Behavioral development in animals undergoing domestication

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Abstract

The process of domestication involves adaptation, usually to a captive environment. Domestication is attained by some combination of genetic changes occurring over generations and developmental mechanisms (e.g., physical maturation, learning) triggered by recurring environmental events or management practices in captivity that influence specific biological traits. The transition from free-living to captive status is often accompanied by changes in availability and/or accessibility of shelter, space, food and water, and by changes in predation and the social environment. These changes set the stage for the development of the domestic phenotype. Behavioral development in animals undergoing domestication is characterized by changes in the quantitative rather than qualitative nature of responses. The hypothesized loss of certain behavior patterns under domestication can usually be explained by the heightening of response thresholds. Increases in response frequency accompanying domestication can often be explained by atypical rates of exposure to certain forms of perceptual and locomotor stimulation. Genetic changes influencing the development of the domestic phenotype result from inbreeding, genetic drift, artificial selection, natural selection in captivity, and relaxed selection. Experiential contributions to the domestic phenotype include the presence or absence of key stimuli, changes in intraspecific aggressive interactions and interactions with humans. Man's role as a buffer between the animal and its environment is also believed to have an important effect on the development of the domestic phenotype. The domestication process has frequently reduced the sensitivity of animals to changes in their environment, perhaps the single-most important change accompanying domestication. It has also resulted in modified rates of behavioral and physical development. Interest in breeding animals in captivity for release in nature has flourished in recent decades. The capacity of domestic animals to survive and reproduce in nature may depend on the extent to which the gene pool of the population has been altered during the domestication process and flexibility in behavioral development. “Natural” gene pools should be protected when breeding wild animals in captivity for the purpose of reestablishing free-living natural populations. In some cases, captive-reared animals must be conditioned to live in nature prior to their release.

Introduction

Animal domestication is best viewed as a process, more specifically, the process by which captive animals adapt to man and the environment he provides. Since domestication implies change, it is expected that the phenotype of the domesticated animal will differ from the phenotype of its wild counterparts. Adaptation to the captive environment is achieved through genetic changes occurring over generations, and environmental stimulation and experiences during an animal's lifetime (Price, 1984). In this sense, domestication can be viewed as both an evolutionary process and a developmental phenomenon.

This review constitutes a summary of the literature dealing with the effects of domestication on the behavior of captive animals. The emphasis in this article is placed on environmentally induced changes in the development of behavior accompanying the domestication of farm, laboratory and companion animals. (See Price, 1998, Darwin, 1868, for a companion review paper focusing on genetic contributions to the domestic phenotype). After defining domestication, the reader is introduced to some of the changes in the animal's environment associated with the transition from nature to captivity. Changes in the quantity and quality of space and shelter available to captive animals, changes in behaviors associated with feeding and drinking, the reduction of predation and changes in the social environment are discussed with respect to their role in behavioral modification and adaptation to the captive environment. While artificial selection and natural selection in captivity facilitate adaptation to the captive environment over generations, the unique experiences of individuals foster adaptation within their lifetime. Environmentally induced changes in the behavior of domestic animals are reflected in the role of key stimuli, changes in intraspecific aggression, interactions with humans, and responsiveness to environmental change as well as in rates of development including neoteny. The review concludes with a short treatment of feralization and concerns associated with rearing animals in captivity for release in nature.

Section snippets

Domestication defined

Darwin (1859), Darwin (1868)suggested that domestication is more than taming, that it includes breeding animals in captivity, is goal-oriented, may occur without conscious effort on the part of man, increases fecundity, may bring about the atrophy of certain body organs, enables animals to achieve greater plasticity, and is facilitated by subjugation to man, the domesticator. Some contemporary definitions postulate that domestication is a condition in which the breeding, care and feeding of

The transition from nature to captivity

Although there are many reported differences between wild and domestic stocks, there is little evidence that domestication has resulted in the loss of behaviors from the species repertoire or that the basic structure of the motor patterns for such behaviors has been changed (Scott and Fuller, 1965; Hale, 1969; Miller, 1977). In nearly all cases, behavioral differences between wild and domestic stocks are quantitative in character and best explained by differences in response thresholds.

Genetic mechanisms

The genetic phenomena with the greatest potential impact on the domestication process are inbreeding, genetic drift, and selection (Price, 1998). Whereas inbreeding and genetic drift produce random changes in gene frequencies, the changes resulting from selection are directional. Price and King (1968)proposed three primary selective phenomena that influence populations of animals undergoing domestication: (1) artificial selection, (2) natural selection in captivity, and (3) relaxation of

Presence or absence of key stimuli

The absence of certain “key” stimuli in the physical environment of captive animals can result in a failure to express certain behavioral patterns. Whereas burrows constructed by wild and domestic Norway rats are similar in every respect, Boice (1977)noted that domestic Norway rats seldom initiate burrowing without an object (e.g., stone) to dig under. Wild rats were less dependent upon such stimulation. Similarly, Huck and Price (1976)reported that a wild stock of Norway rats would exhibit

Feralization as a developmental phenomenon

As in the case of domestication, the process of feralization has meant different things to different people. Some definitions assert that feral animals are merely free-living populations of animals that originated from domestic stock (Pullar, 1950; Shank, 1972). Others suggest that in addition to their free-ranging status, feral animals must be unowned, not intentionally cared for by humans, and not dependent on humans for breeding (McKnight, 1976; Baker and Manwell, 1981). Those with a more

Conclusions

Development of the domestic phenotype is nurtured by the interplay of genetic changes occurring over generations and the experiences of captive animals during their ontogeny. The process of domestication is realized when the gene pool is sufficiently altered and when specific captive environments and management techniques are consistently applied over generations to produce a phenotype that fosters adaptation to the captive environment and human intervention.

Behavioral changes accompanying the

Acknowledgements

I am greatly indebted to the many students and colleagues (too numerous to name) who have contributed either directly or indirectly to the ideas contained in this paper. Parts of this article were published in Price, 1984, Price, 1998.

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