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Chapter 16

Climate and Shelter

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.

©CAB International 2002. Animal Domestication and Behavior

142 (E.O. Price)

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Climate and Shelter 143

Mellen and Sevenich MacPhee (2001) propose a more holistic approach to

environmental enrichment in which enrichment is re-defined for each species and
perhaps each individual. They propose that enrichment should be based on an
animal’s natural and individual histories and the constraints of its living environ-
ment and should include defining the physical and social environments of the
animal, the role of human caretakers (feeding, cleaning, training, etc.) and diet
(type, presentation, variety). It is more than simply adding something to the
animal’s environment that is believed to improve its welfare.
Wemelsfelder (1997) makes the point that the welfare of animals in captivity
may depend on how environmental stimuli are perceived from a functional
standpoint. The specific stimulus properties of an environment in terms of
variability and complexity may not be as important as what they afford the animal
to do. For example, water can be a cool bath, a life-saving drink or a threat to the
animal’s safety. What is perceived is not merely water, but the functional context
of water. Likewise, housing systems can be designed which allow animals to be
functionally creative. This may include the use of artificial objects or devices
which the animal can manipulate and control (Markowitz and Line, 1989).
The ultimate challenge is to create an environment which retains its functional
properties over time, one which does not lose its effectiveness.
The next two chapters provide examples of how the physical environment of
captive wild and domestic animals differs from that of their free-living counter-
parts in nature. These reports also include techniques that have been developed
or are currently being researched to provide more functionally relevant living
environments for captive animals. This chapter begins with a few examples of how
temperature, light and day length influence the maturation, reproduction and
behavior of captive animals. The adaptive significance of shelter is then discussed,
pointing out the possible role of shelter (hiding places) in the development of
avoidance responses of animals toward humans and even maturation rates.

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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Climate and Shelter 145

Protection against extreme heat is more difficult, however, since avoidance of

the direct rays of the sun offers only partial relief. Hansen and Jeppesen (2001a)
propose that access to water for swimming can assist farmed mink (M. vison)
in thermoregulating in response to very high ambient temperatures. Thermo-
regulation is less of a problem for domestic stock tolerant of high temperatures
(Macfarlane, 1968).
Providing the optimal temperature, humidity and photoperiod can be
critical for the survival and reproduction of animals in captivity. Milonas and
Savopoulou-Soultani (2000) studied the effect of (laboratory) temperature on
the development, fecundity and longevity of Colpoclypeus florus, a parasitoid of
Adoxophyes orana, an economically important pest of peach, apple and cherry
orchards in Europe. C. florus were reared at constant temperatures of 15, 17, 20,
25 and 30°C with a photoperiod of 16:8 (L:D) hours. The rate of development
from egg to adult increased as a linear function of temperature from 15 to 30°C
(31.7 ± 0.4 vs. 12.8 ± 0.2 days, respectively) but no adults emerged at 30°C.
Fecundity (total number of eggs per female) was greatest at 17°C (57.4 ± 5.1) and
least at 15°C (30.7 ± 6.2), only 2° lower. The average number of eggs oviposited
per female per day was greatest at 25°C (9.3 ± 0.9) and least at 15°C (2.8 ± 0.3).
Longevity of adult females (days) was greatest at 17°C (11.0 ± 1.2) but not
significantly different from 15 and 20°C. Longevity was least at 25°C (4.5 ± 0.4
days). Studies such as this one highlight the importance of temperature in rearing
invertebrate species such as C. florus in captivity to gain information on their
potential geographical distribution, population dynamics or for their mass
production for field release as a means of biological control of pest species.

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

Reproduction in free-living animal populations is often synchronized to

changing day-length. Constant additional light together with good nutrition in the
laboratory can facilitate year-round breeding by many rodent species that breed
seasonally in nature (Blus, 1971). Constant light–dark cycles may also be a con-
tributing factor to the reproductive failure of many captive animal populations.
Price (1966) observed a sevenfold increase in the reproductive success of captive
woodland deermice (Peromyscus maniculatus gracilis), approximately 15 generations
removed from the wild, by delaying the offset of the daily light cycle in the labora-
tory 1 h per week for 3 successive weeks. Kujawa et al. (1999) described a system
that controls photoperiod and water temperature for keeping fish prior to artificial
spawning. It has been successfully used with a number of commercial and wild
European cyrinid and percid fish species, and can be used to alter spawning time
relative to the natural environment. More opportunistic species are not as affected
by photoperiod and reproduce whenever they are in a positive energy balance
(Bronson, 1998).

