Genetic analysis of growth and maturity in pink salmon (Oncorhynchus gorbuscha)

Variation in growth and sexual maturity was examined for five stocks of pink salmon (Oncorhynchus gorbuscha) spawning at different times in British Columbia. In each stock, four males were mated with eight females in a nested breeding design, and the juveniles were reared for 500 d after fry emergence. Adults in early-spawning northern stocks were smaller than those in late-spawning southern ones, but pink salmon from northern stocks had faster growth rates than those from southern ones. The relative ranking within stocks of family weight remained constant after late winter in the year of maturity. Heritability of weight based upon sire variance components was usually greater than 0.9 after 150 d of rearing. Pink salmon from the earlier-spawning stocks were in a more advanced state of sexual maturity when the experiment was terminated than were those from later-spawning stocks, indicative of a significant genetic component in timing of sexual maturity.     

Plasma insulin levels during the spawning migration of the pink salmon, Oncorhynchus gorbuscha

Plasma insulin concentration was measured by homologous radioimmunoassay in male and female pink salmon. Oncorhynchus gorbuscha , during spawning migration in the Fraser and Thompson Rivers, British Columbia. Although the fish ceased feeding prior to entering fresh water, plasma levels of insulin remained stable (males) or even elevated (females) during the final stages of oogenesis and spermatogenesis, decreasing thereafter. Mean concentrations ranged from 0–69 to 1.24 ng ml⁻¹ in males and from 0.33 to 0.88 ng ml⁻¹ in females. At all stages in the anadromous migration where a significant difference in plasma insulin levels between the sexes was observed, males had higher concentrations than females.     

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.     

Genetic differentiation of pink salmon Oncorhynchus gorbuscha Walbaum in the Asian part of the range

Genetic variation at 19 enzyme (including 11 polymorphic) and 10 microsatellite loci was examined in the population samples of odd-and even-broodline pink salmon from the southern part of Sakhalin Island, Southern Kuril Islands, and the northern coast of the Sea of Okhotsk. The estimates of relative interpopulation component of genetic variation for the allozyme loci, per broodline, were on average 0.43% (G _ST), while over the microsatellite loci it was 0.26% (the ?_ST coefficient, F-statistics based on the allele frequency variance), and 0.90% (the ρ_ST coefficient, R-statistics based on the allele size variance). The values of interlinear component constituted 2.34, 0.31, and 1.05% of the total variation, respectively. Using the allozyme loci, statistically significant intralinear heterogeneity was demonstrated among the regions, as well as among the populations of southern Sakhalin. Multidimensional scaling based on the allozyme data demonstrated regional clustering of the sample groups, representing certain populations during the spawning run or in different years. Most of the microsatellite loci examined were found to be highly polymorphic (mean heterozygosity > 0.880). The estimates of interlinear, interregional, and interpopulation variation over these loci in terms of ?_ST values were substantially lower than in terms of ρ_ST values. Regional genetic differentiation, mostly expressed at the allozyme loci between the populations from the northern Sea of Okhotsk and the Sakhalin and Kuril group of populations, was less expressed at the microsatellite loci. The differentiation between these regions observed can be considered as the evidence in favor of a large-scale isolation by distance characterizing Asian pink salmon. It is suggested that in pink salmon, low genetic differentiation at neutral microsatellite loci can be explained by extremely high heterozygosity of the loci themselves, as well as by the migration gene exchange among the populations (the estimate of the gene migration coefficient inferred from the “private” allele data constituted 2.6 to 3.4%), specifically, by the ancient migration exchange, which occurred during postglacial colonization of the range     

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.     

Genetic changes in pink salmon Oncorhynchus gorbuscha Walbaum during acclimatization in the White sea basin

Genetic parameters of pink salmon introduced into the White Sea basin in 1985 and 1998 were compared to the corresponding parameters of the donor population from the Ola River (Magadan oblast). The detected genetic differences indicate that colonization of a new area is accompanied by impoverishment of the gene pool of the native population. This effect was particularly marked in the odd-year line of pink salmon introduced in 1985. The probable causes of these genetic changes are discussed.     

