How Comparative Psychologists Study Animal Behavior

Comparative Psychology

How Comparative Psychologists Study Animal Behavior

Problems of genesis concern the evolutionary and developmental histories of behavioral patterns. Those of control relate to the immediate factors, both inside and outside of the animal, affecting behavior.

The con­sequences of behavior can be those for the animal itself, other animals, or the environment, but concern espe­cially the reproductive success of the animal displaying the behavioral pattern.

Such reproductive success, or biological fitness, feeds back to affect the evolution of the behavior and the species.

Psychology is sometimes defined as the study of hu­man behavior. In such a context, the notion of animals as subjects of psychological study appears paradoxical. However, animal research has been a part of psychol­ogy since its differentiation as a distinct branch of in­quiry and has contributed much to the field during that time.

The term comparative psychology has created much difficulty. Through much of the history of the field it has been used in reference to all research on nonhuman animals. The term is better used, however, only in reference to a part of that subdiscipline.

Animal re­search targeted strictly at understanding the process of learning and that aimed at physiological mechanisms are important endeavors but are best treated as differ­ent from true comparative psychology.

The term animal psychology can be used as an umbrella for all three.

The term comparative is, however, somewhat mis­leading in the description of the field. Some authors have proposed that in order to qualify as comparative, research must entail overt comparisons among different animal species. This perspective has been generally re­jected.

What is critical for research in comparative psy­chology is the broad biological framework in which the research is placed. Problems are framed in relation to the evolution and development of behavior conceived broadly. Some early pioneers of comparative psychol­ogy, C.

Lloyd Morgan and Linus Kline, proposed that the field be termed zoological psychology. This term is more descriptive of the comparative psychology that has evolved because the problems studied, modes of thinking.

and interpretations of comparative psychol­ogists overlap those of many zoologists interested in an­imal behavior. Comparative psychologists have differed from zoologists in that their problems have often been generated and interpreted in relation to those of the psychology of human behavior.

Thus, comparative psy­chologists have had to tread a narrow line as they work with a vision of psychological breadth. on the one hand. while placing their work in broad biological per­spective, on the other.

History of Comparative Psychology

Thought concerning the relationship between humans and nonhuman species can be seen early on in intel­lectual history.

Aristotle wrote of the Scala naturae, be­lieving that species could be placed along a single con­tinuum from the lowest to the highest, a view that is rejected today.

Prominent among the many philoso­phers who addressed these issues was Rene Descartes, who believed there to be a fundamental dichotomy be­tween the human soul and the reflex mechanisms gov­erning animal behavior.

The true antecedents of comparative psychology can be seen among British scientists of the nineteenth cen­tury. The work of Charles Darwin was critical in estab­lishing the relationship between human and nonhuman species, thus rendering a comparative psychology reasonable.

Darwin’s protégé, George John Romanes, and C. Lloyd Morgan followed Darwin in applying ev­olutionary principles to the study of behavior. They saw continuity between the instincts and intelligent acts of humans and those of other species. Also in England, Douglas A.

Spalding began experimental studies of the factors important in the development of behavior.

Experimental studies in comparative psychology de­veloped in the United States around the turn of the century. Under the influence of psychologists such as William James at Harvard and G. Stanley Hall at Clark University, psychologists such as Edward L.

Thorndike, Robert Yerkes, Linus Kline, and Willard Small conducted experimental studies of animal be­havior in a psychological context. Margaret Floy Washburn (1908), with a Ph.D.

degree from Cornell University, wrote The Animal Mind, which would be­come the standard textbook in the field for 25 years. John B. Watson, with a University of Chicago Ph.D.

degree, worked in the field of developmental psycho-biology, conducted field studies of the behavioral pat­terns of noddy and sooty terns, and eventually devel­oped the school of behaviorism.

After a bit of a lull during and after World War I. comparative psychology flourished during the 1930s.

Robert Yerkes opened the facility that would become the Yerkes Laboratories of Primate Biology in Orange Park, Florida in 1930, It would become the leading fa­cility for research on the great apes and attract scien­tists from all over the world. Among Yerkes’s associates, H. C. Bingham. Henry Nissen, and C.