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.

Adaptive Significance of Shelter

In nature, natural cover or shelter provides protection from climatic events,
privacy for mating and rearing of offspring, and a means of escape from predators
and aggressive social partners. An increase in the quantity or quality of cover
permits higher population densities by reducing the incidence of intraspecific
and interspecific contacts (Jenkins, 1961). Shelter may serve some of these same
functions in captivity. Patterson-Kane et al. (2001) studied the cage preferences of
laboratory rats (R. norvegicus) and discovered that paper towels (nesting material)
and a tin nest box (coffee can) were consistently preferred over a larger social
group (three rats), larger cage space, toys, a plastic tunnel and objects that
provided opportunity for chewing (wooden sticks). Bird species that use nest
cavities for reproduction in nature may require nest boxes for successful breeding
in captivity (Millam et al., 1988). Hansen and Damgaard (1991) reported that the
physiological stress level of farmed mink (M. vison) increased when they were
deprived of nest boxes in their cages. They hypothesized that nest boxes shielded
the animals from stress-provoking external stimulation from the animals in
neighboring cages. Clark and Galef (1977) demonstrated that the tameness and
docility of captive gerbils (M. unguiculatus) can be influenced by the physical design
of their cages. Laboratory-reared gerbils given access to an enclosed hiding place

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Climate and Shelter 147

(e.g. burrow system or shelter) during ontogeny exhibited greater avoidance of a

human-like stimulus than gerbils reared in open laboratory cages. Elicitation of
escape responses (flight) into a concealed chamber was found to be the ‘critical’
experience in the development of this avoidance response. Once flight and con-
cealment responses were established, experience in an open-cage environment
had little influence on avoidance behavior. In addition, Clark and Galef (1980,
1981) reported that gerbils reared in standard open cages vs. cages with shelters
exhibited earlier eye-opening, faster growth, earlier sexual maturity, and a
marked decrease in adrenal size, all of which are traits normally associated with
domestication (King and Donaldson, 1929; Richter, 1949; Clark and Price, 1981).
Harri et al. (1998) reported that farmed blue fox cubs (A. lagopus) provided with a
nest box in their cage were more fearful towards humans than foxes without a nest
box. If nest boxes are offered, foxes take advantage of the opportunity to hide and,
as a result, do not habituate to the everyday activities of farm staff and visitors.
These examples illustrate how environmentally induced developmental events
recurring in each generation can play an important role in the development of an
important component of the domestic phenotype (i.e. reduced fearfulness of
humans).
Nikoletseas and Lore (1981) found that domestic Norway rats (R. norvegicus)
reared in cages with burrows were more aggressive toward strange intruders than
rats reared in standard laboratory cages without shelter. Hubrecht et al. (1992) in a
study relating the behavior of kenneled dogs (C. familiaris) to pen size and housing
conditions noted that the provision of a shelter (coop) not only provided a place
for the dogs to rest but was also used in play and served as a refuge from other
dogs. The shelter provided the dogs with some control (i.e. choice) over their
environment. Hansen and Berthelsen (2000) compared the behavior of domestic
rabbits (O. cuniculus) maintained in conventional cages vs. enriched cage systems
with access to shelter. Behaviors in open-field tests and in response to being
captured (timidity) were recorded. Rabbits housed in conventional cages, espe-
cially females, exhibited more restlessness, excessive grooming, bar-gnawing and
timidity than rabbits in the enriched cages. Only a few rabbits used the shelters for
resting; more often they were observed resting on the roof of the shelter.

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

It is not surprising that captive animals actively use shelter if it is provided.

Their wild ancestors may have needed shelter to survive and bear young.
Although shelter can shield animals from potentially threatening stimuli and
events such as contact with humans and other animals, in captivity it can result in
the development of increased fearfulness towards humans and aggression directed
at penmates. The traditional use of open-cage designs for laboratory rodents and
other captive animals may facilitate habituation to humans and human activity,
thus reducing fearfulness and improving animal welfare. Such practices recurring
in each generation can play an important role in the development of the domestic
phenotype. There is some evidence (in gerbils) that the use of shelter may retard
maturation and growth and result in larger adrenal glands. The generality of this
interesting finding should be studied because of its potential importance to the
domestication process and animal welfare.

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