Evoked magnetic responses of the human auditory cortex to minor pitch changes: localization of the mismatch field

The neuromagnetic source localizations of the auditory M100 and the mismatch field (MMF) were studied using a large-array biomagnetometer. Standard tones of 1000 Hz and deviant tones of 1050 Hz were delivered with 90% and 10% probability, respectively. Wave forms of the derived MMF were computed by examining difference wave forms between the responses to the deviants and the responses to the standards preceding (D-P) and following (D-F) the deviants as well as to all remaining standards (D-A). The subset of standards preceding the deviants was used for a more realistic comparison with the set of deviants (having the same number of epochs and a similar signal-to-noise ratio). while the subset of standards following the deviants served to answer the question whether those standards also elicit an MMF. The MMF deflections were compared with each other, with the “native” MMF occurring in response to the deviants, and with wave M100. (The MMF as it appears in the unprocessed response to the deviants was termed “native” for an easy distinction from the “derived” MMF).Our results demonstrate a distinct MMF deflection, corresponding in latency to the simultaneously recorded fronto-central electrical MMN. Source analysis, using a single moving dipole model, showed the same spatial localization for the native MMF and for the different derived MMFs. The MMF source location turned out to be significantly anterior, medial and inferior relative to the sources of the M100. The present data also demonstrate that a minor frequency deviation may not activate measurably different M100 generators, yet be sufficient to trigger the nearby but spatially distinct mismatch generator.     

Auditory event-related potentials to deviant stimuli during drowsiness and stage 2 sleep

Twelve subjects were tested using a 3-tone auditory oddball paradigm consisting of a standard 1000 Hz tone (P = 80%) and two deviants, namely, a 1200 Hz tone and a 2000 Hz tone (both P = 10%). Testing took place in 3 conditions: (1) attend, in which the subject had to count one of the deviant tones; (2) ignore, in which the subject read a book; and (3) sleep, in which the subject was encouraged to go to sleep during presentation of the tones.In the awake conditions stimulus deviance elicited mismatch negativity (MMN) and P3. During drowsiness, no separate mismatch negativity (MMN) could be detected, but the 2000 Hz tone evoked a broad fronto-central early negative deflection, suggesting an overlap of N₁ and MMN. In the same condition, P210, N330 and P430 appeared, all being sensitive to magnitude of deviance. During stage 2, the P210, N330 and P430 amplitudes increased, most notably to the large deviant.These data indicate that differential processing of auditory inputs is maintained during drowsiness and stage 2 sleep, but do not support the notion that MMN or P3 activity comparable to the waking state occurs to oddball stimuli during this stage. It is hypothesised that during light sleep, scanning of the environment is performed by a different system than in the awake state and that during drowsiness a gradual switch between these two systems takes place.     

The mismatch negativity for duration decrement of auditory stimuli in healthy subjects

The amplitude and latency of the mismatch negativity (MMN) elicited by occasional shorter-duration tones (25 and 50 ms) in a sequence of 75 ms standard tones were studied in 40 healthy subjects (9–84 years). The replicability and age dependence of the MMN-responses were determined. The 25 ms deviant tone evoked a clear response in 39 of the subjects, while the 50 ms deviant tone evoked an observable MMN only in 32 of the subjects. The MMN peak amplitude for the 25 ms deviants was significantly larger than for the 50 ms deviants. There was no significant difference in the peak latencies (measured from stimulus offset). For the 25 ms deviant, the amplitude diminished with increasing age. The MMN curves for the 25 ms deviant, measured on separate days in 14 subjects, looked very replicable. As a result of noise and filtering effect, the product-moment correlations were poor. The results indicate that the signal-to-noise ratio for the MMN to 25 ms deviants, obtained even in a 25 min recording session, is large enough for clinical use and individual diagnostics when undetectable (or very low amplitude) MMN is used as a sign of pathology. However, judged from the low correlation coefficients, despite the good replicability in visual evaluation, better methods for MMN quantification have to be used for clinical follow-up.     