Ray Carpenter pioneered field studies of primate behavior, thus open­ing the door to a productive line of interdisciplinary research. During this period T. C. Schneirla began his research, including numerous field studies, on the be­havior of ants; Frank A.

Beach began his long program of work on the neural and hormonal determinants of instinctive patterns in mammals; and Harry E Harlow began work on such problems as development and learning in monkeys. Also during this period various new textbooks, such as E A. Moss’s (1934) Comparative Psychology, appeared. The field was in full swing.

The leaders who matured during the 1930s pro­duced numerous students who continued to develop the field. Three post-World War II developments greatly af­fected comparative psychology.

The first was the strengthening of ties with European ethology, as rep­resented by Konrad Lorenz, Nikolaas Tinbergen, and their colleagues. These interactions increased compar­ative psychologists’ interest in problems of evolution and adaptive significance and in instinctive behavior.

The second was the development during the 1970s of the field of behavioral ecology/sociobiology. This ap­proach worked from such principles as (i) natural se­lection works at the level of the individual animal, and (2) one’s genetic interests are represented in close kin as well as in one’s self.

The study of animal behavior was revolutionized as a result and helped focus theory in psychology on issues of biological currency. The third was the “cognitive revolution.” which resulted in a weakening of the hold of more reductionistic forms of behaviorism on the field.

Although cognitive studies had always been a part of comparative psychology, they moved from the background to the foreground during this period. Today, comparative psychology flourishes as a rather diverse but productive field.

Representative Research in Comparative Psychology

The main research categories in comparative psychol­ogy are four in number. They begin with efforts to iden­tify the evolution of behavioral patterns and continue with understanding the effects of various stimuli, con­sidering the consequences of behavior, and investigat­ing animal cognition.


The study of evolutionary history entails efforts to trace the history of behavioral patterns through their evolution in different species. For exam­ple, Greene and Burghardt (Science, 1978) traced the probable history of different patterns of constriction in ancient and modern snakes.

The development of behavior entails the continuous, dynamic interaction of genes, environment, and the or­ganism in an epigenetic process. Studies of genetic ef­fects on behavior have traditionally concerned inbred strains, genetic selection experiments, and hybridiza­tion.

More powerful techniques have recently become available with the development of modern molecular genetics. The influence of important early experience and the changing baseline on which that experience acts have been the focus of comparative developmental research.

An active research area concerns the ontog­eny of bird song, where genes and environmental input interact in diverse and complex ways in different spe­cies.


Many influences on behavior are contem­poraneous, or nearly so, with the behavior. Many com­parative psychologists study the effects of particular stimuli, such as the displays and calls of various spe­cies, on the individual of concern.

Comparative psychologists are increasingly interact­ing with scientists from related fields to also study in­ternal mechanisms that control behavior. There is much interest in sensory function within the animal’s skin.

The study of the nervous system as it affects and is affected by behavior has become quite prominent with the development of behavioral neuroscience. Hor­monal determinants and consequences of behavior have received much interest.

An important trend is the ability to monitor and manipulate hormonal levels un­der field conditions so that effects can be examined in their full natural context.


Although many consequences of behavior are important, effects on reproductive success have taken center stage because of their importance to evolutionary change.

Such areas as prey-predator in­teraction and reproductive behavior have received much attention because of the obvious relationship to survival and reproductive success.

In the latter case, modern techniques allow the psychologist to determine which of several possible males is the father of young animals. This enables the study of the tie between ag­gressive, sexual, and parental behavior and successful genetic transmission.

Animal Cognition

Studies of animal cognition have been part of comparative psychology for many years, as in the work of Wolfgang Kohler. Walter Hun­ter, and Robert Yerkes. In recent years, however, there has been a rebirth of interest in cognitive studies.

One group of students of comparative cognition works within a behavioristic context in the study of such phe­nomena as attention, memory, timing, concept forma­tion, and counting. Others. following the lead of ethologist Donald Griffin.

are developing a field of cognitive ethology which advocates a return to mentalistic con­cepts such as consciousness, intention. and mind.

Stud­ies of language learning by chimpanzees, such as those of the Gardners with American sign language, Pre-mack with small plastic “words,” and Rumbaugh and Savage-Rumbaugh with symbols in a “Yerkish” lan­guage, and of other species. such as dolphins. parrots, and sea lions. have been especially influential in the field of animal cognition.

