The emergence of semantic categorization in early visual processing: ERP indices of animal vs. artifact recognition

Background

Neuroimaging and neuropsychological literature show functional dissociations in brain activity during processing of stimuli belonging to different semantic categories (e.g., animals, tools, faces, places), but little information is available about the time course of object perceptual categorization. The aim of the study was to provide information about the timing of processing stimuli from different semantic domains, without using verbal or naming paradigms, in order to observe the emergence of non-linguistic conceptual knowledge in the ventral stream visual pathway. Event related potentials (ERPs) were recorded in 18 healthy right-handed individuals as they performed a perceptual categorization task on 672 pairs of images of animals and man-made objects (i.e., artifacts).

Results

Behavioral responses to animal stimuli were ~50 ms faster and more accurate than those to artifacts. At early processing stages (120–180 ms) the right occipital-temporal cortex was more activated in response to animals than to artifacts as indexed by posterior N1 response, while frontal/central N1 (130–160) showed the opposite pattern. In the next processing stage (200–260) the response was stronger to artifacts and usable items at anterior temporal sites. The P300 component was smaller, and the central/parietal N400 component was larger to artifacts than to animals.

Conclusion

The effect of animal and artifact categorization emerged at ~150 ms over the right occipital-temporal area as a stronger response of the ventral stream to animate, homomorphic, entities with faces and legs. The larger frontal/central N1 and the subsequent temporal activation for inanimate objects might reflect the prevalence of a functional rather than perceptual representation of manipulable tools compared to animals. Late ERP effects might reflect semantic integration and cognitive updating processes. Overall, the data are compatible with a modality-specific semantic memory account, in which sensory and action-related semantic features are represented in modality-specific brain areas.     

Arithmetic mismatch negativity and numerical magnitude processing in number matching

Background

After a prolonged exposure to a paired presentation of different types of signals (e.g., color and motion), one of the signals (color) becomes a driver for the other signal (motion). This phenomenon, which is known as contingent motion aftereffect, indicates that the brain can establish new neural representations even in the adult's brain. However, contingent motion aftereffect has been reported only in visual or auditory domain. Here, we demonstrate that a visual motion aftereffect can be contingent on a specific sound.

Results

Dynamic random dots moving in an alternating right or left direction were presented to the participants. Each direction of motion was accompanied by an auditory tone of a unique and specific frequency. After a 3-minutes exposure, the tones began to exert marked influence on the visual motion perception, and the percentage of dots required to trigger motion perception systematically changed depending on the tones. Furthermore, this effect lasted for at least 2 days.

Conclusions

These results indicate that a new neural representation can be rapidly established between auditory and visual modalities.     

The developmental pattern of stimulus and response interference in a color-object Stroop task: an ERP study

Background

Several studies have shown that Stroop interference is stronger in children than in adults. However, in a standard Stroop paradigm, stimulus interference and response interference are confounded. The purpose of the present study was to determine whether interference at the stimulus level and the response level are subject to distinct maturational patterns across childhood. Three groups of children (6–7 year-olds, 8–9 year-olds, and 10–12 year-olds) and a group of adults performed a manual Color-Object Stroop designed to disentangle stimulus interference and response interference. This was accomplished by comparing three trial types. In congruent (C) trials there was no interference. In stimulus incongruent (SI) trials there was only stimulus interference. In response incongruent (RI) trials there was stimulus interference and response interference. Stimulus interference and response interference were measured by a comparison of SI with C, and RI with SI trials, respectively. Event-related potentials (ERPs) were measured to study the temporal dynamics of these processes of interference.

Results

There was no behavioral evidence for stimulus interference in any of the groups, but in 6–7 year-old children ERPs in the SI condition in comparison with the C condition showed an occipital P1-reduction (80–140 ms) and a widely distributed amplitude enhancement of a negative component followed by an amplitude reduction of a positive component (400–560 ms). For response interference, all groups showed a comparable reaction time (RT) delay, but children made more errors than adults. ERPs in the RI condition in comparison with the SI condition showed an amplitude reduction of a positive component over lateral parietal (-occipital) sites in 10–12 year-olds and adults (300–540 ms), and a widely distributed amplitude enhancement of a positive component in all age groups (680–960 ms). The size of the enhancement correlated positively with the RT response interference effect.

