Temperament describes the idea that individual behavioural differences are repeatable over time and across situations. This common phenomenon covers numerous traits, such as aggressiveness, avoidance of novelty, willingness to take risks, exploration, and sociality. The study of temperament is central to animal psychology, behavioural genetics, pharmacology, and animal husbandry, but relatively few studies have examined the ecology and evolution of temperament traits. This situation is surprising, given that temperament is likely to exert an important influence on many aspects of animal ecology and evolution, and that individual variation in temperament appears to be pervasive amongst animal species. Possible explanations for this neglect of temperament include a perceived irrelevance, an insufficient understanding of the link between temperament traits and fitness, and a lack of coherence in terminology with similar traits often given different names, or different traits given the same name. We propose that temperament can and should be studied within an evolutionary ecology framework and provide a terminology that could be used as a working tool for ecological studies of temperament. Our terminology includes five major temperament trait categories: shyness-boldness, exploration-avoidance, activity, sociability and aggressiveness. This terminology does not make inferences regarding underlying dispositions or psychological processes, which may have restrained ecologists and evolutionary biologists from working on these traits. We present extensive literature reviews that demonstrate that temperament traits are heritable, and linked to fitness and to several other traits of importance to ecology and evolution. Furthermore, we describe ecologically relevant measurement methods and point to several ecological and evolutionary topics that would benefit from considering temperament, such as phenotypic plasticity, conservation biology, population sampling, and invasion biology.

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Integrating animal temperament within ecology and evolution.

EVOLUTION: Of Mice . . .

Biological reactivity to psychological stressors comprises a complex, integrated, and highly conserved repertoire of central neural and peripheral neuroendocrine responses designed to prepare the organism for challenge or threat. Developmental experience plays a role, along with heritable, polygenic variation, in calibrating the response dynamics of these systems, with early adversity biasing their combined effects toward a profile of heightened or prolonged reactivity. Conventional views of such high reactivity suggest that it is an atavistic and pathogenic legacy of an evolutionary past in which threats to survival were more prevalent and severe. Recent evidence, however, indicates that (a) stress reactivity is not a unitary process, but rather incorporates counterregulatory circuits serving to modify or temper physiological arousal, and (b) the effects of high reactivity phenotypes on psychiatric and biomedical outcomes are bivalent, rather than univalent, in character, exerting both risk-augmenting and risk-protective effects in a context-dependent manner. These observations suggest that heightened stress reactivity may reflect, not simply exaggerated arousal under challenge, but rather an increased biological sensitivity to context, with potential for negative health effects under conditions of adversity and positive effects under conditions of support and protection. From an evolutionary perspective, the developmental plasticity of the stress response systems, along with their structured, context-dependent effects, suggests that these systems may constitute conditional adaptations: evolved psychobiological mechanisms that monitor specific features of childhood environments as a basis for calibrating the development of stress response systems to adaptively match those environments. Taken together, these theoretical perspectives generate a novel hypothesis: that there is a curvilinear, U-shaped relation between early exposures to adversity and the development of stress-reactive profiles, with high reactivity phenotypes disproportionately emerging within both highly stressful and highly protected early social environments.

https://cals.arizona.edu/fcs/sites/cals.arizona.edu.fcs/files/D&P%20Boyce%20&%20Ellis%202005.pdf

Biological sensitivity to context: I. An evolutionary–developmental theory of the origins and functions of stress reactivity

The author explores the viability of a comparative approach to personality research. A review of the diverse animal-personality literature suggests that (a) most research uses trait constructs, focuses on variation within (vs. across) species, and uses either behavioral codings or trait ratings; (b) ratings are generally reliable and show some validity (7 parameters that could influence reliability and 4 challenges to validation are discussed); and (c) some dimensions emerge across species, but summaries are hindered by a lack of standard descriptors. Arguments for and against cross-species comparisons are discussed, and research guidelines are suggested. Finally, a research agenda guided by evolutionary and ecological principles is proposed. It is concluded that animal studies provide unique opportunities to examine biological, genetic, and environmental bases of personality and to study personality change, personality-health links, and personality perception.

From mice to men: what can we learn about personality from animal research?

Assessment of positive emotions in animals to improve their welfare.

Information processing by humans can be biased by their emotions — for example, anxious and depressed people tend to make negative judgements about events and to interpret ambiguous stimuli unfavourably1,2,3,4. Here we show that such a 'pessimistic' response bias can also be measured in rats that are housed in unpredictable conditions5,6. Our findings indicate that cognitive bias can be used as an indicator of affective state in animals, which should facilitate progress in animal-welfare studies.

Animal behaviour: Cognitive bias and affective state

Measuring emotional processes in animals: the utility of a cognitive approach

A better understanding of animal emotion is an important goal in disciplines ranging from neuroscience to animal welfare science. The conscious experience of emotion cannot be assessed directly, but neural, behavioural and physiological indicators of emotion can be measured. Researchers have used these measures to characterize how animals respond to situations assumed to induce discrete emotional states (e.g. fear). While advancing our understanding of specific emotions, this discrete emotion approach lacks an overarching framework that can incorporate and integrate the wide range of possible emotional states. Dimensional approaches that conceptualize emotions in terms of universal core affective characteristics (e.g. valence (positivity versus negativity) and arousal) can provide such a framework. Here, we bring together discrete and dimensional approaches to: (i) offer a structure for integrating different discrete emotions that provides a functional perspective on the adaptive value of emotional states, (ii) suggest how long-term mood states arise from short-term discrete emotions, how they also influence these discrete emotions through a bi-directional relationship and how they may function to guide decision-making, and (iii) generate novel hypothesis-driven measures of animal emotion and mood.

https://royalsocietypublishing.org/doi/pdf/10.1098/rspb.2010.0303

Michael Mendl

Papers

Animal behaviour: Cognitive bias and affective state

Measuring emotional processes in animals: the utility of a cognitive approach

An integrative and functional framework for the study of animal emotion and mood

Cognitive bias as an indicator of animal emotion and welfare: Emerging evidence and underlying mechanisms

Revisiting translocation and reintroduction programmes: the importance of considering stress