The spectrally dependent monotic component in the decreasing-loudness aftereffect: implications for dynamic auditory localization

Listeners exposed to a tone increasing in intensity report an aftereffect of decreasing loudness in a steady tone heard afterward. In the present study, the spectral dependence of the monotic decreasing-loudness aftereffect (adapting and testing 1 ear) was compared with (a) the spectral dependence of the interotic decreasing-loudness aftereffect (adapting 1 ear and testing the other ear) and (b) a non-adaptation control condition. The purpose was to test the hypothesis that the decreasing-loudness aftereffect may concern the sensory processing associated with dynamic localization. The hypothesis is based on two premises: (a) dynamic localization requires monaural sensory processing, and (b) sensory processing is reflected in spectral selectivity. Hence, the hypothesis would be supported if the monotic aftereffect were more spectrally dependent and stronger than the interotic aftereffect; A. H. Reinhardt-Rutland (1998) showed that the hypothesis is supported with regard to the related increasing-loudness aftereffect. Two listeners were exposed to a 1-kHz adapting stimulus. From responses of "growing softer" or "growing louder" to test stimuli changing in intensity, nulls were calculated; test carrier frequencies ranged from 0.5 kHz to 2 kHz. Confirming the hypothesis, the monotic aftereffect peaked at around the 1-kHz test carrier frequency. In contrast, the interotic aftereffect showed little evidence of spectrally dependent peaking. Except when test and adaptation carrier frequencies differed markedly, the interotic aftereffect was smaller than the monotic aftereffect.     

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.     

Changing-loudness aftereffects: Slope of response functions and spectral dependence

Aftereffects of azimuthal auditory motion may have two components. A sensory component is inferred from strong aftereffects, because they are spectrally dependent and have shallower response functions than those for non-adaptation. Neither property applies to weak aftereffects, suggesting a cognitive component. Two experiments determined whether changing-loudness aftereffects (CLA) might be understood similarly. In a single-interval forced-choice procedure, listeners responded growing softer or growing louder to test stimuli changing in intensity. In Exp. 1, adapting and test stimuli were diotic and had the same 1-kHz sinusoidal carrier. Although response functions following adaptation were displaced from response functions for non-adaptation — indicating CLA — their slopes were broadly similar. In Exp. 2, stimuli were monotic; adapting frequency was 1 kHz and test frequencies were between 0.5 and 2.0 kHz. CLA was present in most adaptation conditions, but was strongest when the test frequency was 1.0 kHz; functions' slopes again evinced no systematic variation. The two-component hypothesis for CLA is supported by spectral dependence alone. It is argued that the slope of response functions is due to the nulling procedures for measuring auditory aftereffects. The slope depends on whether the adapted property is processed by direct and indirect mechanisms; aftereffects tap direct mechanisms alone, which may affect sensitivity during measurement.     

Top-down gain control in the auditory system: evidence from identification and discrimination experiments

The influence of intensity range in auditory identification and intensity discrimination experiments is well documented and is usually attributed to nonsensory factors. Recent studies, however, have suggested that the stimulus range effect might be sensory in origin. To test this notion, in one set of experiments, we had listeners identify the individual tones in a set. One baseline condition consisted of identifying four 1-kHz, low-intensity tones; the other consisted of identifying four 1-kHz, high-intensity tones. In the experimental conditions, these baseline tone sets were augmented by adding a fifth tone at either 1 or 5 kHz. Added 5-kHz tones had little effect on identification accuracy for the four baseline tones. When an added 1-kHz tone differed substantially in intensity from the four baseline tones, it adversely affected performance, with the addition of a high-intensity tone to a set of low-intensity tones having a more deleterious effect than the addition of a low-intensity tone to a set of high-intensity tones. These and further results, obtained in an exploration of this asymmetrical range effect in a third identification experiment and in two intensity-discrimination experiments, were consistent with the notion of a nonlinear amplifier under top-down control whose functions include protection against sensory overload from loud sounds. The identification data were well described by a signal-detection model using equal-variance Laplace distributions instead of the usual Gaussian distributions.     

