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.

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.

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.

Socio-ecological correlates of neophobia in corvids.

Behavioral responses to novelty, including fear and subsequent avoidance of novel stimuli, i.e., neophobia, determine how animals interact with their environment. Neophobia aids in navigating risk and impacts on adaptability and survival. There is variation within and between individuals and species; however, lack of large-scale, comparative studies critically limits investigation of the socio-ecological drivers of neophobia. In this study, we tested responses to novel objects and food (alongside familiar food) versus a baseline (familiar food alone) in 10 corvid species (241 subjects) across 10 labs worldwide. There were species differences in the latency to touch familiar food in the novel object and novel food conditions relative to the baseline. Four of seven socio-ecological factors influenced object neophobia: (1) use of urban habitat (versus not), (2) territorial pair versus family group sociality, (3) large versus small maximum flock size, and (4) moderate versus specialized caching (whereas range, hunting live animals, and genus did not), while only maximum flock size influenced food neophobia. We found that, overall, individuals were temporally and contextually repeatable (i.e., consistent) in their novelty responses in all conditions, indicating neophobia is a stable behavioral trait. With this study, we have established a network of corvid researchers, demonstrating potential for further collaboration to explore the evolution of cognition in corvids and other bird species. These novel findings enable us, for the first time in corvids, to identify the socio-ecological correlates of neophobia and grant insight into specific elements that drive higher neophobic responses in this avian family group. VIDEO ABSTRACT.

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.

Age affects pigeons' (Columba livia) memory capacity but not representation of serial order during a locomotor sequential-learning task.

Aging affects individuals of every species, with sometimes detrimental effects on memory and cognition. The simultaneous-chaining task, a sequential-learning task, requires subjects to select items in a predetermined sequence, putting demands on memory and cognitive processing capacity. It is thus a useful tool to investigate age-related differences in these domains. Pigeons of three age groups (young, adult and aged) completed a locomotor adaptation of the task, learning a list of four items. Training began by presenting only the first item; additional items were added, one at a time, once previous items were reliably selected in their correct order. Although memory capacity declined noticeably with age, not all aged pigeons showed impairments compared to younger pigeons, suggesting that inter-individual variability emerged with age. During a subsequent free-recall memory test in the absence of reinforcement, when all trained items were presented alongside novel distractor items, most pigeons did not reproduce the trained sequence. During a further forced-choice test, when pigeons were given a choice between only two of the trained items, all three age groups showed evidence of an understanding of the ordinal relationship between items by choosing the earlier item, indicating that complex cognitive processing, unlike memory capacity, remained unaffected by age.

Abstract-concept learning in two species of new world corvids, pinyon jays (Gymnorhinus Cyanocephalus) and California scrub jays (Aphelocoma Californica).

concepts require individuals to identify relationships between novel stimuli. Previous studies have reported that the ability to learn abstract concepts is found in a wide range of species. In regard to a same/different concept, Clark's nutcrackers (Nucifraga columbiana) and black-billed magpies (Pica hudsonia), two corvid species, were shown to outperform other avian and primate species (Wright et al., 2017). Two additional corvid species, pinyon jays (Gymnorhinus cyanocephalus) and California scrub jays (Aphelocoma californica) chosen as they belong to a different clade than nutcrackers and magpies, were examined using the same set-size expansion procedure of the same/different task (the task used with nutcrackers and magpies) to evaluate whether this trait is common across the Corvidae lineage. During this task, concept learning is assessed with novel images after training. Results from the current study showed that when presented with novel stimuli after training with an 8-image set, discrimination accuracy did not differ significantly from chance for pinyon jays and California scrub jays, unlike the magpies and nutcrackers from previous studies that showed partial transfer at that stage. However, concept learning improved with each set-size expansion, and the jays reached full concept learning with a 128-image set. This performance is similar to the other corvids and monkeys tested, all of which outperform pigeons. Results from the current study show a qualitative similarity in full abstract-concept learning in all species tested with a quantitative difference in the set-size functions, highlighting the shared survival importance of mechanisms supporting abstract-concept learning for corvids and primates. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

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.

