70 Comparative Psychology
study of the reactions of frogs to four chlorides; genetics and
development, such as Yerkes and Bloomfield’s (1910) study
of the reactions of kittens to mice; and learning, such as
Coburn and Yerkes’s (1915) study of crows.
Edward Bradford Titchener dominated psychology at
Cornell University. Although he is often portrayed as having
opposed comparative psychology, he conducted a number of
studies in the field early in his career (Dewsbury, 1997). A
prize student at Cornell was his first PhD, Margaret Floy
Washburn, who later became the second woman elected to
the presidency of the APA. Her most notable contribution to
comparative psychology was her book The Animal Mind
(1908), that went through four editions and was the standard
textbook in comparative psychology into the 1930s. Re-
search at Cornell included a study of vision in fish (M. F.
Washburn & Bentley, 1906) and one on learning in parame-
cia (Day & Bentley, 1911). Even Edwin G. Boring (1912),
future historian of psychology, published a study of phototro-
pisms in flatworms.
The pride of the program at the University of Chicago,
directed by Angell, was John Broadus Watson. Although
Watson became famous later in his career for his writings
on behaviorism, he did work in comparative psychology dur-
ing his younger years. His dissertation, Animal Education
(Watson, 1903), was an early study in developmental psy-
chobiology, as Watson tried to correlate the development of
learning in rats with the development of the nervous system.
Watson also studied imitation in monkeys and spent several
summers studying noddy and sooty terns on the Dry Tortugas
Islands off Florida (e.g., Watson & Lashley, 1915). This study
anticipated some later research in ethology. Many psycholo-
gists who know only his writings on behaviorism are
surprised by his earlier thinking on instinctive behavior
(Watson, 1912). Most of the other students in animal psy-
chology at Chicago worked on rats, although Clarence S.
Yoakum (1909) studied learning in squirrels.
Edward L. Thorndike had a brief, but extremely influen-
tial, career in comparative psychology. After conducting
some research with William James at Harvard, Thorndike
moved to Columbia University, where he completed his PhD
under James McKeen Cattell in 1898. After a year at Western
Reserve University, he returned to Columbia, where he spent
the remainder of his career, most of it as an educational psy-
chologist. His dissertation,Animal Intelligence(Thorndike,
1898), was a classic study of cats learning to escape from puz-
zle boxes; Thorndike (1911) later expanded this work with
the addition of several previously published articles. He be-
lieved that cats used simple trial and error to learn to operate
manipulanda to escape from the compartments in which they
had been enclosed; they kept emitting different behavioral
patterns until one was successful. Further, he believed that
virtually all learning in all species followed the same laws of
trial-and-error and reward (the law of effect). This provided
little impetus for comparative analysis. Thorndike’s major
contribution was the development of precise methods for
careful study of learning in the laboratory. In the tradition of
C. L. Morgan, Thorndike generally sought to explain behav-
ior in terms of relatively simple processes and eschewed no-
tions of insight in creative problem solving. T. Wesley Mills
took a very different approach, closer to that of Romanes than
to that of Morgan. This led to a bitter exchange of mutually
critical articles. Mills emphasized the importance of testing
under natural conditions, writing of Thorndike’s puzzle box
experiments that one might “as well enclose a living man in a
coffin, lower him, against his will, into the earth, and attempt
to deduce normal psychology from his conduct” (Mills, 1899,
p. 266). Thorndike (1899) defended his research as the only
way “to give us an explanatory psychology and not fragments
of natural history” (p. 415).
Karl S. Lashley, best known as a physiological psycholo-
gist, also had a lifelong interest in comparative psychology.
He was influenced by Watson at Johns Hopkins and spent one
summer working with him on the tern project. Lashley influ-
enced comparative psychology not only through his research
and integrative writings but also through his students. Harry
M. Johnson was another Hopkins-trained comparative psy-
chologist, as exemplified in his study of visual pattern dis-
crimination in dogs, monkeys, and chicks (Johnson, 1914).
Other comparative psychologists in graduate school dur-
ing this period included John F. Shepard at the University of
Michigan, who did many studies of learning in ants and rats
(see Raphelson, 1980), and William T. Shepherd at George
Washington University, who worked on a variety of species
(e.g., Shepherd, 1915).
Perhaps the most influential foreign-born comparative
psychologist was Wolfgang Köhler, who completed a doctor-
ate at the University of Berlin in 1909. Much of his career
was devoted to the development and promotion of Gestalt
psychology. His major work in comparative psychology was
conducted on the island of Tenerife in the Canary Islands dur-
ing World War I. Köhler’s best-known studies were of prob-
lem solving with chimpanzees. These studies used such tasks
as the stacking of boxes to reach a banana suspended above
the animals’ enclosure and stick problems in which the chim-
panzees had to manipulate sticks of one sort or another to
reach a banana that was placed outside of the enclosure
where it could be reached with a stick but not without it
(Köhler, 1925).
Little original theory was created during this period. The
guiding theoretical framework came from the theory of