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WellBeing International WBI Studies Repository

2005

Chimpanzees in Research: Past, Present, and Future

Kathleen Conlee Studies Repository

Sarah T. Boysen

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Part of the Animal Experimentation and Research Commons, Animal Studies Commons, and the and Medical Ethics Commons

Recommended Citation Conlee, K.M., & Boysen, S.T. (2005). in research: Past, present, and future. In D.J. Salem & A.N. Rowan (Eds.), The state of the III: 2005 (pp.119-133). Washington, DC: Humane Press.

This material is brought to you for free and open access by WellBeing International. It has been accepted for inclusion by an authorized administrator of the WBI Studies Repository. For more information, please contact [email protected]. Chimpanzees in Research: Past, Present, 9CHAPTER and Future

Kathleen M. Conlee and T. Boysen

Introduction himpanzees have been used conservation issues, and pursuit of across equatorial , from Sene- in research in the United their legal rights (Cavalieri and gal, Mali, Sierra Leone, Côte CStates since the 1920s Singer 1993; Wise 2000, 2002). d’Ivoire, Ghana, Nigeria, Cameroon, (Brent 2004), with their breeding Why is there particular interest and Gabon in West Africa; the cen- and use highlighted in the 1980s as in the use of chimpanzees in tral African countries of , a model for acquired immune defi- research? They are the only Equatorial Guinea, the Central ciency syndrome (AIDS) research. (of both great and small) used in African Republic, the Democratic However, the use of chimpanzees in biomedical research and testing in Republic of Congo, Uganda, and harmful research has come to be the United States, and much has Burundi; and Tanzania in east questioned throughout the world, been learned about their emotion- Africa. Chimpanzee social structure based on both ethical and scientific al and over the has been observed to include nearly concerns. Public support for chim- last several decades.1 every type of relationship seen panzee research has been declining Although the welfare of chim- among different , over time (National Science Board panzees encompasses many issues, including multimale or multifemale 2002), costs of using chimpanzees this chapter addresses their use in groups, bachelor groups, male/ in research have been rising, the research, including their historical female breeding pairs, a mother number of chimpanzees in laborato- and current use in the United and her infant, or a female and her ries (including in the United States, ethical and scientific con- offspring of various ages. States) has been declining, and leg- cerns, public opinion, international In general, chimpanzee social islation and policies prohibiting the legislation, and future directions. organization is described as a fis- use of great apes in research have sion-fusion society, with individuals been on the rise internationally. or small groups leaving and then These trends may indicate an end to The Species periodically rejoining the group. the use of chimpanzees in research Chimpanzee Like many primate species, chim- in the United States and abroad in panzees give birth to a single the near future. Other than ( troglodytes) infant, who may nurse for four to increased attention to the use of Chimpanzees are members of the five years, so the offspring have an chimpanzees in research, animal taxonomic and the extended period of maturation and protection groups, conservationists, great (), which learning. Males remain in their lawyers, and others are focusing on also includes (both lowland natal group for their entire , issues related to chimpanzees as and mountain subspecies), orang- while females of reproductive age well, including their use in enter- utans, and (formerly emigrate and take up residence in tainment, of them in the referred to as pygmy chimpanzees). neighboring communities. These wild for food (known as “bush- The natural habitat of the chim- sex-related behavioral strategies meat”) and the pet trade, general panzee is a range of countries thus serve as a natural incest taboo

119 and help maintain • Studies that suggest chim- within and among different chim- Chimpanzee panzees, like , under- panzee groups in a given area. Male Intelligence stand that other chimpanzees chimpanzees maintain order and Cognitive and behavioral research may have the same or differ- position in their groups through a with chimpanzees, including both ent set of beliefs, desires, and and often field studies and captive work over knowledge from their own, a form coalitions of two to three the past forty years in particular, capacity formerly believed to males who co-rule the group. have taught us much about the be unique to humans (e.g., Females, however, are not as social remarkable capabilities chim- Hare, Call, and Tomasello with other females as males are panzees share with humans. These 2001; Tomasello and Call with males, although a dominance include: 1997). structure does exist among them. • An extensive list of some thir- Clearly, the evidence demon- Exceptions have been observed, ty-nine-plus types of use strates that the chimpanzee is a even to the point of a female who in the wild (e.g., Goodall species whose genetic, morphologi- participated in cooperative hunt- 1968; McGrew 1992; Whiten cal, anatomical, neurological, bio- ing with the males of her group, et al. 1999) chemical, behavioral, and cognitive although most of such opportunis- • Complex processing capacities similarity to humans is unique tic predation on small for acquiring concepts such as among all other species living today. (including monkeys such as the “same vs. different” (e.g., red colobus) has typically occurred Premack and Premack 1983) among all-male groups. • Numerical skills, including Chimpanzee Like many nonhuman primates counting abilities, that are Emotions and whose habitats are being en- comparable in chimpanzees’ croached upon, the chimpanzee is development as they are in Motivation listed as “endangered” in the wild young children (e.g., Boysen During the past several decades, under the U.S. Endangered Species and Berntson 1989; Matsuza- much has been learned about the Act. Some estimates are that only wa 1985a) chimpanzee’s motivation and capac- 110,000 animals remain across • Productive use and compre- ity for emotional expression. Empir- Africa. However, unlike any other hension of symbolic - ical studies under controlled condi- species on the list, the chimpanzee like systems of several types, tions in have documented is the only species that is cross-list- including signed English based that the emotional range of chim- ed as “threatened” in captivity, on American , panzees is quite comparable to that thereby given less protection from visual symbol systems such as observed in humans, with consider- certain types of biomedical and plastic shapes that stand for able overlap in facial expressions invasive research. Consequently, words, or graphic symbols that (Parr, Dove, and Hopkins 1998; Parr the “threatened” status of the cap- are computer-interfaced to dis- 2001, 2003). These include expres- tive population permits procedures play the word-like symbols cho- sions exhibited during laughter; and other activities that are not sen and the order in which under conditions of fear, anger, or legally permitted with wild chim- they have been selected (e.g., sadness; and a range of grimaces panzees. If chimpanzees were list- Matsuzawa 1985b; Premack observed in neonates, such ed solely as endangered, the types 1986; Savage-Rumbaugh 1986; as disgust or pleasure in response to of research that are currently Gardner, Gardner, and van odors and/or taste. allowable could simply not be Cantfort 1989) Observations in both wild and done. Currently, only a few coun- • Extensive skills with problem captive settings suggest that chim- tries other than the United States, solving of all kinds observed in panzees are subject to some of the including Gabon, Liberia, and both the wild and under exper- same types of behavioral and emo- Japan (although a ban is in prepa- imental conditions in captivity tional pathologies as have been ration there), permit biomedical (e.g., Matsuzawa 1985b; observed in humans, including research on chimpanzees. Chim- Limongelli, Boysen, and Visal- depression, various neuroses, anxi- panzee research is not permitted berghi 1995; Kuhlmeier and ety, and even to the point of in the United Kingdom, Sweden, Boysen 2002) death (Goodall 1986). It is typical- Australia, New Zealand, or the • Recognition of kin relation- ly easy, especially for young chil- Netherlands (although not formal- ships based on comparing dren, to watch chimpanzees in a ly declared by each country, no photographs alone of chim- or sanctuary and recognize European Union countries conduct panzees and their offspring that the animals are playing tag or research on chimpanzees). (Parr and de Waal 1999a) play-fighting or that a disagree-

