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ANIMAL BEHAVIOR
m
O;
II
I O
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STUDIES IN
ANIMAL BEHAVIOR
S. J. HOLMES, Ph.D.
ASSOCIATE PROFESSOR OF ZOOLOGY, UNIVERSITY OF CALIFORNIA
3
4 Preface
39677
STUDIES IN ANIMAL BEHAVIOR
STUDIES IN ANIMAL
BEHAVIOR
i
LXVI.)
"Sense," he says, "is found in all animals, but ani-
mals other than man have no intellect; which is
proved by this, that they do not work, like intellec-
tual agents, in diverse and opposite ways, but just
as nature moves them to fixedand uniform specific
activities." "Sense iscognizant only of singulars,
but intellect is cognizant of universals. Sensory
Animal Psychology, the Old and the New 15
species.
Without raising the question as to whether the
vitalisticexplanation of adaptation really explains
anything, it might be remarked that, since a teleo-
logically working principle is assumed as the basis
for the acquired adjustments of the individual, the
same principle might also be evoked to guide the
entire course of evolution without appealing to an
last
wary. To
find out what probably goes on in an
animal's mind requires close and continuous obser-
vation, and usually experiments under carefully con-
trolled conditions. The careful experimental work
of Thorndike, Cole, Yerkes, Hobhouse, Small and
many other investigators, has given us a more ex-
act knowledge of the mental activities of the ani-
mals studied than would have been possible through
the collection of any quantity of scattered observa-
tions. The results have sometimes proven disap-
pointing to zealous champions of the high mental
34 Studies in Animal Behavior
ceptions the birds sit upon the eggs until they are
hatched. There has been much speculation concern-
ing the origin of this curious instinct of incubation.
As Whitman has remarked, "the incubation instinct
was supposed to have arisen after the birds had ar-
42 Studies in Animal Behavior
rived and laid their eggs, which would have been left
to rot had not some birds just blundered into cud-
dling over them and rescued the line from extinc-
tion." A little reflection makes it evident that such
an origin is clearly impossible, and that we must go
back probably to the cold-blooded reptilian ances-
tors of the birds for the beginnings of the instinct.A
comparative survey of the behavior of the more
primitive animals toward their eggs makes it prob-
able that the instinct of incubation grew out of the
instinct to remain or near the place where the
in
of evolution. The
suckling of young presupposes a
certain tolerance, if not regard, on the part of the
mother for her offspring. Had there not been among
the ancestors of the mammals a fairly close associa-
tion between the parents and their offspring during
the infancy of the latter there obviously could not
have been evolved the mammary glands and other
adaptations for suckling the young which are among
the most fundamental and distinctive features of
mammalian structure. Instinct and organization are
REFERENCES
BAIN, A. The emotions and the will. 4th ed.,
1899.
DARWIN, C. The descent of man. N. Y., 1874.
FABRE, J. H. Souvenirs entomologiques, T. 9.
FISKE, J. The meaning of infancy. N. Y., 1909.
HOLMES, S. J. Observations on the habits and
natural history of Amphithoe longimana. Biol.
Bull. 2, 165, 1900.
REFERENCES
BOHN, G. ( i ) La naissance de Fintelligence,
Paris, 1909; (2) La nouvelle psychologic animale,
Paris, 1911.
COLE, L. An
experimental study of the im-
J.
REFERENCES
BANCROFT, F. W. Heliotropism, differential sen-
sibility, and galvanotropism in Euglena. Jour. Exp.
Zool. 15, 383, 1913.
BRUNDIN, M. T. Light reactions of terrestrial
amphipods. Jour. Animal Behavior, 3, 344, 1913.
HAKFER, E. H. Reactions to light and mechani-
cal stimuli in the earthworm Perichaeta bermudensis
93
94 Studies in Animal Behavior
ERRATUM.
Page 103, line 5 and line 9,
change phototactic to gcotactic.
n
t< * miwiucu uy many ot the larvae
of Polygordius and by certain species of amphipods,
both of which are changed from positive to negative
at an unusually high temperature. Many forms
which are changed from positive to negative by ex-
posure to light are changed more quickly at a higher
temperature, but the swarm spores of algae appar-
ently form an exception to this rule. There are
many organisms in which the sense of the photo-
tactic reaction remains the same at all intensities of
light, and in these cases temperature usually has no
The Reversal of Tropisms 103
REFERENCES
BANCROFT, F. W. ( i ) The control of galvano-
tropism Paramoecium by chemicals.
in Univ. of
Calif. Pubs. Physiol. 3, 21, 1906; (2) Heliotro-
pism, etc., in Euglena. Jour. Exp. Zool. 15, 383,
of proof is on M. Bohn.
