Having satisfied myself that motor reactions for reaction-time measurements could not be gotten to any ordinary sounds in the laboratory, I tried the effect of the reflex croaking of another frog of the same species. In attempting to get frogs to croak regularly, I tested the effect of removing the brain. The animals are said to croak reflexly after this operation whenever the back is stroked; but for some reason I have never been successful in getting the reaction uniformly. In many cases I was able to make normal animals croak by rubbing the back or flanks, and to this sound the animals under observation occasionally responded by taking what looked like an attitude of attention. They straightened up and raised the head as if listening. In no case have other motor responses been noticed; and the above response was so rare that no reaction-time measurements could be made.
Again, while working with the green frog on habit formation, I one day placed two animals in a labyrinth from which they could escape by jumping into a tank of water. Several times when one frog jumped into the water I noticed the other one straighten up and hold the 'listening' or 'attentive' attitude for some seconds. As the animals could not see one another this is good evidence of their ability to hear the splash made by a frog when it strikes the water.
B. Influence of Sounds in Nature.
In order to learn how far fear and artificial conditions were causes of the inhibition of response to sounds in the laboratory, and how far the phenomenon was indicative of the animal's inability to perceive sounds, I observed frogs in their native haunts.
By approaching a pond quietly, it is easy to get within a few yards of frogs sitting on the banks. In most cases they will not jump until they have evidence of being noticed. Repeatedly I have noted that it is never possible to get near to any frogs in the same region after one has jumped in. In this we have additional proof that they hear the splash-sound. To make sure that sight was not responsible for this on-guard condition in which one finds the frogs after one of their number has jumped into the water, I made observations on animals that were hidden from one another. The results were the same. I therefore conclude that the splash of a frog jumping into the water is not only perceived by other frogs in the vicinity, but that it is a peculiarly significant sound for them, since it is indicative of danger, and serves to put them 'on watch.'
A great variety of sounds, ranging in pitch from a low tone in imitation of the bull frog's croak to a shrill whistle, and in loudness from the fall of a pebble to the report of a pistol, were tried for the purpose of testing their effects upon the animals in their natural environment. To no sound have I ever seen a motor response given. One can approach to within a few feet of a green frog or bull frog and make all sorts of noises without causing it to give any signs of uneasiness. Just as soon, however, as a quick movement is made by the observer the animal jumps. I have repeatedly crept up very close to frogs, keeping myself screened from them by bushes or trees, and made various sounds, but have never succeeded in scaring an animal into a motor response so long as I was invisible. Apparently they depend almost entirely upon vision for the avoidance of dangers. Sounds like the splash of a plunging frog or the croak or pain-scream of another member of the species serve as warnings, but the animals do not jump into the water until they see some sign of an unusual or dangerous object. On one occasion I was able to walk to a spot where a large bull frog was sitting by the edge of the water, after the frogs about it had plunged in. This individual, although it seemed to be on the alert, let me approach close to it. I then saw that the eye turned toward me was injured. The animal sat still, despite the noise I made, simply because it was unable to see me; as soon as I brought myself within the field of vision of the functional eye the frog was off like a flash.
Many observers have told me that frogs could hear the human voice and that slight sounds made by a passer-by would cause them to stop croaking. In no case, however, have such observers been able to assert that the animals were unaffected by visual stimuli at the same time. I have myself many times noticed the croaking stop as I approached a pond, but could never be certain that none of the frogs had seen me. It is a noteworthy fact that when one frog in a pond begins to croak the others soon join it. Likewise, when one member of such a chorus is frightened and stops the others become silent. This indicates that the cessation of croaking is a sign of danger and is imitated just as is the croaking. There is in this fact conclusive evidence that the animals hear one another, and the probability is very great that they hear a wide range of sounds to which they give no motor reactions, since they do not depend upon sound for escaping their enemies.
The phenomenon of inhibition of movement in response to sounds which we have good reason to think the frogs hear, and to which such an animal as a turtle or bird would react by trying to escape, is thus shown to be common for frogs in nature as well as in the laboratory. This inhibition is in itself not surprising, since many animals habitually escape certain of their enemies by remaining motionless, but it is an interesting phenomenon for the physiologist. We have to inquire, for instance, what effects sounds which stimulate the auditory organs and cause the animal to become alert, watchful, yet make it remain rigidly motionless, have on the primary organic rhythms of the organism, such as the heart-beat, respiration, and peristalsis. It is also directly in the line of our investigation to inquire how they affect reflex movements, or the reaction time for any other stimulus—what happens to the reaction time for an electrical stimulus, for example, if a loud noise precede or accompany the electrical stimulus.
For the purpose of determining the range of hearing in the frog, I was driven to study the influence of sounds upon respiration. Although the animals did not make any detectable movement, not even of an eyelid, in response to noises, it seemed not improbable that if the sounds acted as auditory stimuli at all, they would in some degree modify the form or rate of the respiratory movement.