A problem with problem solving: motivational traits, but not cognition, predict success on novel operant foraging tasks

Published onlineRates of innovative foraging behaviours and success on problem-solving tasks are often used to assay differences in cognition, both within and across species. Yet the cognitive features of some problemsolving tasks can be unclear. As such, explanations that attribute cognitive mechanisms to individual variation in problem-solving performance have revealed conflicting results. We investigated individual consistency in problem-solving performances in captive-reared pheasant chicks, Phasianus colchicus, and addressed whether success depends on cognitive processes, such as trial-and-error associative learning, or whether performances may be driven solely via noncognitive motivational mechanisms, revealed through subjects' willingness to approach, engage with and persist in their interactions with an apparatus, or via physiological traits such as body condition. While subjects' participation and success were consistent within the same problems and across similar tasks, their performances were inconsistent across different types of task. Moreover, subjects' latencies to approach each test apparatus and their attempts to access the reward were not repeatable across trials. Successful individuals did not improve their performances with experience, nor were they consistent in their techniques in repeated presentations of a task. However, individuals that were highly motivated to enter the experimental chamber were more likely to participate. Successful individuals were also faster to approach each test apparatus and more persistent in their attempts to solve the tasks than unsuccessful individuals. Our findings therefore suggest that individual differences in problem-solving success can arise from inherent motivational differences alone and hence be achieved without inferring more complex cognitive processesWe thank Rothamsted Research North Wyke for accommodating the rearing and release of the pheasants. We thank Aidan Hulatt, Sebastian Bekker and James Foley for their help with data collection and animal husbandry. Lauren Brent and Rob Heathcote provided advice on analysis. J.R.M. and J.O.v.H. were funded by an ERC consolidator grant (616474)

Serial reversal learning and cognitive flexibility in two species of Neotropical parrots (Diopsittaca nobilis and Pionites melanocephala)

ArticleThis is the author accepted manuscript.Serial reversal learning of colour discriminations was assessed as an index of behavioural flexibility in two captive species of Neotropical parrots. Both species showed similar performances across serial reversals and no between species differences were observed. In a second task subjects’ performances were assessed after they experienced either a low or high pre-reversal learning criterion. If reversal performances improve through processes of associative learning, a high pre-reversal criterion is expected to strengthen previously learned associations and hence impede post-reversal performances. Conversely, highly reinforced associations may facilitate the use of conditional rules that can be generalised across reversals and improve post-reversal performances. We found that high criterion subjects made fewer post-reversal errors and required fewer trials to reach criterion, than low criterion subjects. Red-shouldered macaws and black-headed caiques may therefore demonstrate capacities for solving serial reversal problems by applying conditional rules, rather than learning solely by associative processes. Such performances coincide with findings in great apes, but contrast with findings in monkeys and prosimians, which generally show impaired reversal performances when trained to a highly rigorous pre-reversal criterion. Overall, these findings suggest an evolutionary convergence of behavioural flexibility between parrots and non-human great apes

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

This is the final published version. Available from the Royal Society via the DOI in this record.The dataset associated with this article is located in ORE at: http://hdl.handle.net/10871/30840Dominant 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 individual's 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.ERC Consolidator Award (616474) to J.R.M

Individuals in larger groups are more successful on spatial discrimination tasks

This is the final version of the article. Available from the publisher via the DOI in this record.To understand how natural selection may act on cognitive processes, it is necessary to reliably determine interindividual variation in cognitive abilities. However, an individual's 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 individual's social environment when attempting to characterize cognitive capacities. It also demonstrates the flexibility of an individual's cognitive performance depending on the social context.The work was funded by an ERC Consolidator Award to J.R.M (616474)

The relationship between social rank and spatial learning in pheasants, Phasianus colchicus: Cause or consequence? (article)

This is the final version. Available from PeerJ via the DOI in this record.The dataset associated with this article is in ORE: https://doi.org/10.24378/exe.143Individual differences in performances on cognitive tasks have been found to differ according to social rank across multiple species. However, it is not clear whether an individual's cognitive performance is flexible and the result of their current social rank, modulated by social interactions (social state dependent hypothesis), or if it is determined prior to the formation of the social hierarchy and indeed influences an individual's rank (prior attributes hypothesis). We separated these two hypotheses by measuring learning performance of male pheasants, Phasianus colchicus, on a spatial discrimination task as chicks and again as adults. We inferred adult male social rank from observing agonistic interactions while housed in captive multi-male multi-female groups. Learning performance of adult males was assayed after social rank had been standardised; by housing single males with two or four females. We predicted that if cognitive abilities determine social rank formation we would observe: consistency between chick and adult performances on the cognitive task and chick performance would predict adult social rank. We found that learning performances were consistent from chicks to adults for task accuracy, but not for speed of learning and chick learning performances were not related to adult social rank. Therefore, we could not support the prior attributes hypothesis of cognitive abilities aiding social rank formation. Instead, we found that individual differences in learning performances of adults were predicted by the number of females a male was housed with; males housed with four females had higher levels of learning performance than males housed with two females; and their most recent recording of captive social rank, even though learning performance was assayed while males were in a standardized, non-competitive environment. This does not support the hypothesis that direct social pressures are causing the inter-individual variation in learning performances that we observe. Instead, our results suggest that there may be carry-over effects of aggressive social interactions on learning performance. Consequently, whether early life spatial learning performances influence social rank is unclear but these performances are modulated by the current social environment and a male's most recent social rank.European Research Counci

Differences in social preference between the sexes during ontogeny drive segregation in a precocial species (article)

