Chapter 16
General Considerations
Some domestic animals are typically provided with an environment physically
similar to the habitat of their wild ancestors. Behavioral and physiological adapta-
tions to such an environment will be readily achieved. Very often, however, the
captive environment does not match the ancestral environment and adaptation is
challenged. Bartlett (1985) has identified some of the physical characteristics of
laboratory environments that compromise the ability of insects to survive and
reproduce in captivity.
There is a tendency for animal caretakers to establish captive environments
that are believed to support the least adaptive individuals in the population yet
still convenient and efficient to maintain (Bartlett, 1993). It is not uncommon
for the living space of captive animals to be smaller and less complex than that of
their free-living counterparts. In addition, there is typically less temporal variation
in the physical components of captive environments. Environmental enrichment
is often used to compensate for biologically important deficiencies and to
make captive environments more compatible with species-specific behaviors
and requirements. The term ‘enrichment’ is typically applied to ‘increasing the
physical, social and temporal complexity of captive environments’ (Carlstead and
Shepherdson, 2000) rather than some measure of the outcome of the change on
the animals involved (Newberry, 1995). Changes in physical aspects of the captive
environment do not necessarily improve adaptation. Captive animals may
actively investigate novel objects when first placed in their living space but do not
always retain interest in these objects (Vignes et al., 2001). Enrichment devices
should be biologically relevant to the species and designed to achieve specific goals
(Newberry, 1995). Feeding devices which require animals to work for their food
provide them with perceptual and locomotor stimulation, which may or may not
be similar to that required for procuring food in nature. Nevertheless, the stimula-
tion obtained probably helps to promote normal development and good welfare.
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Climate – Temperature
The wild ancestors of most domesticated animals lived in temperate regions of the
world (Ucko and Dimbleby, 1969) and typically possessed a variety of adaptations
to natural temperature fluctuations. Many wild animals escape temperature
extremes by burrowing underground, hibernating, estivating, migrating or
otherwise seeking out favorable microhabitats. Expression of such temperature-
regulating behaviors is sometimes prevented by the captive environment (Geiser
et al., 1990) and other behavioral and physiological adjustments have to be
employed (Randolph et al., 1990). In contrast, captive animals are sometimes
housed in climate-controlled facilities, where temperature and humidity are
far more uniform than ever experienced in nature. Constant temperature in
the laboratory is responsible for a number of reports of delayed sexual maturation
in migratory locusts (Locusta migratoria) exposed to long-day photoperiods and
reared under crowded conditions (Hasegawa and Tanaka, 1996). Crowded
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144 Chapter 16
female locusts exposed to cool nights (15°C) and warm days (30°C) during the first
two nymphal instars exhibited significantly earlier sexual maturation (i.e. age at
first oviposition) than locusts maintained at a constant 30°C. Most of the experi-
mental group had started oviposition by 30 days, whereas a third of the control
group had not oviposited by 50 days. The suppression of sexual maturation often
reported from laboratory studies may not necessarily occur in the field; free-living
locusts may experience low temperatures during the early nymphal instars.
Initial attempts to spawn white sturgeon (Acipenser transmontanus) broodstock
maintained at constant 15 and 18°C water temperature were largely unsuccessful
due to ovarian regression. Normal ovarian development can be obtained in the
majority of females with intact ovarian follicles by transferring them to cold water
(12 ± 1°C) in the fall or early spring (Webb et al., 2001). There appears to be a
temperature-sensitive phase in ovarian development during the transition from
vitellogenic growth to oocyte maturation and the degree and timing of sensitivity
to environmental temperature are dependent on the female’s endogenous
reproductive rhythm.
Constant temperature conditions may affect the rate of genetic changes
resulting from propagating animal populations in captivity, particularly poikilo-
thermic species (Bartlett, 1984). Economopoulos and Loukas (1986) monitored
genetic changes in allele frequencies at the alcohol dehydrogenase locus of olive
fruit flies (D. oleae) in response to being reared with an artificial larval medium
instead of olives under constant temperature regimens of either 17 or 25°C or a
daily fluctuating temperature ranging from 17 to 25°C. The magnitude of genetic
changes was the same for all three temperature regimens, but the changes
occurred more rapidly at the stable temperatures, particularly at 25°C, which
favored rapid development and high colony productivity.
There is some evidence that domestication may have reduced the capacity of
animals to adapt to extremes in temperature. Richardson et al. (1994) compared
the offspring of wild-caught house mice (M. domesticus) with domestic house mice of
the random-bred ICR strain and their reciprocal hybrids for their ability to
exhibit nonshivering thermogenesis (NST) in a cold environment, a process that
enables small mammals to rapidly increase heat production during cold exposure.
They also examined the mice for depositions of brown adipose tissue, an adapta-
tion to living in a cold environment. Wild and hybrid mice together had higher
(+18.2%) regulatory NST (maximal NST minus basal metabolic rate) and larger
(+21.2%) deposits of interscapular brown adipose tissue than the laboratory
strain suggesting that, during domestication, the laboratory strains may have
experienced relaxed selection for thermoregulatory abilities since intense cold
never occurs in the laboratory environment. Foley et al. (1971) demonstrated that
neonatal feral (wild) hogs (S. scrofa) are more resistant to extreme cold than
domestic neonates because of greater heat production and more pelage. Vincent
(1960) reported that wild-genotype brook trout (S. trutta) could endure higher
water temperatures than domestic stock even though reared under the same
conditions. Domestic farm animals living outdoors seek protection from cold and
wind by using natural vegetation or various types of shelter provided by humans.
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Light
Dou et al. (2000) found that light is the primary factor regulating the feeding
activity of captive-reared Japanese flounder larvae, P. olivaceus, an important fish
for stock enhancement and sea ranching in Asia (Japan, China and Korea).
Tank-reared flounder show a major feeding peak in the morning and a secondary
peak in the afternoon throughout larval development with little or no feeding in
darkness. Maximum feeding rates occurred at 19°C.
Day Length
Day length can affect growth, reproduction and a host of other biological
characteristics. Endal et al. (2000) and others have reported that adding additional
light enhances the growth rate of Atlantic salmon (S. salar) reared in sea cages.
Additional light can also increase or decrease the rate of sexual development of
salmon depending on timing relative to a critical period of development (see
review by Endal et al., 2000).
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146 Chapter 16
Ultraviolet-light Exposure
Lack of sunlight or ultraviolet-light exposure in captivity can pose special prob-
lems for certain species. Gillespie et al. (2000) discuss the deficiencies in vitamin D3
found in Komodo dragons (Varanus komodoensis) kept in captive, indoor exhibits vs.
those with daily ultraviolet-B exposure.
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Conclusions
Many captive wild and domestic animals are housed under constant temperature
(and humidity). Although unnatural, constant temperature does not constitute
a problem for most homeothermic animals providing the temperature is
not extreme. In contrast, constant temperature may be very important for
cold-blooded (poikilothermic) animals that depend on temperature fluctuations
for normal reproduction and maturation. Constant day length is a common
feature of many captive environments and may be a key factor in explaining
reproductive failure in captive populations of seasonally breeding species.
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148 Chapter 16
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