Search

The extension of cognition beyond the brain to the body and beyond the body to the environment is an area of debate in philosophy and the cognitive sciences. Yet, these debates largely overlook olfaction, a sensory modality used by most animals. Here, I use the philosopher's framework to explore the implications of embodiment for olfactory cognition. The philosopher's 4E framework comprises embodied cognition, emerging from a nervous system characterized by its interactions with its body. The necessity of action for perception adds enacted cognition. Cognition is further embedded in the sensory inputs of the individual and is extended beyond the individual to information stored in its physical and social environments. Further, embodiment must fulfill the criterion of mutual manipulability, where an agent's cognitive state is involved in continual, reciprocal influences with its environment. Cognition cannot be understood divorced from evolutionary history, however, and I propose adding evolved, as a fifth term to the 4E framework. We must, therefore, begin at the beginning, with chemosensation, a sensory modality that underlies purposive behavior, from bacteria to humans. The PROUST hypothesis (perceiving and reconstructing odor utility in space and time) describers how olfaction, this ancient scaffold and common denominator of animal cognition, fulfills the criteria of embodied cognition. Olfactory cognition, with its near universal taxonomic distribution as well as the near absence of conscious representation in humans, may offer us the best sensorimotor system for the study of embodiment.

The ability to use environmental geometry when orienting in space reflects an animal's ability to use a global, allocentric framework. Therefore, understanding when and how animal's use geometry relative to other types of cues in the environment has interested comparative cognition researchers for decades. Yet, only two amphibians have been tested to date. We trained the poison frog Dendrobates auratus to find goal shelters in a rectangular arena, in the presence and absence of a feature cue, and assessed the relative influence of the two types of cues using probe trials. We chose D. auratus because the species has complex interactions with their physical and social environments, including parental care that requires navigating to and from distant locations. We found that, like many vertebrates, D. auratus are capable of using geometric information to relocate goals. In addition, the frogs preferentially used the more reliable feature cue when the location of the feature conflicted with the geometry of the arena. The frogs were equally successful at using the feature cue when it was proximal or distal to the goal shelter, consistent with prior studies that found that D. auratus can use distal cues in a flexible manner. Our results provide further evidence that amphibians can use environmental geometry during orientation. Future studies that examine when and how amphibians use geometry relative to other types of cues will contribute to a more complete picture of spatial cognition in this important, yet understudied, group. [ABSTRACT FROM AUTHOR]

How do bottlenose dolphins visually perceive the space around them? In particular, what cues do they use as a frame of reference for left–right perception? To address this question, we examined the dolphin's responses to various manipulations of the spatial relationship between the dolphin and the trainer by using gestural signs for actions given by the trainer, which have different meanings in the left and right hands. When the dolphins were tested with their backs to the trainer (Experiment 1) or in an inverted position underwater (Experiments 2 and 3), correct responses from the trainer's perspective were maintained for signs related to movement direction instructions. In contrast, reversed responses were frequently observed for signs that required different sounds for the left and right hands. When the movement direction instructions were presented with symmetrical graphic signs such as " × " and "●", accuracy decreased in the inverted posture (Experiment 3). Furthermore, when the signs for sounds were presented from either the left or right side of the dolphin's body, performance was better when the side of the sign movement coincided with the body side on which it was presented than when it was mismatched (Experiment 4). In the final experiment, when one eye was covered with an eyecup, the results showed that, as in the case of body-side presentation, performance was better when the open eye coincided with the side on which the sign movement was presented. These results indicate that dolphins used the egocentric frame for visuospatial cognition. In addition, they showed better performances when the gestural signs were presented to the right eye, suggesting the possibility of a left-hemispheric advantage in the dolphin's visuospatial cognition. [ABSTRACT FROM AUTHOR]

The capacity to navigate by layout geometry has been widely recognized as a robust strategy of place-finding. It has been reported in various species, although most studies were performed with vision-based paradigms. In the presented study, we aimed to investigate layout symmetry-based navigation in the house cricket, Acheta domesticus, in the absence of visual cues. For this purpose, we used a non-visual paradigm modeled on the Tennessee Williams setup. We ensured that the visual cues were indeed inaccessible to insects. In the main experiment, we tested whether crickets are capable of learning to localize the centrally positioned, inconspicuous cool spot in heated arenas of various shapes (i.e., circular, square, triangular, and asymmetric quadrilateral). We found that the symmetry of the arena significantly facilitates crickets' learning to find the cool spot, indicated by the increased time spent on the cool spot and the decreased latency in locating it in subsequent trials. To investigate mechanisms utilized by crickets, we analyzed their approach paths to the spot. We found that crickets used both heuristic and directed strategies of approaching the target, with the dominance of a semi-directed strategy (i.e., a thigmotactic phase preceding direct navigation to the target). We propose that the poor performance of crickets in the asymmetrical quadrilateral arena may be explained by the difficulty of encoding its layout with cues from a single modality. [ABSTRACT FROM AUTHOR]

