Zoo Biology 17:333341 (1998)

Positive Reinforcement Training to

Enhance the Voluntary Movement of
Group-housed Chimpanzees Within Their
Enclosures
M.A. Bloomsmith,1* A.M. Stone1, and G.E. Laule2
1

Department of Veterinary Sciences, The University of Texas M. D. Anderson Cancer

Center, Science Park, Bastrop, Texas
2
Active Environments, Lompoc, California

Positive reinforcement techniques were applied to train groups of chimpanzees to move voluntarily into the indoor portions of their enclosures at the
request of trainers and to be briefly restricted to those areas. Subjects were
66 members of eight social groups, including 44 adults (14 males, 30 females), and 22 immatures (eight males, 14 females). Performance of individual animals was recorded during four experimental phases of the project:
baseline, initial training, maintenance of reliable performance, and transfer
of responsibility for training from the original trainers to others on staff. A
mean of 16.1 training sessions was required to reach reliable performance,
defined as the subjects complying with 90% of the requests to move indoors. Analyses of variance indicated that chimpanzee compliance was significantly increased after training. Females required significantly fewer
training sessions to reach reliable performance than did males. Adult males
showed the lowest level of compliance in each experimental phase. Overall,
compliance was not affected by the transfer of responsibility for the procedure from the original trainer to other staff, although there was evidence of a
temporary and small decrement in performance immediately following transfer. These findings indicate that training can improve the voluntary movement
of captive chimpanzees. They also demonstrate that animal training can be
objectively evaluated using systematic study design, data collection, and statistical analysis of data. Zoo Biol 17:333341, 1998. 1998 Wiley-Liss, Inc.
Key words: animal training; behavioral management; great ape behavior; operant conditioning; psychological well-being

Received for publication September 10, 1996; revision accepted June 11, 1998.
*Correspondence to: Dr. Mollie A. Bloomsmith, Zoo Atlanta, 800 Cherokee Ave.S.E., Atlanta, GA 30315.

1998 Wiley-Liss, Inc.

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INTRODUCTION

While behavioral scientists in zoological parks and laboratories are studying

many types of environmental changes designed to promote the psychological wellbeing of captive nonhuman primates [Novak and Petto, 1991; Novak and Suomi,
1988; Segal, 1989; Shepherdson et al., 1997], training of animals has not yet been
thoroughly assessed from this perspective. Training is only one form of interaction
between humans and the nonhuman primates they care for, but it may have a powerful impact on well-being [Bloomsmith et al., 1994].
There is growing interest in documenting the time requirements for, and the animals response to, training procedures related to animal care. Through a variety of training techniques, nonhuman primates have been trained to tolerate human handling and
restraint [Aarons, 1973; Heath, 1989], vaginal or rectal swabbing [Bunyak et al., 1982;
Desmond and Laule, 1987; Moseley and Davis, 1989], blood pressure measurement [Segal,
1989; Turkkan, 1990], and the application of topical drugs [Reinhardt, 1990]. They have
been taught to cooperate while undergoing a physical examination [Paciulli, 1990; Smith,
1981]; offer urine, blood, or fecal samples for collection [Bunyak et al., 1982; Hearn,
1977; Kelley and Bramblett, 1981; Moseley and Davis, 1989; Phillipi-Falkenstein and
Clarke, 1992; Priest, 1991; Reinhardt, 1991; Stone et al., 1994; Vertein and Reinhardt,
1989; Walker et al., 1982]; avoid electric fencing [Ogden et al., 1992]; and move into a
restraining device [Hearn, 1977; Knowles et al., 1995; Luttrell et al., 1994; Moseley and
Davis, 1989; Reinhardt, 1990; Smith, 1981; Walker et al., 1982] or portable transport
cage [Clarke et al., 1988; Heath, 1989; Kessel-Davenport and Gutierrez, 1994]. There
are several reports of training designed to address social problems [Bloomsmith et al.,
1994; Desmond et al., 1987; Joines, 1977; Thorpe, 1989; de Waal and Morris, 1982].
At The University of Texas M. D. Anderson Cancer Centers, Science Park chimpanzee breeding facility in Bastrop, Texas, a broad-scale animal training program uses
positive reinforcement techniques in the daily management of the animals. Positive
reinforcement training has been used to elicit the voluntary cooperation of chimpanzees with a number of husbandry, medical, and research procedures as well as to enrich their environment and improve social relationships [see Laule et al., 1992, for
further details on the program].
One focus of our training has been on improving voluntary movement of the
chimpanzees between portions of their enclosures to facilitate husbandry, veterinary
care, and research. Housing nonhuman primates and other animals in groups and in
spacious enclosures often leads to difficulties in obtaining reliable movement of animals when needed. Several publications reported training macaques to move between portions of their enclosures and to enter smaller transport caging or chute
systems for collecting biological samples [Knowles et al., 1995; Luttrell et al., 1994;
Phillippi-Falkenstein and Clarke, 1992; Smith, 1981; Walker et al., 1982]. These
articles described the use of a combination of positive (e.g., offering food rewards)
and negative reinforcement strategies (e.g., using poles, loud noises, or other techniques) to coerce animals into complying. In all cases except one [Smith, 1981],
animal care staff entered the monkeys enclosures as a part of the training process.
In the current study, positive reinforcement techniques were exclusively applied to train groups of chimpanzees living in indoor/outdoor enclosures to move
voluntarily into the indoor portions of their enclosures and to be restricted to those
areas. This report describes the training effort, the quantitative analysis of the time
requirements of the training, and the response of the animals to the procedures.

