Behaviour represents a reaction to the environment as fish perceive it and is therefore a key element of fish welfare. This review summarises the main findings on how behavioural changes have been used to assess welfare in farmed fish, using both functional and feeling-based approaches. Changes in foraging behaviour, ventilatory activity, aggression, individual and group swimming behaviour, stereotypic and abnormal behaviour have been linked with acute and chronic stressors in aquaculture and can therefore be regarded as likely indicators of poor welfare. On the contrary, measurements of exploratory behaviour, feed anticipatory activity and reward-related operant behaviour are beginning to be considered as indicators of positive emotions and welfare in fish. Despite the lack of scientific agreement about the existence of sentience in fish, the possibility that they are capable of both positive and negative emotions may contribute to the development of new strategies (e.g. environmental enrichment) to promote good welfare. Numerous studies that use behavioural indicators of welfare show that behavioural changes can be interpreted as either good or poor welfare depending on the fish species. It is therefore essential to understand the species-specific biology before drawing any conclusions in relation to welfare. In addition, different individuals within the same species may exhibit divergent coping strategies towards stressors, and what is tolerated by some individuals may be detrimental to others. Therefore, the assessment of welfare in a few individuals may not represent the average welfare of a group and vice versa. This underlines the need to develop on-farm, operational behavioural welfare indicators that can be easily used to assess not only the individual welfare but also the welfare of the whole group (e.g. spatial distribution). With the ongoing development of video technology and image processing, the on-farm surveillance of behaviour may in the near future represent a low-cost, noninvasive tool to assess the welfare of farmed fish.
Individual differences in physiological and behavioural responses to stressors are increasingly recognised as adaptive variation and thus raw material for evolution and fish farming improvements including selective breeding. Such individual variation has been evolutionarily conserved and is present in all vertebrate taxa including fish. In farmed animals, the interest in consistent trait associations, that is coping styles, has increased dramatically over the last years because many studies have demonstrated links to performance traits, health and disease susceptibility and welfare. This study will review (i) the main behavioural, neuroendocrine, cognitive and emotional differences between reactive and proactive coping styles in farmed fish; (ii) the methodological approaches used to identify coping styles in farmed fish, including individual (group) mass-screening tests; and (iii) how knowledge on coping styles may contribute to improved sustainability of the aquaculture industry, including welfare and performance of farmed fish. Moreover, we will suggest areas for future research, where genetic basis (heritability/ epigenetic) of coping styles, and the neuroendocrine mechanisms behind consistent as well as flexible behavioural patterns are pinpointed as central themes. In addition, the ontogeny of coping styles and the influence of age, social context and environmental change in coping styles will also be discussed.
Behaviour represents a reaction to the environment as fish perceive it and is therefore a key element of fish welfare. This review summarises the main findings on how behavioural changes have been used to assess welfare in farmed fish, using both functional and feeling-based approaches. Changes in foraging behaviour, ventilatory activity, aggression, individual and group swimming behaviour, stereotypic and abnormal behaviour have been linked with acute and chronic stressors in aquaculture and can therefore be regarded as likely indicators of poor welfare. On the contrary, measurements of exploratory behaviour, feed anticipatory activity and rewardrelated operant behaviour are beginning to be considered as indicators of positive emotions and welfare in
Despite the use of fish models to study human mental disorders and dysfunctions, knowledge of regional telencephalic responses in nonmammalian vertebrates expressing alternative stress coping styles is poor. As perception of salient stimuli associated with stress coping in mammals is mainly under forebrain limbic control, we tested regionspecific forebrain neural (i.e. mRNA abundance and monoamine neurochemistry) and endocrine responses under basal and acute stress conditions for previously characterised proactive and reactive Atlantic salmon. Reactive fish showed a higher degree of the neurogenesis marker proliferating cell nuclear antigen ( pcna) and dopamine activity under basal conditions in the proposed hippocampus homologue (Dl) and higher post-stress plasma cortisol levels. Proactive fish displayed higher post-stress serotonergic signalling (i.e. higher serotonergic activity and expression of the 5-HT 1A receptor) in the proposed amygdala homologue (Dm), increased expression of the neuroplasticity marker brain-derived neurotropic factor (bdnf ) in both Dl and the lateral septum homologue (Vv), as well as increased expression of the corticotropin releasing factor 1 (crf 1 ) receptor in the Dl, in line with active coping neuro-profiles reported in the mammalian literature. We present novel evidence of proposed functional equivalences in the fish forebrain with mammalian limbic structures.
We investigated the relationship between social interactions, brain serotonergic activity, and two behavioural patterns in juvenile Arctic char (Salvelinus alpinus): feeding and spontaneous swimming activity. Dominant and subordinate individuals were observed during rearing in pairs, followed by rearing in isolation. Throughout the experiment, levels of both food intake and swimming activity remained high in dominant fish. When they were in pairs, food intake was completely inhibited in subordinate fish; thus, dominant fish were able to monopolise food. At the same time, brain serotonergic activity, as indexed by the ratio of 5-hydroxyindoleacetic acid (5-HIAA) to serotonin (5-hydroxytryptamine, 5-HT) was elevated in the hypothalamus and brain stem of subordinate fish compared with dominants. During subsequent rearing in isolation, food intake, but not spontaneous locomotor activity, gradually increased in previously subordinate fish, while serotonergic activity fell to near that of dominants. Thus, appetite inhibition in subordinate fish can be reversed by rearing in isolation, an effect that may be related to the reversal of a stress-induced activation of brain serotonergic neurones accompanying social subordination. Reduced swimming activity is either a long-lasting response to social subordination or reflects permanently different behavioural strategies of subordinate and dominant individuals.
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