Insect navigation: do ants live in the now?

Visual navigation is a critical behaviour formanyanimals, and it has been particularly well studied in ants. Decades of ant navigation research have uncovered many ways in which efficient navigation can be implemented in small brains. For example, ants show us how visual information can drive navigation via procedural rather than map-like instructions. Two recent behavioural observations highlight interesting adaptive ways in which ants implement visual guidance. Firstly, it has been shownthat the systematic nest searches of ants can be biased by recent experience of familiar scenes. Secondly, ants have been observed to show temporary periods of confusion when asked to repeat a route segment, even if that route segment is very familiar. Taken together, these results indicate that the navigational decisions of ants take into account their recent experiences as well as the currently perceived environment

Visual navigation in ants

Les remarquables capacités de navigation des insectes nous prouvent à quel point ces " mini-cerveaux " peuvent produire des comportements admirablement robustes et efficaces dans des environnements complexes. En effet, être capable de naviguer de façon efficace et autonome dans un environnement parfois hostile (désert, forêt tropicale) sollicite l'intervention de nombreux processus cognitifs impliquant l'extraction, la mémorisation et le traitement de l'information spatiale préalables à une prise de décision locomotrice orientée dans l'espace. Lors de leurs excursions hors du nid, les insectes tels que les abeilles, guêpes ou fourmis, se fient à un processus d'intégration du trajet, mais également à des indices visuels qui leur permettent de mémoriser des routes et de retrouver certains sites alimentaires familiers et leur nid. L'étude des mécanismes d'intégration du trajet a fait l'objet de nombreux travaux, par contre, nos connaissances à propos de l'utilisation d'indices visuels sont beaucoup plus limitées et proviennent principalement d'études menées dans des environnements artificiellement simplifiés, dont les conclusions sont parfois difficilement transposables aux conditions naturelles. Cette thèse propose une approche intégrative, combinant 1- des études de terrains et de laboratoire conduites sur deux espèces de fourmis spécialistes de la navigation visuelle (Melophorus bagoti et Gigantiops destructor) et 2- des analyses de photos panoramiques prisent aux endroits où les fourmis naviguent qui permettent de quantifier objectivement l'information visuelle accessible à l'insecte. Les résultats convergents obtenus sur le terrain et au laboratoire permettent de montrer que, chez ces deux espèces, les fourmis ne fragmentent pas leur monde visuel en multiples objets indépendants, et donc ne mémorisent pas de 'repères visuels' ou de balises particuliers comme le ferait un être humain. En fait, l'efficacité de leur navigation émergerait de l'utilisation de paramètres visuels étendus sur l'ensemble de leur champ visuel panoramique, incluant repères proximaux comme distaux, sans les individualiser. Contre-intuitivement, de telles images panoramiques, même à basse résolution, fournissent une information spatiale précise et non ambiguë dans les environnements naturels. Plutôt qu'une focalisation sur des repères isolés, l'utilisation de vues dans leur globalité semble être plus efficace pour représenter la complexité des scènes naturelles et être mieux adaptée à la basse résolution du système visuel des insectes. Les photos panoramiques enregistrées peuvent également servir à l'élaboration de modèles navigationnels. Les prédictions de ces modèles sont ici directement comparées au comportement des fourmis, permettant ainsi de tester et d'améliorer les différentes hypothèses envisagées. Cette approche m'a conduit à la conclusion selon laquelle les fourmis utilisent leurs vues panoramiques de façons différentes suivant qu'elles se déplacent en terrain familier ou non. Par exemple, aligner son corps de manière à ce que la vue perçue reproduise au mieux l'information mémorisée est une stratégie très efficace pour naviguer le long d'une route bien connue ; mais n'est d'aucune efficacité si l'insecte se retrouve en territoire nouveau, écarté du chemin familier. Dans ces cas critiques, les fourmis semblent recourir à une seconde stratégie qui consiste à se déplacer vers les régions présentant une ligne d'horizon plus basse que celle mémorisée, ce qui généralement conduit vers le terrain familier. Afin de choisir parmi ces deux différentes stratégies, les fourmis semblent tout simplement se fier au degré de familiarisation avec le panorama perçu. Cette thèse soulève aussi la question de la nature de l'information visuelle mémorisée par les insectes. Le modèle du " snapshot " qui prédomine dans la littérature suppose que les fourmis mémorisent une séquence d'instantanés photographiques placés à différents points le long de leurs routes. A l'inverse, les résultats obtenus dans le présent travail montrent que l'information visuelle mémorisée au bout d'une route (15 mètres) modifie l'information mémorisée à l'autre extrémité de cette même route, ce qui suggère que la connaissance visuelle de l'ensemble de la route soit compactée en une seule et même représentation mémorisée. Cette hypothèse s'accorde aussi avec d'autres de nos résultats montrant que la mémoire visuelle ne s'acquiert pas instantanément, mais se développe et s'affine avec l'expérience répétée. Lorsqu'une fourmi navigue le long de sa route, ses récepteurs visuels sont stimulés de façon continue par une scène évoluant doucement et régulièrement au fur et à mesure du déplacement. Mémoriser un pattern général de stimulations, plutôt qu'une série de " snapshots " indépendants et très ressemblants les uns aux autres, constitue une hypothèse parcimonieuse. Cette hypothèse s'applique en outre particulièrement bien aux modèles en réseaux de neurones, suggérant sa pertinence biologique. Dans l'ensemble, cette thèse s'intéresse à la nature des perceptions et de la mémoire visuelle des fourmis, ainsi qu'à la manière dont elles sont intégrées et traitées afin de