Heterozygosity and components of fitness in a strain of rainbow trout

We examined the relationship between multilocus heterozygosity and several components of fitness in the Arlee strain of rainbow trout (Salmo gairdneri) . There is a significant negative association between multilocus heterozygosity and standardized rate of oxygen consumption (VO₂) in 168 fish produced from a pooled mating between 55 females and 60 males. Fish which are more heterozygous also tend to be longer, heavier and have a significantly greater condition factor than fish which are less heterozygous. However, no relationship is evident between VO₂, weight, length, condition factor and multilocus heterozygosity within two full-sib families. These results indicate that the larger size of more heterozygous fish may partially be attributed to the increased metabolic efficiency in heterozygotes compared to homozygotes and that these differences may only be detectable at the population level. More heterozygous females of the same age also have larger eggs. Energy conversion may be more efficient in more heterozygous females resulting in the production of larger eggs. The prediction that fish with faster developmental rates are metabolically more efficient and, therefore, have lower VO₂ was also tested. Earlier hatching fish from five families do not have significantly lower VO₂ than their later hatching full-sibs.     

Somatostatin inhibits growth of rainbow trout

Implantation of rainbow trout Oncorhynchus mykiss with somatostatin-14 (SS-14) for 20 days resulted in reduced food conversion as well as significant growth retardation compared to controls. Relative growth as mass was reduced by 20%, whereas relative growth by length was reduced by 45%. A single intraperitoneal injection of SS-14 reduced plasma levels of growth hormone (GH), insulin-like growth factor-1 (IGF-I) and insulin. SS-14 injection also reduced [³⁵S]-sulphate incorporation into gill cartilage compared to saline-injected fish. In addition, in vitro incubation of gill cartilage with SS-14 reduced [³⁵S]-sulphate incorporation in a doserelated manner. These results indicate that SS-14 inhibits growth of rainbow trout and suggests that SS-14, in addition to influencing GH, may play an extra-pituitary role in the modulation of the GH-IGF-I axis.     

Enzyme heterozygosity,metabolism, and developmental stability

Developmental homeostasis, measured as either fluctuating asymmetry or variance of morphological characters, increases with enzyme heterozygosity in many, but not all, natural populations. These results have been reported forDrosophila, monarch butterflies, honeybees, blue mussels, side-blotched lizards, killifish, salmonid fishes, guppies, Sonoran topminnows, herring, rufous-collared sparrows, house sparrows, brown hares, white-tailed deer, and humans. Because heterozygosity at a few loci can not predict heterozygosity of the entiry genome, these loci must be detecting localized zones that influence the developmental environment. Studies of malate dehydrogenase in honeybees,Apis mellifera, and lactate dehydrogenase in killifish,Fundulus heteroclitus, revealed that developmental homeostasis varied with heterozygosity of individual loci. Heterozygotes differed from homozygotes in fluctuating asymmetry, morphological variance, and in correlations between morphological characters. The protein loci in these studies code for enzymes, and therefore do not directly influence morphological characters. However, some enzymatic loci substantially influence metabolism, and contribute to variation in the amount of energy available for development and growth. This argument can be made most convincingly for the LDH polymorphism in killifish. LDH genotypes differ in enzyme kinetic properties that measure differences in physiological efficiency, and these differences produce measurable and predictable differences in physiology and development. Under environmental conditions which impose a stress upon development, genotypes at these loci may have different amounts of energy available for development, and consequently exhibit different levels of developmental homeostasis.     

Developmental rates of heterozygous and homozygous rainbow trout reared at three temperatures

The hatching distributions of rainbow trout (Salmo gairdneri) with different genotypes at eight loci are compared in two experiments with the same strain. Embryos were incubated at temperatures colder (5 and 8°C) and warmer (12°C) than normally experienced by these fish (9.5°C). At hatching, embryos were separated into five hatching groups representing the chronological order of hatching. There is no significant correlation between multilocus heterozygosity and hatching time at any temperature in either experiment. Fish in the middle of the hatching distribution had the highest average heterozygosity. In both experiments, heterozygotes at the majority of loci examined tended to hatch relatively later within the hatching distribution at 12°C than at both 5 and 8°C. Fish with different genotypes atPgm2 andCk1 showed significant differences in hatching time that were consistent between experiments.Ck1 heterozygotes hatched sooner than homozygotes at 8°C but later at 12°C.Pgm2 heterozygotes hatched later than homozygotes at all temperatures and significantly later in four of five cases. At the other loci examined, however, the relative hatching distributions of fish with particular genotypes were not significantly different or repeatable between experiments.This research was supported by National Science Foundation Grant BSR-8300039 awarded to Dr. Fred W. Allendorf. Moira M. Ferguson was supported by a postgraduate scholarship from the Natural Sciences and Engineering Research Council of Canada.     

