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Individual variation in how animals react to stress and environmental change has become a central topic in a wide range of biological disciplines, from evolutionary ecology to biomedicine. Such variation manifests phenotypically as correlated trait-clusters (referred to as coping styles, behavioral syndromes, shyness-boldness, or personality traits). Thresholds for switching from active coping (fight-flight) to inhibition and passive behavior when exposed to stress depend on experience and genetic factors. Comparative research has revealed a range of neuroendocrine-behavioral associations which are conserved throughout the vertebrate subphylum, including factors affecting perception, learning, and memory of stimuli and events. Here we review conserved aspects of the contribution of neurogenesis and other aspects of neural plasticity to stress coping. In teleost fish, brain cell proliferation and neurogenesis have received recent attention. This work reveals that brain cell proliferation and neurogenesis are associated with heritable variation in stress coping style, and they are also differentially affected by short- and long-term stress in a biphasic manner. Routinedependent and inflexible behavior in proactive individuals is associated with limited neural plasticity. These evolutionarily conserved relationships hold the potential to illuminate the biological background for stress-related neurobiological disorders. [ABSTRACT FROM AUTHOR]

Abstract: Physiological and behavioural responses to environmental change are individually variable traits, which manifest phenotypically and are subject to natural selection as correlated trait-clusters (coping styles, behavioural syndromes, or personality traits). Comparative research has revealed a range of neuroendocrine–behavioural associations which are conserved throughout the vertebrate subphylum. Regulatory mechanisms universally mediate a switch between proactive (e.g. active/aggressive) and reactive (e.g. conservation/withdrawal) behaviour in response to unpredictable and uncontrollable events. Thresholds for switching from active coping to behavioural inhibition are individually variable, and depend on experience and genetic factors. Such factors affect physiological stress responses as well as perception, learning, and memory. Here we review the role of an important contributor to neural processing, the set of biochemical, molecular, and structural processes collectively referred to as neural plasticity. We will concentrate on work in teleost fishes, while also elucidating conserved aspects. In fishes, environmental and physiological control of brain cell proliferation and neurogenesis has received recent attention. This work has revealed that the expression of genes involved in CNS plasticity is affected by heritable variation in stress coping style, and is also differentially affected by short- and long-term stress. Chronic stress experienced by subordinate fish in social hierarchies leads to a marked suppression of brain cell proliferation. Interestingly, typically routine dependent and inflexible behaviour in proactive individuals is also associated with low transcription of neurogenesis related genes. The potential for these findings to illuminate stress-related neurobiological disorders in other vertebrates is also discussed. [Copyright &y& Elsevier]

Abstract: Here we use a comparative model to investigate how behavioral and physiological traits correlate with neural plasticity. Selection for divergent post-stress cortisol levels in rainbow trout (Oncorhynchus mykiss) has yielded low- (LR) and high responsive (HR) lines. Recent reports show low behavioral flexibility in LR compared to HR fish and we hypothesize that this divergence is caused by differences in neural plasticity. Genes involved in neural plasticity and neurogenesis were investigated by quantitative PCR in brains of LR and HR fish at baseline conditions and in response to two different stress paradigms: short-term confinement (STC) and long-term social (LTS) stress. Expression of proliferating cell nuclear antigen (PCNA), neurogenic differentiation factor (NeuroD) and doublecortin (DCX) was generally higher in HR compared to LR fish. STC stress led to increased expression of PCNA and brain-derived neurotrophic factor (BDNF) in both lines, whereas LTS stress generally suppressed PCNA and NeuroD expression while leaving BDNF expression unaltered. These results indicate that the transcription of neuroplasticity-related genes is associated with variation in coping style, while also being affected by STC – and LTS stress in a biphasic manner. A higher degree of neural plasticity in HR fish may provide the substrate for enhanced behavioral flexibility. [Copyright &y& Elsevier]

