Multiple-landmark piloting in pigeons (Columba livia): landmark configuration as a discriminative cue

Using the landmark-transformation technique, researchers have shown that pigeons (Columba livia) tend to encode a goal location relative to 1 landmark, even when multiple landmarks are in the vicinity of the goal. The current experiments examined pigeons' ability to use configural information from a set of landmarks by making the arrangement of 4 landmarks a discriminative cue to the location of buried seeds. Results showed that pigeons used information from the 3 consistently placed landmarks to search accurately when 1 landmark was displaced. Findings indicate that pigeons are able to search for a goal using information from multiple landmarks instead of just 1 and that landmark use by these birds may be more flexible than previously theorized.     

Labeling and family resemblance in the discrimination of polymorphous categories by pigeons

Two experiments examined whether pigeons discriminate polymorphous categories on the basis of a single highly predictive feature or overall similarity. In the first experiment, pigeons were trained to discriminate between categories of photographs of complex real objects. Within these pictures, single features had been manipulated to produce a highly salient texture cue. Either the picture or the texture provided a reliable cue for discrimination during training, but in probe tests, the picture and texture cues were put into conflict. Some pigeons showed a significant tendency to discriminate on the basis of the picture cue (overall similarity or family resemblance), whereas others appeared to rely on the manipulated texture cue. The second experiment used artificial polymorphous categories in which one dimension of the stimulus provided a completely reliable cue to category membership, whereas three other dimensions provided cues that were individually unreliable but collectively provided a completely reliable basis for discrimination. Most pigeons came under the control of the reliable cue rather than the unreliable cues. A minority, however, came under the control of single dimensions from the unreliable set. We conclude that cue salience can be more important than cue reliability in determining what features will control behavior when multiple cues are available.     

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.     

Internal sense of direction and landmark use in pigeons (Columba livia)

The relative importance of an internal sense of direction based on inertial cues and landmark piloting for small-scale navigation by White King pigeons (Columba livia) was investigated in an arena search task. Two groups of pigeons differed in whether they had access to visual cues outside the arena. In Experiment 1, pigeons were given experience with 2 different entrances and all pigeons transferred accurate searching to novel entrances. Explicit disorientation before entering did not affect accuracy. In Experiments 2-4, landmarks and inertial cues were put in conflict or tested 1 at a time. Pigeons tended to follow the landmarks in a conflict situation but could use an internal sense of direction to search when landmarks were unavailable.     

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.     

Sex differences and the effect of instruction on reorientation abilities by humans

This study examined whether differences in the amount of information provided to men and women, in the form of verbal instruction, influenced their encoding during a reorientation task. When a navigator needs to orient, featural (e.g., colour or texture) and geometry (e.g., metric information) are used to determine which direction to begin traveling. The current study used a spatial reorientation task to examine how men and women use featural and geometric cues and whether the content of the task’s instructions influenced how these cues were used. Participants were trained to find a target location in a rectangular room with distinctive objects situated at each corner. Once the participants were accurately locating the target, various tests manipulating the spatial information were conducted. We found both men and women encoded the featural cues, and even though the features provided reliable information, participants generally showed an encoding of geometry. However, when participants were not provided with any information about the spatial aspects of the task in the instructions, they failed to encode geometry. We also found that women used distant featural cues as landmarks when the featural cue closest to the target was removed, whereas men did not. Yet, when the two types of cues were placed in conflict, both sexes weighed featural cues more heavily than geometric cues. The content of the task instructions also influenced how cues were relied upon in this conflict situation. Our results have important implications for our understanding of how spatial cues are used for reorientation.     

Optimal combination of environmental cues and path integration during navigation

Navigation is influenced by body-based self-motion cues that are integrated over time, in a process known as path integration, as well as by environmental cues such as landmarks and room shape. In two experiments we explored whether humans combine path integration and environmental cues (Exp. 1: room shape; Exp. 2: room shape, single landmark, and multiple landmarks) to reduce response variability when returning to a previously visited location. Participants walked an outbound path in an immersive virtual environment before attempting to return to the path origin. Path integration and an environmental cue were both available during the outbound path, but experimental manipulations created single- and dual-cue conditions during the return path. The response variance when returning to the path origin was reduced when both cues were available, consistent with optimal integration predicted on the basis of Bayesian principles. The findings indicate that humans optimally integrate multiple spatial cues during navigation. Additionally, a large (but not a small) cue conflict caused participants to assign a higher weight to path integration than to environmental cues, despite the relatively greater precision afforded by the environmental cues.     

