treadmill paradigm with a tethered, walking bee was successful as bees exhibited robust discrimination and preferred the CS+ to the CS- after training. As learning was better in the maze, movement freedom, active vision and behavioral context might be important for visual learning. The nature of the punishment associated with the CS- also affects learning as quinine and distilled water enhanced the proportion of learners. Thus, visual learning is amenable to a controlled environment in which tethered bees learn visual stimuli, a result that is important for future neurobiological studies in virtual reality.

Follower ants in a tandem pair are not always naïve.

In addition to foraging individually several species of ants guide nestmates to a goal by tandem running. We found that the Australian ant, Camponotus consobrinus, forages both individually and by tandem running to head to the same goal, nest-specific native Australian trees on which they forage. While paths of solitary foragers and initial paths of tandem followers showed no differences in heading directions or straightness, tandem followers moved at about half the speed of solitary runs. When leaders were experimentally removed, follower ants initially engaged in a systematic search around the point of interruption, following which they either (a) headed directly towards and successfully reached the foraging trees, or (b) continued searching or (c) returned to the nest. The high incidence of followers that successfully navigated towards the foraging trees on their own provides strong evidence that many tandem followers are in fact experienced foragers. Detailed analysis of the searching behaviour revealed that even seemingly lost followers displayed a directional bias towards the foraging trees in their search path. Our results show that in a foraging context follower ants in a tandem pair are not always naïve.

Multiple sources of celestial compass information in the Central Australian desert ant Melophorus bagoti.

The Central Australian desert ant Melophorus bagoti is known to use celestial cues for compass orientation. We manipulated the available celestial cues for compass orientation for ants that had arrived at a feeder, were captured and then released at a distant test site that had no useful terrestrial panoramic cues. When tested in an enclosed transparent box that blocked some or most of the ultraviolet light, the ants were still well oriented homewards. The ants were again significantly oriented homewards when most of the ultraviolet light as well as the sun was blocked, or when the box was covered with tracing paper that eliminated the pattern of polarised light, although in the latter case, their headings were more scattered than in control (full-cue) conditions. When the position of the sun was reflected 180° by a mirror, the ants headed off in an intermediate direction between the dictates of the sun and the dictates of unrotated cues. We conclude that M. bagoti uses all available celestial compass cues, including the pattern of polarised light, the position of the sun, and spectral and intensity gradients. They average multiple cues in a weighted fashion when these cues conflict.

Does the Australian desert ant Melophorus bagoti approximate a Lévy search by an intrinsic bi-modal walk?

We suggest that the Australian desert ant Melophorus bagoti approximates a Lévy search pattern by using an intrinsic bi-exponential walk and does so when a Lévy search pattern is advantageous. When attempting to locate its nest, M. bagoti adopt a stereotypical search pattern. These searches begin at the location where the ant expects to find the nest, and comprise loops that start and end at this location, and are directed in different azimuthal directions. Loop lengths are exponentially distributed when searches are in visually familiar surroundings and are well described by a mixture of two exponentials when searches are in unfamiliar landscapes. The latter approximates a power-law distribution, the hallmark of a Lévy search. With the aid of a simple analytically tractable theory, we show that an exponential loop-length distribution is advantageous when the distance to the nest can be estimated with some certainty and that a bi-exponential distribution is advantageous when there is considerable uncertainty regarding the nest location. The best bi-exponential search patterns are shown to be those that come closest to approximating advantageous Lévy looping searches. The bi-exponential search patterns of M. bagoti are found to approximate advantageous Lévy search patterns.

Beginnings of a synthetic approach to desert ant navigation.