produire une réponse navigationnelle appropriée. Nos résultats sont aussi discutés dans le cadre de la cognition comparée. Insectes comme vertébrés ont résolu le même problème qui consiste à naviguer de façon efficace sur terre. A la lumière de la théorie de l'évolution de Darwin, il n'y a 'a priori' aucune raison de penser qu'il existe une forme de transition brutale entre les mécanismes cognitifs des différentes espèces animales. Le fossé marqué entre insectes et vertébrés au sein des sciences cognitives pourrait bien être dû à des approches différentes plutôt qu'à de vraies différences ontologiques. Historiquement, l'étude de la navigation de l'insecte a suivi une approche de type 'bottom-up' qui recherche comment des comportements apparemment complexes peuvent découler de mécanismes simples. Ces solutions parcimonieuses, comme celles explorées dans cette thèse, peuvent fournir de remarquables hypothèses de base pour expliquer la navigation chez d'autres espèces animales aux cerveaux et comportements apparemment plus complexes, contribuant ainsi à une véritable cognition comparée.Navigating efficiently in the outside world requires many cognitive abilities like extracting, memorising, and processing information. The remarkable navigational abilities of insects are an existence proof of how small brains can produce exquisitely efficient, robust behaviour in complex environments. During their foraging trips, insects, like ants or bees, are known to rely on both path integration and learnt visual cues to recapitulate a route or reach familiar places like the nest. The strategy of path integration is well understood, but much less is known about how insects acquire and use visual information. Field studies give good descriptions of visually guided routes, but our understanding of the underlying mechanisms comes mainly from simplified laboratory conditions using artificial, geometrically simple landmarks. My thesis proposes an integrative approach that combines 1- field and lab experiments on two visually guided ant species (Melophorus bagoti and Gigantiops destructor) and 2- an analysis of panoramic pictures recorded along the animal's route. The use of panoramic pictures allows an objective quantification of the visual information available to the animal. Results from both species, in the lab and the field, converged, showing that ants do not segregate their visual world into objects, such as landmarks or discrete features, as a human observers might assume. Instead, efficient navigation seems to arise from the use of cues widespread on the ants' panoramic visual field, encompassing both proximal and distal objects together. Such relatively unprocessed panoramic views, even at low resolution, provide remarkably unambiguous spatial information in natural environment. Using such a simple but efficient panoramic visual input, rather than focusing on isolated landmarks, seems an appropriate strategy to cope with the complexity of natural scenes and the poor resolution of insects' eyes. Also, panoramic pictures can serve as a basis for running analytical models of navigation. The predictions of these models can be directly compared with the actual behaviour of real ants, allowing the iterative tuning and testing of different hypotheses. This integrative approach led me to the conclusion that ants do not rely on a single navigational technique, but might switch between strategies according to whether they are on or off their familiar terrain. For example, ants can recapitulate robustly a familiar route by simply aligning their body in a way that the current view matches best their memory. However, this strategy becomes ineffective when displaced away from the familiar route. In such a case, ants appear to head instead towards the regions where the skyline appears lower than the height recorded in their memory, which generally leads them closer to a familiar location. How ants choose between strategies at a given time might be simply based on the degree of familiarity of the panoramic scene currently perceived. Finally, this thesis raises questions about the nature of ant memories. Past studies proposed that ants memorise a succession of discrete 2D 'snapshots' of their surroundings. Contrastingly, results obtained here show that knowledge from the end of a foraging route (15 m) impacts strongly on the behaviour at the beginning of the route, suggesting that the visual knowledge of a whole foraging route may be compacted into a single holistic memory. Accordingly, repetitive training on the exact same route clearly affects the ants' behaviour, suggesting that the memorised information is processed and not 'obtained at once'. While navigating along their familiar route, ants' visual system is continually stimulated by a slowly evolving scene, and learning a general pattern of stimulation rather than storing independent but very similar snapshots appears a reasonable hypothesis to explain navigation on a natural scale; such learning works remarkably well with neural networks. Nonetheless, what the precise nature of ants' visual memories is and how elaborated they are remain wide open question. Overall, my thesis tackles the nature of ants' perception and memory as well as how both are processed together to output an appropriate navigational response. These results are discussed in the light of comparative cognition. Both vertebrates and insects have resolved the same problem of navigating efficiently in the world. In light of Darwin's theory of evolution, there is no a priori reason to think that there is a clear division between cognitive mechanisms of different species. The actual gap between insect and vertebrate cognitive sciences may result more from different approaches rather than real differences. Research on insect navigation has been approached with a bottom-up philosophy, one that examines how simple mechanisms can produce seemingly complex behaviour. Such parsimonious solutions, like the ones explored in the present thesis, can provide useful baseline hypotheses for navigation in other larger-brained animals, and thus contribute to a more truly comparative cognition

