Landmarks, beacons, or panoramic views: What do pigeons attend to for guidance in familiar environments?

Birds and social insects represent excellent systems for understanding visually guided navigation. Both animal groups use surrounding visual cues for homing and foraging. Ants extract sufficient spatial information from panoramic views, which naturally embed all near and far spatial information, for successful homing. Although egocentric panoramic views allow for parsimonious explanations of navigational behaviors, this potential source of spatial information has been mostly neglected during studies of vertebrates. Here we investigate how distinct landmarks, a beacon, and panoramic views influence the reorientation behavior in pigeons (Columba livia). Pigeons were trained to search for a location characterized by a beacon and several distinct landmarks. Transformation tests manipulated aspects of the landmark configuration, allowing for a dissociation among navigational strategies. Quantitative image and path analyses provided support that the panoramic view was used by the pigeons. Although the results from some individuals support the use of beaconing, overall the pigeons relied predominantly on the panoramic view when spatial cues provided conflicting information regarding the goal location. Reorientation based on vector and bearing information derived from distinct landmarks as well as environmental geometry failed to account fully for the results. Thus, the results of our study support that pigeons can use panoramic views for reorientation in familiar environments. Given that the current model for landmark use by pigeons posits the use of different vectors from an object, a global panorama-matching strategy suggests a fundamental change in the theory of how pigeons use surrounding visual cues for localization.

Age-related reduction of hemispheric asymmetry by pigeons: A behavioral and FDG-PET imaging investigation of visual discrimination.

Pigeons are long-lived and slowly aging animals that present a distinct opportunity to further our understanding of age-related brain changes. Generally, for pigeons, the left hemisphere contributes to discrimination of local information, whereas the right contributes to processing of global information. The function of each hemisphere may be examined by covering one eye, as the optic nerves decussate almost completely in birds, directing the majority of visual information to the contralateral hemisphere. Using this eye-capping technique, we investigated pigeons' ability to select grains from among grit while under binocular and monocular viewing conditions, across three different age groups. Prior to the grit-grain discrimination task, pigeons were injected with a radioactive tracer, which was taken up by the brain as the pigeons performed the task. Upon completion of the discrimination task, the pigeons' brains were imaged via [18F] fluorodeoxyglucose positron emission tomography (FDG-PET) scans. This process allowed us to compare hemispheric activity during the discrimination task for each individual within each age group. The Very Old subjects showed significantly worse discrimination performance compared to the Adult and Old subjects, particularly when needing to search primarily with their right hemisphere. Furthermore, the Very Old subjects did not show differences in hemispheric activation when performing the task, whereas the left hemisphere was most active for the Adult and Old groups. To our knowledge, this is the first study to use FDG-PET imaging to evaluate whether the pigeon brain shows evidence of age-related reduction in hemispheric asymmetry during a visual discrimination task.

Does stress run through the leash? An examination of stress transmission between owners and dogs during a walk.

Humans and dogs have co-evolved for over 10,000 years. Recent research suggests that, through the domestication process, dogs have become proficient at responding to human commands, attention and emotional states. However, the extent to which a companion dog responds to human emotions, such as stress, remains to be understood. This study examines whether a companion dog's stress, as measured by cortisol levels and heart rate, increases during a familiar outdoor walk in response to its owner's experience of stress. Sixty-eight owner/dog dyads participated in this study. The dyads were randomly assigned to an Experimental or Control group. Owners in the Experimental group were informed the walk would be digitally recorded for subsequent evaluation of their handling skills, whereas those in the Control group were informed the walk would be digitally recorded for archival purposes (no evaluation). This manipulation was implemented to induce a mild stress response in the owners. Salivary cortisol samples were collected from the owner and their dog before and after the walk. The dyad was also fitted with monitoring devices to record heart rate throughout the walk. Finally, personality information regarding the owner and their dog was collected. We found that cortisol production within the dyad showed a marginal inverse correlation. We also found that owners' Openness to Experience and dogs' Fearfulness influenced the heart rate of the other during the first minute of a walk. These results support that although stress may be detected within a dyad, this does not result in an associated significant change in cortisol or heart rate.

Same/different concept learning by primates and birds.

