The relationships between fish health, metabolic rate, swimming performance and recovery in return-run sockeye salmon, Oncorhynchus nerka (Walbaum)

The repeat swimming ability and oxygen uptake (Mo2) of adult sockeye salmon, Oncorhynchus nerka (Walbaum), were assessed at ambient water temperatures at three field locations along their migration route. Following these measurements, internal and external fish condition was evaluated according to United States Environmental Protection Agency guidelines. Here we report on the physiological characteristics of fish having either moderate or severe levels of disease and injury. Routine oxygen uptake (Mo2) did not differ between healthy fish and those with indices of ill health. In contrast, fish classified as sick, which included conditions of damaged internal organs, an Ichthyophonus spp. heart infection, a Saprolegnia spp. gill infection, and skin wounds, had a lower post-exercise Mo2 and were unable to repeat their critical swim speed (U(crit)) on the second swim test. Moderate levels of disease or injury did not significantly affect either U(crit) or post-exercise Mo2. We conclude that the ability of adult salmon to recover quickly from exercise may be a useful indicator of sublethal pathologies.     

Influence of the marine abundance of pink (Oncorhynchus gorbuscha) and sockeye salmon (O. nerka) on growth of Ozernaya River sockeye

The length and weight of Russian sockeye ( Oncorhynchus nerka ) returning to the Ozernaya River (Kamchatka) was substantially reduced in years when the ocean abundances of Kamchatkan pink ( O. gorbuscha ) and sockeye salmon were high. We found that the density-dependent reduction in sockeye growth on a per-capita basis was greater for sockeye than for pink salmon. However, the overall effect of pink salmon abundance on sockeye growth was greater because of the higher numerical abundance of pink salmon. The strongest statistical relationships were found for sockeye from separate age groups; pooled data combining all age classes were statistically insignificant. We estimate that, if pink salmon were absent, the most strongly affected age group of sockeye salmon (2.1 males) would weigh twice as much at maturity than if pink salmon populations from eastern and western Kamchatka were both simultaneously at peak observed abundances. Trophic competition in the ocean between pink and sockeye salmon can therefore have a significant influence on the productivity of sockeye populations for the most strongly affected age groups. These effects are large enough that they should be explicitly considered in the management of salmon populations.     

Seasonal changes of hormones and muscle enzymes in adult lacustrine masu (Oncorhynchus masou) and sockeye salmon (O. nerka)

The upstream migration of adult anadromous fishes is characterized by physiological changes in responses to reproductive and energetic challenges. This study analyzed the physiological responses of lake-resident anadromous masu salmon (Oncorhynchus masou) and sockeye salmon (O. nerka) to migration in order to determine if these fish might serve as a suitable model for ocean-running populations and to differentiate between physiological responses to reproduction and to exercise-linked aspects of migration. Reproductive (estradiol, testosterone, 11-ketotestosterone, 17α,20β-dihydroxy-4-pregnen-3-one) and metabolically-linked (thyroxine, triiodothyronine) hormones showed similar patterns to ocean-running anadromous populations. White muscle pyruvate kinase, lactate dehydrogenase and malate dehydrogenase decreased with the onset of spawning season while white muscle citrate synthase, β-hydroxyacetyl CoA dehydrogenase, phosphofructokinase and glutamate oxaloacetate transaminase did not, suggesting that the former group of enzymes are responding to reproductive or food intake signals while the second group, which typically change during anadromous migration, may be responding to exercise-linked aspects of migration. This revised version was published online in August 2006 with corrections to the Cover Date.     

Long-term changes in rearing habitat and downstream movement by juvenile sockeye salmon (Oncorhynchus nerka) in an interconnected Alaska lake system

Abstract –  In some populations the phenomenon of partial migration develops where some individuals stay in a given habitat rather than move with the migratory component. Depending on the selective pressures, the individuals that stay may be larger, smaller or similar in size to those that move. Freshwater movements of juvenile sockeye salmon ( Oncorhynchus nerka Walbaum) fry vary among and within populations, and can be complex, especially in interconnected lake systems. We examined variation of movement patterns by a sockeye salmon population in an interconnected lake system during a period of rapid natural habitat change and found that fry migrating downstream were shorter, had lower body condition, and were more likely ill and moribund compared with fish remaining in the lake. However, otolith microstructure measurements indicated that emigrants did not grow significantly slower than residents prior to downstream movement. We show that patterns (i.e., demography of migrants, timing of movement) of downstream movement have changed since the 1970s, corresponding to changes in rearing habitat. Our findings parallel the results with other salmonid species and are generally consistent with the paradigm that density-dependent interactions from declining habitat availability or quality result in the downstream movement of competitively inferior individuals, although the mechanisms governing downstream migration are unclear in this system.     