Ants use a predictive mechanism to compensate for passive displacements by wind

SummaryInsect navigation is a fruitful system for analysing the ingenious and economical mechanisms that can underlie complex behaviour [1]. Past work has highlighted unsuspected navigational abilities in ants and bees, such as accurate path integration, long distance route following or homing from novel locations [2]. Here we report that ants can deal with one of the greatest challenges for any navigator: uncontrolled passive displacements. Foraging ants were blown by a jet of air over 3 meters into a dark pit. When subsequently released at windless unfamiliar locations, ants headed in the compass direction opposite to the one they had been blown away, thus functionally increasing their chance of returning to familiar areas. Ants do not appear to collect directional information during the actual passive displacement, but beforehand, while clutching the ground to resist the wind. During that time window, ants compute and store the compass direction of the wind. This is achieved by integrating the egocentric perception of the wind direction relative to their current body-axis with celestial compass information from their eyes

Route-following ants respond to alterations of the view sequence

Ants can navigate by comparing the currently perceived view with memorised views along a familiar foraging route. Models regarding route-following suggest the views are stored and recalled independently of the sequence in which they occur. Hence, the ant only needs to evaluate the instantaneous familiarity of the current view to obtain a heading direction. This study investigates whether ant homing behaviour is influenced by alterations in the sequence of views experienced along a familiar route, using the frequency of stop-and-scan behaviour as an indicator of the ant's navigational uncertainty. Ants were trained to forage between their nest and a feeder which they exited through a short channel before proceeding along the homeward route. In tests, ants were collected before entering the nest and released again in the channel, which was placed either in its original location or halfway along the route. Ants exiting the familiar channel in the middle of the route would thus experience familiar views in a novel sequence. Results show that ants exiting the channel scan significantly more when they find themselves in the middle of the route, compared to when emerging at the expected location near the feeder. This behaviour suggests that previously encountered views influence the recognition of current views, even when these views are highly familiar, revealing a sequence component to route memory. How information about view sequences could be implemented in the insect brain as well as potential alternative explanations to our results are discussed

Is the relationship among outcome variables shown in randomized trials?

