Assessment of spawning site fidelity in interior Fraser River Coho salmon Oncorhynchus kisutch using otolith microchemistry, in British Columbia, Canada.

Coho Salmon Oncorhynchus kisutch show fidelity to natal spawning watersheds. Fine-scale homing, however, within rivers is not well understood. Interior Fraser Coho (IFC) salmon eggs were incubated at known spawning locations in the Coldwater River, two main stem sites and one-off channel pond site, providing otolith reference data for comparison to otolith signatures for returning adults using laser ablation inductively coupled plasma mass spectrometry. Elemental ratios for Ba:Ca and Sr:Ca in otoliths of juvenile O. kisutch differed significantly among the spawning locations examined. Juvenile otolith data were used to conduct a linear discriminant analysis to assess fine-scale homing in adults. Juvenile data were all assigned to the location where they had been incubated, producing a robust data set used to compare adult otoliths and define natal locations based on elemental signatures in otoliths of adult spawners. Homing and straying were apparent at the reach level; 57.1% of adults returned to their natal spawning locations, while 42.9% strayed to other spawning sites within the Coldwater River. Straying to novel incubation sites at the reach scale demonstrated plasticity in homing within a watershed.

Behavioural changes during the parr-smolt transformation in coho salmon Oncorhynchus kisutch: is it better to be cool?

Behavioural changes that occur during the parr-smolt transformation were investigated in juvenile coho salmon Oncorhynchus kisutch. Fish from two populations were examined from the Fraser River catchment in British Columbia, Canada; a short and a long-distance migrating population. Fish showed a significant decrease in condition factor and significant increase in gill Na+ K+ -ATPase activity during the spring indicating that they became competent smolts, but no difference between populations. Temperature preference trials were conducted using a shuttlebox system throughout the spring. Mean temperature preference did not differ between the two populations, but preferred temperature decreased with development from 16.5 ± 0.3°C for parr to 15.5 ± 0.4°C for smolts. Mean swimming velocity was also greater in smolts than parr, but there was no difference between the two populations. The preference for warmer water temperature observed for parr in early spring may be advantageous for stimulating smolt development. Preference for slightly cooler temperatures observed for smolts would sustain elevated seawater tolerance during the smolt window by a short time and may ensure successful transition to the marine environment.

Is the duration of the smolt window related to migration distance in coho salmon Oncorhynchus kisutch?

Physiological changes during the parr-smolt transformation were investigated in short distance (Chilliwack River) and long-distance (Salmon River) migrating coho salmon Oncorhynchus kisutch populations in British Columbia, Canada. Biochemical and molecular indicators were used to monitor smolt development for fish reared at 10 °C throughout the spring. Fish grew well and developed the physical appearance of competent smolts. Both populations exhibited increases in gill Na+ -K+ -ATPase activity (NKA; an important indicator of seawater tolerance) at the same date and the duration of the increase in enzyme activity did not differ between populations. Gill messenger (m)RNA copies for two isoforms of the NKA α subunit, α1a and α1b, showed significant changes and the pattern was similar between populations. Growth hormone receptor and prolactin receptor mRNA from the gill showed modest changes associated with smolting in the spring for both populations, suggesting that these may not be useful indicators of smolt development in hatchery-reared O. kisutch. Consequently, the duration of the smolt window was not based on the region of origin in the present study.

Temporal and spatial differences in smolting among Oncorhynchus nerka populations throughout fresh and seawater migration.

Physiological changes that occur in the spring are preparatory for salmonid smolts to successfully enter seawater, but variation is likely to exist within species with a wide geographic distribution. Whether differences in development of seawater tolerance exist among populations that differ in distance to the ocean, temporally during the spring, or as fish migrate downstream is not known. Juvenile sockeye salmon Oncorhynchus nerka from four regions in the Fraser River catchment, British Columbia, were intercepted to assess physiological differences among populations and at different times during migration to characterize the parr-smolt transformation. Pre-migratory fish had low levels of gill Na+ -K+ -ATPase (NKA) activity. High gill NKA activities were observed at the start of migration for some populations, but smolts leaving the lake did not consistently have higher gill NKA activity than non-migratory juvenile O. nerka sampled in their natal lakes. Gill NKA activity was highly variable at the start of migration with no relationship with distance from the ocean. Gill NKA activity changes with migration were also highly variable, but consistently smolts in the ocean had the highest enzyme activities. Internal and external factors may influence this variation, but the dynamic nature of smolting was not based on the region of origin, timing during migration or on the year of migration.