The way in which animals grasp objects to perform subsequent action execution allows studying their anticipatory abilities. We examined whether 11 capuchin monkeys (Sapajus spp.) were able to prospectively grasp a magnetic dowel to use it as a tool to retrieve a baited metallic container from a plexiglas box placed in front of them. We investigated whether and how initial dowel orientation (horizontal vs vertical) affected grasping and using the dowel to retrieve the container in two testing conditions: (1) 2-FE condition in which the dowel had two functional magnetic ends; (2) 1-FE condition in which the dowel had only one functional magnetic end. In the 2-FE condition, capuchins had to take into account the initial dowel orientation since both ends were functional, whereas in the 1-FE condition capuchins had also to take into account the initial functional end position when grasping the dowel. Capuchins were trained to grasp the dowel to put one functional end in contact with the metallic container. However, they did not learn to associate the functional end of the 1-FE dowel to successful retrieval. Capuchins showed better anticipatory planning (1) in 2-FE than in 1-FE condition and (2) when the dowel was initially positioned on the horizontal plane than on the vertical one. Moreover, hand preferences affected planning in the 1-FE condition. Results were discussed within the frameworks of primates' abilities to use abstract cues and on their abilities to process functional features and spatial cues and to perform mental rotations. [ABSTRACT FROM AUTHOR]

To increase efficiency in the search for resources, many animals rely on their spatial abilities. Specifically, primates have been reported to use mostly topological and rarely Euclidean maps when navigating in large-scale space. Here, we aimed to investigate if the navigation of wild common marmosets inhabiting a semiarid environment is consistent with a topological representation and how environmental factors affect navigation. We collected 497 h of direct behavioral and GPS information on a group of marmosets using a 2-min instantaneous focal animal sampling technique. We found that our study group reused not only long-route segments (mean of 1007 m) but entire daily routes, a pattern that is not commonly seen in primates. The most frequently reused route segments were the ones closer to feeding sites, distant to resting sites, and in areas sparse in tree vegetation. We also identified a total of 56 clustered direction change points indicating that the group modified their direction of travel. These changes in direction were influenced by their close proximity to resting and feeding sites. Despite our small sample size, the obtained results are important and consistent with the contention that common marmosets navigate using a topological map that seems to benefit these animals in response to the exploitation of clustered exudate trees. Based on our findings, we hypothesize that the Caatinga landscape imposes physical restrictions in our group's navigation such as gaps in vegetation, small trees and xerophytic plants. This study, based on preliminary evidence, raises the question of whether navigation patterns are an intrinsic characteristic of a species or are ecologically dependent and change according to the environment. [ABSTRACT FROM AUTHOR]

We suggest that socio-spatial behavior, which is an interaction between social and spatial cognition, can be viewed as a set of excursions that originate and end in close proximity to another individual(s). We present an extension of earlier studies that perceived spatial behavior in individual animals as a series of excursions originating from a particular location. We measured here the momentary distance between two individuals (social distance) to differentiate among eight possible types of social excursion originating in a state of proximity between excursion-participants. The defined excursion types are based on whether or not the excursion initiator also concludes the excursion, whether or not the excursion starts and ends at the same location, and the dynamics of the distance between excursion participants. We validated this approach to socio-spatial behavior as a set of excursions using it to analyze the behavior of the two sexes in rodents, of normal vs. stereotyped rats, as well as of different rodent species. Each of these groups displays a prevalent excursion type that reflects a distinct social dynamics. Our approach offers a useful and comprehensive tool for studying socio-spatial cognition, and can also be applied to distinguish among different social situations in rodents and other animals. [ABSTRACT FROM AUTHOR]

Animals' cognitive abilities can be tested by allowing them to choose between alternatives, with only one alternative offering the correct solution to a novel problem. Hermit crabs are evolutionarily specialized to navigate while carrying a shell, with alternative shells representing different forms of 'extended architecture', which effectively change the extent of physical space an individual occupies in the world. It is unknown whether individuals can choose such architecture to solve novel navigational problems. Here, we designed an experiment in which social hermit crabs (Coenobita compressus) had to choose between two alternative shells to solve a novel problem: escaping solitary confinement. Using X-ray microtomography and 3D-printing, we copied preferred shell types and then made artificial alterations to their inner or outer shell architecture, designing only some shells to have the correct architectural fit for escaping the opening of an isolated crab's enclosure. In our 'escape artist' experimental design, crabs had to choose an otherwise less preferred shell, since only this shell had the right external architecture to allow the crab to free itself from isolation. Across multiple experiments, crabs were willing to forgo preferred shells and choose less preferred shells that enabled them to escape, suggesting these animals can solve novel navigational problems with extended architecture. Yet, it remains unclear if individuals solved this problem through trial-and-error or were aware of the deeper connection between escape and exterior shell architecture. Our experiments offer a foundation for further explorations of physical, social, and spatial cognition within the context of extended architecture. [ABSTRACT FROM AUTHOR]

The feats of arthropods, and of the well-studied insects and crustaceans in particular, have fascinated scientists and laymen alike for centuries. Arthropods show a diverse repertoire of cognitive feats, of often unexpected sophistication. Despite their smaller brains and resulting lower neuronal capacity, the cognitive abilities of arthropods are comparable to, or may even exceed, those of vertebrates, depending on the species compared. Miniature brains often provide parsimonious but smart solutions for complex behaviours or ecologically relevant problems. This makes arthropods inspiring subjects for basic research, bionics, and robotics. Investigations of arthropod spatial cognition have originally concentrated on the honeybee, an animal domesticated for several thousand years. Bees are easy to keep and handle, making this species amenable to experimental study. However, there are an estimated 5–10 million arthropod species worldwide, with a broad diversity of lifestyles, ecology, and cognitive abilities. This high diversity provides ample opportunity for comparative analyses. Comparative study, rather than focusing on single model species, is well suited to scrutinise the link between ecological niche, lifestyle, and cognitive competence. It also allows the discovery of general concepts that are transferable between distantly related groups of organisms. With species diversity and a comparative approach in mind, this special issue compiles four review articles and ten original research reports from a spectrum of arthropod species. These contributions range from the well-studied hymenopterans, and ants in particular, to chelicerates and crustaceans. They thus present a broad spectrum of glimpses into current research on arthropod spatial cognition, and together they cogently emphasise the merits of research into arthropod cognitive achievements. [ABSTRACT FROM AUTHOR]