BACKGROUND: Randomized controlled trials (RCTs) often have more than one primary outcome and frequently have secondary and harm outcomes. Comparison of outcomes between study arms is the primary focus of RCTs, but there are times when the relation between outcomes is important, such as determining whether an intermediate outcome and a clinical outcome have a strong association. We sought to determine how often reports of RCTs depict the relations among outcomes at the individual patient level and, for those studies that use composite outcomes, how often the relations between component elements are depicted. METHODS: We selected 20 general, specialty and subspecialty medical journals with high impact factors that publish original clinical research. We identified every RCT in the 2011 and 2012 issues and randomly selected 10 articles per journal. For each article we recorded the number of outcomes, the number of composite outcomes and how often the relations between outcomes or elements of composite outcomes were portrayed. RESULTS: All but 16 of the 200 RCTs had more than one outcome. Thus, outcomes could have been related in 92% of studies, but such relations were only reported in 2 (1%). A total of 33 (17%) investigations measured a composite outcome, 32 of which showed data for each component. None, however, showed cross-tabulation of the components. CONCLUSIONS: Readers are rarely shown the relation between outcomes. Mandatory posting of datasets or requirements for detailed appendices would allow readers to see these cross-tabulations, helping future investigators know which outcomes are redundant, which provide unique information and which are most responsive to changes in the independent variables. While not every relationship between outcomes requires depiction, at present such information is seldom portrayed

Opponent processes in visual memories: A model of attraction and repulsion in navigating insects’ mushroom bodies

International audienceSolitary foraging insects display stunning navigational behaviours in visually complex natural environments. Current literature assumes that these insects are mostly driven by attractive visual memories, which are learnt when the insect's gaze is precisely oriented toward the goal direction, typically along its familiar route or towards its nest. That way, an insect could return home by simply moving in the direction that appears most familiar. Here we show using virtual reconstructions of natural environments that this principle suffers from fundamental drawbacks, notably, a given view of the world does not provide information about whether the agent should turn or not to reach its goal. We propose a simple model where the agent continuously compares its current view with both goal and anti-goal visual memories, which are treated as attractive and repulsive respectively. We show that this strategy effectively results in an opponent process, albeit not at the perceptual level-such as those proposed for colour vision or polarisation detection-but at the level of the environmental space. This opponent process results in a signal that strongly correlates with the angular error of the current body orientation so that a single view of the world now suffices to indicate whether the agent should turn or not. By incorporating this principle into a simple agent navigating in reconstructed natural environments, we show that it overcomes the usual shortcomings and produces a step-increase in navigation effectiveness and robust-ness. Our findings provide a functional explanation to recent behavioural observations in ants and why and how so-called aversive and appetitive memories must be combined. We propose a likely neural implementation based on insects' mushroom bodies' circuitry that produces behavioural and neural predictions contrasting with previous models

Landmarks or panoramas: what do navigating ants attend to for guidance?