126 Studies in Animal Behavior
the environment.
The which have been made of primitive
studies
of a lower one.
The step from sensori-motor association to the as-
sociation of ideas is not, I believe, a wide one, and
comes about as a natural consequence of the elabo-
rateness and what Hobhouse has designated as the
"articulateness" of the mental process of adjustment.
It is foreign to our purpose, however, to trace the
ological !
V
The Beginnings of Intelligence 133
physiological state x
stimulus reaction pain inhibition
physiological state y
perience.
Animals in the course of their instinctive responses
encounter stimuli which bring about other responses.
These become associated. According to the nature
of the nervous pathways involved, there may be re-
inforcement of, or interference with the original re-
action. Experience brings about an extension of the
REFERENCES
BAIN, A. The
and the intellect. 3d ed.,
senses
JENNINGS, H. S. (
i
) Contributions to the study
of the behavior of lower organisms. Carnegie
Inst. Pubs.,1904. (2) Modifiability in behavior.
I, Behavior of sea anemones. Jour. Exp. Zool. 2,
447, 1905. (3) Behavior of lower organisms. N.
Y., 1906.
SPFNCER, H. Principles of psychology. N. Y.,
1892.
WASMANN, E. Instinct and intelligence in the
animal kingdom. St. Louis, 1903.
WHITMAN, C. O. Animal behavior. Biol. Lec-
'
I
A
HE essential nature of the process of learning
*-
constitutes a problem of such fundamental im-
are harmful.
"Second: A mechanism in the neurones gives
behavior of the animal as a whole that
results in the
seem beyond mechanism. By their unmodifiable
abandonment of certain specific conditions and re-
tention of others, the animal as a whole can modify
its behavior. Their one rule of conduct causes in
him a countless complexity of habits. The learning
of an animal is an instinct of its neurones."
this is
probably because the incoming stimuli be-
come connected with other neurones than the ones
which gave the original motor expression. In other
cases it may be due to quite different organic ad-
it.
Satisfying and annoying states do not affect the
reactions by which these feelings are immediately
aroused, but other reactions performed in close tem-
poral relation to them.
The various theories put forward to explain the
learning process as an effect of the physiological cor-
relates of agreeable and disagreeable states are all
ganization.
Profiting by experience in an animal of a primi-
tive type of intelligencewe may conceive, then, to
take place as follows: The creature is endowed
with the capacity for responding to beneficial stimuli
Considerations on the Problem of Learning 153
'
I
A
HE concept of trial and error one that has
is
*
played a prominent role in modern writings on
comparatively psychology, and especially those which
concern themselves with the problem of how be-
havior comes to be adaptively modified. The activi-
ties of animals must obviously be so shaped as to
REFERENCES
BAIN, A. The senses and the intellect. 3d ed.,
1894.
BALDWIN, J. M. Mental development. Meth-
ods and processes. 2nd ed. N. Y., 1903.
References 165
*
I
A
HE idea has been recently emphasized that in
*
many cases the structure of an organism is, to
a considerable extent, the effect of its behavior. We
have, of course, long been familiar with the fact
of functionally produced modifications, but it is
only of late that behavior has been brought forward
as a factor of importance in development, regenera-
tion and other modes of form regulation. If the
creasing thickness
in towards the edges and with
a rounded outline. The mechanical conditions re-
sulting from the movement of the wax through the
water are not widely different from those which the
undulating margins of Leptoplana produce. If the
wire axis of the wax be considered as the longi-
tudinal axis the effect of movement through the
water is lateral extension. In Leptoplana the un-
dulating movement is confined chiefly to the lateral
regions in the anterior third of the body and it
side is
gradually stretched out so as to form the
oral side and anterior end of the new individual.
In the process of regeneration here followed the
172 Studies in Animal Behavior
REFERENCES
CHILD, C. M.
(i) Studies on regulation. IV.