The relationship between allozyme and morphometric variation in pink salmon Oncorhynchus gorbuscha from Southern Sakhalin

The relationship between allozyme polymorphism and morphometric variation in pink salmon from Southern Sakhalin was examined in order to determine the effects of individual genes. Dramatic differences were found between individuals of some genotypes. For instance, the difference in average male body length between the two prevailing PGDH* genotypes reached 36sigma, which corresponds to 5% contribution to the total variance of the trait. It would take only several generations for selection in favor of extreme phenotypes to rapidly shift the allele frequencies at loci of such a strong effect, which should be taken into account both in fishery management and in practical hatchery. Long-term selection can result in irreversible genetic changes in the population, which may be hazardous in the context of causing irreparable harm to the biological resources of the Earth.     

Changes in reproductive parameters during the spawning migration of pink salmon, Oncorhynchus gorbuscha (Walbaum)

The hormones 17β-estradiol, 17α-hydroxy-20β-dihydroprogesterone(17α, 20β-P), 11-ketotestosterone, testosterone, gonadotropin and also vitellogenin, were determined during the spawning migration of wild pink salmon in the Fraser and Thompson Rivers in British Columbia. This stock of pink salmon takes approximately 2 weeks to migrate the 333 km upstream to the spawning grounds. Both sexes were at an advanced stage of sexual development when they entered fresh water. In females both the 17β-estradiol and vitellogenin levels fell precipitously during the migration, to be very low at spawning, whereas the 17α,20β-P level rose rapidly, to be highest at arrival on the spawning grounds. The gonadotropin level also rose rapidly during the migration, and was highest in spent fish. Testosterone was at a high level throughout, although this level decreased steadily during migration. In many respects similar endocrine changes were observed in the male. For example, in the case of androgen levels, both testosterone and 11-ketotestosterone fell steadily during migration but were still relatively high at spawning, whereas both gonadotropin and 17α, 20β-P levels rose markedly as migration progress. However, although the qualitative changes were often similar between the sexes, the levels of 17α, 20β-P, testosterone, and gonadotropin were considerably higher throughout in females than in males. It is concluded that this stock of pink salmon is at an advanced stage of sexual development when it enters fresh water. The endocrine changes observed during this study represent those controlling the final stages of reproduction, specifically final oocyte maturation and ovulation in females, and the final stages of spermatogenesis and spermiation in males.     

Developmental progression of gill rakers as a post-hatch developmental marker in pink salmon, Oncorhynchus gorbuscha

We evaluate the usefulness of gill rakers as a post-hatch developmental marker in salmon by tracking development in undisturbed and stressed yolk-bearing salmon embryos. Native pink salmon (Oncorhynchus gorbuscha) from Auke Creek, Juneau, Alaska and genetically stressed outbred hybrids between Auke Creek and Pillar Creek (Kodiak Island) salmon were incubated in ambient-temperature Auke Creek water. Environmentally stressed native embryos were reared in water that was 2 to 4 °C warmer than ambient. The sum of rakers on the first left and right branchial arches of natives reared at ambient temperatures averaged 23.20 (SD?±?1.64) per embryo when post-hatch sampling began. The subsequent increase in raker counts was linear and positively correlated with the accumulation of thermal units until counts reached maxima 223 days after fertilization, which coincided with the complete consumption of yolk reserves. The average maximum raker count was 39.55 (SD?±?1.76) per embryo, which is substantially fewer than the 60 rakers typically observed in adults. Neither raker development nor yolk consumption patterns were affected by hybridization. Elevated incubation temperatures accelerated raker formation and yolk consumption in native embryos, but the number of rakers associated with a given amount of yolk was the same regardless of temperature suggesting that changes in yolk consumption rate and raker development rate did not influence raker counts. These results indicate that rakers are easily observed and counted, grow in a predictable sequence, and are developmentally stable in the face of both genetic and environmental stress, thereby making them potentially reliable post-hatch developmental markers.     

