Incidental encoding of enclosure geometry does not require visual input: evidence from blindfolded adults

Although spatial orientation with respect to the geometric properties of an environment appears to be an ability shared across various species, debate remains concerning potential similarities and differences with respect to the underlying mechanism(s). One prominent theoretical account of orientation with respect to the environment suggests that participants match visual memories to their current visual perception and navigate to reduce the discrepancy between the two. We tested whether visual input was necessary to incidentally encode the geometric properties of an environment, by training disoriented and blindfolded adult participants to search by touch for a target object hidden in one of four locations, marked by distinctive textural cues, located in the corners of a rectangular enclosure. Following training, we removed the distinctive textural cues and probed the extent to which the participants had learned the geometry of the enclosure. Even in the absence of vision and unique textural cues, search behavior was consistent with evidence for the encoding of enclosure geometry. A follow-up experiment in which participants were trained in a rectangular enclosure but were tested in a square enclosure provided converging evidence that search behavior was influenced by the geometric properties of the enclosure. Collectively, these results suggest that even in the absence of vision, participants incidentally encoded the geometric properties of the enclosure, indicating that visual input is not required to encode the geometric properties of an environment.     

Reorientation in a two-dimensional environment: II. Do pigeons (Columba livia) encode the featural and geometric properties of a two-dimensional schematic of a room?

Pigeons (Columba livia) searched for a hidden target area in images showing a schematic rectangular environment. The absolute position of the goal varied across trials but was constant relative to distinctive featural cues and geometric properties of the environment. Pigeons learned to use both of these properties to locate the goal. Transformation tests showed that pigeons could use either the color or shape of the features, but performance was better with color cues present. Pigeons could also use a single featural cue at an incorrect corner to distinguish between the correct corner and the geometrically equivalent corner; this indicates that they did not simply use the feature at the correct corner as a beacon. Interestingly, pigeons that were trained with features spontaneously encoded geometry. The encoded geometric information withstood vertical translations but not orientation transformations.     

Geometric orientation by humans: angles weigh in

Human participants were trained to navigate to two geometrically equivalent corners of a parallelogram-shaped virtual environment. The unique shape of the environment combined three distinct types of geometric information that could be used in combination or in isolation to orient and locate the goals: the angular amplitudes of the corners, the relative wall length relationships, and the principal axis of symmetry. In testing, participants were placed in manipulated versions of the training environment that tested which types of geometry they had encoded and how angular information weighed in against the other two geometric properties. The test environments were (a) a rectangular environment that removed the angular information, (b) a rhombic environment that removed wall length information and drastically reduced the principal axis, and (c) a reverse-parallelogram-shaped environment that placed angular information against both wall length and principal axis information. Participants chose accurately in the rectangular and rhombus environments, despite the removal of one of the cues. In the conflict test, participants preferred corners with the correct angular amplitudes over corners that were correct according to both wall length relationships and the principal axis. These results are comparable to recent findings with pigeons and suggest that angles are a salient orientation cue for humans.     

Finding the best angle: pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>) weight angular information more heavily than relative wall length in an open-field geometry task

Pigeons were trained to locate food in two geometrically equivalent corners of a parallelogram-shaped enclosure. Both the angular amplitude of the corners and the length of the walls alone were sufficient for successfully completing the task. Following training, birds were tested in three separate conditions that manipulated the geometric information available. During tests in both a rectangular-shaped enclosure that preserved the wall length information but not the angular amplitude, and a rhombus-shaped enclosure that did the opposite, pigeons located their goal corners with a high degree of accuracy, indicating an ability to use both types of geometric information in isolation. This result is consistent with prior research with domestic chicks. However, in a conflict test in a reverse parallelogram-shaped enclosure, in which the correct angular location was paired with an incorrect wall length location, birds showed a preference for the correct angular location. This suggests that pigeons weight angles more heavily than wall lengths in this type of navigation task, which differs from findings in a similar task conducted with the domestic chick. Results in the conflict test also suggest that pigeons did not use the principal axis as their main strategy of small-scale navigation.     