120 The State of the Animals III: 2005 ment has occurred between ani- panzee behavior, including a range bers, spinning in giant centrifuges, mals, with resultant real fighting. of observations that included social and use of shock as punishment The overlap among behavioral and interaction, play, sexual activity, while training (Save the Chimps emotional expressions between diet, morphology, anatomy, emo- n.d.). In January 1961 a chim- humans and chimpanzees is quite tional states, facial expressions, panzee named was placed on a dramatic, such that even very vocalizations, and intelligence. ballistic trajectory flight and forced young children are able to interpret Yerkes’s work was critical to the to perform a motor task through- often complex social interactions emergence of primate studies in the out the flight for which he had been among chimpanzees quite accu- United States. His burgeoning labo- trained. In November 1961 a sec- rately. (There are notable excep- ratory moved first to Orange Park, ond chimpanzee, , orbited the tions, however, such as differences Florida, in 1930 and then to Emory twice and was forced to per- in the two species’ respective University in Atlanta, Georgia, in form a more complex task (NASA “smiles”—a chimpanzee “smiling” 1965 where, as the Yerkes National 2004). Unfortunately, through a with upper and lower teeth showing Primate Research Center, it remains malfunction in equipment, Enos is expressing fear, for example.) today (Yerkes National Primate received a shock for every correct Research Center n.d.). In addition maneuver he made, which contra- to his numerous books on apes, dicted the 1,263 hours of training The History of including chimpanzees, Yerkes con- he had undergone (NASA 2004; U.S. Chimpanzee tributed a wealth of scientific papers Save the Chimps n.d.); despite the to the emerging literature. Yerkes’s shocks, Enos continued to com- Research: books and journal articles remain an plete the task correctly. important source for researchers, After some Air Force chimpan- 1920–1979 particularly for those whose inter- zees were sent into space, they were Chimpanzee research began with ests are in chimpanzee cognition reassigned to other projects, such the work of Robert M. Yerkes of Yale and behavior. He was the first to as testing seat belts. In the 1970s University, who established a labo- study many phenomena in chim- the Air Force no longer used chim- ratory at his rural home in the early panzees of great importance to the panzees but did lease them out for 1920s with two purchased chim- field of and is consid- biomedical research studies (Save panzees (Yerkes and Learned ered to be one of the fathers of pri- the Chimps n.d.). In 1975 the Con- 1925). His early writing about matology in the United States. vention on International Trade in these animals, a male and a female, In the 1940s the focus at Yerkes Endangered Species (CITES) was explored a wide range of behavioral National Primate Research Center adopted, which greatly restricted and intellectual capacities ob- shifted from the study of behavior importation of chimpanzees from served both directly and indirectly to the study of infectious disease the wild. This prompted a captive- as the young chimpanzees devel- (Committee on Animal Models in breeding effort within the United oped. He was particularly interest- Biomedical Research 1995). The States, which has been federally ed in and wrote fairly extensively use of chimpanzees for the study of funded since 1986 (Brent 2004). about the differences he noted infectious disease has increased between the two animals and, at ever since, particularly in hepatitis the time, attributed such to sex dif- and human immunodeficiency Chimpanzee ferences. However, it was later con- virus (HIV), and continues at a Research: 1980 firmed that Yerkes actually had one number of facilities (Table 1). chimpanzee (Pan troglodytes) and In the 1950s the U.S. Air Force to the Present one (Pan paniscus), so created a research and breeding many of the differences he attrib- program with sixty-five wild-caught AIDS Research uted to sex may actually have been chimpanzees to determine the in the 1980s species differences. This was partic- effects of space flight on humans During the 1980s there was a dras- ularly notable with respect to dif- (Brent 2004; Save the Chimps tic increase in chimpanzee re- ferences in vocalizations, although n.d.). The aeronautics research search, primarily prompted by the many other behavioral traits were involved subjecting chimpanzees to human AIDS epidemic. A massive also confounded by reporting them a number of stressors during train- breeding effort was launched in as sex rather than species differ- ing as well as the obvious stressors 1986 (National Research Council ences (Yerkes and Learned 1925). associated with being launched 1997), and in 1992 scientists repre- Despite this misguided start, Yerkes into space. These stressors includ- senting and AIDS and his wife contributed several of ed exposure to G forces, loss of con- research interests met to discuss the first descriptions of chim- sciousness in decompression cham-

Chimpanzees in Research: Past, Present, and Future 121 Table 1 U.S. Facilities Housing Chimpanzees: Types of Research and Numbers of Animals

Facility* Location Type Total Number Number of of Research of Chimpanzees NCRR-Supported Chimpanzees3

New Iberia New Iberia, La. Breeding, vaccine research, drug Research Center efficacy 3501 130

Alamogordo Alamogordo, N.M. Behavioral 2751 270 Primate Facility

Southwest San Antonio, Tex. Vaccine and drug testing, hepatitis, 2501 15 National Primate Alzheimer’s, HIV Research Center

Yerkes National Atlanta, Ga. HIV, behavioral, neuroscience, 1971 75 Primate Research reproduction Center

M.D. Anderson Bastrop, Tex. Breeding colony, hepatitis, 1541 105 Cancer Center infectious disease

Primate Mesa, Ariz. Behavioral, reproductive, research 751 74 Foundation supply of Arizona

Bioqual Rockville, Md. Hepatitis, respiratory viruses 632 Not mentioned

Centers for Atlanta, Ga. Hepatitis 142 Not mentioned Disease Control and Prevention

Food and Drug Rockville, Md. 112 Not mentioned Administration

Ohio State Columbus, Ohio Behavioral, cognitive (noninvasive) 111 0 University

Language Decatur, Ga.Behavioral (noninvasive) 42 Not mentioned Research Center, Georgia State University

Chimpanzee and Ellensburg, Wash. Behavioral (noninvasive) 42 Not mentioned Human Commu- nication Institute, Central Washing- ton University

*This is not meant to be an exhaustive list of the types of research being conducted at each facility. 1 According to the International Directory of Primatology. 2 According to Goodall et al. 2003. 3 According to a presentation given by J. Strandberg at the American Association of Laboratory Animal Science (AALAS) conference in 2003. The remaining chimpanzees are not federally owned, but the facilities may still receive federal funding for research.

122 The State of the Animals III: 2005 the use of chimpanzees in human The Humane Society of the Unit- Some grants extended over as immunodeficiency virus (HIV) ed States (HSUS) examined U.S. many as twenty-five years; there- research (van Akker et al. 1993). At Public Health Service (PHS)-fund- fore, data for each year reflect both that time, the group acknowledged ed grants that involved captive ongoing research and newly funded there were some areas of HIV re- chimpanzees in HIV research in projects. In 1980 three PHS-fund- search for which chimpanzees were some way (including breeding for ed studies involved the use of chim- not necessary, such as prevention of HIV research), beginning in 1980. panzees in HIV-related research. maternal-infant transmission and physiological safety tests for vac- cine development. The group advo- Table 2 cated for alternatives, such as using Public Health Service-Funded Grants: monkeys, but it emphasized that HIV Research Involving Captive some of the suggested approaches engendered animal welfare con- Chimpanzees cerns as well. The group considered other factors related to HIV re- Year Number Types of HIV Research search on chimpanzees, such as of Grants housing conditions, and concluded that not allowing chimpanzees in 1980 3 Receptors HIV research to interact socially Vaccine safety Chimpanzee housing with other chimpanzees or humans “is both unnecessary and unethi- 1984 5 Receptors cal” (van Akker et al. 1993). The Vaccine safety Chimpanzee housing group advocated the use of environ- Transmission of HIV mental enrichment (innovative ways to enrich the lives of chim- 1988 17 Receptors panzees that promote natural be- Vaccine safety Chimpanzee housing havior) and housing that allows the Transmission of HIV chimpanzees to express natural Vaccine efficacy locomotor behaviors. Chimpanzee breeding/management It is not known whether HIV sur- 1992 18 Receptors vives in chimpanzees, but we do Vaccine safety know that the animals do not devel- Chimpanzee housing op the AIDS-related complex seen Transmission of HIV Vaccine efficacy in humans (Balls 1995; Nath, Schu- Chimpanzee breeding/management mann, and Boyer 2000). There is, Immune response however, a specific strain that is pathogenic in chimpanzees and 1996 20 Receptors Vaccine safety typically takes up to ten years to Chimpanzee housing progress to AIDS-like symptoms. Transmission of HIV Great controversy has arisen over Vaccine efficacy whether chimpanzees should, in Chimpanzee breeding/management Immune response fact, be challenged with that partic- HIV progression and pathogenesis ular strain (Nath Schumann, and Genetic inoculation Boyer 2000). Some members of the 2000 23 Receptors research community have strongly Vaccine safety opposed the idea, some publicly Chimpanzee housing (Prince et al. 1999). Over time, Transmission of HIV however, it has been determined Vaccine efficacy Chimpanzee breeding/management that the chimpanzee is a poor HIV progression in young chimpanzees model for HIV research, and some Infection with strain most virulent in chimpanzees researchers argue that the use of Cell-based immunotherapy chimpanzees is not likely to lead to 2004 7 Chimpanzee breeding/management a cure for AIDS (Reynolds 1995). expression in infected chimpanzees Despite this, HIV-related research Vaccine development in chimpanzees continues.

