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«A Thesis Presented to The Academic Faculty By Estelle A. Sandhaus In Partial Fulfillment Of the Requirements for the Degree Master of Science in ...»

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A Thesis

Presented to

The Academic Faculty


Estelle A. Sandhaus

In Partial Fulfillment

Of the Requirements for the Degree

Master of Science in Psychology

Georgia Institute of Technology

September, 2004

Variation of Feeding Regimes: Effects on

Giant Panda (Ailuropoda melanoleuca) Behavior

Approved by:

Dr. Terry L. Maple, Advisor School of Psychology Georgia Institute of Technology Dr. Mollie A. Bloomsmith School of Psychology Georgia Institute of Technology Dr. M. Jackson Marr School of Psychology Georgia Institute of Technology Date Approved: September 15, 2004


This thesis is dedicated in loving memory to my grandmother, Eleanor A. O’Gorman, whose unwavering faith will be with me always.


I wish to thank my committee members, Dr.’s Terry Maple, Mollie Bloomsmith, and M. Jackson Marr, for their guidance on this endeavor. I also wish to thank Dr.

Rebecca Snyder for sharing her expertise on giant panda behavior throughout the entire course of this project.

I would like to express my profound gratitude to the directors, researchers, and animal care staff at the Chengdu Research Base of Giant Panda Breeding. I am thankful for having had the rare opportunity to share in your efforts on behalf of the giant panda.





Conservation, Research, and Education in the Zoo 1 The Giant Panda 2 Environmental Enrichment and Psychological Well-Being 4 The Role of Predictability in Animal Welfare 9 Environmental Enrichment for the Giant Panda 12 METHODS 16 Study Site 16 Experiment 1 18 Subjects 18 Procedure 19 Data Collection 22 Data Analysis

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Figure 1 Mean Number of Occurrences of Agonistic Interactions for Each 33 Pair of Females in the Baseline Phase Figure 2 Mean Hourly Rate of Bouts of Stereotypy for Each Experimental 37 Phase.

Figure 3 Mean Hourly Rate of Head-Tosses for Each Experimental Phase. 37

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In zoos, an increased emphasis on conservation, animal welfare, and public education has fueled the drive to create captive environments that encourage the expression of natural patterns of behavior. However, captive environments are inherently less complex and/or more predictable than wild ones (Tudge, 1992). It is not uncommon for a number of abnormal behaviors to arise in environments lacking in complexity and/or predictability (Mason, 1991a, 1991b), which is one issue addressed by the principle of environmental enrichment. The giant panda (Ailuropoda melanoleuca) is a critically endangered species for which conservation and public education efforts are of vital importance. While most of a wild panda’s time is spent foraging, processing, and eating bamboo (Schaller Jinchu, Wenshi, & Jing, 1985), captive giant pandas are typically fed discrete amounts of highly concentrated foods on a fixed schedule (Dierenfeld, Qiu, Mainka, & Liu, 1995). Captive giant pandas in various facilities routinely engage in a number of abnormal behaviors prior to the feedings of these predictable meals. These observations are consistent with the findings that members of many species when fed on fixed schedules exhibit increased arousal and activity just prior to feeding, and this is referred to as feeding anticipatory activity (Mistleberger, 1994). While the animal welfare literature abounds with recommendations to implement more temporally complex feeding schedules, few quantitative assessments of these recommendations have been made.

This study proposed to provide a quantitative analysis of the effects of meal predictability on giant panda behavior. A reversal design was planned in which the animals were to be evaluated as they were fed on their usual predictable schedule, then as

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more predictable schedule. The goal was to allow for the objective evaluation of a management strategy that has been frequently proposed, yet infrequently investigated, in the literature.

Because dietary restrictions were imposed on the female giant pandas (each of which was possibly pregnant at the time of this study), the originally proposed diet manipulations were modified. For the female subjects, the frequency of bamboo feedings was increased (the amount was held constant to baseline levels) in the manipulation phase of the experiment. For the male subjects, the delivery of concentrated meals was, as planned, made more unpredictable in the manipulation phase. Because of housing arrangements, the males were necessarily subject to the increase in bamboo feedings that was arranged for the females.

It was hypothesized that a pattern of feeding anticipatory activity would be present in the 30-minute periods prior to the feeding of concentrated meals. It was expected that rates of stereotypic behavior would be highest in these prefeed periods and that it would be lowest during nonfeeding periods throughout the day. It was also predicted that a more species-appropriate activity budget, with a lower incidence of abnormal and stereotypic behaviors, would be observed in giant pandas when switched to the modified feeding regimes, and that when returned to the less naturalistic regime the activity budget would return to baseline levels.

Across all phases of the study, the females spent significantly more time engaged in door-directed/human-oriented behavior, stereotypic behavior, and non-stereotypic locomotion in the 30-minute periods prior to feeding of a concentrated meal when

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differences were found between study phases for the above-mentioned behaviors of interest, though percentage of time engaged in stereotypy approached statistical significance. We did not find significant differences in behaviors of interest between experimental phases or observation periods in the males’ data. These findings may be attributable in part to the low power inherent in the small sample size. However, some visual trends which may be indicative of increased feeding anticipatory activity were apparent. Thus, it appears that giant pandas, like many other animal species discussed in the literature, are sensitive to periodic feeding regimes. Further study is needed to determine just which modifications to current regimes will be most beneficial to captive giant pandas. Methodology of the current study is examined from an applied perspective with the goal of aiding future research.

