I. ANIMAL BEHAVIOR AND THE SENSES

Since the behavior of an animal is conditioned by its senses, it is extremely important that the comparative psychologist should have accurate and detailed knowledge of the sense-impressions received by his subjects. Knowledge of the so-called "special senses" does not suffice for the satisfactory description of behavior, for there are several other kinds of sense-data of equal or even greater importance than those of the five special senses. As investigation of the subject progresses the banefulness of the notion that all sense-experience is summed up in "the five special senses" becomes more and more evident. For the comparative psychologist the senses are not five, six, eight, or ten, but as numerous as are the kinds of sense-data which condition animal activities. There can be no doubt that many of the lower animals are largely dependent upon senses which are not included in the conventional special sense list. The contention that certain organs which are commonly recognized as sensory in function, the cristæ acusticæ of the ear, for instance, are merely reflex control organs, has little weight in this connection, for every sense-organ is part of a motor control mechanism, and so far as we are able to judge from available evidence each has as an accompaniment of its functioning a mode of sensation. If there are two kinds of peripheral organs in connection with the afferent nerves, namely, those whose functioning has sensation for an accompaniment and those in which motor control is the sole phenomenon, it is high time that the fact were definitely known.

Even a thoroughly accurate knowledge of the general condition of the senses in a particular phylum, genus, or species may be of trifling value in the study of the behavior of a given individual, for within these groups the state of development and relative importance of a sense may differ strikingly. Herrick,[140] in his admirable investigation of the sense of taste in fishes, has rendered comparative psychology an important service by showing that even a highly developed sense may be of markedly different value in the associative life of different species. The cat-fish, according to Professor Herrick's observations, obtains its food primarily by the aid of taste-impressions, the hake by the aid of touch, and the sea-robin chiefly by means of vision. All three of the senses mentioned are possessed by each of the fishes, yet their values differ so widely that an understanding of the habits and associative processes of any one of the species would be impossible except in the light of just such facts as Herrick has discovered. Clearly, then, we must know the relative importance of the various sense-impressions received by an animal before we can discuss its behavior or psychic characteristics intelligently.

Furthermore, if behavior is to be serviceably described in terms of stimuli and physiological conditions it is necessary first of all to recognize that an animal responds to a situation, not to any one independent and isolated stimulus. Every situation, to be sure, may be analyzed into its component simple stimuli, but the influence of each and all of these stimuli is conditioned by the situation. Too often in our accounts of an animal's behavior we name some one stimulus as the condition of the reaction and entirely neglect the situation, without which the stimulus would have been of quite different value to the animal. For any given stimulus other external and internal stimuli constitute an environment. The complete description of a reaction demands knowledge of all the stimuli which enter into the situation and of their mutual relations of interference or supplementation. A frog which in its native habitat and undisturbed by an unusual situation would react violently to the light touch of a stick may give no sign of reaction to the same stimulus when a human being stands nearby. The influence of the tactual stimulus has been changed entirely by the simultaneous appearance of visual, olfactory, and possibly still other sense-data (man). The animal reacts not to the touch alone, but to this stimulus as part of a certain situation. The general effect of a situation we often speak of as excitement, timidity, etc. These are words for which must be substituted in our accounts of animal behavior accurate descriptions of the situations. Experimental studies prove that an animal must become thoroughly accustomed to the general situation in which it is to be observed before the influence of any particular condition can be studied to advantage.

Only a few of the important reactions of an animal to either external or internal stimuli are visible to the casual observer, and many of them can be detected only by the employment of indirect methods. Frequently the lack of a visible motor response to a new situation is good evidence of a fundamentally important reaction. The death-feigning opossum, crustacean, or insect, truly reacts by becoming motionless. As Whitman[141] has shown in the case of the leech it is as hazardous to judge of the degree of sensitiveness of another animal solely on the basis of our own as it is to maintain that lower animals possess only the senses which are ours. Varied, indirect and delicate methods are necessary in the investigation of the senses, as the results of the experiments to be described below help to prove.

It is my purpose in this paper to call attention to and emphasize the importance of studying stimuli in their mutual relations of interference and supplementation. This I shall do by presenting the results of an investigation of the behavior of the green frog. I shall discuss briefly, first, the sense-data received by the animal, their relative importance, significance, and mutual relations, and, next, the phenomena of reënforcement and inhibition.

II. THE SENSORY REACTIONS OF THE GREEN FROG

The following sensory reactions have been observed in the frog, but most of them have not been studied with care: olfactory, temperature, visual, tactual, equilibrational, and auditory. It is my purpose to investigate each of these senses in such fashion that we shall know the receptive capacity of the animal. Thus far I have completed only the work on auditory reactions, but the chemical and temperature senses and vision will be discussed in similar fashion later.

At present there is little known concerning the chemical senses. Unpublished observations made by Mr. Sherwin in this laboratory indicate the existence of olfactory sensitiveness to camphor, iodine, and several other strong stimuli. The reactions to the stimuli were slow, however, and there is no reason to believe that the sense of smell is of great importance to the animal. Of taste barely more is known than that it is present.