This is the final version of the article. Available from the publisher via the DOI in this record.The datasets analysed during the current study are available at the Open Research Exeter repository. http://hdl.handle.net/ 10871/27114.Hypotheses for why animals sexually segregate typically rely on adult traits, such as differences in sexual roles causing differential habitat preferences, or size dimorphism inducing differences in diet or behaviour. However, segregation can occur in juveniles before such roles or size dimorphism is well established. In young humans, leading hypotheses suggest that (1) sexes differ in their activity and the synchronisation of behaviour causes segregation and (2) sexes separate in order to learn and maximise future reproductive roles. We reared pheasants, Phasianus colchicus, from hatching in the absence of adults in a controlled environment. Females aggregated with their own sex from hatching, whereas males initially exhibited random association, but segregation became pronounced with age. The increase in segregation corresponded to an increase in sexual size dimorphism. By standardising habitat availability and diet and by removing predation risk, we could disregard the Predation Risk and the Forage Selection Hypotheses operating at this age. Activity budgets did not differ between the sexes, providing no support for the Behavioural Synchrony or the Activity Budget Hypotheses. Both sexes preferentially approached groups of unfamiliar, same-sex birds in binary choice tests, providing support for the Social Preference Hypothesis. Females may segregate to avoid male aggression. Sexual segregation may become established early in development, especially in precocial species, such as pheasants. A clear understanding of ontogenetic factors is essential to further our understanding of adult assortment patterns. Assortment by sex may not be inherent, but rather emerge as a consequence of social interactions early in life.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

Low survival of strongly footed pheasants may explain constraints on lateralization

This is the final version. Available from Springer Nature via the DOI in this record.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.This work was supported by a European Research Council Consolidator Award (616474) awarded to JRM

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

This is the final version. Available on open access from the Royal Society via the DOI in this recordData accessibility: Data are available in DRYAD: Data from: The quick are the dead: pheasants that are slow to reverse a learned association survive for longer in the wild. http://dx.doi.org/10.5061/dryad.9bn4443Cognitive 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’.European Research Council (ERC

Response learning confounds assays of inhibitory control on detour tasks

This is the final version. Available from the publisher via the DOI in this record.All data are available on Dryad.The ability to inhibit prepotent actions towards rewards that are made inaccessible by transparent barriers has been considered to reflect capacities for inhibitory control (IC). Typically, subjects initially reach directly, and incorrectly, for the reward. With experience, subjects may inhibit this action and instead detour around barriers to access the reward. However, assays of IC are often measured across multiple trials, with the location of the reward remaining constant. Consequently, other cognitive processes, such as response learning (acquisition of a motor routine), may confound accurate assays of IC. We measured baseline IC capacities in pheasant chicks, Phasianus colchicus, using a transparent cylinder task. Birds were then divided into two training treatments, where they learned to access a reward placed behind a transparent barrier, but experienced differential reinforcement of a particular motor response. In the stationary-barrier treatment, the location of the barrier remained constant across trials. We, therefore, reinforced a fixed motor response, such as always go left, which birds could learn to aid their performance. Conversely, we alternated the location of the barrier across trials for birds in the moving-barrier treatment and hence provided less reinforcement of their response learning. All birds then experienced a second presentation of the transparent cylinder task to assess whether differences in the training treatments influenced their subsequent capacities for IC. Birds in the stationary-barrier treatment showed a greater improvement in their subsequent IC performance after training compared to birds in the moving-barrier treatment. We, therefore, suggest that response learning aids IC performance on detour tasks. Consequently, non-target cognitive processes associated with different neural substrates appear to underlie performances on detour tasks, which may confound accurate assays of IC. Our findings question the construct validity of a commonly used paradigm that is widely considered to assess capacities for IC in humans and other animals.European Commissio

Unpredictable environments enhance inhibitory control in pheasants

This is the final version. Available from the publisher via the DOI in this record.All data are available in DryadThe ability to control impulsive actions is an important executive function that is central to the self-regulation of behaviours and, in humans, can have important implications for mental and physical health. One key factor that promotes individual differences in inhibitory control (IC) is the predictability of environmental information experienced during development (i.e. reliability of resources and social trust). However, environmental predictability can also influence motivational and other cognitive abilities, which may therefore confound interpretations of the mechanisms underlying IC. We investigated the role of environmental predictability, food motivation and cognition on IC. We reared pheasant chicks, Phasianus colchicus, under standardised conditions, in which birds experienced environments that differed in their spatial predictability. We systematically manipulated spatial predictability during their first 8 weeks of life, by either moving partitions daily to random locations (unpredictable environment) or leaving them in fixed locations (predictable environment). We assessed motivation by presenting pheasants with two different foraging tasks that measured their dietary breadth and persistence to acquire inaccessible food rewards, as well as recording their latencies to acquire a freely available baseline worm positioned adjacent to each test apparatus, their body condition (mass/tarsus3) and sex. We assessed cognitive performance by presenting each bird with an 80-trial binary colour discrimination task. IC was assessed using a transparent detour apparatus, which required subjects to inhibit prepotent attempts to directly acquire a visible reward through the barrier and instead detour around a barrier. We found greater capacities for IC in pheasants that were reared in spatially unpredictable environments compared to those reared in predictable environments. While IC was unrelated to individual differences in cognitive performance on the colour discrimination task or motivational measures, we found that environmental predictability had differential effects on sex. Males reared in an unpredictable environment, and all females regardless of their rearing environment, were less persistent than males reared in a predictable environment. Our findings, therefore, suggest that an individual’s developmental experience can influence their performance on IC tasks.European Commissio