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.     

The influence of the acoustic context on vertical sound localization in the median plane

The influence of background sounds (frames) on vertical localization of single sound sources (targets) was examined in four experiments. Loudspeakers (five targets and four frames) were positioned in the median plane, ranging from +30 degrees to -30 degrees above and below the subject's ear level. The subjects determined the vertical position of the targets by either verbal judgments or manual pointing. Frame and target sounds were presented concurrently or successively with a 1-sec interval; both consisted of (1) 300-Hz square waves, (2) noise, or (3) targets of noise and frames of 300-Hz square waves. Particularly in the second condition, the subjects consistently shifted the apparent target positions away from the frame locations. This contrast effect persisted even 1 sec after the offset of the frames. No effect was found with different waveforms for the frame and the target. Results are related to recent findings indicating a similar effect in the azimuthal dimension. Possibly the effect is based on a mechanism in which the auditory system adapts to recently heard sound source positions.     

Spatial position versus ear of entry as determinant of the auditory laterality effects: a stereophonic test.

The study concerned discriminating between ear of entry and apparent spatial position as possible determinants of lateral asymmetries in the recall of simultaneous speech messages. Apparent localization to the left or right of the median plane was created either through a time difference (.7 msec), through intensity differences between presentations of the same verbal message at the two ears, or through dichotic presentations. Right-side advantage was observed with the three types of presentation (Experiments 1, 2, and 3). The finding of right-side advantage with stereophony based on a time difference only, in the absence of intensity difference, cannot be accounted for in terms of an ear advantage and shows that apparent spatial separation of the sources can by itself produce a laterality effect. Differences in the degree of lateral asymmetry between the various conditions were also observed. The findings of Experiments 4 and 5 suggest that these differences are better explained in terms of different impressions of localization of the sound sources than in terms of relative intensity at the "privileged" ear.     

Adaptation to auditory motion in the horizontal plane: effect of prior exposure to motion on motion detectability.

Thresholds for auditory motion detectability were measured in a darkened anechoic chamber while subjects were adapted to horizontally moving sound sources of various velocities. All stimuli were 500-Hz lowpass noises presented at a level of 55 dBA. The threshold measure employed was the minimum audible movement angle (MAMA)--that is, the minimum angle a horizontally moving sound must traverse to be just discriminable from a stationary sound. In an adaptive, two-interval forced-choice procedure, trials occurred every 2-5 sec (Experiment 1) or every 10-12 sec (Experiment 2). Intertrial time was "filled" with exposure to the adaptor--a stimulus that repeatedly traversed the subject's front hemifield at ear level (distance: 1.7 m) at a constant velocity (-150 degrees/sec to +150 degrees/sec) during a run. Average MAMAs in the control condition, in which the adaptor was stationary (0 degrees/sec,) were 2.4 degrees (Experiment 1) and 3.0 degrees (Experiment 2). Three out of 4 subjects in each experiment showed significantly elevated MAMAs (by up to 60%), with some adaptors relative to the control condition. However, there were large intersubject differences in the shape of the MAMA versus adaptor velocity functions. This loss of sensitivity to motion that most subjects show after exposure to moving signals is probably one component underlying the auditory motion aftereffect (Grantham, 1989), in which judgments of the direction of moving sounds are biased in the direction opposite to that of a previously presented adaptor.     

A two-choice sound localization procedure for detecting lateralized tinnitus in animals

Rats were trained in a two-choice procedure to respond in the direction of left and right sounds. Silent trials, on which no sound was presented and for which the animals received no feedback, were interspersed among the sound trials to determine each animal’s natural side preference. Following training, the rats were exposed to a loud tone in the ear opposite their side preference. A shift in responding on the silent trials to the side of the exposed ear indicated that the animals were hearing a sound in that ear (i.e., tinnitus). Simulating lateralized tinnitus by presenting a low-level, continuous sound on one side also caused the rats to shift their responding on the silent trials to that side. Sham exposures indicated that halothane/nitrous oxide anesthesia could reinstate tinnitus in animals that had previously tested positive for it. Exposing rats to loud tones of various frequencies indicated that frequencies near the limits of the rat’s hearing range were less likely to cause tinnitus than tones in the midrange.     

