Biomedical subjects

R S Heffner

Publications and source records attributed to R S Heffner.

At least 19 recordsLinked to original sources

PubMed1999

Sound localization in an Old-World fruit bat (Rousettus aegyptiacus): acuity, use of binaural cues, and relationship to vision.

The passive sound-localization acuity of Egyptian fruit bats (Rousettus aegyptiacus) was determined using a conditioned-avoidance procedure. The mean minimum audible angle for left-right discrimination for 3 bats was 11.6 degrees--very near the mean for terrestrial mammals. The bats also were able to localize low- and high-frequency pure tones, indicating that they can use both binaural phase-difference and binaural intensity-difference cues to localize sound. Moreover, they were able to use the binaural phase-difference cue up to at least 5.6 kHz, which is higher than other mammals yet tested. The width of the Egyptian fruit bats' field of best vision was 27 degrees. This value is consistent with the hypothesis that the role of passive sound localization is to direct the eyes for visual scrutiny of sound sources. Thus, the passive localization abilities of these echolocating megachiropteran fruit bats do not deviate from the patterns established for nonecholocating mammals.

Animals↗

PubMed1998

Passive sound-localization ability of the big brown bat (Eptesicus fuscus).

The passive sound-localization ability (i.e. minimum audible angle) of the big brown bat, Eptesicus fuscus, was determined using a conditioned avoidance procedure in which the animals were trained to discriminate left sounds from right sounds. The mean threshold of three bats for a 100-ms broadband noise burst was 14 degrees, a value that is about average for mammals. A similar threshold of 15 degrees was obtained for one animal when it was retested with one of its own recorded echolocation calls as the stimulus. The two bats tested on pure-tone localization were able to localize high-frequency, but not low-frequency tones, even when a low-frequency tone was amplitude modulated, a result indicating that these bats are not able to use binaural time-difference cues for localization. Finally, given the width of the bat's field of best vision, as determined by a count of its ganglion-cell density, its sound-localization acuity is consistent with the hypothesis that the role of passive sound localization is to direct the eyes to the source of a sound.

Acoustic Stimulation↗

PubMed1998

Hearing in a megachiropteran fruit bat (Rousettus aegyptiacus).

The Egyptian fruit bat (Rousettus aegyptiacus) is one of the few megachiropteran bats capable of echolocation. However, it uses rudimentary tongue clicks rather than laryngeally produced echo calls. We determined the audiogram of 2 bats using a conditioned avoidance procedure with fruit puree reward. At an intensity of 60 dB sound pressure level, the bats' hearing extended from 2.25 kHz to 64 kHz, with a region of good sensitivity between 8 kHz and 45 kHz. A dip in sensitivity at 32 kHz appears to be due to pinna directionality. The hearing of Egyptian fruit bats is typical for a mammal of that size and is not as limited as previously reported. Methodological issues, specifically training an animal to listen for low-intensity signals and imposing a significant cost for failing to report signals (i.e., misses), are discussed as the basis for the discrepancy between our results and earlier reports.

Animals↗

PubMed1997

Audiogram of the big brown bat (Eptesicus fuscus).

The audiograms of three big brown bats (Eptesicus fuscus) were determined using a conditioned avoidance procedure. The average audiogram ranged from 0.850 kHz at 106 dB to 120 kHz at 83 dB SPL, with a best threshold of 7 dB at 20 kHz and a distinct decrease in sensitivity at 45 kHz. The results confirm those of a previous study by Dalland (1965a) that the big brown bat has good high-frequency hearing coupled with poor low-frequency hearing. Comparative analysis suggests that the bat's good high-frequency hearing initially evolved for passive sound localization and that it was later coopted for use in echolocation. In addition, the restricted low-frequency hearing of the big brown bat is typical of mammals with good high-frequency hearing.

Acoustic Stimulation↗

PubMed1997

Audiogram of the fox squirrel (Sciurus niger).

The behavioral audiograms of 2 fox squirrels (Sciurus niger) were determined with a conditioned avoidance procedure. The squirrels were able to hear tones ranging from 113 Hz to 49 kHz at a level of 60 dB sound-pressure level or less, with their best sensitivity of 1 dB occurring at 8 kHz. Their ability to hear frequencies below 150 Hz indicates that they have good low-frequency hearing, as do the 2 other members of the squirrel family (black-tailed and white-tailed prairie dogs) for which audiograms are available. This suggests that the ancestral sciurid may also have had good low-frequency hearing.

Animals↗

PubMed1997

Comparative study of sound localization and its anatomical correlates in mammals.

One of the fundamental features of hearing is the ability to localize the sources of sounds, particularly brief sounds, which may warn of nearby animals. Yet not all mammals localize sound equally well with threshold acuity ranging from about 1 degree for elephants and humans to more than 25 degrees for gerbils and horses and a near absence of localization in some subterranean species. During the past decade evidence has accumulated that this variation cannot be accounted for simply by the availability of the physical cues for locus. Nor does it appear to be a function of an animal's lifestyle. Rather sound-localization acuity in mammals appears to be a function of the precision required of the visual orienting response to sound. Thus the neural integration of hearing and vision in cortex, as well as in multimodal subcortical structures, is a reflection of their behavioral integration and evolutionary coupling.

Animals↗

PubMed1996

Sound localization in chinchillas, III: Effect of pinna removal.