Auditory sensitivity in school-age children

Thresholds for octave-band noises with center frequencies of 0.4, 1, 2, 4, and 10 kHz and 1/3-octave-band noises centered at 10 and 20 kHz were obtained from children 6 to 16 years of age. Such thresholds, combined with those obtained previously for infants, preschool children, and adults, provide a detailed picture of developing auditory sensitivity between infancy and maturity. Continuing improvements in sensitivity are evident from infancy through the preschool period, well into the school years. For stimuli with center frequencies of 0.4 and 1 kHz, maximal sensitivity is achieved at about 10 years of age, compared to 8 years for stimuli of 2 and 4 kHz. For 10-kHz stimuli, there is little change beyond 4 or 5 years of age. Finally, 20-kHz stimuli yield maximal sensitivity at about 6 or 8 years of age, followed by a progressive decline to adult levels. These findings are considered in relation to auditory sensitivity in nonhuman species, to structural and functional development of the ear, and to possible changes in the efficiency of neural processing.     

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.     

The additivity of loudness across critical bands: A conjoint measurement approach

Five subjects were required in each trial to compare directly two sounds and to indicate which sound was louder. Each of the 64 sounds employed consisted of a combination of one of eight intensity levels of a 2-kHz tone and one of eight intensities of a 5-kHz tone. If, as Fletcher and Munson (1933) argued, loudness is additive for tone combinations in which the frequencies are widely separated, then subjects’ judgments should reflect the summed loudnesses of the 2- and 5-kHz tones in a two-tone combination. Judgments of individual subjects were shown to satisfy the conditions for an additive structure, and individual loudness scales were constructed. These loudness scales varied from subject to subject. Since this paired comparison procedure minimized response biases, the results suggest substantial individual differences in the sensory representation of sound intensity. The relations among sensory scales derived from other structured sensory judgments, such as binaural loudness, are discussed.     

CS-specific gamma,theta, and alpha EEG activity detected in stimulus generalization following induction of behavioral memory by stimulation of the nucleus basalis

Tone paired with stimulation of the nucleus basalis (NB) induces behavioral memory that is specific to the frequency of the conditioned stimulus (CS), assessed by cardiac and respiration behavior during post-training stimulus generalization testing. This paper focuses on CS-specific spectral and temporal features of conditioned EEG activation. Adult male Sprague-Dawley rats, chronically implanted with a stimulating electrode in the NB and a recording electrode in the ipsilateral auditory cortex, received either tone (6kHz, 70dB, 2s) paired with co-terminating stimulation of the nucleus basalis (0.2s, 100Hz, 80-105 microA, ITI approximately 45s) or unpaired presentation of the stimuli (approximately 200 trials/day for approximately 14 days). CS-specificity was tested 24h post-training by presenting test tones to obtain generalization gradients for the EEG, heart rate, and respiration. Behavioral memory was evident in cardiac and respiratory responses that were maximal to the CS frequency of 6kHz. FFT analyses of tone-elicited changes of power in the delta, theta, alpha, beta1, beta2, and gamma bands in the paired group revealed that conditioned EEG activation (shift from lower to higher frequencies) was differentially spectrally and temporally specific: theta, and alpha to a lesser extent, decreased selectively to 6kHz during and for several seconds following tone presentation while gamma power increased transiently during and after 6kHz. Delta exhibited no CS-specificity and the beta bands showed transient specificity only after several seconds. The unpaired group exhibited neither CS-specific behavioral nor EEG effects. Thus, stimulus generalization tests reveal that conditioned EEG activation is not unitary but rather reflects CS-specificity, with band-selective markers for specific, associative neural processes in learning and memory.     

Simple and contingent aftereffects of perceived duration in vision and audition

After repeated presentations of a long inspection tone (800 or 1,000 msec), a test tone of intermediate duration (600 msec) appeared shorter than it would otherwise appear. A short inspection tone (200 or 400 msec) tended to increase the apparent length of the intermediate test tone. Thus, a negative aftereffect of perceived auditory duration occurred, and a similar aftereffect occurred in the visual modality. These aftereffects, each involving a single sensory dimension, aresimple aftereffects. The following procedures producedcontingent aftereffects of perceived duration. A pair of lights, the first short and the second long, was presented repeatedly during an inspection period. When a pair of test lights of intermediate duration was then presented, the first member of the pair appeared longer in relation to the second. A similar aftereffect occurred in the auditory modality. In these latter aftereffects, the perceived duration of a test light or tone is contingent—dependent—on its temporal order, first or second, within a pair of test stimuli. An experiment designed to test the possibility of cross-modal transfer of contingent aftereffects between audition and vision found no significant cross-modal aftereffects.     