Comparative Psychology References:

  1. R. (1984). From Darwin to behaviourism: Psychology and the minds of animals. Cambridge: Cambridge Uni­versity Press. A study of the history of comparative psychology with an emphasis on learning and behavioristic approaches.
  2. D. A. (1978). Comparative animal behavior. New York: McGraw-Hill. A textbook in comparative psy­chology.
  3. D. A. (1984). Comparative psychology in the twen­tieth century. Stroudsburg. PA: Hutchinson Ross. A comprehensive study of the history of comparative psy­chology.
  4. Dewsbury, D. A. (Ed.). (1990). Contemporary issues in com­parative psychology. Sunderland, MA: Sinauer. A selec­tion of research articles by some leading comparative psychologists.
  5. Dewsbury, D. A. (1992). Triumph and tribulation in the history of American comparative psychology. Journal of Comparative Psychology, 106, 3-19. A survey of the dif­ficulties experienced and the accomplishments of com­parative psychologists in the United States.
  6. Maier, R. (1998). Comparative animal behavior: An evolution­ary and ecological approach. Boston: Allyn & Bacon. A recent textbook in comparative psychology.
  7. Moss, A. (Ed.). (1934). Comparative psychology. New York: Prentice Hall. A comprehensive treatment of compar­ative psychology as it was conceptualized in the 1930s.
  8. H. L. (1987). Introduction to comparative cognition. New York: Freeman. A textbook in comparative cogni­tion.
  9. Washburn, M. F. (1908). The animal mind: A text-book of comparative psychology. New York: Macmillan. A stan­dard textbook in comparative psychology during the early part of the century.
  10. E. A. (1993). Comparative cognition: Begin­ning the second century of the study of animal intel­ligence. Psychological Bulletin, 113, 211-228. A position paper for the study of animal cognition near the end of the twentieth century.


Comparative psychology and ethology

How Comparative Psychologists Study Animal Behavior

Comparative Psychology is a sub-discipline of psychology and ethology of biology By Gary Greenberg, PhD (To appear in the forthcoming Encyclopedia of the Sciences of Learning, Springer)

Comparative Psychology and Ethology are both sciences which study animal behavior, typically non-human behavior, though both have often studied humans.

Comparative Psychology is a sub-discipline of psychology and ethology of biology. Both can trace their roots to the late 19th century. Depending on which history one reads the first comparative psychologist was Pierre Flourens, a protégé of Baron Cuvier or George John Romanes, a friend and student of Charles Darwin.

Flourens' book title represented the first use of the term, Comparative Psychology (Psychologie Comparée, 1864) and pre-dated Romanes' Animal Intelligence (1882).

Both proposed a science which would compare animal and human behavior, Romanes postulating the existence of a gradient of mental processes and intelligence from the simplest animals to man — the comparative approach much in use today. Romanes strengthened his proposal by a vast collection of anecdotal accounts of clever behavior in dozens of animal species.

Though perhaps best known today for the fallacies of his anecdotal method and for his easy assignment of human mental faculties to animals—anthropomorphism—Romanes nevertheless succeeded in establishing his idea of a gradient of mental processes across the ani-mal kingdom as a basic premise of early comparative psychology. Ethology too has a mixed parentage.

Isidore Geoffroy-Saint-Hillaire first used the term in 1859, though Oskar Heinroth, a late 19th century German biologist was one of the first to apply the methods of comparative morphology to animal behavior; he is thus considered to be one of the founders of ethology.

Both disciplines had many adherents in the early and middle parts of 20th century: Comparative Psychology in the United States under the influence of the learning psychologists (e.g., Ivan Pavlov, Edward Thorndike), the behaviorists (e.g., Zing-Yang Kuo, John Watson, B. F. Skinner), and the epigeneticists (e.g., T. C.

Schneirla, Daniel Lehrman, Ethel Tobach, Gilbert Gottlieb); while Ethology became firmly established after WWII in Europe under the influence of biologists such as William Thorpe, Nikko Tinbergen, and Konrad Lorenz.

The latter two, in fact, were awarded the Nobel Prize in medicine (there is no separate prize for behavioral research) in 1972 for their animal behavior studies (they shared this prize with Karl von Frisch, an early 20th century biologist).

