Hemispheric differences in auditory oddball responses during monaural versus binaural stimulation

Hemispheric lateralization of early event-related potentials (ERPs; e.g. N1) is largely based on anatomy of the afferent pathway; lateralization of later auditory ERPs (P2/N2, P250, P3b) is less clear. Using 257-channel EEG, the present study examined hemispheric laterality of auditory ERPs by comparing binaural and monaural versions of an auditory oddball task. N1 showed a contralateral bias over auditory cortex in both hemispheres as a function of ear of stimulation, although right hemisphere sources were activated regardless of which ear received input. Beginning around N1 and continuing through the time of P3b, right hemisphere temporal–parietal and frontal areas were more activated than their left hemisphere counterparts for stimulus evaluation/comparison and target detection. P250 and P3b component amplitudes, topographies, and source estimations were significantly influenced by ear of stimulation, with right hemisphere activity being stronger. This was particularly true for anterior temporal and inferior frontal sources which were more strongly associated with the later, more cognitive components (P250, P3b). Results are consistent with theories of a right hemisphere network that is prominently involved in sustained attention, stimulus evaluation, target detection, and working memory/context updating.     

Age and inter-stimulus interval effects on event-related potentials to frequent and infrequent auditory stimuli.

The aim of this study was to investigate aging effects on non-attended changes of auditory stimulation, by using psychophysiological methods. Event-related potentials (ERPs) were recorded to frequent (standard; 950 Hz, p = 0.9) and infrequent (deviant; 1045 Hz, p = 0.1) auditory stimuli in older (mean age = 60.8 years) and younger (mean age = 21.3 years) subjects. In various blocks the inter-stimulus interval (ISI) was either 800, 2400 or 7200 ms. During the experimental sessions the subjects read books, and ignored the auditory stimuli. As a function of ISI, the amplitude of the N1 and the amplitude and latency of the P2 increased. The P2 amplitude was larger in the younger group. In the 120-180 ms latency range the deviant stimuli elicited more negative ERPs (mismatch negativity, MMN) than the standard stimuli. The amplitude of the MMN did not change as a function of ISI. MMN was larger in the younger group. Thus the younger subjects were more sensitive to the deviant stimuli. In the younger group, at the two shorter ISIs, the MMN was followed by a positive wave (P3a). The emergence of this component is an indication of the increased activity of the orienting system in the younger subjects, in comparison to the older age group.     

Modification of neuromagnetic responses of the human auditory cortex by masking sounds

Summary We have studied the effects of masking sounds on auditory evoked magnetic fields (AEFs) of healthy humans. The AEFs were elicited by 25-ms tones presented randomly to the left or to the right ear, and the responses were recorded over the right auditory cortex. Without masking, the 100-ms deflection (N100m) was of somewhat higher amplitude and of shorter latency for contrathan ipsilateral stimuli. Continuous speech, music, or intermittent noise, delivered to the left ear, dampened N100m to stimulation of both ears without correlated changes in sensation. Intermittent noise had a weaker effect on N100m than speech or music. Continuous noise fed to the left ear dampened both the sensation of and the responses to the left-ear stimuli, with no significant effect on the responses to the right-ear stimuli. The results suggest that the masking effects of continuous noise, seen at the auditory cortex, derive mainly from the periphery whereas the effects of sounds with intensity and frequency modulations take place at more central auditory pathways.     

