There is marked sensitiveness to variations in temperature, as I have demonstrated by preliminary test experiments, but the limits, distribution, and significance of this sensitiveness remain to be investigated. I am not aware that the existence of temperature spots has been determined. In connection with a study of the reactions of frogs to light Torelle[142] discovered that the animals suddenly become inactive and usually attempt to bury themselves when brought into a temperature of 8° to 10° C. This reaction is prompt and definite; its value to an animal which hibernates is evident, yet one would scarcely anticipate the suddenness and regularity with which it occurs.
My studies of habit-formation and reaction-time[143] have revealed the importance of vision in the life of the frog. Perception of movement appears to be of far greater value to the animal than perception of form or color. The spectral colors are discriminated in all probability, for the animals react very differently to those of the blue end than to those of the red. According to Torelle blue is preferred to red. There is evidence that red has a higher stimulating value than blue, and the apparent avoidance of red in Torelle's experiments may be due to this fact. None of the work with which I am familiar demonstrates that the suspected color-reactions are due to stimulation of the eye. They may be due to stimulation of the skin, for Parker[144] has shown that the reactions of Rana pipiens to light are due to stimulation of the sink as well as of the eyes, or they may even be due to intensity instead of color.
The tactual-auditory sense series is better known and also, it would appear, better developed than the chemical series. A large portion of the body surface of the green frog is keenly sensitive to mechanical stimulation, and Steinach[145] by measurement of electrical changes in the nerves of the skin has discovered the existence of "touch spots." His method, which is ingenious, promises to be of considerable value in the objective investigation of the senses, but it involves operations on the subject which inevitably destroy the normal condition of the sense.
According to Steinach we have in the negative variation in the electrical condition of nerves during stimulation a phenomenon which may be used in the determination of the threshold of stimulation as well as in the investigation of irritability. In a previous paper[146] I have discussed the associational rôle of tactual impressions as well as the tactual reaction-time. All my observations lead me to believe that touch is a highly developed and important sense in the green frog.
Of the senses intermediate between touch and hearing that of equilibration has been most discussed. Certainly there is good reason to suppose that the sense-organs of the semicircular canals of the ear furnish the animal with impressions of position, movement, and possibly also of direction. Further study of the tactual-auditory senses of frogs may indicate the existence of conditions similar to those discovered by Parker in certain fishes, in which, as he remarks, "the skin, lateral line organs and ears represent, figuratively speaking, three generations of sense-organs. The oldest is the skin stimulated by varying pressures, such as are produced by irregular currents, and capable of initiating equilibrational responses. From the skin have been derived the lateral line organs stimulated by water vibrations of low rate, and also significant for equilibration. Finally, from the lateral line organs have come the ears stimulated by water vibrations of a high rate and important for equilibration. The ear, unlike the skin and lateral line organs, is differentiated for its two functions, the sacculus for hearing, the utriculus for equilibration."[147]
The sense of hearing remains to be considered. My attention was first drawn to this subject by failure to obtain motor reactions to sounds in the investigation of the time-relations of the neural processes of the green frog. Although a large number of sounds of different qualities, pitches, and intensities were employed, no visible motor reactions were observed. This led me to seek the significance of what appeared to be either a surprising lack of sensitiveness to changes in the environment which would naturally be expected to stimulate the animal, or an interesting and important case of the inhibition of reaction to auditory stimuli. This suggested the question, Are frogs deaf, or do they under certain conditions completely inhibit their usual reactions to sound?
In the literature on the senses and reactions of frogs I have found nothing which contributes importantly to our knowledge of the sense of hearing. Most of the investigations which deal with the ear are concerned with the equilibrational and orientational functions of the labyrinth organs, and have nothing whatever to say about hearing. In the natural histories the existence of a well-developed sense of hearing is usually assumed, and numerous instances of what are supposed to be reactions to sound are cited. It is to be noted, however, that none of the observations in these popular works furnishes satisfactory proof of the exclusion of the influence of visual stimuli. Among the few references to frog audition of which I have knowledge, the only one which seems worthy of special notice is that of Gaupp in his Anatomie des Frosches. Since his few paragraphs sum up the state of our knowledge on the subject, while at the same time furnishing an illustration of the assumption of hearing on the basis of analogy, I present the substance of them in free and slightly abbreviated translation.
"The labyrinth organ has an acoustic and non-acoustic (static) function. For these two functions, according to the leading if not generally accepted view, entirely different portions of the organ are in question, and since the non-acoustic is attributed to the three Cristae acusticae ampullarum and the three Maculae (M. recessus utriculi, M. sacculi, M. lagenae), there remain for the acoustic function only the Papilla basilaris and the Macula neglecta. It is not certain, however, that the non-acoustic organs do not participate in the acoustic function.
"With regard to the acoustic sense of the frog nothing exact is known. That it exists, and that in good development, is certain. The existence of the drum and columella, and the fact that frogs have a voice are unmistakeable proofs of hearing. The participation of the Papilla basilaris in acoustic functions is rendered certain by comparative anatomical studies: the Papilla basilaris is the nerve end-organ from which, in the mammalia, the undoubtedly acoustic organ of Corti arises. From analogy of structure we may also infer an acoustic function in the Macula neglecta: on this, as on the Papilla basilaris, there is a simple tectorial membrane, and further the Pars neglecta, like the Pars basilaris, has a strong thick wall which only in a limited region, namely, where it approaches a part of the perilymphatic space, is markedly thinner." (For fish Breuer (1891) has already stated that if they really hear—which is proved—the Macula neglecta alone can come into consideration in connection with the function, for there is no Papilla basilaris in fishes, and the six other nerve end-organs apparently serve the non-acoustic function.)[148]