Sensitivity to initial conditions in the route-finder model

A model of stimulus generalization [Staddon, J.E.R., Reid, A.K., 1990. On the dynamics of generalization. Psychol. Rev. 97, 576-578] was extended by Reid and Staddon [Reid, A.K., Staddon, J.E.R., 1998. A dynamic route finder for the cognitive map. Psychol. Rev. 105, 585-601] to account for the route-finder issue in the Tolman-Guthrie discussion about cognitive maps. This deterministic model has been described as sensitive to initial conditions in terms of the parameters of a diffusion equation [Reid, A.K., Staddon, J.E.R., 1998. A dynamic route finder for the cognitive map. Psychol. Rev. 105, 585-601]. A simulation study was carried out to test this assertion, evaluating whether the pattern of variation in the model output resembled a chaotic pattern. Results indicate that the model is sensitive to initial conditions, suggesting that the spatial navigation task addressed by this model involves non-linear relationships and complexity beyond the apparent simplicity of the stimulus generalization process.     

How does food distance influence foraging in the ant Lasius niger: the importance of home-range marking

We study the influence of food distance on the individual foraging behaviour of Lasius niger scouts and we investigate which cue they use to assess their distance from the nest and accordingly tune their recruiting behaviour. Globally, the number of U-turns made by scouts increases with distance resulting in longer travel times and duration of the foraging cycle. However, over familiar areas, home-range marking reduces the frequency and thereby the impact of U-turns on foraging times leading to a quicker exploitation of food sources than over unmarked set-ups. Regarding information transfer, the intensity of the recruitment trail reaching the nest decreases with increasing food distance for all set-ups and is even more reduced in the absence of home-range marking. Hence, the probability of a scout continuing to lay a trail changes along the homeward journey but in a different way according to home-range marking. Over unexplored setups, at a given distance from the food source, the percentage of returning trail-laying ants remains unchanged for all tested nest-feeder distances. Hence, the tuning of the trail recruiting signal by scouts was not influenced by an odometric estimate of the distance already travelled by the ants during their outward journey to the food. By contrast, over previously explored set-ups, a distance-related factor – that is the intensity of home-range marking – strongly influences their recruiting behaviour. In fact, over a home-range marked bridge, the probability of returning ants maintaining their trail-laying behaviour increases with decreasing food distance while the gradient of home-range marks even induces ants which have stopped laying a trail to resume this behaviour in the nest vicinity. We suggest that home-range marking laid passively by walking ants is a relevant cue for scouts to indirectly assess distance from the nest but also local activity level or foraging risks in order to adaptively tune trail recruitment and colony foraging dynamics. Received 13 July 2004; revised 26 January and 20 May 2005; accepted 2 July 2005.     

Host plant finding in the specialised leaf beetle Cassida canaliculata: an analysis of small-scale movement behaviour

Abstract.  1. Host plant finding in walking herbivorous beetles is still poorly understood. Analysis of small-scale movement patterns under semi-natural conditions can be a useful tool to detect behavioural responses towards host plant cues.
2. In this study, the small-scale movement behaviour of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) was studied in a semi-natural arena ( r = 1 m). In three different settings, a host ( Salvia pratensis L., Lamiales: Lamiaceae), a non-host ( Rumex conglomeratus Murr., Caryophyllales: Polygonaceae), or no plant was presented in the centre of the arena.
3. The beetles showed no differences in the absolute movement variables, straightness and mean walking speed, between the three settings. However, the relative movement variables, mean distance to the centre and mean angular deviation from walking straight to the centre, were significantly smaller when a host plant was offered. Likewise, the angular deviation from walking straight to the centre tended to decline with decreasing distance from the centre. Finally, significantly more beetles were found on the host than on the non-host at the end of all the trials.
4. It is concluded that C. canaliculata is able to recognise its host plant from a distance. Whether olfactory or visual cues (or a combination of both) are used to find the host plant remains to be elucidated by further studies.     

Influence of visual targets and landmarks on honey bee foraging and waggle dancing

Animals use diverse sensory stimuli to navigate their environment and to recognize rewarding food sources.Honey bees use visual atributes of the targeted food source,such as its color,shape,size,direction and distance from the hive,and the landmarks around it to navigate during foraging.They transmit the location information of the food source to other bees if it is highly rewarding.To investigate the relative importance of these attributes,we trained bees to feeders in two different experiments.In the first experiment,we asked whether bees prefer to land on(a)a similar feeder at a different distance on the same heading or on(b)a visually distinct feeder located at the exact same location.We found that,within a short foraging range,bees relied heavily on the color and the shape of the food source and to a lesser extent on its distance from the hive.In the second experiment,we asked if moving the main landmark or the feeder(visual target)influenced recruitment dancing for the feeder.We found that foragers took longer to land and danced fewer circuits when the location of the food source,or a major landmark associated with it,changed.These results demonstrate that prominent visual atributes of food sources and landmarks are evidently more reliable than distance information and that foraging bees heavily utilize these visual cues at the later stages of their journey.     

