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Role of the Elephant Pinna in Hearing

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Role of the Elephant Pinna in Hearing 628 ANIMAL BEHAVIOUR, 30, 2 198A (Lighthill 1952). Because of the effects of the half-muting Lighthill, M. J. 1952. Oq sound generated aerodynami- operation on the quality of courtship song, we surmise cally. I. General theory. Proc. Roy. Soc. A., 211, that G. chopardi makes use of both of these variables to -564-587. - . - - . produce both hisses and whistles with the same pair of Masters, W. M. 1980. Insect disturbance stridulation: spiracles. The courtship whistles of half-muted males are characterization of airborne and vibrational com- consistently higher in frequency (by p to 1 kHz) than ponents of the sound. J. comp. Physiol., 135, the same males' pre-operative whisJes; they are also 259-263. interrupted frequently by brief lapses into broad-band Nelson, M. C. 1979. Sound production in the cockroach, noise. If males have a fixed motor programme for the Gromphadorhina portentosa : the sound-producing abdominal compression that forces air through the apparatus. J. comp. Physiol., 132, 27-38. tracheal horn, the rate of air flow should be higher than Nelson, M. C. & Fraser, J. 1980. Sound production in the normal through the single spiracle that is capable of cockroach, Gromphadorhina portentosa: evidence opening, causing a shift toward a higher frequency in the for communication by hissing. Behav. Ecol. resulting sound. If whistles are produced with the valve Sociobiol., 6, 305-314. partly closed and hisses are produced with the valve fully Pipher, R. D. & Morris, G. K. 1974. Frequency modu- open, the higher rate of flow in half muted males might lation in Conocephalus nigropleurum, the Black- well lead to an occasional 'blow-out' of the valve, ~idedmeadow katydid (Orthoptera: Tettigoniidae). resulting in abrupt lapses from whistle into hiss (as in Can. Entom., 106,997-1001. Fig. 1, Plate I: 'R.Sp. Blocked' row 2). Young, D. 1972. Neuromuscular mechanisms of sound Why does G. chopardi u$e both whistles and hisses? We production in Australian cicadas. J. comp. suggest two explanations. First, the disturbance hiss, like Physiol., 79, 343-362. many defensive sounds, has a broad frequency spectrum which probably is advantageous to the emitter, since a (Received 19 August 1981 ; revised 17 December 1981 ; broad range of predators can detect the sound (Masters MS. number: AS-153) 1980). Second, preliminary studies show sound produc- tion is used in very similar ways in the courtship of portentosa and chopardi. The great difference in the Role of the Elephant Pinna in Sound Localization frequency characteristics of their courtship sounds may Because of its unique size and mobility, the elephant thus represent a species isolation mechanism (both pinna has been the subject of studies aimed at determining species have been collected from the same regions of its functional significance (for a review, see Buss & Estes Madagascar although direct evidence for sympatry is 1971). The results of these studies have suggested two lacking; Chopard 1950). This unusual divergence in the possible roles for the elephant pinna. First, large differ- sound signalling systems of two such closely related ences in arterio-venous .temperature in the pinna, coup- svecies undoubtedlv owes its emergence to the nature of led with the observation that elephants increase the rate ~rom~hadorhina'smethod of souna production, in which at which they flap their pinnae as ambient temperature the potential for frequency modulation is inherent in the increases, suggest that the pinna plays a role in regula- design. ting body temperature. Second, the observation that We wish to thank the students of the MBL (Woods elephants extend their pinnae rigidly from their heads Hole) Neural Systems & Behavior Course, class of 1979 in alarm or threat situations has been interpreted to (C. Derby and T. Flanagan especially), without whom mean that they use the pinnae to alert nearby elephants this project might never have begun. R. Hoy, R. Murphey, or to warn intruders. Thus, the elephant pinna has been and J. Paton made many helpful comments on earlier implicated in both thermoregulation and communication. drafts of this MS. This research was supported by Recently, we have had the opportunity to test the NINCDS Grant NS16323 to MCN. hearing capacities of a 7-year-old wild-born female JEANFRASER* Indian elephant (Elephas maximus) at the Ralph MARGARETC. NELSON? Mitchell Zoo in Independence, Kansas. The tests given the animal included the determination of absolute, *Department of Biology, sound-localization, and frequency-discrimination thres- Boston University, holds. During these tests it was noted that the Boston, MA 02215. elephant extended its pinnae when attempting to localize brief sounds, but did not extend them during the non- ?Section of Neurobiology & Behavior, localization tests. The purpose of this paper, then, is to Cornell University, describe these observations and to suggest that the ele- Zthaca, NY 14853. phant's pinna may play an important role in the local- ization of sound. - References The details of the behavioural procedure used to test I Chopard, L. 1950. Sur I'anatomie et le dbveloppement the elephant have been described elsewhere (Heffner & d'une blatte vivipare. Znt. Congr. Ent., 8 Heffner 1980. 