Influence of very low doses of mediators on fungal laccase activity - nonlinearity beyond imagination

Background  

Compared to the waveform or spectrum analysis of event-related potentials (ERPs), time-frequency representation (TFR) has the advantage of revealing the ERPs time and frequency domain information simultaneously. As the human brain could be modeled as a complicated nonlinear system, it is interesting from the view of psychological knowledge to study the performance of the nonlinear and linear time-frequency representation methods for ERP research. In this study Hilbert-Huang transformation (HHT) and Morlet wavelet transformation (MWT) were performed on mismatch negativity (MMN) of children. Participants were 102 children aged 8–16 years. MMN was elicited in a passive oddball paradigm with duration deviants. The stimuli consisted of an uninterrupted sound including two alternating 100 ms tones (600 and 800 Hz) with infrequent 50 ms or 30 ms 600 Hz deviant tones. In theory larger deviant should elicit larger MMN. This theoretical expectation is used as a criterion to test two TFR methods in this study. For statistical analysis MMN support to absence ratio (SAR) could be utilized to qualify TFR of MMN.     

Neuromagnetic evidence for early auditory restoration of fundamental pitch

Background

Understanding the time course of how listeners reconstruct a missing fundamental component in an auditory stimulus remains elusive. We report MEG evidence that the missing fundamental component of a complex auditory stimulus is recovered in auditory cortex within 100 ms post stimulus onset.

Methodology

Two outside tones of four-tone complex stimuli were held constant (1200 Hz and 2400 Hz), while two inside tones were systematically modulated (between 1300 Hz and 2300 Hz), such that the restored fundamental (also knows as “virtual pitch”) changed from 100 Hz to 600 Hz. Constructing the auditory stimuli in this manner controls for a number of spectral properties known to modulate the neuromagnetic signal. The tone complex stimuli only diverged on the value of the missing fundamental component.

Principal Findings

We compared the M100 latencies of these tone complexes to the M100 latencies elicited by their respective pure tone (spectral pitch) counterparts. The M100 latencies for the tone complexes matched their pure sinusoid counterparts, while also replicating the M100 temporal latency response curve found in previous studies.

Conclusions

Our findings suggest that listeners are reconstructing the inferred pitch by roughly 100 ms after stimulus onset and are consistent with previous electrophysiological research suggesting that the inferential pitch is perceived in early auditory cortex.     

Memory-based mismatch response to frequency changes in rats

Any occasional changes in the acoustic environment are of potential importance for survival. In humans, the preattentive detection of such changes generates the mismatch negativity (MMN) component of event-related brain potentials. MMN is elicited to rare changes ('deviants') in a series of otherwise regularly repeating stimuli ('standards'). Deviant stimuli are detected on the basis of a neural comparison process between the input from the current stimulus and the sensory memory trace of the standard stimuli. It is, however, unclear to what extent animals show a similar comparison process in response to auditory changes. To resolve this issue, epidural potentials were recorded above the primary auditory cortex of urethane-anesthetized rats. In an oddball condition, tone frequency was used to differentiate deviants interspersed randomly among a standard tone. Mismatch responses were observed at 60-100 ms after stimulus onset for frequency increases of 5% and 12.5% but not for similarly descending deviants. The response diminished when the silent inter-stimulus interval was increased from 375 ms to 600 ms for +5% deviants and from 600 ms to 1000 ms for +12.5% deviants. In comparison to the oddball condition the response also diminished in a control condition in which no repetitive standards were presented (equiprobable condition). These findings suggest that the rat mismatch response is similar to the human MMN and indicate that anesthetized rats provide a valuable model for studies of central auditory processing.     

Paired associative stimulation of the auditory system: a proof-of-principle study

Background

Paired associative stimulation (PAS) consisting of repeated application of transcranial magnetic stimulation (TMS) pulses and contingent exteroceptive stimuli has been shown to induce neuroplastic effects in the motor and somatosensory system. The objective was to investigate whether the auditory system can be modulated by PAS.

