Biomedical subjects

R S Heffner

Publications and source records attributed to R S Heffner.

At least 19 recordsLinked to original sources

PubMed1992

Visual factors in sound localization in mammals.

The ability of mammals to localize sound varies widely among species. During the past decade, evidence has accumulated that this variation cannot be accounted for simply on the basis of the availability of the physical cues for locus. Evidence is presented that a major factor in sound localization is the need to direct the field of best vision to a sound source for further scrutiny. Thus, species with broad fields of best vision (such as visual streaks) require less accurate information regarding the location of a sound source than do species with very narrow fields of best vision (such as foveae). To support this suggestion, data are reported for the width of the field of best vision in the form of retinal ganglion cell isodensity contours for thirteen species of mammals. The possible contribution of other factors including binocular fields, visual acuity, and the degree to which a species is predatory in lifestyle, is also examined.

Animals↗

PubMed1992

Hearing and sound localization in blind mole rats (Spalax ehrenbergi).

Two blind mole rats were tested for their ability to detect and localize sound. The results indicate that blind mole rats have severely limited, and probably degenerate, auditory abilities. Although their 60-dB low-frequency hearing limit of 54 Hz is within the range for other rodents, the highest frequency they can hear at a level of 60 dB SPL is only 5.9 kHz, giving them the poorest high-frequency sensitivity yet observed in any mammal. In addition they have poor sensitivity as indicated by the fact that their lowest threshold is only 32 dB SPL (at 1 kHz). Finally, they are unable to localize brief sounds but retain a rudimentary ability to localize sounds of 0.5 s or longer. These results, combined with those of previous studies of subterranean species (i.e., blind mole rats, naked mole rats, and pocket gophers), suggest that poor auditory sensitivity, the loss of high-frequency hearing, and an inability to localize brief sounds is a degenerate state which may be characteristic of subterranean mammals. Thus it appears that an exclusive adaptation to a subterranean lifestyle (where airborne sound propagates poorly and where directional responses are limited by the tunnels) can result in vestigial auditory abilities just as the absence of light results in vestigial vision.

Acoustic Stimulation↗

PubMed1992

Hearing in large mammals: sound-localization acuity in cattle (Bos taurus) and goats (Capra hircus).

Sound localization acuity of 3 cattle (Bos taurus) and 2 goats (Capra hircus) was determined for brief complex sounds in a two-choice procedure. Thresholds around the median sagittal plane averaged 30 degrees and 18 degrees, respectively. For comparison, thresholds were obtained in the same test apparatus for humans (0.8 degrees) and a dog (8 degrees). Although the relatively poor acuity of cattle and goats compared with most mammals comes as some surprise, given their large interaural distances and the large binaural locus cues available to them, it is not unexpected when other factors are considered. Like other poor localizers (both domesticated and nondomesticated), cattle and goats are prey species with their best vision directed throughout nearly the entire horizon. In contrast to mammals with very narrow foveal fields, they may not need very accurate locus information from their auditory systems to direct their gaze to a sound source.

Adult↗

PubMed1991

Objective auditory threshold estimation using sine-wave derived responses.

A derived response method of acquiring frequency specific auditory evoked potentials that utilizes a pure tone in combination with a toneburst is applied to the measurement of hearing sensitivity in guinea pigs, chinchillas and pocket gophers. Two experiments which demonstrate that thresholds acquired via tone-derived responses are 10 to 15 dB more sensitive than thresholds to solitary tonebursts are described. The derived potentials approximate behaviorally acquired thresholds at frequencies of 0.5 kHz and above. This technique may provide a more rapid means of assessing hearing sensitivity in laboratory animals than by behavioral means.

Acoustic Stimulation↗

PubMed1991

Behavioral hearing range of the chinchilla.

The audiograms of three chinchillas were determined using pure tones ranging from 32 Hz to 45 kHz. The animals were tested with a conditioned avoidance procedure in which their heads were fixed within the sound field by requiring them to place their mouths on a water spout. At a level of 60 dB SPL the average hearing range extended from 50 Hz to 33 kHz with none of the animals able to hear 45 kHz at 89 dB. Overall, the audiogram of the chinchilla appears to resemble the human audiogram more closely than do other rodent audiograms. An analysis of ten published chinchilla audiograms indicates that those procedures which do not fix an animal within the sound field may overestimate their sensitivity.

Animals↗

PubMed1990

Vestigial hearing in a fossorial mammal, the pocket gopher (Geomys bursarius).

