Atlantic salmon Salmo salar, brown trout Salmo trutta (including the anadromous form, sea trout) and Arctic charr Salvelinus alpinus (including anadromous fish) provide important commercial and sports fisheries in Western Europe. As water temperature increases as a result of climate change, quantitative information on the thermal requirements of these three species is essential so that potential problems can be anticipated by those responsible for the conservation and sustainable management of the fisheries and the maintenance of biodiversity in freshwater ecosystems. Part I compares the temperature limits for survival, feeding and growth. Salmo salar has the highest temperature tolerance, followed by S. trutta and finally S. alpinus. For all three species, the temperature tolerance for alevins is slightly lower than that for parr and smolts, and the eggs have the lowest tolerance; this being the most vulnerable life stage to any temperature increase, especially for eggs of S. alpinus in shallow water. There was little evidence to support local thermal adaptation, except in very cold rivers (mean annual temperature <6·5° C). Part II illustrates the importance of developing predictive models, using data from a long-term study (1967-2000) of a juvenile anadromous S. trutta population. Individual-based models predicted the emergence period for the fry. Mean values over 34 years revealed a large variation in the timing of emergence with c. 2 months between extreme values. The emergence time correlated significantly with the North Atlantic Oscillation Index, indicating that interannual variations in emergence were linked to more general changes in climate. Mean stream temperatures increased significantly in winter and spring at a rate of 0·37° C per decade, but not in summer and autumn, and led to an increase in the mean mass of pre-smolts. A growth model for S. trutta was validated by growth data from the long-term study and predicted growth under possible future conditions. Small increases (<2·5° C) in winter and spring would be beneficial for growth with 1 year-old smolts being more common. Water temperatures would have to increase by c. 4° C in winter and spring, and 3° C in summer and autumn before they had a marked negative effect on trout growth.
SUMMARYIn fishes, performance failure at high temperature is thought to be due to a limitation on oxygen delivery (the theory of oxygen and capacity limited thermal tolerance, OCLTT), which suggests that thermal tolerance and hypoxia tolerance might be functionally associated. Here we examined variation in temperature and hypoxia tolerance among 41 families of Atlantic salmon (Salmo salar), which allowed us to evaluate the association between these two traits. Both temperature and hypoxia tolerance varied significantly among families and there was a significant positive correlation between critical maximum temperature (CT max ) and hypoxia tolerance, supporting the OCLTT concept. At the organ and cellular levels, we also discovered support for the OCLTT concept as relative ventricle mass (RVM) and cardiac myoglobin (Mb) levels both correlated positively with CT max (R 2 =0.21, P<0.001 and R 2 =0.17, P=0.003, respectively). A large RVM has previously been shown to be associated with high cardiac output, which might facilitate tissue oxygen supply during elevated oxygen demand at high temperatures, while Mb facilitates the oxygen transfer from the blood to tissues, especially during hypoxia. The data presented here demonstrate for the first time that RVM and Mb are correlated with increased upper temperature tolerance in fish. High phenotypic variation between families and greater similarity among full-and half-siblings suggests that there is substantial standing genetic variation in thermal and hypoxia tolerance, which could respond to selection either in aquaculture or in response to anthropogenic stressors such as global climate change.
The critical thermal maximum for salmon and trout parr was not affected significantly by age or acclimation temperature, and increased asymptotically with the rate of temperature increase. Mean thermal maxima were estimated with poor precision at high and low rates of temperature increase, and high precision at rates of 1 and 2" C h ~ I . 0 1995 The Fisheries Society of the B r m h Isles
Comparative genome scans can be used to identify chromosome regions, but not traits, that are putatively under selection. Identification of targeted traits may be more likely in recently domesticated populations under strong artificial selection for increased production. We used a North American Atlantic salmon 6K SNP dataset to locate genome regions of an aquaculture strain (Saint John River) that were highly diverged from that of its putative wild founder population (Tobique River). First, admixed individuals with partial European ancestry were detected using STRUCTURE and removed from the dataset. Outlier loci were then identified as those showing extreme differentiation between the aquaculture population and the founder population. All Arlequin methods identified an overlapping subset of 17 outlier loci, three of which were also identified by BayeScan. Many outlier loci were near candidate genes and some were near published quantitative trait loci (QTLs) for growth, appetite, maturity, or disease resistance. Parallel comparisons using a wild, nonfounder population (Stewiacke River) yielded only one overlapping outlier locus as well as a known maturity QTL. We conclude that genome scans comparing a recently domesticated strain with its wild founder population can facilitate identification of candidate genes for traits known to have been under strong artificial selection.