Ants are central-place foragers: they always return to the nest, and this requires the ability to remember relationships between features of the environment, or an individual's path through the landscape. The distribution of these cognitive responsibilities within a colony depends on a species' foraging style. Solitary foraging as well as leader-scouting, which is based on information transmission about a distant targets from scouts to foragers, can be considered the most challenging tasks in the context of ants' spatial cognition. Solitary foraging is found in species of almost all subfamilies of ants, whereas leader-scouting has been discovered as yet only in the Formica rufa group of species (red wood ants). Solitary foraging and leader-scouting ant species, although enormously different in their levels of sociality and ecological specificities, have many common traits of individual cognitive navigation, such as the primary use of visual navigation, excellent visual landmark memories, and the subordinate role of odour orientation. In leader-scouting species, spatial cognition and the ability to transfer information about a distant target dramatically differ among scouts and foragers, suggesting individual cognitive specialization. I suggest that the leader-scouting style of recruitment is closely connected with the ecological niche of a defined group of species, in particular, their searching patterns within the tree crown. There is much work to be done to understand what cognitive mechanisms underpin route planning and communication about locations in ants. [ABSTRACT FROM AUTHOR]

Within arthropods, the investigation of navigational aspects including homing abilities has mainly focused on insect representatives, while other arthropod taxa have largely been ignored. As such, scorpions are rather underrepresented concerning behavioral studies for reasons such as low participation rates and motivational difficulties. Here, we review the sensory abilities of scorpions related to navigation. Furthermore, we present an improved laboratory setup to shed light on navigational abilities in general and homing behavior in particular. We tracked directed movements towards home shelters of the lesser Asian scorpion Mesobuthus eupeus to give a detailed description of their departure and return movements. To do so, we analyzed the departure and return angles as well as measures of directness like directional deviation, lateral displacement, and straightness indices. We compared these parameters under different light conditions and with blinded scorpions. The motivation of scorpions to leave their shelter depends strongly upon the light condition and the starting time of the experiment; highest participation rates were achieved with infrared conditions or blinded scorpions, and close to dusk. Naïve scorpions are capable of returning to a shelter object in a manner that is directionally consistent with the home vector. The first-occurring homing bouts are characterized by paths consisting of turns about 10 cm to either side of the straightest home path and a distance efficiency of roughly three-quarters of the maximum efficiency. Our results show that neither chemosensation nor vision, but rather path integration based on proprioception, plays a superior role in the homing of scorpions. [ABSTRACT FROM AUTHOR]

One of the hardest problems in studying animal behaviour is to quantify patterns of social interaction at the group level. Recent technological developments in global positioning system (GPS) devices have opened up new avenues for locating animals with unprecedented spatial and temporal resolution. Likewise, advances in computing power have enabled new levels of data analyses with complex mathematical models to address unresolved problems in animal behaviour, such as the nature of group geometry and the impact of group-level interactions on individuals. Here, we present an information theory-based tool for the analysis of group behaviour. We illustrate its affordances with GPS data collected from a freely interacting pack of 15 Siberian huskies (Canis lupus familiaris). We found that individual freedom in movement decisions was limited to about 4%, while a subject's location could be predicted with 96% median accuracy by the locations of other group members. Dominant individuals were less affected by other individuals' locations than subordinate ones, and same-sex individuals influenced each other more strongly than opposite-sex individuals. We also found that kinship relationships increased the mutual dependencies of individuals. Moreover, the network stability of the pack deteriorated with an upcoming feeding event. Together, we conclude that information theory-based approaches, coupled with state-of-the-art bio-logging technology, provide a powerful tool for future studies of animal social interactions beyond the dyadic level. [ABSTRACT FROM AUTHOR]

Japanese eels (Anguilla japonica) were trained on a Morris-type spatial learning task. There were four tubes in a pool, but the eels could hide in only one of these. The eels learned the position of the open tube, and maintained their performance when the pool was rotated to remove possible intra-maze cues. The eels could not maintain their performance in a dark room, suggesting that spatial learning involved extra-maze visual cues. When the position of the open tube was randomly changed every day, the performance of the eels in finding the open tube did not improve. [ABSTRACT FROM AUTHOR]