Methods

Acoustic stimuli (4 kHz) were paired with TMS of the auditory cortex with intervals of either 45 ms (PAS(45 ms)) or 10 ms (PAS(10 ms)). Two-hundred paired stimuli were applied at 0.1 Hz and effects were compared with low frequency repetitive TMS (rTMS) at 0.1 Hz (200 stimuli) and 1 Hz (1000 stimuli) in eleven healthy students. Auditory cortex excitability was measured before and after the interventions by long latency auditory evoked potentials (AEPs) for the tone (4 kHz) used in the pairing, and a control tone (1 kHz) in a within subjects design.

Results

Amplitudes of the N1-P2 complex were reduced for the 4 kHz tone after both PAS(45 ms) and PAS(10 ms), but not after the 0.1 Hz and 1 Hz rTMS protocols with more pronounced effects for PAS(45 ms). Similar, but less pronounced effects were observed for the 1 kHz control tone.

Conclusion

These findings indicate that paired associative stimulation may induce tonotopically specific and also tone unspecific human auditory cortex plasticity.     

Short-term replicability of the mismatch negativity

The short-term replicability of the mismatch negativity (MMN) between two recording sessions spaced 2 h apart was evaluated at individual and group levels in a sample of 11 healthy adults. Subjects were presented with a random sequence of 1000 Hz standard (92%) and 1100 Hz deviant (8%) tones while they were reading a book. The N1 and P2 exogenous components to standard tones showed a fairly good individual and group replicability. There were no significant differences in the MMN amplitude and latency between the two sessions in the group of subjects as a whole. The individual replicability of the MMN was not as good as for the N1 to standards, reaching significance in only some of the electrodes. This result was, however, similar to that obtained for the N1 after deviant tones. The results indicate that the MMN has good replicability at the group level, and further that at the individual level, MMN replicability is similar to that of the N1 to deviants. This suggests that the number of summations should be increased in order to improve the clinical usefulness of the MMN.     

The phases of visual cortex and hippocampal evoked potentials in rabbit reflect the orienting reaction in case of visual stimuli intensity changes

The oddball paradigm was applied in experiments with waking rabbits using rare (deviant) and frequent (standard) stimuli, which were similar in color but different in intensity, deviant ones being of lower intensity. In addition, the VEPs to the single deviants were averaged. Such single deviants (without standard stimuli) were given at the beginning and at the end of the stimulation. The positivity of cortical and hippocampal visual evoked potentials to deviant stimuli increased in comparison to responses to standards and single deviants. The VEP-peaks P1 and P2 in the visual cortex and the VEP P1, N1 and P2 peaks in hippocampus increased. The most prominent significant changes were demonstrated for cortical VEP P2 (P130) peak. It is suggested that the increase of positivity to oddball deviants is due to the orienting reflex arising in response to rare stimuli. The increase of P2-peak can be connected with the transfer of information signaling changes of light intensity. It was demonstrated that the most clear and contrast differences in the VEPs to deviants and standards took place for the case of minimal distinction in their intensities. This fact may reflect the orienting reactionto threshold stimuli.     

Effects of Presentation Rate and Attention on Auditory Discrimination: A Comparison of Long-Latency Auditory Evoked Potentials in School-Aged Children and Adults

Decoding human speech requires both perception and integration of brief, successive auditory stimuli that enter the central nervous system as well as the allocation of attention to language-relevant signals. This study assesses the role of attention on processing rapid transient stimuli in adults and children. Cortical responses (EEG/ERPs), specifically mismatch negativity (MMN) responses, to paired tones (standard 100–100Hz; deviant 100–300Hz) separated by a 300, 70 or 10ms silent gap (ISI) were recorded under Ignore and Attend conditions in 21 adults and 23 children (6–11 years old). In adults, an attention-related enhancement was found for all rate conditions and laterality effects (L>R) were observed. In children, 2 auditory discrimination-related peaks were identified from the difference wave (deviant-standard): an early peak (eMMN) at about 100–300ms indexing sensory processing, and a later peak (LDN), at about 400–600ms, thought to reflect reorientation to the deviant stimuli or “second-look” processing. Results revealed differing patterns of activation and attention modulation for the eMMN in children as compared to the MMN in adults: The eMMN had a more frontal topography as compared to adults and attention played a significantly greater role in childrens’ rate processing. The pattern of findings for the LDN was consistent with hypothesized mechanisms related to further processing of complex stimuli. The differences between eMMN and LDN observed here support the premise that separate cognitive processes and mechanisms underlie these ERP peaks. These findings are the first to show that the eMMN and LDN differ under different temporal and attentional conditions, and that a more complete understanding of children’s responses to rapid successive auditory stimulation requires an examination of both peaks.     