Behavioral tests of hearing and sound localization in the North American pocket gopher (Geomys bursarius) show that it is unique among mammals. It has a severely attenuated range of hearing and only rudimentary ability to localize sound. In these respects, the hearing of gophers can be properly termed 'vestigial' and suggests that life underground can produce as severe a change in hearing as a light-less world produces in vision or an odorless world produces in olfaction.

Acoustic Stimulation↗

PubMed1990

Hearing in domestic pigs (Sus scrofa) and goats (Capra hircus).

Behavioral audiograms were determined for three pigs and two goats. The hearing of the pigs ranged from 42 Hz to 40.5 kHz with a region of best sensitivity from 250 Hz to 16 kHz. Hearing in goats ranged from 78 Hz to 37 kHz with a well-defined point of best sensitivity at 2 kHz. Because these animals are unable to localize high-frequency tones, it seems unlikely that selective pressure to use the interaural spectral-difference cue for sound localization is behind their high-frequency hearing. Instead, we suggest that these and other hoofed mammals evolved high-frequency hearing in order to use monaural locus cues which prevent front/back locus reversals.

Acoustic Stimulation↗

PubMed1990

Effect of bilateral auditory cortex lesions on absolute thresholds in Japanese macaques.

1. The behavioral audiograms of four Japanese macaques (Macaca fuscata) were assessed before and after receiving two-stage bilateral lesions of auditory cortex. Thresholds were assessed for each ear with the use of insertion earphones. 2. The bilateral lesions resulted in a large initial hearing loss followed by partial recovery that left the animals with a permanent hearing loss in both ears. 3. The initial hearing loss consisted of a total insensitivity to sound in the ear contralateral to the second lesion with limited hearing in the other ear. However, the animal with the most complete lesion was initially unable to hear sound in either ear. Broadband noise was often more effective in eliciting a behavioral response than tones. 4. Partial recovery occurred in all animals and was observed as early as the first week after surgery. Most of this recovery occurred during the first 3-7 wk after surgery. This rapid phase of recovery was sometimes followed by a more gradual phase although thresholds were still elevated after 94 wk. 5. The permanent hearing loss, which averaged from 30 to 44 dB, was not constant across frequency. Threshold shifts were smallest at 63 Hz and progressively increased with frequency to a maximum loss from 8 to 25 kHz with slightly less loss at 32 kHz. 6. Analysis of the psychophysical functions and threshold stability gave no indication of any nonsensory deficits in attention or vigilance. 7. These results, taken with those of previous experiments, indicate that each hemisphere is primarily involved in the detection of sound in the contralateral ear and secondarily involved in detection in the ipsilateral ear. This arrangement differs from that seen in sound localization where each hemisphere is involved with the contralateral hemifield as opposed to the contralateral ear. Thus it appears that the functional organization of auditory cortex for sound localization is different from that for the detection and identification of sound itself.

Animals↗

PubMed1990

Effect of bilateral auditory cortex lesions on sound localization in Japanese macaques.

1. The ability of four Japanese macaques (Macaca fuscata) to localize sound was determined after bilateral ablation of auditory cortex. The animals were given two tests: a "midline" test in which they had to discriminate noise bursts presented from a loudspeaker located to the left from identical noise bursts presented from a loudspeaker located to the right of midline, and a "hemifield" test in which both loudspeakers were located in their right hemifield. 2. Both of the tests were administered by the use of two different behavioral tasks: a conditioned-avoidance task in which the animals were trained to make or break contact with a water spout to indicate the location of a sound source, and a two-choice task that required the animals to walk to the source of the sound. 3. The results of both the conditioned-avoidance and the two-choice tasks demonstrated that the animals were able to perform the midline discrimination although their localization acuity was reduced. However, the animals had great difficulty in learning to walk to the source of a sound in spite of the fact that they had received previous sound-localization training in the conditioned-avoidance task. This difficulty suggested that the monkeys no longer associated the sound with a location in space. 4. The results of both the conditioned-avoidance and the two-choice tasks demonstrated that the animals were unable to discriminate the locus of a sound source when both loudspeakers were located in the same hemifield. 5. Bilateral ablation of auditory cortex results in both sensory and perceptual deficits. The presence of sensory deficits is indicated by the decreased acuity in the left-right discrimination and the inability to discriminate between two loudspeakers located in the same hemifield. The deficit in the perception of the locus of sound is indicated by the difficulty in learning to approach the source of a sound, an ability which normal monkeys exhibit without training. 6. There appear to be species' differences in the effect of auditory cortex lesions on sound localization. Although cortical lesions result in a sound-localization deficit in several species of primates and carnivores, they have little or no effect on rats.

