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ISSN 0889-3667
IJCP8 5(4)165-209(1992)




Published by

The International Society


Comparative Psychology



Ethel Tobach

The American Museum of Natural History

Central Park West at 79th Street

New York, New York 10024-5192 U.S.A.


Nancy K. Innis

University of Western Ontario

London, Ontfirio, Canada, N6A 5C2


Ruben Ardila Marco Poli

University of Colombia University of Milan

Apartado 88754 Via Francesco Sforza 23

Bogota, Colombia, S.A. Milan, Italy 20122

M. Balakrishnan
University of Kereda
Keiriavattom 695581
Trivandrum, India

Abelardo Brenes
University of Costa Rica
San Jose, Costa Rica

David Croft
P.O.B. 1

University of New South Wales
Kensington, N.S.W. 2033

J.D. Delius

Ruhr Universitat Bochum

Universitat sstr. 150

1 Bochum D4630, Germany

Gary Greenberg

The Wichita State University

Wichita, Kansas 67208 U.S.A.

Robert N. Hughes
University of Canterbury
Christchurch, 1, New Zealand

J. Martin Ramirez
Universidad Complutense Madrid
Enrique Larreta 10
28036 Madrid, Spain

A.F. Semiokhina
Moscow State University
Moscow, 119899

Jiao Shao

Department of Psychology

Peking University

Beijing 100871

People's Republic of China

L.C. Simbayi

Department of Psychology
University of Port Elizabeth
P.O. Box 1600
Port Elizabeth 6000
Republic of South Africa

Joseph Terkel
University of Tel Aviv
69 978 Ramat Aviv
Tel Aviv, Israel

S. Koref-Santibanez
Museum f. Naturkunde
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Kiyoko Murofushi
Ritsumeikan University
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Universitiet van Amsterdam
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Charles W. Tolman
University of Victoria
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Canada V8W 2Y2

D.F. Ventura
Instituto de Psicologia
University de Sao Paulo
Cx. Postal, Sao Paulo
(SP) Brazil


Volume 5, Number 4, Summer 1992


Introduction 165

Gary Greenberg
The Cat (Felis catus) as an Example of the Contribution that
Comparative Psychology has made to Human Factors 167

Duncan A. White
Interactive Models of Cognitive Abilities of Monkeys and Humans
(Saimiri sciureus sciureus; S. boliviensus boliviensus; Homo sap-
iens sapiens) 179
Roger K. Thomas
Human Factors with Nonhumans: Factors that Affect Computer-
Task Performance 191
David A. Washburn
Human Factors, Psychological Factors, and Affirmation of
Continuity 205
Duane M. Rumbaugh and Shelly L. Williams

by the International Society for Comparative Psychology, an affiliate of the International
Union of Psychological Sciences. In consonance with the goals and activities of the Society,
it publishes reports of studies in the evolution and development of behavior of all species;
on the use of the comparative method for the understanding of behavioral processes; and
the examination of the adequacy of psychological and evolutionary theories. It accepts
articles that deal with historical and contemporary relationships between humans and
other animals; that elucidate problems of ecological and behavioral processes in resource
management; that explicate fundamental concepts about human evolution; and that present
research results and develop theories about the development and evolution of behavior.
Articles on the contemporary and historical organization of knowledge about comparative
psychology; promoting public education on the evolution and development of behavior;
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letters discussing published articles with replies by authors; notices of new books, journals,
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MANUSCRIPTS should be submitted in triplicate to Ethel Tobach, Editor. See inside
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bitnet address: [email protected] or Fax 61 49 216 902.

INDEXED OR ABSTRACTED IN: Psychological Abstracts.

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COPYRIGHT 1992 by International Society for Comparative Psychology. Published quar-
terly in the Fall, Winter, Spring, and Summer. Send address changes to International
Journal of Comparative Psychology, % Gary Greenberg, Department of Psychology, The
Wichita State University, Wichita, KS 67208-1595, USA.

ISSN 0889-3667 IJCPE8 5(3)165-209(1992)

International Journal of Comparative Psychology, Vol. 5, No. 4, 1992


Gary Green berg
The Wichita State University

Human factors, a rapidly growing discipline since World War II, is
usually defined as the study of human -machine interactions. One also
sees the term "ergonomics" used in defining the field, and this stems
from the discipline's roots in industrial engineering. Most would agree
that the field today includes the study of human-computer interactions,
artificial intelligence, cockpit design, workplace design, information-pro-
cessing and cognitive processes, robotics, occupational stress, human
error and reliability, etc. It seems clear that this is a discipline for the
psychologist whose interests lie primarily with the human species.