Site fidelity of spawning sockeye salmon (Oncorhynchus nerka W.) in the presence and absence of olfactory cues

Abstract –  We examined the site fidelity of spawning adult sockeye salmon ( Oncorhynchus nerka ) by tagging and releasing fish in the same stream reach (controls) and displacing them among different but nearby sites ( c . 50 m away). Three sites – two above a stream junction ('upper' reach and 'pond') and one below ('lower' reach) – allowed us to compare the behavior of salmon in the presence and absence of olfactory cues and habitat similarity. Most controls of both sexes (90%) remained in the immediate vicinity of the tagging and release site. When displaced downstream, where the odors of both the upper reach and the pond were detectable, most salmon returned to their former site (65%). Displaced sockeye were more likely to return to the pond from the lower reach than from the upper one ( P  = 0.05), consistent with olfactory orientation and the hypothesis that salmon prefer certain habitats. Salmon displaced from the upper to the lower reach were much more likely to return than those displaced to the pond ( P  < 0.01), consistent with the role of odors in orientation and inconsistent with the habitat choice hypothesis.     

Oxidative stress in juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

Juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum), were held in 8–11°C freshwater, starved for 3 days and subjected to a low‐water stressor to determine the relationship between the general stress response and oxidative stress. Lipid peroxidation (LPO) levels (lipid hydroperoxides) were measured in kidney, liver and brain samples taken at the beginning of the experiment (0‐h unstressed controls) and at 6, 24 and 48 h after application of a continuous low‐water stressor. Tissue samples were also taken at 48 h from fish that had not been exposed to the stressor (48‐h unstressed controls). Exposure to the low‐water stressor affected LPO in kidney and brain tissues. In kidney, LPO decreased 6 h after imposition of the stressor; similar but less pronounced decreases also occurred in the liver and brain. At 48 h, LPO increased (in comparison with 6‐h stressed tissues) in the kidney and brain. In comparison with 48‐h unstressed controls, LPO levels were higher in the kidney and brain of stressed fish. Although preliminary, results suggest that stress can cause oxidative tissue damage in juvenile chinook salmon. Measures of oxidative stress have shown similar responses to stress in mammals; however, further research is needed to determine the extent of the stress–oxidative stress relationship and the underlying physiological mechanisms in fish.     

An investigation of the biological basis of return variability for sockeye salmon (Oncorhynchus nerka) from Great Central and Sproat lakes,Vancouver Island

We tested whether variations in stock characteristics (spawner and smolt abundance) and biotic conditions (prey variability, predation, competition) during the early marine period explained variations in the return of sockeye salmon (Oncorhynchus nerka) to Great Central and Sproat lakes, adjacent lakes on the west coast of Vancouver Island. There are two freshwater age groups in each lake; fish spend 1 or 2 yrs in freshwater after hatching. We tested the influences of stock and biotic factors on the return of each of the two age groups from each of the two lakes. Results of regression analyses showed that prey biomass variability best explained the variation in return for all lake‐age groups. Euphausiid (Thysanoessa spinifera) and cladoceran (Evadne) prey biomass variability explained between 0.75 and 0.95 (adjusted R²) of the variation in return. There appear to be instances of a mismatch between the seasonality of prey productivity and the apparent critical period of feeding for juvenile sockeye.     

山女鳟早期发育及低温耐性观察

将发育积温242℃.d的山女鳟发眼卵置山溪中进行流水孵化。在日均水温4.8℃环境下,发育积温达382℃.d时开始有少数鱼苗孵出。此后在日均2.3~1.5℃,最低0.2℃环境下历时36d,发育积温达456℃.d时全部孵化出膜,孵化率90.2%。最早出膜的孵化稚鱼在日均水温1.5~3.7℃,最低0.2℃环境下历时93d,发育积温达668℃.d时开始上浮。受精后172d发育积温达762℃.d时全部上浮,上浮率77.3%。驯饲一个月的稚鱼育成率63.2%。雏鱼孵化及形态机能发育正常,表明山女鳟在日均水温4.8~1.5℃,最低0.2℃的低温环境中可以正常地进行早期发育。     

A matter of timing: the role of ocean conditions in the initiation of spawning migration by late‐run Fraser River sockeye salmon (Oncorhynchus nerka)