Same/different abstract-concept learning experiments were conducted with two primate species and three avian species by progressively increasing the size of the training stimulus set of distinctly different pictures from eight to 1,024 pictures. These same/different learning experiments were trained with two pictures presented simultaneously. Transfer tests of same and different learning employed interspersed trials of novel pictures to assess the level of correct performance on the very first time of subjects had seen those pictures. All of the species eventually performed these tests with high accuracy, contradicting the long-accepted notion that nonhuman animals are unable to learn the concept of same/different. Capuchin and rhesus monkeys learned the concept more readily than did pigeons. Clark's nutcrackers and black-billed magpies learned as readily as monkeys, and even showed a slight advantage with the smallest training stimulus sets. Those tests of same/different learning were followed by delay procedures, such that a delay was introduced after the subjects responded to the sample picture and before the test picture. In the sequential same/different task, accuracy was shown to diminish when the stimulus on a previous trial matched the test picture previously shown on a different trial. This effect is known as proactive interference. The pigeons' proactive interference was greater at 10-s delays than 1-s delays, revealing time-based interference. By contrast, time delays had little or no effect on rhesus monkeys' proactive interference, suggesting that rhesus monkeys have better explicit memory of where and when they saw the potential interfering picture, revealing better event-based memory.

Conspecific presence, but not pilferage, influences pinyon jays' (Gymnorhinus cyanocephalus) caching behavior.

Caching species store food when plentiful to ensure availability when resources are scarce. These stores may be at risk of pilferage by others present at the time of caching. Cachers may reduce the risk of loss by using information from the social environment to engage in behaviors to secure the resource-cache protection strategies. Here, we examined whether pinyon jays, a highly social corvid, use information from the social environment to modify their caching behavior. Pinyon jays were provided with pine seeds to cache in two visually distinct trays. The cacher could be observed by a non-pilfering conspecific, a pilfering conspecific, or an inanimate heterospecific located in an adjoining cage compartment, or the cacher could be alone. After caching, the pilfered tray was placed in the adjoining compartment where caches were either pilfered (pilfering conspecific and inanimate heterospecific conditions) or remained intact (non-pilfering conspecific and alone conditions). The safe tray was placed in a visible, but inaccessible, location. Overall, pinyon jays reduced the number of pine seeds cached in the pilfered tray when observed, compared with caching alone. However, their caching behavior did not differ between the pilfering conspecific and the non-pilfering conspecific conditions. These results suggest that either pinyon jays were unable to discriminate between the pilfering and non-pilfering conspecifics, or they generalized their experience of risk from the pilfering conspecific to the non-pilfering conspecific. Thus, we report evidence that pinyon jays use cache protection strategies to secure their resources when observed, but respond similarly when observed by pilfering and non-pilfering conspecifics.

Reorientation by features and geometry: Effects of healthy and degenerative age-related cognitive decline.

The ability to orient is critical for mobile species. Two visual cues, geometry (e.g., distance and direction) and features (e.g., colour and texture) are often used when establishing one's orientation. Previous research has shown the use of these cues, in particular, geometry, may decline with healthy aging. Few studies have examined whether degenerative aging processes show similar time points for the decline of geometry use. The present study examined this issue by training adult and aged mice from two strains, a healthy wild-type and an Alzheimer's model, to search for a hidden platform in a rectangular water maze. The shape of the maze provided geometric information, and distinctive features were displayed on the walls. Following training, manipulations to the features were made to examine whether the mice were able to use the features and geometry, and whether they showed a preference between these two cue types. Results showed that although Alzheimer's transgenic mice were slower to learn the task, overall age rather than strain, was associated with a degradation in use of geometry. However, the presence of seemingly uninformative features (due to their redundancy) facilitated the use of geometry. Additionally, when features and geometry provided conflicting information, only young wild-type mice showed a primary use of features. Our results suggest the failure to use geometry may be a generalized function of aging, and not a diagnostic feature of degeneration for mice. Whether this is also the case for other mammals, such as humans for which the mouse is an important medical model, remains to be examined.

Motoric self-regulation by sled dogs and pet dogs and the acute effect of carbohydrate source in sled dogs.

Inhibitory control is a term used to envelop a collection of processes that allow an organism to refrain from engaging in an inappropriate prepotent or responsive behavior. Studies have examined the propensity of inhibitory control by nonhuman animals, from the cognitively complex processes involved in self-control to potentially less cognitively taxing processes such as motoric self-regulation. Focusing on canines, research has suggested that the domestication process as well as experiences during ontogeny contribute to inhibitory control. Diet may also play an important role in an individual's ability to self-regulate. This study examined this possibility by investigating motoric self-regulation in sled dogs, using three well-established tasks (i.e., A-not-B Bucket, Cylinder, and A-not-B Barrier tasks), performed after consumption of one of three dietary treatments with different glycemic index values. We also compared the performance of sled dogs during these tasks with results previously obtained from pet dogs. Overall, the results show many similarities in the performance of sled dogs and pet dogs on the motoric self-regulation tasks, with the notable exception that sled dogs may have a stronger spatial perseveration during the A-not-B Bucket task. Previous research findings reporting a lack of correlation among these tasks are also supported. Finally, during the early postprandial phase (period after consumption), dietary treatments with different glycemic index values did not influence self-regulatory performance for sled dogs.