But psychology has a long history of work with non-human animals,
often with an eye towards human applications. The question arises whether
there is a role for the comparative psychologist to play in the human
factors field. This question is addressed in the papers in this special issue.
These papers are based on a symposium. Comparative and Neuropsy-
chological Research in Human Factors Psychology, held at the 1991
American Psychological Society meeting in Washington, D.C. Duncan
White addresses the significance of his work on the visual system of cats
for our understanding of human visual process and proposes an applied
physiological psychology; David Washburn discusses his work with non-
human primates, which are used to model human task performance,
including that which is pertinent to space travel; Roger Thomas discusses
issues of comparative intelligence, indicating the value of research with
monkeys in understanding human intelligence and other cognitive pro-
cesses; finally, Duane Rumbaugh and Shelly Williams provide a com-
mentary about these papers by addressing the idea of continuity in
psychology, i.e., to what extent can we generalize from one species to
another, particularly in humans, in psychology.

I organized this symposium in an attempt to develop a broader defi-

Address correspondence to Gary Greenberg, Department of Psychology, The Wichita
State University, Wichita, KS 67208-1595, USA.

© 1992 International Society of Comparative Psychology 1 65


nition of human factors psychology than is currently in use, something
that even the current president of the Human Factors Society recognized
as necessary (Sheridan, 1991). Greg Moran (1987) recently proposed that
we establish a discipline called Applied Comparative Psychology in which
all human-animal interactions would be appropriate for investigation.
Consistent with this is the idea that the study of human factors would
include investigations of problems involving seeing-eye or hearing-ear
dogs, zoo design for improving the educational mission of those facilities,
the study of the behavior of sea animals that military and sports divers
encounter, and so forth. This seems to be crucial as human factors pro-
grams move from their birthplace in engineering departments of new
homes in psychology departments.


Moran, G. (1987). Applied dimensions of comparative psychology. Journal of Compar-
ative Psychology, 101, 277-281.

Sheridan, T. B. (1991). New realities of human factors. Human Factors Society Bulletin,
34, 1-3.

International Journal of Comparative Psychology, Vol. 5, No. 4, 1992





Duncan A. White
Rhode Island College

ABSTRACT: Human factors is an area of psychology, which systematically applies in-
formation about human behavior to designing environments for human use. The contri-
bution that comparative psychology has made to human factors is demonstrated in this
article using the example of the cat, which shows many of the neurophysiological and overt
behaviors observed in humans. The article begins with a summary of exemplary basic
research which illustrates similarities between the cat and humankind. The summary is
followed by a discussion of various applications of these data to improve the human con-

Human factors (ergonomics; engineering psychology) is the systematic
application of information about human behavior. The objectives of this
discipline are to improve the effectiveness and efficiency of human ac-
tivities, and to improve certain desirable human conditions such as health,
safety, and satisfaction (McCormick & Sanders, 1982). Human factors
applies research data in such issues as development, mobility, vision,
and socialization to realistic settings, modifying the environment to max-
imize the efficiency of human performance (Smith, 1990). Knowing how
the human operates (i.e., the characteristics and needs of the "user") is
fundamental to designing environments for human use, optimizing the
relationship between technology and humankind, and facilitating the
improvement of human life.

Comparative psychology is the study of similarities and differences in
adaptive capacities between human and nonhuman animals, in an at-
tempt to discover general laws of behavior in the context of evolution

Address correspondence to Duncan A. White, Department of Psychology, Rhode Island
College, Providence, RI 02908, USA.

1992 International Society of Comparative Psychology 1 67


theory (Hodos & Campbell, 1969). Through this process, comparative
psychology acquires insights into human behavior.

The cat {Felis catus) is a particularly appropriate species for this
purpose. Cats are relatively tractable, amenable to human interaction
(Imada, Tsukahara, & Imada, 1987; Mertens & Turner, 1988), inexpen-
sive to maintain, and demonstrate many neurophysiological and overt
behaviors observed in humans and other animals. As a result, the cat
offers researchers an opportunity to control and manipulate the internal
and/or external "human" environment, in ways that are ethically sound
but not possible with humans of contingency-deficient computer simu-
lations. Though nonhuman animal models of human behavior have their
limitations (Lockery & Stich, 1989), data from neuroanatomical and
neurophysiological investigations of the cat's central and peripheral ner-
vous systems corroborate what is known about the human.

Research employing the cat to understand humankind is both diverse
and voluminous. The specific areas of research overlap extensively, mak-
ing an organized comprehensive overview elusive. On the other hand,
the extent to which these areas interact indicates the richness of the
contribution that cat-related research has made to our understanding of
the human animal.

The purpose of this article is not to provide an exhaustive review of
the cat literature, but instead to demonstrate the contribution that com-
parative psychology has made to human factors. Toward this end, ex-
emplary basic research illustrating the many similarities between the cat
and humankind will be discussed. Then attention will be given to the
application of these data to improve the human condition.


