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.     

Genetic variation of resistance to mercury poisoning in steelhead (Oncorhynchus mykiss) alevins

Newly hatched steelhead alevins were obtained from the factorial breeding of 24 male and 10 female steelhead trout, Oncorhynchus mykiss. Each set of offspring were in a separate cell. They were tested for resistance to intoxication by methylmercuric chloride (CH3HgCl) in water at a nearly constant mercury concentration of 8 microg l(-1). High mortality (81% of the tested alevins) occurred within 2 weeks. Resistance to intoxication, as measured by the time to death, as well as by the survival rate, shared high paternal and maternal variation with negligible interaction. Heritability of time to death was 0.59 +/- 0.17; heritability of survival (all-or-none trait) was lower (0.26 +/- 0.09). Mercury in dead alevins increased with time to death, exhibiting a large environmental variation and (comparatively) negligible genetic influence. At the end of the bioassay, the mercury content in survivors varied widely (3-21 microg g(-1) wet weight). The content was greater than, but correlated with that of dead alevins from the same cells, and it showed little relation with survival rate. Thus, it seems that resistance to poisoning implies a tolerance to high levels of mercury rather than a limitation of its accumulation.     

Genotype — hatch interactions as a source of bias in estimating genetic components of variance for eight-week weight in poultry

Summary Data on 16,557 chicks from 198 sire groups were analyzed to determine the importance of genotype-hatch interactions as sources of bias in the estimation of genetic parameters for eight-week weight in poultry. Estimates of heritability for eight-week weight obtained from a paternal half-sib correlation were .27±.04 and .29±.04 for males and females respectively when the sire component of variance was unconfounded with the sire-hatch interaction variance. If the sire-hatch interaction effects were confounded with the sire effects in an intra-hatch analysis the corresponding estimates of heritability would be .30 and .35 for males and females respectively. No attempt was made to artificially create differences in the hatch environments.The ratio of the dam-hatch interaction variance to the dam component of variance as compared with the sire-hatch interaction variance to the sire component gave a slight indication of a greater degree of importance of the interaction of non-additive genetic-environment effects than additive x environmental effects in the male progeny but not in the female progeny.The authors gratefully acknowledge Nichols, Inc., Brunswick, Maine for providing the data for this study.     

Seawater adaptability of underyearling coho,chinook and chum salmons in the estuary of Avacha River (Kamchatka)

Synopsis The 96 h LC50 tests were conducted with chum, coho and chinook salmon alevins caught in the Avacha River (Kamchatka) estuary. Coho and chinook alevins exhibited very low seawater a adaptability when compared with chum. It is likely that coho and chinook young were pulled downstream by high velocity flow during the summer freshets.     

Lost in the zygote: the dilution of paternal mtDNA upon fertilization

The mechanisms by which paternal inheritance of mitochondrial DNA (mtDNA) (paternal leakage) and, subsequently, recombination of mtDNA are prevented vary in a species-specific manner with one mechanism in common: paternally derived mtDNA is assumed to be vastly outnumbered by maternal mtDNA in the zygote. To date, this dilution effect has only been described for two mammalian species, human and mouse. Here, we estimate the mtDNA content of chinook salmon oocytes to evaluate the dilution effect operating in another vertebrate; the first such study outside a mammalian system. Employing real-time PCR, we determined the mtDNA content of chinook salmon oocytes to be 3.2 x 10(9)+/-1.0 x 10(9), and recently, we determined the mtDNA content of chinook salmon sperm to be 5.73+/-2.28 per gamete. Accordingly, the ratio of paternal-to-maternal mtDNA if paternal leakage occurs is estimated to be 1:5.5 x 10(8). This contribution of paternal mtDNA to the overall mtDNA pool in salmon zygotes is three to five orders of magnitude smaller than those revealed for the mammalian system, strongly suggesting that paternal inheritance of mtDNA per offspring will be much less likely in this system than in mammals.     

The road to extinction is paved with good intentions: negative association of fish hatcheries with threatened salmon

Hatchery programmes for supplementing depleted populations of fish are undergoing a worldwide expansion and have provoked concern about their ramifications for populations of wild fish. In particular, Pacific salmon are artificially propagated in enormous numbers in order to compensate for numerous human insults to their populations, yet the ecological impacts of this massive hatchery effort are poorly understood. Here we test the hypothesis that massive numbers of hatchery-raised chinook salmon reduce the marine survival of wild Snake River spring chinook, a threatened species in the USA. Based on a unique 25-year time-series, we demonstrated a strong, negative relationship between the survival of chinook salmon and the number of hatchery fish released, particularly during years of poor ocean conditions. Our results suggest that hatchery programmes that produce increasingly higher numbers of fish may hinder the recovery of depleted wild populations.     

