A comparison of plasma glucose and plasma cortisol as selection markers for high and low stress-responsiveness in female rainbow trout
Introduction
Repeated or prolonged exposure of salmonid fish to stressful stimuli can result in reduced growth, impaired reproductive performance and immunosuppression (Barton, 1997). A wide range of responsiveness to stressors has been reported both for strains of salmonid fish (Pickering and Pottinger, 1989; McGeer et al., 1991) and for individual fish within strains (Pottinger et al., 1992). Because stress is an unavoidable component of the finfish aquaculture environment, and because of the adverse effects of chronic stress on performance characteristics, it has been suggested that fish which display a lower degree of responsiveness to stressors may perform better under intensive culture conditions than those which display a higher level of responsiveness (Pottinger and Pickering, 1997). It is also acknowledged that the converse may be true; given the adaptive significance of the stress response, high-responding fish may prove to be at a greater advantage, or low-responding fish at a relative disadvantage, overall. The prospect of selectively breeding fish for reduced responsiveness to stress has been addressed with respect to disease resistance (Fevolden et al., 1993) and overall performance (Pottinger et al., 1994). It is clear from these studies and similar work in poultry (Satterlee and Johnson, 1988), that selection of fish on the basis of stress responsiveness is practicable. However, as yet, no thorough evaluation of the performance of fish selected for divergent stress responsiveness exists. The present study was carried out as one of a number of preliminary elements of a selective breeding programme, the aim of which is to generate lines of rainbow trout (Oncorhynchus mykiss) which are divergent for stress-responsiveness. If successful, this will allow the relative performance of fish with high- and low-responsiveness to stressors to be compared and permit an assessment to be made of which trait, if either, is desirable in fish under aquaculture conditions.
The changes in blood cortisol levels which occur in fish following the perception of a stressful stimuli are widely employed as a reliable index of activation of the neuroendocrine stress response (Barton and Iwama, 1991). Furthermore, cortisol itself is closely linked to many of the adverse consequences of chronic stress including effects on growth (Pickering, 1993), reproduction (Pickering et al., 1987), and the immune system (Pickering and Pottinger, 1989). Consequently, the extent of cortisol elevation following exposure to a standardised stressor has been adopted as a trait of physiological significance on which a selection procedure may be based (Fevolden et al., 1991; Pottinger et al., 1994). Blood glucose levels are also increased in fish following exposure to stressful stimuli. In the short-term, hyperglycaemia is mediated primarily as a consequence of the rapid increase in blood catecholamines which occurs during the first phase of the stress response (Wendelaar Bonga, 1997) although in the longer-term, cortisol-dependent gluconeogenesis may be responsible (van Raaij et al., 1996). The measurement of post-stress blood glucose levels therefore offers an indirect alternative marker of the magnitude of the neuroendocrine stress response. The measurement of blood glucose can be carried out rapidly using portable equipment (Morgan and Iwama, 1997), in contrast to the more sophisticated facilities required for the determination of cortisol. This is an important consideration when assessing the practicality of employing either selection trait within a commercial environment.
The aim of this study was to compare the utility of both plasma cortisol and plasma glucose levels as selection markers for stress responsiveness and to assess the reliability of each as identifiers of consistency in stress-responsiveness with time. Because uncertainty exists regarding the relative benefits of a high or low responsiveness to stressors, individuals with divergent responses were sought to permit comparison.
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Fish
During February 1996, 125 2-year old female rainbow trout (Stirling strain) were divided evenly between five 1500-l holding tanks (25 fish/tank), each supplied with a constant flow of lake water (25 l min−1). Each fish was weighed, measured, and individually tagged by two methods. A passive integrated transponder (PIT) tag (Fish Eagle) was inserted into the peritoneal cavity of each fish and a visible implant (VI) tag (Northwest Marine Technology) was inserted under the clear tissue posterior
Selection by post-stress cortisol level
The procedure by which HRC and LRC fish were selected within a single experimental tank, on the basis of their plasma cortisol response to a 3 h confinement stress, is illustrated by Table 1. The mean post-stress plasma cortisol level was determined for each fish from the plasma cortisol values obtained on each of the sample dates (March–July). The mean plasma cortisol values were then ranked and the four fish with lowest mean post-stress cortisol levels and the four fish with highest mean
Discussion
These results confirm and extend earlier observations (Pottinger et al., 1992) that the cortisol response to confinement stress in rainbow trout is an individual characteristic that is stable with time, allowing individual fish with divergent responses to be selected. These data also demonstrate that fish displaying a hyperglycaemic response to confinement, which is consistently higher or lower than the population mean, may be identified. However, selection of individuals on the basis of their
Acknowledgements
This study forms part of a project co-funded by the European Commission (contract FAIR-CT95-0152) and the Natural Environment Research Council, UK.
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