Learning, Memory, and Problem- Solving

Although the cat has not been used extensively as a research model
to address learning, memory, and problem-solving processes, this general
research area is mentioned first for two fundamental reasons. The first
is the historical significance of Thorndike's puzzle box (1911) and K.U.
Smith's 1930's visual discrimination studies. The second reason is that
cat research, apart from neurological investigations, often uses a learning/
performance task. In these cases the cat has been shown to perform in
ways that are strikingly like human performance under similar condi-
tions. For instance, both species are capable of visually discriminating
between the members of at least seven test item pairs randomly presented
in the same test session. In addition, cats and humans demonstrate a
curvilinear rate of learning these discriminations as a function of test
item complexity, with eight-sided figure pairs being the easiest (White
& Ward, 1988). Bourassa and Weiden's (1985) investigations of orienting


responses and detection of thalamic stimulation also demonstrated that
mechanisms of perceptual learning in the cat are consistent with those
reported in humans. Bourassa and Weiden concluded by suggesting that
input to cerebral cortex from sensory organs in either species is not a
sufficient condition to produce conscious experience or discrimination.


The most extensive, and probably the most well-known, use of the cat
as a model of human behavior is in the area of vision research. For
example, behavioral experiments have revealed that cat and human spa-
tial vision are comparable in several important ways. Both species possess
stereoptic vision (Fox & Blake, 1971), both resolve about a five octave
range of spatial frequencies (Blake, Cool, & Crawford, 1974), both show
a trade off" between spatial resolution and temporal resolution (Blake &
Camisa, 1977), and both demonstrate a decline in acuity with retinal
eccentricity (Blake & Bellhorn, 1978). Blake (1988) has noted that many
of the hallmarks of mammalian vision were first discovered and explored
in the cat. Some notable examples are retinal ganglion cells' center/
surround organization (Kuffler, 1953), cortical neurons' orientation se-
lectivity and massed binocularity (Hubel & Wiesel, 1962), and the ex-
istence of parallel pathways originating with retinal X-, Y-, and W-cells
(Shapley & Perry, 1986). Other similarities between the human and cat
visual systems, resulting from analogous neuroanatqmical and neuro-
physiological components, include contour and movement adaption, af-
tereffects, and recovery responses to prolonged stimulation with moving
test items (Vautin & Berkley, 1977). These results continue to encourage
the use of the cat to determine trigger features of human visual neurons.

Similarly, neural mechanisms for velocity discrimination in the cat
have been suggested as a model for human motion perception (Orban,
Kennedy, & Maes, 1981). Responses of feline X-cells of the LGNd to a
small flashing test probe are likewise comparable to results from similar
human psychophysical tests used clinically (Essock, Lehmkuhle, Fras-
cella, & Enoch, 1985). These X-cell responses indicate that the size of a
concentric background can have a pronounced influence on the sensitivity
of both human and cat observers to a small probe.

Still other areas of cat vision research have addressed human depth
perception (Mitchell & Baker, 1973) and flicker perception (Tyler, 1975),
saccade awareness suppression (Riggs, Merton, & Morton, 1974) and
main sequence saccades (Nelken, Heit, & Bridgeman, 1981), motion de-
tection (Pasternak & Merigan, 1980), contrast sensitivity (Albrecht &
Hamilton, 1982), the effects of positive and negative lens aberrations
(Sivak & Kreuzer, 1983), vernier acuity (Swindale & Cynader, 1986),
discrimination ability as a function of texture (Wilkinson, 1986), spatio-
temporal aspects of the visual scene (Stanford, 1987), and subjective


contour discrimination (Orban, De Weerd, & Vandenbussche, 1990). In
all these areas, the cat has displayed visual performance similar to that
of humans.

Principles of visual development in cats also apply to humans. For
example, Mitchell (1989) noted that an abrupt onset of stereopsis, and
possibly vernier acuity, has been observed in both species; and visual
acuity, as assessed with grating, develops gradually in primates and kit-
tens. Similarly the development of cognitive performance, such as object
permanence, has been observed in kittens and the flexibility of this
animal's cognitive ability has been displayed, for example, by novel prob-
lem solving of visible displacement tests (Dumas & Dore, 1989). Though
analogous behavior does not ensure analogous underlying mechanisms
responsible for those behaviors (Innis & Staddon, 1989), similar behaviors
do suggest similar adaptive capabilities and provide useful predictive
models of behavior.


