Use of binaural cues for sound localization in two species of Phyllostomidae: the Greater spear-nosed bat (Phyllostomus hastatus) and the Short-tailed fruit bat (Carollia perspicillata)

Unlike humans, not all mammals use both of the binaural cues for sound localization. Whether an animal uses these cues can be determined by testing its ability to localize pure tones; specifically, low frequencies are localized using time-difference cues, and high frequencies are localized using intensity-difference cues. We determined the ability to use binaural cues in 2 New World bats, Phyllostomus hastatus, large omnivores, and Carollia perspicillata, small frugivores, by testing their tone-localization ability using a conditioned avoidance procedure. Both species easily localized high-frequency tones, indicating that they could use the interaural intensity-difference cue. However, neither species was able to use the phase-difference cue to localize either low-frequency pure tones or amplitude-modulated tones (which provided an envelope for additional time analysis). We now know of 3 bat species that cannot use binaural time cues and 2 that can. Further exploration of localization in bats may provide insight into the neural analysis of time cues in species that do not hear low frequencies.     

Sound localization in a predatory rodent, the northern grasshopper mouse (Onychomys leucogaster)

A comparison of the ability of mammals to localize sound revealed that among the animals examined to date, none of the rodents have been able to localize as accurately as the carnivores. Because all of these rodents are prey animals, the question arises as to whether their poor localization acuity is a phyletic trait of Rodentia or whether it is a trait common to prey species that may be under less selective pressure than predators to localize sound accurately. To answer this question, sound localization acuity was determined in a species that is both predatory and a rodent, the northern grasshopper mouse. Localization thresholds for a single 100-ms noise burst were determined for three grasshopper mice using a conditioned avoidance procedure. Their 50% discrimination threshold of 19 degrees is larger than that of any of the previously tested carnivores and well within the range of other rodents. However, calculations of the binaural sound localization cues available to rodents (based on their head size) suggest that the grasshopper mouse may make more efficient use of the available locus cues than other rodents. Thus, although the grasshopper mouse cannot localize as accurately as carnivores, it appears to be more accurate than predicted for a nonpredatory rodent of its size.     

Neuroanatomical basis of binaural phase-difference analysis for sound localization: a comparative study.

Four varieties of mammals whose medial superior olives range from large to none at all were tested for their ability to localize single, brief tone pips at various frequencies. Although each animal could localize high-frequency tone pips, their ability to localize middle- and low-frequency tone pips corresponded to the size of their medial superior olive (MSO). Since this latter range of frequencies is the one in which binaural phase-difference cues predominate, this anatomical-behavioral correspondence supports the idea that MSO is the chief binaural time-analyzing center for sound localization.     

Control of responding by the location of sound: role of binaural cues.

In auditory localization experiments, where the subject observes from a fixed position, both relative sound intensity and arrival time at the two ears determine the extent of localization performance. The present experiment investigated the role of binaural cues in a different context, the sound-position discrimination task, where the subject is free to move and interact with the sound source. The role of binaural cues was investigated in rats by producing an interaural imbalance through unilateral removal of the middle auditory ossicle (incus) prior to discrimination training. Discrete trial go-right/go-left sound-position discrimination of unilaterally incudectomised rats was then compared with that of normal rats and of rats with the incus of both sides removed. While bilateral incus removal affected binaural intensity and arrival times, the symmetry of sound input between the two ears was preserved. Percentage of correct responses and videotaped observations of sound approach and exploration showed that the unilateral rats failed to localize the sounding speaker. Rats with symmetrical binaural input (normal and bilaterally incudectomised rats) accurately discriminated sound position for the duration of the experiment. Previously reported monaural localization based upon following the intensity gradient to the sound source was not observed in the unilaterally incudectomised rats of the present experiment. It is concluded that sound-position discrimination depends upon the use of binaural cues.     

The influence of stimulus bandwidth on localization of sound in space

The ability of listeners, deprived of prominent interaural time and intensity cues, to locate noise bands differing in width was investigated. To minimize binaural cues, we placed the sound source at various positions in the median sagittal plane. To eliminate binaural cues, we occluded one ear. The stimuli consisted of broadband noise and bands of noise centered at 8.0 kHz. The width of the latter ranged from 1.0 to 6.0 kHz. The results from seven listeners showed that localization proficiency for sounds in the median sagittal plane decreased with decreases in bandwidth for both binaural and monaural listening conditions. This function was less orderly for monaural localization of horizontally positioned sounds. Another consequence of a reduction in bandwidth was an increasing tendency of listeners to select certain loudspeakers over others as the source of the sound. A previous finding showing that localization of sound in the median sagittal plane is more accurate when listening binaurally rather than monaurally was confirmed.     

