Attention to endogenous and exogenous cues affects auditory localization

Three experiments examine attentional differences in auditory localization using either endogenous or exogenous visual cues. Participants were presented with visual cues on a computer screen and asked to localize auditory targets presented through headphones. In conditions in which the auditory and visual stimuli traveled in the same direction, participants showed illusory directional hearing (Hari in Neurosci Lett 189:29–30, 1995) in that the targets were perceived to travel through the head. In conditions in which the directions of the auditory and visual stimuli were conflicting, participants localized the auditory targets as traveling in the direction of the visual cues. These data suggest that visual capture plays a predominate role in the processes of auditory localization that occurs within the head. Additionally, endogenous response times were significantly greater than exogenous response times. We propose this is the result of additional time required to shift one’s attentional window in the endogenous condition.     

Responses of auditory cortical neurons to pairs of sounds: correlates of fusion and localization

When two brief sounds arrive at a listener's ears nearly simultaneously from different directions, localization of the sounds is described by "the precedence effect." At inter-stimulus delays (ISDs) <5 ms, listeners typically report hearing not two sounds but a single fused sound. The reported location of the fused image depends on the ISD. At ISDs of 1-4 ms, listeners point near the leading source (localization dominance). As the ISD is decreased from 0.8 to 0 ms, the fused image shifts toward a location midway between the two sources (summing localization). When an inter-stimulus level difference (ISLD) is imposed, judgements shift toward the more intense source. Spatial hearing, including the precedence effect, is thought to depend on the auditory cortex. Therefore we tested the hypothesis that the activity of cortical neurons signals the perceived location of fused pairs of sounds. We recorded the unit responses of cortical neurons in areas A1 and A2 of anesthetized cats. Single broadband clicks were presented from various frontal locations. Paired clicks were presented with various ISDs and ISLDs from two loudspeakers located 50 degrees to the left and right of midline. Units typically responded to single clicks or paired clicks with a single burst of spikes. Artificial neural networks were trained to recognize the spike patterns elicited by single clicks from various locations. The trained networks were then used to identify the locations signaled by unit responses to paired clicks. At ISDs of 1-4 ms, unit responses typically signaled locations near that of the leading source in agreement with localization dominance. Nonetheless the responses generally exhibited a substantial undershoot; this finding, too, accorded with psychophysical measurements. As the ISD was decreased from ~0.4 to 0 ms, network estimates typically shifted from the leading location toward the midline in agreement with summing localization. Furthermore a superposed ISLD shifted network estimates toward the more intense source, reaching an asymptote at an ISLD of 15-20 dB. To allow quantitative comparison of our physiological findings to psychophysical results, we performed human psychophysical experiments and made acoustical measurements from the ears of cats and humans. After accounting for the difference in head size between cats and humans, the responses of cortical units usually agreed with the responses of human listeners, although a sizable minority of units defied psychophysical expectations.     

Proprioceptive effects on evoked responses to sounds in the cat auditory cortex

Summary With a view to analyse the influence of neck proprioceptors on directional hearing, evoked potentials (EPs) to dichotically or monaurally presented clicks were recorded from the auditory cortex of cats under deep Nembutal anaesthesia with their head pointing to the front, and then to the right or to the left side at 45°.The change in the head position produced considerable changes in the amplitude of the two primary EP components and in their thresholds. The changes were of two kinds: either decrease or increase of the amplitude. At symmetrical points of the auditory cortex they went in the same direction. The also appeared in the associative zone with the same sign.With monaurally presented clicks, the change of the side of stimulation for the most part resulted in a reversal of the sign of the proprioceptive effect.Similar proprioceptive influences were recorded when the clicks were presented not through earphones but in an open acoustic field.     