Movement tactics of a mobile predator in a meta‐ecosystem with fluctuating resources: the arctic fox in the High Arctic

Animal movement is a fundamental process shaping ecosystems at multiple levels, from the fate of individuals to global patterns of biodiversity. The spatio‐temporal dynamic of food resources is a major driver of animal movement and generates patterns ranging from range residency to migration and nomadism. Arctic tundra predators face a strongly fluctuating environment marked by cyclic microtine populations, high seasonality, and the potential availability of sea ice, which gives access to marine resources in winter. This type of relatively poor and highly variable environment can promote long‐distance movements and resource tracking in mobile species. Here, we investigated the winter movements of the arctic fox, a major tundra predator often described as a seasonal migrant or nomad. We used six years of Argos satellite telemetry data collected on 66 adults from Bylot Island (Nunavut, Canada) tracked during the sea ice period. We hypothesized that long‐distance movements would be influenced by spatio‐temporal changes in resource availability and individual characteristics. Despite strong annual and seasonal changes in resource abundance and distribution, we found that a majority of individuals remained resident, especially those located in an area characterized by highly predictable pulse resources (goose nesting colony) and abundant cached food items (eggs). Foxes compensated terrestrial food shortage by commuting to the sea ice rather than using long‐distance tracking or moving completely onto the sea ice for winter. Individual characteristics also influenced movement patterns: age positively influenced the propensity to engage in nomadism, suggesting older foxes may be driven out of their territories. Our results show how these mammalian predators can adjust their movement patterns to favor range residency despite strong spatio‐temporal fluctuations in food resources. Understanding the movement responses of predators to prey dynamics helps identifying the scales at which they work, which is a critical aspect of the functioning and connectivity among meta‐ecosystems.     

Foraging in highly dynamic environments: leaf‐cutting ants adjust foraging trail networks to pioneer plant availability

Major shifts in the availability of palatable plant resources are of key relevance to the ecology of leaf‐cutting ants in human‐modified landscapes. However, our knowledge is still limited regarding the ability of these ants to adjust their foraging strategy to dynamic environments. Here, we examine a set of forest stand attributes acting as modulating forces for the spatiotemporal architecture of foraging trail networks developed by Atta cephalotes L. (Hymenoptera: Formicidae: Attini). During a 12‐month period, we mapped the foraging systems of 12 colonies located in Atlantic forest patches with differing size, regeneration age, and abundance of pioneer plants, and examined the variation in five trail system attributes (number of trails, branching points, leaf sources, linear foraging distance, and trail complexity) in response to these patch‐related variables. Both the month‐to‐month differences (depicted in annual trail maps) and the steadily accumulating number of trails, trail‐branching points, leaf sources, and linear foraging distance illustrated the dynamic nature of spatial foraging and trail complexity. Most measures of trail architecture correlated positively with the number of pioneer trees across the secondary forest patches, but no effects from patch age and size were observed (except for number of leaf sources). Trail system complexity (measured as fractal dimension; Df index) varied from 1.114 to 1.277 along the 12 months through which ant foraging was monitored, with a marginal trend to increase with the abundance of pioneer stems. Our results suggest that some leaf‐cutting ant species are able to generate highly flexible trail networks (via fine‐tuned adjustment of foraging patterns), allowing them to profit from the continuous emergence/recruitment of palatable resources.     