I 1. The chief objective was to determine the upper and lower thermal limits for feeding and survival in the stone loach, Noemacheilus barbatulus, using juveniles (total length 30-45 mm, live weight 0.25-0.80 g) from one population and adults (total length 77-100mm, live weight 3.6-7.9g) from three populations. 2. Fish were acclimatized to constant temperatures of 3, 7, 10, 15, 20, 25 and 27°C; then the temperature was changed at a rate of rC/30min to determine the critical limits for feeding, survival over 7 days (indpient lethal temperature), or survival for 10 min or less (ultimate lethal temperature). The rate of rC/30min was the optimum value from preliminary experiments, using nine rates from 0.5°C/48h to 18''Ch~^ As values for adults were not significantly different between populations, they were pooled to provide arithmetic means (with 95% CL) for the thermal limits at each acclimation temperature. 3. Feeding limits increased with acclimation temperature to upper and lower mean values of 28.0 ± 0.15°C and 5.1 ± 0.55°C for adults, 25.0 ± 0.54°C and 6.1 ± 0.92°C for juveniles. Indpient lethal levels defined a tolerance zone within which stone loach survive for a considerable time; upper limits increased with acclimation temperature to reach a maximum plateau of 29.1 ± 0.18°C for adults and 29.0 ± 0.40°C for juveniles; lower limits also increased from near 0°C to 3.0 ± 0.40°C for adults and juveniles. Upper limits for the ultimate lethal level increased with acclimation temperature to a maximum plateau of 33.5''C for adults (95% CL ± 0.19) and juveniles (95% CL ± 0.40), whilst the lower limits increased from near 0°C to 2.5 ± 0.30°C. At acclimation temperatures below 20°C, upper indpient and ultimate lethal values were significantly lower for juveniles than those for adults. 4. The thermal tolerance of stone loach was higher than that of juvenile Atlantic salmon or brown trout, one or both of these spedes often being dominant in streams with stone loach.
1. The objective was to determine the thermal limits for feeding and sur\'ival in the bullhead, Cottus gobio, using juveniles (total length 20-30 mm, live weight 0.5-1.5 g) from one population and adults (50-70 mm, 3.5-5.5 g) from three populations. 2. Fish were acclimated to constant temperatures (3, 7, 10, 15, 20, 25 or 27 °C) and the temperature was then changed at a rate of 1 °C /30 min to determine the critical limits for feeding, survival over 7 days (incipient lethal temperature), or survival for 10 min or less (ultimate lethal temperature). The rate of 1 °C/30 min was the optimum value from preliminary experiments, using nine rates from 0.5 °C/48 h to 18 °C h"'. As values for adults were not significantly different between populations, they were pooled to provide arithmetic means (with 95% CL) for the thermal limits at each acclimation temperature. 3. Feeding limits increased with acclimation temperature to upper and lower mean values (± 95% CL) of 26.5 ± 0.16 °C and 4.2 ± 0.20 °C for adults, 26.6 ± 0.59 "C and 5.0 ± 0.55 °C for juveniles. Incipient lethal levels defined a tolerance zone within which fish survive indefinitely; upper limits increased with acclimation temperature to a plateau of 27.6 ± 0.22 °C for adults and 27.5 ± 0.47 "C for juveniles, lower limits increased from near 0 °C to 2.5 ± 0.31 °C for adults and 2.7 ± 0.47 °C for juveniles. Ultimate lethal levels increased with acclimation temperature to a plateau of 32.5 ± 0.24 °C for adults and 32.6 ± 0.46 °C for juveniles, whilst the lower limits increased from near 0 to 0.9 ± 0.29 °C. Upper feeding, incipient and ultimate lethal values were significantly lower for juveniles than those for adults at acclimation temperatures < 20, < 20 and < 15 "C, respectively. 4. The thermal tolerance of bullheads was slightly lower than that of stone loach, similar to that of juvenile Atlantic salmon and higher than that of brown trout; the thermal limits for feeding were much wider than those for salmon or trout.
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