The role of spatial disparity and hemifields in audio-visual temporal order judgments

We explored whether sensitivity to audio-visual temporal order judgments (TOJs) was affected by the amount of spatial separation between a sound and light, and by whether the sound and light were presented in the same or in different hemifields. Participants made TOJs about noise bursts and light flashes, and judged whether the stimuli came from the same location or not. Flashes were presented either in the left or right hemifield (at ±10° from central fixation), and sounds either came from the same location as the lights, or at small or large disparities (20 or 40° from the light, respectively), thereby crossing the hemifields or not. TOJs became more accurate (i.e., the just noticeable difference, JND, became smaller) when spatial disparity increased and when hemifields were crossed. Location discrimination of the sound and light was affected similarly. These results demonstrate that audio-visual TOJs are critically dependent on both the relative position from which stimuli are presented and on whether stimuli cross hemifields or not.     

Vision contingent auditory pitch aftereffects

Visual motion aftereffects can occur contingent on arbitrary sounds. Two circles, placed side by side, were alternately presented, and the onsets were accompanied by tone bursts of high and low frequencies, respectively. After a few minutes of exposure to the visual apparent motion with the tones, a circle blinking at a fixed location was perceived as a lateral motion in the same direction as the previously exposed apparent motion (Teramoto et al. in PLoS One 5:e12255, 2010). In the present study, we attempted to reverse this contingency (pitch aftereffects contingent on visual information). Results showed that after prolonged exposure to the audio-visual stimuli, the apparent visual motion systematically affected the perceived pitch of the auditory stimuli. When the leftward apparent visual motion was paired with the high–low-frequency sequence during the adaptation phase, a test tone sequence was more frequently perceived as a high–low-pitch sequence when the leftward apparent visual motion was presented and vice versa. Furthermore, the effect was specific for the exposed visual field and did not transfer to the other side, thus ruling out an explanation in terms of simple response bias. These results suggest that new audiovisual associations can be established within a short time, and visual information processing and auditory processing can mutually influence each other.     

Functional properties of neurons in the temporo-parietal association cortex of awake monkey

Summary The temporo-parietal association cortex around the caudal end of the Sylvian fissure was studied with the single cell recording technique in three awake behaving Macaca speciosa-monkeys. Of the 197 cells isolated, 5% were active only during the monkey's own movements, mostly during head rotation, and 95% were responsive to sensory stimulation: 54% to auditory stimuli, 24% to somatosensory stimuli, 13% to both of these and 4% to visual stimuli. Some cells, classified as responsive to somatosensory stimuli, were activated only by passive rotation of the head on the cervical axis; it is possible that they were driven by vestibular stimuli. Half of the cells were activated by stimuli on both sides of the monkey, and almost all the rest, only by stimuli on the side contralateral to the hemisphere recorded.Of the acoustically drivable cells, 95% responded to natural sounds, such as, rubbing hands together, rustle of clothes, clicks or jingles (sounds with noise spectrum and rapid intensity transitions). Most of these neurons were also examined with pure tones of 0.2–20 kHz: various inhibitory or excitatory responses were elicited in half of them, usually over a wide range of frequencies. The responses of most acoustically drivable cells (62%) depended on the location of the sound source with reference to the monkey's head so that the maximal response was elicited by sounds with a certain angle of incidence, usually on the contralateral side.The present results suggest that the area studied participates in the analysis of the temporal pattern of a sound, the location of the sound source and in spatial control of head movements.     

Representation of interaural time delay in the human auditory midbrain

Interaural time difference (ITD) is a critical cue to sound-source localization. Traditional models assume that sounds leading at one ear, and perceived on that side, are processed in the opposite midbrain. Using functional magnetic resonance imaging we demonstrate that as the ITDs of sounds increase, midbrain activity can switch sides, even though perceived location remains on the same side. The data require a new model for human ITD processing.     

Auditory deprivation affects biases of visuospatial attention as measured by line bisection

In this study, we investigated whether early deafness affects the typical pattern of hemispheric lateralization [i.e., right hemisphere (RH) dominance] in the control of spatial attention. To this aim, deaf signers, deaf non-signers, hearing signers, and hearing non-signers were required to bisect a series of centrally presented visual lines. The directional bisection bias was found to be significantly different between hearing and deaf participants, irrespective of sign language use. Hearing participants (both signers and non-signers) showed a consistent leftward bias, reflecting RH dominance. Conversely, we observed no evidence of a clear directional bias in deaf signers or non-signers (deaf participants overall showing a non-significant tendency to deviate rightward), suggesting that deafness may be associated to a more bilateral hemispheric engagement in visuospatial tasks.     

