Laserfiche WebLink
Data on inbreeding effects in fishes are impressive, and <br />include severe body deformities, growth reduction, behav- <br />ioral changes, and reproductive failures in such forms as <br />convict cichlids, Cichlosorna nigrofasciatunc (Winemiller and <br />Taylor 1982), carp, Cyprinus carpio (Moav and Wolfarth 1963, <br />ref. in Mrakovcic and Haley 1979), zebra fish, Brachydanio <br />rerio (Mrakovcic and Haley 1979), brook trout, Salvelinus fon- <br />tinalis (Cooper 1961), and rainbow, trout, Sahno gairdneri <br />(Aulstad and Kittleson 1971; Gjerde et al. 1983; Kincaid 1983). <br />Although species and populations differ markedlv in their <br />resistance to inbreeding depression (Ralls et al. 1979; Frank- <br />lin 1980), and magnitude of inbreeding effects cannot be <br />predicted beforehand, "... the unavoidable conclusion is <br />that relatively small amounts of inbreeding can do tremen- <br />dous damage to the reproductive potential and productivity <br />of a fish stock." (FAO/UNEP 1981, p. 10). <br />A different conservation genetics problem is the imposi- <br />tion, knowingly or othenvise, of artificial selection upon a <br />wild stock, leading to a state of domestication (Brisbin 1974). <br />That fish respond by genetic change to selection in captivity <br />is well established in salmonids (Vincent 1960; Donaldson <br />1970; Kincaid et al. 1977) and other groups (Moav et al. 1978; <br />Hynes et al. 1981). By relaxing natural selection, or replacing <br />it with random or directional artificial selective forces, the <br />fate of propagated populations may be jeopardized. Assum- <br />ing that natural selection has already optimized most char- <br />acter states of populations in their particular habitats, any <br />deviation from those states, even those advantageous in <br />captivity, would not be beneficial in nature. "... genetically <br />based performance under one set of conditions (i.e., hatch- <br />ery) may not be correlated with performance under a dif- <br />ferent set of conditions (stream, lake, natural area). If the <br />goal is to release stock in a different environment from that <br />where they are bred, then the brood stock selection practices <br />must be designed to avoid unconscious selection and in- <br />breeding." (FAO/UNEP 1981, p. 25). <br />Frankel and Soule (1981) discuss four types of selection <br />in captivity that can affect genetics: selection for increased <br />productivity, selection for a (perfect) type, selection for tract- <br />ability, and non-selection. Selection for increased produc- <br />tivity in fishes can occur when the most fecund females are <br />continually used to develop a breeding stock. Although this <br />is most efficient in a propagation program,.it can be dele- <br />terious in nature, where optimal clutch size is a function of <br />predation risks, growth rates, adult and juvenile mortality <br />schedules, food availability, etc. (Stearns 1976, 1977). Since <br />increased fecundity may be maladaptive in nature, this type <br />of artificial selection should be avoided. <br />It may also be tempting to select for a particular type, such <br />as larger or smaller, more or less aggressive, more colorful, <br />etc. This is analogous to directional or stabilizing selection <br />and can reduce quantitative or qualitative gene pool varia- <br />tion. The third type of artificial selection, for tractability, is <br />really a subset of selection for a type. It occurs when less <br />aggressive animals are used for breeding because they are <br />easier to handle. The practice of non-selection occurs when <br />the survival probability of sick or abnormal individuals is <br />increased by the captive breeding program; this may in- <br />crease the incidence of deleterious genes in the population. <br />Effects of Variance Reduction Within Populations <br />Thus far, I have presented sources of genetic variance <br />reduction within populations, under the implicit assump- <br />tion that loss of variance is undesirable. What specific ge- <br />netic and phenotypic changes, however, will arise from re- <br />duced variance, and what is the evidence that these changes <br />have adverse effects? Loss of genetic variance has three ma- <br />jor effects: increased homozygosity (leading to reduced fit- <br />ness), loss of additive variance, and increase in deleterious <br />recessive alleles. <br />There is little doubt that increased homozygosity can lower <br />an individual's fitness (Beardmore 1983) although actual <br />mechanisms for this reduction are not known. With respect <br />to relative fitnesses of hetero- and homozygotes, there exists <br />a "heterozygosity consensus," which is "... the belief based <br />on extensive laboratory and farm experience that fitness <br />(variability, vigour, fecundity, fertility, etc.) is enhanced by <br />heterozygosity, and that any decrease in genetic variation <br />will be paralleled by a diminution of fitness" (Frankel and <br />Soule 1981, p. 48). Numerous data sets directly or indirectly <br />support the notion that reduction in genetic variance may <br />lower fitness (Wheat et al. 1974; Vrijenhoek and Lerman <br />1982; Leary et al. 1984) and the interested reader is referred <br />to Mitton and Grant (1984, and numerous references therein) <br />for a thorough review. These papers in aggregate demon- <br />strate that increased homozygosity can result in at least slower <br />growth, reduced survivorship, and developmental instabil- <br />ity. Perhaps the most striking example of effects of ho- <br />mozygosity is in the South African cheetah (Acinonyx j. ju- <br />batus), which has a paucity of genetic variance at over 200 <br />structural loci and the major histo-compatibility complex, <br />apparently due to a severe bottleneck (O'Brien et al. 1985). <br />These animals have great difficulty breeding in captivity, <br />high rates of juvenile mortality in wild and captive popu- <br />lations, and an abnormally high susceptibility to an epizootic <br />coronavirus, all attributed to genetic uniformity. <br />With respect to the second problem, additive genetic var- <br />iance is generally lost at the same rate as heterozygosity <br />1 <br />(2Ne) per generation [Franklin 19k]) . The major cause is <br />genetic drift, which of course may be powerful in small <br />populations. The reader is referred to Fisher (1930), Franklin <br />(1980) and Falconer (1981) for more detailed discussions of <br />additive genetic variance. The third point, increase in del- <br />eterious recessive alleles, has been discussed in the section <br />on inbreeding. <br />Among-Population Variance <br />Now we approach a very different problem, that of genetic <br />variation among populations. Spatially isolated populations <br />of organisms, with little to no gene flow among them, will <br />tend to genetically diverge either through different selective <br />forces or random drift (Endler 1977); in its extreme form, <br />this leads to speciation. Any isolated population of fishes is <br />a potentially unique gene pool with characteristics that may <br />differ from all other demes. This is particularly evident in <br />salmonids, which demonstrate the potential for extensive <br />genetic divergence of fish stocks (Donaldson 1971; Busack <br />et al. 1979; Stahl 1981, 1983). Consequently, in cases where <br />endangered fishes exist in more than one natural popula- <br />tion, we have at our disposal a powerful management tool: <br />the potential existence of genetically distinct groups with <br />which to recover the species. This is not a trivial observation, <br />for it allows the manager flexibility in building breeding <br />stocks, reintroduction to field sites, and other management