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.

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.

Why individuals differ in behavioral responses has received intense research attention (particularly in the context of animal personality), and has typically focused on describing variation in boldness, activity, and exploration. Nevertheless, the mechanisms underlying this behavioral variation remain largely unknown. Variation in these behaviors is likely influenced by genetic differences between individuals, with genes in the monoaminergic systems commonly implicated. When examining the link between variation in boldness, activity and exploration and genes, studies have focused on different monoaminergic systems (mainly serotonergic and dopaminergic), and predominantly on mammals and passerine birds. Therefore, to replicate this general approach and examine if genes from these systems are linked to boldness, activity, and exploration, we exposed red junglefowl chicks (Gallus gallus) to behavioral assays (measuring boldness, activity, exploration) before analyzing prefrontal cortex gene expression of several dopaminergic (DRD1, DRD2) and serotonergic genes (TPH, 5HT2A, 5HT2B, 5HT2C, 5HT1B). We observed no relationships between our measured behaviors and gene expression. Together with previous studies, our results suggest that a clear link between boldness, activity and exploration and monoaminergic gene variation is lacking. We, therefore, suggest that this is due to differences among studies (e.g., methodological differences), or that the nature of the relationship between these behaviors and monoaminergic systems is more species-specific, and/or more complex than so far assumed.

Among-individual behavioral differences (i.e. animal personality) are commonly observed across taxa, although the underlying, causal mechanisms of such differences are poorly understood. Animal personality has been correlated with physiological functions as well as fitness-related traits. Variation in many aspects of monoamine systems, such as metabolite levels and gene polymorphisms, has been linked to behavioral variation. Therefore, here we experimentally investigated the potential role of monoamines in explaining individual variation in personality, using two common pharmaceuticals that respectively alter the levels of serotonin and dopamine in the brain: fluoxetine and ropinirole. We exposed three-spined sticklebacks, a species that shows animal personality, to either chemical alone or to a combination of the two chemicals, for 18 days. During the experiment, fish were assayed at four time points for the following personality traits: exploration, boldness, aggression and sociability. To quantify brain gene expression on short- and longer-term scales, fish were sampled at two time points. Our results show that monoamine manipulations influence fish behavior. Specifically, fish exposed to either fluoxetine or ropinirole were significantly bolder, and fish exposed to the two chemicals together tended to be bolder than control fish. Our monoamine manipulations did not alter the gene expression of monoamine or stress-associated neurotransmitter genes, but control, untreated fish showed covariation between gene expression and behavior. Specifically, exploration and boldness were predicted by genes in the dopaminergic, serotonergic and stress pathways, and sociability was predicted by genes in the dopaminergic and stress pathways. These results add further support to the links between monoaminergic systems and personality, and show that exposure to monoamines can causally alter animal personality.

Among-individual behavioral differences (i.e. animal personality) are commonly observed across taxa, although the underlying, causal mechanisms of such differences are poorly understood. Animal personality has been implicated in correlations with physiological functions as well as affecting fitness-related traits. Variation in many aspects of monoamine systems, such as metabolite levels and gene polymorphisms, has been linked to behavioral variation. Therefore, here we investigated the potential role of monoamines in explaining individual variation in personality, using two common pharmaceuticals that respectively alter the levels of serotonin and dopamine in the brain: fluoxetine and ropinirole. We exposed three- spined sticklebacks, a species that shows animal personality, to either chemical alone or to a combination of the two chemicals, for 18 days. During the experiment, fish were assayed at four time points for the following personality traits: exploration, boldness, aggression and sociability. To quantify brain gene expression on short- and longer-term scales, fish were sampled at two time points. Our results show that monoamine manipulations influence fish behavior. Specifically, fish exposed to either fluoxetine or ropinirole were significantly bolder, and fish exposed to the two chemicals together tended to be bolder than control fish. Our monoamine manipulations did not alter the gene expression of monoamine or stress-associated neurotransmitter genes, but control, untreated fish showed covariation between gene expression and behavior. Specifically, exploration and boldness were predicted by genes in the dopaminergic, serotonergic and stress pathways, and sociability was predicted by genes in the dopaminergic and stress pathways. These results add further support to the links between monoaminergic systems and personality, and show that exposure to monoamines can causally alter animal personality.

Cognitive processes are often biased by emotions. In humans, affective disorders are accompanied by pessimistic judgement, while optimistic judgement is linked to emotional stability. Similar to humans, animals tend to interpret ambiguous stimuli negatively after experiencing stressful events, although the long-lasting impact on judgement bias has rarely been investigated. We measure judgement bias in female chicks (Gallus gallus domesticus) after exposure to cold stress, and before and after exposure to additional unpredictable stressors. Additionally, we explore if brain monoamines can explain differences in judgement bias. Chicks exposed to cold stress did not differ in judgement bias compared to controls, but showed sensitivity to additional stressors by having higher motivation for social reinstatement. Environmental complexity reduced stress-induced negative judgement bias, by maintaining an optimistic bias in individuals housed in complex conditions even after stress exposure. Moreover, judgement bias was related to dopamine turnover rate in mesencephalon, with higher activity in individuals that had a more optimistic response. These results demonstrate that environmental complexity can buffer against negative effects of additive stress and that dopamine relates to judgement bias in chicks. These results reveal that both internal and external factors can mediate emotionally biased judgement in animals, thus showing similarities to findings in humans.

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.

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.

The causes of individual variation in behavior are often not well understood, and potential underlying mechanisms include both intrinsic and extrinsic factors, such as early environmental, physiological, and genetic differences. In an exploratory laboratory study, we raised three-spined sticklebacks <i>(Gasterosteus aculeatus)</i> under 4 different environmental conditions (simulated predator environment, complex environment, variable social environment, and control). We investigated how these manipulations related to behavior, brain physiology, and gene expression later in life, with focus on brain dopamine and serotonin levels, turnover rates, and gene expression. The different rearing environments influenced behavior and gene expression, but did not alter monoamine levels or metabolites. Specifically, compared to control fish, fish exposed to a simulated predator environment tended to be less aggressive, more exploratory, and more neophobic; and fish raised in both complex and variable social environments tended to be less neophobic. Exposure to a simulated predator environment tended to lower expression of dopamine receptor DRD4A, a complex environment increased expression of dopamine receptor DRD1B, while a variable social environment tended to increase serotonin receptor 5-HTR2B and serotonin transporter SLC6A4A expression. Despite both behavior and gene expression varying with early environment, there was no evidence that gene expression mediated the relationship between early environment and behavior. Our results confirm that environmental conditions early in life can affect phenotypic variation. However, the mechanistic pathway of the monoaminergic systems translating early environmental variation into observed behavioral responses was not detected.