The learning walks of ants (Hymenoptera: Formicidae)

When transitioning from in-nest duties to their foraging life outside the nest, ants perform a series of highly choreographed learning walks around the nest entrance, before leaving to forage for the first time. These learning walks have been described in detail only for a few species of ants, but a pattern of similarities and differences is emerging that we review here with an emphasis on understanding the functional significance of this learning process for efficient homing in ants. We compare the organization of learning walks in ants with that of the learning flights in bees and wasps and provide a list of key research questions that would need to be tackled if we are to understand the role of learning walks in the acquisition of nest-location information, the evolution of this highly conserved learning process, and how it is controlled.We acknowledge financial support to JZ from the Australian Research Council’s Discovery Program (DP150101172 and DP150102699) and to PNF from a Scientia-Scholarship, University of Würzburg, and the Deutsche Forschungsgemeinschaft (project FL1060/1-1)

Migration from Teleoperation to Autonomy via Modular Sensor and Mobility Bricks

In this thesis, the teleoperated communications of a Remotec ANDROS robot have been reverse engineered. This research has used the information acquired through the reverse engineering process to enhance the teleoperation and add intelligence to the initially automated robot. The main contribution of this thesis is the implementation of the mobility brick paradigm, which enables autonomous operations, using the commercial teleoperated ANDROS platform. The brick paradigm is a generalized architecture for a modular approach to robotics. This architecture and the contribution of this thesis are a paradigm shift from the proprietary commercial models that exist today. The modular system of sensor bricks integrates the transformed mobility platform and defines it as a mobility brick. In the wall following application implemented in this work, the mobile robotic system acquires intelligence using the range sensor brick. This application illustrates a way to alleviate the burden on the human operator and delegate certain tasks to the robot. Wall following is one among several examples of giving a degree of autonomy to an essentially teleoperated robot through the Sensor Brick System. Indeed once the proprietary robot has been altered into a mobility brick; the possibilities for autonomy are numerous and vary with different sensor bricks. The autonomous system implemented is not a fixed-application robot but rather a non-specific autonomy capable platform. Meanwhile the native controller and the computer-interfaced teleoperation are still available when necessary. Rather than trading off by switching from teleoperation to autonomy, this system provides the flexibility to switch between the two at the operator’s command. The contributions of this thesis reside in the reverse engineering of the original robot, its upgrade to a computer-interfaced teleoperated system, the mobility brick paradigm and the addition of autonomy capabilities. The application of a robot autonomously following a wall is subsequently implemented, tested and analyzed in this work. The analysis provides the programmer with information on controlling the robot and launching the autonomous function. The results are conclusive and open up the possibilities for a variety of autonomous applications for mobility platforms using modular sensor bricks

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

How variation in head pitch could affect image matching algorithms for ant navigation

Desert ants are a model system for animal navigation, using visual memory to follow long routes across both sparse and cluttered environments. Most accounts of this behaviour assume retinotopic image matching, e.g. recovering heading direction by finding a minimum in the image difference function as the viewpoint rotates. But most models neglect the potential image distortion that could result from unstable head motion. We report that for ants running across a short section of natural substrate, the head pitch varies substantially: by over 20 degrees with no load; and 60 degrees when carrying a large food item. There is no evidence of head stabilisation. Using a realistic simulation of the ant’s visual world, we demonstrate that this range of head pitch significantly degrades image matching. The effect of pitch variation can be ameliorated by a memory bank of densely sampled along a route so that an image sufficiently similar in pitch and location is available for comparison. However, with large pitch disturbance, inappropriate memories sampled at distant locations are often recalled and navigation along a route can be adversely affected. Ignoring images obtained at extreme pitches, or averaging images over several pitches, does not significantly improve performance

How is an ant navigation algorithm affected by visual parameters and ego-motion?

Ants typically use path integration and vision for navigation when the environment precludes the use of pheromones for trails. Recent simulations have been able to accurately mimic the retinotopic navigation behaviour of these ants using simple models of movement and memory of unprocessed visual images. Naturally it is interesting to test these navigation algorithms in more realistic circumstances, particularly with actual route data from the ant, in an accurate facsimile of the ant world and with visual input that draws on the characteristics of the animal. While increasing the complexity of the visual processing to include skyline extraction, inhomogeneous sampling and motion processing was conjectured to improve the performance of the simulations, the reverse appears to be the case. Examining closely the assumptions about motion, analysis of ants in the field shows that they experience considerable displacement of the head which when applied to the simulation leads to significant degradation in performance. The family of simulations rely upon continuous visual monitoring of the scene to determine heading and it was decided to test whether the animals were similarly dependent on this input. A field study demonstrated that ants with only visual navigation cues can return the nest when largely facing away from the direction of travel (moving backwards) and so it appears that ant visual navigation is not a process of continuous retinotopic image matching. We conclude ants may use vision to determine an initial heading by image matching and then continue to follow this direction using their celestial compass, or they may use a rotationally invariant form of the visual world for continuous course correction

Learning cognitive maps: Finding useful structure in an uncertain world

In this chapter we will describe the central mechanisms that influence how people learn about large-scale space. We will focus particularly on how these mechanisms enable people to effectively cope with both the uncertainty inherent in a constantly changing world and also with the high information content of natural environments. The major lessons are that humans get by with a less is more approach to building structure, and that they are able to quickly adapt to environmental changes thanks to a range of general purpose mechanisms. By looking at abstract principles, instead of concrete implementation details, it is shown that the study of human learning can provide valuable lessons for robotics. Finally, these issues are discussed in the context of an implementation on a mobile robot. © 2007 Springer-Verlag Berlin Heidelberg