Year 1995 marked the start of a major shift to earlier river entry of late‐run Fraser River sockeye salmon (Oncorhynchus nerka) en route to the spawning grounds. Earlier entry has typically been accompanied by considerably greater in‐river mortality. We examine this behavioral change using correlation analyses between the entry timing of the Adams River and Weaver Creek stocks and the surface marine conditions encountered by the stocks during their homing migration from the northeast Pacific several months earlier. For Adams stocks, maximum correlation is between entry timing and offshore winds, such that the weaker the wind stress in the direction of the prevailing surface currents in early July, the earlier the river entry in late summer. For Weaver stocks, maximum correlation is with salinity, such that the lower the surface salinity along the coast in August, the earlier the river entry. We hypothesize that oceanic changes lead to changes in late‐run sockeye physiology which then leads to changes in behavior. Physiological changes are postulated to arise from two types of preconditioning: Type‐1 occurs in the offshore region, whereby the weaker the prevailing currents that normally hinder eastward migration, the more endogenous energy available for maturation and the earlier the river entry. Type‐2 occurs in coastal regions, whereby the lower the salinity, the more rapid the osmoregulatory adaptation to freshwater (and possible susceptibility to water‐borne pathogens) and the earlier the entry. Results suggest that the earlier entry that began in the mid‐1990s is linked to weaker ocean currents and lower coastal salinities.     

Relationship between dietary lipid source, oxidative stress, and the physiological response to stress in sub-yearling chinook salmon (Oncorhynchus tshawytscha)

Relationships between dietary lipid source, stress, and oxidative stress were examined in juvenile chinook salmon (Oncorhynchus tshawytscha). Four different experimental diets were used: menhaden oil (MHO; elevated 20:5n-3 and 22:6n-3), soybean oil (SBO; elevated 18:2n-6), linseed oil (LSO; elevated 18:3n-3), and a mixture of 55% linseed oil and 45% soybean oil (MIX; approximately equal levels of 18:2n-6 and 18:3n-3). Juvenile salmon (initial body weight of 16.0 g) were fed experimental diets for 12 weeks (early March to early June). At the end of feeding, fish subjected to a low-water stressor for 96 h had greater liver and brain lipid peroxidation compared to unstressed controls; peroxidation was not influenced by diet. Diet and stress affected plasma cortisol levels. Stressed fish fed SBO had the greatest cortisol concentrations, followed by MIX, MHO, and LSO (mean concentrations for the SBO and LSO diets differed significantly). The cortisol response to stress may have been influenced by the ratio of prostaglandin 1- and 2-series to prostaglandin 3-series precursor fatty acids provided by the different diets. The results of this study suggest a connection between the physiological response to stress, dietary lipid quality, and oxidative stress. This is the first evidence of such a relationship in fish. Abbreviations: AA - arachidonic acid; ACTH - adrenocorticotropin; BHT - butylated hydroxytoluene; BLPO - brain lipid peroxidation; dGLA - dihomo-γ-linolenic acid; DHA - docosahexanoic acid; EPA - eicosapentanoic acid; FER - feed efficiency ratio; FOX - ferrous oxidation-xylenol orange; GLA -γ-linolenic acid; LA - linoleic acid; LCO3 - long-chain n-3 polyunsaturated fatty acids; LLPO - liver lipid peroxidation; LN - linolenic acid; LPO - lipid peroxidation; LSO - linseed oil; MHO - menhaden oil; MIX - 55% linseed oil + 45% soybean oil; PC - plasma cortisol; PG - prostaglandin(s); PGE₂- prostaglandin E₂; PUFA - polyunsaturated fatty acid; SBO - soybean oil. This revised version was published online in July 2006 with corrections to the Cover Date.     

Carcass selenium loss as an indicator of stress in barge transported chinook salmon (Oncorhynchus tshawytscha Walbaum) smolts

Two salmon smolt barge transport experiments were conducted to measure tissue trace element selenium (Se) loss during the 30 h transport 500 km downriver past seven hydroelectric dams on the Columbia River. Carcass Se was measured before and after the barge trip. Liver glutathione peroxidase (GTPX) activity and total ascorbate concentrations were assayed to correlate Se loss with GTPX levels. Hatchery‐reared smolt chinook salmon, Oncorhynchus tshawytscha (Walbaum), were tested before and after barging. Liver ascorbate concentrations, measured by high‐performance liquid chromatography techniques, showed levels adequate to support GTPX activity. Salmon smolts lost up to 20% of carcass tissue Se during the 30 h barge confinement. Selenium was analysed by analytical polarography. Samples collected after two barging episodes showed Se loss and concomitant elevated liver GTPX activity after the transport. Tissue Se loss may be useful to measure stress in fish, and increasing tissue Se in hatchery salmon smolts prior to release and transport may be warranted.     