1981). Brieflv. the elevhant was brought (Stockholm 1948), 218:222. into a room (6.7 x 5.6 x 4:6 m) and tethered in front Dumortier, B. 1963. The phys~calcharacteristics of sound of a response panel which contained three buttons emissions in arthropods. In: Acoustic Behaviour of mounted in a horizontal row. The animal was trained to Animals (Ed. by R. -G. Busnel), pp. 346-373. make an 'observing' response by pressing the centre London : Elsevier. button with its trunk, a response which initiated a trial Elsner, N. 1974. Neuroethology of sound production in and was followed by the presentation of an auditory cue, Gomphocerine grasshoppers - (Orthoptera : In the sound-localization tests, the animal was then re- ' ' Acrididad I. Sona vatterns and stridulatow quired to press the left or right button if a sound cmca movements. J. corn;. ~h~siol.,88, 67-102. from a loudspeaker located to the left or right of bhe Elsner, N. & Popov, A. 1978. Neuroethology of acoustic animal, respectively. A comt response was rew&dacll communication. Adv. Insect Physiol., 13, 229-355. by dispensing 30 ml of a fruit-flavoured sugar s01t.doim SHORT COMA into a small trough located below the centre button. In signs of reduced motivation by occasionally ceasing to the absolute-threshold tests, the animal was required to [perform and playfully pushing on the fenq behind press the centre button and then press the left button if which the loudspeakers were mounted. Though it wn- a sound was presented from a loudspeaker and to press tinued to extend its pinnae on most of the trials, it oc- the right button if no sound was presented. Frequency- casionally failed to extend them or extended only one of discrimination texts were conducted in a similar man- them. Thus, while the poorer performance observed in ner in which, after pressing the centre button, the animal this situation may have been due to reduced motivation, was required to press the left button if a train of tone it is of interest to note that failure to extend the pinnae pulses emitted from a loudspeaker were all of the same was accompanied by poorer performance in localizing 1 frequency and to press the right button if the tone pulses sound. alternated between two different frequencies. Test In summary, the elephant was observed to extend its 1 sessions were conducted during the quiet early morning pinnae during sound-localization tests, but not during hours shortly before the animal's morning feeding time. tests of absolute sensitivity or frequency discrimination. Following initial training, the elephant's absolute Pinna extension was observed to occur for a variety of r thresholds for pure tones from 16 Hz to 12 kHz were different sounds and at both large and small angles. ,, determined. During these tests it was noted that the ele- However, when the animal was insufficiently motivated . phant's pinnae were maintained in the relaxed position to work at a steady pace, it did not always extend its back against its head and neck, with only occasional pinnae and its error rate increased. These observations flapping movements such as to remove flies. However, suggest that elephants extend their pinnae incorder to when the sound-localization tests were given, a very dif- localize sound accurately. I ferent pattern of pinnae movements emerged. During Though it is widely believed that movabb pinnae, A sound-localization testing, the animal extended its such as the elephant's, play an important role in sound pinnae nearly perpendicular to its head just before it localization, it is not clear just how pinna extension en- pressed the centre button. After the centre button was hances the ability to localize sound. While the function pressed and a brief sound had been presented, the ani- of the human pinna has been studied in detail, little is mal then moved its trunk to press either the left or right known concerning the contribution of large movable button, during which time it usually returned its ears to pinnae to sound localization (for reviews, see Butler the normal relaxed position against its head. Following 1975; Shaw 1974). One possibility is that extension of the localization tests, the elephant was tested on fre- the pinnae may enhance the difference between the fre- quency discrimination and was re-tested on its absolute quency-intensity spectra of the sounds reaching the sensitivity to 4 kHz and 8 kHz. In both of these sub- two ears, thereby providing a more distinct localization sequent tests, the animal maintained its pinnae in the cue. Another is that it may reduce the intensity of echoes relaxed position.
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