Many species orient towards specific locations to reach important resources using different cognitive mechanisms. Some of these, such as path integration, are now well understood, but the cognitive orientation mechanisms that underlie movements in non-human primates remain the subject of debate. To investigate whether movements of chacma baboons are more consistent with Euclidean or topological spatial awareness, we investigated whether baboons made repeated use of the same network of pathways and tested three predictions resulting from the hypothesized use of Euclidean and topological spatial awareness. We recorded ranging behaviour of a group of baboons during 234 full days and 137 partial days in the Soutpansberg Mountains, South Africa. Results show that our baboons travelled through a dense network of repeated routes. In navigating this route network, the baboons did not approach travel goals from all directions, but instead approached them from a small number of the same directions, supporting topological spatial awareness. When leaving travel goals, baboons' initial travel direction was significantly different from the direction to the next travel goal, again supporting topological spatial awareness. Although we found that our baboons travelled with similar linearity in the core area as in the periphery of their home range, this did not provide conclusive evidence for the existence of Euclidean spatial awareness, since the baboons could have accumulated a similar knowledge of the periphery as of the core area. Overall, our findings support the hypothesis that our baboons navigate using a topological map. [ABSTRACT FROM AUTHOR]

Strategies of navigation have been shown to play a critical role when animals revisit resource sites across large home ranges. The habitual route system appears to be a sufficient strategy for animals to navigate while avoiding the cognitive cost of traveling using the Euclidean map. We hypothesize that wild elephants travel more frequently using habitual routes to revisit resource sites as opposed to using the Euclidean map. To identify the elephants' habitual routes, we created a python script, which accounted for frequently used route segments that constituted the habitual routes. Results showed elephant navigation flexibility traveling at Kruger National Park landscape. Elephants shift strategies of navigation depend on the familiarity of their surroundings. In the core area of their home range, elephants traveled using the Euclidean map, but intraindividual differences showed that elephants were then converted to habitual routes when navigating within the less familiar periphery of their home range. These findings are analogous to the recent experimental results found in smaller mammals that showed that rats encode locations according to their familiarity with their surroundings. In addition, as recently observed in monkeys, intersections of habitual routes are important locations used by elephants when making navigation decisions. We found a strong association between intersections and new segment usage by elephants when they revisit resource sites, suggesting that intersection choice may contribute to the spatial representations elephants use when repeatedly revisiting resource sites. [ABSTRACT FROM AUTHOR]

There is evidence that wild animals are able to recall key locations and associate them with navigational routes. Studies in primate navigation suggest most species navigate through the route network system, using intersections among routes as locations of decision-making. Recent approaches presume that points of directional change may be key locations where animals decide where to go next. Over four consecutive years, we observed how a wild group of bearded capuchin monkeys used a route network system and Change Point locations (CPs) in the Brazilian ecotone of Cerrado-Caatinga. We built 200 daily routes of one wild bearded capuchin group. We used ArcGIS, the Change Point Test, Spatial Analysis in Macroecology (SAM), and statistical models to test the hypothesis that wild bearded capuchins use CPs located along routes in a different fashion than they use the CPs located at intersections of routes. A logistic regression model was used to determine the landscape variables affecting capuchins’ directional changes at intersections or along routes. CPs at intersections were important points of travel path changes, whereas CPs along routes represented a zig-zag movement along the routes following the landscape features. CPs at intersections were associated with steeper terrains and shorter distances from important resources, along with better visibility of the home range. Our results support the hypothesis that intersections among routes in the route network system are located at points where monkeys have the best visibility available to make decisions on where to visit next. [ABSTRACT FROM AUTHOR]

Spatial perseveration has been documented for domestic animals such as mules, donkeys, horses and dogs. However, evidence for this spatial cognition behavior among other domestic species is scarce. Alpacas have been domesticated for at least 7000 years yet their cognitive ability has not been officially reported. The present article used an A-not-B detour task to study the spatial problem-solving abilities of alpacas (Vicugna pacos) and to identify the perseveration errors, which refers to a tendency to maintain a learned route, despite having another available path. The study tested 51 alpacas, which had to pass through a gap at one end of a barrier in order to reach a reward. After one, two, three or four repeats (A trials), the gap was moved to the opposite end of the barrier (B trials). In contrast to what has been found in other domestic animals tested with the same task, the present study did not find clear evidence of spatial perseveration. Individuals’ performance in the subsequent B trials, following the change of gap location, suggests no error persistence in alpacas. Results suggest that alpacas are more flexible than other domestic animals tested with this same task, which has important implications in planning proper training for experimental designs or productive purposes. These results could contribute toward enhancing alpacas’ welfare and our understanding of their cognitive abilities. [ABSTRACT FROM AUTHOR]

We used a modified version of the Do as I Do paradigm to investigate dogs' preference and flexibility in the acquisition of different types of spatial information in social learning situations. When required to match the location of the demonstration, dogs ( N = 16) preferentially relied on allocentric information, i.e., the relationship between the location of the demonstration and the various objects surrounding it. However, when allocentric cues were inadequate to solve the task, dogs learned to rely on egocentric information, i.e., the direction-left/right-taken by the human demonstrator. The ease of resorting to the non-preferred egocentric strategy was sex-dependent with males acquiring the egocentric strategy in fewer trials than females. This study shows that dogs rely preferentially on allocentric cues when recalling socially acquired spatial information. However, they are impressively flexible in switching to egocentric strategies according to the task requirements. Whether preference for the allocentric strategy in processing spatial information is embedded in the nature of social learning or restricted to our paradigm is an open question. This study also supports the idea that sex differences in cognitive domains are widespread among mammals and calls for further investigations aimed at shedding light on the evolution, function and mechanisms of such differences. [ABSTRACT FROM AUTHOR]

