The auditory motion aftereffect: its tuning and specificity in the spatial and frequency domains

In this paper, the auditory motion aftereffect (aMAE) was studied, using real moving sound as both the adapting and the test stimulus. The sound was generated by a loudspeaker mounted on a robot arm that was able to move quietly in three-dimensional space. A total of 7 subjects with normal hearing were tested in three experiments. The results from Experiment 1 showed a robust and reliable negative aMAE in all the subjects. After listening to a sound source moving repeatedly to the right, a stationary sound source was perceived to move to the left. The magnitude of the aMAE tended to increase with adapting velocity up to the highest velocity tested (20 degrees/sec). The aftereffect was largest when the adapting and the test stimuli had similar spatial location and frequency content. Offsetting the locations of the adapting and the test stimuli by 20 degrees reduced the size of the effect by about 50%. A similar decline occurred when the frequency of the adapting and the test stimuli differed by one octave. Our results suggest that the human auditory system possesses specialized mechanisms for detecting auditory motion in the spatial domain.     

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.     

Apparent velocity of motion aftereffects in central and peripheral vision

Adapting to a drifting grating (temporal frequency 4 Hz, contrast 0.4) in the periphery gave rise to a motion aftereffect (MAE) when the grating was stopped. A standard unadapted foveal grating was matched to the apparent velocity of the MAE, and the matching velocity was approximately constant regardless of the visual field position and spatial frequency of the adapting grating. On the other hand, when the MAE was measured by nulling with real motion of the test grating, nulling velocity was found to increase with eccentricity. The nulling velocity was constant when scaled to compensate for changes in the spatial 'grain' of the visual field. Thus apparent velocity of MAE is constant across the visual field, but requires a greater velocity of real motion to cancel it in the periphery. This confirms that the mechanism underlying MAE is spatially-scaled with eccentricity, but temporally homogeneous. A further indication of temporal homogeneity is that when MAE is tracked, by matching or by nulling, the time course of temporal decay of the aftereffect is similar for central and for peripheral stimuli.     

Effect of luminance contrast on the motion aftereffect

The effects of luminance contrast and spatial frequency on the motion aftereffect were investigated. The point of subjective equality for velocity was measured as an index of the motion aftereffect. The largest effect was observed when a low contrast grating (5%) was presented as a test stimulus after adaptation to a high contrast grating (100%) in the low spatial frequency condition (0.8 cycle deg.-1). On the whole, the effect increased with increasing adapting contrast and with decreasing test contrast or spatial frequency. Small effects were observed at high test contrasts. These results were inconsistent with those of Keck, Palella, and Pantle in 1976. Analysis showed that there was no saturation on velocity of the motion aftereffect above 5% of the contrast although Keck, et al. (1976) found that the incremental increases of the effect above 3% adapting contrast were small.     

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.     

Adaptation to spiral motion in crowding condition

When a single, moving stimulus is presented in the peripheral visual field, its direction of motion can be easily distinguished, but when the same stimulus is flanked by other similar moving stimuli, observers are unable to report its direction of motion. In this condition, known as 'crowding', specific features of visual stimuli do not access conscious perception. The aim of this study was to investigate whether adaptation to spiral motion is preserved in crowding conditions. Logarithmic spirals were used as adapting stimuli. A rotating spiral stimulus (target spiral) was presented, flanked by spirals of the same type, and observers were adapted to its motion. The observers' task was to report the rotational direction of a directionally ambiguous motion (test stimulus) presented afterwards. The directionally ambiguous motion consisted of a pair of spirals flickering in counterphase, which were mirror images of the target spiral. Although observers were not aware of the rotational direction of the target and identified it at chance levels, the direction of rotation reported by the observers during the test phase (motion aftereffect) was contrarotational to the direction of the adapting spiral. Since all contours of the adapting and test stimuli were 90 degrees apart, local motion detectors tuned to the directions of the mirror-image spiral should fail to respond, and therefore not adapt to the adapting spiral. Thus, any motion aftereffect observed should be attributed to adaptation of global motion detectors (ie rotation detectors). Hence, activation of rotation-selective cells is not necessarily correlated with conscious perception.     

