The effect of substrate rearing on growth, aerobic scope and physiology of larval white sturgeon Acipenser transmontanus.

The effect of substratum on growth and metabolic rate was assessed in larval white sturgeon Acipenser transmontanus. Yolk-sac larvae (YSL) were reared in bare tanks or tanks with gravel as substratum from hatch until approximately 16 days post hatch (dph). The effect of an artificial substratum was also evaluated on growth alone. Substratum had a significant effect on mass, with larvae reared in gravel and artificial substrata being larger than those reared without substratum. Routine metabolic rates were significantly lower and relative aerobic scope (the difference between maximum and routine metabolic rate) was significantly higher for YSL and feeding larvae (FL) reared in gravel relative to those reared in bare tanks, particularly before fish started feeding exogenously. Furthermore, gravel-reared larvae had higher whole-body glycogen concentrations relative to bare-tank-reared larvae. Routine factorial scope (maximum metabolic rate divided by routine metabolic rate) was relatively low in all treatments (< 1·7) indicating a limited ability to elevate metabolic rate above routine early in development and mass exponents for metabolic rate exceeded 1. Taken together, these data indicate that YSL reared without substratum may divert more of their energy to non-growth related processes impairing growth. This finding underscores the importance of adequate rearing substratum for growth of A. transmontanus and may provide support for habitat restoration and alternative hatchery rearing methods associated with sturgeon conservation.

Freshwater movement patterns by juvenile Pacific salmon Oncorhynchus spp. before they migrate to the ocean: Oh the places you'll go!

Juvenile movement patterns for coho salmon Oncorhynchus kisutch and Chinook salmon Oncorhynchus tshawytscha from two large interior rivers of British Columbia, Canada, were examined. Otoliths from post-spawned fishes were collected on spawning grounds and elemental signatures were determined through transects from sectioned otoliths using laser ablation-inductively coupled plasma-mass spectrometry (LA-ICP-MS). Large variations in otolith elemental signatures were found during the freshwater life stage indicative of movement downstream to rivers and tributaries that differed in elemental signature. This study highlights that downstream movements occur before migration to the ocean during the parr-smolt transformation. Extensive downstream movements of parr appear to be a successful life-history strategy based on variations observed in the otolith elemental signatures of spawners. Movements downstream in parr and the remarkable homing ability of adults also suggest that imprinting to natal streams must occur prior to the parr-smolt transformation.

Routine and active metabolic rates of migrating adult wild sockeye salmon (Oncorhynchus nerka Walbaum) in seawater and freshwater.

We present the first data on the differences in routine and active metabolic rates for sexually maturing migratory adult sockeye salmon (Oncorhynchus nerka) that were intercepted in the ocean and then held in either seawater or freshwater. Routine and active oxygen uptake rates (MO2) were significantly higher (27%-72%) in seawater than in freshwater at all swimming speeds except those approaching critical swimming speed. During a 45-min recovery period, the declining postexercise oxygen uptake remained 58%-73% higher in seawater than in freshwater. When fish performed a second swim test, active metabolic rates again remained 28%-81% higher for fish in seawater except at the critical swimming speed. Despite their differences in metabolic rates, fish in both seawater and freshwater could repeat the swim test and reach a similar maximum oxygen uptake and critical swimming speed as in the first swim test, even without restoring routine metabolic rate between swim tests. Thus, elevated MO2 related to either being in seawater as opposed to freshwater or not being fully recovered from previous exhaustive exercise did not present itself as a metabolic loading that limited either critical swimming performance or maximum MO2. The basis for the difference in metabolic rates of migratory sockeye salmon held in seawater and freshwater is uncertain, but it could include differences in states of nutrition, reproduction, and restlessness, as well as ionic differences. Regardless, this study elucidates some of the metabolic costs involved during the migration of adult salmon from seawater to freshwater, which may have applications for fisheries conservation and management models of energy use.

Physiological and molecular endocrine changes in maturing wild sockeye salmon, Oncorhynchus nerka, during ocean and river migration.