Interstimulus interval and auditory event-related potentials in children: evidence for multiple generators

In the present study, the component structure of auditory event-related potentials (ERP) was studied in children of 7–9 years old by presenting stimuli with different interstimulus intervals (ISI). A short-term auditory sensory memory, as reflected by ISI effects on ERPs, was also studied. Auditory ERPs were recorded to brief unattended 1000 Hz frequent, `standard' and 1100 Hz rare, `deviant' (probability 0.1) tone stimuli with ISIs of 350, 700 and 1400 ms (in separate blocks). With the 350 ms-ISI, the ERP waveform to the standard stimulus consisted of P100-N250 peaks. With the two longer ISIs, in addition, the frontocentral N160 and N460 peaks were observed. Results suggested that N160, found with the longer ISIs, is a correlate of the adult auditory N1. In difference waves, obtained by subtracting ERP to standard stimuli from ERP to deviant stimuli, two negativities were revealed. The first was the mismatch negativity (MMN), which is elicited by any discriminable change in repetitive auditory input. The MMN data suggested that neural traces of auditory sensory memory lasted for at least 1400 ms, probably considerably longer, as no MMN attenuation was found across the ISIs used. The second, later negativity was similar to MMN in all aspects, except for the scalp distribution, which was posterior to that of the MMN.     

DURATION DISCRIMINATION IN NORTHERN BOBWHITE QUAIL

ABSTRACT

We examined the ability of northern bobwhite quail Colinus virginianus to discriminate along the duration dimension because of its suspected importance in their ability to identify individual covey members. If quail are using variability along this dimension to identify individuals we would expect their auditory resolving powers to be sufficiently refined in that dimension to make appropriate discriminations. Three 2-year old male bobwhite quail were trained using a positive reinforcement technique to discriminate between a 1000 Hz standard (correct) tone 600 ms in duration (100 ms rise/fall time) and a 1000 Hz comparison tone 300 ms in duration (also 10 ms rise/fall time). Quail were then required to discriminate between pairs of tones in which the duration of the comparison tone was progressively increased from 400 ms to 590 ms. Because of the categorical nature of the analysis we concluded that the two tones must differ in duration by 20 ms or more if they are to be discriminated correctly. Our study has shown that bobwhites are able to detect a 3% difference in duration around a standard tone 600 ms in duration, which compares favourably with results obtained from earlier studies with parakeets Melopsittacus undulatus and pigeons Columbia livia. Such a finely tuned duration discrimination ability also suggests that it is possible bobwhites may be using individual variability along the duration dimension of the separation call to identify individual covey members.     

Correlation between auditory thalamic area evoked responses and species-specific call characteristics

Evoked potentials were recorded from the posterior dorsal thalamus of green treefrogs (Hyla cinerea) in response to single tones and combinations of two and three tones. 1. The responses to two tones were largest when one of the component tones was 500 Hz and when the second component was between 2000 and 4000 Hz (Fig.3). 2. The response to 500 + 3000 Hz showed nonlinear facilitation; i.e., the amplitude of the response was greater than the sum of the responses to the component tones alone (Figs. 4, 5). This result provides evidence that cells functioning as 'AND' gates will be found in this center. 3. When a third tone around 1200 Hz was added to a stimulus of 500 + 3000 Hz a 65% decrease in the evoked response amplitude occurred (Fig. 6). 4. The largest evoked response amplitude to a two-tone stimulus (500 + 3000 Hz) occurred when the rise-time was less than 50 ms (Fig. 7). 5. The two-tone tuning was found to be temperature dependent. The optimal lower frequency tone shifted downward with decreasing temperatures (Fig. 8). 6. When the temperatures of the neurophysiological and the behavioral experiments are matched, the optimal stimuli for evoking a large response are closely correlated to the parameters of the acoustic stimuli preferred by gravid H. cinerea females in discrimination tests. This center therefore appears to be very important for the processing of complex species-specific sounds.     