Though less attention has been given to the cat as a model for human
audition, useful similarities exist. For example, comparable cat and hu-
man auditory vertex potentials have been found on a number of neu-
roanatomical and neurophysiological dimensions (Buchwald, Hinman,
Norman, Huang, & Brown, 1981) and suggest that scalp-recorded fre-
quency-following responses could be used to ascertain low-frequency
hearing sensitivity in uncooperative humans (Gardi, Merzenich, & Mc-
Kean, 1979). Similarly, psychophysical methods investigating pitch per-
ception of the cat have produced results which parallel human data
(Chung & Colavita, 1976). Other auditory research employing the cat as
a model for human behavior has been conducted in areas of binaural
interaction (Hoppe & Langford, 1974), interaural intensive and temporal
disparities (Wakeford & Robinson, 1974), sound localization (Kuwada,
Yin, & Wickesberg, 1979), and auditory nerve-fiber responses to spoken-
stop and nasal consonant-vowel syllables presented in four diff'erent lev-
els of speech-shaped noise (Geisler & Gamble, 1989). This latter study
found that consonants, being of smaller amplitude, are more affected by
noise than are vowels.

Peripheral Nervous System

The peripheral nervous system of the cat has been found to be a valid
model for neural mechanisms of human tactile and vibrotactile sensation
(Hamalainen, 1983). The cat has also been used as a model to reconstruct
the functional events occurring in nerves at the site of stimulation when
human subjects reported pain relief (Swett & Law, 1983). In addition,
the cat model has been used in kinesthetic research to acquire insights


into information regarding body part position, location, motion, speed,
and direction (Burgess, Wei, Clark, & Simon, 1982). Such information
is thought to contribute to body image (Gregory, Morgan, & Proske,
1988) and cognitive map construction (Mergner, Anastasopoulos, Becker,
& Deecke, 1981).

Other Areas

Other areas of cat research, resulting in a better understanding of
human (i.e., "user") characteristics, include: gustation (Boudreau, Ora-
vec, & White, 1981), motor function (Frederickson, Smylie, Howell, &
Lenig, 1978), split-brain behavior (Lepore, Ptito, Provencal, & Bedard,
1985), stroboscopic motion perception (de Bruyn & Orban, 1989), time
perception (Macar, Vitton, & Requin, 1984), emotions (Bear, Rosen-
baum, & Norman, 1986; Piazza, Crescimanno, Benigno, & Amato, 1986),
sleep (Koridze & Nemsadze, 1982; Sinton & Petitjean, 1989), and evo-
lution (Rush, 1988).

Comparative Research Summary

This overview of exemplary basic research illustrating similarities be-
tween cats and humans shows how the cat is a rich and varied source of
knowledge about human neural functioning and behavior. As such, the
cat is a valuable comparative model for suggesting useful modifications
in environmental contingencies that result in improved human perfor-
mance. To date, medicine has benefited most from the cat as a research
model. These applications will be discussed along with other issues such
as psychological, social and ecological welfare.


Medical Models

The similarities between cat and human capabilities and behavior have
been exploited to create better human conditions. For example, the cat
is used extensively as a medical model to gain insight into the nature
and treatment of human physiological and behavioral abnormalities. To
illustrate, the cat is a valuable clinical model for understanding the neural
mechanisms of cardiovascular disorders that may commonly accompany
psychological stress (Tashiro, Tanaka, Fukumoto, Hirata, & Nakao, 1986).
It has also been used to assess recovery following mild to moderate head
trauma (Hayes, Clifton, & Kreutzer, 1989) and plasticity following gross
and selective insult to the nervous system (Burgess, Villablanca, & Le-
vine, 1986; Cornwell, Herbein, Corso, Eskew, & Warren, 1989). Other
examples of the cat's substantial clinical impact are in developmental


monocular deprivation. The regimens of part-time reverse occlusion, which
optimize recovery from the visual deficits induced by monocular depri-
vation in kittens, are similar to the patching of a good eye in order to
induce the strengthening of a "lazy eye." This therapy is also like the
patching regimens that promote the development of good vision in human
infants following early corrective surgery for congenital unilateral cat-
aracts (Mitchell, 1989). In addition, the cat has been used to develop a
visual prosthesis for blind humans (Pollen, 1977) based on analogous
cellular activity underlying visual perceptual events. Furthermore, sim-
ilar visual pathways (i.e., "M" for fast-moving course visual forms and
"P" for spatial detail of stationary or slow-moving visual forms) have
clinical implications for diseases like glaucoma, Alzheimer's, and aniso-
metropic amblyopia (Bassi & Lehmkuhle, 1990). Other clinical appli-
cations take advantage of the fact that P300, a measure of specific cortical
activity, which is characteristic of both cats and humans, is a key to
aging and disease processes. For instance, P300 changes functionally with
age and is missing in Alzheimer's (Harrison & Buchwald, 1985).

Similar neurological investigations using cats have provided insights
into other diseases such as "locked-in" syndrome (Zernicki, 1986) and
hyperkinetic syndromes like Parkinson's Disease, Huntington's Chorea,

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