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.     

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.     

Use of position and feature cues in discrimination learning by the whiptail lizard (Cnemidophorus inornatus)

Animals use a variety of cue types to locate and discriminate objects. The ease with which particular cue types are learned varies across species and context. An enormous literature contains comparisons of spatial cue use to use of other cue types, but few experiments examine the ease with which various nonspatial cues are learned. In addition, few studies have examined cue use in reptiles. Thus, the authors compared whiptail lizards' (Cnemidophorus inornatus) ability to learn and reverse a discrimination using either position (left or right) or visual feature cues. Lizards learned and reversed the task using position cues faster and with greater accuracy than using feature cues.     

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.     

Temperature sensitivity: One subjective continuum or two?

An experiment is reported on the effect of six consecutive days of exposure to distortions of binaural time and intensity cues on the localization function for tones of 350, 1000, and 4000 Hz. Tests were conducted under “free-field” conditions. The changes in auditory spatial orientation indicate that adjustments to binaurally distorted temporal and intensive cues can occur. Reorientation does not appear to depend upon the existence of veridical auditory 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.     

Negating the effects of binaural cues: competition between auditory streaming and contralateral induction

When a pair of monaural pure tones, A and B, are repeatedly alternated in one ear, with noise bursts presented in synchrony with B in the other ear, the noise sometimes delateralizes B. This is presumably a case of Warren and Bashford's (1976) contralateral induction effect. However, the present experiment shows that the degree of contralateral induction is proportional to the separation in frequency between A and B. It was also found that the degree to which the noise bursts influenced B's timbre was proportional to the separation in frequency between A and B. The combined results suggest that cues that govern the sequential organization of sounds influence the use of binaural cues not only during the assignment of position to auditory events but during the assignment of timbre.     

Cuing mechanisms in autitory signal detection

Detection of auditory signals under frequency uncertainty can be improved by presenting cues to the listeners. Since various cues have been found to differ in effectiveness, three conceivable mechanisms were considered which might account for these differences. Cuing might reduce the number and/or width of the employed auditory filters or listening bands. Also, cues could modulate the precision of frequency tuning of the filters. Psychometric functions were collected in a detection experiment with frequency uncertainty employing three kinds of cues: pure tones whose frequency was identical to that of the signal (iconic cues), complex tones with a missing fundamental equal to the signal (complex cues), and pure tones with a certain frequency relation to the signal (relative cues). Compared with a no-cue condition, all cue types improved detection performance. Fitting models to the data suggests that in the no-cue condition as well as the complex-cue condition, multiple bands were utilized, and that the iconic and relative cues induced single-band listening. There is no indication that accuracy of frequency tuning was responsible for cue-efficiency differences.     

Learning and limits of use of eye gaze by capuchin monkeys (Cebus apella) in an object-choice task

The ability of 3 capuchin monkeys (Cebus apella) to use experimenter-given cues to solve an object-choice task was assessed. The monkeys learned to use explicit gestural and postural cues and then progressed to using eye-gaze-only cues to solve the task, that is, to choose the baited 1 of 2 objects and thus obtain a food reward. Increasing cue-stimulus distance and introducing movement of the eyes impeded the establishment of effective eye-gaze reading. One monkey showed positive but imperfect transfer of use of eye gaze when a novel experimenter presented the cue. When head and eye orientation cues were presented simultaneously and in conflict, the monkeys showed greater responsiveness to head orientation cues. The results show that capuchin monkeys can learn to use eye gaze as a discriminative cue, but there was no-evidence for any underlying awareness of eye gaze as a cue to direction of attention.     

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.     

Attributing attention: the use of human-given cues by domestic horses (Equus caballus)

Recent research has shown that domestic dogs are particularly good at determining the focus of human attention, often outperforming chimpanzees and hand-reared wolves. It has been suggested that the close evolutionary relationship between humans and dogs has led to the development of this ability; however, very few other domestic species have been studied. We tested the ability of 36 domestic horses to discriminate between an attentive and inattentive person in determining whom to approach for food. The cues provided were body orientation, head orientation or whether the experimenters’ eyes were open or closed. A fourth, mixed condition was included where the attentive person stood with their body facing away from the subjects but their head turned towards the subject while the inattentive person stood with their body facing the subject but their head turned away. Horses chose the attentive person significantly more often using the body cue, head cue, and eye cue but not the mixed cue. This result suggests that domestic horses are highly sensitive to human attentional cues, including gaze. The possible role of evolutionary and environmental factors in the development of this ability is discussed.     

Flexibility of cue use in the fox squirrel (Sciurus niger)

Recent work on captive flying squirrels has demonstrated a novel degree of flexibility in the use of different orientation cues. In the present study, we examine to what extent this flexibility is present in a free-ranging population of another tree squirrel species, the fox squirrel. We trained squirrels to a rewarded location within a square array of four feeders and then tested them on transformations of the array that either pitted two cue types against one cue type, the majority tests, or all cue types against each other, the forced-hierarchy test. In Experiment 1, squirrels reoriented to the two-cue-type location in all majority tests and to the location indicated by the visual features of the feeders in the forced-hierarchy test. This preference for visual features runs contrary to previous studies that report the use of spatial cues over visual features in food-storing species. In Experiments 2-5 we tested squirrels with different trial orders (Experiments 2 and 3), a different apparatus (Experiment 4) and at different times of the year (Experiment 5) to determine why these squirrels had chosen to orient using visual features in the first experiment. Like captive flying squirrels, free-ranging fox squirrels showed a large degree of flexibility in their use of cues. Furthermore, their cue use appeared to be sensitive both to changes in the test apparatus and the season in which we tested. Altogether our results suggest that the study of free-ranging animals over a variety of conditions is necessary for understanding spatial cognition.     