Theoretical Background

Given the biological roots of both comparative psychology and ethology, evolution was seen to play an important role in behavioral origins by both disciplines, though in different ways. Comparative psychology, strongly influenced by early 20th century Functionalists (e.g.

, William James, John Dewey), believed behavior allowed organisms to adapt to their environments (i.e., Darwinism); behavior itself was not an evolved phenomenon, though the organism was. Thus, as organisms changed through evolution, new or different behavioral potentials arose.

Ethologists, on the other hand, understood behavior itself to be an evolved process, the route being genes—instincts, or inherited behaviors. In later years this one-way route, from genes to behavior, became to be known as the Central Dogma of Molecular Biology.

Additionally, while comparative psychology tended to engage primarily in laboratory research, ethology emphasized the significance and importance of studying behavior outside the laboratory, in the natural settings.

These two fundamentally different approaches to the study of behavior lead to a serious intellectual and theoretical ―war around the 1950s.

Ethology advocated the position that behavior was a biological phenomenon, determined, and not merely influenced by the organism's genotype; much animal behavior was thus believed to be in-stinctive. Indeed, Lorenz, whose mentor was Oskar Heniroth, and Tinbergen spelled out the full meaning of what instinctive behavior was.

The clearest statement of this is found in Tinbergen's book, The Study of Instinct (1951), Comparative psychologists, on the other hand, tended to take an epigenetic approach, stressing the importance of development, experience, and other psychological proc-esses.

The differences were summarized in an important paper by Daniel Lehrman (1953), which today still represents one of the best critiques of instinct theory. While healthy, the ensuing debates settled little.

It was an important 1966 book by Robert Hinde (Animal behaviour: A synthesis of ethology and comparative psychology) that seemed to resolve the differences between these two opposing views. Indeed, a later 1981 book by the ethologist S. A. Barnett (Modern ethology: The science of animal behavior) was able to discuss the discipline without resorting to instinct explanations.

Important Scientific Research and Open Questions

The two disciplines historically sparred over the nature-nurture issue: Was behavior a biological or a psychological phenomenon? Endless debates over this issue have yet to see it formally resolved. Contemporary reports of the discovery of a gene for a behavior are routinely retracted following failures to replicate such findings — but the search continues.

This is as true in psychology as it is in biology, though many in both camps understand behavior to be a biopsychosocial phenomenon. The significance of both psychological and biological development, long ignored, is now seen to be crucial to a full understanding of behavioral origins.

And, though studied now for well over 100 years, there are still new developments to be found in the area of learning.

Current Status

While Comparative Psychology grew in America, ethol-ogy remained somewhat stagnant in Europe. Many still identi-fied with the discipline, though it was clear that they had abandoned the hard nosed biological determinism of the classical ethologists.

Beginning in 1944 with the initiation of the Ameri-can Psychological Association's divisional structure, comparative psychology had a home in Division 6, Physiological Psychology and Comparative Psychology.

In the 1990s, in an effort to attract new members, the division entered into discussion of a name change — the important point for the present discussion was the retention of ―comparative psychology in the new name adopted at the 1995 APA meeting, Behavioral Neuroscience and Comparative Psychology.

While membership in Division 6 was falling, Comparative Psychology as a field of study remained healthy as illustrated by the appearance of several comparative psychology societies in the clos-ing years of the 20th century: The Southwestern Comparative Psychology Association (founded in 1983 by Michael Domjan, Del Thiessen, Steve Davis and Gary Greenberg); the Comparative Cognition Society (founded in 1994 by Ron Weisman, Mark Bouton, Marcia Spetch and Ed Wasserman; and the International Society for Comparative Psychology (founded in 1983 by Ethel Tobach and Gary Greenberg). An even earlier group, the International Society for Developmental Psychobiology, was founded in 1967 by George Collier, Norman Spear, Bryon Campbell, John Paul Scott and others. The an-nual and biennial meetings of these societies attract animal behavior researchers from several disciplines across the globe; their membership is also international. There are, of course, several other such societies in countries around the world.

The picture was not so rosy for ethology which seemed to languish in the same period. This was ly because,―The simple truth is that ethology never did deliver as a science of comparative behavior…(Plotkin, 2004, p. 105). Indeed, in 1989 ethology was declared…dead, or at least senescent.

