Journal of Comparative Psychology Copyright 1987 by ihe American Psychological Association, Inc.

1987, Vol. 101, No. 3,277-281 07 35-70 36/87/SOO. 75

Applied Dimensions of Comparative Psychology

Greg Moran
University of Western Ontario, London, Canada

Relatively little attention has been paid to potential applications of comparative psychology
during the recent review and reevaluation of the discipline. Yet, problems arising from the
human-animal relationship most often demand solutions that call upon both a knowledge of
naturalistic behavior patterns and familiarity with the principles of learning and behavioral
analysis, a combination uniquely characteristic of comparative psychology. Applications of
comparative psychology are well established in zoos, in attempts to preserve endangered species,
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

small animal veterinary practice, in agriculture, and in animal welfare. Applied comparative
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psychology is likely to establish itself as a valuable adjunct to more traditional activities in all of
these areas.

In the decade prior to this celebration of the one hundredth studying learning processes is likely to be skilled in the latter
volume of the Journal of Comparative Psychology, we have two. However, only the comparative psychologist can offer
seen much debate over the definition of the boundaries of the expertise in all three areas. In the remainder of this article, I
discipline, a kind of academic territorial marking based on will describe some of the problems posed in areas of applied
the word processor rather than pheromones. In the end, most animal behavior and illustrate work already in progress. I
would agree that it matters little if so-and-so's work is classified hope that this review will support the contention that the
as "comparative psychology," "animal psychology" or "eth- comparative psychologist is best suited to fulfill a central role
ology." However, the scholarly endeavor of searching for the in this relatively new and growing field.
content and processes of the discipline has been of great value
to all of us. At the conclusion of his personal search, Dewsbury
Zoos
(1984a) characterized comparative psychology as a subset of
animal psychology, excluding the study of animal learning Essentially all problems associated with the maintenance of
processes and much of physiological psychology. More sub- a variety of species in a zoological collection have a behavioral
stantively, it typically involves the study of the naturalistic dimension. These problems demand a broad perspective and
behavior of species other than the white rat and the pigeon, knowledge base. Issues of housing or aggression will call on
performed in the laboratory or the field, using both experi- an understanding of species-specific behavioral repertoires,
mental and observational methods. This research can be whereas problems in day-to-day management of veterinary
explicitly comparative, but need not be, and is embedded in handling often benefit from the application of learning prin-
an appreciation of evolution and natural selection. ciples. Animal behaviorists have long been applying their
Given these characteristics, there is at least one area for skills in the zoo, most often displaying this unique blend of
which the comparative psychologist seems eminently suited the ethological and psychological that characterizes compar-
and, yet, which has evaded substantial notice in the many ative psychology. An illustration of such an application is
excellent recent reviews of the field (e.g., Burghardt. 1985; provided by the innovative work of Markowitz and his col-
Dewsbury, 1984a, 1984b). This area is applied animal behav- laborators (e.g., Markowitz, 1982; Markowitz & LaForse,
ior, those activities defined by behavioral problems and issues 1987; Markowitz & Woodworth, 1978; Mellen, Stevens, &
arising from the maintenance of animals by humans. This Markowitz, 1981). Markowitz has applied the experimental
husbandry relationship is found in zoological collections, analysis of behavior, more typical of the animal learning
aspects of wildlife conservation, the keeping of pets, and in laboratory, to the enrichment of the captive environment. He
agriculture. The solution of animal behavior problems inev- argues that the essential constraints of captivity preclude the
itably involves the application of three distinct knowledge creation of a natural environment for display animals. In-
bases: first, an understanding of the natural behavior of a stead, he suggests that the principles of operant conditioning
particular species; second, an understanding of the principles and behavioral engineering be used to create an "unnatural"
and technology of animal learning; and finally, expertise in environment that nonetheless provides the animal with an
the experimental analysis of behavior. The ethologist is well opportunity to exercise more of its behavioral repertoire than
qualified in the first of these areas, and the animal psychologist is typically the case in captivity. One of the most recent
applications of these techniques has been with servals (Fells
serval) at the San Francisco zoo (Markowitz & LaForse,
Preparation of this manuscript was supported by Grant A7046
from the Natural Sciences and Engineering Research Council of 1987). A small hairy "rodent" was propelled through a Plex-
Canada to Greg Moran. iglas tube on the floor of the enclosure contingent upon
Correspondence concerning this article should be addressed to exploration of one of two stumps by the servals. If the animals
Greg Moran, Department of Psychology, University of Western On- pursued and pounced on the "prey," they were rewarded by
tario, London, Ontario, Canada .N6A 5C2. the delivery of food. Thus naturally occurring stimuli and

