The effects of depth and salinity on juvenile Chinook salmon Oncorhynchus tshawytscha (Walbaum) habitat choice in an artificial estuary

The energetic cost for juvenile Chinook salmon Oncorhynchus tshawytscha to forage in habitats of different salinity and depth was quantified using a behavioural titration based on ideal free distribution theory. When given a choice between freshwater habitats of different depths (>0·83 or <0·83 m), a greater proportion of fish used the deeper habitat. When the deeper habitat was saltwater, the proportion of fish using it increased. When food was added to both the shallow freshwater and deep saline habitats, however, fish distribution returned to that observed when both habitats were fresh water. This indicates that the preference for deep saline habitats during the stratified phase was driven by some benefit associated with residency in deeper water, rather than salinity. The low perceived cost of low salinity might be in part due to the fish's ability to minimize this cost by only making brief forays into the alternate freshwater habitat. When the food ration delivered to the more costly, shallow habitat was 50% greater than that delivered to the less costly, deep habitat, fish distributed themselves equally between the two habitats, presumably because of equal net benefits. This study demonstrates that juvenile Chinook salmon prefer deep saline habitat to shallow freshwater habitats but will make brief forays into the freshwater habitat if food availability is sufficiently high.     

Development of seawater tolerance and subsequent downstream migration in wild and stocked young‐of‐the‐year derived Atlantic salmon Salmo salar smolts

This study investigated the development of hypo‐osmoregulatory capacity and timing of downstream migration in wild Atlantic salmon Salmo salar smolts from the River Stjørdalselva and stocked young‐of‐the‐year (YOY), derived S. salar smolts from the tributary River Dalåa. Both wild and stocked S. salar smolts developed seawater (SW) tolerance in early May, persisting through June, measured as their ability to regulate plasma osmolality and chloride following 24 h SW (salinity = 35) exposure. Although the majority of downstream migration among the stocked S. salar smolts occurred later than observed in their wild counterparts, the development of SW tolerance occurred concurrently. The wild S. salar from Stjørdalselva and stocked YOY smolts from the River Dalåa started to migrate on the same cumulative day‐degrees (D°). The study revealed no downstream migration before development of SW tolerance. This emphasizes the importance of incorporating physiological status when studying environmental triggers for downstream migration of S. salar smolts. Overall, these findings suggest that the onset of smolt migration in stocked S. salar smolts was within the smolt window from an osmoregulatory point of view.     

The effects of paternal reproductive tactic and rearing environment on juvenile variation in growth as mediated through aggression and foraging behaviours of Chinook salmon (Oncorhynchus tshawytscha)

In species with indeterminate growth, differential growth rates can lead to animals adopting alternative reproductive tactics such as sneak–guard phenotypes, which is partially predicted by variation in growth during the juvenile life‐history stage. To investigate sources of growth variation, we examined the independent and joint effects of paternal reproductive tactic (G) and rearing environment (E) on juvenile growth in Chinook salmon (Oncorhynchus tshawytscha), hypothesizing G and E effects are partially mediated through differences in behaviour such as aggressive interactions and resulting foraging behaviours. We created maternal half‐sibling families with one‐half of the female's eggs fertilized by the milt of a sneaker “jack” and the other half by a guarder “hooknose”. At the exogenous feeding stage, each split‐clutch family was then divided again and reared in a rationed diet or growth‐promotion diet environment for approximately 6 months, during which growth parameters were measured. Before saltwater transfer at 9 months of age, social interactions were observed in groups of six fish of various competitor origins. We found ration restricts growth rate and juvenile mass, and evidence of genetic effects on growth depensation, where jack‐sired individuals grew less uniformly over time. These growth‐related differences influenced an individual's level of aggression, with individuals raised on a restricted diet and those whose families experienced greatest growth being most aggressive. These individuals were more likely to feed than not and feed most often. Jack‐sired individuals were additionally aggressive in the absence of food, and when raised on a rationed diet outcompeted others to feed most. These results show how individuals may achieve higher growth rates via intrinsic (G) or induced (E) aggressive behavioural phenotypes, and eventually attain the threshold body size necessary during the saltwater phase to precociously sexually mature and adopt alternative reproductive phenotypes.     