Induced loudness reduction as a function of frequency difference between test tone and inducer

When a high-intensity tone (inducer) is followed by a moderate-intensity tone (test tone), the loudness of the latter is reduced. This phenomenon, called induced loudness reduction (ILR), depends on the frequency separation of the two tones; as the difference in frequency increases, the amount of ILR decreases. However, the precise course of this decrease is not well known. This article presents two experiments that address this question. In the first experiment, the amount of loudness reduction produced by a 2.5-kHz 80-dB-SPL inducer was measured with the frequency of the test tone swept from 800 Hz to 6 kHz. In the second experiment, the amount of ILR was measured with the same inducer and with test tones set at 2, 2.5, 3, and 4 kHz. Both experiments show that some ILR occurs at frequency separations as wide as four critical bands.     

Spectral frequency and the modulation of the acoustic startle reflex by background noise

The rat's (Long-Evans) acoustic startle reflex to a high-frequency tone burst (10.5 kHz) was depressed by intense high-frequency band-pass noise (8-16 kHz) but enhanced by low frequency noise (1-2 kHz). However, contrary to the hypothesis that the depression of startle in intense background noise is produced by sensory masking, the reflex to a low-frequency tone burst (at 1 kHz) was depressed by both high- and low-frequency band-pass noise. Two additional hypotheses are offered to supplement sensory masking in order to explain the asymmetry in these data. The first is that the intratympanic reflex, which acts as a high pass filter on acoustic input, is elicited in intense backgrounds. The second is that acoustic startle reflexes elicited by intense low-frequency tones are in part elicited by their high-frequency distortion products and that these distortion products are then masked by high-frequency background noise.     

Pulling faces: an investigation of the face-distortion aftereffect

After adaptation to a face distorted to look unnaturally thin or fat, a normal face appears distorted in the opposite direction (Webster and MacLin 1999 Psychonomic Bulletin & Review 6 647-653). When the adapting face is oriented 45 degrees from vertically upright and the test face 45 degrees in the opposite direction, the axis of perceived distortion changes with the orientation of the face. The magnitude of this aftereffect shows a reduction of approximately 40% from that found when both adapting and test faces are tilted identically. This finding suggests that to a large degree the aftereffect is mediated not by low-level retinotopic (image-based) visual mechanisms but at a higher level of object-based processing. Aftereffects of a similar magnitude are obtained when adapting and test images are both either upright or inverted, or for an upright adapter and an inverted test; but aftereffects are smaller when the adapter is inverted and the test upright. This pattern of results suggests that the face-distortion aftereffect is mediated by object-processing mechanisms including, but not restricted to, configurational face-processing mechanisms.     

Behavior-Based Assessment of the Auditory Abilities of Brushtail Possums

Brushtail possums (Trichosurus vulpecula) were trained to press a right lever when a tone was presented (a tone‐on trial) and a left lever when a tone was not presented (a tone‐off trial) to gain access to food. During training the tone was set at 80 dB(A), with a frequency of 0.88 kH for 3 possums and of 4 kH for the other 2. Once accuracy was over 90% correct across five consecutive sessions, a test session was conducted where the intensity of the tone was reduced by 8 dB(A) over blocks of 20 trials until accuracy over a block fell below 60%. After each test session, training sessions were reintroduced and continued until accuracy was again over 90%, when another test session was conducted. This process continued until there were at least five test sessions at that tone frequency. The same procedure was then used with frequencies of 0.20, 0.88, 2, 4, 10, 12.5, 15, 20, 30, and 35 kHz. Percentage correct and d' decreased approximately linearly for all possums as tone intensity reduced. Both sets of lines were shallowest at the higher frequencies and steepest at the lower frequencies. Hit and false alarm rates mirrored each other at high frequencies but were asymmetric at lower frequencies. Equal d' contours showed that sensitivity increased from 2 to 15 kHz and continued to be high over 20 to 35 kHz. The possums remained sensitive to the 20 to 35 kHz tones even at low intensities. The present study is the first to report the abilities of possum to detect tones over this range of frequencies and its results support the findings of a microelectrode mapping survey of possums' auditory cortex.     