Training for Voluntary Movement

335

METHODS
Subjects and Housing

Subjects were 66 members of eight social groups of chimpanzees (Pan troglodytes). They included 44 adults (14 males, 30 females) and 22 immatures (eight
males, 14 females). Group size ranged from six to 16 animals. Groups were composed of one to four adult males, three to six adult females, and their offspring.
Subjects 210 years old were considered immature; those older were considered adult.
Groups also contained animals younger than 2 years that were not subjects of this
study. The data on four immature subjects who became adults during the study were
deleted from the analysis once they reached 11 years old. All these subjects completed at least the baseline and initial training phases of the study.
All subjects were housed in indoor/outdoor enclosures at the Science Park chimpanzee facility in Bastrop, Texas. The outdoor portion of each enclosure was octagonal, 22 m in diameter, and contained natural substrates, climbing structures, and
manipulable objects. Chimpanzees had nearly continuous access to the indoor portions of their enclosures (5.8 1.9 m); access was restricted to outdoors only during
periods of cleaning or maintenance of the indoor dens. All subjects participated in a
comprehensive daily environmental enrichment program, and many of the subjects
had prior exposure to the positive reinforcement training program.
Training Method

Positive reinforcement techniques were used exclusively to train all subjects to

enter the indoor portion of their enclosures. A single training attempt was accomplished by opening the guillotine doors separating each indoor and outdoor enclosure, giving the verbal command inside, and waiting for the animals to enter. After
~90 sec, the doors were closed and locked. If chimpanzees were still moving toward
the inside, closing the door was delayed slightly. After the door was locked, subjects
inside were reinforced with a preferred food item (e.g., fruit juice, fruit, peanuts).
The reinforcers were not visible to the animals prior to their delivery. The subjects
were not deprived of food. Those animals that were already inside when the training
sessions began and that remained inside were also reinforced. Animals that did not
cooperate by moving to the indoor enclosure were simply not reinforced; there was
no other consequence to their lack of cooperation. After the cooperating animals
were reinforced, the door was opened. In some cases, multiple attempts were made
immediately following one another; this set of attempts completed sequentially is
referred to hereafter as a training session.
A total of eight people worked with the chimpanzees during this program of
voluntary movement training. Two individuals served sequentially as the primary
trainers; six members of the caregiving staff took over responsibility from the two
trainers for maintaining this behavior after the subjects were performing reliably.
The transfer of responsibility was accomplished by individual caregivers observing
the primary trainer during several training sessions and then being instructed in completing the appropriate records. The caregiver was then observed by the trainer during several sessions as the caregiver completed the process independently and was
given additional instruction as needed. After the transfer of responsibility for the
behavior to the caregiver, the primary trainer occasionally observed the caregiver to
monitor the entire process for consistency.

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Bloomsmith et al.