Sex-specific genetic variances in life-history and morphological traits of the seed beetle Callosobruchus maculatus

Knowledge of heritability and genetic correlations are of central importance in the study of adaptive trait evolution and genetic constraints. We use a paternal half-sib-full-sib breeding design to investigate the genetic architecture of three life-history and morphological traits in the seed beetle, Callosobruchus maculatus. Heritability was significant for all traits under observation and genetic correlations between traits (r(A)) were low. Interestingly, we found substantial sex-specific genetic effects and low genetic correlations between sexes (r(MF)) in traits that are only moderately (weight at emergence) to slightly (longevity) sexually dimorphic. Furthermore, we found an increased sire ([Formula: see text]) compared to dam ([Formula: see text]) variance component within trait and sex. Our results highlight that the genetic architecture even of the same trait should not be assumed to be the same for males and females. Furthermore, it raises the issue of the presence of unnoticed environmental effects that may inflate estimates of heritability. Overall, our study stresses the fact that estimates of quantitative genetic parameters are not only population, time, environment, but also sex specific. Thus, extrapolation between sexes and studies should be treated with caution.     

Consequences to fitness‐related traits of hybridization between farmed and wild Atlantic salmon, Salmo salar

Increasing concern has been expressed about the genetic effects of cultured salmonid fishes on natural populations. Avoidance of extreme negative outcomes was one reason for the establishment of a genetic management policy for the State of Alaska. However, domestication within the hatchery may still cause divergence from the wild donor population. This divergence could potentially lead to adverse impacts on wild stocks through straying and introgression. This study examines potential domestication in two Alaskan chinook salmon stocks. The Little Port Walter (LPW) Hatchery Chickamin River stock resulted from a small collection of wild broodstock in 1976. The LPW Unuk stock was founded with a larger number of individuals in 1976 and has had subsequent infusion of wild gametes. These lines have been maintained at LPW through ocean ranching of tagged smolts. Comparisons are made between the hatchery lines, progeny of wild chinook collected from the Chickamin and Unuk Rivers, and hybrids between the hatchery and wild groups. Mature ocean‐ranched female chinook salmon returning to the facility were periodically graded for ripeness and spawned. Body size and meristic measurements were collected from these mature spawners. Maturation timing, fecundity, and individual egg size of these fourth generation hatchery fish are compared with that of offspring of wild fish from the same donor stock. Stock of origin is confirmed for all spawners and offspring using microsatellite DNA analysis.     

A Genetic Test of Sexual Size Dimorphism in Pre-Emergent Chinook Salmon

Sex differences in early development may play an important role in the expression of sexual size dimorphism at the adult stage. To test whether sexual size dimorphism is present in pre-emergent chinook salmon (Oncorhynchus tshawytscha), alevins were reared at two temperatures (10 °C and 15 °C) and sexed using the OtY1 marker on the Y-chromosome. Linear mixed models were used to test for sex differences in alevin size within families while controlling for the random effects of sire and dam nested within sire. Males and females did not differ in weight at 10 °C but males were heavier than females at 15 °C. Sex accounted for 2% of the within-family variance in weight. In addition, at 15°C, the relationship between weight and sex was greater in families with larger eggs. Whereas male-biased sexual size dimorphism was present at the juvenile stage, female-biased sexual size dimorphism was present at sexual maturity. Males were also younger than females at sexual maturity. A head start on growth by males may underlie their earlier maturation at a smaller size, thus leading to female-biased SSD at the adult stage.     