Relation of sound intensity and accuracy of localization

Tests were carried out on 17 subjects to determine the accuracy of monaural sound localization when the head is not free to turn toward the sound source. Maximum accuracy of localization for a constant-volume sound source coincided with the position for maximum perceived intensity of the sound in the front quadrant. There was a tendency for sounds to be perceived more often as coming from a position directly toward the ear. That is, for sounds in the front quadrant, errors of localization tended to be predominantly clockwise (i.e., biased toward a line directly facing the ear). Errors for sounds occurring in the rear quadrant tended to be anticlockwise. The pinna's differential effect on sound intensity between front and rear quadrants would assist in identifying the direction of movement of objects, for example an insect, passing the ear.     

Pigeons' performance under differential reinforcement of low rate schedules depends upon the operant

Pigeons were reinforced with grain for pecking a key or depressing a foot treadle according to differential reinforcement of low rate (DRL) schedules. Birds which depressed a treadle performed efficiently on DRL schedules as high as DRL 35-sec; while birds reinforced for keypecking showed low efficiency under DRL 14-sec. While treadle pressing and keypecking differ along a number of dimensions (including force requirement of the operant and differences in temporal distributions of responses), the present results are consistent with an interpretation based on differences in the degree to which these two responses are elicited by periodic presentations of food.     

Detour behaviour in three species of birds: quails (Coturnix sp.), herring gulls (Larus cachinnans) and canaries (Serinus canaria)

Detour behaviour is the ability of an animal to reach a goal stimulus by moving round any interposed obstacle. It has been widely studied and has been proposed as a test of insight learning in several species of mammals, but few data are available in birds. A comparative study in three species of birds, belonging to different eco-ethological niches, allows a better understanding of the cognitive mechanism of such detour behaviour. Young quails (Coturnix sp.), herring gulls (Larus cachinnans) and canaries (Serinus canaria), 1 month old, 10–25 days old and 4–6 months old, respectively, were tested in a detour situation requiring them to abandon a clear view of a biologically interesting object (their own reflection in a mirror) in order to approach that object. Birds were placed in a closed corridor, at one end of which was a barrier through which the object was visible. Four different types of barrier were used: vertical bar, horizontal bar, grid and transparent. Two symmetrical apertures placed midline in the corridor allowed the birds to adopt routes passing around the barrier. After entering the apertures, birds could turn either right or left to re-establish social contact with the object in the absence of any local sensory cues emanating from it. Quails appeared able to solve the task, though their performance depended on the type of barrier used, which appeared to modulate their relative interest in approaching the object or in exploring the surroundings. Young herring gulls also showed excellent abilities to locate spatially the out-of-view object, except when the transparent barrier was used. Canaries, on the other hand, appeared completely unable to solve the detour task, whatever barrier was in use. It is suggested that these species differences can be accounted for in terms of adaptation to a terrestrial or aerial environment.     

Landmark use by Clark’s nutcrackers (Nucifraga columbiana): influence of disorientation and cue rotation on distance and direction estimates

Many species have been shown to encode multiple sources of information to orient. To examine what kinds of information animals use to locate a goal we manipulated cue rotation, cue availability, and inertial orientation when the food-storing Clark’s nutcracker (Nucifraga columbiana) was searching for a hidden goal in a circular arena. Three groups of birds were used, each with a different goal–landmark distance. As the distance between the goal and the landmark increased, nutcrackers were less accurate in finding the correct direction to the goal than they were at estimating the distance (Experiment 1). To further examine what cues the birds were using to calculate direction, the featural cues within the environment were rotated by 90° and the birds were either oriented when searching (Experiments 2 and 3) or disoriented (Experiment 3). In Experiment 4, all distinctive visual cues were removed (both internal and external to the environment), a novel point of entry was used and the birds were either oriented or disoriented. We found that disorienting the nutcrackers so that they could not use inertial cues did not influence the birds’ total search error. The birds relied heavily but not completely on cues within the environment, as rotating available cues caused them to systematically shift their search behavior. In addition, the birds also relied to some extent on Earth-based cues. These results show the flexible nature of cue use by the Clark’s nutcracker. Our study shows how multiple sources of spatial information may be important for extracting multiple bearings for navigation.     