Genetic signals of artificial and natural dispersal linked to colonization of South America by non‐native Chinook salmon (Oncorhynchus tshawytscha)

Genetics data have provided unprecedented insights into evolutionary aspects of colonization by non‐native populations. Yet, our understanding of how artificial (human‐mediated) and natural dispersal pathways of non‐native individuals influence genetic metrics, evolution of genetic structure, and admixture remains elusive. We capitalize on the widespread colonization of Chinook salmon Oncorhynchus tshawytscha in South America, mediated by both dispersal pathways, to address these issues using data from a panel of polymorphic SNPs. First, genetic diversity and the number of effective breeders (N_b) were higher among artificial than natural populations. Contemporary gene flow was common between adjacent artificial and natural and adjacent natural populations, but uncommon between geographically distant populations. Second, genetic structure revealed four distinct clusters throughout the Chinook salmon distributional range with varying levels of genetic connectivity. Isolation by distance resulted from weak differentiation between adjacent artificial and natural and between natural populations, with strong differentiation between distant Pacific Ocean and Atlantic Ocean populations, which experienced strong genetic drift. Third, genetic mixture analyses revealed the presence of at least six donor geographic regions from North America, some of which likely hybridized as a result of multiple introductions. Relative propagule pressure or the proportion of Chinook salmon propagules introduced from various geographic regions according to government records significantly influenced genetic mixtures for two of three artificial populations. Our findings support a model of colonization in which high‐diversity artificial populations established first; some of these populations exhibited significant admixture resulting from propagule pressure. Low‐diversity natural populations were likely subsequently founded from a reduced number of individuals.     

Effects of early experience on the reproductive performance of Atlantic salmon

To examine the link between early experience and subsequentreproductive performance, we experimentally manipulated theearly experience of Atlantic salmon (Salmo salar). Salmon ofa common genetic background were reared as juveniles eithernaturally in the river or artificially in a hatchery (sea ranched),depriving them of river experience, and then allowed to growto maturity naturally in the ocean. At maturity, the competitiveand reproductive abilities of these fish were compared in fourexperiments quantifying reproductive success. Although levelsof aggressive behavior were similar, the experience-deprived,sea-ranched males were involved in more prolonged aggressiveencounters and incurred greater wounding and mortality thanwild males. Furthermore, sea-ranched males were less able tomonopolize spawnings and as a result obtained 51% the reproductivesuccess of wild males across the experiments. This reproductiveinferiority varied directly with the male density and bias inthe sex ratio, reflecting the intensity of male breeding competition.A lower intensity of female than male competition was likelyresponsible for the lack of differences in breeding performancebetween sea-ranched and wild females. Sea-ranched females, however,produced smaller eggs than wild females, apparently in responseto their higher juvenile growth rate. Differences in migratorybehavior were also apparent, as sea-ranched males and femalesascended the River Imsa later than wild fish. Our results indicatethat early experience has implications for subsequent adultreproductive performance, affecting the development of specializedskills and traits important not only for early life, but alsolater life.     

Quantitative genetic and translocation experiments reveal genotype‐by‐environment effects on juvenile life‐history traits in two populations of Chinook salmon (Oncorhynchus tshawytscha)

Understanding the genetic architecture of phenotypic plasticity is required to assess how populations might respond to heterogeneous or changing environments. Although several studies have examined population‐level patterns in environmental heterogeneity and plasticity, few studies have examined individual‐level variation in plasticity. Here, we use the North Carolina II breeding design and translocation experiments between two populations of Chinook salmon to detail the genetic architecture and plasticity of offspring survival and growth. We followed the survival of 50 800 offspring through the larval stage and used parentage analysis to examine survival and growth through freshwater rearing. In one population, we found that additive genetic, nonadditive genetic and maternal effects explained 25%, 34% and 55% of the variance in larvae survival, respectively. In the second population, these effects explained 0%, 24% and 61% of the variance in larvae survival. In contrast, fry survival was regulated primarily by additive genetic effects, which indicates a shift from maternal to genetic effects as development proceeds. Fry growth also showed strong additive genetic effects. Translocations between populations revealed that offspring survival and growth varied between environments, the degree of which differed among families. These results indicate genetic differences among individuals in their degree of plasticity and consequently their ability to respond to environmental variation.     