Chimpanzees in Research: Past, Present, and Future 123 Figure 1 Chimpanzee Research Grants, 2000–September 2004

This number increased to five Chimpanzees and ties in the United States as of grants in 1984 and jumped to sev- Research Facilities 2005. Table 1 provides a list of re- enteen in 1988. The next few years search facilities that as of 2005 resulted in an increase in these in the United States housed chimpanzees, some areas grants, to twenty-three in 2000, According to Stephens (1995), there of research conducted at each but this number fell to seven were approximately 1,800 chimp- facility, and the number of chim- grants in 2004 (Table 2). As of anzees in fourteen biomedical and panzees (if known) at each facility. 2001 150 chimpanzees had been behavioral research facilities in the The majority of captive research infected with various strains of HIV, United States in 1993. In 2001 a chimpanzees are housed at six bio- but only four had had evidence of National Institutes of Health (NIH) medical facilities. Information re- “progressive HIV infection,” and report to Congress identified 1,584 garding the number of chim- one of the four had progressed to chimpanzees, including 614 who panzees and chimpanzee research AIDS (Muchmore 2001). AIDS were government owned, who may facilities in the United States was research on chimpanzees (includ- have been used in federally support- also supported by a census con- ing colony maintenance) has been ed or conducted research and were ducted and reported by the Great conducted primarily at Yerkes housed in thirteen biomedical and Ape Project (Goodall et al. 2003). National Primate Research Center behavioral research facilities in the A review of the literature pub- (Atlanta, Georgia), Southwest United States (National Center for lished during 2001 and included in National Primate Research Center Research Resources 2001). Since the National Library of Medicine (San Antonio, Texas), New Iberia that time approximately 266 chim- and PrimateLit databases revealed Primate Research Center (New panzees formerly owned by a bio- that of the 4,411 studies worldwide Iberia, Louisiana), and the M.D. medical research facility in Alam- involving nonhuman primate re- Anderson Cancer Center Science ogordo, New Mexico, were search, nine involved the use of apes Park (Bastrop, Texas) (Table 2). transferred and are now being cared (Carlsson et al. 2004). Overall, it for by a sanctuary organization was estimated that 41,000 primates based in Florida. It was estimated were used, although the specific that there were approximately number of great apes represented 1,300 chimpanzees in twelve facili-

124 The State of the Animals III: 2005 by these studies is unknown, partic- can also cause and distress. mals live in large social groups of ularly because not all publications Invasive research, in general, raises eight to 20 individuals. The type of specify the number of animals used particular concerns regarding housing used depends on the partic- (Carlsson et al. 2004). Some stud- chimpanzee welfare in captivity. ular institution and the type of ies, particularly those from private- research being conducted. Chim- sector organizations such as phar- Chimpanzee Housing panzees who live in groups also can maceutical companies, are not and Care be separated for a period and placed published (Carlsson et al. 2004) at Individuals who have worked closely on research protocols that involve all. These data suggest that a review with chimpanzees in research single housing. The likelihood of of the published literature may not report that those used in many inva- this depends on several factors, in- produce reliable information about sive protocols are typically housed cluding the specific institution, the the actual number of chimpanzees alone in cages required by USDA type of research conducted there used in research, consequently standards to be only five feet by five (whether study animals could infect requiring reliance on other sources feet by seven feet, with twenty-five others if they were housed togeth- of information. square feet of floor space. This can er), and precedents within the insti- be compared to the interior of an tution that may not be necessary Research in Which elevator (Figure 2). Cages are typi- for the specific study but instead Chimpanzees Are Used cally constructed from steel and, in reflect the culture of the institution. Chimpanzees are most commonly some cases, include a perch for rest- An analysis of chimpanzee re- used for hepatitis (particularly hep- ing or sleeping. Many cages also search for the years 2000 to mid- atitis C) and HIV/AIDS research. A have a “squeeze back,” a moveable 2002 conducted by The HSUS total of 334 federally funded grants interior wall that can be pulled from revealed that information about the between 2000 and 2004 involved the back of the cage toward the types of housing provided in publi- the use of live chimpanzees, with front and can press or hold a chim- cations or in federal grant abstracts approximately 29 percent related panzee closer to the front of the was lacking (Conlee, Hoffeld, and to hepatitis research and 16 per- cage so that a technician, veterinar- Stephens 2004). Among 189 publi- cent related to HIV/AIDS research. ian, or researcher can administer cations 24 percent mentioned so- Stephens (1995) reported that injections or perform other proce- cial housing and 76 percent did not approximately 80 percent of dures without anesthetizing the mention any specific housing type. research conducted on chim- chimp. Under some conditions, Overall, information regarding the panzees in the early ’90s was relat- housing areas do not have any natu- specific number of chimpanzees ed to hepatitis and HIV/AIDS. ral light, and the animals live under maintained in each type of housing Therefore, these types of biomed- artificial lighting (light/dark) cyc- (individual vs. social) was not readi- ical research with chimpanzees are les at all times. ly available. Housing and environ- not as prevalent as they are in the In the wild, chimpanzees are very mental conditions, however, can recent past, although such invasive social and live in complex groups of have significant effects on research studies continue. varying sizes. Therefore, social Other areas of research for which housing is almost certainly the sin- chimpanzees are currently used gle most important factor for chim- include cognitive and behavioral panzee psychological well-being studies, as models for human repro- (National Research Council 1997). duction, malaria, gene therapy, res- Individual housing can lead to pro- piratory viruses, Crohn’s disease, found depression, increased aggres- drug and vaccine testing, and a vari- sion, psychological withdrawal, ety of other infectious diseases (Fig- extreme frustration, and self-muti- ure 1). Experiments in some of lation, such as physical wounding, these areas, such as studies of cer- hair plucking, rocking, and other tain strains of HIV, can lead to psychotic-like behaviors. Chim- severe appetite and weight loss, panzees who are not being used in lethargy, diarrhea, severe illness, active research protocols typically infections, and/or eventual death. are housed in pairs or social groups. Procedures such as major surgery, The physical environment for social liver biopsies (required for some housing can range from a cage that protocols in hepatitis research and is slightly larger than the individual FAUNA FOUNDAITON Figure 2. involving multiple biopsies), fre- cage depicted in Figure 2 to large A typical laboratory cage for individually quent blood sampling, and restraint outdoor enclosures where the ani- housed chimpanzees. Chimpanzees in Research: Past, Present, and Future 125 results, so such information should tooth and the chimp’s loud alarm be included in all publications. Ethical calls drew the other chimps to the Regardless of whether housing Questions and scene, the group’s response—rau- information is available, Balls cous calls and all members peering (1995) raises an important point: it Responsibilities at the tiny white tooth on the may be impossible to provide hous- The United States currently uses ground—clearly suggested that ing in laboratories that truly meets more chimpanzees in biomedical the group interpreted the pain and the physiological and behavioral research than any other country in blood loss as caused by the tooth needs of chimpanzees under captive the world. The U.S. government pro- itself as an animate object. conditions. vides more funding for the study of One author (S.B.) and her stu- chimpanzee cognition and behavior dents have observed their subjects Funding for Research than does any other country. Results readily sharing food with younger The HSUS analysis of federally fund- from studies over the past four chimps, assisting older animals ed great ape research found that decades in particular have provided having difficulty moving from place $20 million to $25 million dollars of a wealth of scientific evidence show- to place in the facility, and re- federal funding per year is devoted ing that chimpanzees and humans sponding with “reverence” to the to chimpanzee research and care bear striking similarities. While we body of a group member who had (Conlee, Hoffeld, and Stephens have known for up to two hundred died of natural causes. In the last 2004) (Figure 3). Hepatitis re- years that the anatomy, physiology, instance, the dead chimp’s cage search accounts for $4.2 million of morphology, biochemistry, and gen- mate picked up a blanket, covered this funding each year, and HIV etic overlap between chimpanzees the dead chimp’s head, and then research accounts for approximate- and humans is overwhelming, it has placed a second blanket over her ly $500,000. The amount of private- only been within the last forty years body. A videotaped record of these sector funding for chimpanzee that demonstrations of chimpanzee events leads an observer to the research is not available to the pub- cognitive abilities and behavior, conclusion that the “friend’s” lic; however, the use of chimpanzees including a wide range of emotions response was intentional and by the private sector may be on the evoked by chimpanzees and human empathetic (S.B., personal obser- rise. A chimpanzee researcher sit- beings in similar situations, have vation 2003). Goodall (1968) ting on a panel at the 2003 Ameri- been reported from field studies reports similar behaviors to those can Association of Laboratory Ani- (e.g., Goodall 1968) and captive described above among wild mal Science conference indicated work (e.g., Washburn and Rum- chimps, suggesting that captive that 75 percent of private-sector baugh 1992; Brown and Boysen chimpanzees are not acquiring growth (particularly pharmaceuti- 2000). Recent technological ad- behaviors unseen in the wild. Long- cal companies) at the New Iberia vances have allowed direct compar- term observations of chimps in the Research Center was due to re- isons at the neuroanatomical level field and captivity have increasing- quests for chimpanzee use. between the two species, with ly complemented and confirmed a It is estimated that it costs notable correspondence between a range of comparable behaviors $20–$30 a day to care for a chim- significant number of neuroanatom- that are seen in humans as well as panzee in the laboratory and $15 a ical structures that likely support in the chimpanzee. Observations day to care for one—better—in a the same functions (e.g., Cantalupo, of behaviors of this level of sophis- sanctuary. Compare the $9.5–$14.2 Pilcher, and Hopkins 2003; Hopkins tication and complexity raise diffi- million a year to care for the United and Cantalupo 2004). cult ethical and moral questions States’ 1,300 chimpanzees in a lab- With more than thirty years of about the types of research on oratory to the $7.1-million-a-year direct interactions with chim- chimpanzees that are permitted in cost of sanctuary care. It is impor- panzees as part of a comparative the United States. tant to emphasize that the sanctu- cognition project, one author More detailed studies of the sim- ary setting not only costs less per (S.B.) (2000) reports that her ilarities between human and chim- chimpanzee per day, but also can chimps have shown a number of panzee behavioral and emotional provide a much more naturalistic behaviors suggesting that they responses are even more telling. and stimulating environment. were responding to natural events Parr and de Waal (1999b) provided such as wind or thunderstorms captive chimpanzees with photo- with great fear. A similar response graphs of chimpanzees they didn’t was likely felt by early humans, who know and found that the chim- subsequently created myths and panzees were not only able to legends to explain these phenome- match two different photographs na. When a chimpanzee lost a of the same individual, but also to

