Organizing sound sequences in the human brain: the interplay of auditory streaming and temporal integration

The present study examined the relationship between two of the early brain processes of sound organization: auditory streaming and the temporal window of integration (TWI). Presented at a fast stimulus delivery rate, two tones alternating in frequency are perceived as separate streams of high and low sounds. However, when two sounds are presented within a ca. 200 ms temporal window, they are often processed as a single auditory event. Both stream segregation and temporal integration occur even in the absence of focused attention as was shown by their effect on the mismatch negativity (MMN) event-related potential. The goal of the present study was to determine the precedence between these two sound organization processes by using the stimulus-omission MMN paradigm. Infrequently omitting one stimulus from a homogeneous tone sequence only elicits an MMN when the stimulus onset asynchrony separating successive tones is shorter than 170 ms. This demonstrates the effect of the TWI. Magnetic brain responses elicited by infrequent stimulus omissions appearing in a sequence of two alternating tones were recorded. The magnetic MMN was elicited by tone omission when the alternating tones formed a single stream (with no or only small frequency separation between the two tones) but not when separate high and low streams emerged in perception (large frequency separation between the two alternating tones). This result shows that auditory streaming takes precedence over the processes of temporal integration.     

Fast vigilance decrement in closed head injury patients as reflected by the mismatch negativity (MMN)

Event-related potentials (ERPs) were measured from 24 chronic closed head injury (CHI) patients and 18 age- and education-matched controls. The oddball paradigm was applied while subjects were watching a silent movie. The standard (p=0.8) sound of 75 ms duration had a basic frequency of 500 Hz with harmonic partials of 1000 Hz and 1500 Hz, whereas these frequencies for the pitch deviant were each 10% higher. The frequencies of the duration deviant matched with those of the standard but was 25 ms in duration. The MMN (mismatch negativity), generated by the brain's automatic auditory change-detector mechanism, was elicited by both deviants. No significant differences in the MMN latency or amplitude for either pitch or duration deviants were found between the groups. However, the MMN amplitude for the pitch deviant decreased in the patient group during the experiment considerably faster than in controls, suggesting a faster vigilance decrement in the patients.     

The mismatch negativity to frequency deviant stimuli during natural sleep

Eight subjects spent a single night in the sleep laboratory. Event-related potentials (ERPs) were recorded during the presentation of two auditory ‘oddball’ stimulus conditions in which tonal frequency was manipulated. In the first condition, 1000 Hz ‘standard’ and 2000 Hz ‘deviant’ tones were presented. In the second condition, the deviant tone was reduced to 1050 Hz. In both conditions, deviant probability was 0.2. Stimuli were presented every 600 ms during wakefulness and stages 2, 4, and REM of sleep. A distinctive N1 wave was visible in both stimulus conditions when the subject was awake. The deviant stimuli elicited a ‘mismatch negativity’ (MMN) that inverted in polarity at the mastoid. In REM sleep, an N1 and a MMN were also elicited in both conditions. In the large deviance condition, the MMN had a slightly attenuated amplitude and was shorter in duration while in the small deviant condition, its peak latency was unusually early. Neither the N1 nor the MMN could be recorded in non-REM sleep.     

Independent component analysis on the mismatch negativity in an uninterrupted sound paradigm

We compared the efficiency of the independent component analysis (ICA) decomposition procedure against the difference wave (DW) and optimal digital filtering (ODF) procedures in the analysis of the mismatch negativity (MMN). The comparison was made in a group of 54 children aged 8-16 years. The MMN was elicited in a passive oddball protocol presenting uninterrupted auditory stimulation consisting of two frequent alternating tones (600 and 800 Hz) of 100 ms duration each. Infrequently, one of the 600 Hz tones was shortened to 50 or 30 ms. The event related potentials (ERPs) were decomposed into the MMN-like and non-MMN-like independent components (ICs) through the FastICA algorithm. The ICA decomposition procedure extracted a cleaner MMN compared to the ODF or DW procedures. It extracted the MMN, whose characteristics concurred with the substantial number of publications demonstrating a significantly larger peak amplitude and shorter latency of the MMN in response to the more deviant stimulus (30 ms) compared to the less deviant stimulus (50 ms). The MMN to these two deviant stimuli did not differ in the peak amplitude or latency when it was extracted through the other two procedures. The ICA decomposition and ODF procedures, similarly, significantly improved the single trial signal-to-noise ratio (SNR) of the MMN compared to the DW procedure. Due to this improvement, the proposed ICA decomposition procedure might allow shortening of the recording session and could be used to study the MMN in paradigms similar to this with small modifications.     