Increased heterozygosity and child growth in an isolated subsistence agricultural community in the valley of Oaxaca, Mexico

Gene flow and rate of inbreeding (delta F) were calculated from demographic data for a community previously reported to be isolated from outside genetic influences of immigration. Significant child growth differences caused by gene flow among children born to native parents (n = 287) and offspring of native-immigrant matings (n = 38) were found in fatness (triceps skinfold), body proportions (sitting height ratio), and size (leg length). No differences were found between the two groups in height, weight, sitting height, and arm circumference. Variation in absolute and relative leg length in this population parallels previously reported differences in adult body size and proportion associated with increased heterozygosity caused by gene flow in other populations in southern Mexico.     

Recovery growth following periods of restricted rations and starvation in rainbow trout Salmo gairdneri Richardson

Fingerling rainbow trout (6.9 to 11.6 cm fork length) maintained at 12° C were either: fed restricted rations (3% of dry body weight/day) for 16 weeks, or starved for 3 weeks (14.5% weight loss), or for 13 weeks (32.5% weight loss). Whole body wet weights and dry weights (as % of wet weight), and wet and dry weights of tissues (heart, spleen, liver, skin, gonad, gut, visceral fat, and 'carcase') resulting from these treatments were compared to corresponding weights of an 'initial sample'. Visceral fat was completely utilized during both short and long-term starvation and gut was significantly reduced in the extended starvation group.
During subsequent recovery growth on full rations, following food deprivation, fish grew at approximately the same rate as fully-fed controls, which had not suffered food deprivation, as regards body wet weight and condition ( K ), but tended to surpass controls in % dry weight.
Heart, liver, gonad, gut and visceral fat also increased during recovery growth in a manner suggestive of overcompensation in response to former food deprivation. Gut, skin and carcase dry weights were less in the control, and in the 14.5% weight loss groups following recovery growth than in those in 3% rations and in the severely starved group, indicating that the effects of slow growth resulting from limited rations resemble those of severe starvation more than those of limited starvation.     

Uptake and accumulation of zinc in juvenile rainbow trout,Salmo gairdneri

Synopsis The toxicity of zinc to rainbow trout was determined and the 72 h median lethal concentration was found to be 2.00 mg l^–1 in freshwater, hardness 7.50 mg l^–1 as calcium. An insignificant increase in zinc concentration of internal tissues occurred in fish exposed to 1.52 mg l^–1 in freshwater for 72 h. However, there was a significant uptake of zinc by gills and the body surface. Fish exposed to 10 mg l^–1 zinc for 72 h in two-thirds sea water showed significant zinc uptake by liver, rectum and muscle, when compared to control fish. Drinking rate decreased from 1.43 to 0.26 ml kg^–1 h^–1 when zinc sulphate was added to freshwater. Trout adapted to two-thirds sea water showed no decrease in drinking, about 7 ml kg^–1 h^–1 when zinc was added to the water.     

Relationship between heterosis and heterozygosity at marker loci: a theoretical computation

Summary In this paper we have studied the linear correlation between a genetic distance index between two parent lines (based on marker loci information) and the heterosis observed in the F₁ hybrid from the two lines, for a quantitative character (determined by several loci, or QTL). Theoretical computations of the correlation coefficient () between the distance index and the heterosis were made, assuming the biallelic model (defined by Fisher). When the alleles at both marker loci and QTL are equally distributed among the whole population of considered lines, the coefficient is a function of the squares of linkage disequilibria between alleles at marker loci and alleles at QTL. The QTL that are not marked by marker loci and marker loci that do not mark any QTL play symmetrical roles and can decrease greatly. We conclude that the prediction of F₁ hybrid heterosis based on marker loci would be more efficient if these markers were selected for their relationship to the alleles implicated in the heterotic traits considered.     

Deviation from morphological intermediacy in interstrain hybrids of rainbow trout,Salmo gairdneri

Synopsis Deviations from morphological intermediacy in six first generation hybrids between three hatchery strains of rainbow trout, raised in a common environment, are reported. Hybrids have higher mean counts of four meristic characters than their maternal parental strain in a significantly greater number of cases (18 out of 24). Furthermore, eight of eleven hybrid indices are not intermediate. These results are discussed in reference to several mechanisms and models proposed to account for observed responses of meristic characters to environmental and genetic influences.     

