Risk-taking behavior in weight-compensating coho salmon, Oncorhynchus kisutch

The effects of food restriction on predation risk-taking behaviorwere studied in juvenile coho salmon, Oncorhyndtus kisutch,during the period before seawater migration in the spring. Aradiographic technique enabled the study of individual foodintake in a safe and risky food patch before and after exposureto a piscivorous predator. The study revealed a significantincrease in food intake and specific growth rate in the weeksfollowing deprivation, resulting in compensatory growth anda recovery of the weight loss caused by dietary restriction.The increase in food intake resulted from a change in risk-takingbehavior. Following a period of diet restriction, the fish habituatedfalter after predator exposure, and the proportion of fish inthe risky patch was significantly higher than before deprivation.Deprived fish took greater shares of the group meal comparedwith control fish, and the pattern of the individual share ofthe group meal after food restriction indicated a change inthe social hierarchy. This study indicates that risk-takingbehavior is state dependent and changes temporarily to compensatefor a period of food restriction. The results are discussedin terms of tradeoffs between food and risk in a period leadingup to an ontogenetic habitat shift.     

Behaviour of growth hormone transgenic coho salmon Oncorhynchus kisutch in marine mesocosms assessed by acoustic tag telemetry

Underwater acoustic tag telemetry was used to assess behavioural differences between juvenile wild‐type (i.e. non‐transgenic, NT) and growth hormone (GH) transgenic (T) coho salmon Oncorhynchus kisutch in a contained simulated ocean environment. T O. kisutch were found across days to maintain higher baseline swimming speeds than NT O. kisutch and differences in response to feeding were detected between T and NT genotypes. This is the first study to assess behaviour of GH transgenic salmonids in a marine environment and has relevance for assessing whether behavioural effects of GH overexpression seen in freshwater environments can be extrapolated to oceanic phases of the life cycle.     

Oxygen uptake of growth hormone transgenic coho salmon during starvation and feeding

Oxygen uptake of growth hormone transgenic coho salmon Oncorhynchus kisutch was measured in individual fish with a closed-system respirometer and was compared with that of similar-sized non-transgenic control coho salmon during starvation and when fed a fixed ration or to satiation. Transgenic and control fish did not differ in their standard oxygen uptake after 4 days of starvation, although control fish had a higher routine oxygen uptake, scope for spontaneous activity and initial acclimation oxygen uptake. During feeding, transgenic fish ate significantly more than control fish, and had an overall oxygen uptake that was 1·7 times greater than control fish. When fish that had eaten the same per cent body mass were compared, transgenic fish had an oxygen uptake that was 1·4 times greater than control fish. Differences in oxygen uptake in growth hormone transgenic coho salmon and non-transgenic fish appear to be due to the effects of feeding, acclimation and activity level, and not to a difference in basal metabolism.     

Disease resistance and health parameters of growth-hormone transgenic and wild-type coho salmon,Oncorhynchus kisutch

To extend previous findings regarding fish health and disease susceptibility of growth-enhanced fish, hematological and immunological parameters have been compared between growth hormone (GH) transgenic and wild-type non-transgenic coho salmon (Oncorhynchus kisutch). Compared to non-transgenic coho salmon, transgenic fish had significantly higher hematocrit (Hct), hemoglobin (Hb), mean cellular hemoglobin (MCH), mean cellular volume (MCV), and erythrocyte numbers, and lower white cell numbers. In addition, resistance to the bacterial pathogen Aeromonas salmonicida (causal agent of furunculosis) has been assessed between the strains. Higher susceptibility of transgenic fish to this disease challenge was observed in two separate year classes of fish. The present findings provide fundamental knowledge of the disease resistance on GH enhanced transgenic coho salmon, which is of importance for assessing the fitness of transgenic strains for environmental risk assessments, and for improving our understanding effects of growth modification on basic immune functions.     

Intestinal morphology in growth hormone transgenic coho salmon

In two GH transgenic coho salmon Oncorhynchus kisutch , the surface area of the intestine was 2·2 times that of control salmon and the growth rate was about twice that of controls. It seems likely that the enhanced intestinal surface area is a compensatory feature that is manifested in fast growing salmonids.     