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.     

Binaural and monaural localization of sound in two-dimensional space

Two experiments were conducted. In experiment 1, part 1, binaural and monaural localization of sounds originating in the left hemifield was investigated. 104 loudspeakers were arranged in a 13 x 8 matrix with 15 degrees separating adjacent loudspeakers in each column and in each row. In the horizontal plane (HP), the loudspeakers extended from 0 degrees to 180 degrees; in the vertical plane (VP), they extended from -45 degrees to 60 degrees with respect to the interaural axis. Findings of special interest were: (i) binaural listeners identified the VP coordinate of the sound source more accurately than did monaural listeners, and (ii) monaural listeners identified the VP coordinate of the sound source more accurately than its HP coordinate. In part 2, it was found that foreknowledge of the HP coordinate of the sound source aided monaural listeners in identifying its VP coordinate, but the converse did not hold. In experiment 2, part 1, localization performances were evaluated when the sound originated from consecutive 45 degrees segments of the HP, with the VP segments extending from -22.5 degrees to 22.5 degrees. Part 2 consisted of measuring, on the same subjects, head-related transfer functions by means of a miniature microphone placed at the entrance of their external ear canal. From these data, the 'covert' peaks (defined and illustrated in text) of the sound spectrum were extracted. This spectral cue was advanced to explain why monaural listeners in this study as well as in other studies performed better when locating VP-positioned sounds than when locating HP-positioned sounds. It is not claimed that there is inherent advantage for localizing sound in the VP; rather, monaural localization proficiency, whether in the VP or HP, depends on the availability of covert peaks which, in turn, rests on the spatial arrangement of the sound sources.     

3-D localization of virtual sound sources: Effects of visual environment, pointing method, and training

The ability to localize sound sources in three-dimensional space was tested in humans. In Experiment 1, naive subjects listened to noises filtered with subject-specific head-related transfer functions. The tested conditions included the pointing method (head or manual pointing) and the visual environment (VE; darkness or virtual VE). The localization performance was not significantly different between the pointing methods. The virtual VE significantly improved the horizontal precision and reduced the number of front-back confusions. These results show the benefit of using a virtual VE in sound localization tasks. In Experiment 2, subjects were provided with sound localization training. Over the course of training, the performance improved for all subjects, with the largest improvements occurring during the first 400 trials. The improvements beyond the first 400 trials were smaller. After the training, there was still no significant effect of pointing method, showing that the choice of either head- or manual-pointing method plays a minor role in sound localization performance. The results of Experiment 2 reinforce the importance of perceptual training for at least 400 trials in sound localization studies.     

Development of the auditory orientation response in the albino rat (Rattus norvegicus)

The development of head orientation to auditory stimulation was examined in rat pups at Postnatal Days 8, 11, 14, 17, and 20. The animals were tested in a quiet environment with single bursts of 65 dB (SPL) broad-band noise. A reflexive head turn toward the sound was first seen on Postnatal Day 14 and subsequently on Days 17 and 20. This result demonstrates that the onset of directional auditory responses occurred between Day 11 and Day 14. The role of binaural cues in early sound orientation was examined in 17-day-old pups with monaural ligation of the external meatus. These animals were unable to localize a sound source and consistently turned toward the side of the unligated ear regardless of the position of the stimulus. Thus binaural cues were shown to be important for head orientation to sound in early development. In a separate study, head orientation to high and low frequency tone pips was examined. Directional responses were first seen on Day 12 for a 16-kHz tone and Day 14 for a 2-kHz tone. These results indicate an earlier onset for orientation to high frequency sounds in the rat.     

Infants’ understanding of auditory events

Infants’ reaching‐in‐the‐dark was studied in a sample of normal 7.5–11‐month‐olds to determine whether infants can use sound cues to localize and recognize the action and objects of complex events. Infants were shown an event in which a moving, sounding object rotated clockwise through the infant's reaching space in the light and dark. Infrared recorded videotapes were later coded for reaching behaviour. Results showed that infants were able to localize the object on most trials in the dark but were slower and less efficient than in the light. Infants grasped the object at first contact and contacted the object near its salient feature in the dark, suggesting recognition of the object. Further, contact time was 1.7 s less when infants grasped the object at first contact in the dark (recognition) than when they touched the object, suggesting that recognition of the object improves reaching efficiency. There were no age and gender differences. In sum, the results support the use of the reaching‐in‐the‐dark method to demonstrate auditory localization of moving sounds and to reveal infants capacity to use represented information to guide subsequent action. Copyright © 1999 John Wiley & Sons, Ltd.     