Food distance and its effect on nutrient balancing in a mobile insect herbivore

We investigated how the distance between foods of differing nutrient content affects macronutrient (protein and digestible carbohydrate) regulation and the patterns of food acquisition, movement and feeding activity in fifth-instar nymphs of Locusta migratoria (L.) (Orthoptera: Acrididae). We placed individual insects into one of three differently sized circular arenas (20, 40 or 80 cm diameter) that contained four dishes of chemically defined synthetic food. One of these dishes contained high-protein, low-carbohydrate food (P), and the other three dishes contained low-protein, high-carbohydrate food (C). Alone, these foods are nutritionally unbalanced, but together they are complementary. Regardless of arena size, locusts regulated their protein-carbohydrate intake to similar points, and in all three arenas they ate preferentially from the dish containing the P-food. We also recorded the patterns of foraging behaviour for 12 h on days 1 and 4 of the experiment, which allowed us to determine the effect of distance between foods and whether behaviour was modified with experience in the different arenas. Locusts' foraging behaviour in small arenas was similar on days 1 and 4, and the time they spent in different parts of the arena did not differ from that predicted by the simplest random probability model. In contrast, the foraging behaviour of locusts in the medium and large arenas changed between days 1 and 4, including spending more time in the area of the arena containing the P-food and visiting fewer of the C-food dishes. We discuss how distance between foods influences foraging strategies, and the possible role of learning in its development. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Context-dependent navigation in a collectively foraging species of ant, Messor cephalotes

More than 100 years of scientific research has provided evidence for sophisticated navigational mechanisms in social insects. One key role for navigation in ants is the orientation of workers between food sources and the nest. The focus of recent work has been restricted to navigation in individually foraging ant species, yet many species do not forage entirely independently, instead relying on collectively maintained information such as persistent trail networks and/or pheromones. Harvester ants use such networks, but additionally, foragers often search individually for food either side of trails. In the absence of a trail, these ‘off-trail’ foragers must navigate independently to relocate the trail and return to the nest. To investigate the strategies used by ants on and off the main trails, we conducted field experiments with a harvester ant species, Messor cephalotes, by transferring on-trail and off-trail foragers to an experimental arena. We employed custom-built software to track and analyse ant trajectories in the arena and to quantitatively compare behaviour. Our results indicate that foragers navigate using different cues depending on whether they are travelling on or off the main trails. We argue that navigation in collectively foraging ants deserves more attention due to the potential for behavioural flexibility arising from the relative complexity of journeys between food and the nest.     

Motor control of voluntary arm movements

The motor control of pointing and reaching-to-grasp movements was investigated using two different approaches (kinematic and modelling) in order to establish whether the type of control varies according to modifications of arm kinematics. Kinematic analysis of arm movements was performed on subjects' hand trajectories directed to large and small stimuli located at two different distances. The subjects were required either to grasp and to point to each stimulus. The kinematics of the subsequent movement, during which subject's hand came back to the starting position, were also studied. For both movements, kinematic analysis was performed on hand linear trajectories as well as on joint angular trajectories of shoulder and elbow. The second approach consisted in the parametric identification of the black box (ARMAX) model of the controller driving the arm movement. Such controller is hypothesized to work for the correct execution of the motor act. The order of the controller ARMAX model was analyzed with respect to the different experimental conditions (distal task, stimulus size and distance). Results from kinematic analysis showed that target distance and size influenced kinematic parameters both of angular and linear displacements. Nevertheless, the structure of the motor program was found to remain constant with distane and distal task, while it varied with precision requirements due to stimulus size. The estimated model order of the controller confirmed the invariance of the control law with regard to movement amplitude, whereas it was sensitive to target size.     

Foraging arena size and structural complexity affect the dynamics of food distribution in ant colonies

Food acquisition by ant colonies is a complex process that starts with acquiring food at the source (i.e., foraging) and culminates with food exchange in or around the nest (i.e., feeding). While ant foraging behavior is relatively well understood, the process of food distribution has received little attention, largely because of the lack of methodology that allows for accurate monitoring of food flow. In this study, we used the odorous house ant, Tapinoma sessile (Say) to investigate the effect of foraging arena size and structural complexity on the rate and the extent of spread of liquid carbohydrate food (sucrose solution) throughout a colony. To track the movement of food, we used protein marking and double-antibody sandwich enzyme-linked immunosorbent assay, DAS-ELISA. Variation in arena size, in conjunction with different colony sizes, allowed us to test the effect of different worker densities on food distribution. Results demonstrate that both arena size and colony size have a significant effect on the spread of the food and the number of workers receiving food decreased as arena size and colony size increased. When colony size was kept constant and arena size increased, the percentage of workers testing positive for the marker decreased, most likely because of fewer trophallactic interactions resulting from lower worker density. When arena size was kept constant and colony size increased, the percentage of workers testing positive decreased. Nonrandom (clustered) worker dispersion and a limited supply of food may have contributed to this result. Overall, results suggest that food distribution is more complete is smaller colonies regardless of the size of the foraging arena and that colony size, rather than worker density, is the primary factor affecting food distribution. The structural complexity of foraging arenas ranged from simple, two-dimensional space (empty arenas) to complex, three-dimensional space (arenas filled with mulch). The structural complexity of foraging arenas had a significant effect on food distribution and the presence of substrate significantly inhibited the spread of food. Structural complexity of foraging arenas and the resulting worker activity patterns might exert considerable influence on socioecological processes in ants and should be considered in laboratory assays.     