Encoding multiple cues can improve the accuracy and reliability of navigation and goal localization. Problems may arise, however, if one cue is displaced and provides information which conflicts with other cues. Here we investigated how pigeons cope with cue conflict by training them to locate a goal relative to two landmarks and then varying the amount of conflict between the landmarks. When the amount of conflict was small, pigeons tended to integrate both cues in their search patterns. When the amount of conflict was large, however, pigeons used information from both cues independently. This context-dependent strategy for resolving spatial cue conflict agrees with Bayes optimal calculations for using information from multiple sources. [ABSTRACT FROM AUTHOR]

Place learning is thought to be an adaptive and flexible facet of navigation. Due to the flexibility of this learning, it is thought to be more complex than the simpler strategies such as learning a particular route or navigating through the use of cues. Place learning is crucial in a familiar environment as it allows an individual to successfully navigate to the same endpoint, regardless of where in the environment the journey begins. Much of the research to date focusing on different strategies employed for navigation has used human subjects or other mammals such as rodents. In this series of experiments, the spatial memory of four different species of fish (goldfish, killifish, zebrafish and Siamese fighting fish) was analysed using a plus maze set-up. Results suggest that three of the species showed a significant preference for the adoption of a place strategy during this task, whereas zebrafish showed no significant preference. Furthermore, zebrafish took significantly longer to learn the task than the other species. Finally, results suggest that zebrafish took the least amount of time (seconds) to complete trials both during training and probe. [ABSTRACT FROM AUTHOR]

Although small-scale navigation is well studied in a wide range of species, much of what is known about landmark use by vertebrates is based on laboratory experiments. To investigate how vertebrates in the wild use landmarks, we trained wild male rufous hummingbirds to feed from a flower that was placed in a constant spatial relationship with two artificial landmarks. In the first experiment, the landmarks and flower were 0.25, 0.5 or 1 m apart and we always moved them 3-4 m after each visit by the bird. In the second experiment, the landmarks and flower were always 0.25 m apart and we moved them either 1 or 0.25 m between trials. In tests, in which we removed the flower, the hummingbirds stopped closer to the predicted flower location when the landmarks had been closer to the flower during training. However, while the distance that the birds stopped from the landmarks and predicted flower location was unaffected by the distance that the landmarks moved between trials, the birds directed their search nearer to the predicted direction of the flower, relative to the landmarks, when the landmarks and flower were more stable in the environment. In the field, then, landmarks alone were sufficient for the birds to determine the distance of a reward but not its direction. [ABSTRACT FROM AUTHOR]

Evolutionary theories suggest that ecology is a major factor shaping cognition in primates. However, there have been few systematic tests of spatial memory abilities involving multiple primate species. Here, we examine spatial memory skills in four strepsirrhine primates that vary in level of frugivory: ruffed lemurs ( Varecia sp.), ring-tailed lemurs ( Lemur catta), mongoose lemurs ( Eulemur mongoz), and Coquerel's sifakas ( Propithecus coquereli). We compare these species across three studies targeting different aspects of spatial memory: recall after a long-delay, learning mechanisms supporting memory and recall of multiple locations in a complex environment. We find that ruffed lemurs, the most frugivorous species, consistently showed more robust spatial memory than the other species across tasks-especially in comparison with sifakas, the most folivorous species. We discuss these results in terms of the importance of considering both ecological and social factors as complementary explanations for the evolution of primate cognitive skills. [ABSTRACT FROM AUTHOR]

Arthropods are renowned for their navigational capabilities, with numerous examples known from insects and crustaceans. Early studies of amblypygids (Class Arachnida, Order Amblypygi) also suggest complex nocturnal navigation, despite their apparent lack of visual adaptations to the low-light conditions of a tropical understory. In a series of two studies, we use the tropical amblypygid, Phrynus pseudoparvulus, to assess their nocturnal homing ability. Our first experiment displaced and tracked resident and nonresident individuals. Resident individuals, displaced up to 4.5 m from their home refuges and released onto their home tree, were more likely to return to their previously occupied refuge than were nonresident individuals that were collected from trees outside the study area and released at the same locations. In a follow-up study, we displaced amblypygids longer distances (6-8.7 m) from their home trees and tracked them by telemetry. These individuals returned to home trees, typically within 1-3 nights, often via indirect paths. Taken together, our results provide evidence that P. pseudoparvulus are able to navigate home, often taking indirect routes, and can do so through a mechanism other than path integration. [ABSTRACT FROM AUTHOR]

Understanding animals' spatial perception is a critical step toward discerning their cognitive processes. The spatial sense is multimodal and based on both the external world and mental representations of that world. Navigation in each species depends upon its evolutionary history, physiology, and ecological niche. We carried out foraging experiments on wild vervet monkeys ( Chlorocebus pygerythrus) at Lake Nabugabo, Uganda, to determine the types of cues used to detect food and whether associative cues could be used to find hidden food. Our first and second set of experiments differentiated between vervets' use of global spatial cues (including the arrangement of feeding platforms within the surrounding vegetation) and/or local layout cues (the position of platforms relative to one another), relative to the use of goal-object cues on each platform. Our third experiment provided an associative cue to the presence of food with global spatial, local layout, and goal-object cues disguised. Vervets located food above chance levels when goal-object cues and associative cues were present, and visual signals were the predominant goal-object cues that they attended to. With similar sample sizes and methods as previous studies on New World monkeys, vervets were not able to locate food using only global spatial cues and local layout cues, unlike all five species of platyrrhines thus far tested. Relative to these platyrrhines, the spatial location of food may need to stay the same for a longer time period before vervets encode this information, and goal-object cues may be more salient for them in small-scale space. [ABSTRACT FROM AUTHOR]