Disease resistance, stress response and effects of triploidy in growth hormone transgenic coho salmon

Diploid and triploid coho salmon Oncorhynchus kisutch transgenic for growth hormone (GH) and control coho salmon were compared for differences in disease resistance and stress response. Resistance to the bacterial pathogen Vibrio anguillarum was not affected in transgenic fish relative to their non‐transgenic counterparts when they were infected at the fry stage, but was lower in transgenic fish when infected near smolting. Vaccination against vibriosis provided equal protection to both transgenic and non‐transgenic fish. Triploid fish showed a lower resistance to vibriosis than their diploid counterparts. Diploid transgenic fish and non‐transgenic fish appeared to show similar physiological and cellular stress responses to a heat shock. These studies provide information useful for both performance and ecological risk assessments of growth‐accelerated coho salmon.     

The blood vasculature of the gastrointestinal tract in chinook, Oncorhynchus tshawytscha (Walbaum), and coho, O. kisutch (Walbaum), salmon

The venous and arterial vasculature of the chinook and coho salmon gastrointestinal tract were examined using corrosion casts and India ink injection techniques. Observations derived from 28 individuals of various sizes and of both sexes were used to construct simplified venous and arterial plans. Examination of the blood vasculature revealed the presence of a variety of anastomoses hitherto undescribed in teleosts.     

Interaction of growth hormone overexpression and nutritional status on pituitary gland clock gene expression in coho salmon,Oncorhynchus kisutch

Clock genes are involved in generating a circadian rhythm that is integrated with the metabolic state of an organism and information from the environment. Growth hormone (GH) transgenic coho salmon, Oncorhynchus kisutch, show a large increase in growth rate, but also attenuated seasonal growth modulations, modified timing of physiological transformations (e.g. smoltification) and disruptions in pituitary gene expression compared with wild-type salmon. In several fishes, circadian rhythm gene expression has been found to oscillate in the suprachiasmatic nucleus of the hypothalamus, as well as in multiple peripheral tissues, but this control system has not been examined in the pituitary gland nor has the effect of transgenic growth modification been examined. Thus, the daily expression of 10 core clock genes has been examined in pituitary glands of GH transgenic (T) and wild-type coho salmon (NT) entrained on a regular photocycle (12L: 12D) and provided either with scheduled feeding or had food withheld for 60?h. Most clock genes in both genotypes showed oscillating patterns of mRNA levels with light and dark cycles. However, T showed different amplitudes and patterns of expression compared with wild salmon, both in fed and starved conditions. The results from this study indicate that constitutive expression of GH is associated with changes in clock gene regulation, which may play a role in the disrupted behavioural and physiological phenotypes observed in growth-modified transgenic strains.     

Effects of temperature and growth hormone on individual growth trajectories of wild‐type and transgenic coho salmon Oncorhynchus kisutch

In this study, individual growth patterns of wild‐type and growth‐enhanced coho salmon Oncorhynchus kisutch at 8, 12 and 16° C water temperature were followed. Despite large differences among individuals in growth rates, there was generally little variation in the shape of the growth curves among O. kisutch individuals of both genotypes and at all temperatures. Typically, individuals that were relatively large initially were also relatively large at the end of the growth period. The limitation in variation was more pronounced in the growth‐enhanced O. kisutch than in the wild type, where the relative size of some individuals reared at 12 and 8° C changed by the end of the trial. As a warmer temperature seems to decrease the plasticity of growth trajectories in wild‐type fish, it is possible that global warming will influence the ability of wild fish to adapt their growth to changing conditions.     