The development of memory trace depending on the number of the standard stimuli.

The mismatch negativity (MMN) component of the event-related potentials reflects the automatic detection mechanism of sound change. MMN is elicited by a neuronal mismatch process between deviant (infrequent) auditory input and the sensory memory trace of the standard (frequent) stimuli. Although many previous studies have investigated MMN to reveal the sensory memory mechanism, the development of memory representation still remains unclear, in particular, the topographical aspect of the trace-development in sensory memory has not been clarified. We measured the frontal and the temporal MMN components, respectively, when the sound trace was developed as the number of standard stimuli was changed to 1, 3, 5 or 7. In this experiment, the inter-train interval was 15 sec. The stimulus train with the different frequency of 800 Hz, 900 Hz, or 1000 Hz was repeatedly presented. Thus, we reduced the influence of the previous train. For the first time, we found not only the enhanced amplitude but also the shortened latency for both MMN components when the number of standard stimuli was increased. These findings indicate that both frontal and temporal MMN components reflect the development of memory trace depending on the number of standard stimuli.     

Do tone duration changes that elicit the mismatch negativity also affect the preceding middle latency responses?

The human brain can automatically detect sound changes. Previous studies have reported that rare sounds presented within a sequence of repetitive sounds elicit the mismatch negativity (MMN) in the absence of attention in the latency range of 100–250 ms. On the other hand, a previous study discovered that occasional changes in sound location enhance the middle latency response (MLR) elicited in the latency range of 10–50 ms. Several studies have reported an increase in the amplitude of the MLR within the frame of oddball paradigms such as frequency and location changes. However, few studies have been conducted on paradigms employing a duration change. The purpose of the present study was to examine whether the peak amplitudes of the MLR components are enhanced by a change in duration. Twenty healthy Japanese men (age: 23.9 ± 2.9 years) participated in the present study. We used an oddball paradigm that contained standard stimuli with a duration of 10 ms and deviant stimuli with a duration of 5 ms. The peak amplitudes of the MLR for the deviant stimuli were then compared with those for the standard stimuli. No changes were observed in the peak amplitude of the MLR resulting from a duration change, whereas a definite MMN was elicited. The amplitude of the MLR was increased within the frame of oddball paradigms such as frequency and location changes. By contrast, the amplitude of the MLR was not changed within the duration change oddball paradigm that elicited the MMN.     

An electrophysiological and behavioral investigation of involuntary attention towards auditory frequency,duration and intensity changes

We measured behavior and event-related brain potentials (ERPs) in 12 subjects performing on an audio-visual distraction paradigm to investigate the cerebral mechanisms of involuntary attention towards stimulus changes in the acoustic environment. Subjects classified odd/even numbers presented on a computer screen 300 ms after the occurrence of a task-irrelevant auditory stimulus, by pressing the corresponding response button. Auditory stimuli were standard tones (600 Hz, 200 ms, 85 dB; P=0.8) or deviant tones (P=0.2), these differing from the standard either in frequency (700 Hz), duration (50 ms) or intensity (79 dB), in separate blocks. In comparison to performance to visual stimuli following the standard tones, reaction time increased by 24 ms (F(1,11)=10.91, P<0.01) and hit rate decreased by 4.6% (F(1,11)=35.47, P<0.001) to visual stimuli following the deviant tones, indicating behavioral distraction. ERPs revealed the mismatch negativity (MMN) elicited to deviant tones, which was larger for the duration deviant than for the frequency and intensity deviants (F(2,22)=19.43, P<0.001, epsilon =0.83), and which had different scalp distribution for all three deviant conditions (F(16,176)=2.40, P<0.05, epsilon =0.12). As the shorter duration and softer intensity deviant tones were unlikely to engage fresh neurons responding to their specific physical features, the present results indicate that a genuine change detection mechanism is involved in triggering attention switching towards sound changes, and suggest a largely distributed neural network of the auditory cortex underlying such involuntary attention switching.     