Auditory spatial responses of young guinea pigs (Cavia porcellus) during and after ear blocking

Newborn guinea pigs were tested to determine their ability to approach an auditory stimulus early in development. Observations of the behavior of 1-4-day-old animals in a circular eight-choice maze revealed a pronounced tendency to orient toward and approach a tape-recorded signal of guinea pig vocalizations. The occurrence of approach responses was reduced to chance in animals tested with one ear occluded by wax ear plugs which attenuated but did not totally eliminate sound. The effect of monaural ear blocks was more severe than binaural blocks, which reflects the importance of binaural cues in the maintenance of approach responses to sound. In a second study, the ability of older animals, 11-31 days of age, was examined. Directional approach responses to sound were also evident at this age, and ear plugs disrupted performance only under monaural conditions. Furthermore, in animals raised from birth with monaural ear blocks but tested without ear plugs, there was a subsequent disruption of performance for at least 21 days. These results indicate the importance of binaural cues in the development of early auditory spatial responses and suggest the need for appropriate binaural experience for subsequent localization of sounds.     

Congruent auditory display and confusion in sound localization: Case of elderly drivers

Elderly people may suffer from age-related hearing loss. They often report difficulties to perceive and localize sound sources in noisy environment. How this can be a driving safety issue? This study investigates the effect of hearing impairment, on the driver behavior to localize external sound sources such as emergency vehicles sirens. Subjective tests show that localization confusion appears to be a common problem. Thus, we focus on how to assist the driver, taking into consideration the age-related hearing impairment, to better localize emergency vehicle siren. The hypothesis was based on the stimulus-response compatibility using an effective congruent auditory display, for attentional guidance toward the direction of arrival (DOA) of the external alarm. The proposed approach aims to reduce front-back confusion and enhance the sound localization accuracy of the driver, which is very important for elderly driver, subject to older age cognitive decline. A localization test was performed in lab and in vivo on a test track, where a group of drivers was asked to identify the DOA of an emergency vehicle siren, with and without a dedicated embedded system set up to assist them on sound localization task.     

What neuroimaging and brain localization can do, cannot do and should not do for social psychology

Interest in bridging social psychology and neuroscience has seen a significant upsurge. Much of this interest has centered on brain localization--the attempt to relate psychological events to locations of brain events. Although many articles have sought to localize brain activity that supports social behavior, scant attention has been paid to the specific methods to be used in integrating brain localization data into psychological theory. The authors describe 4 strategies psychologists can use to integrate brain localization data and psychological theory, and they consider whether social psychology presents special considerations in the use of these strategies. They conclude that brain localization offers a useful tool for some but not all problems in social psychology, and they discuss the types of problems for which it may and may not prove useful.     

Localization of sound in the vertical plane with and without high-frequency spectral cues.

Binaural localization of 3.0-kHz high- and lowpass noise presented in the median vertical plane (MVP) and lateral vertical plane (LVP) was investigated. We anticipated superior performance when localizing the highpass noise by virtue of the availability of pinna cues. The viability of this supposition was strengthened by monaural localization tests in which performance proficiency for the highpass noise exceeded that for the lowpass noise (p less than .01). The main result showed that binaural localization of proficiency for highpass noise surpassed that for lowpass noise for all listening conditions (p less than .01). However, the importance of binaural temporal and level differences in vertical-plane localization was demonstrated by the highly respectable performances when the lowpass noise was presented in the LVP. Data from binaural localization in the MVP and monaural localization in the LVP suggested that the influence of pinna cues diminishes for source elevations above 45 degrees.     

Tactile localization of the direction and distance of sounds

In a previous paper, experiments were reported which demonstrated that human subjects can judge accurately the azimuthal direction of sounds using a tactile localization device. It was also demonstrated that a tactile analogue of selective auditory attention was possible with this system. Three additional experiments, reported here, indicate that subjects are also able to judge the distance of the sound source and can concurrently judge both the azimuthal direction and distance of a source. Comparisons were made between conditions where head movements were permitted lactive~ and conditions where the head was held still Ipassive, and between normal auditory judgments and tactile judgments. Active tactile performance was essentially similar to auditory performance. Active performance was superior to passive in both directional and distance judgment, but different components of the motor-sensory complex were found to contribute to active superiority in the two tasks. The implication of these experiments for the design of auditory prosthetic devices is discussed.     

An automated sound-localization chamber

A completely automated free-field sound-localization apparatus is described along with a behavioral technique that provides several advantages over previously available methods. With the apparatus, the ability to localize the source of a sound has been tested in a wide variety of mammals including cats, rats, squirrel monkeys, hedgehogs, and tree shews.     