Conclusion

Although processes of stimulus interference control as measured with the color-object Stroop task seem to reach mature levels relatively early in childhood (6–7 years), development of response interference control appears to continue into late adolescence as 10–12 year-olds were still more susceptible to errors of response interference than adults.     

The function of Shp2 tyrosine phosphatase in the dispersal of acetylcholine receptor clusters

Background

Tone languages such as Thai and Mandarin Chinese use differences in fundamental frequency (F₀, pitch) to distinguish lexical meaning. Previous behavioral studies have shown that native speakers of a non-tone language have difficulty discriminating among tone contrasts and are sensitive to different F₀ dimensions than speakers of a tone language. The aim of the present ERP study was to investigate the effect of language background and training on the non-attentive processing of lexical tones. EEG was recorded from 12 adult native speakers of Mandarin Chinese, 12 native speakers of American English, and 11 Thai speakers while they were watching a movie and were presented with multiple tokens of low-falling, mid-level and high-rising Thai lexical tones. High-rising or low-falling tokens were presented as deviants among mid-level standard tokens, and vice versa. EEG data and data from a behavioral discrimination task were collected before and after a two-day perceptual categorization training task.

Results

Behavioral discrimination improved after training in both the Chinese and the English groups. Low-falling tone deviants versus standards elicited a mismatch negativity (MMN) in all language groups. Before, but not after training, the English speakers showed a larger MMN compared to the Chinese, even though English speakers performed worst in the behavioral tasks. The MMN was followed by a late negativity, which became smaller with improved discrimination. The High-rising deviants versus standards elicited a late negativity, which was left-lateralized only in the English and Chinese groups.

Conclusion

Results showed that native speakers of English, Chinese and Thai recruited largely similar mechanisms when non-attentively processing Thai lexical tones. However, native Thai speakers differed from the Chinese and English speakers with respect to the processing of late F₀ contour differences (high-rising versus mid-level tones). In addition, native speakers of a non-tone language (English) were initially more sensitive to F₀ onset differences (low-falling versus mid-level contrast), which was suppressed as a result of training. This result converges with results from previous behavioral studies and supports the view that attentive as well as non-attentive processing of F₀ contrasts is affected by language background, but is malleable even in adult learners.     

Asymmetrical hippocampal connectivity in mesial temporal lobe epilepsy: evidence from resting state fMRI

Background

Recent studies have shown that the human right-hemispheric auditory cortex is particularly sensitive to reduction in sound quality, with an increase in distortion resulting in an amplification of the auditory N1m response measured in the magnetoencephalography (MEG). Here, we examined whether this sensitivity is specific to the processing of acoustic properties of speech or whether it can be observed also in the processing of sounds with a simple spectral structure. We degraded speech stimuli (vowel /a/), complex non-speech stimuli (a composite of five sinusoidals), and sinusoidal tones by decreasing the amplitude resolution of the signal waveform. The amplitude resolution was impoverished by reducing the number of bits to represent the signal samples. Auditory evoked magnetic fields (AEFs) were measured in the left and right hemisphere of sixteen healthy subjects.

Results

We found that the AEF amplitudes increased significantly with stimulus distortion for all stimulus types, which indicates that the right-hemispheric N1m sensitivity is not related exclusively to degradation of acoustic properties of speech. In addition, the P1m and P2m responses were amplified with increasing distortion similarly in both hemispheres. The AEF latencies were not systematically affected by the distortion.

Conclusions

We propose that the increased activity of AEFs reflects cortical processing of acoustic properties common to both speech and non-speech stimuli. More specifically, the enhancement is most likely caused by spectral changes brought about by the decrease of amplitude resolution, in particular the introduction of periodic, signal-dependent distortion to the original sound. Converging evidence suggests that the observed AEF amplification could reflect cortical sensitivity to periodic sounds.     

Perceptual interactions in the loudness of combined auditory and vibrotactile stimuli

The loudness of auditory (A), tactile (T), and auditory-tactile (A+T) stimuli was measured at supra-threshold levels. Auditory stimuli were pure tones presented binaurally through headphones; tactile stimuli were sinusoids delivered through a single-channel vibrator to the left middle fingertip. All stimuli were presented together with a broadband auditory noise. The A and T stimuli were presented at levels that were matched in loudness to that of the 200-Hz auditory tone at 25 dB sensation level. The 200-Hz auditory tone was then matched in loudness to various combinations of auditory and tactile stimuli (A+T), and purely auditory stimuli (A+A). The results indicate that the matched intensity of the 200-Hz auditory tone is less when the A+T and A+A stimuli are close together in frequency than when they are separated by an octave or more. This suggests that A+T integration may operate in a manner similar to that found in auditory critical band studies, further supporting a strong frequency relationship between the auditory and somatosensory systems.     