Frequency specific impairment of automatic pitch change detection by fMRI acoustic noise: an MEG study

The loud acoustic noise produced by the magnetic resonance scanner is a major source of interference in auditory fMRI research. Whole-head magnetoencephalography (MEG) was used to investigate the interaction between the frequency range of auditory stimulation and fMRI acoustic noise. Pure tones and 3-harmonic complexes varying between 240 and 1240 Hz in frequency were presented while participants attended to a silent subtitled film. Continuous fMRI acoustic noise was presented during half of the blocks. The activity in six regions of interest was analyzed in 100-200 and 200-300 ms time windows to evaluate the magnetic counterparts of the mismatch negativity (MMN) and P3a brain responses. The results suggested that fMRI noise significantly reduced the amplitude of these responses. The effect of the noise on the automatic processing of the tones was more prominent for the tones with frequencies higher than 500 Hz. It is recommended that in the MMN protocols using continuous fMRI acquisition the sound stimuli should be spectrally separated from the fMRI scanner noise spectrum.     

High-frequency gamma activity (80-150Hz) is increased in human cortex during selective attention.

OBJECTIVE: To study the role of gamma oscillations (>30Hz) in selective attention using subdural electrocorticography (ECoG) in humans. METHODS: We recorded ECoG in human subjects implanted with subdural electrodes for epilepsy surgery. Sequences of auditory tones and tactile vibrations of 800 ms duration were presented asynchronously, and subjects were asked to selectively attend to one of the two stimulus modalities in order to detect an amplitude increase at 400 ms in some of the stimuli. RESULTS: Event-related ECoG gamma activity was greater over auditory cortex when subjects attended auditory stimuli and was greater over somatosensory cortex when subjects attended vibrotactile stimuli. Furthermore, gamma activity was also observed over prefrontal cortex when stimuli appeared in either modality, but only when they were attended. Attentional modulation of gamma power began approximately 400 ms after stimulus onset, consistent with the temporal demands on attention. The increase in gamma activity was greatest at frequencies between 80 and 150 Hz, in the so-called high-gamma frequency range. CONCLUSIONS: There appears to be a strong link between activity in the high-gamma range (80-150 Hz) and selective attention. SIGNIFICANCE: Selective attention is correlated with increased activity in a frequency range that is significantly higher than what has been reported previously using EEG recordings.     

Hemispheric differences in processing tone frequency and amplitude modulations.

Transient frequency and amplitude modulations (FMs, AMs) of sound are requisite to speech recognition. We recorded whole-head magnetoencephalographic signals from seven subjects to binaural 620 ms 667 Hz tones, with 3, 30, or 300 ms FMs or AMs in the beginning or middle of the tone. Responses were significantly larger and earlier for FMs than AMs, for rapid than slow modulations, and for modulations at the beginning (BEG) than in the middle (MID) of the sound. BEG 3 ms FMs elicited strongest signals in the left and MID 3 ms FMs in the right hemisphere. Fast MID modulations produced significantly stronger responses in the right than left hemisphere. These differences may reflect different functions of the left and right hemisphere in speech perception.     

Frequency discrimination at different frequency levels as indexed by electrophysiological and behavioral measures

The present study systematically compared the neural and behavioral accuracy of discriminating a frequency change ("deviant") in a repetitive tone ("standard") across a frequency range of 250-4000 Hz. The sound structure (pure sinusoidal vs. harmonically rich tones) and the magnitude of frequency change (2.5%, 5%, 10%, 20%) were also varied. The accuracy of neural frequency-change detector was determined by comparing the auditory event-related potentials (ERP) elicited by deviant and standard stimuli in the absence of attention. In a separate behavioral task, subjects were to indicate when they noticed a frequency change. The ranges of the across-subject means of ERP parameters across the conditions were: the mismatch negativity (MMN) amplitude -0.9 to -4.9 microV, latency 125-218 ms, the P3a amplitude 0.3-3.2 microV, latency 239-304 ms. The ERP latency was shortest for the standard-stimulus frequency from 1000 to 2000 Hz suggesting that automatic frequency discrimination was the most accurate in that range. The ERP latencies and amplitudes correlated with the hit rate (HR) and reaction time (RT), with highest correlation found between the MMN amplitude and the HR (r=0.8). The harmonical tones elicited MMN and P3a with shorter latencies and larger amplitudes, than did pure sinusoidal tones in all frequency bands. The results may have implication to pitch-perception theories.     