Characteristics of No-go-P300 component during somatosensory Go/No-go paradigms

The present study investigated one of the characteristics of No-go-related brain activity during somatosensory Go/No-go paradigms, by manipulating the stimulus site and response hand. Somatosensory event-related potentials (ERPs) were recorded in ten right-handed subjects. Electrical stimulation was delivered to the second and fifth digit of one hand, and the subjects had to respond to a Go stimulus by pushing a button with the thumb contralateral to the stimulated side as quickly as possible. We focused on the peak amplitude and latency of Som-Go-P300 (P300 evoked by somatosensory Go stimuli) and Som-No-go-P300 (P300 evoked by somatosensory No-go stimuli) components. The amplitude of Som-No-go-P300, which is very similar to No-go-P300 components following visual and auditory stimulation, was significantly larger than that of Som-Go-P300 at fronto-central electrodes, indicating ‘anteriorization’ of the No-go-P300. The amplitude of Som-No-go-P300 was significantly larger in right than left hemispheres during right hand response conditions, but this difference was not found under left hand conditions. In addition, the difference in amplitude between Som-Go- and Som-No-go-P300, which is frequently described as ‘the Go/No-go effect’ on P300, was significant in the left hemisphere under right hand response conditions, whereas a significant effect was found in both the left and the right hemispheres under left hand response conditions. Our findings suggested that the anteriorization of No-go-P300 was independent of stimulus modalities such as visual, auditory, and somatosensory, and the amplitude of No-go-P300 and Go/No-go effects on P300 was affected by the response hand.     

Reaction Time in a Visual 4-Choice Reaction Time Task: ERP Effects of Motor Preparation and Hemispheric Involvement

Reaction time (RT), the most common measure of CNS efficiency, shows intra- and inter-individual variability. This may be accounted for by hemispheric specialization, individual neuroanatomy, and transient functional fluctuations between trials. To explore RT on these three levels, ERPs were measured in a visual 4-choice RT task with lateralized stimuli (left lateral, left middle, right middle, and right lateral) in 28 healthy right-handed subjects. We analyzed behavioral data, ERP microstates (MS), N1 and P3 components, and trial-by-trial variance. Across subjects, the N1 component was contralateral to the stimulation side. N1-MSs were stronger over the left hemisphere, and middle stimulation evoked stronger activation than lateral stimulation in both hemispheres. The P3 was larger for the right visual field stimulation. RTs were shorter for the right visual hemifield stimulation/right hand responses. Within subjects, covariance analysis of single trial ERPs with RTs showed consistent lateralized predictors of RT over the motor cortex (MC) in the 112–248 ms interval. Decreased RTs were related to negativity over the MC contralateral to the stimulation side, an effect that could be interpreted as the lateralized readiness potential (LRP), and which was strongest for right side stimulation. The covariance analysis linking individual mean RTs and individual mean ERPs showed a frontal negativity and an occipital positivity correlating with decreased RTs in the 212–232 ms interval. We concluded that a particular RT is a composite measure that depends on the appropriateness of the motor preparation to a particular response and on stimulus lateralization that selectively involves a particular hemisphere.     

Event-related potentials (ERPs) to hemifield presentations of emotional stimuli: differences between depressed patients and healthy adults in P3 amplitude and asymmetry.

Depression may involve dysfunction of right parietotemporal cortex, a region activated during perception of affective stimuli. To further test this hypothesis, event-related brain potentials (ERPs) were measured in a paradigm previously shown to produce ERP asymmetries to affective stimuli over parietal sites in healthy adults. Pictures of patients with dermatological diseases showing disordered or healed facial areas before (negative) or after (neutral) surgical treatment were briefly exposed for 250 ms to either the left or right hemifield. ERPs of 30 unmedicated, unipolar depressed patients and 16 healthy adults, all right-handed, were recorded from 30 electrodes. A principal components analysis extracted factors which closely corresponded to distinctive ERP components previously reported for this task (N1, N2, early P3, late P3, slow wave). Significant effects of emotional content, i.e. enhanced amplitudes to negative than neutral stimuli, were found for early and late P3. Control subjects showed significant hemispheric asymmetries of emotional processing for late P3 (peak latency 460 ms), with the largest emotional content effects over the right parietal region. In striking contrast to control subjects, depressed patients did not show an increase in late P3 for negative compared to neutral stimuli over either hemisphere and had smaller late P3 amplitude than control subjects. Patients did, however, show larger early P3 (peak latency 330 ms) to negative than neutral stimuli. Results suggest intact early discrimination but abnormal late appraisal of affective content in depression, which may arise from selective inhibition of right parietal regions integral for perceiving and evaluating emotional stimuli.     