Reorientation in diamond-shaped environments: encoding of features and angles in enclosures versus arrays by adult humans and pigeons (Columbia livia)

Although geometric reorientation has been extensively studied in numerous species, most research has been conducted in enclosed environments and has focused on use of the geometric property of relative wall length. The current studies investigated how angular information is used by adult humans and pigeons to orient and find a goal in enclosures or arrays that did not provide relative wall length information. In enclosed conditions, the angles formed a diamond shape connected by walls, whereas in array conditions, free-standing angles defined the diamond shape. Adult humans and pigeons were trained to locate two geometrically equivalent corners, either the 60° or 120° angles. Blue feature panels were located in the goal corners so that participants could use either the features or the local angular information to orient. Subsequent tests in manipulated environments isolated the individual cues from training or placed them in conflict with one another. In both enclosed and array environments, humans and pigeons were able to orient when either the angles or the features from training were removed. On conflict tests, female, but not male, adult humans weighted features more heavily than angular geometry. For pigeons, angles were weighted more heavily than features for birds that were trained to go to acute corners, but no difference in weighting was seen for birds trained to go to obtuse corners. These conflict test results were not affected by environment type. A subsequent test with pigeons ruled out an interpretation based on exclusive use of a principal axis rather than angle. Overall, the results indicate that, for both adult humans and pigeons, angular amplitude is a salient orientation cue in both enclosures and arrays of free-standing angles.     

The impact of landmark properties in shaping exploration and navigation

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.     

Representation of two geometric features of the environment in the domestic chick (<Emphasis Type="Italic">Gallus gallus</Emphasis>)

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.     

Coding location in enclosed spaces: is geometry the principle?

Both animals and human toddlers can find an object in a rectangular enclosure after they have been disoriented. They use geometric cues (relative lengths of walls) to discriminate among different corners (e.g. long wall to the left, short to the right). It has been claimed that this ability is 'modular', i.e. exclusively geometric. The present study demonstrates that the ability toddlers exhibit is a more general one, namely, an ability to discriminate relative quantity. Using a square enclosure, we show that toddlers use the relative sizes of the figures on different walls to characterize different corners. We also show that they do not use simple non-relative features to distinguish different corners. Possible reasons for differences in the ability to use relative versus non-relative cues are discussed.     

Is there an innate geometric module? Effects of experience with angular geometric cues on spatial re-orientation based on the shape of the environment

Non-human animals and human children can make use of the geometric shape of an environment for spatial reorientation and in some circumstances reliance on purely geometric information (metric properties of surfaces and sense) can overcome the use of local featural cues. Little is known as to whether the use of geometric information is in some way reliant on past experience or, as would likely be argued by advocates of the notion of a geometric module, it is innate. We tested the navigational abilities of newborn domestic chicks reared in either rectangular or circular cages. Chicks were trained in a rectangular-shaped enclosure with panels placed at the corners to provide salient featural cues. Rectangular-reared and circular-reared chicks proved equally able to learn the task. When tested after removal of the featural cues, both rectangular- and circular-reared chicks showed evidence that they had spontaneously encoded geometric information. Moreover, when trained in a rectangular-shaped enclosure without any featural cues, chicks reared in rectangular-, circular-, or c-shaped cages proved to be equally able to learn and perform the task using geometric information. These results suggest that effective use of geometric information for spatial reorientation does not require experience in environments with right angles and metrically distinct surfaces, thus supporting the hypothesis of a predisposed geometric module in the animal brain.     

Reorientation in a two-dimensional environment: I. Do adults encode the featural and geometric properties of a two-dimensional schematic of a room?

Adults searched for a goal in images of a rectangular environment. The goal's position was constant relative to featural and geometric cues, but the absolute position changed across trials. Participants easily learned to use the featural cues to find the target, but learning to use only geometric information was difficult. Transformation tests revealed that participants used the color and shape of distinct features to encode the goal's position. When the features at the correct and geometrically equivalent corners were removed, participants could use distant features to locate the goal. Accuracy remained above chance when a single distant feature was present, but the feature farthest from the goal yielded lower accuracy than one closer. Participants trained with features spontaneously encoded the geometric information. However, this representation did not withstand orientation transformations.     

Modularity as a fish (Xenotoca eiseni) views it: conjoining geometric and nongeometric information for spatial reorientation

When disoriented in a closed rectangular tank, fish (Xenotoca eiseni) reoriented in accord with the large-scale shape of the environment, but they were also able to conjoin geometric information with nongeometric properties such as the color of a wall or the features provided by panels located at the corners of the tank. Fish encoded geometric information even when featural information sufficed to solve the spatial task. When tested after transformations that altered the original arrangement of the panels, fish were more affected by those transformations that modified the geometric relationship between the target and the shape of the environment. Finally, fish appeared unable to use nongeometric information provided by distant panels. These findings show that a reorientation mechanism based on geometry is widespread among vertebrates, though the joint use of geometric and nongeometric cues by fish suggest that the degree of information encapsulation of the mechanism varies considerably between species.     