Plasma somatolactin levels in coho salmon (Oncorhynchus kisutch) during smoltification and sexual maturation

Somatolactin (SL) is a novel pituitary hormone recently characterized in several fish species. Structural analyses have shown that SL belongs to the growth hormone/prolactin family, and that it is a highly conserved protein. SL is synthesized by the periodic acid/Schiff-positive cells in the pars intermedia, but has an as yet unidentified function(s). We have recently developed a homologous radioimmunoassay for coho salmon SL and measured plasma levels of SL during two stages of the coho salmon life cycle, smoltification and sexual maturation. During smoltification, plasma levels of SL changed almost in parallel with plasma levels of thyroxine; levels increased as morphological indices of smoltification appeared and decreased as smoltification was completed. Following this period, SL levels remained low until the spring prior to spawning. In a separate study, plasma levels of SL were measured in sexually maturing coho salmon that remained in fresh water throughout their life cycle. During the year of sexual maturation, plasma levels of SL gradually increased from the spring onward, reaching peak levels at the time of spawning in November and December. These data are similar to those previously reported for sexually maturing coho salmon that were maintained in seawater prior to spawning (Rand-Weaver et al. 1992). Therefore, increases in plasma SL levels occurred in sexually maturing fish irrespective of whether they were maintained in fresh water or seawater. Peak levels at spawning were higher than those observed during smoltification. Possible roles for SL in metabolism and reproduction are discussed.     

Effects of temperature on disease progression and swimming stamina in Ichthyophonus-infected rainbow trout, Oncorhynchus mykiss (Walbaum)

Rainbow trout, Oncorhynchus mykiss , were infected with Ichthyophonus sp. and held at 10 °C, 15 °C and 20 °C for 28 days to monitor mortality and disease progression. Infected fish demonstrated more rapid onset of disease, higher parasite load, more severe host tissue reaction and reduced mean-day-to-death at higher temperature. In a second experiment, Ichthyophonus -infected fish were reared at 15 °C for 16 weeks then subjected to forced swimming at 10 °C, 15 °C and 20 °C. Stamina improved significantly with increased temperature in uninfected fish; however, this was not observed for infected fish. The difference in performance between infected and uninfected fish became significant at 15 °C ( P  =   0.02) and highly significant at 20 °C ( P  =   0.005). These results have implications for changes in the ecology of fish diseases in the face of global warming and demonstrate the effects of higher temperature on the progression and severity of ichthyophoniasis as well as on swimming stamina, a critical fitness trait of salmonids. This study helps explain field observations showing the recent emergence of clinical ichthyophoniasis in Yukon River Chinook salmon later in their spawning migration when water temperatures were high, as well as the apparent failure of a substantial percentage of infected fish to successfully reach their natal spawning areas.     

Oxidative stress parameters during starvation and refeeding periods in Adriatic sturgeon (Acipenser naccarii) and rainbow trout (Oncorhynchus mykiss)

This work analyses the changes in the redox balance in two fish species: Adriatic sturgeon ( Acipenser naccarii ) and rainbow trout ( Oncorhynchus mykiss ) during starvation and refeeding period. The starvation period raised the lipid peroxidation (thiobarbituric-acid-reacting substances) levels in liver and blood, while a decline occurred in the antioxidant enzymatic activities of superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPX) and glutathione reductase (GR) in both fish species. In liver, after the refeeding period, SOD activity recovered in both species, whereas CAT activity recovered only in trout. Furthermore, in both tissues of the two species, the lipid peroxidation levels remained high after 2 months of refeeding. In white muscle and heart, the lipid peroxidation levels indicate that these tissues did not undergo oxidative stress during the 72-day period. During starvation, in the muscle of both fish the fall in the lipid peroxidation level coincided with a rise in CAT, GPX and GR. The refeeding period in this tissue raised the lipid peroxidation level, and the enzymatic activities reached the values of the first point of starvation. In heart, no oxidative damage was detected during starvation in either species. The CAT and SOD activities increased during the starvation period only in trout.     

Responses of circulating arginine vasotocin, isotocin, and melatonin to osmotic and disturbance stress in rainbow trout (Oncorhynchus mykiss)

In teleost fish exposed to stressors AVT has been implicated in activation of the hypothalamo-pituitary-interrenal axis. Mel has been shown to counteract several behavioural and endocrine consequences of stress in mammals. These studies were undertaken to investigate the effects of either disturbance or osmotic stress, applied either separately or simultaneously, on plasma AVT, IT and melatonin in rainbow trout (Oncorhynchus mykiss). Hormones were determined in plasma by high-performance liquid chromatography preceded by solid-phase extraction. The results showed that both forms of stress caused significant increase in plasma AVT concentration, although more pronounced elevation was observed in physically disturbed fish. Conversely, neither osmotic nor disturbance stress affected plasma IT concentration. The apparent difference in response to stress by the two close related neurohypophysial nonapeptides suggests independent mechanisms controlling their synthesis and/or release and supports the idea that only AVT plays a role in physiological response to stress. Plasma Mel level was depressed in fish subjected to disturbance stress and to both stresses applied simultaneously, an effect possibly associated with the elevation of plasma AVT concentration. Results are discussed in relation to physiological interactions between hormones.     