Motion aftereffects with horizontally moving sound sources in the free field

A horizontally moving sound was presented to an observer seated in the center of an anechoic chamber. The sound, either a 500-Hz low-pass noise or a 6300-Hz high-pass noise, repeatedly traversed a semicircular arc in the observer's front hemifield at ear level (distance: 1.5 m). At 10-sec intervals this adaptor was interrupted, and a 750-msec moving probe (a 500-Hz low-pass noise) was presented from a horizontal arc 1.6 m in front of the observer. During a run, the adaptor was presented at a constant velocity (-200 degrees to +200 degrees/sec), while probes with velocities varying from -10 degrees to +10 degrees/sec were presented in a random order. Observers judged the direction of motion (left or right) of each probe. As in the case of stimuli presented over headphones (Grantham & Wightman, 1979), an auditory motion aftereffect (MAE) occurred: subjects responded "left" to probes more often when the adaptor moved right than when it moved left. When the adaptor and probe were spectrally the same, the MAE was greater than when they were from different spectral regions; the magnitude of this difference depended on adaptor speed and was subject-dependent. It is proposed that there are two components underlying the auditory MAE: (1) a generalized bias to respond that probes move in the direction opposite to that of the adaptor, independent of their spectra; and (2) a loss of sensitivity to the velocity of moving sounds after prolonged exposure to moving sounds having the same spectral content.     

Auditory motion aftereffects

Observers were adapted to simulated auditory movement produced by dynamically varying the interaural time and intensity differences of tones (500 or 2,000 Hz) presented through headphones. At lO-sec intervals during adaptation, various probe tones were presented for 1 sec (the frequency of the probe was always the same as that of the adaptation stimulus). Observers judged the direction of apparent movement (“left” or “right”) of each probe tone. At 500 Hz, with a 200-deg/sec adaptation velocity, “stationary” probe tones were consistently judged to move in the direction opposite to that of the adaptation stimulus. We call this result an auditory motion aftereffect. In slower velocity adaptation conditions, progressively less aftereffect was demonstrated. In the higher frequency condition (2,000 Hz, 200-deg/sec adaptation velocity), we found no evidence of motion aftereffect. The data are discussed in relation to the well-known visual analog-the “waterfall effect.” Although the auditory aftereffect is weaker than the visual analog, the data suggest that auditory motion perception might be mediated, as is generally believed for the visual system, by direction-specific movement analyzers.     

Changing-loudness aftereffect following simulated movement: implications for channel hypotheses concerning sound level change and movement.

Listening to a tone changing unidirectionally in sound level causes an illusion of changing loudness in a steady tone afterward. This aftereffect may indicate channels for detecting the feature of change in sound level, which would primarily concern dynamic sound localization. Three subjects, one of whom was the author, participated in this study. The author predicted that opposite adaptation of the ears (the adapting stimulus is heard to move from one ear to the other) should lead to a movement aftereffect. This was not reported by the subjects. However, the subjects did report a changing-loudness aftereffect in a monaural test stimulus, and the characteristics of the changing-loudness aftereffect (such as its magnitude) were consistent with previous data, suggesting a two-stage channel hypothesis: Output from channels for several features, including sound-level change, simultaneously stimulate movement channels.     

Localization of moving sound

The final position of a moving sound source usually appears to be displaced in the direction of motion. We tested the hypothesis that this phenomenon, termed auditory representational momentum, is already emerging during, not merely after, the period of motion. For this purpose, we investigated the localization of a moving sound at different points in time. In a dark anechoic environment, an acoustic target moved along the frontal horizontal plane. In the initial, middle, or final phase of the motion trajectory, subjects received a tactile stimulus and determined the current position of the moving target at the moment of the stimulus by performing either relative-judgment or pointing tasks. Generally, in the initial phase of the auditory motion, the position was perceived to be displaced in the direction of motion, but this forward displacement disappeared in the further course of the motion. When the motion stimulus had ceased, however, its final position was again shifted in the direction of motion. The latter result suggests that representational momentum in spatial hearing is a phenomenon specific to the final point of motion. Mental extrapolation of past trajectory information is discussed as a potential source of this perceptual displacement.     

Localization of moving sound

The final position of a moving sound source usually appears to be displaced in the direction of motion. We tested the hypothesis that this phenomenon, termed auditory representational momentum, is already emerging during, not merely after, the period of motion. For this purpose, we investigated the localization of a moving sound at different points in time. In a dark anechoic environment, an acoustic target moved along the frontal horizontal plane. In the initial, middle, or final phase of the motion trajectory, subjects received a tactile stimulus and determined the current position of the moving target at the moment of the stimulus by performing either relative-judgment or pointing tasks. Generally, in the initial phase of the auditory motion, the position was perceived to be displaced in the direction of motion, but this forward displacement disappeared in the further course of the motion. When the motion stimulus had ceased, however, its final position was again shifted in the direction of motion. The latter result suggests that representational momentum in spatial hearing is a phenomenon specific to the final point of motion. Mental extrapolation of past trajectory information is discussed as a potential source of this perceptual displacement.     