277
 278 GREG MORAN

contingencies of the hunt are replaced with analogs compati- foster mother. The second method can be successful only if
ble with the captive environment. Perhaps the most significant the ontogeny of important species behavioral characteristics
distinction between this approach and others aimed at envi- is relatively insensitive to the changes associated with rearing
ronmental enrichment is that it focuses on the provision of among individuals of a different species. Clearly such factors
natural contingencies between an animal's behavior and a will vary according to species-specific attachment patterns
responsive environment. The institution of this technology and developmental plasticity (Gibbons & Durrani, 1987).
increased the overall activity of the animals and led to the In any case, the potential problems suggest lhal Ihe successful
exhibition of elements of their behavioral repertoire previ- use of embryo transplantation will require systematic inves-
ously unseen in the zoo. At the same time as these techniques tigation for each particular embryo-hosl pair. Such study
promote the welfare of the animals, they provide a more involves expertise in behavioral assessment and management
educational and entertaining display. Similar approaches to as well as a thorough knowledge of species-specific behavior.
behavioral enrichment have been demonstrated successfully
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with a wide variety of species, ranging from gibbons through

Domestic Pets
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harbor seals (Markowitz & Woodworth, 1978) to Indian

elephants (Mellen et al., 1981). Our image of the "pet psychologist" too often comes from
Animal behaviorists have made other less immediately television talk shows and, as a result, lends towards lhal of a
visible but equally important contributions to display design charlatan oul to make the most of neurotic preoccupations
and assessment (e.g., Maple & Finlay, 1987; Myers, 1978; and dependencies. However, Ihe solution of behavioral prob-
Schassburger, 1987) and general applied research (e.g., Ra- lems with pets can be importanl to both Ihe pel owner and
paport & King, 1987; Schilder & Boer, 1987). The mutual Ihe welfare of the animal itself. A few comparative psycholo-
benefits of a general behavioral research and teaching program gists have demonstrated in recent years lhal small animal
as a valuable adjunct to more traditional zoo functions have behavior iherapy can be approached successfully and wilh
been reviewed extensively elsewhere (see Eisenberg, 1975; professional integrity. The sludy of applied animal behavior
Maple, 1981; Moran & Sorensen, 1984/1985; Rumbaugh, wilh particular relevance lo clinical behavior problems wilh
1971, 1972). household pels is becoming an accepted component of vet-
erinary education and research. Perhaps Ihe foremosl repre-
Reproduction and Reintroduction of Endangered senlalives of this approach are Benjamin and Lynette Hart
Species (Hart, 1985; Hart & Hart, 1985). Here, once again, Ihe
practice of Ihe animal behavior therapist is marked by a
Another quite different application, most often also occur- synlhesis of behavioral analysis and a knowledge of Ihe ani-
ring in the zoo, is the attempt to promote the preservation of mal's naturalistic patterns of behavior. For example, Hart and
endangered species (Conway, 1980) by reproduction in cap- Hart's (1985) behavioral assessment of an animal patient