Animals↗

PubMed1989

Cortical deafness cannot account for the inability of Japanese macaques to discriminate species-specific vocalizations.

Bilateral ablation of the superior temporal gyrus in Japanese macaques results in a significant hearing loss (cortical deafness) as well as in an inability to discriminate between two types of their "coo" vocalizations. A two-part investigation was conducted to determine whether the hearing loss may itself affect the ability to discriminate vocalizations. First, four normal Japanese macaques were tested for their ability to discriminate coos which were filtered to simulate the effect of a cortical hearing loss. Second, four Japanese macaques with bilateral superior temporal gyrus lesions were tested for their ability to discriminate coos which were amplified and equalized to compensate for each animal's hearing loss. All four normal macaques were able to discriminate the filtered coos easily whereas compensating for the operated monkeys' hearing losses did not improve their performances. It appears that the inability of monkeys with bilateral superior temporal gyrus lesions to discriminate conspecific vocalizations is not simply due to the accompanying hearing loss, but is a separate auditory disorder.

Animals↗

PubMed1989

Effect of restricted cortical lesions on absolute thresholds and aphasia-like deficits in Japanese macaques.

The effect of small bilateral cortical lesions on pure-tone audiograms and on the ability to discriminate between two types of Japanese macaque coo vocalizations was determined in four Japanese macaques (Macaca fuscata). A lesion that included the middle portion of the superior temporal gyrus of both hemispheres, that is, the primary and secondary auditory areas, resulted in a partial hearing loss as well as an inability to discriminate the vocalizations. Lesions that included the ventral portions of the superior temporal gyrus of both hemispheres but spared auditory cortex on one side also resulted in a partial hearing loss but had either a small effect or no effect on the ability to discriminate the vocalizations. Bilateral ablation of the dorsal superior temporal gyrus and adjacent parietal and occipital areas did not appear to result in a hearing loss and had no effect on the ability to discriminate the vocalizations. These results suggest that a hearing loss may be produced by lesions that involve small portions of the ventral two-thirds of the superior temporal gyrus bilaterally although the resulting loss is not as great as that observed with larger lesions. However, the aphasia-like deficit appears to result from a lesion of primary and/or secondary auditory cortex.

Acoustic Stimulation↗

PubMed1989

Unilateral auditory cortex ablation in macaques results in a contralateral hearing loss.

1. The behavioral audiograms of four Japanese macaques (Macaca fuscata) were assessed before and after unilateral ablation of auditory cortex. The tones were presented via insertion earphones so that each ear could be tested separately. 2. Each animal had a hearing loss in the ear contralateral to the lesion, whereas the ipsilateral ear showed no change in sensitivity. The hearing loss initially appeared as a large shift in thresholds followed by rapid but incomplete recovery during the first 3-5 wk after surgery. The initial hearing loss ranged as high as 68 dB at some frequencies, although thresholds at other frequencies were occasionally unchanged. A threshold shift could be demonstrated with broadband noise as well as with tones. Although thresholds for some tones returned to normal within a few weeks, most were still elevated 16 wk after surgery when testing was discontinued. The largest long-term hearing losses occurred at frequencies from 4 to 25 kHz. 3. Analysis of the animals' psychophysical functions suggested that the hearing loss resembled a sensory deficit, as opposed to a nonsensory deficit in attention or vigilance. 4. Testing with binaural stimuli indicated that the hearing loss could best be described as a contralateral "ear" deficit, as opposed to a contralateral "auditory field" deficit. 5. It is suggested that a similar hearing loss occurs in humans after unilateral damage to auditory cortex.

Animals↗

PubMed1989

Sound localization, use of binaural cues and the superior olivary complex in pigs.

Noise localization thresholds and the ability to localize pure tones at 60 degrees separation were determined for three domestic pigs using a conditioned avoidance procedure. The average threshold for localizing a brief noise burst was 4.5 degrees which is much more accurate than the thresholds of other hoofed mammals, such as horses, cattle and goats. The ability of pigs to localize low-frequency tones indicates that they can use the binaural phase-difference cue. However, they were unable to localize tones of 4 kHz and higher, indicating that, like other hoofed mammals, their ability to use binaural intensity cues is greatly restricted if not completely absent. An examination of the superior olivary complex of pigs revealed that in relative size, shape and cell density it is more like that of cats than that of other hoofed mammals.

Animals↗

PubMed1988

Sound localization acuity in the cat: effect of azimuth, signal duration, and test procedure.