Comparison of maturation timing,egg size and fecundity between hatchery lines of chinook salmon and their wild donor stocks

Increasing concern has been expressed about the genetic effects of cultured salmonid fishes on natural populations. Avoidance of extreme negative outcomes was one reason for the establishment of a genetic management policy for the State of Alaska. However, domestication within the hatchery may still cause divergence from the wild donor population. This divergence could potentially lead to adverse impacts on wild stocks through straying and introgression. This study examines potential domestication in two Alaskan chinook salmon stocks. The Little Port Walter (LPW) Hatchery Chickamin River stock resulted from a small collection of wild broodstock in 1976. The LPW Unuk stock was founded with a larger number of individuals in 1976 and has had subsequent infusion of wild gametes. These lines have been maintained at LPW through ocean ranching of tagged smolts. Comparisons are made between the hatchery lines, progeny of wild chinook collected from the Chickamin and Unuk Rivers, and hybrids between the hatchery and wild groups. Mature ocean‐ranched female chinook salmon returning to the facility were periodically graded for ripeness and spawned. Body size and meristic measurements were collected from these mature spawners. Maturation timing, fecundity, and individual egg size of these fourth generation hatchery fish are compared with that of offspring of wild fish from the same donor stock. Stock of origin is confirmed for all spawners and offspring using microsatellite DNA analysis.     

Responses of alloplasmic (cytoplasm=Triticum timopheevii) and euplasmic wheats (Triticum aestivum) to photoperiod and vernalization

Summary Heritability estimated from sire family variance components, ignoring dams, pools conventional paternal and maternal half sib estimates, in a way which is biased upward, and sub-optimal for minimizing the sampling variance. Standard error of a sire family estimate will be smaller than that of the equivalent paternal half sib estimate, but not as small as that of an estimate obtained by optimal pooling of paternal and maternal half sib estimates. If only additive genetic variance components are significant, the bias may be removed by use of a computed average genetic relationship for sire families, in place of a nominal R = 0.25. Average genetic relationship may be computed from mean and variance of dam family size within sire families. If dominance, epistatic, or maternal components are significant, this simple correction is not appropriate. In situations likely to be encountered in large domestic species such as sheep and cattle (dam family size small and uniform) bias will be negligible. The method could be useful where cost of dam identification is a limiting factor.     

Temporal and Spatial Occurrence of Female Chinook Salmon Carrying a Male-Specific Genetic Marker in the Columbia River Watershed

Recent declines in many chinook salmon, Oncorhynchus tshawytscha, populations within the Columbia River watershed have prompted an examination of their reproductive biology. In a previous study many female fall chinook salmon collected in 1999 from the Hanford Reach of the Columbia River tested positive for a male-specific DNA marker (OtY1) found on the Y chromosome. The purpose of this study was to determine if females testing positive for the OtY1 marker could be found in other populations of fall chinook salmon from the Columbia River, and to assess the prevalence of OtY1 incidence in different female cohorts. Post-spawned male and female fall chinook salmon from three different naturally spawning populations (Hanford Reach, Yakima River and Ives Island) and one hatchery population (Priest Rapids Hatchery) on the Columbia River were tested in 2000 and 2001 for the OtY1 marker. Among naturally spawning populations, 57.4% of the females tested positive from the Hanford Reach, 33.3% tested positive from the Yakima River, and 32.5% tested positive from Ives Island. Of the Priest Rapids Hatchery fish, 62.5% of the females tested positive, and significant differences were detected between the 1995–1996 and 1997–1998 female cohorts from this population. No significant differences were detected between any of the female cohorts from the naturally spawning populations. All male chinook salmon samples, tested positive for OtY1.     

Probability of Obtaning Negative Estimates of Heritability from Full-Sib Analysis under a General Model of Gene Action

Formulae are derived for the probability of obtaining negative estimates of heritability (h²) in full-sib analysis under the additive-dominance-epistasis model of gene action. Evaluation of the probabilities was undertaken for several combinations of sire/dam number, h² and proportions of dominance and additive X additive epistatic variances, assuming two full-sibs per mating and that the dominance related epistasis is absent. In the light of the results, minimum sample sizes for obtaining admissible estimates from sire, dam components and their combination have been prescribed.     

Genetic variation in mortality of chinook salmon during a bloom of the marine alga Heterosigma akashiwo

Mortality in a netpen-reared population of 3-year-old chinook salmon Oncorhynchus tshawytscha during an extensive 1997 bloom of the alga Heterosigma akashiwo in Puget Sound, Washington, was low (7·2%), and corresponded to a reduction in variance effective population size of 9·4%. Under a liability threshold model, the heritability of mortality (± S.E.), based on paternal half-sibs, was estimated at 0·15 ± 0·04. No significant genetic variation was detected for date of death. Despite the low overall mortality, the consequences for variation in family size underscore the importance of maximizing genetic variation in cultured fish populations later released to the wild as a precaution against mortality and losses of genetic variation over the life cycle. The pattern of family variation in response to this algal bloom provides evidence for potentially selective mortality of anadromous salmonids in the marine environment during natural perturbations.     