Comparing cognition by integrating concept learning, proactive interference, and list memory.

This article describes an approach for training a variety of species to learn the abstract concept of same/different, which in turn forms the basis for testing proactive interference and list memory. The stimulus set for concept-learning training was progressively doubled from 8, 16, 32, 64, 128 . . . to 1,024 different pictures with novel-stimulus transfer following learning. All species fully learned the same/different abstract concept: capuchin and rhesus monkeys learned more readily than pigeons; nutcrackers and magpies were at least equivalent to monkeys and transferred somewhat better following initial training sets. A similar task using the 1,024-picture set plus delays was used to test proactive interference on occasional trials. Pigeons revealed greater interference with 10-s than with 1-s delays, whereas delay time had no effect on rhesus monkeys, suggesting that the monkeys' interference was event based. This same single-item same/different task was expanded to a 4-item list memory task to test animal list memory. Humans were tested similarly with lists of kaleidoscope pictures. Delays between the list and test were manipulated, resulting in strong initial recency effects (i.e., strong 4th-item memory) at short delays and changing to a strong primacy effect (i.e., strong 1st-item memory) at long delays (pigeons 0-s to 10-s delays; monkeys 0-s to 30-s delays; humans 0-s to 100-s delays). Results and findings are discussed in terms of these species' cognition and memory comparisons, evolutionary implications, and future directions for testing other species in these synergistically related tasks.

Individual performance across motoric self-regulation tasks are not correlated for pet dogs.

Inhibitory control, the ability to restrain a prepotent but ineffective response in a given context, is thought to be indicative of a species' cognitive abilities. This ability ranges from "basic" motoric self-regulation to more complex abilities such as self-control. During the current study, we investigated the motoric self-regulatory abilities of 30 pet dogs using four well-established cognitive tasks - the A-not-B Bucket task, the Cylinder task, the Detour task, and the A-not-B Barrier task - administered in a consistent context. One main goal of the study was to determine whether the individual-level performance would correlate across tasks, supporting that these tasks measure similar components of motoric self-regulation. Dogs in our study were quite successful during tasks requiring them to detour around transparent barriers (i.e., the Cylinder and Detour tasks), but were less successful with tasks requiring the production of a new response (i.e., A-not-B Bucket and A-not-B Barrier tasks). However, individual dog performance did not correlate across tasks, suggesting these well-established tasks likely measure different inhibitory control abilities, or are strongly influenced by differential task demands. Our results also suggest other aspects such as perseveration or properties of the apparatus may need to be carefully examined in order to better understand canine motoric self-regulation or inhibitory control more generally.

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.

Corvids Outperform Pigeons and Primates in Learning a Basic Concept.

Corvids (birds of the family Corvidae) display intelligent behavior previously ascribed only to primates, but such feats are not directly comparable across species. To make direct species comparisons, we used a same/different task in the laboratory to assess abstract-concept learning in black-billed magpies ( Pica hudsonia). Concept learning was tested with novel pictures after training. Concept learning improved with training-set size, and test accuracy eventually matched training accuracy-full concept learning-with a 128-picture set; this magpie performance was equivalent to that of Clark's nutcrackers (a species of corvid) and monkeys (rhesus, capuchin) and better than that of pigeons. Even with an initial 8-item picture set, both corvid species showed partial concept learning, outperforming both monkeys and pigeons. Similar corvid performance refutes the hypothesis that nutcrackers' prolific cache-location memory accounts for their superior concept learning, because magpies rely less on caching. That corvids with "primitive" neural architectures evolved to equal primates in full concept learning and even to outperform them on the initial 8-item picture test is a testament to the shared (convergent) survival importance of abstract-concept learning.

How Clark's nutcrackers (Nucifraga columbiana) weigh geometric cues depends on their previous experience.

Following passive disorientation, Clark's nutcrackers (Nucifraga columbiana) learned to search for a hidden food reward located in one corner of a rectangular-shaped enclosure that contained either identical or distinct features in each corner. Identical features allowed for explicit learning of geometric cues, whereas distinct features allowed for both explicit learning of featural cues and incidental learning of geometric cues. Birds that only learned about geometry incidentally (group Distinct) weighed features greater than geometry when the two cues were placed in conflict. However, birds that received explicit training with geometry, in addition to feature training (groups Distinct-Identical and Identical-Distinct), weighed geometry heavier relative to features. Cue preference by the birds also depended on the order in which learning was experienced; if explicit training with geometry followed that of features (group Distinct-Identical), then both geometry and features were weighed equally, but if explicit training with geometry training preceded that of features (group Identical-Distinct), the birds weighed geometry greater than features. Results suggest both a heightened sensitivity to geometric cues by Clark's nutcrackers relative to other species of birds and an increased sensitivity to any spatial cue (either features or geometry) that has proven both stable and reliable.