The ability of chinchillas to make left/right, front/back, and vertical locus discriminations was determined before and after surgical removal of the pinnae. The animals were tested behaviorally using a conditioned avoidance procedure. In the left/right localization tests, removal of both pinnae had no effect on localization acuity for broadband noise but did result in a small decrement in performance when localizing low-pass filtered noise. In the front/back localization tests, removal of a single pinna resulted in a small but consistent decrement in performance when the sound sources were located in the hemifield on the same side as the intact pinna, and a greater decrement when the sound sources were located in the hemifield on the side of the missing pinna; removal of both pinnae resulted in the largest decrement in performance. Finally, vertical localization acuity and performance when localizing low-pass filtered noise were greatly impaired following removal of both pinnae. These results demonstrate the importance of the pinnae in performing front/back and vertical localization tasks in which binaural cues are not available.

Acoustic Stimulation↗

PubMed1995

Sound localization in chinchillas. II. Front/back and vertical localization.

The ability of chinchillas to make front/back and vertical locus discriminations was examined behaviorally using a conditioned avoidance procedure. Their minimum audible angle for localizing single broadband noise bursts was 36 degrees for front/back localization and 23 degrees for vertical localization. Sound localization tests using filtered noise demonstrated that the signal must contain high frequencies in order for chinchillas to make front/back and vertical locus judgements and that frequencies in their highest audible octave (i.e., above 16 kHz) contribute to localization. These results support the view that a major selective advantage of high-frequency hearing in mammalian evolution was its utility for monaural as well as binaural sound localization.

Acoustic Stimulation↗

PubMed1994

Sound localization in chinchillas. I: Left/right discriminations.

The ability of chinchillas to localize sound was examined behaviorally using a conditioned avoidance procedure in which the animals were trained to discriminate left from right sound sources. Their minimum audible angle was 15.6 degrees for 100-ms broadband noise making them one of the more accurate rodents, although they are not as accurate as primates and carnivores. Thresholds obtained for filtered noise stimuli demonstrated that chinchillas are equally accurate in localizing either low- or high-frequency noise. Further, they are able to use both interaural phase-difference and interaural intensity-difference cues as demonstrated by their ability to localize both low- and high-frequency pure tones. Finally, analysis of the chinchilla retina supports the hypothesis that the role of auditory localization in directing the eyes to sound sources played a role in the evolution of auditory spatial perception.

Acoustic Stimulation↗

PubMed1994

Hearing in prairie dogs: transition between surface and subterranean rodents.

Behavioral audiograms were determined for four black-tailed and one white-tailed prairie dogs (Cynomys ludovicianus and C. leucurus) using a conditioned avoidance procedure. The hearing of black-tailed prairie dogs ranges from 29 Hz to 26 kHz and that of the white-tailed prairie dog from 44 Hz to 26 kHz (at sound pressure levels of 60 dB). Both species have good low-frequency hearing, especially black-tailed prairie dogs which can hear as low as 4 Hz and are more sensitive than any other rodent yet tested at frequencies below 63 Hz. In contrast, prairie dogs are relatively insensitive in their midrange and have poor high-frequency hearing. It is suggested that the reduced midrange sensitivity and high-frequency hearing are related to their adaptation to an underground lifestyle with its reduced selective pressure for sound localization. In this respect they appear to be intermediate between the more exclusively subterranean rodents (such as gophers and mole rats) and surface dwellers (such as chinchillas and kangaroo rats).

Acoustic Stimulation↗

PubMed1994

Audiogram of the hooded Norway rat.

The behavioral audiogram of the hooded Norway rat was determined for frequencies from 250 Hz to 70 kHz. The resulting audiogram is virtually identical to the albino rat audiogram obtained by Kelly and Masterton (1977), indicating that there is no detectable effect of albinism on the audiogram of the Norway rat. The two audiograms also indicate the degree of replicability that can be obtained with current behavioral techniques.

Acoustic Stimulation↗

PubMed1993

Degenerate hearing and sound localization in naked mole rats (Heterocephalus glaber), with an overview of central auditory structures.

Behavioral tests of absolute sensitivity and sound localization in African naked mole rats show that, despite their communal social structure and large vocal repertoire, their hearing has degenerated much like that of other subterranean species. First, their ability to detect sound is limited, with their maximum sensitivity being only 35 dB (occurring at 4 kHz). Second, their high-frequency hearing is severely limited, with their hearing range (at 60 dB sound pressure level [SPL]) extending from 65 Hz to only 12.8 kHz. Third, determination of the effect of duration on noise thresholds indicates that, compared with other animals, mole rats require a sound to be present for a much longer duration before reaching asymptotic threshold. Finally, they are unable consistently to localize sounds shorter than 400 ms and cannot accurately localize sounds of longer duration, raising the possibility that they are unable to use binaural locus cues. Thus, it seems that the essentially one-dimensional burrow system of a subterranean habitat produces severe changes in hearing comparable to the changes in vision that result from the absence of light. To explore the relation between vision and sound-localization acuity, retinal ganglion cell densities were determined. The results indicate that naked mole rats have a broad area of best (albeit poor) vision, with maximum acuity estimated at 44 cycles/degree. That mammals with wide fields of best vision have poorer sound-localization acuity than those with narrower fields is consistent with the thesis that a major function of sound localization is to direct the gaze to the source of a sound. However, the fact that subterranean mammals have little use for vision in a lightless environment suggests that they represent an extreme case in this relationship and may explain the fact that, unlike surface-dwelling mammals, they have virtually lost the ability to localize brief sounds. Finally, despite their very limited auditory abilities, the major brainstem auditory nuclei, although relatively small, appear to be present.

Animals↗

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↗