Maturing adult sockeye salmon Oncorhynchus nerka were intercepted while migrating in the ocean and upstream in freshwater over a combined distance of more than 1,300 km to determine physiological and endocrine changes associated with ionoregulation. Sockeye migrating through seawater and freshwater showed consistent declines in gill Na+/K+ -ATPase (NKA) activity, plasma osmolality and plasma chloride concentration. In contrast, plasma sodium concentration became elevated in seawater as fish approached the river mouth and was then restored after sockeye entered the river. Accompanying the movement from seawater to freshwater was a significant increase in mRNA for the NKA α1a subunit in the gill, with little change in the α1b subunit. Potential endocrine signals stimulating the physiological changes during migration were assessed by measuring plasma cortisol and prolactin (Prl) concentrations and quantifying mRNA extracted from the gill for glucocorticoid receptors 1 and 2 (GR1 and GR2), mineralocorticoid receptor (MR), growth hormone 1 receptor (GH1R), and prolactin receptor (PrlR). Plasma cortisol and prolactin concentrations were high in seawater suggesting a preparatory endocrine signal before freshwater entry. Generally, the mRNA expression for GR1, GR2 and MR declined during migration, most notably after fish entered freshwater. In contrast, PrlR mRNA increased throughout migration, particularly as sockeye approached the spawning grounds. A highly significant association existed between gill PrlR mRNA and gill NKA α1a mRNA. GH1R mRNA also increased significantly, but only after sockeye had migrated beyond tidal influence in the river and then again just before the fish reached the spawning grounds. These findings suggest that cortisol and prolactin stimulate ionoregulation in the gill as sockeye salmon adapt to freshwater.

Effects of experimental manipulations of salinity and maturation status on the physiological condition and mortality of homing adult sockeye salmon held in a laboratory.

Relatively little is known about the physiological response and mortality consequences of the return of anadromous fish to freshwater (FW). We explored the consequences of the return to FW by collecting maturing sockeye salmon from the marine waters off the mouth of the Fraser River and holding approximately 50 sockeye in each of five treatments: saltwater (SW; salinity = 28 ppt), iso-osmotic water (ISO; 13 ppt), FW (0 ppt), SW + gonadotropin-releasing hormone (SW + GnRH), and FW + GnRH. Exogenous GnRH treatments were intended to accelerate maturation. Results demonstrate that gill Na(+),K(+) ATPase activity, sex steroid concentrations, and cortisol levels were highly responsive to experimental manipulations and followed predicted trajectories (i.e., FW + GnRH sockeye were the most mature and FW adapted). There were few among-treatment differences in hematocrit and plasma concentrations of lactate, glucose, Na(+), Cl(-), and plasma osmolality among sockeye that survived to the end of treatments, indicating that sockeye rigorously maintain internal homeostatic conditions while alive. There were large among-treatment differences in mortality (SW+GnRH > SW> FW+GnRH > FW=ISO), and each treatment experienced a notable increase in mortality rate around the fifth day of treatment. Our results indicate that salinity represented a modestly larger challenge to the experimental sockeye than did the artificially accelerated sexual maturation. Our results also suggest that maturing sockeye either successfully acclimate to FW within 5 d of exposure or perish. These findings are consistent with the predictions of the theory of anadromy, in suggesting that the return of adults to FW can be physiologically challenging and can represent a period of significant natural mortality.

Physiological condition differentially affects the behavior and survival of two populations of sockeye salmon during their freshwater spawning migration.

Recently, a segment of the Adams-Shuswap sockeye salmon (Oncorhynchus nerka) population initiated freshwater migration several weeks earlier than historically recorded, resulting in high mortality rates. The comigrating Chilko population maintained their historic river entry timing and did not experience elevated mortality. To test the hypothesis that population-specific differences in physiological condition would differentially influence behavior and survival when exposed to fisheries capture stress, we physiologically sampled individuals from both populations at the onset of the freshwater phase of their reproductive migration and tracked the remainder of their migrations using radio telemetry. Adams-Shuswap individuals had slower migration rates and were less likely to reach natal subwatersheds relative to Chilko individuals. Metabolic and osmoregulatory impairment was related to mortality for Adams-Shuswap individuals but not for Chilko individuals. Similarly, physiological condition correlated with migration rate for Adams-Shuswap but not Chilko fish. Survival to natal subwatersheds was 1.9 times higher for Chilko relative to Adams-Shuswap, a result that did not emerge until individuals approached natal subwatersheds several days after the stressor was applied. We conclude that physiological condition differentially affects the behavior and survival of these two populations, which may be a consequence of the early-entry phenomenon by a segment of the Adams-Shuswap population.

Dosage effects on heritability and maternal effects in diploid and triploid Chinook salmon (Oncorhynchus tshawytscha).