That is, if you think of ethology in the narrow sense – the study of animal behavior as elaborated by Konrad Lorenz, Nikolas Tinbergen, and Karl von Frisch. It has been quiescent for some time. No exciting ideas were emerging, and data gathering on key issues had lost its direction…

[Barlow, 1989, p. 2].

However, the biological study of animal behavior has thrived well into the 21st century. Ethology was reborn in the early 1970s as a new science, that of Sociobiology (Wilson 1975), the goal of which was to biologicize the social sciences.

But this blatant attempt at understanding animal and human behavior as a purely biological phenomenon was met with scathing criticism (Hull, 1988; Lustig, Richards & Ruse, 2004) from numerous quarters.

The main point of contention centered around the continuing nature/nurture issue and the question of whether behavior, especially human behavior, was the result of genetic and biological determinism. To many opponents of sociobiology, psychology was not a biological science at all, but a uniquely psychological science (e.g., Greenberg, 2007).

The intellectual sparks flew for years, well into the end of the 20th century which witnessed the appearance of a still new iteration of ethology, Evolutionary Psychology.

This approach focuses primarily on human behavior and posited that we owe our universal nature to evolutionary adaptations faced by our Pleistocene ancestors that we have inherited in our genomes. A good source for reviewing the tenets and the research conducted in this field is The Handbook of evolutionary psychology (Buss, 2005).

With evolutionary psychology, instincts are once again in vogue. As with ethology and sociobiology, evolutionary psychology is not without its critics (e.g, Honycutt & Lickliter, 2003). It is not the application of evolution to behavior that is at question, but the manner in which it is understood to apply to behavioral origins.

Evolutionary Psychology, though seen by many to be seriously flawed, is a rather popular orientation in the contemporary behavioral sciences. After all, what serious scientist in 2010 can object to the significance of evolution to psychology?

There has also been new life breathed into ethology and sociobiology. The sociobiological idea of the genetic basis of human altruism has recently been somewhat retracted by one of its earliest proponents, E. O. Wilson.

While this is comforting news to many non-reductionistic comparative psychologists and other animal behaviorists, it doesn't sit well with all students of behavior (Marshall, 2010) attesting to the staying power of the classical ideas of ethology.

In a recent analysis Salzen (2010) makes a case for interpreting the ideas of ethology in modern neuroscientific terms. There is in fact a discipline known as―Neuroethology,‖ which describes animal behavior in terms of how the nervous system works.

As a comparative psychologist, I take comfort in the staying power of my discipline. Its history has been long, though not nearly as tumultuous as that of ethology.


  • Barlow, G.W. (1989). Has sociobiology killed ethology or revitalized it? In P. P. G. Bateson and P. H. Klopfer (Eds). Perspectives in ethology. Volume 8. Whither ethology? (pp. 1-45). New York: Plenum Press.

  • Buss, D. M. (Ed.) (2005). The handbook of evolutionary psychology. Hoboken, NJ: John Wiley.

  • Greenberg, G. (2007). Why psychology is not a biological science: Gilbert Gottlieb and probabilistic epigenesis. European Journal of Developmental Science, 1, 111-121. Hull, D. (1988). Science as a process. Chicago: University of Chicago Press.

  • Lehrman, D. S. (1953). A critique of Konrad Lorenz's theory of instinct. Quarterly Review of Biology, 28, 337-363.

  • Lickliter, R. & Honeycutt, H. (2003). Developmental dynamics: toward a biologically plausible evolutionary psychology. Psychological Bulletin, 129, 819-835.

  • Lustig, A., Richards, R. J. & Ruse, M. (Eds.) (2004). Darwinian heresies. Cambridge, UK: Cambridge University Press.

  • Marshall. M. (2010). Sparks fly over origin of altruism. New Scientist, No. 2780, October 2, 8-9.

  • Salzen, E. (2010). Whatever happened to ethology? The case for the fixed action pattern in psychology. History and Philosophy of Psychology, 12

  • Tinbergen, N. (1951). The study of instinct. Oxford, UK: Oxford University Press.

  • Wilson, E. O. (1975). Sociobiology: The new synthesis. Cambridge, MA: Harvard University Press.

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