Is the somatosensory N250 related to deviance discrimination or conscious target detection?

Effects of attention to, and probability of sudden changes in, repetitive stimuli on somatosensory evoked potentials (SEP) were studied. Low- (30 Hz) and high-frequency (140 Hz) vibratory stimuli were delivered in random order to the middle finger of the left hand with different presentation probabilities in different blocks. Also ignore conditions were administered.In the ignore conditions, the probability had no effect on SEPs. However, when the standard stimuli were omitted, the “deviants” elicited small N140 and P300 deflections not observed in response to deviants when standards were also present. In the attention conditions, deviant stimuli (targets) elicited large N250 and P300 deflections which increased in amplitude with a decreased target probability. However, when subjects counted infrequently presented “deviants” alone (standards omitted) the enhanced N140 and the P300 with shortened latency were elicited, but no N250 wave could be found. At the ipsilateral side, a distinct N200 deflection was seen which could be the N250 with a shorter latency because of an easier task (detection instead of discrimination). The results might be interpreted as suggesting that the somatosensory N250 is related to conscious detection of target stimuli.     

Discrimination of timbre in early auditory responses of the human brain

Background

The issue of how differences in timbre are represented in the neural response still has not been well addressed, particularly with regard to the relevant brain mechanisms. Here we employ phasing and clipping of tones to produce auditory stimuli differing to describe the multidimensional nature of timbre. We investigated the auditory response and sensory gating as well, using by magnetoencephalography (MEG).

Methodology/Principal Findings

Thirty-five healthy subjects without hearing deficit participated in the experiments. Two different or same tones in timbre were presented through conditioning (S1) – testing (S2) paradigm as a pair with an interval of 500 ms. As a result, the magnitudes of auditory M50 and M100 responses were different with timbre in both hemispheres. This result might support that timbre, at least by phasing and clipping, is discriminated in the auditory early processing. The second response in a pair affected by S1 in the consecutive stimuli occurred in M100 of the left hemisphere, whereas both M50 and M100 responses to S2 only in the right hemisphere reflected whether two stimuli in a pair were the same or not. Both M50 and M100 magnitudes were different with the presenting order (S1 vs. S2) for both same and different conditions in the both hemispheres.

Conclusions/Significances

Our results demonstrate that the auditory response depends on timbre characteristics. Moreover, it was revealed that the auditory sensory gating is determined not by the stimulus that directly evokes the response, but rather by whether or not the two stimuli are identical in timbre.     

Stimulus-specific adaptation and deviance detection in the rat auditory cortex

Stimulus-specific adaptation (SSA) is the specific decrease in the response to a frequent ('standard') stimulus, which does not generalize, or generalizes only partially, to another, rare stimulus ('deviant'). Stimulus-specific adaptation could result simply from the depression of the responses to the standard. Alternatively, there may be an increase in the responses to the deviant stimulus due to the violation of expectations set by the standard, indicating the presence of true deviance detection. We studied SSA in the auditory cortex of halothane-anesthetized rats, recording local field potentials and multi-unit activity. We tested the responses to pure tones of one frequency when embedded in sequences that differed from each other in the frequency and probability of the tones composing them. The responses to tones of the same frequency were larger when deviant than when standard, even with inter-stimulus time intervals of almost 2 seconds. Thus, SSA is present and strong in rat auditory cortex. SSA was present even when the frequency difference between deviants and standards was as small as 10%, substantially smaller than the typical width of cortical tuning curves, revealing hyper-resolution in frequency. Strong responses were evoked also by a rare tone presented by itself, and by rare tones presented as part of a sequence of many widely spaced frequencies. On the other hand, when presented within a sequence of narrowly spaced frequencies, the responses to a tone, even when rare, were smaller. A model of SSA that included only adaptation of the responses in narrow frequency channels predicted responses to the deviants that were substantially smaller than the observed ones. Thus, the response to a deviant is at least partially due to the change it represents relative to the regularity set by the standard tone, indicating the presence of true deviance detection in rat auditory cortex.     