Induced triploidy (3N) in salmon results from a blockage of maternal meiosis II, and hence provides a unique opportunity to study dosage effects on phenotypic variance. Chinook salmon families were bred using a paternal half-sib breeding design (62 females and 31 males) and half of each resulting family was treated to induce triploidy. The paired families were used to test for dosage effects (resulting from triploidy) on (1) the distribution and magnitude of phenotypic variation, (2) narrow-sense heritability and (3) maternal effects in fitness-related traits (i.e., survival, size-at-age, relative growth rate and serum lysozyme activity). Quantitative genetic analyses were performed separately for diploid and triploid family groups. Triploidization resulted in significantly higher levels of phenotypic variance and substantial differences in patterns of variance distribution for growth and survival-related traits, although the patterns were reversed for lysozyme activity. Triploids exhibited higher narrow sense heritability values relative to diploid Chinook salmon. However, maternal effects estimates were generally lower in triploids than in diploids. Thus, the dosage effects resulting from adding an extra set of chromosomes to the Chinook salmon genome are primarily additive. Somewhat counterintuitively, however, the relative magnitude of the combined effects of dominance, epistasis and maternal effects is not affected by dosage. Our results indicate that inheritance of fitness-related quantitative traits is profoundly affected by dosage effects associated with induced triploidy, and that triploidization can result in unpredictable performance and fitness outcomes.

Ionoregulatory changes in different populations of maturing sockeye salmon Oncorhynchus nerka during ocean and river migration.

We present the first data on changes in ionoregulatory physiology of maturing, migratory adult sockeye salmon Oncorhynchus nerka. Fraser River sockeye were intercepted in the ocean as far away as the Queen Charlotte Islands (approximately 850 km from the Fraser River) and during freshwater migration to the spawning grounds; for some populations this was a distance of over 700 km. Sockeye migrating in seawater toward the mouth of the Fraser River and upriver to spawning grounds showed a decline in gill Na+,K+-ATPase activity. As a result, gill Na+,K+-ATPase activity of fish arriving at the spawning grounds was significantly lower than values obtained from fish captured before entry into freshwater. Plasma osmolality and chloride levels also showed significant decreases from seawater values during the freshwater migration to spawning areas. Movement from seawater to freshwater increased mRNA expression of a freshwater-specific Na+,K+-ATPase isoform (alpha1a) while having no effect on the seawater-specific isoform (alpha1b). In addition, gill Na+,K+-ATPase activity generally increased in active spawners compared with unspawned fish on the spawning grounds and this was associated with a marked increase in Na+,K+-ATPase alpha1b mRNA. Increases in gill Na+,K+-ATPase activities observed in spawners suggests that the fish may be attempting to compensate for the osmotic perturbation associated with the decline in plasma chloride concentration and osmolality.

Downregulation of corticosteroid receptors in gills of coho salmon due to stress and cortisol treatment.

The effect of stress and cortisol treatment on corticosteroid receptors (CRs) in the gills of coho salmon (Oncorhynchus kisutch) was examined. Plasma cortisol levels were elevated acutely by intraperitoneal injection of cortisol 21-hemisuccinate and chronically by implanting miniosmotic pumps filled with cortisol or by repeated daily handling stresses. CR concentration and affinity were measured by radioreceptor assay employing 3H-labeled triamcinolone acetonide as ligand. Acute administration of cortisol resulted in a reduction in CR numbers for 72 h with no change in affinity. Chronic cortisol treatment resulted in a decrease in CR concentration and affinity. The change in affinity occurred only while plasma cortisol levels remained elevated, but CR population remained significantly reduced for at least 10 days after cessation of hormone treatment. Repeated handling stresses resulted in a similar reduction in CR numbers but without an apparent change in affinity. The chronic or repeated elevation in plasma cortisol downregulates the sensitivity of the gills to cortisol by a persistent reduction in CR concentration, despite the return to nonstress levels of circulating cortisol.

Regulation of gill cytosolic corticosteroid receptors in juvenile Atlantic salmon: interaction effects of growth hormone with prolactin and triiodothyronine.

The potential effects of growth hormone (GH), prolactin (Prl), and triiodothyronine (T3) on gill Na+,K+-ATPase activity and corticosteroid receptor (CR) concentration (Bmax) and dissociation constant (Kd) were examined in juvenile Atlantic salmon (Salmo salar). Compared to controls, fish injected with GH (ovine, 5.0 microgram g-1) had significantly greater gill Na+,K+-ATPase activity after 7 and 14 days. Gill CR Bmax and Kd were significantly elevated on day 7, but not day 14. T3 also significantly increased CR Bmax. The effect of GH on CR Bmax was also additive with T3 (5.0 microgram g-1) treatment. There was a synergistic effect on CR Bmax when purified coho salmon GH (csGH, 0.1 microgram g-1) was injected in combination with T3 (1.6 microgram g-1). Prl (ovine, 5.0 microgram g-1; purified coho salmon, 0.1 microgram g-1) did not significantly alter gill CR Bmax. Although Prl limited the increase in CR Bmax by GH, the effect was not signicant. T3 and Prl did not have an effect on Kd. GH significantly increased gill Na+,K+-ATPase activity, T3 administration did not have a significant effect, and Prl-treated fish had significantly lower gill Na+,K+-ATPase activity. The results indicate that T3 acts additively with GH, while Prl has no effect in regulating CR Bmax. An increase in cytosolic CR by GH and T3, but not Prl, may regulate gill responsiveness to cortisol and be an important mechanism in the endocrine control of physiological changes during the parr-smolt transformation.