Abstract: Compared to mammals, teleost fish have a very high rate of adult brain cell proliferation. Still little is known about how this process is regulated in comparative models, and what its functional implications are. We investigated the effect of stressful social interaction on brain cell proliferation in size matched rainbow trout pairs after the formation of stable social hierarchies. After 4 days of interaction, socially subordinate fish displayed common signs of chronic stress including reduced feeding behaviour, elevated plasma cortisol levels, and up-regulated brain stem 5-HT activity. The number of newborn cells in the telencephalon was quantified using immunohistochemistry for the exogenously administered S-phase marker BrdU. Subordinate fish had 40% fewer BrdU-positive telencephalic cells compared to isolated controls, while dominant individuals showed a non-significant tendency towards reduced cell proliferation. Cell proliferation in subordinate animals correlated negatively with aggression received immediately after hierarchy formation, indicating that the level of cell division suppression is related to the severity of the social stressor. These findings are comparable to findings in mammalian models of psychosocial stress, indicating that the suppressive effect of social stress on brain cell proliferation is conserved, and thus likely confers adaptive benefits throughout the vertebrate subphylum. [Copyright &y& Elsevier]

Abstract: The fish brain grows throughout life, and new cells are added continuously in all major brain areas. As in mammals, the rate of adult brain cell proliferation in fish can be regulated by external factors including environmental complexity and interaction with conspecifics. We have recently demonstrated that the stress experienced by subordinate rainbow trout in social hierarchies leads to a marked suppression of brain cell proliferation in the telencephalon, and that this is accompanied by an increase in plasma levels of cortisol. Corticosteroid hormones are known to suppress adult neurogenesis in mammals, and to investigate whether this is also the case in fish, rainbow trout were fed feed containing either a low or a high dose of cortisol for 6days. Compared to control animals receiving regular feed, both cortisol treated groups had significantly elevated cortisol levels 24h after the last feeding, with the high group having levels comparable to those previously reported in socially stressed fish. To quantify cell proliferation, immunohistochemistry for proliferating cell nuclear antigen (PCNA) was performed to identify actively cycling cells. The density of PCNA-positive nuclei in the telencephalon was reduced by about 50% in both cortisol treated groups. The effect of cortisol on brain cell proliferation did not reflect a general down regulation of growth, as only the high cortisol group had reduced growth rate, and there was no correlation between brain cell proliferation and growth rate in any group. These results indicate that the reduced proliferative activity seen in brains of socially stressed fish is mediated by cortisol, and that there is a similar suppressive effect of cortisol on brain cell proliferation in the teleost forebrain as in the mammalian hippocampus. [Copyright &y& Elsevier]

Abstract: Comparative studies on neural plasticity in non-mammalian vertebrates are increasingly promoted as an important complement to mammalian models. In teleost fishes the number of brain cells increases with age, body weight, and body length throughout life. Neurogenesis persists to a large degree, and both neuron replacement and net brain growth occur during adulthood. Whether environmental factors affect brain cell proliferation has however been scarcely investigated in this animal group. In the current study adult male zebrafish were kept in social isolation in different environments (enriched vs. barren) for one week. Telencephalic cell proliferation was investigated by proliferating cell nuclear antigen (PCNA) immunohistochemistry. Higher numbers of PCNA positive nuclei and significantly increased inter-individual variability was observed in fish kept in aquaria enriched with artificial plants and gravel. Zebrafish rapidly regained feed intake after transfer to social isolation. Whole-body cortisol levels were also generally low in isolated fish, although slightly elevated in fish from enriched environments. In summary, this study demonstrates that environmental alterations can rapidly alter cell cycle dynamics in the zebrafish brain. Furthermore, the results support the idea that mild short-term stressors and concomitant small increases in corticosteroid exposure stimulate brain cell proliferation. [Copyright &y& Elsevier]

Salmonid fishes such as the rainbow trout (Oncorhynchus mykiss) are frequently used to study behavioral and neuroendocrine effects of socially induced stress. A predictable aggressive response to territorial intrusion, a well described neuroanatomy, and many essential similarities in the stress response in fishes and other vertebrates are among the advantages of this comparative model. One conspicuous difference when compared to mammals, however, is that in telost fish and other non-mammalian vertebrates, neurogenesis persists into adulthood to a much higher degree. Very little is known about the functional significance of individual differences in the rate of brain cell proliferation in fish, or whether structural changes in the fish brain are influenced by the social environment. In this paper we discuss the observation that brain cell proliferation is reduced in subordinate fish, focusing in particular on whether such individual variation reflects a difference in coping style or is indeed a response to social interactions. Copyright © 2007 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]