Spatial and environmental effects on Coho Salmon life history trait variation.

Adult size, egg size, fecundity, and mass of gonads are affected by trade-offs between reproductive investment and environmental conditions shaping the evolution of life history traits among populations for widely distributed species. Coho salmon Oncorhynchus kisutch have a large geographic distribution, and different environmental conditions are experienced by populations throughout their range. We examined the effect of environmental variables on female size, egg size, fecundity, and reproductive investment of populations of Coho Salmon from across British Columbia using an information theoretic approach. Female size increased with latitude and decreased with migration distance from the ocean to spawning locations. Egg size was lowest for intermediate intragravel temperature during incubation, decreased with migration distance, but increased in rivers below lakes. Fecundity increased with latitude, warmer temperature during the spawning period, and river size, but decreased in rivers below lakes compared with rivers with tributary sources. Relative gonad size increased with latitude and decreased with migration distance. Latitude of spawning grounds, migratory distance, and temperatures experienced by a population, but also hydrologic features-river size and headwater source-are influential in shaping patterns of reproductive investment, particularly egg size. Although, relative gonad size varied with latitude and migration distance, how gonadal mass was partitioned gives insight into the trade-off between egg size and fecundity. The lack of an effect of latitude on egg size suggests that local optima for egg size related to intragravel temperature may drive the variation in fecundity observed among years.

Are we missing a mineralocorticoid in teleost fish? Effects of cortisol, deoxycorticosterone and aldosterone on osmoregulation, gill Na+,K+ -ATPase activity and isoform mRNA levels in Atlantic salmon.

It has long been held that cortisol, acting through a single receptor, carries out both glucocorticoid and mineralocorticoid actions in teleost fish. The recent finding that fish express a gene with high sequence similarity to the mammalian mineralocorticoid receptor (MR) suggests the possibility that a hormone other than cortisol carries out some mineralocorticoid functions in fish. To test for this possibility, we examined the effect of in vivo cortisol, 11-deoxycorticosterone (DOC) and aldosterone on salinity tolerance, gill Na(+),K(+)-ATPase (NKA) activity and mRNA levels of NKA alpha 1a and alpha 1b in Atlantic salmon. Cortisol treatment for 6-14 days resulted in increased, physiological levels of cortisol, increased gill NKA activity and improved salinity tolerance (lower plasma chloride after a 24h seawater challenge), whereas DOC and aldosterone had no effect on either NKA activity or salinity tolerance. NKA alpha 1a and alpha 1b mRNA levels, which increase in response to fresh water and seawater acclimation, respectively, were both upregulated by cortisol, whereas DOC and aldosterone were without effect. Cortisol, DOC and aldosterone had no effect on gill glucocorticoid receptor GR1, GR2 and MR mRNA levels, although there was some indication of possible upregulation of GR1 by cortisol (p=0.07). The putative GR blocker RU486 inhibited cortisol-induced increases in salinity tolerance, NKA activity and NKA alpha 1a and alpha 1b transcription, whereas the putative MR blocker spironolactone had no effect. The results provide support that cortisol, and not DOC or aldosterone, is involved in regulating the mineralocorticoid functions of ion uptake and salt secretion in teleost fish.

Mechanistic basis of individual mortality in Pacific salmon during spawning migrations.

Reproductive-based migration is a challenging period for many animals, but particularly for Pacific salmonids, which must navigate from the high seas to freshwater natal streams. For the first time, we attempt to answer the question as to why some migratory adult Pacific salmon die en route to spawning grounds. Summer-run sockeye salmon (Oncorhynchus nerka) were used as a model, and the migration behavior of 301 fish was followed by intercepting them in the ocean about 215 km from the mouth of the Fraser River, British Columbia, Canada, and implanting a gastric radio transmitter. Before release, telemetered fish were also bio-sampled, which included drawing a blood sample, collecting a gill biopsy, and quantifying energetic status with a microwave energy meter. We tested the predictions that the fish that died prematurely would be characterized by low energy reserves, advanced reproductive development, elevated indicators of stress, and low osmoregulatory preparedness compared with fish that completed their river migration. Just over half (52.3%) of the sockeye tagged were subsequently detected in the Fraser River. Salmon that failed to enter the river had exhibited indicators of stress (e.g., elevated plasma lactate, glucose, and cortisol). Contrary to our prediction, fish that failed to enter the river tended to have higher gross somatic energy and be larger at the time of sampling in the ocean than fish that successfully entered the river. Of the fish that were detected in the river (i.e., 134 fish excluding fishery removals), 9.7% did not migrate beyond the lower reaches (approximately 250 km from ocean), and a further 14.2% reached the upper reaches but failed to reach natal sub-watersheds, whereas the remainder (76.1%) reached natal sub-watersheds. Of these, fish unsuccessful in the lower reaches tended to have a high plasma osmolality in the ocean, whereas fish failing in the upper reaches had lower levels of reproductive hormones in the ocean.