126 The State of the Animals III: 2005 match mothers and sons. This moral responsibilities to chimps man health problems.” In 2002 demonstrates that chimpanzees (National Research Council 1997). (the most recent survey results are capable of identifying similari- Unlike humans who participate in available as of 2005), 52 percent of ties in the faces of related individu- biomedical research, chimpanzees adults opposed or strongly op- als who were unfamiliar to them. are incapable of giving informed posed this statement. When the In another test by Parr and De consent. Therefore, it is clearly same statement was used in a 1985 Waal, chimpanzees were presented time for society to reappraise the survey, only 30 percent of adults with sample head-shot photo- status of humankind’s closest pri- voiced opposition (National Sci- graphs of chimpanzees. The sub- mate relative. ence Board 2002) (Table 3). jects recognized the emotional expressions of the chimpanzees in the sample photographs and Public Opinion: U.S. Overview matched them to photographs of Driving Change novel chimpanzees showing facial Recent Issues Increasing public concern has large- expressions that depicted the same ly driven international efforts to end Over the last twenty years, major emotional state. The subjects the use of chimpanzees in research. changes in the use of chimpanzees chose the photograph that best According to a recent opinion poll in research have taken place. The matched the sample chimpanzee’s conducted by Zogby International rush to increase breeding for HIV picture, based on the underlying for the Doris Day Animal League in research in the 1980s was followed meaning of the facial features and 2001 (in Conlee 2003), 90 percent by a significant decrease in the num- configuration, since the perceptual of Americans believe it is unaccept- ber of facilities housing chim- and physical features were not pre- able to confine chimpanzees in gov- panzees as well as in the number of cisely the same. ernment-approved cages (Figure 2), chimpanzees at each facility in sub- Such trials underscore chim- 54 percent believe it is unaccept- sequent years. Three large chimp- panzees’ capacity for empathetic able for chimpanzees to “undergo anzee research laboratories have responses. Such responses, coupled research which causes them to suf- closed since 1995, and many of their with the cognitive capacities hu- fer for human benefit,” and 65 per- chimpanzees are now permanently mans demonstrably share with cent say it is unacceptable to kill retired at sanctuaries throughout chimps, indicate that, under cir- them for research. the United States. In 1995 New York cumstances in which a human A 2002 opinion poll by Penn, University decided to close its Labo- being might experience emotional Schoen, and Berland Associates for ratory of Experimental Medicine distress or trauma, chimpanzees The Humane Society of the United and Surgery in Primates (LEMSIP). respond similarly under comparable States (HSUS n.d.) found that 79 Approximately half of the LEMSIP conditions. One example would be percent of the U.S. public supports chimpanzees were sent to various for a chimpanzee to be housed in creation of a government-sponsored retirement facilities, but the other isolation, with no physical or social sanctuary system to provide life- half were sent to the Coulston Foun- contact with other chimpanzees, as time care to chimpanzees no longer dation, Alamagordo, New Mexico, well as with only minimal daily con- used in research. This and other the largest chimpanzee colony in tact with caregivers. There is a rea- survey findings indicate that not the world at that time, which had a son that similar housing conditions only does the public oppose the suf- poor record of compliance with the in our nation’s prisons, that is, soli- fering of chimpanzees in research, Animal Welfare Act (AWA). tary confinement, are considered to but it also is willing to financially The second large closure was that be the worst conditions for inmates support a significant commitment of the chimpanzee colony at the to endure. (Indeed, solitary confine- to chimpanzees, who can live to be Holloman Air Force base, also in ment of human prisoners is consid- sixty years old in captivity. New Mexico, in 1997. This colony of ered by some to be “cruel and The National Science Board, 141 chimpanzees who were used by unusual punishment.”) which conducts surveys of public the space program was released These findings suggest that the attitudes toward scientific re- from the Air Force. In a controver- range and of invasive search every three years, included sial decision, all but thirty chim- research in the United States repre- the following statement in its 1985 panzees were sent to the Coulston sents unethical and, indeed, im- survey: “Scientists should be al- Foundation instead of to sanctuar- moral actions. In its 1997 report, lowed to do research that causes ies that had volunteered to take in a the National Research Council that pain and injury to animals like number of them. (Those requests examined the status of chim- dogs and chimpanzees if it pro- had been denied by the Air Force.) panzees in research facilities in the duces new information about hu- One of those sanctuaries was the United States noted the ethical and Center for Captive Chimpanzee