Study Design and Data Collection

Information was systematically recorded during four experimental phases. In

the first phase, baseline performance was recorded for each group during 15 attempts in which no reinforcement was provided, but animals were called inside. In
the second phase, initial training, training was initiated and continued until the subjects
cooperated with the request to move inside 9 of 10 times. This 90% compliance score
was our operational definition of reliable performance. In the third phase, maintenance,
reliable performance was maintained by one of the two original trainers, so the animals
had met our definition of reliable performance, and the primary trainers were still working with them. In the fourth phase, transfer, responsibility for the procedure was transferred from the original trainers to other people on the caregiving staff.
The data recorded were simply whether each subject entered before the door
was closed. This was recorded for each training attempt. Other information recorded
included the date, time of day, trainer, reinforcer given to the subjects, and sex and
age class of each subject. The frequency of training sessions per week was not controlled, so the duration between training sessions was not consistent.
In most cases, responsibility for performing the procedure was transferred from
the trainer to the caregiving staff members only after all members of a subject group
had reached the 90% reliability level. Four subjects (one representative of each age/
sex class) did not follow this usual pattern. Although each of these animals became
reliable during the course of the study, it was after, rather than before, the transfer of
responsibility to caregiving staff. For practical concerns, we continued with the transfer, even though these individuals had not yet met the criterion performance. Data
for these four subjects were analyzed for the number of training sessions and for
attempts to reach reliable performance, but they were deleted from analysis in the
maintenance and transfer phases of the study because they did not meet the criteria
for those phases (i.e., they were not transferred as reliable performers to the responsibility of another person acting as a trainer).
Statistical Analysis

Analysis of variance (ANOVA) was applied to measure differences in training

performance, with sex and age class as grouping factors. One ANOVA compared the
number of training sessions required to reach reliable performance, and a second
ANOVA compared the number of training attempts required to reach reliable performance. A third ANOVA for repeated measures compared the percentage of compliance scores across the four phases of the study. This analysis included scores for 58
of the 66 subjects; the remaining scores were deleted because of either a change in a
subjects age status or a failure to reach reliability prior to transfer, as described
above. Planned comparisons were then completed to make particular comparisons
among phases of the study. A fourth ANOVA for repeated measures was completed
on a subset of the data containing only the 20 attempts immediately prior to and the
55 attempts immediately after the transfer of responsibility for the procedure from
the original trainer to other members of the caregiving staff.
RESULTS

A total of 1,412 training sessions (range per group, 104342) and 1,835 training attempts (range per group, 126492) were recorded among the eight subject groups

Training for Voluntary Movement

337

(mean per group, 176.5 sessions, 229.4 attempts). A total of 12,702 attempts for
individual animals was included in this analysis. Data from each group were gathered between 26 and 54 months (mean, 36.9). Table 1 gives the number of training
attempts completed during each experimental phase.
A variety of food reinforcers was used, with bananas being used the most often
(39.5% of attempts), followed by fruit juice (18.1%), combinations of two or more
reinforcers (11.6%), and apples (7.1%). Other reinforcers given 2% of the time
included grapes, oranges, raisins, and peanuts.
During the baseline phase, before any training, the subjects cooperated with the
request to move indoors a mean of 66.1% of the time. This score peaked at 91.7% in the
maintenance phase. A mean of 16.1 training sessions (standard error, 17.8; range, 493)
and a mean of 22.0 training attempts (standard error, 28.8; range, 9.0154) were required
to reach reliable performance (i.e., subject complying with 90% of the requests). Since
each training attempt took at most 5 min of personnel time, this amounts to a maximum
of 110 min of personnel time to train a group of chimpanzees in this behavior. Table 1
provides a summary of compliance scores across phases of the study. Individual differences were evident in response to the training process as subjects required between nine
and 154 attempts to reach reliability.
The ANOVA results for the number of training sessions and training attempts
required to reach reliable performance indicated that the subjects performance was
influenced by sex. Females required significantly fewer training sessions (F = 6.0; df
= 1; P < 0.02) than did males (female mean, 11.6; male mean, 25.0). Females also
required fewer training attempts than did males (F = 4.7; df = 1; P = 0.04) (female
mean, 15.4; male mean, 35.2). There were no age effects or age-by-sex interactions.
The percentage of time the chimpanzees complied with the request to move to
the indoor portion of their enclosures was dependent on the phase of the study as
indicated by a significant value for the repeated-measures ANOVA (F = 31.7; df = 3;
P < 0.001). Between-subject effects also indicated a significant interaction between
the sex and age class of the subjects (F = 4.9; df = 1; P = 0.03). There was no
significant phase-by-sex-by-age interaction, indicating that the age-by-sex effect was
consistent among the four phases of the study. These findings are explained by the
finding that the adult males showed the poorest performance in each experimental
phase (see Table 1).
TABLE 1. Compliance Scores Across Study Phases
Experimental Phase