Maturation in chinook salmon, Oncorhynchus tshawytscha (Walbaum): early identification based on the development of a bimodal weight-frequency distribution

Male chinook salmon, orhynchus tshawytscha (Walbaum), are known to mature sexually after one summer in sea water, one year earlier than any of the females. Two year classes of first generation domestic chinook salmon stock reared to sexual maturity on a commercial salmon farm in British Columbia, Canada were sampled for wet weight at various times during the spring after their first winter in the sea. Frequency distributions of wet weight were developed for both year classes. The weight-frequency distributions for both cohorts became statistically bimodal in May of their first year in the sea. The fish from the two modal groups for both cohorts were sorted and kept separate over the summer. The difference in the mean weight for the two groups increased throughout the summer in both years. The higher weight group in each cohort proved to be >95% early maturing males, and represented 64–46% of all the early maturing males evident in the September following the sort. The potential use of this information for commercial salmon growers and fisheries scientists interested in sexual maturation of the chinook salmon is discussed.     

Lake‐specific variation in growth,migration timing and survival of juvenile sockeye salmon Oncorhynchus nerka: separating environmental from genetic influences

Time series on juvenile life‐history traits obtained from sockeye salmon Oncorhynchus nerka were analysed to assess lake‐specific environmental influences on juvenile migration timing, size and survival of fish from a common gene pool. Every year for the past two decades, O. nerka have been spawned at a hatchery facility, and the progeny released into two lakes that differ in average summer temperatures, limnological attributes and growth opportunities. Juveniles reared in the warmer, more productive Crosswind Lake were larger and heavier as smolts compared to those from the cooler, less productive Summit Lake and had higher in‐lake and subsequent marine survival. Crosswind Lake smolts migrated from the lake to sea slightly earlier in the season but the migration timing distributions overlapped considerably across years. Fry stocking density had a negative effect on smolt length for both lakes, and a negative effect on in‐lake survival in Summit Lake. Taken together, the results revealed a strong effect of lake‐rearing environment on the expression of life‐history variation in O. nerka. The stocking of these lakes each year with juveniles from a single mixed‐source population provided a large‐scale reverse common‐garden experiment, where the same gene pool was exposed to different environments, rather than the different gene pools in the same environment approach typical of evolutionary ecology studies. Other researchers are encouraged to seek and exploit similar serendipitous situations, which might allow environmental and genetic influences on ecologically important traits to be distinguished in natural or semi‐natural settings.     

Integration of Random Forest with population‐based outlier analyses provides insight on the genomic basis and evolution of run timing in Chinook salmon (Oncorhynchus tshawytscha)

Anadromous Chinook salmon populations vary in the period of river entry at the initiation of adult freshwater migration, facilitating optimal arrival at natal spawning. Run timing is a polygenic trait that shows evidence of rapid parallel evolution in some lineages, signifying a key role for this phenotype in the ecological divergence between populations. Studying the genetic basis of local adaptation in quantitative traits is often impractical in wild populations. Therefore, we used a novel approach, Random Forest, to detect markers linked to run timing across 14 populations from contrasting environments in the Columbia River and Puget Sound, USA. The approach permits detection of loci of small effect on the phenotype. Divergence between populations at these loci was then examined using both principle component analysis and F_ST outlier analyses, to determine whether shared genetic changes resulted in similar phenotypes across different lineages. Sequencing of 9107 RAD markers in 414 individuals identified 33 predictor loci explaining 79.2% of trait variance. Discriminant analysis of principal components of the predictors revealed both shared and unique evolutionary pathways in the trait across different lineages, characterized by minor allele frequency changes. However, genome mapping of predictor loci also identified positional overlap with two genomic outlier regions, consistent with selection on loci of large effect. Therefore, the results suggest selective sweeps on few loci and minor changes in loci that were detected by this study. Use of a polygenic framework has provided initial insight into how divergence in a trait has occurred in the wild.     

Upper thermal tolerance of wild‐type,domesticated and growth hormone‐transgenic coho salmon Oncorhynchus kisutch

In coho salmon Oncorhynchus kisutch, no significant differences in critical thermal maximum (c. 26·9° C, C_Tmax) were observed among size‐matched wild‐type, domesticated, growth hormone (GH)‐transgenic fish fed to satiation, and GH‐transgenic fish on a ration‐restricted diet. Instead, GH‐transgenic fish fed to satiation had significantly higher maximum heart rate and Arrhenius breakpoint temperature (mean ± s.e. = 17·3 ± 0·1° C, T_AB). These results provide insight into effects of modified growth rate on temperature tolerance in salmonids, and can be used to assess the potential ecological consequences of GH‐transgenic fishes should they enter natural environments with temperatures near their thermal tolerance limits.     