Modeling Drosophila Positional Preferences in Open Field Arenas with Directional Persistence and Wall Attraction

In open field arenas, Drosophila adults exhibit a preference for arena boundaries over internal walls and open regions. Herein, we investigate the nature of this preference using phenomenological modeling of locomotion to determine whether local arena features and constraints on movement alone are sufficient to drive positional preferences within open field arenas of different shapes and with different internal features. Our model has two components: directional persistence and local wall force. In regions far away from walls, the trajectory is entirely characterized by a directional persistence probability, , for each movement defined by the step size, , and the turn angle, . In close proximity to walls, motion is computed from and a local attractive force which depends on the distance between the fly and points on the walls. The directional persistence probability was obtained experimentally from trajectories of wild type Drosophila in a circular open field arena and the wall force was computed to minimize the difference between the radial distributions from the model and Drosophila in the same circular arena. The two-component model for fly movement was challenged by comparing the positional preferences from the two-component model to wild type Drosophila in a variety of open field arenas. In most arenas there was a strong concordance between the two-component model and Drosophila. In more complex arenas, the model exhibits similar trends, but some significant differences were found. These differences suggest that there are emergent features within these complex arenas that have significance for the fly, such as potential shelter. Hence, the two-component model is an important step in defining how Drosophila interact with their environment.     

Individual rules for trail pattern formation in Argentine ants (Linepithema humile)

We studied the formation of trail patterns by Argentine ants exploring an empty arena. Using a novel imaging and analysis technique we estimated pheromone concentrations at all spatial positions in the experimental arena and at different times. Then we derived the response function of individual ants to pheromone concentrations by looking at correlations between concentrations and changes in speed or direction of the ants. Ants were found to turn in response to local pheromone concentrations, while their speed was largely unaffected by these concentrations. Ants did not integrate pheromone concentrations over time, with the concentration of pheromone in a 1 cm radius in front of the ant determining the turning angle. The response to pheromone was found to follow a Weber's Law, such that the difference between quantities of pheromone on the two sides of the ant divided by their sum determines the magnitude of the turning angle. This proportional response is in apparent contradiction with the well-established non-linear choice function used in the literature to model the results of binary bridge experiments in ant colonies (Deneubourg et al. 1990). However, agent based simulations implementing the Weber's Law response function led to the formation of trails and reproduced results reported in the literature. We show analytically that a sigmoidal response, analogous to that in the classical Deneubourg model for collective decision making, can be derived from the individual Weber-type response to pheromone concentrations that we have established in our experiments when directional noise around the preferred direction of movement of the ants is assumed.     

Local versus landscape determinants of butterfly movement behaviors

Large-scale patterns of animal distributions and abundances may be determined by mechanisms that act at local or landscape scales. We studied the movement behaviors of four species of bottomland butterflies in a natural setting to examine the determinants of movement behavior across different scales. We tested the relative importance of three landscape attributes: drainage slope, boundary type, and stream proximity, and local habitat attributes related to food plants and plants that influence habitat structure. Across species, we tested the relative importance of organism size and habitat specificity to explain response variation. In general, butterfly responses to landscape features were more universal than responses to local features. Specifically, results from this study showed that drainage slope did not influence movement behaviors but boundary type, stream proximity, and host plant abundance all influenced movement patterns. Responses to local features varied by species and often complemented landscape effects on movement. Responses to all features were not related to butterfly size, but did vary in accordance with butterfly host plant specificity. These behaviors help to explain landscape-level variation in population distribution among species.     