Safety in numbers is thought to be the principal advantage of living in groups for many species. The group can only provide protection against predators, however, when group cohesion is maintained. Vocalisations are used to monitor inter-individual distances, especially under conditions of poor visibility, but should be avoided in the presence of predators. Mentally tracking the movements of silent and invisible group members would allow animals foraging in dense vegetation to stay close to their group members while reducing the use of vocal contact. We tested the socio-spatial cognitive abilities of wild vervet monkeys ( Chlorocebus pygerythrus) by comparing their reactions to plausible and implausible displacements of group members simulated by sound playbacks. Our methods are comparable to those used in studies of 'object permanence' and 'invisible displacements' of inanimate objects. Our results show that vervets can track the whereabouts of invisibly and silently moving group members, at least over short periods of time. [ABSTRACT FROM AUTHOR]

Some researchers have recently argued that humans may be unusual among primates in preferring to use landmark information when reasoning about some kinds of spatial problems. Some have explained this phenomenon by positing that our species' tendency to prefer landmarks stems from a human-unique trait: language. Here, we test this hypothesis-that preferring to use landmarks to solve such tasks is related to language ability-by exploring landmark use in a spatial task in one non-human primate, the brown capuchin monkey ( Cebus apella). We presented our subjects with the rotational displacement task, in which subjects attempt to relocate a reward hidden within an array of hiding locations which are subsequently rotated to a new position. Over several experiments, we varied the availability and the salience of a landmark cue within the array. Specifically, we varied (1) visual access to the array during rotation, (2) the type of landmark, (3) the consistency of the landmark qualities, and (4) the amount of exposure to the landmark. Across Experiments 1 through 4, capuchins did not successfully use landmarks cues, suggesting that non-linguistic primates may not spontaneously use landmarks to solve some spatial problems, as in this case of a small-scale dynamic spatial task. Importantly, we also observed that capuchins demonstrated some capacity to learn to use landmarks in Experiment 4, suggesting that non-linguistic creatures may be able to use some landmarks cues in similar spatial tasks with extensive training. [ABSTRACT FROM AUTHOR]

The task of determining an optimal route to several locations is called the traveling salesperson problem (TSP). The TSP has been used recently to examine spatial cognition in humans and non-human animals. It remains unclear whether or not the decision process of animals other than non-human primates utilizes rigid rule-based heuristics, or whether non-human animals are able to flexibly 'plan' future routes/behavior based on their knowledge of multiple locations. We presented pigeons in a One-way and Round-Trip group with TSPs that included two or three destinations (feeders) in a laboratory environment. The pigeons departed a start location, traveled to each feeder once before returning to a final destination. Pigeons weighed the proximity of the next location heavily, but appeared to plan ahead multiple steps when the travel costs for inefficient behavior appeared to increase. The results provide clear and strong evidence that animals other than primates are capable of planning sophisticated travel routes. [ABSTRACT FROM AUTHOR]

Investigations made into the cognitive decision making of honeybees ( Apis mellifera) traveling from one flower patch to another flower patch (interpatch foraging) are few. To facilitate such research, we present methods to artificially emulate interpatch foraging and quantify the immediate decision making of honeybees (within 50 cm) choosing to fly an interpatch path. These 'Interpatch Methods' are validated, applied, and shown to produce novel information for the field of honeybee spatial cognition. Generally, we demonstrate that a single foraging cohort of honeybees is shown to be capable of making decisions based upon different sets of learned cues, in the exact same context. Specifically, both terminal beacon orientation cues and compass navigation cues can guide the cognitive decision making of interpatch foraging honeybees; our bees chose both cues equally. Finally, the theoretical importance of decision making for interpatch paths is compared with the other foraging paths (outward and homing) with respect to the information available to recruited foragers and scout foragers. We conclude that the ability to analyze interpatch foraging is critical for a more complete understanding of honeybee foraging cognition and that our methods are capable of providing that understanding. [ABSTRACT FROM AUTHOR]

We investigated whether chimpanzees, bonobos, and orangutans encoded the location of a reward hidden underneath one of three identical cups in relation to (1) the other cups in the array-i.e., the relative position of the baited cup within the array; or (2) the landmarks surrounding the cups-e.g., the edge of the table. Apes witnessed the hiding of a food reward under one of three cups forming a straight line on a platform. After 30 s, they were allowed to search for the reward. In three different experiments, we varied the distance of the cups to the edge of the platform and the distance between the cups. Results showed that both manipulated variables affected apes' retrieval accuracy. Subjects' retrieval accuracy was higher for the outer cups compared with the Middle cup, especially if the outer cups were located next to the platform's edge. Additionally, the larger the distance between the cups, the better performance became. [ABSTRACT FROM AUTHOR]