The glutathione antioxidant system is enhanced in growth hormone transgenic coho salmon (<Emphasis Type="Italic">Oncorhynchus kisutch</Emphasis>)

Insertion of a growth hormone (GH) transgene in coho salmon results in accelerated growth, and increased feeding and metabolic rates. Whether other physiological systems within the fish are adjusted to this accelerated growth has not been well explored. We examined the effects of a GH transgene and feeding level on the antioxidant glutathione and its associated enzymes in various tissues of coho salmon. When transgenic and control salmon were fed to satiation, transgenic fish had increased tissue glutathione, increased hepatic glutathione reductase activity, decreased hepatic activity of the glutathione synthesis enzyme γ-glutamylcysteine synthetase, and increased intestinal activity of the glutathione catabolic enzyme γ-glutamyltranspeptidase. However, these differences were mostly abolished by ration restriction and fasting, indicating that upregulation of the glutathione antioxidant system was due to accelerated growth, and not to intrinsic effects of the transgene. Increased food intake and ability to digest potential dietary glutathione, and not increased activity of glutathione synthesis enzymes, likely contributed to the higher levels of glutathione in transgenic fish. Components of the glutathione antioxidant system are likely upregulated to combat potentially higher reactive oxygen species production from increased metabolic rates in GH transgenic salmon.     

Transient serotonin-immunoreactive neurons coincide with a critical period of neural development in coho salmon (Oncorhynchus kisutch)

Summary In coho salmon (Oncorhynchus kisutch), smolt transformation has been shown to be associated with sequential surges of neurotransmitters in the brain. In order to determine if the surge of serotonin (5-HT) is correlated with structural changes, we have used immunocytochemistry to observe changes in the serotonin immunoreactivity before, during and after the 5-HT surge. The following stages were studied: 12-month-old freshwater presmolts, 17-month-old freshwater presmolts, 18-month-old saltwater smolts, 19-month-old saltwater postsmolt, 24-month-old postsmolt, and adult spawners. In the 17-month-old samples, but not at any other stage, we found a set of transient (serotonin-immunoreactive) 5-HT-immunoreactive neurons in the lateral preoptic area, as well as a discrete population of 5-HT-immunoreactive neurons in the lateral part of the dorsal right habenular nucleus. In addition, a higher density of serotonergic fibers was found in the telencephalon at this stage compared to the following two stages. Since the transient 5-HT-immunoreactive structures presented here do not appear simultaneously with the 5-HT total brain concentration surge, we conclude that they are unlikely to be the source of the 5-HT surge, but are probably related to other developmental changes in the brain associated with smolt transformation.     

The energetic equivalence of cover to juvenile coho salmon (Oncorhynchus kisutch): ideal free distribution theroy applied

Cover is often thought to be an important habitat characteristicfor juvenile stream salmonida. In addition to providing protectionfrom predators, cover may be associated with reduced food availability.Thus, an individual's use of cover is likely to reflect a trade-offbetween the conflicting demands of growth and survival. We measuredthe influence of cover on foraging-site selection in groupsof eight juvenile coho salmon (Oncorhynchus kisutch) by examiningtheir distribution across two stream channel patches, one providingaccess to cover but little food (the "poor" patch), the otherproviding more food but no cover (the "good" patch). Becausefish distributions in the absence of cover conformed to an idealfree distribution (IFD) for unequal competitors (i.e., the distributionof competitive abilities matched the distribution of food),we used IFD theory to quantify the energetic equivalence ofcover to the fish. In the presence of cover and a model avianpredator, use of the poor patch increased relative to the predictionsof the IFD model. Using this observed deviation from an IFD,we calculated how much extra food must be added to the goodpatch to return the distribution of fish to the previously observedIFD of unequal competitors. As predicted, adding this amountof food caused the fish to return to their previous distribution,demonstrating that IFD theory can be used to relate energy intakeand risk of predation in a common currency     

Selection on increased intrinsic growth rates in coho salmon, Oncorhynchus kisutch