Genetic linkage mapping of allozyme loci in even- and odd-year pink salmon (Oncorhynchus gorbuscha)

We constructed genetic linkage maps of allozyme loci in even- and odd-year pink salmon (Oncorhynchus gorbuscha), using the total of 320 families (each female was crossed with two different males, and 80 females and 160 males were used for each of even year and odd year). The maps include eight linkage groups involving 22 loci. We observed substantial variation in recombination frequencies among different families within broodline and between sexes within broodlines. In the linkage analysis between sAAT-3* and sMDH-B1,2*, two even-year families and one odd-year family exhibited evidence of association, but two even-year and one odd-year families did not. Recombination rate tends to be reduced in males in pink salmon. The ratio of recombination rate (female/male), which ranged from 1.7 to infinity, averaged 2.8 in the even-year crosses and 3.2 in the odd-year crosses. The linkage groups (LG) I and II involving sAAT and mAH loci, which probably duplicated in the recent tetraploidization event, and the orders of loci in the LGs I (sAAT-3* --> mAH-4*) and II (mAH-3* --> sAAT-4*) were reversed, suggesting the possible paracentric inversion during salmonid evolution after the duplication.     

Use of a genetic marker to examine genetic interaction among subpopulations of pink salmon (Oncorhynchus gorbuscha)

In 1979 and 1981, a genetic marker was bred into one of the five identifiable subpopulations of pink salmon [Oncorhynchus gorbuscha (Walbaum)] in the Auke Lake drainage in Southeast Alaska. As a result of the marking effort, the frequencies of two malate dehydrogenase (MDH-B1, 2^*) alleles were changed in the marked subpopulation, but not in other subpopulations that spawn at different times or places. Between 1983 and 1989, the marker allele frequencies were monitored in many of these subpopulations and in early- and late-run pink salmon spawning in nearby Waydelich Creek, located approximately 1km away. Changes in allele frequencies at MDH-B1, 2^*, used to obtain direct estimates of average migration rates (m) from the marked to the unmarked subpopulations, revealed little or no introgression into early subpopulations or into nearby Waydelich Creek. Moreover, spatially distinct late-run Auke Creek subpopulations were not immediately overrun by the more abundant marked subpopulation. These observations suggest that genetic isolation exists between temporally distinct spawning runs and that small temporal and spatial (or ecological) differences contribute to population structure. These observations should be considered in taking actions that affect conservation and harvest management or extensive culture of salmonids.     

Physiological consequences of the salmon louse (Lepeophtheirus salmonis) on juvenile pink salmon (Oncorhynchus gorbuscha): implications for wild salmon ecology and management, and for salmon aquaculture

Pink salmon, Oncorhynchus gorbuscha, are the most abundant wild salmon species and are thought of as an indicator of ecosystem health. The salmon louse, Lepeophtheirus salmonis, is endemic to pink salmon habitat but these ectoparasites have been implicated in reducing local pink salmon populations in the Broughton Archipelago, British Columbia. This allegation arose largely because juvenile pink salmon migrate past commercial open net salmon farms, which are known to incubate the salmon louse. Juvenile pink salmon are thought to be especially sensitive to this ectoparasite because they enter the sea at such a small size (approx. 0.2 g). Here, we describe how 'no effect' thresholds for salmon louse sublethal impacts on juvenile pink salmon were determined using physiological principles. These data were accepted by environmental managers and are being used to minimize the impact of salmon aquaculture on wild pink salmon populations.     

Effect of hunger on the daily pattern of feeding rates in juvenile pink salmon, Oncorhynchus gorbuscha Walbaum

Juvenile pink salmon, Oncorhynchus gorbuscha , in two test groups of differing hunger level fed continuously on live copepods, maintained at high densities, throughout 12 h feeding periods in the laboratory. Analysis of video films showed that mean feeding rates were initially (first 10 min) between 51.5 and 63.8 prey-capture attempts (snaps) fish^-1 10 min^-1, but declined abruptly with increasing satiation to relatively constant levels of about 5.0 and 8.0 snaps fish^-1 10 min^-1, which were maintained during the subsequent 11 h of feeding. The data suggests that after the initial filling of their stomachs with food, juvenile pink salmon keep their stomachs full by feeding at a rate that balances the gastric evacuation rate of 18.6 mg prey h^-1 at 11° C. Less than 15% of the fish's stomach contents need be evacuated apparently for spontaneous feeding to resume or occur. Increased hunger level (by increasing a pre-test food deprivation period from 24 to 72 h) resulted in fish increasing their average feeding rate and thus their ration consumed from 23.6 to 39.8% dry body weight per 12 h.     

On the methodology of studying population structure using genetic markers (with reference to pink salmon Oncorhynchus gorbuscha)

General principles of studies on the species population structure are discussed. The most common errors are outlined in regard to the population-genetic analysis.