Total number of
attempts for individuals
Adults
Females
Males
Immatures
Females
Males
All subjects

Baseline
(n = 66)

Initial
training
(n = 66)

Maintenance
(n = 58)

Transfer
(n = 58)

942

1,451

2,181

8,128

77.9%
41.6%

88.7%
70.0%

95.8%
84.7%

94.2%
76.4%

65.4%
66.1%
66.1%

81.9%
78.9%
82.1%

87.7%
93.1%
91.7%

87.2%
84.2%
87.9%

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Bloomsmith et al.

Three planned, post hoc comparisons of compliance scores were computed to

allow comparisons of performance between particular phases of interest. First, we
found that chimpanzee compliance was significantly increased after the training as
measured by comparison of the baseline and maintenance phase scores (F = 16.2; df
= 4; P < 0.001). Second, a similar comparison of the baseline and the transfer phase
scores indicated that this increase in compliance was maintained after other members of the animal care staff were again responsible for the procedure (F = 15.7; df =
4; P < 0.001). Third, chimpanzee performance was not affected by the transfer of the
procedure from the original trainer to other members of the animal care staff as
measured by comparison of the maintenance and transfer phase scores (F = 1.6; df =
4; P > 0.05).
Another repeated-measure ANOVA was performed to compare chimpanzee performance immediately before and immediately after the transfer, rather than on data
from the entire phase, to identify whether there may have been a more short-term
change in performance that was not measurable in the overall test. This test revealed
a statistically significant reduction in the level of compliance after transfer (F = 9.1;
df = 1; P = 0.004). The mean compliance score in the tests before transfer was 94.2
versus 89.8% after transfer.
When the data from the maintenance and transfer phases were compiled, it was
determined that entire social groups of chimpanzees cooperated with the request to
move indoors during 56% of the attempts. It should be noted that if the entire group
failed to cooperate with an attempt, another attempt immediately following the first
was usually successful. Group differences were evident here with the percentage of
attempts with 100% compliance ranging from a mean of 3082% per group. Groups
with very low compliance tended to contain one uncooperative member who repeatedly accounted for the less than 100% compliance.
DISCUSSION

Our findings indicate that training can be effective in improving the movement
of captive chimpanzees, much as has been shown for macaques [Knowles et al.,
1995; Luttrell et al., 1994; Phillippi-Falkenstein and Clarke, 1992]. Positive reinforcement training techniques were successful in increasing the chimpanzees level
of compliance with the request to move into, and to be restricted to, the indoor portions of their enclosures. There were no consequences to the chimpanzees if they
chose not to cooperate with the procedure, but after the training phase was completed, they continued to cooperate a mean of 87.9% of the time. The improved level
of voluntary movement achieved through this training program is important for allowing personnel safe access to animal enclosures for maintenance, to move animals
to other quarters, to facilitate veterinary or research procedures, and to gain fast
access to chimpanzees in emergency situations. Since these activities are performed
frequently in most colonies, the practical impact of the training in terms of time
savings is great. The procedure is brief, usually requiring only a few minutes, so it
can be readily incorporated into husbandry routines. In practice, we have found we
can call the animals to move indoors several times in a day without any obvious
decrement in performance. Because we are employing only positive techniques and
are not forcing uncooperative animals to participate, we have modified our expectation that every animal will respond every time we ask. We have found that an unco-