Consistent trophic patterns among fishes in lagoon and channel habitats of a tropical floodplain river: Evidence from stable isotopes

The relationship between food web dynamics and hydrological connectivity in rivers should be strongly influenced by annual flood pulses that affect primary production dynamics and movement of organic matter and consumer taxa. We sampled basal production sources and fishes from connected lagoons and the main channel of a low-gradient, floodplain river within the Orinoco River Basin in Venezuela. Stable isotope analysis was used to model the contribution of four basal production sources to fishes, and to examine patterns of mean trophic position during the falling-water period of the annual flood cycle. IsoSource, a multi-source mixing model, indicated that proportional contributions from production sources to fish assemblages were similar in lagoons and the main channel. Although distributions differed, the means for trophic positions of fish assemblages as well as individual species were similar between the two habitats. These findings contradict recent food web studies conducted in temperate floodplain rivers that described significant differences in trophic positions of fishes from slackwater and floodplain versus main channel habitats. Low between-habitat trophic variation in this tropical river probably results from an extended annual flood pulse (ca. 5 mo.) that allows mixing of sestonic and allochthonous basal production sources and extensive lateral movements of fishes throughout the riverscape.     

Negative Feedback Enables Fast and Flexible Collective Decision-Making in Ants

Positive feedback plays a major role in the emergence of many collective animal behaviours. In many ants pheromone trails recruit and direct nestmate foragers to food sources. The strong positive feedback caused by trail pheromones allows fast collective responses but can compromise flexibility. Previous laboratory experiments have shown that when the environment changes, colonies are often unable to reallocate their foragers to a more rewarding food source. Here we show both experimentally, using colonies of Lasius niger, and with an agent-based simulation model, that negative feedback caused by crowding at feeding sites allows ant colonies to maintain foraging flexibility even with strong recruitment to food sources. In a constant environment, negative feedback prevents the frequently found bias towards one feeder (symmetry breaking) and leads to equal distribution of foragers. In a changing environment, negative feedback allows a colony to quickly reallocate the majority of its foragers to a superior food patch that becomes available when foraging at an inferior patch is already well underway. The model confirms these experimental findings and shows that the ability of colonies to switch to a superior food source does not require the decay of trail pheromones. Our results help to resolve inconsistencies between collective foraging patterns seen in laboratory studies and observations in the wild, and show that the simultaneous action of negative and positive feedback is important for efficient foraging in mass-recruiting insect colonies.     

Tradeoff Between Travel Speed and Olfactory Food Detection in Ring‐Tailed Coatis (Nasua nasua)

The distance at which an animal can detect food has important ramifications for foraging behavior. Although some studies have investigated the factors which affect visual food detection, very little is known about what influences olfactory food detection abilities in wild animals. The food discovery behavior of ring‐tailed coatis (Nasua nasua) was studied using experimental fruit plots. Coatis primarily used olfaction to detect these new food sources, and visual food discovery appeared plausible in only five of 148 trials. Coatis detected the fruit from longer distances when traveling compared with when foraging for invertebrates in the leaf litter. Travel speed had a negative effect on discovery distance. Coatis traveling slowly detected the fruit plots from further away, which demonstrated a tradeoff between speed and food detection. If this tradeoff is biologically important, slower groups should have visited more fruit trees per day, so data taken during full‐day coati group follows were analyzed to determine whether this pattern occurred. Slower moving groups visited more fruit trees than faster groups once confounding factors such as daily travel distance, group identity, group spread, and year were controlled for. These results are consistent with the hypothesis that coatis exhibit a speed‐accuracy tradeoff for olfactory food detection. This tradeoff appears to be an important factor influencing the movement ecology of animal groups.     

Compass Orientation During Dispersal of Freshwater Hatchling Snapping Turtles (Chelydra serpentina) and Blanding's Turtles (Emydoidea blandingii)

Freshwater turtle hatchlings primarily use visual cues for orientation while dispersing from nests; however, hatchlings rapidly develop a relationship between a sun or geomagnetic compass and a dispersal target that allows them to maintain an established direction of movement when target habitats are not visible. We examined dispersal patterns of hatchling snapping turtles (Chelydra serpentina) and Blanding's turtles (Emydoidea blandingii) dispersing in large arenas in a mowed field and in dense corn. The dispersal of three categories of hatchlings were examined: (1) naïve individuals (no previous dispersal experience), (2) arena‐experienced (limited dispersal experience in arenas in natural habitat), and (3) natural‐experienced hatchling Blanding's turtles (captured after extensive experience dispersing W in natural habitats toward wetlands). Experienced hatchlings were assigned to treatments consisting of having a magnet or a non‐magnetic aluminum sham or nothing glued to their anterior carapace before release in the corn arena. Dispersal patterns of naïve hatchlings of both species were strongly directional in the field arena with visible target horizons and primarily random in the corn arena where typical target horizons were blocked. When released in corn, dispersal patterns were similar for arena‐experienced hatchlings with magnets or shams attached and differed from their prior dispersal headings in the field arena as naïve hatchlings. Natural‐experienced hatchling Blanding's turtles with and without magnets were able to accurately maintain their prior headings to the WNW while dispersing in the field or corn arenas (i.e., the presence of a magnet did not disrupt their ability to maintain their prior heading). Based on the assumption that no other type of compass exists in hatchlings, we conclude that they were not using a geomagnetic compass, but by default were using sun compass orientation to maintain dispersal headings in dense corn where no typical target habitats were visible.     