Some experimental modifications of form regula-
tion in Leptoplana. Jour. Exp. Zool. i, 95, 1904.
(2) The regulatory change of shape in Planaria
dorotocephala. Biol. Bull. 16, 277, 1909. (3)
176 Studies in Animal Behavior
behavior. In many
crustaceans, fishes, amphibians
and reptiles thepigment cells appear at times to
be richly provided with branched processes, and
at other times nearly all the pigment appears to be
concentrated in a rounded mass. These changes
often produce marked changes in the color of the
animal. They are to a certain extent under the
control of the nervous system, but they may also
take place independently of nervous influence. The
writer has succeeded in isolating pigment cells from
goal?
One widely accepted theory of the development
of the nerve fiber is that it represents an outgrowth
from the nerve cell. Harrison by cultivating in
REFERENCES
'
I
A
HE so-called instinct of feigning death is one
-*-
which is very widely distributed in the animal
tras, the first day they emerged from the egg and
while their appendages were still soft and easily
bent, showed the same death feigning instinct as
the adults, although they did not persist in it for
so long a time.
The water-bug Belostoma usually feigns with the
legs closely pressed to the thorax or else held folded
at right angles to the body. In Nepa, on the other
hand, the attitudes assumed seem to depend mainly
on the position of the legs just previous to the
death feint, so that it is difficult to distinguish a
REFERENCES
male casts off her skin, when the sperms are depos-
The Recognition of Sex 223
tinued my observations."
That the male amphipods do not distinguish the
females by sight was shown by blacking over the eyes
of several males and then placing them in a dish with
females. It was not long before each male had
however, the
if possibility was sufficiently consid-
ered that sex recognition may be a result of the
behavior of the two sexes, much as it was found to
be in amphipods. Despite the interesting experi-
ments of Goltz the matter requires further investi-
gation before a decided conclusion can safely be
drawn.
In the birds the sexes may often be easily dis-
tinguished by sight, and in many species each sex
is
sight. A
certain familiarity with the habits of street
curs will convince one, I think, that the element
of behavior, as in some of the cases previously de-
scribed, plays a certain role also.
The recognition of sex has been little analyzed
in the mammals. The problem is more complex than
in lower forms owing to the higher development of
the mammalian mind, and the fact that several dif-
ferent senses are usually involved.
REFERENCES
ANDREWS, E. A. Conjugation in the crayfish,
Cambarus affinis. Jour. Exp. Zool. 9, 235, 1911.
BANTA, A. M. Sex recognition and the mating
behavior in the wood frog, Rana sylvatica. Biol.
Bull. 26, 171, 1914.
/
I
A
HE reason for the existence of sex is one of
*
those biological problems whose solution seems
as remote as it did a century ago. Many remark-
able discoveries have been made in regard to the
239
240 Studies in Animal Behavior
nail and got the apple, as she did also at the next
trial.
and let the nut drop out. She was too busy trying
to reach it with her fingers to get it by the easiest
method. Even after she had come to get the nut
rather quickly she often spent considerable time in
attempting to reach the nut when the bottle was held
254 Studies in Animal Behavior
let the nut drop out unobserved. When the nut was
replaced Lizzie tried to get it by biting the glass;
in the course of her turning the bottle around, the
nut dropped out and she picked it up. When an-
other nut was put in Lizzie reached in at once and
got it. Another nut was obtained in the same way,
but at the next trial she turned the jar around in
various ways until the nut fell out; and in numerous
and teeth, and then reached in and got the nut, hold-
ing the jar upright with her feet. The next trial
resulted in practically the same way. The cap of
the jar was then screwed -on farther. Lizzie at-
tacked the jar industriously and finally removed the
cover, although working quite unsystematically. The
cover was put on as before and Lizzie worked at
it about fifteen minutes, getting more and more ex-
put the food through the door and she opened the
door at once and got the food. She did the same
at the second trial, after which, when the food was
turning the box over and over until she became dis-
couraged. I recalled her to the task by tapping on
the box, but evoked only feeble efforts. When I
opened and closed the door Lizzie observed me and
went at once to the door and got the apple. Then
I replaced the apple and closed the door and put the
box in another position. Lizzie attacked the box
in various places and then desisted. Soon she looked
at the box, went to it as if an idea struck her, and
tried to pull the door open by using hands and teeth ;
263
264 Index