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.     

Flexible learning and use of multiple-landmark information by pigeons (Columba livia) in a touch screen-based goal searching task

Three touch screen-based experiments were conducted to investigate whether pigeons would learn to use configural information about a goal's location in relation to a multiple-landmark array. In Experiment 1, 4 pigeons (Columba livia) were trained to peck a computer monitor at a location that constituted the third vertex of a hypothetical triangle defined by 2 different landmarks. The landmarks appeared in 3 orientations during the training, and the pigeons' goal-searching ability easily transferred to the landmarks presented in 3 novel orientations. Each landmark was asymmetric, so we next examined whether the pigeons used (a) the small-scale, local orientation information that could be inferred from each landmark individually, or (b) the large-scale, configural information that could be inferred from the spatial arrangement of multiple landmarks taken as a whole. Even when each single landmark appeared by itself, the pigeons were able to locate the goal accurately, suggesting that the large-scale, configural information was not essential. However, when 1 landmark locally pointed to a location that was consistent with the triangular configuration and the other landmark locally pointed to a different location, the pigeons predominantly pecked at the configurally array-consistent location. These results suggest that the pigeons redundantly learned both the large-scale, configural strategy and the local, single-landmark strategy, but they mainly used the latter information, and used the former information solely to disambiguate conflicts when the 2 landmarks pointed toward different targets. Such flexible learning and use of redundant information may reflect the pigeons' adaptation to unstable wild environments during their evolutionary history.     

Landmark use by Clark’s nutcrackers (Nucifraga columbiana): influence of disorientation and cue rotation on distance and direction estimates

Many species have been shown to encode multiple sources of information to orient. To examine what kinds of information animals use to locate a goal we manipulated cue rotation, cue availability, and inertial orientation when the food-storing Clark’s nutcracker (Nucifraga columbiana) was searching for a hidden goal in a circular arena. Three groups of birds were used, each with a different goal–landmark distance. As the distance between the goal and the landmark increased, nutcrackers were less accurate in finding the correct direction to the goal than they were at estimating the distance (Experiment 1). To further examine what cues the birds were using to calculate direction, the featural cues within the environment were rotated by 90° and the birds were either oriented when searching (Experiments 2 and 3) or disoriented (Experiment 3). In Experiment 4, all distinctive visual cues were removed (both internal and external to the environment), a novel point of entry was used and the birds were either oriented or disoriented. We found that disorienting the nutcrackers so that they could not use inertial cues did not influence the birds’ total search error. The birds relied heavily but not completely on cues within the environment, as rotating available cues caused them to systematically shift their search behavior. In addition, the birds also relied to some extent on Earth-based cues. These results show the flexible nature of cue use by the Clark’s nutcracker. Our study shows how multiple sources of spatial information may be important for extracting multiple bearings for navigation.     

Learning your way around town: how virtual taxicab drivers learn to use both layout and landmark information

By having subjects drive a virtual taxicab through a computer-rendered town, we examined how landmark and layout information interact during spatial navigation. Subject-drivers searched for passengers, and then attempted to take the most efficient route to the requested destinations (one of several target stores). Experiment 1 demonstrated that subjects rapidly learn to find direct paths from random pickup locations to target stores. Experiment 2 varied the degree to which landmark and layout cues were preserved across two successively learned towns. When spatial layout was preserved, transfer was low if only target stores were altered, and high if both target stores and surrounding buildings were altered, even though in the latter case all local views were changed. This suggests that subjects can rapidly acquire a survey representation based on the spatial layout of the town and independent of local views, but that subjects will rely on local views when present, and are harmed when associations between previously learned landmarks are disrupted. We propose that spatial navigation reflects a hierarchical system in which either layout or landmark information is sufficient for orienting and wayfinding; however, when these types of cues conflict, landmarks are preferentially used.     

Influence of distal and proximal cues in encoding geometric information

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.     

Landmarks as beacons and associative cues: Their role in route learning

Relatively little is known about how people use the landmarks in their environment to learn routes. Landmarks are commonly regarded as associative cues--stimuli that enable recall of directional responses that lead closer to the navigator's goal. We contrast the function of landmark as associative cue with that of a beacon-a landmark near enough to a goal that moving toward it leads the navigator closer to his or her goal. In five experiments, participants learned a route through a simple desktop virtual environment. In the first three experiments, routes were learned better when their landmarks served as beacons than as associative cues. Two additional experiments showed that the acquired route knowledge depends on the function that landmarks serve during learning. Beacon-based route knowledge is less enduring and relatively less likely to involve knowledge of directions in the environment than is the route knowledge formed from landmarks that serve as associative 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.     