Whither geometry? Troubles of the geometric module

In rectangular arenas, rats often confuse diagonally opposite corners, even when distinctive cues differentiate them. This led to the postulation that rats rely preferentially on the geometry of space, encoded in a dedicated geometric module. Recent research casts doubt on this idea. Distinctive featural cues such as entire walls of a distinct color can hinder or aid the learning of geometry. In one situation in which using geometry would help greatly, rats had trouble learning the task. An associative model has been developed to capture these different learning processes, and view-based matching has been proposed as an alternative to the explicit coding of geometric cues. Considerations about how cues interact in learning are crucial in a recent theory of human spatial cognition.     

Pigeons encode absolute distance but relational direction from landmarks and walls

In recent studies, researchers have examined animals' use of absolute or relational distances in finding a hidden goal. When trained with an array of landmarks, most animals use the default strategy of searching at an absolute distance from 1 or more landmarks. In contrast, when trained in enclosures, animals often use the relationship among walls. In the present study, pigeons were trained to find the center of an array of landmarks or a set of short walls that did not block external cues. Expansion tests showed that both groups of pigeons primarily used an absolute distance strategy. However, on rotational tests, pigeons continued to search in the center of the array, suggesting that direction was learned in relation to array.     

Multiple landmarks, the encoding of environmental geometry and the spatial logics of a dual brain

A series of place learning experiments was carried out in young chicks (Gallus gallus) in order to investigate how the geometry of a landmark array and that of a walled enclosure compete when disoriented animals could rely on both of them to re-orient towards the centre of the enclosure. A square-shaped array (four wooden sticks) was placed in the middle of a square-shaped enclosure, the two structures being concentric. Chicks were trained to ground-scratch to search for food hidden in the centre of the enclosure (and the array). To check for effects of array degradation, one, two, three or all landmarks were removed during test trials. Chicks concentrated their searching activity in the central area of the enclosure, but their accuracy was inversely contingent on the number of landmarks removed; moreover, the landmarks still present within the enclosure appeared to influence the shape of the searching patterns. The reduction in the number of landmarks affected the searching strategy of chicks, suggesting that they had focussed mainly on local cues when landmarks were present within the enclosure. When all the landmarks were removed, chicks searched over a larger area, suggesting an absolute encoding of distances from the local cues and less reliance on the relationships provided by the geometry of the enclosure. Under conditions of monocular vision, chicks tended to rely on different strategies to localize the centre on the basis of the eye (and thus the hemisphere) in use, the left hemisphere attending to details of the environment and the right hemisphere attending to the global shape.This contribution is part of the special issue “Animal Logics” (Watanabe and Huber 2006).     

Enclosure size and the use of local and global geometric cues for reorientation

Multiple spatial cues are utilized to orient with respect to the environment, but it remains unclear why feature (i.e., objects in the environment) and geometric (i.e., shape of the environment) cues are differentially influenced by enclosure size, and the extent to which local (i.e., wall lengths and corner angles) and global (i.e., principal axis of space) geometric cues are influenced by enclosure size. In the present study, we investigated the extent to which environmental size influenced the use of corner angle (i.e., a local geometric cue) and the principal axis of space (i.e., a global geometric cue) for reorientation. We developed an orientation task that allowed the manipulation of enclosure size during training and the isolation of the use of the principal axis of space during testing. Participants were trained to respond to a location in either a small or a large trapezoid-shaped enclosure uniquely specified by both local (i.e., wall lengths and corner angles) and global (i.e., principal axis of space) geometric cues. During testing, we presented both groups with a small and large rectangle (to assess the use of principal axis of space) and a small and large parallelogram (to asses relative use of corner angles and the principal axis of space when in conflict). Enclosure size influenced the relative use of corner angles but not of the principal axis of space. Results suggest that corner angles function like features and that changes in the use of feature cues are the source of the relative reliance on feature and geometric cues during changes of enclosure size.     

Experience and geometry: controlled-rearing studies with chicks

Animals can reorient making use of the geometric shape of an environment, i.e., using sense and metric properties of surfaces. Animals reared soon after birth either in circular or in rectangular enclosures (and thus affording different experiences with metric properties of the spatial layout) showed similar abilities when tested for spatial reorientation in a rectangular enclosure. Thus, early experience in environments with different geometric characteristics does not seem to affect animals’ ability to reorient using sense and metric information. However, some results seem to suggest that when geometric and non-geometric information are set in conflict, rearing experience could affect the relative dominance of featural (landmark) and geometric information. In three separate experiments, newborn chicks reared either in circular- or in rectangular-shaped home-cages were tested for spatial reorientation in a rectangular enclosure, with featural information provided either by panels at the corners or by a blue-coloured wall. At test, when faced with affine transformations in the arrangement of featural information that contrasted with the geometric information, chicks showed no evidence of any effect of early experience on their relative use of geometric and featural information for spatial reorientation. These findings suggest that, at least for this highly precocial species, the ability to deal with geometry seems to depend more on predisposed mechanisms than on learning and experience after hatching.     