Influence of growth rate on white muscle dynamics in rainbow trout and brook trout

In trout, fast growth stimulated white muscle fibre hypertrophy ( P 0·001) and hyperplasia ( P <0·01) in outer fibres but not in deep fibres. Glycogen was most prevalent in outer fibres ( P <0·01) and in brook trout ( P <0·01) that on average had three to four times larger fibres than rainbow trout.     

Biochemical and morphological correlates of growth in diploid and triploid rainbow trout

Diploid and triploid rainbow trout contained the same amount of RNA per unit weight of white muscle tissue at a given growth rate. This finding, along with a previous finding that the protein content of triploid muscle tissues is not different from that of the diploids, indicates that triploid nuclei produce more RNA than their diploid counterparts. This increased production of RNA is probably due to the expression of the extra set of chromosomes in triploid nuclei. RNA-based growth correlates and liver somatic index ( I _H) were valid predictors of growth for both triploid and diploid fish. RNA concentration appeared to be better than RNA/DNA and RNA/protein in predicting growth.     

Muscle fibre growth dynamics in diploid and triploid rainbow trout

The effect of triploidy on muscle fibre growth was determined by comparing hyperplasia and hypertrophy of white muscle fibres in all-female, diploid and triploid rainbow trout Oncorhynchus mykiss (100–400 mm total length). Conventional morphometry and protein and DNA concentrations were used to assess muscle fibre hyperplasia and hypertrophy in white muscle samples derived from an anterio-dorsal location. Muscle fibre distributions were significantly different between triploids and diploids in trout <300 mm. The proportion of fibres <20 μm was higher in diploids than in triploids and the proportion of fibres in the 20–40 μm category was higher in triploids than in diploids. This indicates that the hyperplastic fibres of triploids are larger than those of diploids. Larger hyperplastic fibres in triploids are probably due to the combined effect of increased nuclear size in triploids and the relatively high nucleus: cell ratio observed in small muscle fibres. These larger fibres may be less favourable to cellular metabolic exchange because of their smaller surface area to volume ratios, and perhaps account for reduced viability and growth observed in triploids during early life stages. On the other hand, the lack of difference in the distribution of fibres <20 μm between diploids and triploids at larger body size ranges (301–400 mm) imply that triploid trout may have higher rates of new fibre recruitment and growth capacity at these sizes. There was no difference between diploid and triploid trout in the mean size of muscle fibres; however, the number of fibres per unit area was reduced by 10% in triploids. No differences were observed in protein or DNA concentrations in muscle tissues between the two genetic groups. Since triploid nuclei have 1·5 times more DNA than diploid nuclei, this deviation from the expected muscle DNA concentration (1·3–1·4 times more DNA in triploids when the 10% reduction in fibre density is considered) suggests that the number of nuclei per muscle fibre is reduced. In both diploids and triploids, mean fibre size increased with body length while fibre density decreased. Similarly, protein concentration in the muscle tissue increased and DNA concentration declined with increasing body length. Protein/DNA ratio was strongly and positively correlated with fibre size. These results demonstrate that changes in DNA and protein concentrations can be used to assess hyperplasia and hypertrophy in muscle tissues. However, the morphometric procedure provides better insight into muscle fibre growth as it enables the direct visualization and analysis of muscle fibre distribution patterns.     

Rhythmic pattern of growth and nutrient retention in response to feeding time in the rainbow trout

The effect of six different feeding times was tested on feed intake, growth performance, proximate body composition and nutrient retention in the rainbow trout Oncorhynchus mykiss . Using a non‐linear regression model, a significant rhythmic pattern over a 24 h period was observed for feed gain ratio and nutrient retention responses to feeding time. Specific growth rate and protein growth rate responses were also rhythmic but the trends were not significant. There was no clear effect of feeding time on feed intake and proximate body composition. The study suggested that feed intake, at least under the experimental conditions encountered, was synchronized to feeding time while some physiological rhythms involved in nutrient metabolism were probably synchronized to photoperiod.     