Effect of frequency on the loudness of short sounds: comparison between 2 near frequencies, 500 and 1000 Hz]

Loudness egalizations between short tone (1 to 400 ms) and a long reference tone (800 ms) were made. Two frequencies, 500 and 1000 Hz, were employed. Loudness comparisons were made at 20, 40 and 80 dB SL. Results show a frequency effect on the loudness of short tones. Some partial recuperation of loudness appears, it is greater for the high level.     

Expectations Modulate the Magnitude of Attentional Capture by Auditory Events

What determines the magnitude of attentional capture by deviant sound events? We combined the cross-modal oddball distraction paradigm with sequence learning to address this question. Participants responded to visual targets, each preceded by tones that formed a repetitive cross-trial standard sequence. In Experiment 1, with the standard tone sequence …-660-440-660-880-… Hz, either the 440 Hz or the 880 Hz standard was occasionally replaced by one of two deviant tones (220 Hz and 1100 Hz), that either differed slightly (by 220 Hz) or markedly (by 660 Hz) from the replaced standard. In Experiment 2, with the standard tone sequence …-220-660-440-660-880-660-1100-… Hz, the 440 Hz and the 880 Hz standard was occasionally replaced by either a 220 Hz or a 1100 Hz pattern deviant. In both experiments, a high-pitch deviant was more captivating when it replaced a low-pitch standard, and a low-pitch deviant was more captivating when it replaced a high-pitch standard. These results indicate that the magnitude of attentional capture by deviant sound events depends on the discrepancy between the deviant event and the expected event, not on perceived local change.     

Fast detection of unexpected sound intensity decrements as revealed by human evoked potentials

The detection of deviant sounds is a crucial function of the auditory system and is reflected by the automatically elicited mismatch negativity (MMN), an auditory evoked potential at 100 to 250 ms from stimulus onset. It has recently been shown that rarely occurring frequency and location deviants in an oddball paradigm trigger a more negative response than standard sounds at very early latencies in the middle latency response of the human auditory evoked potential. This fast and early ability of the auditory system is corroborated by the finding of neurons in the animal auditory cortex and subcortical structures, which restore their adapted responsiveness to standard sounds, when a rare change in a sound feature occurs. In this study, we investigated whether the detection of intensity deviants is also reflected at shorter latencies than those of the MMN. Auditory evoked potentials in response to click sounds were analyzed regarding the auditory brain stem response, the middle latency response (MLR) and the MMN. Rare stimuli with a lower intensity level than standard stimuli elicited (in addition to an MMN) a more negative potential in the MLR at the transition from the Na to the Pa component at circa 24 ms from stimulus onset. This finding, together with the studies about frequency and location changes, suggests that the early automatic detection of deviant sounds in an oddball paradigm is a general property of the auditory system.     

Limited plasticity of the rabbit visual cortex and hippocampal neurons in the oddball test

The activity of 41 visual cortex and 20 hippocampal neurons from field CA1 was registered in experiments using oddball-stimulation with different color stimuli varied in intensity. 34% cortical and 37% hippocampal neurons demonstrated plasticity reactions. The significant increase of latest phases of neuronal activity (200-500 and 200-1000 ms after stimulation for cortical neurons and 300-550 ms for hippocampal neurons) was shown in responses to rare deviant stimuli, which had a less intensity than frequently standards. The quantity of the earliest neuronal phase of activity (40-120 ms after stimulation) was stabilized in responses to deviants and standards during the experiment. We propose that such increase of the latest phases of neuronal activity (the limited plasticity) may reflect the mechanisms of orienting reaction.