Temporal integration of acoustic signals by the budgerigar (Melopsittacus undulatus)

Avoidance conditioning and a modified method of limits psychophysical procedure were used to study temporal integration of tone and noise signals in the budgerigar (Melopsittacus undulatus). Integration of both tone and noise signals can be described by a negative exponential function with a time constant of about 200 ms. At very short durations there were differences in the integration of tone and noise signals. These data are similar to those reported for a number of other vertebrates, including man. Thresholds for two complex natural vocalizations of the budgerigar are similar to those of pure tones of equivalent duration.     

Thresholds for hearing mistuned partials as separate tones in harmonic complexes

When a low harmonic in a harmonic complex tone is mistuned from its harmonic value by a sufficient amount it is heard as a separate tone, standing out from the complex as a whole. This experiment estimated the degree of mistuning required for this phenomenon to occur, for complex tones with 10 or 12 equal-amplitude components (60 dB SPL per component). On each trial the subject was presented with a complex tone which either had all its partials at harmonic frequencies or had one partial mistuned from its harmonic frequency. The subject had to indicate whether he heard a single complex tone with one pitch or a complex tone plus a pure tone which did not "belong" to the complex. An adaptive procedure was used to track the degree of mistuning required to achieve a d' value of 1. Threshold was determined for each ot the first six harmonics of each complex tone. In one set of conditions stimulus duration was held constant at 410 ms, and the fundamental frequency was either 100, 200, or 400 Hz. For most conditions the thresholds fell between 1% and 3% of the harmonic frequency, depending on the subject. However, thresholds tended to be greater for the first two harmonics of the 100-Hz fundamental and, for some subjects, thresholds increased for the fifth and sixth harmonics. In a second set of conditions fundamental frequency was held constant at 200 Hz, and the duration was either 50, 110, 410, or 1610 ms. Thresholds increased by a factor of 3-5 as duration was decreased from 1610 ms to 50 ms. The results are discussed in terms of a hypothetical harmonic sieve and mechanisms for the formation of perceptual streams.     

Transcranial Magnetic Stimulation for the treatment of tinnitus: Effects on cortical excitability

Background

Low frequency repetitive transcranial magnetic stimulation (rTMS) has been proposed as an innovative treatment for chronic tinnitus. The aim of the present study was to elucidate the underlying mechanism and to evaluate the relationship between clinical outcome and changes in cortical excitability. We investigated ten patients with chronic tinnitus who participated in a sham-controlled crossover treatment trial. Magnetic-resonance-imaging and positron-emission-tomography guided 1 Hz rTMS were performed over the auditory cortex on 5 consecutive days. Active and sham treatments were separated by one week. Parameters of cortical excitability (motor thresholds, intracortical inhibition, intracortical facilitation, cortical silent period) were measured serially before and after rTMS treatment by using single- and paired-pulse transcranial magnetic stimulation. Clinical improvement was assessed with a standardized tinnitus-questionnaire.

Results

We noted a significant interaction between treatment response and changes in motor cortex excitability during active rTMS. Specifically, clinical improvement was associated with an increase in intracortical inhibition, intracortical facilitation and a prolongation of the cortical silent period. These results indicate that intraindividual changes in cortical excitability may serve as a correlate of response to rTMS treatment.

Conclusion

The observed alterations of cortical excitability suggest that low frequency rTMS may evoke long-term-depression like effects resulting in an improvement of subcortical inhibitory function.     

Electrophysiological evidence for an early processing of human voices

Background  

Previous electrophysiological studies have identified a "voice specific response" (VSR) peaking around 320 ms after stimulus onset, a latency markedly longer than the 70 ms needed to discriminate living from non-living sound sources and the 150 ms to 200 ms needed for the processing of voice paralinguistic qualities. In the present study, we investigated whether an early electrophysiological difference between voice and non-voice stimuli could be observed.     