Training for Voluntary Movement

339

operative animal during one attempt may very well choose to cooperate during the
next, even with very little intervening time, so we try to give a second chance.
Aversive techniques could always be called on in an emergency situation if required.
Adult males displayed the lowest level of compliance during all phases of the
training. Females required fewer training sessions and training attempts to reach reliable performance than did males. However, every subject cooperated often enough
to reach reliable performance (a 90% compliance score) at some point in the study.
Therefore, although age and/or sex effects were statistically significant, this study
indicates that training of any sex/age class of chimpanzees can be successful.
When the two individuals who originally trained the chimpanzees transferred
responsibility for the procedure to other members of the caregiving staff, animal
compliance was stable when we examined the entire data set. However, there was a
reduction in compliance when we compared the period immediately before with that
immediately after the transfer. Although statistically significant, this difference in
compliance (94.2 vs. 89.8%) was relatively small. Even the lower score is well above
that of the baseline performance and is equal to our established operational definition of reliable performance, so the practical significance of this temporary decrement in performance is likely to be small. The generalizability of the animals
performance indicates that it was not specific to the original trainer, a complaint
sometimes voiced as a limitation of training. The success of this transfer of responsibility to other care staff exemplifies the value of the training as it can be applied by
all people working with the animals.
Before beginning our more formal training program, the chimpanzees were
moved to their indoor areas when needed, but aversive methods were sometimes
used. We changed this sometimes aversive experience into a positive one with the
fairly minor modification of reinforcing the behavior of animals who chose to cooperate. Previously, the procedure of moving the chimpanzees indoors was perceived
by those working with the chimpanzees to be more stressful to the animals than it
now appears. This subjective impression should be objectively tested. Because it no
longer seems to be a stressful procedure for the animals, we move the chimpanzees
indoors for brief periods more often. This has given us more frequent access to the
corral portions of the enclosures, which allows, among other things, more routine
stocking of enrichment devices in the corral.
Most previously published reports on training nonhuman primates for movement describe the use of a combination of positive and negative reinforcement
[Knowles et al., 1995; Luttrell et al., 1994; Phillippi-Falkenstein and Clarke, 1992;
Reinhardt, 1990]. In contrast, the success of relying solely on positive reinforcement
techniques is reported here and in a report by Kessel-Davenport and Gutierrez [1994],
who trained chimpanzees to move into transport caging. Avoiding the use of aversive techniques may be especially practical when dealing with apes and other large
animals because it is not generally possible to enter enclosures and force them into
cooperating with a procedure. Using only positive techniques may also provide other
benefits that have a broader impact on animal well-being [Laule et al., 1992]. Positive reinforcement allows increased choice as the animals volunteer to participate or
not. It may improve relationships between people and the animals with whom they
work, reduce stress associated with certain procedures or events, give animals an
opportunity to work for food, and give them a chance to experience the stimulation
of learning. Each of these possibilities should be objectively tested. By relying strictly

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Bloomsmith et al.

on positive training methods, it may take longer to accomplish a goal, but if some of
these other benefits are accrued, the additional time would be well justified.
This study also demonstrates that the training of animals can be objectively
evaluated using systematic study design, data collection, and statistical analysis of
data. For example, to support the conclusion that training is responsible for a reported change in behavior, it is necessary to have baseline data prior to initiating
training procedures, or some other form of experimental control. Such information is
rarely reported but is a part of the current study.
CONCLUSIONS

1. Positive reinforcement training techniques were successful in significantly

increasing chimpanzees level of compliance with the request to move and be locked
into the indoor portions of their enclosures.
2. Females chimpanzees required fewer training sessions than did males.
3. There was a significant age-by-sex interaction in performance, with adult
males complying less than other age/sex classes, but even these males reached reliable performance.
4. Animal compliance with this procedure was temporarily and only slightly
reduced when the individuals who originally trained the chimpanzees transferred responsibility for the procedure to other people.
ACKNOWLEDGMENTS

The authors thank Tom Beck, Bob Thurston, Dr. Pat Alford, and Dr. Michale
Keeling for their help with the project. We also thank Caroyl Maliniemi for manuscript preparation and Jude Richard for manuscript editing. This project was supported by National Institutes of Health/National Center for Research Resources Grants
U42-RR03589 and R01-RR03578. Animals are maintained in facilities approved by
the American Association for Accreditation of Laboratory Animal Care and in accordance with current United States Department of Agriculture, Health and Human Services, and National Institutes of Health regulations and standards.
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