Cross-modal training induces changes in spatial representations early in the auditory processing pathway

In the ventriloquism aftereffect, brief exposure to a consistent spatial disparity between auditory and visual stimuli leads to a subsequent shift in subjective sound localization toward the positions of the visual stimuli. Such rapid adaptive changes probably play an important role in maintaining the coherence of spatial representations across the various sensory systems. In the research reported here, we used event-related potentials (ERPs) to identify the stage in the auditory processing stream that is modulated by audiovisual discrepancy training. Both before and after exposure to synchronous audiovisual stimuli that had a constant spatial disparity of 15°, participants reported the perceived location of brief auditory stimuli that were presented from central and lateral locations. In conjunction with a sound localization shift in the direction of the visual stimuli (the behavioral ventriloquism aftereffect), auditory ERPs as early as 100 ms poststimulus (N100) were systematically modulated by the disparity training. These results suggest that cross-modal learning was mediated by a relatively early stage in the auditory cortical processing stream.     

Contrast and frequency competition for orientation-contingent color aftereffects

A “competition” paradigm was developed to examine separately the effects of pattern contrast and spatial frequency characteristics on the strength of orientation-contingent color aftereffects (McCollough effects). After adapting to alternately presented red/black and green/black square-wave gratings (one horizontal, one vertical), 11 subjects viewed seven different kinds of test patterns. Unlike Standard McCollough effect test stimuli, the present patterns had variable luminance profiles running both horizontally and vertically within each test pattern area. Forced choice responses were used to determine which aftereffect color (red or green) appeared, as characteristics of vertical and horizontal luminance profiles were varied separately among test stimulus types. We conclude that pattern contrast and human contrast sensitivity account for aftereffect colors in such stimuli. When contrast is taken into consideration, aftereffects are not predicted by similarity between adaptation and test pattern Fourier characteristics, nor are they predicted by the width, per se, of pattern elements.     

Retinal and extraretinal information in movement perception: how to invert the Filehne illusion

During a pursuit eye movement made in darkness across a small stationary stimulus, the stimulus is perceived as moving in the opposite direction to the eyes. This so-called Filehne illusion is usually explained by assuming that during pursuit eye movements the extraretinal signal (which informs the visual system about eye velocity so that retinal image motion can be interpreted) falls short. A study is reported in which the concept of an extraretinal signal is replaced by the concept of a reference signal, which serves to inform the visual system about the velocity of the retinae in space. Reference signals are evoked in response to eye movements, but also in response to any stimulation that may yield a sensation of self-motion, because during self-motion the retinae also move in space. Optokinetic stimulation should therefore affect reference signal size. To test this prediction the Filehne illusion was investigated with stimuli of different optokinetic potentials. As predicted, with briefly presented stimuli (no optokinetic potential) the usual illusion always occurred. With longer stimulus presentation times the magnitude of the illusion was reduced when the spatial frequency of the stimulus was reduced (increased optokinetic potential). At very low spatial frequencies (strongest optokinetic potential) the illusion was inverted. The significance of the conclusion, that reference signal size increases with increasing optokinetic stimulus potential, is discussed. It appears to explain many visual illusions, such as the movement aftereffect and center-surround induced motion, and it may bridge the gap between direct Gibsonian and indirect inferential theories of motion perception.     

A comparison of visual and auditory representational momentum in spatial tasks

Similarities have been observed in the localization of the final position of moving visual and moving auditory stimuli: Perceived endpoints that are judged to be farther in the direction of motion in both modalities likely reflect extrapolation of the trajectory, mediated by predictive mechanisms at higher cognitive levels. However, actual comparisons of the magnitudes of displacement between visual tasks and auditory tasks using the same experimental setup are rare. As such, the purpose of the present free-field study was to investigate the influences of the spatial location of motion offset, stimulus velocity, and motion direction on the localization of the final positions of moving auditory stimuli (Experiment 1 and 2) and moving visual stimuli (Experiment 3). To assess whether auditory performance is affected by dynamically changing binaural cues that are used for the localization of moving auditory stimuli (interaural time differences for low-frequency sounds and interaural intensity differences for high-frequency sounds), two distinct noise bands were employed in Experiments 1 and 2. In all three experiments, less precise encoding of spatial coordinates in paralateral space resulted in larger forward displacements, but this effect was drowned out by the underestimation of target eccentricity in the extreme periphery. Furthermore, our results revealed clear differences between visual and auditory tasks. Displacements in the visual task were dependent on velocity and the spatial location of the final position, but an additional influence of motion direction was observed in the auditory tasks. Together, these findings indicate that the modality-specific processing of motion parameters affects the extrapolation of the trajectory.     

