The tympanal organ of insects consists of a group of scolophores associated with a thin, horny (chitinous) membrane at the surface of the body, one on each side. Usually the scolophores are attached at one end by a spinous process to the tympanic membrane (eardrum); the other ends rest on an immobile part of the body structure. When the membrane moves back and forth in response to the alternating pressures of sound waves, the nerve fibre from the ganglion cell of the scolophore transmits impulses to the central nervous system. Because the tympanic membrane is activated by the pressure of sound waves, this is a pressure type of ear.

Simple tympanal organs, such as those found in moths, contain only two or four elements, or scolophores. In cicadas, on the other hand, these organs are highly developed; they include a sensory body (a number of scolophores in a capsule) that may contain as many as 1,500 elements.

With 80 to 100 scolophores, the grasshopper ear, which has been studied more thoroughly than any other insect ear, is structurally between that of moths and cicadas. Ordinarily, the tympanic membrane is hidden beneath the base of the insect’s wing cover. A bundle of auditory nerve fibres runs from one side of the sensory body, which lies on the inner surface of the membrane, and joins other nerve fibres of the region to form a large nerve extending to a ganglion (nerve centre) in the thorax.

Behavioral observations

That the insect ear serves an auditory purpose has been proved by a large number of experimental observations, particularly those that have dealt most extensively with katydids and crickets. Males of these groups produce sounds by stridulation, which usually involves rubbing the covers of the wings together in a particular way. One wing has a serrated surface (a “file”) that runs along an enlarged vein; the other wing has a sharp edge over which the file is scraped. The scraping causes the wing surfaces to vibrate; the natural resonances of the vibrations and the particular rhythm and repetition rate of the scraping movements determine the nature of the song, which varies with each species. Most females are silent, but those of a few species have a poorly developed stridulatory apparatus, and weak sounds have been reported. Both males and females have tympanal organs for sound reception.

The observation that the males of many insect species produce repeated stridulatory sounds during the mating season led to the inference that the primary purpose of these noises was to attract a female. That this is indeed the case was first established by the extensive observations of the Yugoslavian entomologist Ivan Regen, who worked over the period 1902–30 mostly with a few species of katydids and crickets. In one of his earliest experiments, Regen proved (1913–14) that a male katydid of the species Thamnotrizon apterus responds to the sound of another male by chirping. The first male responds in turn to the second male’s chirp, and the two insects then set up an alternating pattern of chirping. Although this pattern had been observed earlier, Regen was the first to prove by a series of experiments that it depends upon the sense of hearing. After removing the forelegs, on which the tympanal organs are located, of certain males, he found that even though these insects continued to stridulate, they did so only in individual rhythms that were not affected by the sounds of other males. Any alternation of chirping between deafened males, or between a deafened and a normal male, occurred only rarely, for brief times, and by chance.

A long series of check experiments by Regen showed that other stimuli, such as light, odours, and surface vibrations, did not affect the chirping behaviour. In these experiments the insects were placed in separate rooms, and their sounds were transmitted by telephone.

Further experiments carried out by Regen on field crickets (Liogryllus campestris) demonstrated the reactions of females to chirping males. In the most elaborate of these experiments, 1,600 sexually receptive females were released around the periphery of a large enclosed area in the middle of which had been placed a cage containing one or more chirping males. Precise data concerning the frequency with which the females moved toward the cage were obtained by surrounding the cage site with an array of traps in which the females were caught as they moved inward. The results were statistically significant. Normal females (those with intact tympanal organs) moved toward the cage and eventually reached it. The removal of one foreleg and its tympanal organ, however, caused difficulty; the movements were more random and the approaches fewer, although some females did succeed in reaching the cage. When both tympanal organs were removed or if the male failed to chirp, the performance of the females was reduced to chance. They also failed to exhibit the seeking performances if the male’s stridulatory organ was modified, as by removing the file, so that little or no sound was produced.

In 1926 Regen returned to his study of the alternating chirping pattern of katydids and succeeded in having males react to an artificial sound, one that Regen himself produced. He also found that the alternation could be demonstrated with a suitably active male by using a variety of sounds—whistles, percussion noises, and sounds made with his mouth. It was never altogether clear, however, what changes Regen had made in his signals that finally brought success; probably the secret lay in the particular rhythm and timing of the signals. At any rate, this method made possible a study of the general nature of the auditory sensitivity of these insects and the range of sound frequencies to which they responded. It was shown that katydids are most sensitive to the very high frequencies, those that are beyond the limit of the human ear. The instruments available to Regen at the time, however, did not permit a precise measurement of intensity thresholds. (A threshold is the lowest point at which a particular stimulus will cause a response in an organism.)

Although the work of Regen and others established the basic character of sound reception in insects and its role in communication and mating, other details had to await the introduction of electrophysiological methods in this field as well as the development of electronic methods for the precise production, control, and measurement of sound stimuli.