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Chapter 7
Artificial Selection
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44 Chapter 7
is often applied to hatchery-raised broodstock, since they produce young that are
larger at traditional release times or can be released earlier in the spring.
These early-released fish may have higher survival rates providing unpredictable
(undesirable) springtime water conditions do not occur.
Inadvertent artificial selection may also account for an acceleration in sexual
development and improved sexual vigor in a long-standing laboratory population
of oriental fruit flies (Dacus dorsalis). In a comparison of a domestic stock of
D. dorsalis maintained in the laboratory for about 330 generations with wild
D. dorsalis collected from three islands in Hawaii, Wong et al. (1982) found that
100% of the domestic females had mated by day 12 after eclosion whereas 90% of
the wild females were not mated until day 25. In addition, the mating speed
(proportion of flies mating per hour) of male domestic flies was about four times
that of the wild males and the domestic females mated about twice as fast as the
wild females. The authors concluded that the laboratory rearing system employed
over the 28 years of propagating the laboratory population had selected for early
sexual development and rapid mating behavior (i.e. sexual vigor). It should be
noted that the wild flies used in this study were older than the laboratory flies
(28–30 days vs. 9–10 days), since it took them longer to reach sexual maturity.
Could mating speed have been influenced by the different ages of the two stocks at
testing? To what extent are mating speed and early sexual maturation related in
this species? It is also possible that light intensity could have confounded the
results of the mating speed experiment. Copulations were monitored for 4 h
starting at 10.00 and 11.00. Wild fruit flies mate more frequently under low
light conditions, which may correspond with the cooler and more humid times
of the day. Selection for sensitivity to light intensity may have been relaxed
in long-standing laboratory populations of fruit flies because of the constant
temperature and humidity of the laboratory environment. Hence, domestic stocks
may be more active in the middle of the day. Koyama et al. (1986) reported that
sexual activity in a laboratory strain of melon flies (Dacus cucurbitae) was initiated
earlier in the afternoon (i.e. at higher light intensity) than in a comparable wild
population.
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Artificial Selection 45
The variety of physical and behavioral traits seen in the many breeds of domestic
dogs (C. familiaris; Scott and Fuller, 1965) reflect the genetic changes that can be
realized in a species through conscious artificial selection and hybridization. Dog
breeds are more diverse in shape and conformation than any other domestic
animal species.
It is estimated that selective breeding of dogs for specific functions and
appearance began about 5000 years ago (Case, 1999). Dogs resembling today’s
greyhounds represent one of the oldest breeds and were often shown on paintings
and pottery in Egypt and western Asia as early as 2900 BC. The ancient Romans
were also heavily involved in the systematic breeding of dogs. Written records
from the fifth century BC describe dogs used for herding, sport, war, arena
fighting, scent hunting and sight hunting. The aristocracy also kept smaller ‘house’
dogs. Mastiff breeds were developed primarily in the area of Tibet and were later
used in war by the Babylonians, Assyrians, Persians and Greeks. Wolf-like breeds
similar to today’s spitz were developed early and are the likely ancestors of
modern-day huskies, Keeshond and other arctic breeds. Pointers were developed
early on to hunt small game and most of the sheepdog breeds probably originated
in Europe and were selected to herd livestock.
The majority of exisiting purebred dog breeds have their origins in some type
of working animal (Case, 1999). The greatest era for the proliferation of dog
breeds was in the Middle Ages in western Europe, spanning the 13th to 15th
centuries AD. Notable during this period were the many breeds (e.g. deerhounds,
wolfhounds, otterhounds, bloodhounds, etc.) established to hunt different species
of game (Menache, 1997).
Artificial selection has not only been used to establish various breeds within
domestic animal species but it has been used to establish populations differing in
very specific characteristics. For example, Bernon and Siegel (1983) demonstrated
the extent to which sexual performance in male chickens (G. domesticus) can be
artificially selected in 20 generations (Table 7.1). Koteja et al. (1999) selected
laboratory mice (M. domesticus) for activity wheel use and, after 13 generations,
reported a 2.2-fold increase in number of revolutions per day. Interestingly, the
increase was in running speed not time spent in the wheels and the selection-line
mice ‘traversed’ 12.1 km per day. Especially noteworthy is the work on selecting
silver foxes (V. vulpes) for nonaggressive behavior towards man (i.e. tameness)
which began in 1959 at the Institute of Cytology and Genetics in Novosibirsk,
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46 Chapter 7
Siberia (Belyaev, 1979). This selection has resulted in a line of foxes that show little
fear of people and resemble domestic dogs in their behavior (Trut, 1999; Fig. 7.1).
The unselected control population still exhibits wild-type behavior, including
strong defensive responses toward humans.
Domestic farm animals artificially selected for production traits have shown
remarkable gains in recent decades (see review by Rauw et al., 1998). Growth
rate of broiler chickens (G. domesticus) has increased from 10 g day−1 in 1960 to
45 g day−1 in 1996. Growth in Norwegian meat-type pigs (S. scrofa) has increased
from 629 g day−1 in 1960 to 962 g day−1 in 1996 due to selection. Milk yield of
dairy cows (Bos taurus) has typically doubled since the early 1950s.
Table 7.1. Mean (± SE) total number of mounts and completed matings
exhibited by male chickens after 20 generations of artificial selection for high
and low levels of sexual performance.a (From Bernon and Siegel, 1983.)
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Artificial Selection 47
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48 Chapter 7
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Artificial Selection 49
Fig. 7.2. Selection for high and low levels of backfat thickness in Duroc pigs
over 16 generations (Hetzer and Miller, 1972).
are not selecting breeding stock prior to reproduction. Similarly, the environment
of animals is also artificially altered when translocating them from nature to
captivity. Except when man chooses potential breeding partners, differential
reproduction in captivity is the hallmark of a natural selection process.
Conclusions
Artificial selection is the only genetic mechanism unique to the domestication
process. It is almost always effective in producing phenotypic change when
consciously applied. In some respects, artificial selection serves as a showcase for
how natural selection works. It has provided insights on how evolution proceeds
and how genes are expressed. Phenotypic changes can be rapid when artificial
selection is intensely applied. Artificial selection differs from natural selection
by the fact that it is applied prior to reproduction whereas natural selection
is measured after reproduction, by an individual’s contributions to the next
generation. Artificial selection can be a double-edged sword, since it is typically
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50 Chapter 7
applied to one or a few specific traits rather than the entire phenotype. Selection
for specific traits may result in unexpected effects on correlated characters, some
of which may counter breeding objectives or jeopardize animal welfare. It has also
been hypothesized that artificial selection for specific characters may sometimes
break up co-adapted gene complexes (i.e. groups of genes that work well together).
No matter what the outcome, artificial selection studies on captive wild and
domestic animals have proved to be the ‘grand experiment’ that has greatly
enhanced our understanding of individual inheritance.
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