Rapid adaptation to auditory-visual spatial disparity

The so-called ventriloquism aftereffect is a remarkable example of rapid adaptative changes in spatial localization caused by visual stimuli. After exposure to a consistent spatial disparity of auditory and visual stimuli, localization of sound sources is systematically shifted to correct for the deviation of the sound from visual positions during the previous adaptation period. In the present study, this aftereffect was induced by presenting, within 17 min, 1800 repetitive noise or pure-tone bursts in combination with synchronized, and 20° disparate flashing light spots, in total darkness. Post-adaptive sound localization, measured by a method of manual pointing, was significantly shifted 2.4° (noise), 3.1° (1 kHz tones), or 5.8° (4 kHz tones) compared with the pre-adaptation condition. There was no transfer across frequencies; that is, shifts in localization were insignificant when the frequencies used for adaptation and the post-adaptation localization test were different. It is hypothesized that these aftereffects may rely on shifts in neural representations of auditory space with respect to those of visual space, induced by intersensory spatial disparity, and may thus reflect a phenomenon of neural short-term plasticity.     

Perceptual asymmetries associated with changing-loudness aftereffects

Listening to decreasing sound level leads to an increasing-loudness aftereffect, whereas listening to increasing sound level leads to a decreasing-loudness aftereffect. Measuring the aftereffects by nulling them in short test stimuli reveals that increasing-loudness aftereffects are greater than decreasing-loudness aftereffects. However, this perceptual asymmetry may be due to another illusion--the growing-louder effect: In the absence of any adaptation, short steady stimuli are heard as growing louder. In an experiment in which the duration of test stimuli varied from 1.0 to 2.5 sec, the growing-louder effect did not occur in the longer test stimuli, but the asymmetry in changing-loudness aftereffects remained. The aftereffect asymmetry is therefore independent of the growing-louder effect. The aftereffect asymmetry is consistent with other psychophysical and physiological evidence that is believed to concern potential collision: An approaching sound-source elicits increasing sound level. In addition, the aftereffect asymmetry parallels a well-known asymmetry regarding aftereffects of visual motion, which is also attributed to potential collision.     

Perceptual load influences auditory space perception in the ventriloquist aftereffect

A period of exposure to trains of simultaneous but spatially offset auditory and visual stimuli can induce a temporary shift in the perception of sound location. This phenomenon, known as the ‘ventriloquist aftereffect’, reflects a realignment of auditory and visual spatial representations such that they approach perceptual alignment despite their physical spatial discordance. Such dynamic changes to sensory representations are likely to underlie the brain’s ability to accommodate inter-sensory discordance produced by sensory errors (particularly in sound localization) and variability in sensory transduction. It is currently unknown, however, whether these plastic changes induced by adaptation to spatially disparate inputs occurs automatically or whether they are dependent on selectively attending to the visual or auditory stimuli. Here, we demonstrate that robust auditory spatial aftereffects can be induced even in the presence of a competing visual stimulus. Importantly, we found that when attention is directed to the competing stimuli, the pattern of aftereffects is altered. These results indicate that attention can modulate the ventriloquist aftereffect.     

Visual motion influences the contingent auditory motion aftereffect

In this study, we show that the contingent auditory motion aftereffect is strongly influenced by visual motion information. During an induction phase, participants listened to rightward-moving sounds with falling pitch alternated with leftward-moving sounds with rising pitch (or vice versa). Auditory aftereffects (i.e., a shift in the psychometric function for unimodal auditory motion perception) were bigger when a visual stimulus moved in the same direction as the sound than when no visual stimulus was presented. When the visual stimulus moved in the opposite direction, aftereffects were reversed and thus became contingent upon visual motion. When visual motion was combined with a stationary sound, no aftereffect was observed. These findings indicate that there are strong perceptual links between the visual and auditory motion-processing systems.     

Aftereffect arising from unidirectional change of sound level in a tone: frequency specificity with a complex adaptor

Subjects adapted to square-wave adaptors, stimuli containing odd harmonics of the fundamental, and, to provide baseline data, sinusoidal adaptors matching the square-wave's fundamental. Nulls were obtained for various frequencies of the test stimulus. At any given frequency, the baseline null was subtracted from the null for square-wave adaptation. These corrected nulls indicated frequency-specific aftereffects at the third and fifth harmonics. The evidence is consistent with previous attempts to link the aftereffect of changing sound level in a tone with the auditory movement aftereffect because the latter may also show frequency specificity.