Fish live in three-dimensional environments, through which they swim with three translational and three rotational degrees of freedom. Navigating through such environments is recognised as a difficult problem, yet fish, and other animals that swim and fly, achieve this regularly. Despite this, the vast majority of research has considered how animals navigate horizontally from place to place and has ignored the vertical component. Here, we test the importance of the vertical axis of space for fish solving a three-dimensional spatial cognition task. We trained banded tetras ( Astyanax fasciatus) to learn the route towards a goal in a rotating Y-maze in which the arms led either up and left or down and right in an environment that allowed access to visual landmarks providing horizontal and vertical information. Our results revealed that the landmarks increased navigational efficiency during training. However, these landmarks were ignored when the horizontal and vertical components were placed in conflict with each other by rotating the maze 90° during testing. From this surprising result, we conclude that the cues that are present in the vertical axis (presumably hydrostatic pressure) override landmark cues that have been shown to be salient in experiments that only consider the horizontal component of space. [ABSTRACT FROM AUTHOR]

We investigated reasoning about spatial relational similarity in three great ape species: chimpanzees, bonobos, and orangutans. Apes were presented with three spatial mapping tasks in which they were required to find a reward in an array of three cups, after observing a reward being hidden in a different array of three cups. To obtain a food reward, apes needed to choose the cup that was in the same relative position (i.e., on the left) as the baited cup in the other array. The three tasks differed in the constellation of the two arrays. In Experiment 1, the arrays were placed next to each other, forming a line. In Experiment 2, the positioning of the two arrays varied each trial, being placed either one behind the other in two rows, or next to each other, forming a line. Finally, in Experiment 3, the two arrays were always positioned one behind the other in two rows, but misaligned. Results suggested that apes compared the two arrays and recognized that they were similar in some way. However, we believe that instead of mapping the left-left, middle-middle, and right-right cups from each array, they mapped the cups that shared the most similar relations to nearby landmarks (table's visual boundaries). [ABSTRACT FROM AUTHOR]

Vertebrates use geometric and featural information for spatial navigation. When both geometric and featural cues are available, animals can use a variety of spatial strategies based on this information. To examine the nature of these strategies, we manipulated the spatial relationship between a conspicuous cue and the position of the goal when goldfish (Carassius auratus) were searching for the exit of a rectangular environment with one distinctive wall. Two groups of fish were used, one with the distinctive wall close to the goal and the other with the distinctive wall on the other end of the enclosure. Results showed that fish encoded featural and geometric information in both conditions but the spatial relationship between the goal and the distinctive wall influences the characteristics of the encoding of the spatial cues and the strategy used to locate the goal. These results suggest that fish in both procedures use the local featural cues associated with the goal instead of the whole set of spatial cues as previous studies propose. [ABSTRACT FROM AUTHOR]

Knowing that objects continue to exist after disappearing from sight and tracking invisible object displacements are two basic elements of spatial cognition. The current study compares dogs and apes in an invisible transposition task. Food was hidden under one of two cups in full view of the subject. After that both cups were displaced, systematically varying two main factors, whether cups were crossed during displacement and whether the cups were substituted by the other cup or instead cups were moved to new locations. While the apes were successful in all conditions, the dogs had a strong preference to approach the location where they last saw the reward, especially if this location remained filled. In addition, dogs seem to have especial difficulties to track the reward when both containers crossed their path during displacement. These results confirm the substantial difference that exists between great apes and dogs with regard to mental representation abilities required to track the invisible displacements of objects. [ABSTRACT FROM AUTHOR]

Wild animals face the challenge of locating feeding sites distributed across broad spatial and temporal scales. Spatial memory allows animals to find a goal, such as a productive feeding patch, even when there are no goal-specific sensory cues available. Because there is little experimental information on learning and memory capabilities in free-ranging primates, the aim of this study was to test whether grey mouse lemurs ( Microcebus murinus), as short-term dietary specialists, rely on spatial memory in relocating productive feeding sites. In addition, we asked what kind of spatial representation might underlie their orientation in their natural environment. Using an experimental approach, we set eight radio-collared grey mouse lemurs a memory task by confronting them with two different spatial patterns of baited and non-baited artificial feeding stations under exclusion of sensory cues. Positional data were recorded by focal animal observations within a grid system of small foot trails. A change in the baiting pattern revealed that grey mouse lemurs primarily used spatial cues to relocate baited feeding stations and that they were able to rapidly learn a new spatial arrangement. Spatially concentrated, non-random movements revealed preliminary evidence for a route-based restriction in mouse lemur space; during a subsequent release experiment, however, we found high travel efficiency in directed movements. We therefore propose that mouse lemur spatial memory is based on some kind of mental representation that is more detailed than a route-based network map. [ABSTRACT FROM AUTHOR]

Previous studies in our laboratory have shown that when presented with a sudden stimulus simulating an oncoming predator, Mongolian gerbils can compute the optimal trajectory to a safe refuge, taking into account the position of the threat, the location of a clearly visible refuge, and several other contextual variables as well. In the present studies, the main goal was to explore the abilities of gerbils to use mental representations of spaces that were visually occluded by opaque barriers to compute efficient escape trajectories. In all studies, gerbils were placed into a round open field containing a single refuge. On each trial, an overhead visual stimulus was caused to ‘fly’ overhead, eliciting robust escape movements from the gerbils. By manipulating the shape and position of a series of opaque barriers that were interposed between the gerbils and the refuge, we were able to show that gerbils can compute the shortest route to an invisible target, even when the available routes to the target are made complex by using elaborate barrier shapes. These findings suggest that gerbils can maintain representations of their locations with respect to salient environmental landmarks and refuges, even when such locations are not continuously visible. [ABSTRACT FROM AUTHOR]