Parameter-specific modulation of the mismatch negativity to duration decrement and increment: Evidence for asymmetric processes

ObjectiveThe mismatch negativity (MMN) component of event-related potentials (ERPs) reflects a change-detection process in the brain. The present study investigated whether stimulus parameters (sound type and duration) exert a differential influence on the MMN for a duration decrement and increment of an equal magnitude. Some asymmetries were reported in the previous studies; yet no systematical study has been conducted.MethodsERPs were recorded from 16 healthy adults presented with repetitive standard sounds interspersed with duration changes (deviant sounds). In separate sequences, stimuli were vowels, music chord, sinusoid, or band-pass filtered white noise. The stimulus durations (standard/deviant) were either 200/120 ms or 400/240 ms for decrements, and vice versa for increments.ResultsThe MMN for the increments was abolished in the 400/240 ms condition, whereas the MMN for decrements was significant irrespective of the sound duration. The amplitude of the increment MMN paralleled with the spectral complexity of the stimulus sound, whereas that of the decrement MMN was larger for natural sounds than artificial sounds.ConclusionsThe observed interactions demonstrated asymmetries in the MMN for duration increment and decrement.SignificanceThe present findings suggest that the effects of stimulus parameters should be taken into account when comparing different studies, especially where clinical populations are involved, with one another.     

Effects of sequential and temporal probability of deviant occurrence on mismatch negativity

The mismatch negativity (MMN) increases in amplitude as the probability of deviant occurrence decreases. It is unclear whether the determining variable is sequential probability (i.e. the probability of a deviant within a number of standards) or temporal probability (i.e. the probability of a deviant within a period of time). Eight subjects heard a train of frequently occurring 1000 Hz standard tones. The probability of a 1100 Hz pitch deviant was manipulated. In one condition the stimulus-onset-asynchrony (SOA) was 150 ms, with temporal probability of deviant occurrence being either 1/9.00, 1/4.50, 1/2.25, or 1/1.125 s (sequential probability being 1/60, 1/30, 1/15, or 1(deviant)/7.5(standards), respectively). In another condition the SOA was 600 ms, with temporal probability being either 1/9.00, 1/4.50, or 1/2.25 s (sequential probability being 1/15, 1/7.5, or 1/3.75, respectively). In a final condition, the SOA was 2400 ms with temporal probability being 1/9.00 s (sequential probability 1/3.75). Both sequential and temporal probabilities had a marked effect on the MMN. When a deviant occurred every 2.25, 4.50, or 9.00 s, the MMN increased as temporal probability decreased. When a deviant occurred once every 7.5 or 15 standards, the MMN was larger for lower sequential probability, but the effect was not significant. Nevertheless, when temporal probability was held constant at 1/9.00 s, the MMN increased as sequential probability decreased. At rapid rates of stimulus presentation, the MMN was largest. However, it was attenuated when the probability of deviant occurrence was very high perhaps due to the refractoriness of its generator. At the slowest rate, the MMN was diminished perhaps due to memory decay for the standard stimuli.     

Auditory mismatch negativity deficits in long-term heavy cannabis users

Mismatch negativity (MMN) is an auditory event-related potential indicating auditory sensory memory and information processing. The present study tested the hypothesis that chronic cannabis use is associated with deficient MMN generation. MMN was investigated in age- and gender-matched chronic cannabis users (n = 30) and nonuser controls (n = 30). The cannabis users were divided into two groups according to duration and quantity of cannabis consumption. The MMNs resulting from a pseudorandomized sequence of 2 × 900 auditory stimuli were recorded by 32-channel EEG. The standard stimuli were 1,000 Hz, 80 dB SPL and 90 ms duration. The deviant stimuli differed in duration (50 ms) or frequency (1,200 Hz). There were no significant differences in MMN values between cannabis users and nonuser controls in both deviance conditions. With regard to subgroups, reduced amplitudes of frequency MMN at frontal electrodes were found in long-term (≥8 years of use) and heavy (≥15 joints/week) users compared to short-term and light users. The results indicate that chronic cannabis use may cause a specific impairment of auditory information processing. In particular, duration and quantity of cannabis use could be identified as important factors of deficient MMN generation.     