Correspondence between brain ERP and behavioral asymmetries in a dichotic complex tone test

Electrophysiologic correlates of perceptual asymmetry for dichotic pitch discrimination were investigated in 20 normal subjects. Brain event-related potentials (ERPs) elicited by dichotic pairs and binaural probe tones in the Complex Tone Test (Sidtis, 1981) were recorded from homologous scalp locations over left and right hemispheres (F3, F4; C3, C4; P3, P4; O1, O2). Baseline-to-peak amplitudes were measured for N100, P200, and a late positive complex consisting of P350, P550, and slow wave. A left ear advantage (LEA) was evident in 70% of the subjects, and hemispheric asymmetries related to this behavioral asymmetry were found for P350 and P550 amplitudes to probe stimuli. Subjects with a strong LEA had greater amplitudes over the right hemisphere than the left, whereas subjects with little or no LEA showed a nonsignificant trend toward the opposite hemispheric asymmetry. Hemispheric asymmetry of these late ERPs at parietal and occipital sites was highly correlated with behavioral asymmetry. These findings suggest the utility of electrophysiological measures in assessing hemispheric asymmetries for processing complex pitch information.     

The Effect of Stimulus Significance on Skin Conductance Recovery

The relationship between skin conductance (SC) recovery time and stimulus meaning was investigated. Previous research has shown that a strong relationship between prestimulus electrodermal activity (prior activity) and recovery time makes it necessary to control prior activity before the relationship of recovery time and stimulus meaning can be evaluated properly. In this study prior activity was controlled by experimental design. We examined the effects of stimulus meaning on skin conductance recovery time and amplitude in 55 teenage and young adult subjects in a continuous performance paradigm. Tones were presented monaurally to left and right ears. Subjects were required to make a pedal press after any tone in the left or right ear that followed a right-ear tone. Recovery time was computed for left-ear and right-ear tones not requiring a press because they had been preceded by a left-ear tone. SC recovery was longer for signal (right-ear) than for nonsignal (left-ear) tones with the effects of prior SC activity and amplitude removed. These findings replicate those of a pilot study (Janes, 1982). We conclude that stimulus significance can affect SC recovery time and that in this paradigm differences in prior SC activity cannot account for the recovery time differences observed.     

The effects of interstimulus interval on sensory gating and on preattentive auditory memory in the oddball paradigm: Can magnitude of the sensory gating affect preattentive auditory comparison process?

P50, and mismatch negativity (MMN) are components of event-related potentials (ERP) reflecting sensory gating and preattentive auditory memory, respectively. Interstimulus interval (ISI) is an important determinant of the amplitudes of these components and N1. In the present study the interrelation between stimulus gating and preattentive auditory sensory memory were investigated as a function of ISI in 1.5, 2.5 and 3.5 s in 15 healthy volunteered participants. ISI factor affected the N1 peak amplitude significantly. MMN amplitude in 2.5 s ISI was significantly smaller compared to 1.5 and 3.5 s ISI. ISI X stimuli interaction on P50 amplitude was statistically significant. P50 amplitudes to deviant stimuli in 2.5 s ISI were larger than the P50 amplitudes in other ISIs. P50 difference (P50d) waveform amplitude correlated significantly with MMN amplitude. The results suggest that: (i) auditory sensory gating could affect preattentive auditory sensory memory by supplying input to the comparator mechanism; (ii) 2.5 s ISI is important in displaying the sensory gating and preattentive auditory sensory memory relation.     