Experimental production of gastric dilation and its association with osmoregulatory stress and biogenic amines in chinook salmon, Oncorhynchus tshawytscha (Walbaum)

Chinook salmon smolt in fresh water fed a commercial diet known to produce minimal gastric dilation and air sacculitis (GDAS) were randomly assigned to four experimental tanks with flow-through sea water. All four groups were acclimatized to sea water for 3 weeks and fed a diet of minced fresh seafood. After 3 weeks the groups were fed either; seafood as before, a different commercial pelleted diet associated with the development of GDAS on farms, or either diet supplemented with 500 mg L(-1) putrescine, 300 mg L(-1) cadaverine and 250 mg L(-1) tyramine. Gastric dilation was produced in fish fed the commercial diet for 1 month but not by feeding a diet of minced seafood. The addition of putrescine, cadaverine and tyramine to either diet had no significant effect on the development of gastric dilation. Fish fed the commercial diet had significantly (P < 0.0001) wider weight-adjusted stomach widths, less prominent longitudinal stomach folds (P < 0.0001) and lower (P < 0.0001) stomach-width ratios than fish fed the fresh seafood diet. There was no significant difference in serum osmolality or sodium concentration between fish from groups with or without gastric dilation or fed biogenic amines.     

Plasma growth hormone levels in fed and fasted rainbow trout (Oncorhynchus mykiss) are decreased following handling stress

Plasma growth hormone concentrations of rainbow trout,Oncorhynchus mykiss, fasted for six weeks, were significantly (p < 0.01) higher than in comparable fed animals; in the fasted fish, the levels fell progressively following acute stress (by displacing the fish within their home aquarium), with significant differences from pre-stressed fish evident between one and thirty-two hours after application of the stressor. Plasma growth hormone concentrations also fell significantly in the fed group, but differences were evident only between two and eight hours after stressor application.Plasma cortisol concentrations in pre- and post-stressed fed and fasted fish were similar. There was a bimodal response to stressor application in both groups, with significantly higher values relative to the pre-stressed sample evident one and eight hours after disturbance, but not after two, four or thirty-two hours. The changes in plasma cortisol levels between the initial (09:00h) sample and the sample taken eight hours later resembles the diet pattern seen in trout given access to self-demand feeders.Plasma glucose concentrations in pre-stressed fed animals were higher than in pre-stressed fasted fish. This relationship was also evident between one and four hours and thirty-two hours after stressor application. The post-stress rise in plasma glucose concentration was evident between one and four hours in the fed group, and between four and eight hours in fasted fish.The diel changes in plasma growth hormone and glucose concentrations could not be attributed to normal circadian patterns, and there was no apparent correlation between changes in plasma growth hormone and cortisol concentrations. There was a significant inverse correlation between plasma glucose and growth hormone concentrations when the total data set were analyzed, but these correlations were not apparent when the treatment groups were analyzed separately.     

The ability of clove oil and MS-222 to minimize handling stress in rainbow trout (Oncorhynchus mykiss Walbaum)

The effects of 60‐mg L^?1 clove oil and 60‐mg L^?1 tricaine methanesulphonate (MS‐222) on the blood chemistry of rainbow trout were compared after exposure to handling stress via caudal puncture blood sampling. Fish sampled by caudal puncture and subsequently exposed to anaesthetics showed a typical handling stress response over a 48‐h period. There were no significant differences between the responses of fish exposed to equal concentrations of clove oil and MS‐222, with the following exceptions: the blood glucose at full anaesthesia, and lactate at full recovery increased significantly in the clove oil‐exposed fish. In a subsequent experiment, the stress response observed in fish sampled by caudal puncture and exposed to clove oil and MS‐222 was compared with a non‐anaesthetized control group. The increases in plasma cortisol levels were significantly lower at recovery in fish treated with either anaesthetic compared with the control fish. Fish exposed to MS‐222 had significantly higher cortisol levels at 1 h. These findings show that few differences exist between the anaesthetic effects of clove oil and MS‐222 on the physiological response of fish to stress. However, clove oil is more effective at reducing the short‐term stress response induced by handling and blood sampling, and is recommended as an alternative fish anaesthetic.