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.     

Gender-Based Navigation Stereotype Improves Men’s Search for a Hidden Goal

While a general stereotype exists that men are better at navigating than women, experimental evidence indicates that men and women differ in their use of spatial strategies, and this preference determines gender-differences. When both environmental geometry and landmark cues are available, men appear to learn to navigate using both types of cues, while women show a preference for using landmarks. Using a computer-generated task, 80 undergraduate students from North-East England learned to navigate to a hidden goal. Activating the general navigation stereotype improved the performance of men, compared to the control condition, both when only geometric cues and only landmark cues were present (stereotype lift), suggesting that activating a general stereotype can affect tasks both with (geometry) and without (landmark) established gender-differences in preference. In addition, in the test trial (hidden goal removed) women who learned to navigate using only landmarks spent longer in the correct location of the hidden goal than those who learned to navigate using only geometry. In contrast, the opposite result was found for men, suggesting that when only one cue-type is available, gender-differences still occur, with women better able to navigate using landmarks than geometry, while men seemed to learn more about the location of the goal with reference to geometric than landmark cues.     

Goldfish (Carassius auratus) matching geometric and featural cues: a reinterpretation of some of the data of Vargas, López, Salas, and Thinus-Blanc (2004)

Vargas, López, Salas, and Thinus-Blanc showed that goldfish (Carassius auratus) can use both geometric and featural cues in relocating a target corner in a rectangular enclosure. When featural cues (arrangement of striped walls) were put in conflict with geometric cues, results differed according to target location during training. Vargas, López, et al. explained the results of their cue conflict in terms of 2 different strategies: mapping and cue guidance. I provide an alternative, more parsimonious interpretation in which the same strategy of attempting to match as many cues as possible applies to both cases.     

Environment size and the use of feature and geometric cues for reorientation

We tested associative-based accounts of orientation by investigating the influence of environment size on the use of feature and geometric cues for reorientation. Two groups of participants were trained in dynamic three-dimensional virtual rectangular environments that differed in size to find a distinctly colored bin located at one of the four corners. Subsequently, we probed the reliance on feature and geometric cues for reorientation during test trials by presenting six trial types: Small Geometry Only, Large Geometry Only, Small Cue Conflict, Large Cue Conflict, Small Distal, and Large Distal. During Geometry Only test trials, all bins were black; thus, all distinctive featural information was removed leaving only geometric cues. For Cue Conflict test trials, all colored bins were shifted counter-clockwise one corner; thus, the geometric cues from the trained corner and the trained color were in direct conflict. During Distal test trials, the bin in the geometrically incorrect corner farthest from the trained corner was colored the same as during training; the remaining three bins were black. Thus, only this distant feature cue could be used to determine the location of the goal bin. Results suggested that geometric cues were used across changes in environment size, featural cues exerted greater influence when in conflict with geometric cues, and the far featural cue was used to disambiguate the correct from the rotationally equivalent location. In short, both feature and geometric cues were used for reorientation, and environment size influenced the relative use of feature and geometric cues. Collectively, our results provide evidence against associative-based accounts of orientation.     

Use of slope and feature cues in pigeon (Columba livia) goal-searching behavior

Terrain slope provides a directional frame of reference for reorientation and navigation, similar to cardinal directions. Previous studies have shown that, in a goal location task, slope is a very salient cue and that pigeons tend to rely on it even if it is not the most informative cue. Such a strong dependence on one type of information, when there are more effective predictors of reward, is a key premise for a modular view of information processing. Here we tested the provocative hypothesis of a "slope module" for reorientation in slanted environments. Pigeons had to solve a goal location task using slope or another, theoretically salient cue: a beacon feature. Overall, searching behavior was controlled almost equally by the two cues. The fact that, for the first time, slope failed to capture most of the associative strength allows us to reject a strong modularity view and suggests instead that there is competition between cues based on salience. As an interesting additional finding, the reliance on slope and the feature was affected by training location (uphill vs. downhill), suggesting the possibility of a modulatory role of effort on the cue-weighting mechanism of reorientation.