A genetic analysis of body size in pink salmon (Oncorhynchus gorbuscha)

Two small-sized and two large-sized male pink salmon (Oncorhynchus gorbuscha) were mated to each of four females, producing eight families sired by small males and eight sired by large males. The juveniles were reared for 500 d after fry emergence. Juvenile weight in the two male size classes was similar until the spring of the year of maturity, when juveniles sired by large males grew faster than those sired by small ones. Heritability estimates of weight based upon the dam component of variance increased during 500 d of rearing from 0.4 to 0.8. Heritability of weight based upon the sire component of variance generally ranged between 0.1 and 0.3. The large variation in male body size in spawning pink salmon populations may have resulted from different male breeding strategies.     

Probability of Inadmissible Estimates of Heritability from Full-Sib Analysis Under a General Model of Gene Action

Evaluation of probabilities of the heritability (h²) estimates in full-sib analysis falling outside the permissible limits [0,1] was undertaken for several combinations of sire/dam numbers, h² and proportions of dominance and additive × additive epistatic variances, assuming two full-sibs per mating and the dominance related epistasis as negligible. In the light of the results use of sire component and sire-dam combination has been recommended in the estimation of heritability of lowly and highly heritable traits respectively.     

Comparative developmental biology of pink salmon, Oncorhynchus gorbuscha, in southern British Columbia

Eggs and alevins from 21 families of pink salmon, Oncorhynchus gorbuscha , from five odd-year broodline stocks spawning in southern British Columbia were incubated under controlled water temperatures of 4° C, 8° C and 12° C. There were significant differences in egg survival among stocks and among families within stocks at all incubation temperatures, but the differences were greatest at 4° C. Alevin survival was at least 97% for each stock at each temperature. The most northern spawning stocks had higher egg survival at 4° C than did the others. Hatching time of the alevins and emergence time of the fry were similar for all five stocks. Alevins hatching at 8° C were longer than those hatching at 4°C or 12°C, but there were no stock differences in alevin length or tissue weight. Stocks with larger eggs produced alevins of greater total weight and more yolk. Emergent fry from Vancouver Island stocks had the greatest tissue weight at 12° C, but Fraser River fry were heaviest at 8° C. There were significant differences among families within stocks for alevin and fry size parameters, suggesting that family variation should be accounted for in studies of salmonid developmental biology.     

Different prevalences of Renibacterium salmoninarum detected by ELISA in Alaskan chinook salmon Oncorhynchus tshawytscha spawned from freshwater and seawater

Soluble antigen of Renibacterium salmoninarum (Rs) was detected by a polyclonal enzyme-linked immunosorbent assay (ELISA) at significantly higher prevalences in adult chinook salmon Oncorhynchus tshawytscha that matured in freshwater than in the same cohort of fish spawned after maturation in seawater. The cumulative results were consistent during 4 yr of comparison at the Little Port Walter Hatchery on Baranof Island, Alaska, USA. Possible causes for this difference are discussed. Maturation of chinook salmon broodstock in seawater has become a practical strategy at this hatchery to reduce the prevalence of Rs-positive parent fish and the numbers of culled eggs.     

Evidence for genetic control of nematode egg-shedding rates in calves

Differences in nematode fecal egg counts from 167 purebred Angus calves born in 1986 and 133 born in 1987 that had been naturally exposed to infection on pasture were analyzed for sire effects. The calves were sired by 26 different bulls, all of which were related. Differences among fecal egg counts were attributable to sires (P less than 0.0002), to age of the calf (P less than 0.0007), to age of dam (P less than 0.05), and to collection year (P less than 0.0001). No variation in fecal egg count was attributable to calf sex (P greater than 0.23). After accounting for the effects of year, age of calf, and age of dam, heritability was estimated to be 29% (SE = 18%). It was concluded that a paternal contribution to the genome of each calf played an important role in determining average fecal egg output of the sire group. With heritability at 29%, selection against higher fecal egg counts or selection for lower fecal egg counts would be expected to produce measurable genetic change in a population.     

Delimiting the frequency of paternal leakage of mitochondrial DNA in chinook salmon

We analyzed embryos of a wild-return hatchery population of chinook salmon for the presence of paternal mtDNA. None of the 10,082 offspring examined revealed paternally transmitted DNA, delimiting the maximum frequency of paternal leakage in this system to 0.03% (power of 0.95) and 0.05% (power of 0.99).