Salience in a social landscape: electrophysiological effects of task-irrelevant and infrequent vocal change

In a dynamically changing social environment, humans have to face the challenge of prioritizing stimuli that compete for attention. In the context of social communication, the voice is the most important sound category. However, the existing studies do not directly address whether and how the salience of an unexpected vocal change in an auditory sequence influences the orientation of attention. In this study, frequent tones were interspersed with task-relevant infrequent tones and task-irrelevant infrequent vocal sounds (neutral, happy and angry vocalizations). Eighteen healthy college students were asked to count infrequent tones. A combined event-related potential (ERP) and EEG time–frequency approach was used, with the focus on the P3 component and on the early auditory evoked gamma band response, respectively. A spatial-temporal principal component analysis was used to disentangle potentially overlapping ERP components. Although no condition differences were observed in the 210–310 ms window, larger positive responses were observed for emotional than neutral vocalizations in the 310–410 ms window. Furthermore, the phase synchronization of the early auditory evoked gamma oscillation was enhanced for happy vocalizations. These findings support the idea that the brain prioritizes the processing of emotional stimuli, by devoting more attentional resources to salient social signals even when they are not task-relevant.     

Effects of sleep onset on the mismatch negativity (MMN) to frequency deviants using a rapid rate of presentation

This study examined the effects of sleep onset-the transition from a waking, conscious state to one of sleep and unconsciousness-on the mismatch negativity (MMN) following frequency deviants when a rapid rate of stimulus presentation is employed. The MMN is thought to reflect a brief-lasting sensory memory. Rapid rates of stimulus presentation should guard the sensory memory from fading. A 1,000 Hz standard stimulus was presented every 150 ms. At random, on 6.6% of the trials, the standard was changed to either a large 2,000 or a small 1,100 Hz deviant. During alert wakefulness (when subject ignored the stimuli and read a book), the large deviant elicited a larger deviant related negativity (DRN) than did the small deviant. This negativity may be a composite of both N1 and MMN activity while that following the small deviant is probably a 'true' MMN. The large deviant continued to elicit a DRN in relaxed wakefulness (eyes closed) and Stages 1 and 2 of sleep, although it was much reduced in amplitude. A significant MMN was recorded for the small deviant only in alert wakefulness. The failure to observe an MMN to small deviance and the attenuation of the DRN to large deviance at sleep onset therefore is probably not due to a decay of sensory memory. It is more likely that cortical encoding of both the standard and deviant is weakened during sleep onset because of prior thalamic inhibition of sensory input.     

A modified oddball paradigm "cross-modal delayed response" and the research on mismatch negativity

A modified oddball paradigm was developed to facilitate the focus of attention and to minimize target effects on deviant-related components of auditory and visual event-related potentials (ERPs) elicited with long interstimulus intervals. Subjects were required to focus on either the visual or auditory stimulus in each stimulus block. Deviant-related components were obtained by subtracting ERPs of the standard stimulus from that of the deviant stimulus for each modality with each stimulus condition. Results showed that auditory mismatch negativity (MMN) and a visual early deviant related negativity (DRN1) were elicited both when stimuli were attended and unattended. In contrast, N2b and P3 were produced only under the attended condition. In comparison of attended MMN and unattended MMN at three time windows (100-150 ms, 150-200 ms, and 200-250 ms) of MMN zone, different scalp distributions were shown, depending on the time windows. This result suggests that the attended auditory MMN is a mixed wave, consisting of genuine MMN, N2b, and possible P165. The effect of attention on MMN may stem from the contamination of these overlapping components. With the present paradigm, at least three sensory memory traces have to be maintained simultaneously in multiple sensory modalities to support automatic processing.     