We investigated whether two dogs that had been specially trained to retrieve objects by their names were able to integrate information about the identity ( What) as well as the location ( Where) of those objects so that they could plan their search accordingly. In a first study, two sets of objects were placed in two separate rooms and subjects were asked to retrieve the objects, one after the other. Both dogs remembered the identity of the objects as they reliably retrieved the correct objects. One of the dogs was also able to integrate information about the object’s location as he chose the correct location in which the object had been placed. Further investigation of the second dog’s behavior revealed that she followed a more stereotyped search strategy. Despite this variation in performance, this study provides evidence for the memory of What and Where in a domestic dog and shows the prospective use of such information in a search task. [ABSTRACT FROM AUTHOR]

This study was aimed at uncovering physical and geometric properties that make a particular landmark a target of exploration and navigation. Rats were tested in a square open-field arena with additional portable corners featuring the same properties as the arena corners. It was found that the routes of progression converged upon the added corners, whether located at the arena wall or the arena center. Route convergence upon the added corners involved numerous visits to these corners. However, time spent at the added corners was relatively short compared with the arena corners, including that from which rats were introduced into the arena. There was no differential effect of testing rats in light or dark, or with a low versus a high portable corner. It is suggested that the added corners were distinct against the background of the arena enclosure, whereas the four arena corners and walls were encoded by the rats as one geometric module. This distinctness, together with the greater accessibility of the added corners, made them salient landmarks and a target of exploration. Thus, the impact of a landmark extended beyond its specific self-geometry to include accessibility and distinctness, which are contextual properties. In addition to the contextual impact on locomotor behavior there was also a temporal effect, with security initially dominating the rats’ behavior but then declining along with an increased attraction to salient landmarks. These spatiotemporal patterns characterized behavior in both lit and dark arenas, indicating that distal cues were secondary to local proximal cues in shaping routes. [ABSTRACT FROM AUTHOR]

The nature of spatial information used for memorizing and recalling places is largely unclear. Earlier studies tested integration of geometric and feature information mostly during reorientation in artificial environments without including time as a memory-critical component. Here, we tested a harbour seal in a delayed matching-to-sample task (DMTS) in a familiar environment under two spatial multiple-choice conditions. The feature condition consisted of a DMTS task with four comparison stimuli presented on fixed positions in a classic matching apparatus and was designed to make stimulus features the most prominent information. The landscape condition consisted of a DMTS task in a familiar environment with four places marked by comparison stimuli and allowed the use of all available spatial information including geometrical and feature information. The seal’s performance was impaired by delays of 3, 6, 9 or 12 s only in the feature condition; a delay of 12 s resulted in chance level performance. Replacing the comparison stimuli at the apparatus with identical spheres resulted in impaired performance. Performance in the landscape condition was impaired neither by delays nor by replacing comparison stimuli with spheres. Landscape information obviously was encoded redundantly and could be recalled more reliably and longer than feature information, which reveals feature information to be a less valuable type of spatial information for memorizing and recalling places. [ABSTRACT FROM AUTHOR]

Human subjects represent the location of a point in 2D space using two independent dimensions (x-y in Euclidean or radius-angle in polar space), and encode location in memory along these dimensions using two levels of representation: a fine-grain value and a category. Here we determined whether monkeys possessed the ability to represent location with these two levels of coding. A rhesus monkey was trained to reproduce the location of a dot in a circle by pointing, after a delay period, on the lo- cation where a dot was presented. Five different delay periods (0.5-5 s) were used. The results showed that the monkey used a polar coordinate system to represent the fine-grain spatial coding, where the radius and angle of the dots were encoded independently. The variability of the spatial response and reaction time increased with longer delays. Furthermore, the animal was able to form a categorical representation of space that was delay-dependent. The responses avoided the circumference and the center of the circle, defining a categorical radial prototype around one third of the total radial length. This radial category was observed only at delay durations of 3-5 s. Finally, the monkey also formed angular categories with prototypes at the obliques of the quadrants of the circle, avoiding the horizontal and vertical axes. However, these prototypes were only observed at the 5-s delay and on dots lying on the circumference. These results indicate that monkeys may possess spatial cognitive abilities similar to humans. [ABSTRACT FROM AUTHOR]

We report experiments based on a novel test in domestic chicks (Gallus gallus), designed to examine the encoding of two different geometric features of an enclosed environment: relative lengths of the walls and amplitude of the corners. Chicks were trained to search for a food reward located in one corner of a parallelogram-shaped enclosure. Between trials, chicks were passively disoriented and the enclosure was rotated, making reorientation possible only on the basis of the internal spatial structure of the enclosure. In order to reorient, chicks could rely on two sources of information: the relative lengths of the walls of the enclosure (associated to their left-right sense order) and the angles subtended by walls at corners. Chicks learned the task choosing equally often the reinforced corner and its rotational equivalent. Results of tests carried out in novel enclosures, the shapes of which were chosen ad hoc (1) to induce reorientation based only on the ratio of walls lengths plus sense (rectangular enclosure), or (2) to induce reorientation based only on corner angles (rhombus-shaped enclosure), suggested that chicks encoded both features of the environment. In a third test, in which chicks faced a conflict between these geometric features (mirror parallelogram-shaped enclosure), reorientation seemed to depend on the salience of corner angles. These results shed light on the elements of the environmental geometry which control spatial reorientation, and broaden the knowledge on the geometric representation of space in animals. [ABSTRACT FROM AUTHOR]