Chimpanzees in Research: Past, Present, and Future 127 out retiring them under the Table 3 CHIMP Act (see below). This is an Public Opinion on Using Chimpanzees unfortunate development. and Dogs in Painful and Injurious U.S. CHIMP Act Research The large chimpanzee breeding effort launched in the United States in 1986 exceeded expecta- Survey Statement: Scientists should be allowed to do research that causes tions at the same time it was pain and injury to animals like dogs and chimpanzees if it produces new information. determined that the chimpanzee was not a critical model for HIV Year Supporting/Strongly Opposing/Strongly research after all. This created a Supporting Animal Research Opposing Animal Research “surplus” of chimpanzees for 1985 63 30 research. As a result, the National Institutes of Health called on the 1988 53 42 National Research Council (NRC) 1990 50 44 to provide input on key issues, including the number of chim- 1992 53 42 panzees required to support 1995 50 46 research needs and how to address the long-term needs of the animals 1997 46 51 who had been produced. The NRC found (l) that euthanasia is not 1999 50 47 considered by the public to be an 2001 44 52 acceptable means of addressing the surplus issue (as previously Source: National Science Board 1985–2001. noted); (2) a five-year breeding Number of adults surveyed varied per year and ranged from 904 to 2,041. moratorium should be adopted; and (3) sanctuaries should be Care (the CCCC—now known as By 2002 the company had col- established for the long-term care Save the Chimps), an organization lapsed financially and divested of retired chimpanzees (National that ultimately sued to obtain cus- itself of 266 chimpanzees, selling Research Council 1997). tody of twenty-one of the chim- them to Save the Chimps, which Following the NRC report, lobby- panzees. The CCCC entered into an purchased the land and facilities ing efforts began for the creation agreement with the Coulston Foun- from the company. of a national chimpanzee sanctu- dation in October 1999 that Despite the decrease in the num- ary system through what became brought those chimpanzees to live ber of chimpanzee laboratories and known as the Chimpanzee Health at the Save the Chimps’ sanctuary the retirement of a significant Improvement, Maintenance and in Florida. number of chimpanzees, there are Protection Act (CHIMP Act). The The most recent laboratory clos- signs that some aspects of chim- animal protection coalition devot- ing was that of the Coulston Foun- panzee research have been grow- ed to passage of the CHIMP Act dation in 2002. Approximately one ing. In addition to 75 percent of was known as the National Chim- year before closing, Coulston trans- private-sector growth at the New panzee Research Retirement Task ferred three hundred chimpanzees Iberia Research Center coming Force (NCRRTF). It consisted of to the Alamogordo Primate Facility, from requests for use of chim- The HSUS, the American Anti-Vivi- currently run under contract by panzees in research, New Iberia and section Society, the American Soci- Charles River Laboratories, to set- the Southwest National Primate ety for the Prevention of Cruelty to tle violations of the AWA. The chim- Research Center have each re- Animals, the Society for Animal panzees at the Alamogordo Primate ceived funds from the National Protective Legislation, and the Facility were not being used for Institutes of Health to expand their National Anti-Vivisection Society, research at that facility as of mid- chimpanzee-holding facilities. The with the support of an advisory 2005, but they could be transferred abstract of the grant for New Iberia board of numerous primatologists. elsewhere for research (Brent specifies that such a facility will The CHIMP Act was sponsored and 2004). In 2001 the National Insti- allow other laboratories to hold introduced in the House of Repre- tutes of Health stopped funding the their chimpanzees within the bio- sentatives (H.R. 3514) by Rep. Coulston Foundation (Brent 2004). medical research community with- James Greenwood (R-PA) on

128 The State of the Animals III: 2005 November 22, 1999; a companion while holding them in less expensive 20, 2000, by President Bill Clinton. bill, sponsored by Sens. Richard housing in the interim. Some pro-animal groups pursued a Durbin (D-IL) and Bob Smith (R- The final legislative language repeal of the CHIMP Act, but they NH), was introduced in the Senate specified that various requirements were unsuccessful. One important (S. 2725) on June 13, 2000. A leg- be met before any individual chim- and positive result of the CHIMP islative hearing was held on May panzee could be removed from the Act was a shift in thinking and pol- 18, 2000, with key individuals tes- system, thereby greatly reducing icy related to the use of chim- tifying, including of the chances that animals would be panzees in research. the . (John moved back into the laboratory. Since passage of the legislation, Strandberg of the National Center These requirements included: various efforts have been underway for Research Resources, National • Researchers could subject the to create the national sanctuary Institutes of Health [NIH], provid- chimpanzee and his or her system. The NIH published a ed the only oral testimony against social group to only minimal “sources sought” notice in 2001 the bill). pain, distress, and disturbance (Federal Register, April 19, 2001) The CHIMP Act incited a fair (as determined by the board of and, on September 30, 2002, grant- amount of controversy when then- directors of the sanctuary). ed the nonprofit Chimp Haven, in House Commerce Committee • Special circumstances related Shreveport, Louisiana, the contract Chairman Thomas Bliley (R-VA) pro- to the particular chimpanzee’s to run the entire system. Chimp posed amendments that would have medical history might make Haven’s mission is to provide life- provided the research community him or her uniquely needed for time care to chimpanzees previous- with limited access to chimpanzees research. ly used in research, as pets, or for after they were sent into the sanctu- • The technology to be used was entertainment (Brent 2004). ary system. When this amendment not available when the chim- The sanctuary contract stipulates was proposed, the animal protection panzee entered the sanctuary that the federal government will community, including NCRRTF, system. provide $19 million for the care of became divided, and its support for • The research is essential to an initial two hundred chimpanzees the legislation declined. Some address an important public for ten years, with Chimp Haven pro- groups decided to continue work on health need, and that the appli- viding matching funds of $4 million the legislation to ensure that any cant has not violated the AWA. (Brent 2004). The government will opportunity to remove chimpanzees • The proposal is subject to also provide $10 million in construc- from the sanctuary system was as public scrutiny through a tion costs, and Chimp Haven is narrow and difficult as possible, fear- sixty-day formal notice and expected to match 10 percent of ing that the bill ultimately would comment process. those funds (Brent 2004). allow the research community to The CHIMP Act (P.L. 106–551) The Chimp Haven facility in have easy access to chimpanzees was signed into law on December Shreveport will house two hundred chimpanzees at the outset and eventually expand to house a total Figure 3 of three hundred. At least two Public Health Service Funding other sites will hold groups of sev- enty-five or more. Chimp Haven for Chimpanzee Research, can also contract care out to other 2000–2002 facilities, but it will ultimately be responsible for all of the chim- panzees in the system—a maxi- mum of nine hundred individuals (Brent 2004). The first phase of construction at Chimp Haven has been completed, and chimpanzees began to arrive on April 1, 2005 (personal communication, Chimp Haven representative, with S.B., April 22, 2005). The U.S. government has asked laboratories and government enti- ties holding chimpanzees to pre- pare lists of animals no longer

