How Inhibitory Control Relates to Positive and Negative Affective States in Red Junglefowl.

Individual differences in inhibitory control, an aspect of cognition, are found in many species. How this variation links to affective states is not much explored, and could be relevant for welfare. As less fearful, more optimistic, individuals may act more impulsively, inhibitory control could link to less negative, more positive, affective states. Alternatively, poorer inhibitory control could associate with more negative, less positive, affective states, as poorer inhibitory control can result in individuals being less able to adapt to changing environments and more likely to show stereotypies. We here explored in three cohorts (N = 209) of captive red junglefowl, the ancestor of domestic chickens, how inhibitory control associated with affective states. Specifically, we measured inhibitory control with a detour task, and negative and positive affective states with a tonic immobility test and a cognitive judgement bias test, respectively. Cognition and behaviour can differ between ages and sexes. Therefore, we investigated how inhibitory control related to affective states in younger chicks (≈2.5 weeks old), older chicks (≈5 weeks old) and sexually mature adults (≈28 weeks old) of both sexes. In younger chicks, poorer inhibitory control associated with less negative, more positive, affective states. We found no relationship between inhibitory control and affective states in older chicks or adults, nor sex differences regarding how inhibitory control related to affective states. Overall, our results suggest that inhibitory control can link to affective states and that the nature of these links can change over ontogeny.

Variation in inhibitory control does not influence social rank, foraging efficiency, or risk taking, in red junglefowl females.

Individual variation in cognition, seen in many taxa, is not well understood, despite its potential evolutionary consequences. Inhibitory control is an aspect of cognition which differs between individuals. However, how selection could act on this variation remains unclear. First, individual consistency over time of behaviours affected by inhibitory control, and how these behaviours relate to each other, is not well understood. Second, consequences in ecologically relevant contexts of variation in behaviours affected by  inhibitory control, are scarcely investigated. Therefore, we explored the temporal consistency and inter-relatedness of two behaviours influenced by inhibitory control (impulsive action and persistence) and how these link to social rank, foraging efficiency, and risk taking in adult female red junglefowl (Gallus gallus). We measured impulsive action in a detour test, and persistence in both a detour test and a foraging test. Impulsive action and persistence, measured in a detour test, were moderately consistent over time, and positively correlated. This implies that selection could act on inhibitory control via these behaviours, and selection on one behaviour could affect the other. However, we found no evidence of links between inhibitory control and social rank, foraging efficiency, or risk taking. This implies that selection may not act on inhibitory control via these measures, and that, in general, there may be a lack of strong selection on inhibitory control. This, in turn, could help explain individual variation in this aspect of cognition. Future research should explore the specificity of when inhibitory control has implications for individuals, and continue to investigate how variation in cognitive traits influences how individuals behave in contexts with potential evolutionary implications.

Manipulating monoamines reduces exploration and boldness of Mediterranean field crickets.

Despite the prevalence and research interest of animal personality, its underlying mechanisms are not yet fully understood. Due to the essential role of monoamines in modulating behaviour, we manipulated the monoaminergic systems of Mediterranean field crickets (Gryllus bimaculatus) to explore whether this altered behavioural responses commonly used to describe animal personality. Previous work has shown that both serotonin and dopamine manipulations can alter cricket behaviour, although results differ depending on the drug in focus. Here, we investigate the effect of Fluphenazine, a dopamine antagonist which also interacts with serotonin receptors, on activity, exploration, boldness, and aggression. These results are compared with those of our earlier work that investigated the effect of drugs that more specifically target serotonin or dopamine systems (Fluoxetine and Ropinirole, respectively). Due to limited research on dose-effects of Fluphenazine, we created dose-response curves with concentrations ranging from those measured in surface waters up to human therapeutic doses. We show that compared to control animals, Fluphenazine manipulation resulted in lower levels of both exploration and boldness, but did not affect activity nor aggression. The effect on explorative behaviour contradicts our previous results of serotonin and dopamine manipulations. These results together confirm the causal role of monoamines in explaining variation in behaviour often used to describe animal personality, effects that can be both dose- and behaviour-dependent. Further, our results suggest that previous results assigned specifically to the dopaminergic system, may at least partly be explained by effects of the serotonergic system. Thus, future studies should continue to investigate the explicit underlying roles of specific monoamines in explaining behavioural variation.

The relationship between monoaminergic gene expression, learning, and optimism in red junglefowl chicks.

