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Chapter 6

Genetic Drift

The last chapter described how inbreeding can cause random fluctuations in gene
frequencies in captive animal populations. The present chapter describes another
mechanism which can bring about random changes in gene frequencies, namely
genetic drift. First, genetic drift is defined. Then, methods of measuring drift and
ways to avoid drift are discussed. Lastly, the point is made that drift may lead to
increased genetic variability between groups of animals even when variability is
eroded within populations.

Genetic Drift Defined

Genetic drift is the random fixation or loss of genes in the population gene pool.
It is typically manifested by the loss of genetic variability in relatively small
populations of animals. Alleles are found in pairs on chromosomes and, by
chance, certain alleles may be randomly ‘fixed’ or lost to the gene pool when there
is only a small number of breeding individuals in the population, in much the
same manner that one is likely to get all ‘heads’ or all ‘tails’ in just a few tosses of
a coin. Drift is common when populations are founded with relatively few
individuals (‘founder effect’). For example, it is believed the first captive colony of
Syrian golden hamsters (Mesocricetus auratus) was established by a single littermate
pair (Murphy, 1985). Mongolian gerbils (Meriones unguiculatus) distributed in
laboratories worldwide are believed to have descended from a founder population
of 20 pairs of animals trapped in Manchuria in 1935 and brought to Japan for
breeding (Rich, 1968; Neumann et al., 2001). Eleven pairs were imported into the
USA in 1954 and of these animals, five females and four males were used to start
the breeding colony at Tumblebrook Farms Ltd, which has provided the breeding
stock for laboratories in the USA and Europe (Schwentker, 1963; Marston, 1972).
Drift may also occur when an existing population experiences a severe
reduction in numbers (bottleneck). Likewise, every time a small daughter colony

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38 (E.O. Price)

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Genetic Drift 39

is established, the phenomenon of genetic drift may be repeated because of the

limited number of breeding animals.

Measuring the Impact of Genetic Drift

In this context, genetic variation is typically estimated by: (i) measuring the degree
of heterozygosity or the proportion of individuals in a population which are
heterozygous at a given gene locus; (ii) measuring the number of alleles present
at a locus; and (iii) determining the percentage of loci which are polymorphic
(Allendorf, 1986; Agnese et al., 1995; Richards and Leberg, 1996). The possible
loss of heterozygosity and allelic diversity should be considered in the design
of breeding systems for captive animals. In general, heterozygosity provides a
good measure of the ability of a population to respond to selection immediately
following a reduction in numbers (i.e. bottleneck). The number of alleles remain-
ing after a bottleneck is important for the long-term response to selection and for
the survival of the population (Allendorf, 1986). Heterozygosity affects the ability
of a population to adapt to short-term changes in its environment, whereas allelic
diversity is more important in the long-term adaptation of populations. Allelic
diversity provides the flexibility for populations to make major shifts in phenotypic
traits over generations. Allendorf (1986) and Fuerst and Maruyama (1986) point
out that allelic diversity is lost much more rapidly than heterozygosity during
bottleneck and founding events. Hence, a larger founder population is needed to
protect allellic diversity than heterozygosity. Of course, ‘founders’ can be added
to the population periodically to protect it against the loss of genetic variation by
genetic drift (Odum, 1994; Ballou and Foose, 1996).
Since founder populations of captive wild and domestic animals often
consist of relatively few individuals, it is not unusual for the genetic variability of
these populations to soon decline due to drift and inbreeding (Verspoor, 1988).
Neumann et al. (2001), working with captive-reared wild and domestic Mongolian
gerbils (M. unguiculatus), found a marked reduction in heterozygosity and allele
number at nine polymorphic dinucleotide repeat loci in the laboratory stock.
Mean (± SE) heterozygosity was only 0.14 (± 0.06) in the domestic gerbils
compared to 0.76 (± 0.02) in the wild animals. Mean (± SE) allele number was 1.78
(± 0.28) at the nine loci in the domestic population and 9.20 (± 0.57) in wild stock.
The 45 domestic gerbils used in the study represented five different strains from
the USA and Europe and the 40 wild animals used were the second-generation
descendents of gerbils trapped in their native habitat (Mongolia) at six different
locations. Ryman and Stahl (1980) compared gene frequencies at two poly-
morphic loci in three ‘domestic’ stocks of brook trout (S. trutta) with gene
frequencies at the same loci in two natural populations of trout from the point of
origin of the hatchery stock. The polymorphism present at one locus in one of the
natural stocks was lost in the corresponding hatchery population. At the other
locus, there was a marked shift in gene frequencies. Verspoor (1988) reported that
mean heterozygosity in a relatively small population of Atlantic salmon (S. salar)

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40 Chapter 6

was reduced 26% in one generation of captive breeding. Mason et al. (1987) found
that a population of soybean loopers (Pseudoplusia includens) maintained in the labo-
ratory for 24 generations over a 2-year period showed a 62% reduction in
heterozygosity at six enzyme loci.
In a computer simulation program used to examine interacting effects of
genetic drift, mutation, immigration, selection and population subdivision on the
loss of genetic variability from small, managed populations, genetic drift was the
overriding factor controlling the loss of genetic variation (Lacy, 1987). As in
inbreeding, the loss in genetic variability due to genetic drift can negatively impact
fitness-determining traits. Ringo et al. (1986) reported that drift in two captive
populations of fruit flies, Drosophila simulans, reduced the propensity of flies to
mate and the number of offspring produced relative to an outcrossed base-line
population.