Chimpanzees in Research: Past, Present, and Future 129 needed for research. These lists United Kingdom since 1986 (U.K. will be shared among the facilities International Animal Procedures Committee so that laboratories can share and Activities 1998, 2001). undertake research on chim- Some countries already prohibit or In 2000 New Zealand placed strin- panzees if desired, but the lists strongly restrict the use of chim- gent restrictions on the use of non- had not been made available to panzees in research. In 1997 the human hominids (nonhuman great the public as of mid-2005. Table 4 United Kingdom announced that apes—which include chimpanzees, provides a timeline of events relat- licenses to conduct research on bonobos, gorillas, and ) ed to the creation of the national great apes would no longer be within its Animal Welfare Act sanctuary system. granted, although great apes have (www.maf.govt.nz/biosecurity/ not been used in research in the legislation/animal-welfare-act/

Table 4 National Chimpanzee Sanctuary System: Timeline of Events

Date Action

April 15, 1999 A coalition that includes representatives from the research, animal-protection, zoo, and sanctuary communities writes a letter regarding the issue of chimpanzee “retirement” and submits it to U.S. Rep. J.E. Porter (R-IL) and U.S. Sen. A. Specter (R-PA).

November 22, 1999 H.R. 3514, the Chimpanzee Health Improvement, Maintenance, and Protection (CHIMP) Act, is introduced in the U.S. House of Representatives by Rep. J. Greenwood (R-PA). This bill will require the federal government to provide for permanent “retirement” of chimpanzees who are identified as no longer needed for research.

May 18, 2000 The House Committee on Commerce holds a hearing on H.R. 3514. Those presenting testimony include J. Goodall (Jane Goodall Institute), J. Strandberg (NIH), T. Nelson (National Chimpanzee Research Retirement Task Force), and A. Prince (New York Blood Center).

June 14, 2000 S. 2725, the Chimpanzee Health Improvement Maintenance and Protection (CHIMP) Act, is introduced in the U.S. Senate by Sens. R. Smith (R-NH) and R. Durbin (D-IL).

September 20, 2000 S. 2725 gains approval by the Senate Health, Education, Labor, and Pensions Committee.

October 24, 2000 The House passes H.R. 3514 with the Bliley amendments (see section, entitled Legislation: United States and International).

December 6, 2000 The Senate passes S. 2725 unanimously.

December 20, 2000 President Clinton signs the CHIMP Act into public law (P.L. 106–551).

April 16, 2001 The National Center for Research Resources (NCRR), part of NIH, publishes a “sources sought” notice to determine whether there is an existing nonprofit that fulfills the requirements of the CHIMP Act and is interested in serving as the “contractor” of the sanctuary system.

September 28, 2001 NIH publishes a Request for Proposal for an entity to operate and maintain a sanctuary system via the CHIMP Act.

December 20, 2001 The departments of Labor, Education, Health and Human Services and related agencies’ 2002 Appropriations Act (H.R. 3061) allocates $5 million to begin construction of the national chimpanzee sanctuary facilities.

January 10, 2002 President G.W. Bush signs H.R. 3061 into public law, including $5 million toward construction of the national sanctuary system.

September 30, 2002 NIH announces the award of a contract to Chimp Haven to establish and operate a chimpanzee sanctuary, pursuant to the CHIMP Act.

May 1, 2003 Chimp Haven, the contractor of the national chimpanzee sanctuary system, breaks ground on its Shreveport, La., facility.

January 11, 2005 NIH publishes a notice of proposed rule making regarding standards of care for chimpanzees held in the national chimpanzee sanctuary system.

130 The State of the Animals III: 2005 guide/awguide.pdf, 24). The coun- try’s director-general can approve Table 5 the use of nonhuman hominids, but International Legislation, Policies, he or she must first consult the National Advisory and Regulations Related to Committee; the use of these Chimpanzees in Research species must be in the best interest of the individual animal or the species; and the benefit must out- Country Type Year Comments weigh the harm. At the time this of Action Enacted ban was implemented, no great United Policy 1997* Licenses to conduct research apes were being used in research, Kingdom on nonhuman great apes will but the action sent a strong mes- no longer be granted sage about the ethics of such use. New Zealand Legislation 2000 Stringent restrictions on the When the Netherlands finalized use of nonhuman great apes an amendment to the Dutch Law in research on Animal Experiments in 2002 United States Legislation 2000 Chimpanzees determined no that prohibits the use of great apes (P.L. 106-551) longer needed in research are in biomedical experiments (Con- transferred to a national lee, Hoffeld, and Stephens 2004), sanctuary system six chimpanzees being used in hep- Netherlands Legislation: 2002 The use of great apes in atitis research already underway an amendment biomedical experiments were exempted from the ban. At to the Dutch is prohibited the time of the amendment, the Law on Animal only chimpanzees in the European Experiments Union were located at the Biomed- Sweden Regulations 2003 The use of apes in research is ical Primate Research Centre prohibited (BPRC) in the Netherlands. In Japan Unknown Unknown Invasive research on great October 2002 the Dutch minister apes is prohibited (Goodman of education and the director of and Check 2002) the BPRC signed an agreement for the transfer of ownership of fifty- *Although the United Kingdom has had its policy in place since 1997, great apes have not been used in research in that country since 1986. nine chimpanzees to the AAP Sanctuary for Primates and other Exotic Animals (Anonymous 2003). 2002), but it appears that nonin- HSUS, to end invasive research on AAP suffered various delays but vasive research is still allowed. chimpanzees in the United States had secured a site for the sanctu- Table 5 provides a summary of in the coming years. ary and expected construction to international legislation, regula- Regardless of legislative efforts, begin in mid-2005 (AAP Sanctuary tions, and policies. the drastic decline in chimpanzee for Exotic Animals 2005). research in the United States over In June 2003 Sweden’s National the past twenty years is the result Board for Laboratory Animals The Future of of various factors, including the established new regulations that Chimpanzee high cost of keeping chimpanzees ban the use of apes (great apes and in laboratories, public pressure, ) in research (Anonymous Research and evidence of the physical and 2003). The only exception is for Trends in international legislation psychological similarities between the conduct of noninvasive behav- strongly suggest that additional chimpanzees and humans. Trends ioral studies. As was the case in countries will adopt legislation to suggest that the use of chim- New Zealand, great apes were not restrict or end the use of chim- panzees in research in the United being used in research in Sweden panzees (and other apes) in bio- States will continue to decline. when these regulations were being medical research and testing. The Additional efforts to protect chim- implemented, but the rules would U.S. CHIMP Act of 2000 acknowl- panzees, such as legislation to pre- prohibit any such use in the future. edged the special status of chim- vent private ownership of chim- Japan has also taken steps by panzees and human responsibility panzees, legal work to gain banning invasive research on for their lifetime care. There are for chimpanzees, and great apes (Goodman and Check current efforts, including by The inclusion of chimpanzees and