Heritability and Y-chromosome influence in the jack male life history of chinook salmon (Oncorhynchus tshawytscha).

Jacking in chinook salmon (Oncorhynchus tshawytscha) is an alternative reproductive strategy in which males sexually mature at least 1 year before other members of their year class. We characterize the genetic component of this reproductive strategy using two approaches; hormonal phenotypic sex manipulation, and a half-sib breeding experiment. We 'masculinized' chinook salmon larvae with testosterone, reared them to first maturation, identified jacks and immature males based on phenotype, and genotyped all fish as male ('XY') or female ('XX') using PCR-based Y-chromosome markers. The XY males had a much higher incidence of jacking than the XX males (30.8% vs 9.9%). There was no difference in body weight, gonad weight, and plasma concentrations of testosterone and 17beta-estradiol between the two jack genotypes, although XY jacks did have a higher gonadosomatic index (GSI) than XX jacks. In the second experiment, we bred chinook salmon in two modified half-sib mating designs, and scored the number of jacks and immature fish at first maturation. Heritability of jacking was estimated using two ANOVA models: dams nested within sires, and sires nested within dams with one-half of the half-sib families common to the two models. The sire component of the additive genetic variance yielded a high heritability estimate and was significantly higher than the dam component (h(2)(sire) = 0.62 +/- 0.21; h(2)(dam) = -0.14 +/- 0.12). Our experiments both indicated a strong sex-linked component (Y-chromosome) to jacking in chinook salmon, although evidence for at least some autosomal contribution was also observed.

The presence of high-affinity, low-capacity estradiol-17beta binding in rainbow trout scale indicates a possible endocrine route for the regulation of scale resorption.

High-affinity, low-capacity estradiol-17beta (E(2)) binding is present in rainbow trout scale. The K(d) and B(max) of the scale E(2) binding are similar to those of the liver E(2) receptor (K(d) is 1.6 +/- 0.1 and 1.4 +/- 0.1 nM, and B(max) is 9.1 +/- 1.2 and 23. 1 +/- 2.2 fmol x mg protein(-1), for scale and liver, respectively), but different from those of the high-affinity, low-capacity E(2) binding in plasma (K(d) is 4.0 +/- 0.4 nM and B(max) is 625.4 +/- 63. 1 fmol x mg protein(-1)). The E(2) binding in scale was displaced by testosterone, but not by diethylstilbestrol. Hence, the ligand binding specificity is different from that of the previously characterized liver E(2) receptor, where E(2) is displaced by diethylstilbestrol, but not by testosterone. The putative scale E(2) receptor thus appears to bind both E(2) and testosterone, and it is proposed that the increased scale resorption observed during sexual maturation in both sexes of several salmonid species may be mediated by this receptor. No high-affinity, low-capacity E(2) binding could be detected in rainbow trout gill or skin.

Increases in gill cytosolic corticosteroid receptor abundance and saltwater tolerance in juvenile coho salmon (Oncorhynchus kisutch) treated with growth hormone and placental lactogen.

Juvenile coho salmon (Oncorhynchus kisutch) were injected with one of two recombinant bovine hormones, growth hormone (bGH; 5.0 and 0.5 micrograms.g-1 body wt) or placental lactogen (bPL; 5.0, 0.5, and 0.05 micrograms.g-1 body wt) to determine the effect on growth, plasma cortisol concentration, cytosolic corticosteroid receptors (CR) in the gills, and the development of hypoosmoregulatory ability. One week following a single injection or six weekly injections of bGH or bPL, the fish were measured and sampled for CR concentration and Na+,K(+)-ATPase activity in the gills. Fish were also challenged with salt water (salinity 25%) for 24 hr to determine saltwater tolerance at the end of the 6-week treatment. Treatment with bPL and bGH significantly increased weight and length of the fish. The 0.05-micrograms bPL dose significantly elevated plasma cortisol concentration, whereas all other hormone treatments did not affect cortisol levels. bPL and bGH also significantly increased CR concentration and Na+,K(+)-ATPase activity in the gills. The perturbation in plasma sodium concentration was least in animals receiving the highest dose of bPL and the bGH-treated animals following transfer to seawater. An increase in cytosolic CR by bGH and bPL may increase responsiveness of the gills to cortisol and partially account for the increase in Na+,K(+)-ATPase activity and greater ability to regulate plasma sodium in seawater as exhibited by the experimental groups.