Avoiding Genetic Drift

Genetic variability can be maintained when founder populations are relatively
large and efforts are made to avoid inbreeding (Ferguson et al., 1991). Lengthening
the generation interval (time between birth of individuals and birth of their
offspring) will also help to preserve genetic diversity. The longevity of animals
living in captivity (Conway, 1978) allows animal breeders to preserve genetic
variability in this manner. Of course, longer generation intervals slow down
(in real time) the genetic adaptation of populations to the captive environment.
The traditional doctrine that founder events reduce genetic variation may not
always be correct. A founder event (population bottleneck) followed by a rapid
increase in population size (‘founder-flush’) may actually increase the genetic
variation available to selection due to significant shifts in the organization of the
gene pool, such as in the rapid increase in frequencies of previously rare alleles
(Carson, 1990; Meffert, 1999). Meffert and Bryant (1991) demonstrated no loss in
mating activity in four of six lines of houseflies (M. domestica) that experienced five
successive founder-flush bottlenecks to a single male–female pair. Reductions in
genetic variance within populations due to genetic drift are most likely to result
from a repeated succession of bottlenecks without recovery of population size.

Genetic Variation Between Populations

While genetic variance within populations may decrease due to drift, variability
between populations may not be affected or even increase (Dobzhansky and
Pavlovsky, 1957; Wang et al., 1990). Dobzhansky and Pavlovsky (1957) tested the
hypothesis that the outcome of natural selection should be more variable among
small than among large populations. They established ten populations of fruit flies
(Drosophila pseudoobscura) consisting of 4000 flies each and ten populations founded
by 20 flies each. At the start of the experiment, the PP gene arrangement was

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Genetic Drift 41

present on 50% of the third chromosomes examined from the source population.
At the end of the study, 19 generations later, the PP gene arrangement was
present on 27.4% of the third chromosomes examined from the ten large
populations and 32.7% of the third chromosomes from the ten small populations.
This difference was not statistically significant. However, the percentage of PP
chromosomes found among the ten large populations ranged from 20.3 to 34.7%,
while among the small populations, the percentage varied from 16.3 to 47.3%
(Fig. 6.1). This difference in variation was statistically significant and evidently due
to the different size of the foundation stocks.
One way to preserve genetic diversity in captive populations is to establish a
number of subpopulations under the assumption that if drift occurs, the genes
fixed or lost will not be the same in the various groups. In a similar manner,
genetic diversity may be preserved in natural populations by geographical and/or
behavioral isolation of subpopulations (Danielsdottir et al., 1997), an important
consideration when restocking captive-reared animals in nature. Fiumera et al.
(2000) report on the loss of genetic diversity in captive-bred populations of
a Lake Victoria cichlid (Prognathochromis perrieri) and the importance of captive
subpopulations located at a number of institutions in maintaining genetic diversity
in the population as a whole. The original captive population was founded by

Fig. 6.1. Range of percentage frequencies of the homozygous PP gene

arrangement on the third chromosome of individuals in ten populations of
Drosophila pseudoobscura founded by 4000 individuals (left) and ten populations
founded by 20 individuals (right) over 19 successive generations of laboratory
breeding (Dobzhansky and Pavlovsky, 1957).

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42 Chapter 6

10–20 adult fish. By the fourth generation of captive breeding, approximately

19% of the original alleles were lost and heterozygosity was reduced by 25%. At
the same time, approximately 33% of the total genetic diversity was maintained
solely between subpopulations. The authors speculate that the polygynous
mating system of the fish studied may have contributed to the relatively rapid
loss in genetic diversity. Rather than all fish contributing offspring for the next
generation, reproductive success was highly variable. A breeding program that
attempted to equalize family size in each generation would have helped to
preserve genetic diversity. Of course, equalizing family size to reduce genetic
change under captive rearing is only effective if it correlates with equal reproduc-
tive success within and between families (Waples, 1999).
Populations of social animals in captivity typically possess a number of
demographic characteristics that naturally increase the magnitude of genetic
drift and inbreeding relative to other evolutionary forces. Small populations,
polygynous mating, female philopatry and constraints on gene flow between
populations tend to reduce genetic variation within subpopulations and increase
genetic variation between populations (Storz, 1999).

Conclusions
Like inbreeding, genetic drift is most likely to occur in relatively small populations
of animals, whether in nature or captivity. A large majority of captive populations
of animals founded by humans are relatively small in numbers and thus
are subject to genetic drift. Genetic heterozygosity and allelic diversity within
populations of a species are typically reduced when drift occurs, thus potentially
compromising the ability of each population to adapt to changes in its environ-
ment. This may be most important to populations in the early generations in
captivity, particularly if the species is not well pre-adapted to the captive environ-
ment. Since the effects of genetic drift are random, genetic variability between
populations of a species may persist, offering animal breeders the opportunity to
maintain genetic variation by periodically crossbreeding animals from different
populations. Unlike inbreeding, genetic drift is not typically associated with a loss
of physical vigor or vitality.

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