Jayden O. van Horik

Differential participation in cognitive tests is driven by personality, sex, body condition and experience

Failure to participate in a cognitive test may result in sampling biases when measuring inter-individual variation in cognitive performances in both captive and wild populations. This would be problematic if particular classes of individuals consistently fail to participate, skewing data and making generalisations or comparisons difficult. We presented 144 pheasant chicks, raised under standardised conditions, with a battery of cognitive tests to investigate whether sex, body condition or personality traits, measured by differences in latencies to explore a novel object, novel environment or unknown conspecific, predicted individual variation in voluntary participation across 37 test sessions. In general, participation increased across testing sessions, yet patterns of participation differed with sex and body condition. Males with a high body condition were more likely to participate in early test sessions compared to males with a low body condition or females. While participation among males in high body condition was consistent across sessions, males with a low body condition and females, regardless of body condition, were more likely to participate in later, rather than earlier sessions. Individuals also showed repeatable behaviours across time and different contexts, revealing not only that the exploration of novelty, but also that the order that subjects entered the testing arena and their latencies to acquire a freely available meal-worm reward may be considered valid proxies for different personality traits. During each test session, those individuals that were among the first to voluntarily enter the testing arena were more likely to participate in subsequent trials. Moreover, when isolated in the testing arena, individuals that rapidly acquired a freely available meal-worm, positioned on the testing apparatus, were also more likely to participate in a cognitive test. Our findings therefore reveal that sex, body condition and personality traits, along with habituation to the testing paradigms, all play important roles in determining whether or not particular individuals participate in cognitive tests. Sampling biases may therefore misrepresent our understanding of variation in cognitive performance in wild and captive populations, making individual differences in cognition difficult to interpret.

Evolution of cognition

Renewed interest in the field of comparative cognition over the past 30 years has led to a renaissance in our thinking of how cognition evolved. Here, we review historical and comparative approaches to the study of psychological evolution, focusing on cognitive differences based on evolutionary divergence, but also cognitive similarities based on evolutionary convergence. Both approaches have contributed to major theories of cognitive evolution in humans and non-human animals. As a result, not only have we furthered our understanding of the evolution of the human mind and its unique attributes, but we have also identified complex cognitive capacities in a few large-brained species, evolved from solving social and ecological challenges requiring a flexible mind. WIREs Cogni Sci 2011 2 621-633 DOI: 10.1002/wcs.144 For further resources related to this article, please visit the WIREs website.

Do detour tasks provide accurate assays of inhibitory control

Transparent Cylinder and Barrier tasks are used to purportedly assess inhibitory control in a variety of animals. However, we suspect that performances on these detour tasks are influenced by non-cognitive traits, which may result in inaccurate assays of inhibitory control. We therefore reared pheasants under standardized conditions and presented each bird with two sets of similar tasks commonly used to measure inhibitory control. We recorded the number of times subjects incorrectly attempted to access a reward through transparent barriers, and their latencies to solve each task. Such measures are commonly used to infer the differential expression of inhibitory control. We found little evidence that their performances were consistent across the two different Putative Inhibitory Control Tasks (PICTs). Improvements in performance across trials showed that pheasants learned the affordances of each specific task. Critically, prior experience of transparent tasks, either Barrier or Cylinder, also improved subsequent inhibitory control performance on a novel task, suggesting that they also learned the general properties of transparent obstacles. Individual measures of persistence, assayed in a third task, were positively related to their frequency of incorrect attempts to solve the transparent inhibitory control tasks. Neophobia, Sex and Body Condition had no influence on individual performance. Contrary to previous studies of primates, pheasants with poor performance on PICTs had a wider dietary breadth assayed using a free-choice task. Our results demonstrate that in systems or taxa where prior experience and differences in development cannot be accounted for, individual differences in performance on commonly used detour-dependent PICTS may reveal more about an individuals prior experience of transparent objects, or their motivation to acquire food, than providing a reliable measure of their inhibitory control.

Vocal ethology of the North Island kaka (Nestor meridionalis septentrionalis)

Abstract We conducted acoustic and behavioural observations on wild New Zealand North Island kaka (Nestor meridionalis septentrionalis) to assess the behavioural context of their most common calls. We distinguished several call types by ear in the field and then quantitatively evaluated our call type classifications using spectrographic analyses. Next, we established the behavioural context of each call type during 500 h of field observations. We observed five distinctive call types that were clearly segregated in subsequent spectrographic analyses. Behavioural observations showed that each call type was generally associated with particular behaviours used by birds separated by different distances. Some call types were used by distantly‐separated solitary birds that were foraging or preening, while others were used mostly during copulation. Overall results indicate that kaka have a range of distinctive call‐types for communication under different spatial and social circumstances.

