Chapter 10
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populations of their wild counterparts. The current ease with which animals and
semen can be transported between populations has also facilitated efforts to
increase and maintain genetic variation both within and between captive animal
populations. These factors affecting genetic variability must be understood before
genetic improvement of economically important traits can be made (Hedgecock
et al., 1976). Documentation of genetic variation within rare breeds is important in
developing and administering conservation programs (Kantanen et al., 2000;
Giovambattista et al., 2001).
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otherwise have been lost or selected against in nature (Ginsburg and Hiestand,
1992). The fact that selection is still applied to preserve breed characteristics
suggests that sufficient genetic variability exists to permit further modification,
including reversion to behaviors previously selected against.
The retention of genetic variability, per se, during the process of domestication
is no guarantee that a population has retained its adaptive fitness. Domestication
may alter the distribution of such variability as well as the arrangement of
genotypes (Bartlett, 1985). Genetic variation has been found in wild house mice
(M. musculus) that is not present in laboratory populations. In contrast, there are
hundreds of mutant strains of mice held in laboratories (e.g. nude, obese, muscular
dystrophy strains) that could not survive in the wild. It could be argued that, taken
as a whole, laboratory mice are substantially more variable than their wild
ancestors in much the same way that the domesticated dog is now more variable
than its wild wolf-like ancestors (Festing and Lovell, 1981).
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76 Chapter 10
for at least 300 generations (15 years). Gibert et al. (1998) measured wing length
and body pigmentation in female wild-living fruit flies (D. melanogaster) and their
ninth-generation laboratory-reared descendants. Phenotypic variability for wing
length was 8.8 times greater in wild-living females than in the laboratory-reared
stock. Poor feeding conditions and higher developmental temperature were likely
explanations for the greater variability in wing length of the wild-living flies.
Phenotypic variability for pigmentation was similar for the two stocks. Shikano
et al. (2000) studied the relationship between genetic variation and salinity
tolerance (survival time after transfer from fresh water to 35 ppt artificial seawater)
in six wild populations and 14 domestic strains of the guppy (P. reticulata), a fish
native to northeastern South America. The domestic strains had been maintained
in captivity for 20–30 generations (10 years). Salinity tolerance and mean genetic
heterozygosity, estimated from five polymorphic and 23 monomorphic allozyme
loci, were greater in the wild populations than in the domestic strains. A positive
correlation was found between mean heterozygosity and salinity tolerance among
wild and domestic populations but not within populations (individual scores).
A loss in heterozygosity due to inbreeding and consequent inbreeding depression
(Chiyokubo et al., 1998) may have been responsible for the reduced salinity
tolerance of the domestic stocks. Dupont-Nevit et al. (2001) studied changes
in genetic variability in a population of the edible snail, Helix aspersa, over six
successive generations of laboratory breeding. The wild-caught generation
consisted of 500 snails sampled from 20 different geographical areas. No artificial
selection was applied during the first three generations but in generations four
through six, snails were selected for increased adult weight or they remained in
an unselected control line. Genetic variability was described using genealogical
parameters such as inbreeding, effective number of founders and ancestors,
effective number of remaining genomes and additive genetic variance. Large
decreases in all parameters of genetic variability were observed over the six
generations consistent with predictions of relatively strong natural selection. For
example, the effective number of ancestors (animals which contributed highly
to the population studied) decreased from 97.4 in the first laboratory-reared
generation to 34.1 in the sixth generation. Similarly, the effective number of
remaining founder genomes, which takes into account the random losses of genes
during reproduction, decreased from 96.2 to 34.1 between the first and sixth
generations. The authors concluded that natural selection was most intense
on families that did not adapt well to artificial rearing or that for other reasons
experienced poor reproduction. Some snails failed to mate under the laboratory
conditions provided. Wimmers et al. (2000) evaluated the genetic variability of
local chicken populations (Gallus) from Bolivia, India, Nigeria and Tanzania
with 22 microsatellites. Between two and 11 alleles per locus were detected and all
populations showed relatively high levels of heterozygosity ranging between 45
and 67%. Levels of heterozygosity are generally lower in commercial strains of
domestic chickens (G. domesticus; Crooijmans et al., 1996).
Norris et al. (2001) compared the microsatellite polymorphism between two
laboratory colonies and a large field population of Anopheles gambiae, the primary
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78 Chapter 10
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stock. Slater and Clayton hypothesize that breeders may have selected individuals
with more complex songs (e.g. more elements per second) for their breeding stock
and thus reduced the proportion of call notes in the song.
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82 Chapter 10
food among individuals and, consequently, more uniform growth rates. It would
be interesting to determine if growth rate in newly captive populations of animals
is greater when selection for growth rate is coupled with selection for reduced
behavioral reactivity.
Conclusions
The assumption held by many that domestication and captive-rearing results in a
reduction in genetic variation is only partly true. Recent investigations have
shown that considerable genetic variation has been retained in many populations
of captive and domestic animals. Studies revealing correlated effects of artificial
selection have demonstrated the interconnectedness of many traits. The proposed
‘resource allocation theory’ may be particularly relevant to the domestication
process since it hypothesizes a reallocation of available resources accompanying
shifts in selective pressures. With a limited pool of resources, exploiting a relatively
large proportion of the pool to support one trait results in fewer resources
available to support other functions. In captivity, environmental resources
normally used for survival in nature may become available for reproduction
and artificially selected production traits. The animal’s behavior can be an
integral part of this process.
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