Intra-species cognitive variation is commonly observed, but explanations for why individuals within a species differ in cognition are still understudied and not yet clear. Cognitive processes are likely influenced by genetic differences, with genes in the monoaminergic systems predicted to be important. To explore the potential role of these genes in association with individual variation in cognition, we exposed red junglefowl (Gallus gallus) chicks to behavioural assays measuring variation in learning (discriminative learning, reversal learning, and cognitive flexibility) and optimism (measured in a cognitive judgement bias test). Following this, we analysed prefrontal cortex gene expression of several dopaminergic and serotonergic genes in these chicks. Of our explored genes, serotonin receptor genes 5HT2A and 5HT2B, and dopaminergic receptor gene DRD1 were associated with measured behaviour. Chicks that had higher 5HT2A were less flexible in the reversal learning task, and chicks with higher 5HT2B also tended to be less cognitively flexible. Additionally, chicks with higher DRD1 were more optimistic, whilst chicks with higher 5HT2A tended to be less optimistic. These results suggest that the serotonergic and dopaminergic systems are linked to observed cognitive variation, and, thus, individual differences in cognition can be partially explained by variation in brain gene expression.

Experimental manipulation of monoamine levels alters personality in crickets.

Animal personality has been described in a range of species with ecological and evolutionary consequences. Factors shaping and maintaining variation in personality are not fully understood, but monoaminergic systems are consistently linked to personality variation. We experimentally explored how personality was influenced by alterations in two key monoamine systems: dopamine and serotonin. This was done using ropinirole and fluoxetine, two common human pharmaceuticals. Using the Mediterranean field cricket (Gryllus bimaculatus), we focused on the personality traits activity, exploration, and aggression, with confirmed repeatability in our study. Dopamine manipulations explained little variation in the personality traits investigated, while serotonin manipulation reduced both activity and aggression. Due to limited previous research, we created a dose-response curve for ropinirole, ranging from concentrations measured in surface waters to human therapeutic doses. No ropinirole dose level strongly influenced cricket personality, suggesting our results did not come from a dose mismatch. Our results indicate that the serotonergic system explains more variation in personality than manipulations of the dopaminergic system. Additionally, they suggest that monoamine systems differ across taxa, and confirm the importance of the mode of action of pharmaceuticals in determining their effects on behaviour.

Heritabilities and co-variation among cognitive traits in red junglefowl.

Natural selection can act on between-individual variation in cognitive abilities, yet evolutionary responses depend on the presence of underlying genetic variation. It is, therefore, crucial to determine the relative extent of genetic versus environmental control of these among-individual differences in cognitive traits to understand their causes and evolutionary potential. We investigated heritability of associative learning performance and of a cognitive judgement bias (optimism), as well as their covariation, in a captive pedigree-bred population of red junglefowl (Gallus gallus, n > 300 chicks over 5 years). We analysed performance in discriminative and reversal learning (two facets of associative learning), and cognitive judgement bias, by conducting animal models to disentangle genetic from environmental contributions. We demonstrate moderate heritability for reversal learning, and weak to no heritability for optimism and discriminative learning, respectively. The two facets of associative learning were weakly negatively correlated, consistent with hypothesized trade-offs underpinning individual cognitive styles. Reversal, but not discriminative learning performance, was associated with judgement bias; less optimistic individuals reversed a previously learnt association faster. Together these results indicate that genetic and environmental contributions differ among traits. While modular models of cognitive abilities predict a lack of common genetic control for different cognitive traits, further investigation is required to fully ascertain the degree of covariation between a broader range of cognitive traits and the extent of any shared genetic control.This article is part of the theme issue 'Causes and consequences of individual differences in cognitive abilities'.

Sophisticated Fowl: The Complex Behaviour and Cognitive Skills of Chickens and Red Junglefowl.

The world's most numerous bird, the domestic chicken, and their wild ancestor, the red junglefowl, have long been used as model species for animal behaviour research. Recently, this research has advanced our understanding of the social behaviour, personality, and cognition of fowl, and demonstrated their sophisticated behaviour and cognitive skills. Here, we overview some of this research, starting with describing research investigating the well-developed senses of fowl, before presenting how socially and cognitively complex they can be. The realisation that domestic chickens, our most abundant production animal, are behaviourally and cognitively sophisticated should encourage an increase in general appraise and fascination towards them. In turn, this should inspire increased use of them as both research and hobby animals, as well as improvements in their unfortunately often poor welfare.

Medical students' attitudes towards general practice and factors affecting career choice: a questionnaire study.

Background The current issue of general practice recruitment is a significant challenge and concern. In order to address this, it is vital to understand medical students' attitudes towards general practice and what influences their choice of intended career. Method We used a questionnaire study to examine these attitudes across all years at Imperial College Medical School and to understand what a group of London medical students' current intended career choices were. Results We found that only 13% of students ranked general practice as their first choice career despite having a generally realistic and positive attitude towards the speciality. They highlighted that the main influence on future career choice was interest in the specific speciality and that lifestyle factors did not seem to be so important. Conclusion Exposure to general practice, primary care research and student GP societies might play some part in increasing interest, but more work is needed to understand why students are rejecting general practice and what we can do collectively to attract students into choosing a career in primary care.