Chimpanzees in Research: Past, Present, and Future 131 humans in the same , are 114: 392–400. 2001. Do chimpanzees know likely continue or expand. In the Cantalupo, C., D. Pilcher, and W.D. what conspecifics know? Animal meantime, the likelihood of prima- Hopkins. 2003. Are asymmetries Behaviour 61: 139–151. tologists providing even more evi- in sylvian fissure length associat- Hopkins, W.D., and C. Cantalupo. dence of the intelligence and emo- ed with the planum temporale? 2004. Handedness in chim- tional capabilities of chimpanzees Neuropsychologia 41: 1975–1981. panzees is associated with asym- will further support the argument Carlsson, H., S.J. Schapiro, I. metries in the primary motor that their use in biomedical Farah, and J. Hau. 2004. Use of but not with homologous lan- research and testing should come primates: A global overview. guage areas. Behavioral Neuro- to an end. American Journal of Primatology science 118: 1176–1183. The authors thank Jennifer Ball, 63: 225–237. Kuhlmeier, V.A., and S.T. Boysen. Leah Nickle, and Stephany Harris for Cavalieri, P., and P. Singer, eds. 2002. Chimpanzees (Pan research assistance for this chapter. 1993. The : troglodytes) recognize spatial Equality beyond humanity. Lon- and object correspondences Note don: Fourth Estate Limited. between a scale model and its 1 Other apes, including gorillas, orangutans, Committee on Animal Models in referent. Psychological Science, and gibbons, were used in the research labora- Biomedical Research. 1995. January: 60–63. tory at one time, but chimps successfully breed in captivity and as adults are smaller Aping science: A critical analysis Limongelli, L., S.T. Boysen, and E. and easier to handle than either gorillas or of research at the Yerkes Regional Visalberghi. 1995. Comprehen- orangutans. Primate Research Center. New sion of cause-effect relations in a York: Medical Research Modern- tool-using task by chimpanzees ization Committee. (Pan troglodytes). Journal of Literature Cited Conlee, K.M. 2003. Great ape Comparative Psychology 109: AAP Sanctuary for Exotic Animals. research: Exposing the hard 18–26. 2005. Breakthrough in Spain: facts and pursuing a ban. AV Matsuzawa, T. 1985a. Use of num- Villena Town Council says yes to Magazine 111: 12–14. bers by a chimpanzee. Nature establishment of Primadomus. Conlee, K.M., E.H. Hoffeld, and 315: 57–59. Electronic newsletter of AAP M.L. Stephens. 2004. A demo- ————. 1985b. Color naming Sanctuary for Exotic Animals. graphic analysis of primate and classification in a chim- March 9. research in the United States. panzee (Pan troglodytes). Jour- Anonymous. 2003. Last European Alternatives to Laboratory Ani- nal of Human 14: research chimps to retire. Labo- mals 32 (Supplement 1): 283–291. ratory Primate Newsletter 42(1): 315–322. McGrew, W.C. 1992. Material cul- 26. January. Gardner, R.A., B.T. Gardner, and ture in chimpanzees. Cambridge: Balls, M. 1995. Chimpanzee med- T.E. van Cantfort. 1989. Teaching Cambridge University Press. ical experiments: Moral, legal, sign language to chimpanzees. Muchmore, E. 2001. Chimpanzee and scientific concerns. Alterna- Albany: State University of New models for human disease and tives to Laboratory Animals 23: York Press. immunobiology. Immunological 607–614. Goodall, J. 1968. The behaviour of Reviews 183: 86–93. Boysen, S.T. 2000. The tooth mon- free-living chimpanzees in the National Aeronautics and Space ster. In The smile of the dolphin, Gombe Stream Reserve. Animal Administration (NASA). 2004. ed. M. Bekoff, 82–83. New York: Behaviour Monographs 1: Ballistic and Discovery Books. 161–311. Orbital Chimpanzee Flights Boysen, S.T., and G.G. Berntson. ————. 1986. The chimpanzees (CHIMP). http://lsda.jsc.nasa. 1989. Numerical competence in of Gombe: Patterns of behavior. gov/scripts/cf/exper.cfm?exp_ a chimpanzee (Pan troglodytes). Cambridge, Mass.: Belknap, Har- index=907. Journal of Comparative Psychol- vard University Press. Nath, B.M., K.E. Schumann, and ogy 103: 23–31. Goodall, J., R. Fouts, F. Patterson, J.D. Boyer. 2000. The chim- Brent, L. 2004. Solutions for E. Katz, M. Bekoff, and B. panzee and other non-human research chimpanzees. Lab Ani- Galdikas. 2003. The Great Ape primate models in HIV-1 vaccine mal 33(1): 37–43. Project census. Portland, Ore.: research. Trends in Microbiology Brown, D.A., and S.T. Boysen. Great Ape Project Books. 8(9): 426–431. 2000. Spontaneous discrimina- Goodman, S., and E. Check. 2002. National Center for Research Re- tion of natural stimuli by chim- The great primate debate. sources (NCRR). 2001. Report to panzees (Pan troglodytes). Jour- Nature 417: 684–687. Congress regarding number of nal of Comparative Psychology Hare, B., J. Call, and M. Tomasello. chimpanzees and funding for care

132 The State of the Animals III: 2005 of chimpanzees, Public Law 106- 23: 621–625. McGrew, T. Nishida, V. Reynolds, 551, Chimpanzees Health Im- Savage-Rumbaugh, E.S. 1986. Ape Sugiyama, Y., Tutin, C.E.G., provement, Maintenance, and language: From conditioned Wrangham, R.W., and Boesch, C. Protection Act. Bethesda, Md.: response to symbol. New York: 1999. Culture in chimpanzees. NCRR. Columbia University Press. Nature 399: 682–685. National Research Council. 1997. Save the Chimps n.d. History: Sav- Wise, S. 2000. Rattling the cage. Chimpanzees in research: Strate- ing space chimps. http:// Cambridge, Mass.: Perseus gies for their ethical care, man- www.savethechimps.org/about_ Books Group. agement, and use. Washington, history.asp. ————. 2002. Drawing the line: D.C.: National Academy Press. Stephens, M. 1995. Chimpanzees Science and the case for animal National Science Board 2002. Sci- in laboratories: Distribution and rights. Cambridge, Mass.: ence and engineering indicators types of research. Alternatives to Perseus Books Group. 2002: Appendix table 7–27. Laboratory Animals 23: 579– Yerkes, R.M., and B.W. Learned. Parr, L.A. 2001. Cognitive and 583. 1925. Chimpanzee intelligence physiological markers of emo- The Humane Society of the United and its vocal expressions. Balti- tional awareness in chimpanzees States (HSUS). n.d. Statement more: Williams and Wilkins Co. (Pan troglodytes). Animal Cogni- of The HSUS on ending biomed- Yerkes National Primate Research tion 4: 223–229. ical research on nonhuman apes. Center. n.d. Innovation and sci- ————. 2003. The discrimina- http://www.hsus.org/animals_in ence: The history of Yerkes. tion of faces and their emotional _research/monkeys_and_apes_in http://www.yerkes.emory.edu/ content by chimpanzees (Pan _research/ending_biomedical_ about_history.html. troglodytes). Annals of the New research_on_nonhuman_apes/ York Academy of Sciences 1000: statement_of_the_hsus_on_ 56–78. ending_biomedical_research_on_ Parr, L.A., and F.B.M. de Waal. nonhuman_apes.html. 1999a. Like mother—like son: Tomasello, M., and J. Call. 1997. Visual kin recognition in chim- . New York: panzees. American Journal of Oxford University Press. Primatology 49: 84–85. United Kingdom Animal Proce- ————. 1999b. Visual kin recog- dures Committee. 1998. Govern- nition in chimpanzees. Nature ment announces end to cosmetic 399: 647–648. testing on animals. Press release Parr, L.A., T. Dove, and W.D. Hop- http://www.apc.gov.uk/press_ kins. 1998. Why faces may be releases/981126b.htm. special: Evidence of the inver- ————. 2001. Report of the Ani- sion effect in chimpanzees. Jour- mal Procedures Committee for nal of Cognitive Neuroscience 10: 2001. London: The Stationary 615–622. Office. http://www.apc.gov.uk/ Premack, D. 1986. Gavagai! or the reference/annrep2001.pdf. future history of the animal lan- van Akker, R., M. Balls, J.W. Eich- guage controversy. Cambridge, berg, J. Goodall, J.L. Heeney, Mass.: MIT Press. A.D.M.E. Osterhaus, A.M. Prince, Premack, D., and A. Premack. and I. Spruit. 1993. Chim- 1983. The mind of an ape. New panzees in AIDS research: A bio- York: Norton. medical and bioethical perspec- Prince, A.M., J. Allan, L. Andrus, B. tive. Journal of Medical Brotman, J. Eichber, R. Fouts, J. Primatology 22: 330–392. Goodall, P. Marx, K.K. Murthy, S. Washburn, D.A., and D.M. Rum- McGreal, C. Noon. 1999. Viru- baugh. 1992. Comparative lent HIV strains, chimpanzees, assessment of psychomotor per- and trial vaccines. Science 283 formance: Target prediction by (5405): 1117–1118. humans and (Macaca Reynolds, V. 1995. Moral issues in mulatta). Journal of Experimen- relation to chimpanzee field tal Psychology: General 121: studies and experiments. Alter- 305–312. natives to Laboratory Animals Whiten, A., J. Goodall, W.C.