Effects of body size on biochemical characteristics of trabecular cardiac muscle and plasma of rainbow trout (Oncorhynchus mykiss)

The objective of this study was to characterize biochemical indices of oxidative metabolism in trabecular cardiac muscle of female rainbow trout over a 200-fold size range. We also examined scaling effects on plasma concentrations of protein, albumin, and non-esterified fatty acids (NEFA). Animals were sampled from four size categories (<20 g, 100–200 g, 300–400 g and >2 kg). Ventricle mass relative to body mass was size-dependent, with the smallest trout having smaller hearts. Total protein in cardiac tissue was 20% higher in animals weighing over 300 g compared to fish less than 200 g. Plasma albumin and protein were increased 50–70% in trout over 100 g compared to fish smaller than 20 g. NEFA in plasma were similar for all animals. Activities of carnitine palmitoyltransferase and cytochrome oxidase were elevated 53% and 38%, respectively, in trabecular cardiac muscle of the largest trout compared to the smallest animals. Citrate synthase activity was independent of heart size, suggesting that the increase in oxidative capacity was not due to an increase in mitochondrial density. The increased capacity to bind and transport fatty acids in blood and the higher oxidative capacity of cardiac muscle may reflect a metabolic adaptation for increased oxidation of fatty acids and ventricular performance in larger female trout. These findings are not consistent with the scaling paradigm of oxidative metabolism and may reflect changes in ventricular architecture with ontogeny.     

Effect of changes in population size on the correlation between mutation rate and heterozygosity

Summary The effect of changes in population size on the correlation between mutation rate and heterozygosity was studied by using two models: sudden change in population size and gradual change. It was shown that the results for the two models are close to each other, unless the rate of change for the latter is exceedingly slow. Thus, in many cases, the former model, which is much simpler than the latter, can be used to treat the present problem. Numerical computations showed that the correlation in a population that is expanding or has expanded in the recent past is stronger while the correlation in a population that is decreasing or has experienced a population reduction or bottleneck in the recent past is weaker than that for an equilibrium population with the same mean heterozygosity. However, regardless of whether the population is at equilibrium or not, the proportion of variation in heterozygosity that is attributable to variation in molecular weight over loci is rather small if the mean heterozygosity of the population is low, say of the order 0.05 or smaller.     

The effect of antioxidants on the quality of cryopreserved semen in two salmonid fish, the brook trout (Salvelinus fontinalis) and the rainbow trout (Oncorhynchus mykiss)

Until now the supplementation of cryopreservation extenders with antioxidants has not been examined in teleost fish. Therefore, the present study investigated whether addition of antioxidants (catalase, superoxide dismutase, peroxidase, reduced glutathione, reduced methione, mixtures of reduced and oxidized glutathione or methionine) to the cryopreservation extenders could increase the quality of frozen-thawed semen of brook trout, Salvelinus fontinalis, and rainbow trout, Oncorhynchus mykiss. In brook trout and rainbow trout semen post-thaw fertility and motility were evaluated and in brook trout additionally the membrane integrity, DNA integrity, and sperm lipid peroxidation were evaluated. The tested antioxidants affected the motility parameters, DNA integrity, and fertility of cryopreserved semen, but not the membrane integrity. Most of the observed effects were negative and only minor positive effects were found. In brook trout 1.5 mmol/l reduced methionine and a mixture of 1.5 mmol/l oxidized and reduced glutathione increased the swimming velocity of frozen-thawed semen. One hundred U/l catalase, 1.5 mmol/l reduced glutathione, and 1.5 mmol/l reduced methionine slightly, but not statistically significantly increased the semen post-thaw fertility. However, these effects were not detectable in rainbow trout. Antioxidative stress or damage seems to play no role during cryopreservation, as also in the lipid peroxidation test no differences were obtained between fresh and cryopreserved semen. Therefore, for routine cryopreservation extender supplementation with antioxidants is not recommended in brook trout and rainbow trout.     

Genetic studies of laboratory reared mussels, Mytilus edulis: heterozygote deficiencies, heterozygosity and growth

Offspring derived from several single pair matings and a mass mating of Mytilus edulis were cultured up to the juvenile stage in the laboratory. Samples were collected from the cultures at an early post- settlement ('spat') stage and again at the juvenile stage and were scored at up to nine enzyme loci using cellulose acetate and starch gel electrophoresis. An overall significant deficiency of heterozygotes was evident at the juvenile stage but not at the spat stage. Selection against heterozygotes (underdominance) is proposed as the cause.
For each mating, shell lengths of juvenile mussels were measured and individuals were classed according to the number of loci at which they were heterozygous. No significant correlations were evident between heterozygosity class and either growth (shell length), or coefficient of variation of shell length. It is argued that these results provide support for the 'associative overdominance' rather than the 'overdominance' hypothesis as an explanation for heterozygosity/growth correlations in marine bivalves.
These data also show that although heterozygosity and growth co-occur in most data sets from natural populations, they may have distinctly differing causes.