Random movement pattern of the sea urchin Strongylocentrotus droebachiensis

We describe the fine-scale movement of the sea urchin Strongylocentrotus droebachiensis based on analyses of video recordings of undisturbed individuals in the two habitats which mainly differed in food availability, urchin barrens and grazing front. Urchin activity decreased as urchin density increased. Individuals alternated between moving and being stationary and their behaviour did not appear to be affected by either current velocity (within the range from 0 to 15 cm s^− 1) and temperature (2.3 to 6.0 °C). Movement of individuals at each location was compared to that predicted by a random walk model. Mean move length (linear distance between two stationary periods), turning angle and net squared displacement were calculated for each individual. The distribution of turning angles was uniform at each location and there was no evidence of a relationship between urchin density and either move length or urchin velocity. The random model predicted a higher dispersal rate at locations with low urchin densities, such as barrens habitats. However, the movement was sometimes greater or less than predicted by the model, suggesting the influence of local environmental factors. The deviation of individual paths from the model revealed that urchins can be stationary or adopt a local (displacement less than random), random or directional movement. The net daily distance displaced on the barrens, predicted by a random walk model, was similar to the observed movement recorded in our previous study of tagged urchins at one site, but less than that observed at a second site. We postulate that the random dispersal of urchins allows individuals on barrens to reach the kelp zone where food is more abundant although the time required to reach the kelp zone may be considerable (months to years). Urchins decrease their rate of dispersal once they reach the kelp zone so that they likely remain close to this abundant food sources for long periods.     

Distributed leadership and adaptive decision-making in the ant Tetramorium caespitum

In the ant species Tetramorium caespitum, communication and foraging patterns rely on group-mass recruitment. Scouts having discovered food recruit nestmates and behave as leaders by guiding groups of recruits to the food location. After a while, a mass recruitment takes place in which foragers follow a chemical trail. Since group recruitment is crucial to the whole foraging process, we investigated whether food characteristics induce a tuning of recruiting stimuli by leaders that act upon the dynamics and size of recruited groups. High sucrose concentration triggers the exit of a higher number of groups that contain twice as many ants and reach the food source twice as fast than towards a weakly concentrated one. Similar trends were found depending on food accessibility: for a cut mealworm, accessibility to haemolymph results in a faster formation of larger groups than for an entire mealworm. These data provide the background for developing a stochastic model accounting for exploitation patterns by group-mass recruiting species. This model demonstrates how the modulations performed by leaders drive the colony to select the most profitable food source among several ones. Our results highlight how a minority of individuals can influence collective decisions in societies based on a distributed leadership.     

A neural-network reinforcement-learning model of domestic chicks that learn to localize the centre of closed arenas

Previous experiments have shown that when domestic chicks (Gallus gallus) are first trained to locate food elements hidden at the centre of a closed square arena and then are tested in a square arena of double the size, they search for food both at its centre and at a distance from walls similar to the distance of the centre from the walls experienced during training. This paper presents a computational model that successfully reproduces these behaviours. The model is based on a neural-network implementation of the reinforcement-learning actor - critic architecture (in this architecture the 'critic' learns to evaluate perceived states in terms of predicted future rewards, while the 'actor' learns to increase the probability of selecting the actions that lead to higher evaluations). The analysis of the model suggests which type of information and cognitive mechanisms might underlie chicks' behaviours: (i) the tendency to explore the area at a specific distance from walls might be based on the processing of the height of walls' horizontal edges, (ii) the capacity to generalize the search at the centre of square arenas independently of their size might be based on the processing of the relative position of walls' vertical edges on the horizontal plane (equalization of walls' width), and (iii) the whole behaviour exhibited in the large square arena can be reproduced by assuming the existence of an attention process that, at each time, focuses chicks' internal processing on either one of the two previously discussed information sources. The model also produces testable predictions regarding the generalization capabilities that real chicks should exhibit if trained in circular arenas of varying size. The paper also highlights the potentialities of the model to address other experiments on animals' navigation and analyses its strengths and weaknesses in comparison to other models.