tive populations. Criticism of such efforts often focus on includes Ihe following sleps: an analysis of Ihe context in
claims that reproduction in relatively small, fixed populations which the problem behavior occurs and of situations where,
ultimately are doomed to failure because of the deleterious in conlrasl, Ihe animal's behavior is appropriate; an explora-
effects of inbreeding. In a thoughtful review, Ricker, Skoog, tion of the animal's early history and genetics; galhering
and Hirsch (1987) acknowledge this possibility, but argue information on Ihe clienl-pet interaction; and direcl obser-
that the loss of genetic variance in small populations is not vations of Ihe animal's behavior.
inevitable. Their behavior-genetic arguments also stress that The behavior therapeutic assessment and Irealment ap-
the effect of captivity on individual development is likely to proaches lo aggression in dogs further illuslrales Ihe applica-
vary greatly across species. In many species, deviations in the tion of behavior modification techniques wilhin Ihe conlext
behavior of captive populations from their wild counterparts of the animal's naturalistic behavior patients. Hart and Hart
may be inconsequential or a product of altered eliciting (1985) have elaborated upon an elhological classification to
conditions rather than of qualitative developmental change distinguish patterns of aggression according to their causal
(see Price, 1984). These considerations are central to any basis. For aggression against humans, categories include dom-
plans to reintroduce captive animals into natural habitats and inance-related attacks, competitive aggression towards chil-
underline the importance of behavioral-genetic involvement dren, fear-related, pain-induced, and territorial aggression,
in the solution of such problems. and idiopalhic attacks. Assessment are based bolh on the
A related application concerns the use of embryo transfer conlexl and nalure of Ihe aggressive pattern. Thus fear-related
techniques with endangered species. In recent years, scientists aggression includes those attacks followed by attempts at
at zoos have begun to use females of related species as the escape and featuring the facial and body postures character-
recipients of the embryos of exotic animals (see Gibbons & istic of fear and submission in all canids. Treatment ap-
Durrani, 1987, for a review). Many potential problems proaches often include ethologically based interventions. Hart
remain even once the issues of reproductive physiology have and Hart (1985) recommend that overly timid owners estab-
been addressed. After successful transfer and birth, two alter- lish clear physical dominance over their pet to conlrol domi-
natives are open to the production of competent adults of real nance-related attacks. However, in almosl all instances such
value in wildlife preservation. Such individuals might be interventions are supplemented with learning-based regimens.
integrated into a group of conspecifics or, alternatively, left to In the case of competitive aggression against children, clients
be reared in the altered environment offered by a host mother. are counselled to deferentially provide attention, affection,
The first option raises issues of acceptance by a conspecific and even food, only when the children are presenl. Here Ihe
 APPLIED DIMENSIONS OF COMPARATIVE PSYCHOLOGY 279