Intermittent contralateral and ipsilateral hemiretinal stimulation and its effect on the phasic stretch reflex

The effect of visual stimulation on the phasic stretch reflex, measured isometrically by way of EMG recordings, was investigated with normal human subjects. Three experimental treatments were applied, namely 3 flashes of light presented as hemiretinal contralateral stimulation prior to tendon taps; similar hemiretinal stimulation presented ipsilaterally; and no flashes of light as a control condition. The analysis of variance showed that the MAP means and standard deviations for both of the lights conditions were significantly greater than for the control condition. Eleven out of 12 subjects showed a larger reflex response to stimulation of the brain ipsilateral to the tendon stimulated. The findings are discussed in terms of visual projections differentially affecting alpha and gamma motoneurons.     

Temporal ventriloquism: crossmodal interaction on the time dimension. 1. Evidence from auditory-visual temporal order judgment.

In the well-known visual bias of auditory location (alias the ventriloquist effect), auditory and visual events presented in separate locations appear closer together, provided the presentations are synchronized. Here, we consider the possibility of the converse phenomenon: crossmodal attraction on the time dimension conditional on spatial proximity. Participants judged the order of occurrence of sound bursts and light flashes, respectively, separated in time by varying stimulus onset asynchronies (SOAs) and delivered either in the same or in different locations. Presentation was organized using randomly mixed psychophysical staircases, by which the SOA was reduced progressively until a point of uncertainty was reached. This point was reached at longer SOAs with the sounds in the same frontal location as the flashes than in different places, showing that apparent temporal separation is effectively longer in the first condition. Together with a similar one obtained recently in a case of tactile-visual discrepancy, this result supports a view in which timing and spatial layout of the inputs play to some extent inter-changeable roles in the pairing operation at the base of crossmodal interaction.     

Visual acceleration and spatial distortion in right brain-damaged patients

A subset of right brain-damaged patients shows leftward overextension in the line extension task. It has been argued that this deficit can be attributed to a distortion of the metric structure of perceived space (spatial anisometry). We investigated whether spatial distortion of static stimuli is associated with a corresponding misperception of perceived acceleration of moving stimuli. Seven right brain-damaged patients with spatial anisometry and two control groups were presented with stimuli moving leftwards or rightwards along the horizontal axis at different rates of acceleration. They were asked to estimate whether the target accelerated or decelerated. The anisometric group judged the perceived acceleration of leftward motions as less than that of rightward motions. The magnitude of the misperception of acceleration correlated positively with relative left overextension in the line extension task and with rightward displacement error in the line bisection task. This directional difference is in line with the predictions of the spatial anisometry hypothesis.     

Tinnitus and hearing loss in hamsters (Mesocricetus auratus) exposed to loud sound

Hamsters were trained to go left and right to sounds on their left and right sides, respectively. Silent trials were occasionally given in which no sound was presented. Hamsters exposed to a loud 2- or 10-kHz tone in 1 ear often shifted their responding on the silent trials to the side of the exposed ear, suggesting that they perceived a sound in that ear (i.e., tinnitus). The degree of tinnitus was related to the degree of the accompanying hearing loss (estimated by the auditory brainstem response). However, a conductive hearing loss (plugging 1 ear) did not cause a hamster to test positive for tinnitus. Tinnitus could be demonstrated within minutes following tone exposure, indicating an immediate onset, as occurs in humans.     