Effects of hatchery rearing on Florida largemouth bass Micropterus floridanus resource allocation and performance under semi‐natural conditions

This study examined the growth, activity, metabolism and post‐release survival of three groups of Florida largemouth bass Micropterus floridanus: wild‐caught fish, hatchery fish reared according to standard practice (hatchery standard) and hatchery fish reared under reduced and unpredictable food provisioning (hatchery manipulated). Hatchery‐standard fish differed from wild‐caught fish in all measured variables, including survival in semi‐natural ponds. Hatchery‐standard and hatchery‐manipulated fish showed higher activity levels, faster growth and lower standard metabolic rates than wild‐caught fish in the hatchery. Fish reared under the manipulated feeding regime showed increased metabolic rates and increased post‐release growth, similar to wild‐caught fish. Their activity levels and post‐release survival, however, remained similar to those of hatchery‐standard fish. Activity was negatively correlated with post‐release survival and failure of the feed manipulation to reduce activity may have contributed to its failure to improve post‐release survival. Activity and post‐release survival may be influenced by characteristics of the rearing environment other than the feeding regime, such as stock density or water flow rates.     

Effect of restricted feeding on early maturation in female and male amago salmon, Oncorhynchus mason ishikawae

Incidence of early maturation of female amago salmon was reduced by >32% by feed restriction beginning in May, 5 months prior to spawning. Male early maturation was unaffected by this feed restriction regime.     

Effect of additional food and water on house mice in a semi‐arid agricultural environment in Australia

Abstract We tested the hypothesis that providing high‐quality food and water would increase reproduction, survival and population size in house mice living in a semi‐arid cropping environment in the mallee region of western Victoria, Australia, where outbreaks of house mice (Mus domesticus) occur irregularly. We employed a factorial design and applied treatments along internal fencelines (16 sites), adjacent to cropping areas, from November 2003 to July 2004. Population abundance was low during the experiment (0–26 mice per site), and the summer rainfall was below average. We confirmed that mice used the supplementary food and water through a reduction in weight of food containers over time and for water through the presence of Rhodamine B in blood samples and positive bands in whiskers. Abundant food was also available through grain spilt on the ground after harvest of the wheat and barley crops. There was some evidence of increased breeding on sites where water was added, but no effect of food or food and water in combination. Sites where free water was available had marginally higher populations over summer (～2 more mice on average; P = 0.07). This difference was well below any biologically meaningful effect. Mice were 0.9 g heavier on sites where water was added (P = 0.04), and were in better condition (P = 0.03). The addition of high‐quality food did not affect mouse population dynamics, and the addition of water resulted in only marginal responses for some demographic characteristics. We conclude that other factors appear to be important for limiting mouse population growth in summer and autumn.     

Effects of an autocorrelated stochastic environment and fisheries on the age at maturity of Chinook salmon

Chinook salmon (Oncorhynchus tshawytscha) reproduce only once in their lifetime, and their age at reproduction varies among individuals (indeterminate semelparous). However, the factors that determine their spawning age still remain uncertain. Evidence from recent studies suggests that individual growth and reproduction of Chinook salmon are affected by the rate of coastal upwelling, which is shown to be positively autocorrelated between years. Therefore, the serially autocorrelated environmental is expected to play an important role in determining their spawning age. In the present study, I demonstrate the advantage of an indeterminate maturation strategy under a stochastic environment. I then present theoretical evidence for the advantage of adjusting the maturation probability based on the environment they experienced and demonstrate that fisheries reduce the fitness of the strategy to delay maturation. The results presented herein emphasize the importance of incorporating detailed life-history strategies of organisms when undertaking population management.     

Persistent parental effects on the survival and size, but not burst swimming performance of juvenile sockeye salmon Oncorhynchus nerka

Sockeye salmon Oncorhynchus nerka were used as a model in an artificial fertilization experiment to investigate the relationships between individual adult O. nerka and their offspring. Survival, size and burst swimming ability were assessed in fry of known parentage (adult spawners from the Weaver Creek population, British Columbia, Canada). Maternal identity significantly affected the survival rate of eggs at hatch time, though this effect did not extend to fry life stages. The results were also suggestive of a paternal effect on both egg and fry survival, though this could not be separated from the experimental block design. After 4 months of exogenous feeding, fry mass remained under significant maternal influence, though fork length did not, despite having a high correlation with mass. Burst swimming performance was highly variable among individuals, and was not significantly influenced by maternal identity or individual fry size. Collectively, the findings presented here suggest that maternal, and possibly paternal, effects can be integral components of population dynamics in the early life stages of O. nerka . A good understanding of these factors will be essential for scientists and fisheries managers in developing a more holistic view of population-level spawning success and fry survival.     