Substantial evidence from the animal kingdom shows that there is a trade-off between benefits and costs associated with rapid somatic growth. One would therefore expect growth rates under natural conditions to be close to an evolutionary optimum. Nevertheless, natural selection in many salmonid species appears to be toward larger size and earlier emergence from spawning redds, indicating a potential for increased growth rate to evolve. We tested how selection for genetic variants (growth hormone transgenic coho salmon, Oncorhynchus kisutch, with more than doubled daily growth rate potential relative to wild genotypes) depended on predator timing and food abundance during the early period of life (fry stage). In artificial redds, fry of the fast-growing genotypes showed a highly significant developmental shift, emerging from gravel nests approximately two weeks sooner, but with an 18.6% reduced survival, relative to wild-genotype fry. In seminatural streams, fry of the fast-growing genotypes suffered higher predation than those of wild genotypes when predators were present at the time of fry emergence, but this difference was less pronounced when food was scarce. In streams where predators were introduced after emergence, fry survived equally well regardless of food availability. Surviving fry grew faster in habitats provided with more food, and fast-growing genotypes also grew faster than wild genotypes when predators arrived late and food was abundant. Fewer fish migrated downstream past a waterfall when food availability was high and in the presence of predators, and wild-genotype fry were more likely to migrate than fry of the fast-growing genotypes. After being returned to the experimental streams after migration, fast-growing genotypes survived equally well as those of the same genotypes that did not migrate, whereas migrating wild genotypes experienced higher mortality relative to those of the same genotypes that did not migrate. Comparisons of growth rates between siblings retained under hatchery conditions and those from habitats with the fastest growth in the experimental stream revealed that growth rates were similar for wild genotypes in both environments, whereas the fast-growing genotypes in the streams only realized 90% of their growth potential. The present study has shown that a major shift in developmental timing can alter critical early stages affecting survival and can have a significant effect on fitness. Furthermore, ecological conditions such as food abundance and predation pressure can strongly influence the potential for fast-growing variants to survive under natural conditions. The large-scale removal of many predatory species around the world may augment the evolution of increased intrinsic growth rates in some taxa.     

Genetic assessment of the effects of streamscape succession on coho salmon Oncorhynchus kisutch colonization in recently deglaciated streams

Measures of genetic diversity within and among populations and historical geomorphological data on stream landscapes were used in model simulations based on approximate Bayesian computation (ABC) to examine hypotheses of the relative importance of stream features (geomorphology and age) associated with colonization events and gene flow for coho salmon Oncorhynchus kisutch breeding in recently deglaciated streams (50–240 years b.p .) in Glacier Bay National Park (GBNP), Alaska. Population estimates of genetic diversity including heterozygosity and allelic richness declined significantly and monotonically from the oldest and largest to youngest and smallest GBNP streams. Interpopulation variance in allele frequency increased with increasing distance between streams (r = 0·435, P < 0·01) and was inversely related to stream age (r = –0·281, P < 0·01). The most supported model of colonization involved ongoing or recent (<10 generations before sampling) colonization originating from large populations outside Glacier Bay proper into all other GBNP streams sampled. Results here show that sustained gene flow from large source populations is important to recently established O. kisutch metapopulations. Studies that document how genetic and demographic characteristics of newly founded populations vary associated with successional changes in stream habitat are of particular importance to and have significant implications for, restoration of declining or repatriation of extirpated populations in other regions of the species' native range.     

A histological description of gonadal development and sex differentiation in the coho salmon (Oncorhynchus kisutch) for both untreated and oestradiol immersed fry

Gonadal development is described in detail for coho salmon ( Oncorhynchus kisutch ) between hatching and 1000 degree-days, post-hatch, to aid sex reversal research. Germ cell morphology and sequence, vascular and reproductive duct development, and gross characteristics of the gonads are presented. Sex differentiation occurs by 380 degree-days, post-hatch (800 degree-days, post-fertilization) and is direct to male and female. Oocytes proliferate rapidly after differentiation while the testes enter a period of quiescence. Fry immersed for short durations in oestradiol (200 μg ⁻¹) are also examined. Hormone immersion advanced sex differentiation by 70 degree-days. The immersions were applied early, at 20 and 90 degree-days, post-hatch, yet still altered the sex ratio and timing of differentiation. Definitive germ cells, which are abundant during this period, may be the type most receptive to steroid treatment.     