Sound-Induced Flash Illusion is Resistant to Feedback Training

A single flash accompanied by two auditory beeps tends to be perceived as two flashes (Shams et al. Nature 408:788, 2000, Cogn Brain Res 14:147–152, 2002). This phenomenon is known as ‘sound-induced flash illusion.’ Previous neuroimaging studies have shown that this illusion is correlated with modulation of activity in early visual cortical areas (Arden et al. Vision Res 43(23):2469–2478, 2003; Bhattacharya et al. NeuroReport 13:1727–1730, 2002; Shams et al. NeuroReport 12(17):3849–3852, 2001, Neurosci Lett 378(2):76–81, 2005; Watkins et al. Neuroimage 31:1247–1256, 2006, Neuroimage 37:572–578, 2007; Mishra et al. J Neurosci 27(15):4120–4131, 2007). We examined how robust the illusion is by testing whether the frequency of the illusion can be reduced by providing feedback. We found that the sound-induced flash illusion was resistant to feedback training, except when the amount of monetary reward was made dependent on accuracy in performance. However, even in the latter case the participants reported that they still perceived illusory two flashes even though they correctly reported single flash. Moreover, the feedback training effect seemed to disappear once the participants were no longer provided with feedback suggesting a short-lived refinement of discrimination between illusory and physical double flashes rather than vanishing of the illusory percept. These findings indicate that the effect of sound on the perceptual representation of visual stimuli is strong and robust to feedback training, and provide further evidence against decision factors accounting for the sound-induced flash illusion.     

Click evoked neurogenic vestibular potentials (NVESTEPs): a method of assessing the function of the vestibular system

OBJECTIVES: To obtain neurogenic vestibular evoked potentials (NVESTEPs) with surface scalp recording using high intensity auditory clicks. The same stimulus is used in myogenic vestibular evoked potentials which has been shown to evoke potentials in the vestibular division of the vestibulocochlear nerve. METHODS: A whole head recording with surface EEG electrodes was performed using high intensity clicks in one normal volunteer to determine the best recording position for vestibular evoked potentials. The results were compared to responses at moderate click intensities used for brainstem auditory evoked potentials (BAEPs). The difference in the location of the two responses on the scalp was assumed to be from the vestibular system. RESULTS: Responses specific to the high intensity clicks were best obtained in the parietal areas, with no reproducible responses obtained in the same area with moderate intensity clicks normally used in BAEPs. Recordings in neurologically normal volunteers showed a consistent response with a negative polarity at around 3 ms, which we therefore called N3. Two case studies are presented. The first case is a patient with unilateral sensorineural hearing loss with NVESTEPs present, suggesting that NVESTEPs is not a cochlear response. The second case is a patient with multiple sclerosis with demyelinating lesions in the pons and an unobtainable NVESTEP response. CONCLUSION: NVESTEPs is a possible new diagnostic technique that may be specific for the vestibular pathway. It has potential use in patients with symptoms of dizziness, subclinical symptoms in multiple sclerosis, and in disorders specific for the vestibular nerve.     

Audiotactile interactions beyond the space and body parts around the head

Recent research has reported that spatial modulation effects of audiotactile interactions tend to be limited to the space and body parts around the head. The present study investigated the generality of this finding by manipulating body parts stimulated and spatial relationships between the body parts and sounds. In Experiment 1, tactile stimuli were presented randomly to either left or right cheek, hand (palm or back) placed near the head, and knee while auditory stimuli were presented to either the same or opposite side from loudspeakers close to the head. Participants made speeded spatial discrimination responses regarding the side (left versus right) of the tactile stimulation. For any body part stimulated, the performance was worse when the auditory stimuli were presented from the opposite side rather than from the same side. Experiment 2 demonstrated that the spatial modulation effects for the palm or the back of the hand occurred irrespective of hand position (near or far from the head) and sound position (near or far from the head). The sounds delivered from near the head exerted a greater influence on tactile spatial discrimination performance as compared with the sound delivered from far from the head. Furthermore, the back of the hand was more influenced by the auditory stimuli than the palm when the hands were placed near the sounds. These results suggest that the spatial modulation effects of audiotactile interactions can occur beyond the space and body surface around the head.     

Perceived timing of vestibular stimulation relative to touch,light and sound

Different senses have different processing times. Here we measured the perceived timing of galvanic vestibular stimulation (GVS) relative to tactile, visual and auditory stimuli. Simple reaction times for perceived head movement (438 ± 49 ms) were significantly longer than to touches (245 ± 14 ms), lights (220 ± 13 ms), or sounds (197 ± 13 ms). Temporal order and simultaneity judgments both indicated that GVS had to occur about 160 ms before other stimuli to be perceived as simultaneous with them. This lead was significantly less than the relative timing predicted by reaction time differences compatible with an incomplete tendency to compensate for differences in processing times.     