child is presented as a discriminative stimulus signaling re- goals of humans in their relationship with captive animals. It
ward in the hope that this association will diminish aggressive is appropriate then that the final area of application discussed
tendencies. The potential for further applications in small concern the welfare of the animals themselves. There is now
animal veterinary practice is great, indeed. Systematic applied a broad recognition of the central role that the animal behav-
research, including rigorous treatment outcome studies, is iorist can play in the debate surrounding the ethical use of
particularly called for in the early stages of this field. animals (see, for example, Dawkins, 1980; Fraser, 1985; Hart
& Hart, 1985; Macdonald & Dawkins, 1981; Moran, 1987).
There is a pressing need for the development and application
Agriculture of techniques to assess animal suffering and for intervention
to minimize such suffering. Once again, the comparative
The term "applied animal behavior" may have first been
psychologist seems best suited to this twofold task.
used to describe the application of the principles of animal
Surprisingly, some animal welfare advocates have argued
behavior to agriculture. Such applications continue to be most
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

that the application of animal behavior science can only

prominent (see, for example, the majority of articles in Ap-
This document is copyrighted by the American Psychological Association or one of its allied publishers.

further contribute to the abuse of animals (see for example,

plied Animal Behaviour Science and two recent major reviews
Fox, 1986). Psychologists, in fact, have received the particular
by Craig, 1981, and Hart & Hart, 1985). The emphasis in
attention of the more radical animal rights groups (see review
agriculture has been on the relation of an animal's naturalistic
and response in Gallup & Suarez, 1985). However, it should
patterns of behavior to the conditions of husbandry with
be clear from even the very brief review included here that
relatively little application of learning-based interventions. In
the applications of comparative psychology have the potential
areas including reproduction, feeding, and housing, animal
to significantly enhance the animals' circumstance and to
behaviorists have used controlled observational studies and a
provide a systematic basis for making judgment regarding
knowledge of the domestic species' natural patterns of behav-
their welfare. A few brief examples should serve to make the
ior to optimize conditions of production. For example, several
point.
studies have focused on the behavioral signs of estrous and
Dawkins (1980) reviews several sources of information that
their evocation (e.g., Veeckman & Odberg, 1978). These signs
have been used as criteria for the assessment of animal suffer-
are important to promote optimal reproduction, particularly
ing. One is the supposed deprivation that any animal under-
where expensive techniques of artificial insemination are used.
goes in captivity. Although her review acknowledges that
Other studies have shown that manipulations aimed at in-
conditions of captivity often can result in serious suffering,
creasing production such as the early weaning of piglets can
she also cites several systematic studies of animals in the wild
result in deleterious side effects if the behavioral limits of the
that suggest that the zoo or farm resident does not always
normal sow-piglet relationship are surpassed (Fraser, 1978).
come out on the short end of the stick. For example, if
More explicitly experimental procedures also have been
longevity is used as a standard, the average robin or song
employed to examine the effects on behavior and physiology
sparrow might live 11 years in the captivity, but only 1 or 2
of different husbandry practices. Hemsworth and his col-
years in the wild (Lack, 1954, cited in Dawkins, 1980). The
leagues, for example, have conducted a series of experiments
central point is that a rigorous study of animal life in the wild
examining the effect of differing conditions of human inter-
reveals a picture of a great deal of hardship and suffering
action on factors important to the farming of pigs. Their work
rather than of romantic peace and harmony. As Darwin
suggests that the behavioral characteristics of an approach to
(1965) put it in a letter to J. D. Hooker: "What a book a
a pig by a human can significantly affect the animal's reaction
devil's chaplain might write on the clumsy, wasteful, blunder-
(Hemsworth, Gonyou, & Dziuk, 1986). Furthermore, the
ing, low and horribly cruel works of nature!" The natural
quality of early contact with humans appears to have a lasting
world should not be used to justify cruelty or abuse of animals,
effect on subsequent behavioral reactions to humans and on
but neither does it serve as a reliable basis for the argument
the animals reproduction and chronic stress level (Hems-
that all captivity is necessarily cruel by comparison.
worth, Barnett, & Hansen, 1986; Hemsworth, Barnett,
Comparative psychology can make important contribu-
Hansen, & Gonyou, 1986). Clearly all of these relations have
tions to the empirical assessment of animal suffering. Dawkins
important implications for farm practices.
(1977) pioneered the use of choice tests for the assessment of
Applied behavioral research also has an important role to
caging preferences in chickens. Some of her recent research
play in attempts at using genetic and hormonal manipulations
clearly demonstrates that some operant responses can be
to increase productivity (see Craig, 1981; Hart & Hart, 1985).
inadequate assays of motivation and preference (Dawkins &
Such manipulations are most often designed to achieve met-
Beardsley, 1986). She has made a preliminary application of
abolic changes that are of direct benefit to the economy of
consumer demand theory to the assessment of "ethological
production, for example, increased rate of growth. However,
need" in making decisions regarding animal welfare (Dawk-
these benefits may have unforseen behavioral side effects on
ins, 1983). Her preliminary results throw some doubt on the
characteristics such as social competence, which will provide
notion that hens might regard access to litter as a "need" in
new problems for the farmer.
anything like the sense that food or water might be. However,
she is quick to stress the pressing need for further empirical
Ethics and the Keeping of Animals research into the behavior of animals in the often severely
restricted environment of agriculture. Such research promises
In each of the areas discussed, I have stressed primarily the to provide something beyond human inferences regarding the
role that comparative psychology can play in furthering the personal preferences and stresses of captive animals.
 280 GREG MORAN