Left-side dominance for song discrimination in Bengalese finches (Lonchura striata var. domestica)

Male Bengalese finches are left-side dominant for the motor control of song in the sensorimotor nucleus (the high vocal center, or HVc) of the telencephalon. We examined whether perceptual discrimination of songs might also be lateralized in this species. Twelve male Bengalese finches were trained by operant conditioning to discriminate between a Bengalese finch song and a zebra finch song. Before training, the left HVc was lesioned in four birds and the right HVc was lesioned in four other birds. The remaining four birds were used as controls without surgery. Birds with a left HVc lesion required significantly more time to learn to discriminate between the two songs than did birds with a right HVc lesion or intact control birds. These results suggest that the left HVc is not only dominant for the motor control of song, but also for the perceptual discrimination of song. Accepted after revision: 11 September 2001 Electronic Publication     

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.     

Adaptation to auditory motion in the horizontal plane: Effect of prior exposure to motion on motion detectability

Thresholds for auditory motion detectability were measured in a darkened anechoic chamber while subjects were adapted to horizontally moving sound saurces of various-velocities. All stimuli were 500-Hz lowpass noises presented at a level of 55 dBA. The threshold measure employed was the minimum audible movement angle(MAMA)—that is, the minimum angle a horizontally moving sound must traverse to be just discriminable from a stationary sound. In an adaptive, two-interval forced-choice procedure, trials occurred every 2-5 sec (Experiment 1) or every 10–12 sec (Experiment 2). Intertrial time was “filled” with exposure to the adaptor—a stimulus that repeatedly traversed the subject’s front hemifield at ear level (distance: 1.7 m) at a constant velocity (?150°/secto + 150°/sec)during a run. Average MAMAs in the control condition, in which the adaptor was stationary (0°/sec), were 2.4° (Experiment 1) and 3.0° (Experiment 2). Three out of 4 subjects in each experiment showed significantly elevated MAMAs (by up to 60%), with some adaptors relative to the control condition. However, there were large intersubject differences in the shape of the MAMA versus adaptor velocity functions. This loss of sensitivity to motion that most subjects show after exposure to moving signals is probably one component underlying the auditory motion aftereffect (Grantham, 1989), in which judgmentsof the direction-afmoving sounds are biased in the direction opposite to that of a previously presented adaptor.     

The role of selection in generating auditory negative priming

The importance of selecting between a target and a distractor in producing auditory negative priming was examined in three experiments. In Experiment 1, participants were presented with a prime pair of sounds, followed by a probe pair of sounds. For each pair, listeners were to identify the sound presented to the left ear. Under these conditions, participants were especially slow to identify a sound in the probe pair if it had been ignored in the preceding prime pair. Evidence of auditory negative priming was also apparent when the prime sound was presented in isolation to only one ear (Experiment 2) and when the probe target was presented in isolation to one ear (Experiment 3). In addition, the magnitude of the negative priming effect was increased substantially when only a single prime sound was presented. These results suggest that the emergence of auditory negative priming does not depend on selection between simultaneous target and distractor sounds.     

Mechanisms underlying speech sound discrimination and categorization in humans and zebra finches

Speech sound categorization in birds seems in many ways comparable to that by humans, but it is unclear what mechanisms underlie such categorization. To examine this, we trained zebra finches and humans to discriminate two pairs of edited speech sounds that varied either along one dimension (vowel or speaker sex) or along two dimensions (vowel and speaker sex). Sounds could be memorized individually or categorized based on one dimension or by integrating or combining both dimensions. Once training was completed, we tested generalization to new speech sounds that were either more extreme, more ambiguous (i.e., close to the category boundary), or within-category intermediate between the trained sounds. Both humans and zebra finches learned the one-dimensional stimulus–response mappings faster than the two-dimensional mappings. Humans performed higher on the trained, extreme and within-category intermediate test-sounds than on the ambiguous ones. Some individual birds also did so, but most performed higher on the trained exemplars than on the extreme, within-category intermediate and ambiguous test-sounds. These results suggest that humans rely on rule learning to form categories and show poor performance when they cannot apply a rule. Birds rely mostly on exemplar-based memory with weak evidence for rule learning.     

Preference for spatial cues in a non-storing songbird species

Male mammals typically outperform their conspecific females on spatial tasks. A sex difference in cues used to solve the task could underlie this performance difference as spatial ability is reliant on appropriate cue use. Although comparative studies of memory in food-storing and non-storing birds have examined species differences in cue preference, few studies have investigated differences in cue use within a species. In this study, we used a one-trial associative food-finding task to test for sex differences in cue use in the great tit, Parus major. Birds were trained to locate a food reward hidden in a well covered by a coloured cloth. To determine whether the colour of the cloth or the location of the well was learned during training, the birds were presented with three wells in the test phase: one in the original location, but covered by a cloth of a novel colour, a second in a new location covered with the original cloth and a third in a new location covered by a differently coloured cloth. Both sexes preferentially visited the well in the training location rather than either alternative. As great tits prefer colour cues over spatial cues in one-trial associative conditioning tasks, cue preference appears to be related to the task type rather than being species dependent.     