Lateralization of narrow-band noise by blind and sighted listeners

The effects of varying interaural time delay (ITD) and interaural intensity difference (IID) were measured in normal-hearing sighted and congenitally blind subjects as a function of eleven frequencies and at sound pressure levels of 70 and 90 dB, and at a sensation level of 25 dB (sensation level refers to the pressure level of the sound above its threshold for the individual subject). Using an 'acoustic' pointing paradigm, the subject varied the IID of a 500 Hz narrow-band (100 Hz) noise (the 'pointer') to coincide with the apparent lateral position of a 'target' ITD stimulus. ITDs of 0, +/-200, and +/-400 micros were obtained through total waveform delays of narrow-band noise, including envelope and fine structure. For both groups, the results of this experiment confirm the traditional view of binaural hearing for like stimuli: non-zero ITDs produce little perceived lateral displacement away from 0 IID at frequencies above 1250 Hz. To the extent that greater magnitude of lateralization for a given ITD, presentation level, and center frequency can be equated with superior localization abilities, blind listeners appear at least comparable and even somewhat better than sighted subjects, especially when attending to signals in the periphery. The present findings suggest that blind listeners are fully able to utilize the cues for spatial hearing, and that vision is not a mandatory prerequisite for the calibration of human spatial hearing.     

Directional responses to sounds in young gerbils (Meriones unguiculatus)

Three experiments were conducted to determine the ability of infant gerbils to approach an auditory stimulus. In the first experiment, gerbil pups, 16-23 days of age, were tested in a circular apparatus with a central start area and a movable sound source located at one of eight positions around the perimeter. Stimuli included high- and low-intensity presentations of a tape-recorded gerbil social call, a broad-band white noise stimulus, and a no-stimulus control condition. The subjects showed a strong tendency to approach the low-intensity social call and a less pronounced tendency to approach the white noise. In the second experiment, gerbil pups were tested in the same apparatus with or without ear blocks to determine the role of binaural cues in directional approach responding. The tendency to approach a low-intensity vocalization was disrupted by obstruction of one ear but not by blocking both ears. Thus, binaural balance was shown to be important for early sound localization. In the third experiment, the tendency to approach a social call was compared at different ages, 12-15, 16-19, 20-23, and 24-27 days after birth. Approach responses were first seen at 16-19 days. The responses continued during the 20-23-day period but began to wane at 24-27 days of age.     

Localization of sound in the vertical plane with and without high-frequency spectral cues

Binaural localization of 3.0-kHz high- and lowpass noise presented in the median vertical plane (MVP) and lateral vertical plane (LVP) was investigated. We anticipated superior performance when localizing the highpass noise hy virtue of the availability of pinna cues. The viability of this supposition was strengthened by monaural localization tests in which performance proficiency for the highpass noise exceeded that for the lowpass noise (p < .01). The main result showed that binaural localization of proficiency for highpass noise surpassed that for lowpass noise for all listening conditions (p < .01). However, the importance of binaural temporal and level differences in vertical-plane localization was demonstrated by the highly respectable performances when the lowpass noise was presented in the LVP. Data from binaural localization in the MVP and monaural localization in the LVP suggested that the influence of pinna cues diminishes for source elevations above 45°.     

Learning where to feed: the use of social information in flower-visiting Pallas’ long-tongued bats (<Emphasis Type="Italic">Glossophaga soricina</Emphasis>)

Social learning is a widespread phenomenon among vertebrates that influences various patterns of behaviour and is often reported with respect to foraging behaviour. The use of social information by foraging bats was documented in insectivorous, carnivorous and frugivorous species, but there are little data whether flower-visiting nectarivorous bats (Phyllostomidae: Glossophaginae) can acquire information about food from other individuals. In this study, we conducted an experiment with a demonstrator-observer paradigm to investigate whether flower-visiting Pallas’ long-tongued bats (Glossophaga soricina) are able to socially learn novel flower positions via observation of, or interaction with, knowledgeable conspecifics. The results demonstrate that flower-visiting G. soricina are able to use social information for the location of novel flower positions and can thereby reduce energy-costly search efforts. This social transmission is explainable as a result of local enhancement; learning bats might rely on both visual and echo-acoustical perception and are likely to eavesdrop on auditory cues that are emitted by feeding conspecifics. We additionally tested the spatial memory capacity of former demonstrator bats when retrieving a learned flower position, and the results indicate that flower-visiting bats remember a learned flower position after several weeks.