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.     

Effect of rearing environment and dietary zinc oxide on the response of group-housed weaned pigs to enterotoxigenic Escherichia coli O149 challenge

A 2 × 2 factorial experiment was conducted to determine the effects of rearing environment (indoor (In) v. outdoor (Out)) and dietary zinc oxide (ZnO) supplementation (0 (-Zn) v. 3100 (+Zn) mg/kg feed) on the response of weaned pigs to a challenge infection with enterotoxigenic Escherichia coli (ETEC). Pigs from the two rearing environments were weaned onto trial diets at 4 weeks of age, moved into conventional accommodation and infected 3 days later with 109 CFU ETEC per os. Faecal ETEC shedding was determined before and after challenge. After 7 days of ETEC infection, all pigs were euthanized for gut lactic acid bacteria (LAB)-to-coliform ratio, pH and small intestine morphological measurements. Both ZnO and outdoor rearing reduced ETEC excretion, and these effects were additive. Outdoor rearing increased small intestine and colon tissue weight. ZnO increased villus height and goblet cell number in the upper small intestine, LAB-to-coliform ratio (through reduced coliforms) in the lower small intestine and proximal colon, and improved growth performance. There were interactive effects of rearing environment and ZnO supplementation on upper small intestine villus height and daily gain, as outdoor rearing conferred advantages on these variables only with ZnO dietary supplementation. Daily gains were 233, 174, 277 and 347 (s.e.m. 27.2) g/day for the In - Zn, Out - Zn, In + Zn and Out + Zn, respectively. These results suggest different, but complementary mechanisms of intestinal health and performance in outdoor-reared pigs and those offered ZnO supplemented diets. The results indicate that the benefits of ZnO to the weaned pig extend beyond suppression of ETEC and appear mediated through altered development of the small intestine mucosa.     

Assessing the effects of repeated handling on the physiology and condition of semi‐precocial nestlings

Repeated exposure to elevated levels of glucocorticoids during development can have long‐term detrimental effects on survival and fitness, potentially associated with increased telomere attrition. Nestling birds are regularly handled for ecological research, yet few authors have considered the potential for handling‐induced stress to influence hormonally mediated phenotypic development or bias interpretations of subsequent focal measurements. We experimentally manipulated the handling experience of the semi‐precocial nestlings of European Storm Petrel Hydrobates pelagicus to simulate handling in a typical field study and examined cumulative effects on physiology and condition in late postnatal development. Neither baseline corticosterone (the primary glucocorticoid in birds), telomere length nor body condition varied with the number of handling episodes. The absence of a response could be explained if Storm Petrels did not perceive handling to be stressful or if there is dissociation of the hypothalamic–pituitary–adrenal axis from stressful stimuli in early life. Eliciting a response to a stressor may be maladaptive for cavity‐dwelling young that are unable to escape or defend themselves. Furthermore, avoiding elevated overall glucocorticoid exposure may be particularly important in a long‐lived species, in which accelerated early‐life telomere erosion could impact negatively upon longevity. We propose that the level of colony‐wide disturbance induced by investigator handling of young could be important in underlining species‐specific responses. Storm Petrel nestlings appear unresponsive to investigator handling within the limits of handling in a typical field study and handling at this level should not bias physiological and morphological measurements.     

Relationship of winter concealment habitat quality on pool use by juvenile spring Chinook salmon (<Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis>) in the Grande Ronde River Basin,Oregon USA

Winter concealment habitat quality was assessed and its use by juvenile spring Chinook salmon (Oncorhynchus tshawytscha) quantified in three hatching areas of the Grande Ronde River Basin, Oregon USA. Fish densities were significantly higher in pools with a higher winter concealment habitat index than pools with a lower index. The mean fork length and mean growth rate of fish did not differ between pools with a higher or lower winter concealment habitat index, even though residual fish were significantly larger than fish that emigrated. Biomass–density was significantly higher in pools with a higher winter concealment habitat index than pools with a lower index in all three hatching areas. Biomass–density was positively associated with the amount of cobble substrate (10–24.9 cm/m²) in all three hatching areas, and inversely associated with embeddedness in two of the hatching areas. Results of this study indicate that enhancing winter concealment habitat could improve habitat quality resulting in increased carrying capacity and winter usage by juvenile spring Chinook salmon. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. A. Cambray