The quick are the dead: pheasants that are slow to reverse a learned association survive for longer in the wild

Cognitive abilities probably evolve through natural selection if they provide individuals with fitness benefits. A growing number of studies demonstrate a positive relationship between performance in psychometric tasks and (proxy) measures of fitness. We assayed the performance of 154 common pheasant (Phasianus colchicus) chicks on tests of acquisition and reversal learning, using a different set of chicks and different set of cue types (spatial location and colour) in each of two years and then followed their fates after release into the wild. Across all birds, individuals that were slow to reverse previously learned associations were more likely to survive to four months old. For heavy birds, individuals that rapidly acquired an association had improved survival to four months, whereas for light birds, slow acquirers were more likely to be alive. Slow reversers also exhibited less exploratory behaviour in assays when five weeks old. Fast acquirers visited more artificial feeders after release. In contrast to most other studies, we showed that apparently ‘poor’ cognitive performance (slow reversal speed suggesting low behavioural flexibility) correlates with fitness benefits in at least some circumstances. This correlation suggests a novel mechanism by which continued exaggeration of cognitive abilities may be constrained. This article is part of the theme issue ‘Causes and consequences of individual differences in cognitive abilities’.

Transfer of physical understanding in a non-tool-using parrot.

Physical cognition has generally been assessed in tool-using species that possess a relatively large brain size, such as corvids and apes. Parrots, like corvids and apes, also have large relative brain sizes, yet although parrots rarely use tools in the wild, growing evidence suggests comparable performances on physical cognition tasks. It is, however, unclear whether success on such tasks is facilitated by previous experience and training procedures. We therefore investigated physical comprehension of object relationships in two non-tool-using species of captive neotropical parrots on a new means-end paradigm, the Trap-Gaps task, using unfamiliar materials and modified training procedures that precluded procedural cues. Red-shouldered macaws (Diopsittaca nobilis) and black-headed caiques (Pionites melanocephala) were presented with an initial task that required them to discriminate between pulling food trays through gaps while attending to the respective width of the gaps and size of the trays. Subjects were then presented with a novel, but functionally equivalent, transfer task. Six of eight birds solved the initial task through trial-and-error learning. Four of these six birds solved the transfer task, with one caique demonstrating spontaneous comprehension. These findings suggest that non-tool-using parrots may possess capacities for sophisticated physical cognition by generalising previously learned rules across novel problems.

Erratum to: Differences in social preference between the sexes during ontogeny drive segregation in a precocial species

The work was jointly funded by the University of Exeter, the Game and Wildlife Conservation Trust, a Royal Society Small Project Grant awarded to JRM and an ERC Consolidator Grant (616474) awarded to JRM.

Group social rank is associated with performance on a spatial learning task

Dominant individuals differ from subordinates in their performances on cognitive tasks across a suite of taxa. Previous studies often only consider dyadic relationships, rather than the more ecologically relevant social hierarchies or networks, hence failing to account for how dyadic relationships may be adjusted within larger social groups. We used a novel statistical method: randomized Elo-ratings, to infer the social hierarchy of 18 male pheasants, Phasianus colchicus, while in a captive, mixed-sex group with a linear hierarchy. We assayed individual learning performance of these males on a binary spatial discrimination task to investigate whether inter-individual variation in performance is associated with group social rank. Task performance improved with increasing trial number and was positively related to social rank, with higher ranking males showing greater levels of success. Motivation to participate in the task was not related to social rank or task performance, thus indicating that these rank-related differences are not a consequence of differences in motivation to complete the task. Our results provide important information about how variation in cognitive performance relates to an individuals social rank within a group. Whether the social environment causes differences in learning performance or instead, inherent differences in learning ability predetermine rank remains to be tested.

Individuals in larger groups are more successful on spatial discrimination tasks

To understand how natural selection may act on cognitive processes, it is necessary to reliably determine interindividual variation in cognitive abilities. However, an individuals performance in a cognitive test may be influenced by the social environment. The social environment explains variation between species in cognitive performances, with species that live in larger groups purportedly demonstrating more advanced cognitive abilities. It also explains variation in cognitive performances within species, with larger groups more likely to solve novel problems than smaller groups. Surprisingly, an effect of group size on individual variation in cognitive performance has rarely been investigated and much of our knowledge stems from impaired performance of individuals reared in isolation. Using a within-subjects design we assayed individual learning performance of adult female pheasants, Phasianus colchicus, while housed in groups of three and five. Individuals experienced the group sizes in a different order, but were presented with two spatial discrimination tasks, each with a distinct cue set, in a fixed order. We found that across both tasks individuals housed in the large groups had higher levels of success than individuals housed in the small groups. Individuals had higher levels of success on their second than their first task, irrespective of group size. We suggest that the expression of individual learning performance is responsive to the current social environment but the mechanisms underpinning this relationship require further investigation. Our study demonstrates that it is important to account for an individuals social environment when attempting to characterize cognitive capacities. It also demonstrates the flexibility of an individuals cognitive performance depending on the social context.

Low survival of strongly footed pheasants may explain constraints on lateralization

Brain lateralization is considered adaptive because it leads to behavioral biases and specializations that bring fitness benefits. Across species, strongly lateralized individuals perform better in specific behaviors likely to improve survival. What constrains continued exaggerated lateralization? We measured survival of pheasants, finding that individuals with stronger bias in their footedness had shorter life expectancies compared to individuals with weak biases. Consequently, weak, or no footedness provided the highest fitness benefits. If, as suggested, footedness is indicative of more general brain lateralization, this could explain why continued brain lateralization is constrained even though it may improve performance in specific behaviors.