Finally, some of the examples cited earlier in this article search in zoos and acauariums (pp. 12-18). Washington, DC:
argue that the comparative psychologist can do far more than National Academy of Sciences.
Fox, M. W. (1986, June). The trouble with zoos. The Animal's
simply assess the stressful aspects of the relationship between
Agenda, 8-12.
man and animals. The work of Markowitz, for example, is
Fraser, D. (1978). Observations of the behavioural development of
clearly illustrative of how environments can be significantly
suckling and early-weaned piglets during the first six weeks alter
enhanced by manipulation based on a thorough knowledge birth. Animal Behaviour, 26, 22-30.
of both natural patterns of behavior and the principles of the Fraser, A. F. (1985). Social animal science. Applied Animal Behaviour
control of behavior. The same is true in agriculture, where Science, 14, 1-10.
work such as that of Dawkins might usefully form the basis Gallup, G. G., & Suarez, S. D. (1985). Alternatives to the use of
of the design of caging that strikes a balance between concern animals in psychological research. American Psychologist, 40,
for animal welfare and the needs of production. 1104-1111.
Gibbons, E. F., Jr., & Durrani, B. S. (1987). The behavioural devel-
Summary opment of offspring from inteispecies embryo transfer: Theoretical
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

issues. Applied Animal Behaviour Science, 18, 105-118.

This document is copyrighted by the American Psychological Association or one of its allied publishers.

Comparative psychology has traditionally been concerned Hart, B. L. (1985). The behavior of domestic animals. New York: W.
with furthering our understanding of the basic mechanisms H. Freeman.
Hart, B. L., & Hart, L. A. (1985). Canine and feline behavior therapy.
of behavior. For many, a movement to solving practical
Philadelphia: Lea and Febiger.
problems arising in the widespread relationships between
Hemsworth, P. H., Barnett J. L., & Hansen, C. (1986). The influence
humans and animals may have little appeal. However, for
of handling by humans on the behaviour, reproduction and corti-
those who feel otherwise, there is much room for growth. costeroids of male and female pigs. Applied Animal Behaviour
Furthermore, although I have emphasized the pragmatic Science, 15, 303-314.
problem-solving aspects of the application here, the applied Hemsworth, P. H., Barnett, J. L., Hansen, C., & Gonyou, H. W.
comparative psychologist need not totally divorce him or (1986). The influence of early contact with humans on subsequent
herself from basic research. Captive animals can provide an behavioural response of pigs to humans. Applied Animal Behaviour
excellent basis for the study of questions of traditional interest Science, 15, 55-64.
to the psychologist and ethologist (see Moran & Sorensen, Hemsworth, P. H., Gonyou, H. W., & Dziuk, P. J. (1986). Human
communication with pigs: The behavioural response of pigs to
1984/1985; Rickeretal., 1987).
specific human signals. Applied Animal Behaviour Science, 15,45-
The potential for the growth of applied comparative psy-
54.
chology has interesting parallels for those of us who have
Macdonald, D., & Dawkins, M. S. (1981). Ethology—The science
watched the spreading inclusion of the psychologist into areas and the tool. In D. Sperlinger (Ed.), Animals in research (pp. 203-
of human application such as business, education, and med- 223). New York: John Wiley & Sons.
icine. The principle involved is the same in both cases. Where Maple, T. L. (1981, August/September). A zoo story: Confessions of
animals are involved, be they human or otherwise, we neglect a zoo psychologist. APA Monitor, 20.
behavior at our peril. Maple, T. L., & Finlay, T. W. (1987). Postoccupancy evaluation in
the zoo. Applied Animal Behaviour Science, 18, 5-18.
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Ricker, J. P., Skoog, L. A., & Hirsch, J. (1987). Domestication and Schilder, M. B. H., & Boer, P. L. (1987). Ethological investigations
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This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
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Call for Nominations

The Publications and Communications Board has opened nominations for the editorship of
the Journal of Experimental Psychology: General for the years 1990-1995. Sam Glucksberg
is the incumbent editor. Candidates must be members of APA and should be available to
start receiving manuscripts in early 1989 to prepare for issues published in 1990. Please note
that the P&C Board encourages more participation by women and ethnic minority men and
women in the publication process, and would particularly welcome such nominees. To
nominate candidates, prepare a statement of one page or less in support of each candidate.
Submit nominations no later than February 15, 1988 to

Donald J. Foss
Department of Psychology
University of Texas
Austin, Texas 78712