Spectral and temporal stimulus characteristics in the processing of abstract auditory features

The processing of abstract stimulus features in the human brain was studied by presenting the subjects with frequent standard tone pairs and infrequent deviant tone pairs. Both pairs varied randomly over a wide frequency and/or intensity range, there being no physically constant standard stimulus. The common feature of the standard pairs was the direction of change within the pair, e.g. the second tone was louder in intensity and/or higher in frequency than the first tone. Deviant pairs, having opposite feature-change direction, elicited the mismatch-negativity (MMN) event-related potential component. MMN was similar to deviations in the direction of frequency and intensity changes and showed no additivity for simultaneous changes in both feature directions. Moreover, MMN was elicited even when the within-pair interval exceeded the 200 ms limit of auditory temporal integration. Results demonstrate that extraction of abstract features is not limited to frequency-based rules, nor is it dependent on temporal integration mechanisms. The lack of MMN additivity between violations of multiple abstract rules suggests that the processing of higher-order invariances differs from that of simple physical features.     

Acute effects of Delta9-tetrahydrocannabinol and standardized cannabis extract on the auditory evoked mismatch negativity

Reduced amplitudes of auditory evoked mismatch negativity (MMN) have often been found in schizophrenic patients, indicating deficient auditory information processing and working memory. Cannabis-induced psychotic states may resemble schizophrenia. Currently, there are discussions focusing on the close relationship between cannabis, the endocannabinoid and dopaminergic system, and the onset of schizophrenic psychosis. This study investigated the effects of cannabis on MMN amplitude in 22 healthy volunteers (age 28+/-6 years, 11 male) by comparing Delta(9)-tetrahydrocannabinol (Delta(9)-THC) and standardized cannabis extract containing Delta(9)-THC and cannabidiol (CBD) in a prospective, double-blind, placebo-controlled cross-over design. The MMNs resulting from 1000 auditory stimuli were recorded by 32 channel EEG. The standard stimuli were 1000 Hz, 80 dB SPL, and 100 ms duration. The deviant stimuli differed in frequency (1500 Hz). Significantly greater MMN amplitude values at central electrodes were found under cannabis extract, but not under Delta(9)-THC. There were no significant differences between MMN amplitudes at frontal electrodes. MMN amplitudes at central electrodes were significantly correlated with 11-OH-THC concentration, the most important psychoactive metabolite of Delta(9)-THC. Since the main difference between Delta(9)-THC and standardized cannabis extract is CBD, which seems to have neuroprotective and anti-psychotic properties, it can be speculated whether the greater MMN amplitude that may imply higher cortical activation and cognitive performance is related to the positive effects of CBD. This effect may be relevant for auditory cortex activity in particular because only MMN amplitudes at the central, but not at the frontal electrodes were enhanced under cannabis.     

Event-related potential features indexing central auditory discrimination by newborns

Behavioral research has produced little evidence on sound feature discrimination in neonates. Sensory processes underlying sound perception can be studied using the mismatch negativity (MMN) component of auditory event-related potentials (ERPs), which is not contingent on conscious perception and response. Thus, MMN is suitable for studying newborns, who are difficult to obtain behavioral responses from. The present study thus utilized spectrally rich sounds, known to elicit the most replicable MMN in adults, to investigate newborns' preattentive analysis of sound duration and frequency changes. An attempt was also made to control for the obligatory ERP effects on the MMN. Three-partial harmonic tones were presented in Duration and in Frequency oddball conditions to 55 newborns. In the other two, Equiprobable duration and Equiprobable frequency, conditions frequency and duration deviants of the oddball paradigms were presented with equal probabilities among sounds of other durations and frequencies. MMN was elicited in 81% of newborns in Frequency oddball condition and in 78% of newborns in Duration oddball condition. No significant amplitude differences between the duration and frequency MMNs were found, but MMN latency was delayed in Duration condition. The obligatory components seemed to contribute significantly to the deviant-standard difference in Duration but not in Frequency condition. The majority of neonates appear to possess effective sound frequency and duration discrimination mechanisms. Their preattentive sound discrimination is facilitated by spectrally rich sound content. The present findings support a change-detection nature of MMN in neonates; however, sound duration-related obligatory effects need to be taken into account in infant MMN studies.