Chimpanzees in Research: Past, Present, and Future 133 About the Contributors

Sarah Boysen received her Ph.D. in Jennifer M. Felt graduated from Kristin Kaschner received her 1984 from The Ohio State Universi- the University of Vermont in 1999 M.Sc. from Albert-Ludwigs-Univer- ty. Her current research interests and served for two and a half years sität of Freiburg, Germany, in 1997. are , particularly in the Peace Corps in Honduras. Based at the Underwater Acoustics the acquisition of counting abilities She was program manager for Group at Loughborough University, and numerical competence in non- Latin America and the Caribbean Leicestershire, England, she devel- human primates; cognitive develop- for Humane Society International oped an acoustic analysis tech- ment in the great apes, including (HSI), the international arm of The nique to study the behavior of small attribution, self-recognition, and HSUS, until June 2004, when she cetaceans around midwater trawl intentional behavior; and social became deputy director for trade nets. She joined the Marine Mam- behavior and tool use in captive capacity building at The HSUS. mal Research Unit at the Fisheries lowland gorillas. She is currently Centre at the University of British consulting editor for the Journal of Stephanie Edwards is a 2003 grad- Columbia (UBC) in 1998 and has Comparative Psychology. uate of the University of Maryland been a member of the Sea Around and program assistant and Web Us Project, based at UBC and Kathleen Conlee is director of pro- content manager for HSI. devoted to studying the impact of gram management, Animal Re- fisheries on the world’s marine search Issues, at The HSUS and is Katherine C. (Kasey) Grier is asso- ecosystems, since 1999. Ms. responsible for the organization’s ciate professor in the Department Kaschner was an invited participant work related to nonhuman primates of History, University of South Car- at the International Whaling Com- in research, the use of animals in olina. She is the author of Culture mission Scientific Committee education, and the HSUS Pain and and Comfort: Parlor Making and workshop on bycatch mitigation Distress campaign. She previously Middle Class Identity and the forth- and acoustic deterrents in 1999 worked for seven years at a large coming Pets in America: A History. and is a member of the Cetacean nonhuman primate breeding facili- Her current research focuses on Bycatch Task Force. She has been a ty and a year and a half at a chim- the history of animal-human inter- FishBase collaborator since 2000, panzee and sanctuary. action. She serves as guest curator compiling information about the for an exhibition on the history of acoustical behavior of fish. pet keeping in the United States originating at McKissick Museum, the University of South Carolina, in 2005 and traveling for three years thereafter.

135 Randall Lockwood received his Daniel Pauly acquired his doctorate J.F. Reece received his B.Sc.(Hons.) Ph.D. from Washington University in fisheries biology in 1979 from the in biology from University of York in St. . He is senior vice pres- University of Kiel in Germany. He is and was qualified (B.V.Sc.) from Liv- ident for anti-cruelty initiatives a former division director of the erpool University Veterinary School and training for the American International Center for Living in 1994. He worked for more than Society for the Prevention of Cruel- Aquatic Resources Management three years in rural, large-animal ty to Animals and former vice pres- (ICLARM) in Manila and taught veterinary practice in Devon, Eng- ident/Research and Educational fisheries sciences at the University land. Since 1998 he has been asso- Outreach for The HSUS. He is the of the Philippines. In 1994 he joined ciated with the work of Help in Suf- co-editor of and the Fisheries Centre, University of fering, an animal welfare charity in Interpersonal Violence and co- British Columbia, while remaining Jaipur, India, as a volunteer veteri- author (with Frank Ascione) of ICLARM’s principal science advisor nary surgeon. Since 2002 he has “Cruelty to Animals: Changing Psy- until 1997 and the science advisor been in charge of the ongoing ABC chological, Social, and Legislative of its FishBase project until 2000. Extension Project, sponsored by Perspectives,” which appeared in Since 1999 he has been principal HSI, at Help in Suffering Jaipur. The State of the Animals: 2001. investigator for the Sea Around Us Project. In 2001 he received the Beth Rosen has worked in the Kelly O’Meara is program manager, Murray Newman Award for Excel- HSUS Government Affairs and Eval- Africa and Asia, for HSI. She has lence in Marine Conservation uation and Planning departments promoted humane slaughter prac- Research and the Oscar E. Sette since 2001. She received her mas- tices to government representatives Award of the Marine Fisheries Sec- ter’s degree in public administra- in Indonesia and Vietnam; initiated tion, American Fisheries Society. He tion from New York University. and organized first-of-their-kind was named an honorary professor at workshops on stray dog/street ani- Kiel University in 2002 and elected Andrew N. Rowan is executive vice mal control in Moscow and St. a Fellow of the Royal Society of president, operations, for The Petersburg; and managed a two-year Canada (Academy of Science) in HSUS. He is the author of Of Mice, street dog control program on the 2003. Models, and Men; co-author of The island of Abaco in the Bahamas. A Animal Research Controversy: graduate of the University of - Nancy Peterson, a registered vet- Protest, Process, and Public Policy; chusetts, she holds certificates from erinary technician, is an issues spe- and coeditor of Humane Society the Royal Society for the Prevention cialist in the Companion Animals Press’s State of the Animals series. of Cruelty to Animals (RSPCA) in section of The HSUS and coordina- large- and from tor of the Pets for Life Training Stephanie Shain is director of Bristol University, England, for ani- Centers. Before joining The HSUS, companion animal outreach for mal welfare officer training. She is Ms. Peterson worked in small-ani- The HSUS and one of the organiza- co-author of the HSI report Dogs on mal veterinary hospitals and as a tion’s leading spokespeople on Abaco Island, the Bahamas: A Case trainer of dogs for people with dis- pet-related topics. From 1995 to Study. abilities. Her articles promoting 2000 she worked as assistant pet-friendly rental housing have director of programs for the Amer- been published in numerous hous- ican Anti-Vivisection Society. ing magazines. She was a member of the HSUS staff who collaborated on The Humane Society of the Unit- ed States Complete Guide to Cat Care (co-authored with Wendy Christensen).

136 The State of the Animals III: 2005 Margaret R. Slater is a veterinarian Neil Trent is executive director of and associate professor of epidemi- HSI. A graduate of the law enforce- ology in the departments of Veteri- ment division of the RSPCA, he nary Anatomy and Public Health worked in a number of capacities and Small Animal Medicine and for the RSPCA in England, the Surgery in the College of Veteri- Bahamas, and Australia. He is a for- nary Medicine at Texas A&M Uni- mer field officer and field services versity in College Station, Texas. director for the World Society for The author of Community Ap- the Protection of Animals. He is co- proaches to Feral Cats: Problems, author of “The State of Meat Pro- Alternatives, and Recommenda- duction in Developing Countries: tions, she is frequently invited to 2002,” which appeared in The speak on feral cat issues at profes- State of the Animals II: 2003. sional conferences nationwide.

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