The discrimination of temporal fine structure in call-like harmonic sounds by birds

Thresholds for discriminating changes in the temporal fine structure of call-like, harmonic sounds were measured in zebra finches (Taeniopygia guttata) and budgerigars (Melopsittacus undulatus). Birds could detect changes in periods as short as 1.225 ms at near 100% accuracy even when spectral and envelope cues were identical, as in time-reversed stimuli. Humans performed poorly on such stimuli, paralleling results from previous studies. Bird thresholds were in the range of those reported in neurophysiological studies of the songbird high vocal center (HVC) to temporally modified conspecific songs. Taken together, these results show that birds can hear differences in temporal fine structure in their natural vocalizations that go beyond human capabilities, but whether these abilities have communicative relevance remains to be seen.     

Responses of urban crows to con- and hetero-specific alarm calls in predator and non-predator zoo enclosures

Urban animals and birds in particular are able to cope with diverse novel threats in a city environment such as avoiding novel, unfamiliar predators. Predator avoidance often includes alarm signals that can be used also by hetero-specifics, which is mainly the case in mixed-species flocks. It can also occur when species do not form flocks but co-occur together. In this study we tested whether urban crows use alarm calls of conspecifics and hetero-specifics (jackdaws, Corvus monedula) differently in a predator and a non-predator context with partly novel and unfamiliar zoo animal species. Birds were tested at the Tiergarten Schönbrunn in the city of Vienna by playing back con- and hetero-specific alarm calls and control stimuli (great tit song and no stimuli) at predator (wolf, polar bear) and non-predator (eland antelope and cranes, peccaries) enclosures. We recorded responses of crows as the percentage of birds flying away after hearing the playback (out of those present before the playback) and as the number of vocalizations given by the present birds. A significantly higher percentage of crows flew away after hearing either con- or hetero-specific alarm calls, but it did not significantly differ between the predator and the non-predator context. Crows treated jackdaw calls just as crow calls, indicating that they make proper use of hetero-specific alarm calls. Responding similarly in both contexts may suggest that the crows were uncertain about the threat a particular zoo animal represents and were generally cautious. In the predator context, however, a high percentage of crows also flew away upon hearing the great tit control song which suggests that they may still evaluate those species which occasionally killed crows as more dangerous and respond to any conspicuous sound.     

Performance and reaction times in monaural localization of first names in the horizontal plane

Many studies have been conducted to measure monaural azimuthal sound localization performance with different sounds varying in frequency and complexity, but few have used linguistic stimuli. The present experimental design used subjects' first names in a monaural azimuthal localization task. Analysis of response accuracy showed that subjects are not more accurate in localizing their own first name than in localizing other first names and that there was no significant advantage of one ear over another. Reaction times were shorter when the subjects localized their own first name than when they localized any other first names and there was no significant ear advantage, but localizing other first names took more time with the right than with the left ear. All stimuli were better and more quickly localized on the side of the open ear, and there was no difference in acuity or velocity of localization with the two different speaker voices used. These results suggest that first names are processed through the controlateral auditory pathway and can be analyzed in the right hemisphere.     

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.     

Infants' localization of sounds in the median vertical plane: estimates of minimum audible angle

Infants 6, 9, 12, 15, and 18 months of age were seated in a dark room directly facing an array of nine loudspeakers positioned along the median vertical plane. One loudspeaker was positioned at ear level, 0 degree, and four others each were positioned above and below 0 degree. To examine infants' resolution of auditory space in the median vertical plane we sought to determine the smallest angular shift in the vertical location of a sound that infants could reliably detect (i.e., minimum audible angle). A two-alternative forced-choice procedure was used in which a sequence of white noise bursts was presented initially at 0 degree, and then shifted vertically (i.e., above or below 0 degree) and continued to be presented until the infant made a directional response; correct responses were visually reinforced. The smallest angular shift in vertical location that was reliably detected systematically decreased with increasing age between 6 months (15 degrees) and 18 months (4 degrees), suggesting a finer partitioning of auditory space along the vertical axis over this age range. By 18 months infants' performance matched that of a group of adults tested under the same circumstances.