In a synthetic approach to studying navigational abilities in desert ants, we review recent work comparing ants living in different visual ecologies. Those living in a visually rich habitat strewn with tussocks, bushes, and trees are compared to those living in visually barren salt pans, as exemplified by the Central Australian Melophorus bagoti and the North African Cataglyphis fortis, respectively. In bare habitats the navigator must rely primarily on path integration, keeping track of the distance and direction in which it has travelled, while in visually rich habitats the navigator can rely more on guidance by the visual panorama. Consistent with these expectations, C. fortis performs better than M. bagoti on various measures of precision at path integration. In contrast, M. bagoti learned a visually based associative task better than C. fortis, the latter generally failing at the task. Both these ants, however, exhibit a similar pattern of systematic search as a 'back up' strategy when other navigational strategies fail. A newly investigated salt-pan species of Melophorus (as yet unnamed) resembles C. fortis more, and its congener M. bagoti less, in its path integration. The synthetic approach would benefit from comparing more species chosen to address evolutionary questions. This article is part of a Special Issue entitled: CO3 2013.

Information content of visual scenes influences systematic search of desert ants.

Many animals - including insects - navigate visually through their environment. Solitary foraging desert ants are known to acquire visual information from the surrounding panorama and use it to navigate along habitual routes or to pinpoint a goal such as the nest. Returning foragers that fail to find the nest entrance engage in searching behaviour, during which they continue to use vision. The characteristics of searching behaviour have typically been investigated in unfamiliar environments. Here we investigated in detail the nest-searching behaviour of Melophorus bagoti foragers within the familiar visual environment of their nest. First, by relating search behaviour to the information content of panoramic (360 deg) images, we found that searches were more accurate in visually cluttered environments. Second, as observed in unfamiliar visual surrounds, searches were dynamic and gradually expanded with time, showing that nest pinpointing is not rigidly controlled by vision. Third, contrary to searches displayed in unfamiliar environments, searches observed here could be modelled as a single exponential search strategy, which is similar to a Brownian walk, and there was no evidence of a Lévy walk. Overall, our results revealed that searching behaviour is remarkably flexible and varies according to the relevance of information provided by the surrounding visual scenery.

Visual cue learning and odometry in guiding the search behavior of desert ants, Melophorus bagoti, in artificial channels.

Terrestrial panoramic cues, path integration and search behavior are the main navigational strategies used by ants to locate food and find their way back to the nest. Searching becomes important when the other navigational cues are either not available or cannot provide sufficient information to pinpoint the goal. When searching in one-dimensional channels Melophorus bagoti ants exhibit a systematic drift in the starting-point-to-goal direction as they turn back and forth, sometimes past the goal location (Narendra et al., 2008). Here we show that this drift in channels is not a stereotypical part of the search behavior in these ants. It rather depends on the conditions of training. In experiments in which the nest entrance is located not at the end but at the side of the channel, forward drift is not always part of the nest search. Experiments on food searches showed that with the food source at the end of the channel, ants performed a linear drift in the starting-point-to-food direction. With food at the side of the channel, they showed a less pronounced drift toward the food source. In this constrained environment, especially with the goal at the end of the channel, ants seem to learn a routine such as 'run along the channel', and mix this routine with their usual strategy of turning back and forth in search.

Views, landmarks, and routes: how do desert ants negotiate an obstacle course?

The Australian desert ant Melophorus bagoti often follows stereotypical routes through a cluttered landscape containing both distant panoramic views and obstacles (plants) to navigate around. We created an artificial obstacle course for the ants between a feeder and their nest. Landmarks comprised natural objects in the landscape such as logs, branches, and tussocks. Many ants travelled stereotypical routes home through the obstacle course in training, threading repeatedly the same gaps in the landmarks. Manipulations altering the relations between the landmarks and the surrounding panorama, however, affected the routes in two major ways. Both interchanging the positions of landmarks (transpositions) and displacing the entire landmark set along with the starting position of the ants (translations) (1) reduced the stereotypicality of the route, and (2) increased turns and meanders during travel. The ants might have used the entire panorama in view-based travel, or the distal panorama might prime the identification and use of landmarks en route. Despite the large data set, both options (not mutually exclusive) remain viable.