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.

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.

Individual exploratory responses are not repeatable across time or context for four species of food-storing corvid.

Exploration is among one of the most studied of animal personality traits (i.e., individual-level behavioural responses repeatable across time and contexts). However, not all species show clear evidence of this personality trait, and this is particularly so for members of the Corvidae family. We assessed the exploratory behaviour of four food-caching corvid species: pinyon jays (Gymnorhinus cyanocephalus), Clark's nutcrackers (Nucifraga columbiana), California scrub jays (Aphelocoma californica), and black-billed magpies (Pica hudsonia). Contextual repeatability was assessed through examining behavioural measures during the Novel Environment task and the Novel Object task, whereas temporal repeatability was assessed by examining changes in these measures over repeated trials. Our results suggest that, for corvids, an individual's exploratory behaviour was not repeatable across contexts or over time. Hence, we found no evidence that exploration constitutes a personality trait for these species of corvid. We did find differences in exploratory behaviour, at a species level, that may be explained by relative reliance on cached food.

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.

Older Adults Show Less Flexible Spatial Cue Use When Navigating in a Virtual Reality Environment Compared With Younger Adults.

Daily life requires accurate navigation, and thus better understanding of aging on navigational abilities is critical. Importantly, the use of spatial properties by older and younger adults remains unclear. During this study, younger and older human adults were presented with a virtual environment in which they had to navigate a series of hallways. The hallways provided 2 general types of spatial information: geometric, which included distance and directional turns along a learned route, and featural, which included landmarks situated along the route. To investigate how participants used these different cue types, geometric and/or landmark information was manipulated during testing trials. Data from 40 younger (20 women) and 40 older (20 women) adults were analyzed. Our findings suggest that (1) both younger and older adults relied mostly on landmarks to find their way, and (2) younger adults were better able to adapt to spatial changes to the environment compared with older adults.

Use of medial axis for reorientation by the Clark's nutcracker (Nucifraga columbiana).

Many animals are challenged with the task of reorientation. Considerable research over the years has shown a diversity of species extract geometric information (e.g., distance and direction) from continuous surfaces or boundaries to reorient. How this information is extracted from the environment is less understood. Three encoding strategies that have received the most study are the use of principal axes, medial axis or local geometric cues. We used a modeling approach to investigate which of these three general strategies best fit the spatial search data of a highly-spatial corvid, the Clark's nutcracker (Nucifraga columbiana). Individual nutcrackers were trained in a rectangular-shaped arena, and once accurately locating a hidden goal, received non-reinforced tests in an L-shaped arena. The specific shape of this arena allowed us to dissociate among the three general encoding strategies. Furthermore, we reanalyzed existing data from chicks, pigeons and humans using our modeling approach. Overall, we found the most support for the use of the medial axis, although we additionally found that pigeons and humans may have engaged in random guessing. As with our previous studies, we find no support for the use of principal axes.

Effects of sun compass error on spatial search by Clark's nutcrackers.

Animals employ compasses during navigation, but little attention has been paid to how accuracy is maintained in the face of compass error, which is inevitable in biological systems. The use of multiple landmarks may minimize the effect of compass error. We allowed Clark's nutcrackers to cache seeds in an outdoor aviary with either one or four landmarks present, and subsequently subjected them to small clock-shifts mimicking the effects of compass error. As predicted, the results showed a significant decrease in search accuracy following the clock-shift when one landmark was present but not when four landmarks were present. These results support that nutcrackers encode information from the sun as well as terrestrial landmarks, and these spatial cues are used in a flexible manner. Overall, our results are important as they support the hypothesis that multiple landmarks may be used during situations where the sun compass has even a small amount of error.

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.