Domestication has been extensively used in agricultural animals to modify phenotypes such as growth rate. More recently, transgenesis of growth factor genes [primarily growth hormone (GH)] has also been explored as a rapid approach to accelerating performance of agricultural species. Growth rates of many fishes respond dramatically to GH gene transgenesis, whereas genetic engineering of domestic mammalian livestock has resulted in relatively modest gains. The most dramatic effects of GH transgenesis in fish have been seen in relatively wild strains that have undergone little or no selection for enhanced growth, whereas genetic modification of livestock necessarily has been performed in highly domesticated strains that already possess very rapid growth. Such fast-growing domesticates may be refractory to further stimulation if the same regulatory pathways are being exploited by both genetic approaches. By directly comparing gene expression in wild-type, domestic, and GH transgenic strains of coho salmon, we have found that domestication and GH transgenesis are modifying similar genetic pathways. Genes in many different physiological pathways show modified expression in domestic and GH transgenic strains relative to wild-type, but effects are strongly correlated. Genes specifically involved in growth regulation (IGF1, GHR, IGF-II, THR) are also concordantly regulated in domestic and transgenic fish, and both strains show elevated levels of circulating IGF1. Muscle expression of GH in nontransgenic strains was found to be elevated in domesticated fish relative to wild type, providing a possible mechanism for growth enhancement. These data have implications for genetic improvement of existing domesticated species and risk assessment and regulation of emerging transgenic strains.
Production of transgenic animals has raised concern regarding their potential ecological impact should they escape or be released to the natural environment. This concern has arisen mainly from research on laboratory-reared animals and theoretical modeling exercises. In this study, we used biocontained naturalized stream environments and conventional hatchery environments to show that differences in phenotype between transgenic and wild genotypes depend on rearing conditions and, critically, that such genotype-by-environment interactions may influence subsequent ecological effects in nature. Genetically wild and growth hormone transgenic coho salmon (Oncorhynchus kisutch) were reared from the fry stage under either standard hatchery conditions or under naturalized stream conditions. When reared under standard hatchery conditions, the transgenic fish grew almost three times longer than wild conspecifics and had (under simulated natural conditions) stronger predation effects on prey than wild genotypes (even after compensation for size differences). In contrast, when fish were reared under naturalized stream conditions, transgenic fish were only 20% longer than the wild fish, and the magnitude of difference in relative predation effects was much reduced. These data show that genotype-by-environment interactions can influence the relative phenotype of transgenic and wild-type organisms and that extrapolations of ecological consequences from phenotypes developed in the unnatural laboratory environment may lead to an overestimation or underestimation of ecological risk. Thus, for transgenic organisms that may not be released to nature, the establishment of a range of highly naturalized environments will be critical for acquiring reliable experimental data to be used in risk assessments.coho salmon ͉ phenotypic plasticity ͉ risk-assessment
Selective breeding for enhanced growth in Pacific salmon Oncorhynchus spp. and other fish typically involves use of the largest mature individuals to breed for future generations of aquaculture broodstock. Owing to an altered selection regime, faster‐growing fish may not be as adapted to the natural environment as wild fish. To increase understanding of the genetic changes underlying selection for enhanced growth that results in phenotypic differentiation of farmed from wild Pacific salmon, multiple generations of pure and hybrid families were generated for coho salmon O. kisutch, including pure farm (D), pure native (Ch; a natural strain propagated by wild and hatchery production), F1 and F2 hybrids, and F1 × wild backcross (BCh) genotypes. The family groups were reared in the laboratory under controlled conditions as (1) individual genotypic groups, (2) mixed groups under culture conditions, and (3) mixed groups under enriched (seminatural) conditions. The growth of the fish was tracked until smoltification. There was a significant genotype effect on growth performance (mass and length), with rankings as follows: D > F2 > F1 > BCh > Ch. This ranking remained the same in all three rearing environments. Behavioral differences were observed among the families, the fast‐growing domesticated families showing a reduced antipredator response relative to the slow‐growing wild families. Expression of the phenotypic differences in the hybrids and backcrosses, together with the results from a joint‐scale analysis on line means, suggests that additive genetic effects contribute significantly to the divergence between the fast‐ and slow‐growing strains. As phenotypic differences between strains are largely a consequence of additive gene action, the phenotypic effects of domestication are largely diluted within two generations of backcrossing to wild salmon. Knowledge of the genetic changes responsible for altered growth rates is crucial to our ability to predict the consequences of introgression of domestic strains into wild populations of salmon.
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