Domestic dogs (Canis familiaris) use human gestures but not nonhuman tokens to find hidden food

The authors examined the ability of domestic dogs to use human body cues (gestures) and equivalent-sized nonhuman cues to find hidden food in an object choice paradigm. In Experiment 1 the authors addressed the importance of the human element of the cue, and the effects of size, topography, and familiarity on dogs' success in using cues. Experiment 2 further explored the role of the human as cue-giver, and the impact of a change in the experimenter's attentional state during cue presentation. This included a systematic test of the role inanimate tokens play as cues apart from human placement. Our results indicate that dogs are more sensitive to human cues than equivalent nonhuman cues, and that the size of the cue is a critical element in determining dogs' success in following it.     

Pure-tone perception and ear advantages in dichotic listening

A same-different matching task was used to investigate how subjects perceived a dichotic pair of pure tones. Pairs of stimulus tones in four frequency ranges (center frequencies of 400–1,700 Hz), with separations between 40 and 400 Hzt were tested. Five types of test tones were matched to the stimulus pair: the stimulus pair presented again (control) or crossed over (same tones, different ears), the geometric mean of the two tones, or a binaural tone of the low or high tone of the pair. In the lowest frequency range and the highest with maximum separation, the crossed-over test tones were perceived as different from the same stimulus tones. A bias for perceiving the higher tone of a pair was evident in the frequency ranges with separations of 40-200 Hz. In the lowest frequency range, the bias was for perceiving the higher tone in the right ear. This restricted ear advantage in the perception of pure tones was not significantly related to the right-ear advantage in dichotic word monitoring.     

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.     

Auditory grouping

Although our subjective experience of the world is one of discrete sound sources, the individual frequency components that make up these separate sources are spread across the frequency spectrum. Listeners. use various simple cues, including common onset time and harmonicity, to help them achieve this perceptual separation. Our ability to use harmonicity to segregate two simultaneous sound sources is constrained by the frequency resolution of the auditory system, and is much more effective for low-numbered, resolved harmonics than for higher-numbered, unresolved ones. Our ability to use interaural time-differences (ITDs) in perceptual segregation poses a paradox. Although ITDs are the dominant cue for the localization of complex sounds, listeners cannot use ITDs alone to segregate the speech of a single talker from similar simultaneous sounds. Listeners are, however, very good at using ITD to track a particular sound source across time. This difference might reflect two different levels of auditory processing, indicating that listeners attend to grouped auditory objects rather than to those frequencies that share a common ITD.     

Use of human visual attention cues by olive baboons (Papio anubis) in a competitive task

The ability of 4 olive baboons (Papio anubis) to use human gaze cues during a competitive food task was investigated. Three baboons used head orientation as a cue, and 1 individual also used eye direction alone. As the baboons did not receive prior training with gestural cues, their performance suggests that the competitive paradigm may be more appropriate for testing nonhuman primates than the standard object-choice paradigm. However, the baboons were insensitive to whether the experimenter could actually perceive the food item, and therefore the use of visual orientation cues may not be indicative of visual perspective-taking abilities. Performance was disrupted by the introduction of a screen and objects to conceal food items and by the absence of movement in cues presented.     

Effects of redundant auditory stimuli on reaction time

Auditory redundancy gains were assessed in two experiments in which a simple reaction time task was used. In each trial, an auditory stimulus was presented to the left ear, to the right ear, or simultaneously to both ears. The physical difference between auditory stimuli presented to the two ears was systematically increased across experiments. No redundancy gains were observed when the stimuli were identical pure tones or pure tones of different frequencies (Experiment 1). A clear redundancy gain and evidence of coactivation were obtained, however, when one stimulus was a pure tone and the other was white noise (Experiment 2). Experiment 3 employed a two-alternative forced choice localization task and provided evidence that dichotically presented pure tones of different frequencies are apparently integrated into a single percept, whereas a pure tone and white noise are not fused. The results extend previous findings of redundancy gains and coactivation with visual and bimodal stimuli to the auditory modality. Furthermore, at least within this modality, the results indicate that redundancy gains do not emerge when redundant stimuli are integrated into a single percept.     

Invalid cues impair auditory motion sensitivity

Compelling lateral motion can be experienced when intensity differences between the two cars change over time. Whether our sensitivity to this dynamic interaural stimulation could be influenced by directional cues was the focus of the present study. On each trial, amplitude-modulated pure tones were presented either diotically (no-motion condition) or dichotically (motion condition), and participants indicated whether lateral motion was present or absent. Randomly across trials, the stimuli were preceded by a valid directional cue, an invalid directional cue, or no cue, while the motion to be detected was identical across these cue conditions. The data indicate that motion sensitivity was comparable in the valid-cue and no-cue conditions. Relative to each of those conditions, however, motion sensitivity was significantly lower in the invalid-cue condition, and motion was reported significantly less often. The results provide evidence that our sensitivity to dynamic interaural intensity differences can be significantly affected by a non-sensory factor, namely cue validity.