Orientation in trapezoid-shaped enclosures: implications for theoretical accounts of geometry learning

Human participants learned to select 1 of 4 distinctively marked corners in a rectangular virtual enclosure. After training, control and test trials were interspersed with training trials. On control and test trials, all markers were equivalent in color, but only during test trials was the shape of the enclosure manipulated. Specifically, for each test trial, a single long wall or short wall of the enclosure increased twice as long as or decreased half as long as that present in the training enclosure. These manipulations produced 8 unique trapezoid-shaped enclosures. Participants were allowed to select 1 corner during control and test trials. Performance during control trials revealed that participants selected the correct and rotationally equivalent locations. Performance during test trials revealed that participants selected locations in trapezoid-shaped enclosures that were consistent with those predicted by global geometry (i.e., principal axis of space) but were inconsistent with those predicted by local geometry (i.e., proportion of rewarded training features present at a location). Results have implications for theoretical accounts of geometry learning.     

Context specificity of operant discriminative performance in pigeons: II. Necessary and sufficient conditions

Six experiments were performed to explore the necessary and sufficient conditions for producing context specificity of discriminative operant performance in pigeons. In Experiment 1, pigeons learned a successive discrimination (red S+/blue S−) in two chambers that had a particular odor present and between which they were frequently switched. The birds subsequently learned the reversal (blue S+/ red S−) in one of these chambers with a different odor present. When switched to the alternative chamber, although the odor and the reinforcement contingency were still appropriate to the reversal, performance appropriate to the original discrimination recurred in subjects for which the houselights were on during training and testing but not for those for which the houselights were off. This indicated the importance of visual contextual cues in producing context specificity. Experiment 2 showed that the frequent switching between boxes in initial training was of no consequence, presumably because the apparatus cues were highly salient to the subjects. Experiment 3 showed significantly less context specificity when odor cues were omitted. Experiment 4 showed that simply using a different reinforced stimulus in each phase of training was ineffective in producing context specificity. Experiment 5 showed that the generalization test procedure used in Experiment 4 was sensitive to context specificity when discrimination-reversal training was used with different odors in the two training phases. Experiment 6 replicated the results of Experiment 4, but then showed that when different odors accompanied the two training phases, context specificity was obtained with the single-stimulus paradigm. Thus in both single-stimulus and discrimination-reversal paradigms, redundant odor cues potentiated learning about apparatus cues.     

Spatial and feature-based memory representation in free-flying homing pigeons

The memory-based recognition of a goal is a capacity well demonstrated in birds, and understanding this ability often involves determining the relative importance of spatial and feature information in representing the properties of a goal. However, surprisingly little avian research has examined goal recognition in a field setting. Here, we demonstrate that homing pigeons can be successfully trained outdoors to fly to and land on a goal platform located in an array of other platforms at a distance on the order of 100 m. They can do so under conditions when the properties of the goal are stable in time as well as when the properties of the goal periodically change; the latter condition indicating that homing pigeons can rapidly adapt their memory representations to take into account changing environmental conditions. When probed for preferential use of either spatial (location) or feature-based (color) information, the pigeons demonstrated an indifferent capacity to use both independent of task demands. The homing pigeon memory systems that support goal recognition appear to be opportunistic with respect to the information exploited, using spatial and feature information equally to guide their behavior. Therefore, and despite strong natural and artificial selection for navigational ability, spatial information is no more important than feature cues in representing a goal location for homing pigeons flying outdoors.     

The contribution of monocular depth cues to scene perception by pigeons

The contributions of different monocular depth cues to performance of a scene perception task were investigated in 4 pigeons. They discriminated the sequential depth ordering of three geometric objects in computer-rendered scenes. The orderings of these objects were specified by the combined presence or absence of the pictorial cues of relative density, occlusion, and relative size. In Phase 1, the pigeons learned the task as a direct function of the number of cues present. The three monocular cues contributed equally to the discrimination. Phase 2 established that differential shading on the objects provided an additional discriminative cue. These results suggest that the pigeon visual system is sensitive to many of the same monocular depth cues that are known to be used by humans. The theoretical implications for a comparative psychology of picture processing are considered.