Genetic versus rearing-environment effects on phenotype: hatchery and natural rearing effects on hatchery- and wild-born coho salmon.

With the current trends in climate and fisheries, well-designed mitigative strategies for conserving fish stocks may become increasingly necessary. The poor post-release survival of hatchery-reared Pacific salmon indicates that salmon enhancement programs require assessment. The objective of this study was to determine the relative roles that genotype and rearing environment play in the phenotypic expression of young salmon, including their survival, growth, physiology, swimming endurance, predator avoidance and migratory behaviour. Wild- and hatchery-born coho salmon adults (Oncorhynchus kisutch) returning to the Chehalis River in British Columbia, Canada, were crossed to create pure hatchery, pure wild, and hybrid offspring. A proportion of the progeny from each cross was reared in a traditional hatchery environment, whereas the remaining fry were reared naturally in a contained side channel. The resulting phenotypic differences between replicates, between rearing environments, and between cross types were compared. While there were few phenotypic differences noted between genetic groups reared in the same habitat, rearing environment played a significant role in smolt size, survival, swimming endurance, predator avoidance and migratory behaviour. The lack of any observed genetic differences between wild- and hatchery-born salmon may be due to the long-term mixing of these genotypes from hatchery introgression into wild populations, or conversely, due to strong selection in nature--capable of maintaining highly fit genotypes whether or not fish have experienced part of their life history under cultured conditions.

Is fishing selective for physiological and energetic characteristics in migratory adult sockeye salmon?

There is extensive evidence that fishing is often selective for specific phenotypic characteristics, and that selective harvest can thus result in genotypic change. To date, however, there are no studies that evaluate whether fishing is selective for certain physiological or energetic characteristics that may influence fish behaviour and thus vulnerability to capture. Here, adult sockeye salmon (Oncorhynchus nerka) were used as a model to test the null hypothesis that fishing is not selective for specific physiological or energetic traits. Fish were intercepted during their spawning migrations, implanted with a gastric radio transmitter, and biopsied (i.e., non-lethally sampled for blood, gill tissue and quantification of energetic status). In both 2003 and 2006, we tagged and biopsied 301 and 770 sockeye salmon, respectively, in the marine environment en route to their natal river system to spawn. In 2006 an additional 378 individuals were tagged and biopsied in freshwater. We found that 23 (7.6%) of the marine fish tagged in 2003, 78 (10.1%) of the marine fish tagged in 2006 and 57 (15.1%) of the freshwater fish tagged in 2006 were harvested by one of three fisheries sectors that operate in the coastal marine environment and the Fraser River (i.e. commercial, recreational or First Nations fisheries between the site of release and Hell's Gate in the Fraser River, approximately 250 km upriver and 465 km from the ocean tagging site). However, fisheries were not open continually or consistently in different locations and for different fisheries sectors necessitating a paired analytical approach. As such, for statistical analyses we paired individual fish that were harvested with another fish of the same genetic stock that was released on the same date and exhibited similar migration behaviour, except that they successfully evaded capture and reached natal spawning grounds. Using two-tailed Wilcoxon matched pairs signed-rank tests, we revealed that the physiological and energetic characteristics of harvested fish did not differ from those of the successful migrants despite evaluating a number of biochemical (e.g. plasma metabolites, cortisol, plasma ions, gill Na(+)/K(+)-ATPase) and energetic (e.g. gross somatic energy density) variables (P's all >0.10). However, for some analyses we suffered low statistical power and the study design had several shortcomings that could have made detection of differences difficult. We suggest that additional research explore the concept of fishing-induced selection for physiological characteristics because physiology is closely linked to three traits where fisheries-induced selection does occur (i.e. life-history, behaviour and morphology).

Individual variation in migration speed of upriver-migrating sockeye salmon in the Fraser River in relation to their physiological and energetic status at marine approach.

