SOUND RECEPTION

response of an organism's aural mechanism, the ear, to a specific form of energy change, or sound waves. Sound waves can be transmitted through gases, liquids, or solids, but the hearing function of each species is particularly (though not exclusively) sensitive to stimuli from one medium. If an animal possessing an auditory mechanism comes in suitable contact with a medium vibrating at a frequency and intensity within its range of aural (hearing) sensitivity, it may hear the sound. For land animals, the usual vibrating medium is the air; for fishes and other aquatic creatures, it commonly is the water. Yet, under suitable conditions, all hearing animals can perceive sound waves transmitted by media other than the one in which they live; thus, humans can hear noise while underwater. (Additional information is contained in the article sound.) In the course of evolution, animals have developed a variety of sense organs that respond to mechanical stimuli. There are at least 10 of these mechanoreceptors in vertebrates and perhaps as many in advanced invertebrates. Not all of these structures respond to sound, however, for among them are the simple touch endings of the skin and the motion receptors that serve (mediate) bodily equilibrium. Although the different ways of registering mechanical changes in the environment or within the body represent various structural specializations, it is not feasible to identify any one of them simply in terms of its structure; many different mechanisms, cells, or organs may perform similar functions. Ears, for example, take many forms in the lower animals and often have little resemblance to these organs in man and other higher vertebrates. Yet the service that they perform in sound reception is similar enough that they may be called ears. Although there is no fossil record of the origin and development of auditory structures, in animals with ears the evolutionary process in every instance appears to have been a conversion to an auditory function of structures that previously mediated a simpler form of mechanoreception. Indeed, any mechanoreceptor, even though best adapted to respond to some other form of mechanical stimulation, will respond to vibrations within some region of the sound frequency range if the vibrations have a sufficiently high level of intensity. Many attempts have been made to define hearing, often with indifferent success. The task is difficult, and in certain respects the lines of distinction are arbitrary. The ear cannot be identified by any standard structure, nor can it be identified in terms of the stimulus as simply a receiver of sound vibrations. As noted above, mechanical receptor organs will respond to sound vibrations within some region of the frequency range if a sufficiently high level of intensity is provided. Moreover, the ear cannot be characterized in terms of the physical principles by which it operates because these principles vary among the ears of different animal species. A definition of hearing, therefore, must be sought in terms of the ear's specialization of function and the relative effectiveness with which it performs this function. Thus, hearing may be characterized as the reception of sound vibrations by an organ, the ear, that has developed for this particular purpose and that has reception of sound as its primary function. This definition excludes the reception of sound vibrations by touch (tactual) endings in the skin, for example, because these structures respond most readily to direct pressure. Before such receptors will respond to sound waves, the vibrational intensity of the sound must be relatively great. Also excluded are the hair sensilla, of which arthropods have many types, whenever it can be shown that these organs respond with greater sensitivity to another stimulus (most often a simple direct deflection of the central hair). Theoretically, several aspects of vibration might serve in its detection by an ear. These characteristics include the amplitude (extent) of the motion of particles (e.g., molecules) in a medium, the velocity and acceleration of the motion, the pressure exerted upon an obstacle in the path of the sound waves, and temperature changes occasioned by the vibrations. All of these manifestations have been utilized in attempts to design microphones for the detection and measurement of sound, but only two (pressure and velocity effects) have proved to be of any practical value. Thus, those devices that employ these two effects are known as pressure and velocity microphones. It seems more than coincidence that these same two aspects of sound, pressure and velocity, are the only stimulus characteristics on which the evolution of ears appears to have been based. Moreover, just as the pressure microphone is the most practical type designed by man, among ears the pressure type is the most widespread and the most highly developed. Ears that distinguish changes in velocity have appeared only in a few lower animalsas an elaborated hair organ in some insects and perhaps spiders and in two special forms among fishes. All other ears are pressure receptors that have taken two lines of evolutionary development, one in most of the insects and another in vertebrates above fishes. Considering the usefulness of the sense of hearing to such highly organized animals as man, it may seem surprising that this sense is so limited in its appearance and development among animals. It is found only in two major groups of animals: arthropods (e.g., insects and crabs) and vertebrates (e.g., amphibians, birds, and mammals). The condition that probably limited the development of hearing in other species was the lack of sufficient advancement and flexibility of the nervous system. In those animals with auditory structures, hearing serves purposes of great biological value: in its more primitive forms, it is used to sense danger and enemies, to detect prey, and to identify prospective mates; at a more complex level, hearing is involved in communication within social groups and in emotional expressions of various kinds. The cry of an infant mouse that has strayed from the nest elicits a response by the mother to retrieve it. The singing of a male thrush asserts a claim to its territory, attracts a female to the area, and warns off other males. Among higher mammals ( e.g., monkeys and apes) vocalizations show even greater variety and express a range of meanings that may be interpreted in human terms as expressions of such concepts as danger, aggression, love, and the availability of food. In man, the elaborations of auditory communication can be even more symbolically complex, extending to speech and music (see speech). The significant features in complicated sounds that people perceive and differentiate correspond to the physical dimensions of frequency (the number of waves, cycles, or vibrations per second), intensity, phase, complexity of wave form, and temporal pattern. The variety of distinguishable acoustic forms is enormous. Among the most highly refined applications of the auditory sense are those found in such animals as bats and dolphins. These creatures are able to discern objects around them by a process called echolocation; the animal sends out a cry and, by the nature of the echo, is informed of the presence of obstacles or potential prey. For these animals, the sense of hearing provides a service in the dark that closely approaches the reliability of vision in the perception of objects and spatial relations. the response of an aural mechanism, the ear, to a specific form of energy change, or sound waves. Sound waves can be transmitted through gases, liquids, or solids, but the hearing function of each species is particularly (though not exclusively) sensitive to stimuli from one medium. In spiders, certain insects, and some fishes, sound reception is based on detection of a vibration's velocity, while in most animals, hearing is caused by pressure on an obstacle in a vibration's path. Members of two animal groups are capable of sound reception: arthropods (e.g., insects and crabs) and vertebrates (e.g., reptiles, birds, and mammals). In more primitive species, hearing can enable an animal to sense danger, locate food, and find mates, and in more complex creatures it is important for communication and emotional expression. Additional reading There are no general texts on sound reception. Included below are some of the specialized references dealing with this subject in a technical manner.J. Schwartzkopff, Mechanoreception, in Morris Rockstein (ed.), The Physiology of Insecta, vol. 1, pp. 509561 (1964); M.L. Wolbarsht, Electrical Characteristics of Insect Mechanoreceptors, J. Gen. Physiol., 44:105122 (1960); E.G. Gray, The Fine Structure of the Insect Ear, Phil. Trans. R. Soc., Series B, 243:7594 (1960); C. Walcott and W.G. van der Kloot, The Physiology of the Spider Vibration Receptor, J. Exp. Zool., 141:191244 (1959); E.G. Wever and J.A. Vernon, The Auditory Sensitivity of Orthoptera, Proc. Natn. Acad. Sci. U.S.A., 45:413419 (1959); H. Frings and M. Frings, Uses of Sounds by Insects, A. Rev. Ent., 3:87106 (1958); R.J. Pumphrey, Hearing in Insects, Biol. Rev., 15:107132 (1940); Sensory Organs: Hearing, in A.J. Marshall (ed.), Biology and Comparative Physiology of Birds, vol. 2. (1961). See also Brian Lewis, Bioacoustics: A Comparative Approach (1983); William C. Stebbins, The Acoustic Sense of Animals (1983); James F. Willott (ed.), The Auditory Psychobiology of the Mouse (1983). Ernest Glen Wever