Auditory stream segregation in monkey auditory cortex: effects of frequency separation, presentation rate, and tone duration

Auditory stream segregation refers to the organization of sequential sounds into "perceptual streams" reflecting individual environmental sound sources. In the present study, sequences of alternating high and low tones, "...ABAB...," similar to those used in psychoacoustic experiments on stream segregation, were presented to awake monkeys while neural activity was recorded in primary auditory cortex (A1). Tone frequency separation (AF), tone presentation rate (PR), and tone duration (TD) were systematically varied to examine whether neural responses correlate with effects of these variables on perceptual stream segregation. "A" tones were fixed at the best frequency of the recording site, while "B" tones were displaced in frequency from "A" tones by an amount = delta F. As PR increased, "B" tone responses decreased in amplitude to a greater extent than "A" tone responses, yielding neural response patterns dominated by "A" tone responses occurring at half the alternation rate. Increasing TD facilitated the differential attenuation of "B" tone responses. These findings parallel psychoacoustic data and suggest a physiological model of stream segregation whereby increasing delta F, PR, or TD enhances spatial differentiation of "A" tone and "B" tone responses along the tonotopic map in A1.     

Mapping the brain's orchestration during speech comprehension: task-specific facilitation of regional synchrony in neural networks

Background

How does the brain convert sounds and phonemes into comprehensible speech? In the present magnetoencephalographic study we examined the hypothesis that the coherence of electromagnetic oscillatory activity within and across brain areas indicates neurophysiological processes linked to speech comprehension.

Results

Amplitude-modulated (sinusoidal 41.5 Hz) auditory verbal and nonverbal stimuli served to drive steady-state oscillations in neural networks involved in speech comprehension. Stimuli were presented to 12 subjects in the following conditions (a) an incomprehensible string of words, (b) the same string of words after being introduced as a comprehensible sentence by proper articulation, and (c) nonverbal stimulations that included a 600-Hz tone, a scale, and a melody. Coherence, defined as correlated activation of magnetic steady state fields across brain areas and measured as simultaneous activation of current dipoles in source space (Minimum-Norm-Estimates), increased within left- temporal-posterior areas when the sound string was perceived as a comprehensible sentence. Intra-hemispheric coherence was larger within the left than the right hemisphere for the sentence (condition (b) relative to all other conditions), and tended to be larger within the right than the left hemisphere for nonverbal stimuli (condition (c), tone and melody relative to the other conditions), leading to a more pronounced hemispheric asymmetry for nonverbal than verbal material.

Conclusions

We conclude that coherent neuronal network activity may index encoding of verbal information on the sentence level and can be used as a tool to investigate auditory speech comprehension.

     

Modulation of auditory evoked responses to spectral and temporal changes by behavioral discrimination training

Background  

Due to auditory experience, musicians have better auditory expertise than non-musicians. An increased neocortical activity during auditory oddball stimulation was observed in different studies for musicians and for non-musicians after discrimination training. This suggests a modification of synaptic strength among simultaneously active neurons due to the training. We used amplitude-modulated tones (AM) presented in an oddball sequence and manipulated their carrier or modulation frequencies. We investigated non-musicians in order to see if behavioral discrimination training could modify the neocortical activity generated by change detection of AM tone attributes (carrier or modulation frequency). Cortical evoked responses like N1 and mismatch negativity (MMN) triggered by sound changes were recorded by a whole head magnetoencephalographic system (MEG). We investigated (i) how the auditory cortex reacts to pitch difference (in carrier frequency) and changes in temporal features (modulation frequency) of AM tones and (ii) how discrimination training modulates the neuronal activity reflecting the transient auditory responses generated in the auditory cortex.     

Rejection of randomly coinciding events in ZnMoO_4 scintillating bolometers

Random coincidence of events (particularly from two neutrino double beta decay) could be one of the main sources of background in the search for neutrinoless double beta decay with cryogenic bolometers due to their poor time resolution. Pulse-shape discrimination by using front edge analysis, mean-time and \(\chi ^2\) methods were applied to discriminate randomly coinciding events in ZnMoO \(_4\) cryogenic scintillating bolometers. These events can be effectively rejected at the level of 99 % by the analysis of the heat signals with rise-time of about 14 ms and signal-to-noise ratio of 900, and at the level of 92 % by the analysis of the light signals with rise-time of about 3 ms and signal-to-noise ratio of 30, under the requirement to detect 95 % of single events. These rejection efficiencies are compatible with extremely low background levels in the region of interest of neutrinoless double beta decay of \(^{100}\) Mo for enriched ZnMoO \(_4\) detectors, of the order of \(10^{-4}\)  counts/(y keV kg). Pulse-shape parameters have been chosen on the basis of the performance of a real massive ZnMoO \(_4\) scintillating bolometer. Importance of the signal-to-noise ratio, correct finding of the signal start and choice of an appropriate sampling frequency are discussed.     