<p>Abstract</p> <p>Background</p> <p>Insects are known to rely on terrestrial landmarks for navigation. Landmarks are used to chart a route or pinpoint a goal. The distant panorama, however, is often thought not to guide navigation directly during a familiar journey, but to act as a contextual cue that primes the correct memory of the landmarks.</p> <p>Results</p> <p>We provided <it>Melophorus bagoti </it>ants with a huge artificial landmark located right near the nest entrance to find out whether navigating ants focus on such a prominent visual landmark for homing guidance. When the landmark was displaced by small or large distances, ant routes were affected differently. Certain behaviours appeared inconsistent with the hypothesis that guidance was based on the landmark only. Instead, comparisons of panoramic images recorded on the field, encompassing both landmark and distal panorama, could explain most aspects of the ant behaviours.</p> <p>Conclusion</p> <p>Ants navigating along a familiar route do not focus on obvious landmarks or filter out distal panoramic cues, but appear to be guided by cues covering a large area of their panoramic visual field, including both landmarks and distal panorama. Using panoramic views seems an appropriate strategy to cope with the complexity of natural scenes and the poor resolution of insects' eyes. The ability to isolate landmarks from the rest of a scene may be beyond the capacity of animals that do not possess a dedicated object-perception visual stream like primates.</p

Crucial role of ultraviolet light for desert ants in determining direction from the terrestrial panorama

Ants use the panoramic skyline in part to determine a direction of travel. A theoretically elegant way to define where terrestrial objects meet the sky is to use an opponent-process channel contrasting green wavelengths of light with ultraviolet (UV) wavelengths. Compared with the sky, terrestrial objects reflect relatively more green wavelengths. Using such an opponent-process channel gains constancy in the face of changes in overall illumination level. We tested the use of UV wavelengths in desert ants by using a plastic that filtered out most of the energy below 400 nm. Ants, Melophorus bagoti, were trained to home with an artificial skyline provided by an arena (experiment 1) or with the natural panorama (experiment 2). On a test, a homing ant was captured just before she entered her nest, and then brought back to a replicate arena (experiment 1) or the starting point (the feeder, experiment 2) and released. Blocking UV light led to deteriorations in orientation in both experiments. When the artificial skyline was changed from opaque to transparent UV-blocking plastic (experiment 3) on the other hand, the ants were still oriented. We conclude that UV wavelengths play a crucial role in determining direction based on the terrestrial surround.10 page(s

How do field of view and resolution affect the information content of panoramic scenes for visual navigation? A computational investigation

The visual systems of animals have to provide information to guide behaviour and the informational requirements of an animal’s behavioural repertoire are often reflected in its sensory system. For insects, this is often evident in the optical array of the compound eye. One behaviour that insects share with many animals is the use of learnt visual information for navigation. As ants are expert visual navigators it may be that their vision is optimised for navigation. Here we take a computational approach in asking how the details of the optical array influence the informational content of scenes used in simple view matching strategies for orientation. We find that robust orientation is best achieved with low-resolution visual information and a large field of view, similar to the optical properties seen for many ant species. A lower resolution allows for a trade-off between specificity and generalisation for stored views. Additionally, our simulations show that orientation performance increases if different portions of the visual field are considered as discrete visual sensors, each giving an independent directional estimate. This suggests that ants might benefit by processing information from their two eyes independently

Running paths to nowhere: repetition of routes shows how navigating ants modulate online the weights accorded to cues

Ants are expert navigators, keeping track of the vector to home as they travel, through path integration, and using terrestrial panoramas in view-based navigation. Although insect learning has been much studied, the learning processes in navigation have not received much attention. Here, we investigate in desert ants (Melophorus bagoti) the effects of repeating a well-travelled and familiar route segment without success. We find that re-running a homeward route without entering the nest impacted subsequent trips. Over trips, ants showed more meandering from side to side and more scanning behaviour, in which the ant stopped and turned, rotating to a range of directions. In repeatedly re-running their familiar route, ants eventually gave up heading in the nestward direction as defined by visual cues and turned to walk in the opposite direction. Further manipulations showed that the extent and rate of this path degradation depend on (1) the length of the vector accumulated in the direction opposite to the food-to-nest direction, (2) the specific visual experience of the repeated segment of the route that the ants were forced to re-run, and (3) the visual panorama: paths are more degraded in an open panorama, compared with a visually cluttered scene. The results show that ants dynamically modulate the weighting given to route memories, and that fits well with the recent models, suggesting that the mushroom bodies provide a substrate for the reinforcement learning of views for navigation