Little research has examined individual variation in migration speeds of Pacific salmon (Oncorhynchus spp.) in natural river systems or attempted to link migratory behavior with physiological and energetic status on a large spatial scale in the wild. As a model, we used three stocks of summer-run sockeye salmon (Oncorhynchus nerka) from the Fraser River watershed, British Columbia, to test the hypothesis that individual variation in migration speed is determined by a combination of environmental factors (i.e., water temperature), intrinsic biological differences (sex and population), and physiological and energetic condition. Before the freshwater portion of the migration, sockeye salmon (Quesnel, Chilcotin, and Nechako stock complexes) were captured in Johnstone Strait ( approximately 215 km from river entry), gastrically implanted with radio transmitters, and sampled for blood, gill tissue, and energetic status before release. Analyses focused solely on individuals that successfully reached natal subwatersheds. Migration speeds were assessed by an extensive radiotelemetry array. Individuals from the stock complex that migrated the longest distance (Nechako) traveled at speeds slower than those of other stock complexes. Females traveled slower than males. An elevated energetic status of fish in the ocean was negatively correlated with migration speed in most river segments. During the transition from the ocean to the river, migration speed was negatively correlated with mean maximum water temperature; however, for the majority of river segments, it was positively correlated with migration speed. Physiological status measured in the ocean did not explain among-individual variability in river migration speeds. Collectively, these findings suggest that there could be extensive variation in migration behavior among individuals, sexes, and populations and that physiological condition in the ocean explained little of this variation relative to in-river environmental conditions and energetic status. Interestingly, individual fish generally retained their rank in swimming speed across different segments, except when transiting a challenging canyon midway during the migration.

Census vs. effective population size in chinook salmon: large- and small-scale environmental perturbation effects.

Population viability has often been assessed by census of reproducing adults. Recently this method has been called into question and estimation of the effective population size (Ne) proposed as a complementary method to determine population health. We examined genetic diversity in five populations of chinook salmon (Oncorhynchus tshawytscha) from the upper Fraser River watershed (British Columbia, Canada) at 11 microsatellite loci over 20 years using DNA extracted from archived scale samples. We tested for changes in genetic diversity, calculated the ratio of the number of alleles to the range in allele size to give the statistic M, calculated Ne from the temporal change in allele frequency, used the maximum likelihood method to calculate effective population size (NeM), calculated the harmonic mean of population size, and compared these statistics to annual census estimates. Over the last two decades population size has increased in all five populations of chinook examined; however, Ne calculated for each population was low (81-691) and decreasing over the time interval measured. Values of NeM were low, but substantially higher than Ne calculated using the temporal method. The calculated values for M were generally low (M < 0.70), indicating recent population reductions for all five populations. Large-scale historic barriers to migration and development activities do not appear to account for the low values of Ne; however, available spawning area is positively correlated with Ne. Both Ne and M estimates indicate that these populations are potentially susceptible to inbreeding effects and may lack the ability to respond adaptively to stochastic events. Our findings question the practice of relying exclusively on census estimates for interpreting population health and show the importance of determining genetic diversity within populations.

Effects of an advanced temperature cycle on smolt development and endocrinology indicate that temperature is not a zeitgeber for smolting in Atlantic salmon.

Atlantic salmon (Salmo salar) juveniles were reared under simulated conditions of normal photoperiod (LDN) or short days (LD 9:15) and ambient temperature (AMB: normal temperature increases in April) or an advanced temperature cycle (ADV: temperature increases in February). Under both photoperiod conditions, the timing of increased and peak levels of gill Na(+),K(+)-ATPase activity were not altered by temperature, although the rate of increase was initially greater under ADV. ADV/LD 9:15 resulted in peak gill Na(+),K(+)-ATPase activity that was half of that seen under normal photoperiod and temperature conditions. Plasma growth hormone (GH) levels increased threefold in late March under ADV/LDN, but not under ADV/LD 9:15, indicating that there is a photoperiod-dependent effect of temperature on levels of this hormone. Plasma insulin-like growth factor I (IGF-I) increased in spring in all groups, with increases occurring significantly earlier in the ADV/LDN group. In each photoperiod condition, the advanced temperature cycle resulted in large decreases in plasma thyroxine (T(4)) levels in March, which subsequently recovered, whereas plasma 3,5,3'-triiodo-L-thyronine (T(3)) levels were not substantially affected by either photoperiod or temperature. There was no consistent pattern of change in plasma cortisol levels. The results do not provide support for the role of temperature as a zeitgeber, but do indicate that temperature has a role in the timing of smolting by affecting the rate of development and interacting with the photoperiod.