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Last modified
7/14/2009 5:01:48 PM
Creation date
5/20/2009 11:07:33 AM
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UCREFRP
UCREFRP Catalog Number
9684
Author
Soule, M. E.
Title
Chapter 9 - Thresholds For Survival
USFW Year
n.d.
USFW - Doc Type
Maintaining Fitness and Evolutionary Potential.
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<br /> <br />were ignorant of the lethal dose. That is, enough is known about inbreed- <br />ing to justify concern that any increase in homozygosity reduces absolute <br />vigor and fecundity, but so far we have no way of deciding how much is <br />tolerable. We can not even pinpoint with any certainty the cause of in- <br />breeding depression, although we suspect that this loss of fitness is <br />mostly attributable to a load of deleterious recessive genes and to altered <br />gene interactions rather than to a loss of single gene heterosis concomi- <br />tant with decreasing heterozygosity (Eberhart, 1977; Comstock, 1977; <br />Falconer, 1977; Singh and Zouros, 1978). Finally, we know that a partial <br />antidote for the accumulation of such deleterious genes during inbreeding <br />is the winnowing effect of natural selection. Our problem is that we lack <br />data on the relative strengths of the poison (genetic load) and the anti- <br />dote (natural selection). <br />We cannot look to population genetic theory and models for help be- <br />cause the models for finite populations require several assumptions <br />(about mutation rates, gene frequencies, selection coefficients) about <br />which we can only guess. In addition, one must decide the mode of selec- <br />tion (dominance, overdominance, or even neutrality, a paradoxical <br />choice). Clearly, some more down to earth approach is needed. Perhaps <br />the best way to proceed is to side-step theory and resort to empiricism. <br />By trial and error, animal breeders have discovered how much in- <br />breeding can be tolerated by domestic animals before the lines begin to <br />decline in performance and fertility. Their rule of thumb is that the per <br />generation rate of inbreeding should not be higher than two or three per- <br />cent (Dickerson et aI., 1954; Stephenson et aI., 1953). Higher rates of in- <br />breeding fix deleterious genes too rapidly for selection to eliminate them <br />from the line. I prefer a slightly more conservative value, an upper limit <br />of one percent inbreeding per generation. The reasons are (1) domesti- <br />cated stocks have been partially purged of deleterious genes over the mil- <br />lenia, so they can tolerate higher rates of inbreeding than can <br />outbreeding species fresh from the wild, and (2) animal breeders can <br />safely ignore some classes of phenotypic change resulting from inbreeding <br />and genetic drift, whereas conservationists wish to preserve the "wild- <br />type". That is, conservationists may have to practice selection on more <br />traits than do animal breeders in order to preserve the original pheno- <br />type, and the rate of inbreeding must be decreased if the number of traits <br />being selected increases. <br />How does this basic rule (the one percent rule) translate into popula- <br />tion size? The rate ofloss of heterozygosity (increase in homozygosity) or <br />f per generation is equal to 1/2~, where ~ is the effective population <br />size. Thus, Ne must equal 50 or more if the inbreeding rate is to be kept <br />below the one percent threshold. <br />Even at the rate of one percent, however, the loss of genetic variation <br />is appreciable after a few generations. That is, even though natural selec- <br />tion may be able to nullify the most hannful inbreeding effects at this <br /> <br />160 <br />
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