The sound localization acuity of cats was determined at 0 degrees, 30 degrees, 60 degrees, and 90 degrees from the median sagittal plane for four durations of noise bursts using two behavioral procedures. Similar thresholds were also obtained for humans. The cats' average thresholds for a 40 ms noise burst ranged from 4.8 degrees at 0 degrees azimuth to 9.0 degrees at 90 degrees azimuth. Comparable thresholds for humans ranged from 1.3 degrees at 0 degrees to 9.7 degrees at 90 degrees. The fact that humans and cats had similar thresholds at 90 degrees azimuth suggests that the mobile pinnae of the cat are not an advantage in localizing sound in the lateral fields. Varying the duration of the stimulus from 10 ms to 'continuous' had little effect on threshold. No difference was found between the results of the two behavioral procedures.

Adolescent↗

PubMed1988

Sound localization and use of binaural cues by the gerbil (Meriones unguiculatus).

Noise-localization thresholds and the ability to localize pure tones at 60 degrees separation were determined for gerbils. The gerbils were trained using a two-choice procedure with observing response in which the gerbils made a left or right response to sounds emanating from their left or right side in order to obtain food. The average 75% correct localization threshold of 7 gerbils for a 100-ms noise burst was 27 degrees with chance performance (p greater than .01) reached at 12 degrees. The ability of 4 gerbils to localize both low- and high-frequency pure tones indicates that gerbils are able to use both phase- and intensity-difference locus cues. The frequency at which tone localization was poorest was 2.8 kHz, well below the theoretical frequency of ambiguity of the phase cue but within the frequency range at which phase locking declines in the mammalian auditory system. The sound localization ability of gerbils is typical of small rodents, and there is no obvious sign that it is affected by the degenerative disorder of the central auditory system which has been recently discovered in gerbils.

Animals↗

PubMed1987

Localization of noise, use of binaural cues, and a description of the superior olivary complex in the smallest carnivore, the least weasel (Mustela nivalis).

Cats and dogs have relatively good sound-localization acuity, and the question arises as to whether this trait is a characteristic of all carnivores or whether it is due to the fact that they have large heads and correspondingly large binaural localization cues available to them. The localization acuity of the least weasel, the smallest extant carnivore, was found to be less accurate than larger carnivores but more accurate than other small mammals. This suggests that carnivores may be under strong selective pressure to localize accurately but that interaural distance may be a limiting factor. The least weasel is capable of using both binaural phase differences and intensity differences to localize, but has a relatively broad mid-frequency range for which neither cue is optimal. Finally, the superior olivary complex of the least weasel is well developed and resembles that of larger carnivores more than that of small rodents.

Animals↗

PubMed1986

Localization of tones by horses: use of binaural cues and the role of the superior olivary complex.

The ability of horses to use binaural time and intensity difference cues to localize sound was assessed in free-field localization tests by using pure tones. The animals were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30 degrees to the left and right of the animals' midline (60 degrees total separation). Three animals were tested with a two-choice procedure; 2 additional animals were tested with a conditioned avoidance procedure. All 5 animals were able to localize 250 Hz, 500 Hz, and 1 kHz but were completely unable to localize 2 kHz and above. Because the frequency of ambiguity for the binaural phase cue delta phi for horses in this test was calculated to be 1.5 kHz, these results indicate that horses can use binaural time differences in the form of delta phi but are unable to use binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional horses, which showed that horses correctly orient to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural delta I.

Animals↗

PubMed1986

Hearing loss in Japanese macaques following bilateral auditory cortex lesions.

The hearing ability of five Japanese macaques (Macaca fuscata) was assessed following two-stage bilateral auditory cortex lesions. The animals were tested using a shock-avoidance procedure with a conditioned-suppression procedure used for comparison in two cases. The animals initially were unable to respond to sound, and the first signs of hearing appeared as late as 13 wk after surgery. Hearing levels improved gradually over time, with maximal recovery reached at 24-35 wk after surgery. Recovery was most pronounced for low frequencies (63-250 Hz) and very high frequencies (32 kHz), which generally returned to normal or near-normal levels. However, the monkeys appeared to have suffered a permanent hearing loss throughout most of their hearing range, especially in the midfrequency range, where they are normally most sensitive. A review of the animal literature reveals little support for the previous view that bilateral auditory cortex lesions have little or no effect on absolute sensitivity in primates and carnivores. Most previous studies did not conduct detailed hearing tests, and those that did often noted a hearing loss. The hearing loss found in monkeys is similar to that noted in human cases following bilateral auditory cortex lesions. The current findings thus provide experimental verification of the clinical phenomenon of cortical deafness.

Adaptation, Physiological↗