Discovery and characterization of single nucleotide polymorphisms in coho salmon,Oncorhynchus kisutch

Molecular population genetic analyses have become an integral part of ecological investigation and population monitoring for conservation and management. Microsatellites have been the molecular marker of choice for such applications over the last several decades, but single nucleotide polymorphism (SNP) markers are rapidly expanding beyond model organisms. Coho salmon (Oncorhynchus kisutch) is native to the north Pacific Ocean and its tributaries, where it is the focus of intensive fishery and conservation activities. As it is an anadromous species, coho salmon typically migrate across multiple jurisdictional boundaries, complicating management and requiring shared data collection methods. Here, we describe the discovery and validation of a suite of novel SNPs and associated genotyping assays which can be used in the genetic analyses of this species. These assays include 91 that are polymorphic in the species and one that discriminates it from a sister species, Chinook salmon. We demonstrate the utility of these SNPs for population assignment and phylogeographic analyses, and map them against the draft trout genome. The markers constitute a large majority of all SNP markers described for coho salmon and will enable both population‐ and pedigree‐based analyses across the southern part of the species native range.     

Characterization of 19 single nucleotide polymorphism markers for coho salmon

We report 39 single nucleotide polymorphisms (SNPs) observed in 23 nuclear DNA sequences in coho salmon Oncorhynchus kisutch. High‐throughput genotyping assays based on the 5′‐nuclease reaction were developed for 17 of these nuclear SNPs and for two previously published mitochondrial DNA SNPs. Minor allele frequency differences (Δq) among collections were between 5.2% and 51.2%, resulting in per locus F_ST estimates of 0.00–0.24 with an average of 0.09.     

Riverine, estuarine and marine migratory behaviour and physiology of wild and hatchery-reared coho salmon Oncorhynchus kisutch (Walbaum) smolts descending the Campbell River, BC, Canada

Eighty coho salmon Oncorhynchus kisutch smolts (40 wild and 40 hatchery-reared) were surgically implanted with acoustic transmitters and released into the Quinsam River over 2 days. Differences in physiology, travel time and migratory behaviour were examined between wild and hatchery-reared fish. In addition, tagged and control fish of both wild and hatchery-reared stock were raised for 3 months following surgery to compare survival and tag retention. Detection ranges of the acoustic receivers were tested in the river, estuary and ocean in a variety of flow conditions and tide levels. Receivers were placed in the river, estuary and up to 50 km north and south from the river mouth in the marine environment. Wild smolts were significantly smaller by mass, fork length and condition factor than hatchery-reared smolts and exhibited significantly higher levels of sodium, potassium and chloride in their blood plasma than hatchery-reared smolts. The gill Na⁺K⁺-ATPase activity was also significantly higher in the wild coho smolts at the time of release. Ninety-eight per cent of wild and 80% of hatchery-reared fish survived to the estuary, 8 km downstream of the release site. No difference was found in migration speed, timing or survival between smolts released during daylight and those released after dark. Wild smolts, however, spent less time in the river and estuary, and as a result entered the ocean earlier than hatchery-reared smolts. Average marine swimming speeds for wild smolts were double those of their hatchery-reared counterparts. While hatchery smolts dispersed in both a northward and southward direction upon entering the marine environment, the majority of wild smolts travelled north from the Campbell River estuary. The wild coho salmon smolts were more physiologically fit and ready to enter sea water than the hatchery-reared smolts, and as a result had higher early survival rates and swimming speeds.     

Gonadotropin release following oral delivery of luteinizing hormone-releasing hormone and its superactive analogue (des-Gly10 [D-Ala6] LHRH ethylamide) to 17β-oestradiol-primed coho salmon, Oncorhynchus kisutch (Walbaum)

The permeability of the coho salmon, Oncorhynchus kisutch (Walbaum), gut to orally administered LHRH and LHRHa (des-Gly¹⁰ [D-Ala⁶] LHRH ethylamide) was examined over a 360 min time-course. Both forms of LHRH were detected in the blood plasma of treated fish within 30 min of delivery. Retained biological activity of the absorbed LHRH was also assessed. Oral delivery of LHRH or LHRHa to 17β-oestradiol-primed coho resulted in a significant ( P < 0.01) release of gonadotropin (GtH) when compared to saline intubated, 17β-oestradiol-primed animals. The dose response of 17β-oestradiol-primed coho salmon to orally delivered LHRHa revealed that maximal GtH release occurred at doses between 2.0 and 20 μg LHRHa per g body weight.