Virtual pitch integration for asynchronous harmonics

This experiment examined the generation of virtual pitch for harmonically related tones that do not overlap in time. The interval between successive tones was systematically varied in order to gauge the integration period for virtual pitch. A pitch discrimination task was employed, and both harmonic and nonharmonic tone series were tested. The results confirmed that a virtual pitch can be generated by a series of brief, harmonically related tones that are separated in time. Robust virtual pitch information can be derived for intervals between successive 40-ms tones of up to about 45 ms, consistent with a minimum estimate of integration period of about 210 ms. Beyond intertone intervals of 45 ms, performance becomes more variable and approaches an upper limit where discrimination of tone sequences can be undertaken on the basis of the individual frequency components. The individual differences observed in this experiment suggest that the ability to derive a salient virtual pitch varies across listeners.     

Schwann cells migrate along axons in the absence of GDNF signaling

Background

Emotionally salient information in spoken language can be provided by variations in speech melody (prosody) or by emotional semantics. Emotional prosody is essential to convey feelings through speech. In sensori-neural hearing loss, impaired speech perception can be improved by cochlear implants (CIs). Aim of this study was to investigate the performance of normal-hearing (NH) participants on the perception of emotional prosody with vocoded stimuli. Semantically neutral sentences with emotional (happy, angry and neutral) prosody were used. Sentences were manipulated to simulate two CI speech-coding strategies: the Advance Combination Encoder (ACE) and the newly developed Psychoacoustic Advanced Combination Encoder (PACE). Twenty NH adults were asked to recognize emotional prosody from ACE and PACE simulations. Performance was assessed using behavioral tests and event-related potentials (ERPs).

Results

Behavioral data revealed superior performance with original stimuli compared to the simulations. For simulations, better recognition for happy and angry prosody was observed compared to the neutral. Irrespective of simulated or unsimulated stimulus type, a significantly larger P200 event-related potential was observed for happy prosody after sentence onset than the other two emotions. Further, the amplitude of P200 was significantly more positive for PACE strategy use compared to the ACE strategy.

Conclusions

Results suggested P200 peak as an indicator of active differentiation and recognition of emotional prosody. Larger P200 peak amplitude for happy prosody indicated importance of fundamental frequency (F0) cues in prosody processing. Advantage of PACE over ACE highlighted a privileged role of the psychoacoustic masking model in improving prosody perception. Taken together, the study emphasizes on the importance of vocoded simulation to better understand the prosodic cues which CI users may be utilizing.     

Vocal Fold Vibration and Phonation Start in Aspirated, Unaspirated, and Staccato Onset

Objectives/Hypotheses

Singers learn to produce well-controlled tone onsets by accurate synchronization of glottal adduction and buildup of subglottal pressure. Spectrographic analyses have shown that the higher spectrum partials are present also at the vowel onset in classically trained singers’ performances. Such partials are produced by a sharp discontinuity in the waveform of the transglottal airflow, presumably produced by vocal fold collision.

Study Design

After hearing a prompt series of a triad pattern, six singer subjects sang the same triad pattern on the vowel /i/ (1) preceded by an aspirated /p/, (2) preceded by an unaspirated /p/, and (3) without any preceding consonant in staccato.

Methods

Using high-speed imaging we examined the initiation of vocal fold vibration in aspirated and unaspirated productions of the consonant /p/ as well as in the staccato tones.

Results

The number vibrations failing to produce vocal fold collision were significantly higher in the aspirated /p/ than in the unaspirated /p/ and in the staccato tones. High frequency ripple in the audio waveform was significantly delayed in the aspirated /p/.

Conclusions

Initiation of vocal fold collision and the appearance of high-frequency ripple in the vowel /i/ are slightly delayed in aspirated productions of a preceding consonant /p/.     