Influence of duty cycle and off time of comparison-tone pulse trains on the measurement of perstimulatory loudness adaptation

An experiment was conducted to determine if duty cycle and off time of tone pulses presented to the comparison ear influence adaptation measured at the opposite (test) ear. Eight subjects were adapted for 5 min to a 1-kHz pure tone at 60 dB SPL. Using a tracking procedure, adaptation was measured under five comparison-signal conditions, each comprised of 1-kHz pulse trains having different on/off times. The on/off times (in milliseconds) were: 200/800 (20% duty cycle); 500/500, 200/200, and 800/800 (50% duty cycle); and 800/200 (80% duty cycle). Adaptation was found to increase as the duty cycle of the comparison tones increased from 20% to 80%. This was evident even when attempts were made to account for the extent to which pulse trains are perceived as softer than continuous signals at the same level (the so-called LOT effect). For the 50%-duty-cycle conditions, similar amounts of adaptation were measured whether the on/off times of the signals were 200, 500, or 800 msec.     

Ear differences in simple reaction time: The influence of attentional factors

Simple motor reaction times (right-handed) to tape-recorded consonant-vowel syllables were obtained from 12 subjects under two conditions of monotic stimulation: “expected” presentation (subject informed as to ear of presentation) and “random” presentation (ear of presentation varied randomly). Significantly faster left ear reaction times were obtained in the “expected” condition. The rank order of subjects' standard dichotic listening task scores correlated negatively with reaction time ear differences in the “expected” presentation condition. Results are discussed in terms of existing theories of response lateralization in simple reaction time tasks, and an expanded hypothesis is offered. Specifically, it is suggested that two or more distinct but potentially overlapping mechanisms may be responsible for observed asymmetries in monotic auditory perception. Both an attentional or lateralized motor response bias and an inherent lateralization of function may operate side by side, differentially activated by task demands, mode of stimulus presentation, and nature of stimuli.     

The spatial attributes of stimulus frequency and their role in monaural localization of sound in the horizontal plane

Listeners, whose right ears were blocked, located low-intensity sounds originating from loudspeakers placed 15 deg apart along the horizontal plane on the side of the open, or functioning, ear. In Experiment 1, the stimuli consisted of noise bursts, 1.0 kHz wide and centered at 4.0 through 14.0 kHz in steps of .5 kHz. We found that the apparent location of the noise bursts was governed by their frequency composition. Specifically, as the center frequency was increased from 4.0 to about 8.0 kHz, the sound appeared to move away from the frontal sector and toward the side. This migration pattern of the apparent sound source was observed again when the center frequency was increased from 8.0 to about 12.0 kHz. Then, with center frequencies of 13.0 and 14.0 kHz, the sound appeared once more in front. We referred to this relation between frequency composition and apparent location in terms of spatial referent maps. In Experiment 2, we showed that localization was more proficient if the frequency content of the stimulus served to connect adjacent spatial referent maps rather than falling within a single map. By these means, we have further elucidated the spectral cues utilized in monaural localization of sound in the horizontal plane.     

The monaural localization of tonal stimuli

With one ear occluded, 17 listeners were asked to locate tone bursts, .25, 4, .6, .9. l.4, 2.0, 3.2, 4.8, and 7.2 kHz, generated by a loudspeaker concealed from view. The S’s response was to callout that number, from a series of numbers arranged horizontally, behind which he thought the tone bursts originated. The listeners perceived the sounds as emanating from the side of the unoccluded ear, but their judgments bore no consistent relation to the actual location of the sound source. Rather, the listeners showed a strong tendency to locate a tone burst, within the range of .9 through 7.2 kHz, in a fixed spatial relation to the next higher- and lower-pitched tone burst. Distorting the pinna of the unoccluded ear failed to modify the perceptual pattern. It was suggested that the perceived spatial relations among the various frequencies was a by-product of the tonotopic organization of the auditory nervous system.     

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.     

Color—luminance relationships and the McCollough effect

The McCollough effect is an orientation-specific color aftereffect induced by adapting to colored gratings. We examined how the McCollough effect depends on the relationships between color and luminance within the inducing and test gratings and compared the aftereffects to the color changes predicted from selective adaptation to different color—luminance combinations. Our results suggest that the important contingency underlying the McCollough effect is between orientation and color—luminance direction and are consistent with sensitivity changes within mechanisms tuned to specific color—luminance directions. Aftereffects are similar in magnitude for adapting color pairs that differ only in S cone excitation or L and M cone excitation, and they have a similar dependence on spatial frequency. In particular, orientation-specific aftereffects are induced for S cone colors even when the grating frequencies are above the S cone resolution limit. Thus, the McCollough effect persists even when different cone classes encode the orientation and color of the gratings.