Superior abstract-concept learning by Clark's nutcrackers (Nucifraga columbiana).

The ability to learn abstract relational concepts is fundamental to higher level cognition. In contrast to item-specific concepts (e.g. pictures containing trees versus pictures containing cars), abstract relational concepts are not bound to particular stimulus features, but instead involve the relationship between stimuli and therefore may be extrapolated to novel stimuli. Previous research investigating the same/different abstract concept has suggested that primates might be specially adapted to extract relations among items and would require fewer exemplars of a rule to learn an abstract concept than non-primate species. We assessed abstract-concept learning in an avian species, Clark's nutcracker (Nucifraga columbiana), using a small number of exemplars (eight pairs of the same rule, and 56 pairs of the different rule) identical to that previously used to compare rhesus monkeys, capuchin monkeys and pigeons. Nutcrackers as a group (N = 9) showed more novel stimulus transfer than any previous species tested with this small number of exemplars. Two nutcrackers showed full concept learning and four more showed transfer considerably above chance performance, indicating partial concept learning. These results show that the Clark's nutcracker, a corvid species well known for its amazing feats of spatial memory, learns the same/different abstract concept better than any non-human species (including non-human primates) yet tested on this same task.

Relational complexity influences analogical reasoning ability.

Human language without analogy is like a zebra without stripes. The ability to understand analogies, or to engage in relational reasoning, has been argued to be an important distinction between the cognitive abilities of human and non-human animals. Current studies have failed to robustly show that animals can perform more complex, relational discriminations, in part because such tests rely on linguistic or symbolic experiences, and therefore are not suitable for evaluating analogical reasoning in animals. We report on a methodological approach allowing for direct comparisons of analogical reasoning ability across species. We show that human participants spontaneously make analogical discriminations with minimal verbal instructions, and that the ability to reason analogically is affected by analogical complexity. Furthermore, performance on our task correlated with participants' fluid intelligence scores. These results show the nuance of analogical reasoning abilities by humans, and provide a means of robustly comparing this capacity across species.

Discrimination of geometric angles by adult humans.

Men and women learned to discriminate between two different size angles presented to them as objects within a real-world task. During Experiment 1, participants in group 50 were trained to choose a 50° angle and participants in group 75 were trained to choose a 75° angle. During testing, both groups were provided with a choice between their training angle and one of a set of test angles that was either smaller or larger than the training angle. Results showed a generalized pattern of responding, with group 50 showing increased responding to test angles smaller than 50° and group 75 showing increased responding to test angles larger than 75°. Further analysis of the response patterns revealed that participants in group 50 showed evidence of absolute learning, whereas participants in group 75 showed evidence of relational learning. During Experiment 2, a third group of participants (group 25) trained to choose a smaller angle (25°) was included in addition to group 50 and group 75. Participants were trained with three angles present and tested with just two, one being their training angle and the other being one of a set of novel test angles. Similar to the participants from Experiment 1, group 75 showed evidence of relational learning. Group 50, for which no relational rule could be applied during training, showed an absolute learning pattern with no response shift to test angles smaller or larger than their training angle. Group 25 showed evidence of absolute responding that was more pronounced than that found for the smallest training angle during Experiment 1. These findings suggest differential learning of geometric angles based on amplitude with smaller angles perceived as more distinct and thus more resistant to broader generalization than larger angles. Implications of these results are that certain geometric properties may be subject to different learning processes based on the specific magnitude of that property.

Spatial reorientation with a geometric array of auditory cues.

A visuocentric bias has dominated the literature on spatial navigation and reorientation. Studies on visually accessed environments indicate that, during reorientation, human and non-human animals encode the geometric shape of the environment, even if this information is unnecessary and insufficient for the task. In an attempt to extend our limited knowledge on the similarities and differences between visual and non-visual navigation, here we examined whether the same phenomenon would be observed during auditory-guided reorientation. Provided with a rectangular array of four distinct auditory landmarks, blindfolded, sighted participants had to learn the location of a target object situated on a panel of an octagonal arena. Subsequent test trials were administered to understand how the task was acquired. Crucially, in a condition in which the auditory cues were indistinguishable (same sound sample), participants could still identify the correct target location, suggesting that the rectangular array of auditory landmarks was encoded as a geometric configuration. This is the first evidence of incidental encoding of geometric information with auditory cues and, consistent with the theory of functional equivalence, it supports the generalisation of mechanisms of spatial learning across encoding modalities.