Temporal discrimination for single components of nonspeech auditory patterns

This paper extends previous research on listeners' abilities to discriminate the details of brief tonal components occurring within sequential auditory patterns (Watson et al., 1975, 1976). Specifically, the ability to discriminate increments in the duration delta t of tonal components was examined. Stimuli consisted of sequences of ten sinusoidal tones: a 40-ms test tone to which delta t was added, plus nine context tones with individual durations fixed at 40 ms or varying between 20 and 140 ms. The level of stimulus uncertainty was varied from high (any of 20 test tones occurring in any of nine factorial contexts), through medium (any of 20 test tones occurring in ten contexts), to minimal levels (one test tone occurring in a single context). The ability to discriminate delta t depended strongly on the level of stimulus uncertainty, and on the listener's experience with the tonal context. Asymptotic thresholds under minimal uncertainty approached 4-6 ms, or 15% of the duration of the test tones; under high uncertainty, they approached 40 ms, or 10% of the total duration of the tonal sequence. Initial thresholds exhibited by inexperienced listeners are two-to-four times greater than the asymptotic thresholds achieved after considerable training (20,000-30,000 trials). Isochronous sequences, with context tones of uniform, 40-ms duration, yield lower thresholds than those with components of varying duration. The frequency and temporal position of the test tones had only minor effects on temporal discrimination. It is proposed that a major determinant of the ability to discriminate the duration of components of sequential patterns is the listener's knowledge about "what to listen for and where." Reduced stimulus uncertainty and extensive practice increase the precision of this knowledge, and result in high-resolution discrimination performance. Increased uncertainty, limited practice, or both, would allow only discrimination of gross changes in the temporal or spectral structure of the sequential patterns.     

Integration of spectral and temporal cues separated in time and frequency

Subjects discriminated a "standard" pair of tone bursts (T1, T2) from a "comparison" pair (T1 + delta t, T2 + delta f), containing increments in the duration delta t of the first burst and/or the frequency delta f of the second burst. The threshold (d' = 2.0) for delta t was measured as a function of delta f, and the threshold for delta f as a function of delta t. The integration of increments in duration and frequency was studied as a function of the spectral and temporal separation between T1 and T2. A trade-off between the values of delta t and delta f required for d' = 2.0 performance was observed. This integration takes place when delta t, delta f occur simultaneously in the same spectral region, and when they occur separated by up to 120 ms, or by up to a full octave. The efficiency of integration was similar for all conditions of temporal and spectral separation studied, because the discriminability of delta t and of delta f is also nearly uniform across experimental conditions. The results from all experimental conditions are adequately described by a vector summation model derived from TSD. In a subsidiary experiment, subjects categorized pure tones varying in duration and frequency as "high" or "low" in pitch and "long" or "short" in duration. It was found that combined variations in duration and frequency result in essentially independent perceptual processes, although pitch has a small effect upon the perceived duration. It is concluded that spectral-temporal integration is a general ability operating in a variety of stimulus conditions.(ABSTRACT TRUNCATED AT 250 WORDS)     

The role of spectral and periodicity cues in auditory stream segregation, measured using a temporal discrimination task.

In a previous paper, it was shown that sequential stream segregation could be based on both spectral information and periodicity information, if listeners were encouraged to hear segregation [Vliegen and Oxenham, J. Acoust. Soc. Am. 105, 339-346 (1999)]. The present paper investigates whether segregation based on periodicity information alone also occurs when the task requires integration. This addresses the question: Is segregation based on periodicity automatic and obligatory? A temporal discrimination task was used, as there is evidence that it is difficult to compare the timing of auditory events that are perceived as being in different perceptual streams. An ABA ABA ABA... sequence was used, in which tone B could be either exactly at the temporal midpoint between two successive tones A or slightly delayed. The tones A and B were of three types: (1) both pure tones; (2) both complex tones filtered through a fixed passband so as to contain only harmonics higher than the 10th, thereby eliminating detectable spectral differences, where only the fundamental frequency (f0) was varied between tones A and B; and (3) both complex tones with the same f0, but where the center frequency of the spectral passband varied between tones. Tone A had a fixed frequency of 300 Hz (when A and B were pure tones) or a fundamental frequency (f0) of 100 Hz (when A and B were complex tones). Five different intervals, ranging from 1 to 18 semitones, were used. The results for all three conditions showed that shift thresholds increased with increasing interval between tones A and B, but the effect was largest for the conditions where A and B differed in spectrum (i.e., the pure-tone and the variable-center-frequency conditions). The results suggest that spectral information is dominant in inducing (involuntary) segregation, but periodicity information can also play a role.