Automatic auditory information processing in sleep

STUDY OBJECTIVES: The mismatch negativity (MMN) component of the event-related potentials reflects the automatic detection of sound change. Only a few researchers have investigated the MMN elicitation during sleep in adult human and some of them reported that MMN amplitude was decreased in sleep compared to in waking. However, it is not clear that the decrease of MMN amplitude was due to increased drowsiness or long-term response decrement. Two experiments were conducted to clarify whether or not the MMN was elicited in each sleep stage. We presented auditory stimuli to subjects continuously from waking until sleep state (Experiment 1). Using the same experimental condition, we examined whether or not MMN amplitude was influenced by long-term stimulus presentation (80min.) and by vigilance level (Experiment 2). DESIGN: N/A SETTING: N/A PARTICIPANTS: N/A INTERVENTIONS: N/A MEASUREMENTS & RESULTS: Experiment 1: MMN was obtained in both drowsiness and REM sleep. MMN was significantly smaller in amplitude and shorter in latency in the both stages than in the waking state. However, MMN was not found in another sleep stage. Experiment 2: Amplitudes were no different among 0-20 min., 20-40 min., 60-80 min. But it was decreased in 40-60 min. and power value of alpha-wave was decreased in 40-60 min. CONCLUSIONS: To obtain the reliable data, by using the automatic spectral analysis, we confirmed that MMN was elicited in REM sleep. MMN was not influenced by long-term stimulation. The result suggested that auditory stimuli could be processed in the pre-attentive sensory memory even in REM sleep.     

Brain event-related potentials to complex tones in depressed patients: Relations to perceptual asymmetry and clinical features

Brain event-related potentials (ERPs) to probe tones in a dichotic complex tone test were recorded from right-handed depressed patients (n= 44) and normal subjects (n= 19) at homologous sites over left and right hemispheres (F3, F4; C3, C4; P3, P4; O1, O2). There were no differences between groups in N1 or P2 amplitude, but patients had smaller P3 amplitude than did normal subjects. Depressed patients failed to show either the left ear advantage or behavior-related hemispheric asymmetry of P3 seen for normal subjects. Depressed patients also showed less differences in hemispheric asymmetry between same and different judgments. These findings indicate that the abnormal behavioral asymmetry for dichotic pitch discrimination in depressed patients reflects a reduction in hemispheric asymmetry and is related to relatively late stages of cognitive processing.     

Selectivity of N170 for visual words in the right hemisphere: Evidence from single‐trial analysis

Neuroimaging and neuropsychological studies have identified the involvement of the right posterior region in the processing of visual words. Interestingly, in contrast, ERP studies of the N170 typically demonstrate selectivity for words more strikingly over the left hemisphere. Why is right hemisphere selectivity for words during the N170 epoch typically not observed, despite the clear involvement of this region in word processing? One possibility is that amplitude differences measured on averaged ERPs in previous studies may have been obscured by variation in peak latency across trials. This study examined this possibility by using single‐trial analysis. Results show that words evoked greater single‐trial N170s than control stimuli in the right hemisphere. Additionally, we observed larger trial‐to‐trial variability on N170 peak latency for words as compared to control stimuli over the right hemisphere. Results demonstrate that, in contrast to much of the prior literature, the N170 can be selective to words over the right hemisphere. This discrepancy is explained in terms of variability in trial‐to‐trial peak latency for responses to words over the right hemisphere.     

Interhemispheric Difference for Upright and Inverted Face Perception in Humans: An Event-Related Potential Study

We recorded event-related potentials (ERPs) to investigate the interhemispheric difference of the N170 component for upright and inverted face perception in detail in fifteen healthy subjects. This is the first ERP study focusing on interhemispheric differences for face perception by showing faces in the hemifield. The face inversion effect, the prolonged latency and enhanced amplitude were found in both hemispheres. We found that the peak latency of the N170 following both upright and inverted face stimulation showed no significant difference between each hemisphere, though the N170 latency for the inverted face in the left hemisphere was shorter than that in the right hemisphere. The N170 recorded from the hemisphere ipsilateral to the stimulated hemifield showed unique findings. The interhemispheric time difference of the N170 between the right and the left hemispheres when the inverted face was presented in the left hemifield was significantly shorter than in the other three conditions. This unique finding may indicate that the conduction time from the right to the left for inverted face perception is faster than the other conditions, or that the left hemisphere specifically processed the inverted face very rapidly after receiving signals from the right hemisphere. If the N170 was generated by some, at least two, temporally overlapping activities, the different style of a summation of these activities may cause the unique findings found in this study. In conclusion, by presenting face stimuli in the hemifields, we could identify several new findings regarding the N170 component related to the face inversion effect.     