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Conservation, Research, and Education in the Zoo Modern zoos have become increasingly involved with wildlife conservation and public education efforts. For instance, the American Zoo and Aquarium Association (AZA) now describes itself as a “professional organization dedicated to the advancement of North American zoos and aquariums through conservation, education, scientific studies, and recreation.” The mission statement of Zoo Atlanta expresses similar sentiments: “...to exhibit, interpret, study and care for wildlife in superior environments, to conserve biodiversity throughout the world, to educate, entertain, and enlighten the public....” This increased emphasis on conservation and education of late has fueled the drive to create captive environments that encourage the expression of natural patterns of behavior. In terms of education, this is significant in that the public may derive little educational benefit from the study of animals that do not behave similarly to their wild counterparts. The implications for conservation, too, are great. Zoo animals “...must be encouraged to retain enough of their natural behaviour to make it possible for them to go back to the wilderness; or enough at least of their native wit to enable them to relearn the necessary skills” (p. 193, Tudge, 1992). The ultimate challenge before zoos, then, is to provide “...sufficiently rich environments to allow the performance and maintenance of the species-typical behaviors necessary for survival in the wild” (Shepherdson, 1988).

This task is inherently difficult; captive environments rarely match the wilderness in complexity and unpredictability (Tudge, 1992). To do this effectively, we must objectively evaluate the strategies that we implement. Maple and Finlay (1989)

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We maintain that behavioral scientists are obliged to play an active movement to improve captive environments. Solutions to the problems of animal housing and husbandry must be cost effective, and we should not waste our time and resources on techniques that have not been objectively evaluated. Further progress depends upon a sustained program of applied research. (p. 102).

The Giant Panda The giant panda (Ailuropoda melanoleuca) is an animal for which issues of conservation and public education hold great urgency. While the giant panda has long held significance in Chinese culture (Schaller, Jinchu, Wenshi, & Jing, 1985), it has only recently become recognized worldwide as a precious resource. The giant panda has had significant public exposure as the icon of the World Wildlife Fund, an organization whose self-proclaimed goal is “...to stop, and eventually reverse, the worsening degradation of the planet's natural environment, and build a future in which humans live in harmony with nature.” The round, black-and-white face of the giant panda is all too appropriate a symbol for the struggles of this organization. The giant panda is a critically endangered species; the most recent peer-reviewed estimate indicates that less than one thousand still live in the wild (Tougard, Chaimane, Suteethorn, Triamwichanon, & Jaeger, 1996).

After extensive DNA analysis, it has generally been agreed upon that the giant panda is a member of the bear family, Ursidae (Ledge & Arnason, 1996; Nash, Weinberg, Ferguson-Smith, Menninger, & O’Brien, 1988; Talbot & Shields, 1996;

Waits, Sullivan, O’Brien, & Ward, 1999). Although bamboo comprises more than 99% of its diet, the giant panda has essentially retained the digestive tract of a carnivore: it has

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retain food, and lacks symbiotic microbes to ferment cellulose into available nutrients.

(Schaller et al., 1985). However, the giant panda does possess several morphological traits that facilitate the consumption and digestion of a bamboo diet, such as relatively flat molars and posterior premolars suited to crushing stems, and an enlarged radial sesamoid and sharply curved claws that facilitate the manipulation of bamboo (Schaller et al., 1985). Adaptations to reduce energy expenditure include large body size, a thick coat with oily, springy hairs, and sparing use of calorically expensive activities (Schaller et al., 1985). Giant pandas are largely solitary, and speculations have been made that its bold black and white coloration, which is cryptic only in the snow, helps them to easily spot one another in the forest and thus avoid unwanted contact (Schaller et al., 1995).

Habitat destruction and fragmentation have caused a rapid decline in the giant panda in recent decades by separating a “...once well-integrated giant panda population into many sub-populations of small size.” These small populations are susceptible to a loss of genetic diversity through inbreeding, and are particularly vulnerable to greater reduction in numbers due to further habitat loss and poaching (Zhou & Pan, 1997).

During the periodic synchronous flowering and die-off that is characteristic of many temperate bamboo species, food availability can drop below carrying capacity, resulting in the starvation of giant pandas in the isolated blocks of habitat (Reid, Jinchu, Sai, Wei, & Yan, 1989). Furthermore, individuals suffer from higher mortality rates when attempting to move from one isolated block of habitat to the next (Reid et al., 1989).

Small litter sizes and high infant mortality rates further contribute to the grave status of the giant panda. Giant pandas usually give birth to one or two highly altricial

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part of a comprehensive management plan for this species. Unfortunately, successful captive breeding remains a challenge. Some individuals are reluctant to breed, and there has been some evidence of sub-optimal maternal care in captivity (Gittleman, 1994). One tool that may be used to improve the overall physiological and psychological welfare, and subsequently the reproductive success of captive giant pandas is that of environmental enrichment.

Environmental Enrichment and Psychological Well-Being Environmental enrichment is defined as “...an animal husbandry principle that seeks to enhance the quality of captive animal care by identifying and providing the environmental stimuli necessary for optimal psychological and physiological well-being” (Shepherdson, 1998). While precisely defining the term “psychological well-being” is problematic for both practical and theoretical reasons, several factors are generally considered as indicative of psychological well-being. It should be emphasized that these factors are not necessarily independent of one another.

The absence of stress has been proposed as an indicator of psychological wellbeing (Moberg, 1985, cited by Novak and Suomi, 1988). Certainly, chronic or inappropriately high levels of stress can be detrimental to the physical and mental health of an organism. For instance, prolonged exposure to stress or to the adrenal steroids secreted during stress have been shown to have detrimental effects on the rodent hippocampus, and more recent findings indicate that a similar phenomenon, associated with neuropsychiatric disorders, occurs in the human hippocampus (Sapolsky, 2000).

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