Distortions of auditory space during rapid head turns

Auditory localisation was examined using brief broadband sounds presented during rapid head turns to visual targets in the peripheral field. Presenting sounds during a rapid head movement will “smear” the acoustic cues to the sound’s location. During the early stages of a head turn, sound localisation accuracy was comparable to a no-turn control condition. However, significant localisation errors occurred when the probe sound was presented during the later part of a head turn. After correcting for head position, the estimate of lateral angle (horizontal position) in the front hemisphere was generally accurate. However, lateral angle estimates for positions in the rear hemisphere exhibited systematic errors that were especially large around the midline. Polar angle (elevation) perception remained robust, being comparable to no-turn controls whether tested early or late in the head turn. The results are interpreted in terms of a ‘multiple look’ strategy for calculating sound location, and the allocation of attention to the hemisphere containing the head-turn target.     

No effect of auditory–visual spatial disparity on temporal recalibration

It is known that the brain adaptively recalibrates itself to small (∼100 ms) auditory–visual (AV) temporal asynchronies so as to maintain intersensory temporal coherence. Here we explored whether spatial disparity between a sound and light affects AV temporal recalibration. Participants were exposed to a train of asynchronous AV stimulus pairs (sound-first or light-first) with sounds and lights emanating from either the same or a different location. Following a short exposure phase, participants were tested on an AV temporal order judgement (TOJ) task. Temporal recalibration manifested itself as a shift of subjective simultaneity in the direction of the adapted audiovisual lag. The shift was equally big when exposure and test stimuli were presented from the same or different locations. These results provide strong evidence for the idea that spatial co-localisation is not a necessary constraint for intersensory pairing to occur.     

Distortions of perceived auditory and visual space following adaptation to motion

Adaptation to visual motion can induce marked distortions of the perceived spatial location of subsequently viewed stationary objects. These positional shifts are direction specific and exhibit tuning for the speed of the adapting stimulus. In this study, we sought to establish whether comparable motion-induced distortions of space can be induced in the auditory domain. Using individually measured head related transfer functions (HRTFs) we created auditory stimuli that moved either leftward or rightward in the horizontal plane. Participants adapted to unidirectional auditory motion presented at a range of speeds and then judged the spatial location of a brief stationary test stimulus. All participants displayed direction-dependent and speed-tuned shifts in perceived auditory position relative to a ‘no adaptation’ baseline measure. To permit direct comparison between effects in different sensory domains, measurements of visual motion-induced distortions of perceived position were also made using stimuli equated in positional sensitivity for each participant. Both the overall magnitude of the observed positional shifts, and the nature of their tuning with respect to adaptor speed were similar in each case. A third experiment was carried out where participants adapted to visual motion prior to making auditory position judgements. Similar to the previous experiments, shifts in the direction opposite to that of the adapting motion were observed. These results add to a growing body of evidence suggesting that the neural mechanisms that encode visual and auditory motion are more similar than previously thought.     

Elderly hearing-impaired persons’ coping behavior

Little is known about the coping behavior associated with hearing impairment. In this study, 24 elderly hearing-impaired participants participated in a structured video interview including communication faults (provocations) presented in association with sounds. The participants also completed measures of self-perceived hearing handicap, optimism, and depression. Several behaviors were coded independently by two observers using a rating scale that reached average Cohen's kappa of .75. A more conservative estimate resulted in a kappa of .50. Results showed that the provocations affected behavior differently. Especially problematic for the participants was a provocation in which the interviewer talked with his hand in front of his mouth. With one exception, no gender differences were found. Few significant relations between the assessed behaviors and self-assessed handicap, optimism, and depression were found. Hearing measurement (pure tone average and speech recognition) were not clearly related with the coping behaviors assessed. Behavioral observation techniques designed for the elderly hearing impaired should be further studied.