Mismatch negativity in dichotic listening: Evidence for interhemispheric differences and multiple generators

The characteristics of mismatch negativity (MMN) elicited by dichotic stimulation were examined using frequency-deviant stimuli presented to the right, to the left, or to both sides. The experiment was run twice, once using earphones and once using loudspeakers in free field. With both modes of stimulation, deviants presented in the left, right, or both ears, or tones that were switched between ears, elicited comparable MMNs, with a peak latency of about 180 ms. With earphones, the amplitude of the MMN was bigger at the frontal-lateral right hemisphere sites than at the homologous left-hemisphere sites for all deviance conditions. Scalp current density analysis revealed that deviance in the right side elicited bilaterally equivalent frontal current sinks and a trend towards stronger contralateral current sources at the mastoid sites. In contrast, left side deviance elicited frontal sinks and temporal current sources stronger over the right hemiscalp. These results are compatible with the multiple-generator model of MMN. The attention-related role of the MMN is discussed, suggesting comparable attention mechanisms for vision and audition.     

The auditory evoked magnetic fields to very high frequency tones

We studied the auditory evoked magnetic fields (AEFs) in response to pure tones especially at very high frequencies (from 4000 Hz to 40,000 Hz). This is the first systematic study of AEFs using tones above 5000 Hz, the upper audible range of humans, and ultrasound. We performed two experiments. In the first, AEFs were recorded in 12 subjects from both hemispheres under binaural listening conditions. Six types of auditory stimulus (pure tones of five different frequencies: 4000 Hz, 8000 Hz, 10,000 Hz, 12,000 Hz, 14,000 Hz, and a click sound as the target stimulus) were used. In the second experiment, we used 1000 Hz, 15,000 Hz, and two ultrasounds with frequencies of 20,000 Hz and 40,000 Hz. The subjects could detect all stimuli in the first experiment but not the ultrasounds in the second experiment.We analyzed N1m, the main response with approximately 100 ms in peak latency, and made the following findings. (1) N1m responses to the tones up to 12,000 Hz were clearly recorded from at least one hemisphere in all 12 subjects. N1m for 14,000 Hz was identified in at least one hemisphere in 10 subjects, and in both hemispheres in six subjects. No significant response could be identified to ultrasounds over 20,000 Hz. (2) The amplitude of the N1m to the tones above 8000 Hz was significantly smaller than that to 4000 Hz in both hemispheres. There was a tendency for the peak latency of the N1m to be longer for the tones with higher frequencies, but no significant change was found. (3) The equivalent current dipole (ECD) of the N1m was located in the auditory cortex. There was a tendency for the ECD for the tones with higher frequencies to lie in more medial and posterior areas, but no significant change was found. (4) As for the interhemispheric difference, the N1m amplitude for all frequency tones was significantly larger and the ECDs were estimated to be located more anterior and medial in the right hemisphere than the left. The priority of the right hemisphere, that is the larger amplitude, for very high frequency tones was confirmed. (5) The orientation of the ECD in the left hemisphere became significantly more vertical the higher the tones. This result was consistent with previous studies which revealed the sensitivity of the frequency difference in the left hemisphere.From these findings we suggest that tonotopy in the auditory cortex exists up to the upper limit of audible range within the small area, where the directly air-conducted ultrasounds are not reflected.     

Neurophysiological evidence for cortical discrimination impairment of prosody in Asperger syndrome

Asperger syndrome (AS), belonging to the autism spectrum of disorders, is one of the pervasive developmental disorders. Individuals with AS usually have normal development of formal speech but pronounced problems in perceiving and producing speech prosody. The present study addressed the discrimination of speech prosody in AS by recording the mismatch negativity (MMN) and behavioural responses to natural utterances with different emotional connotations. MMN responses were abnormal in the adults with AS in several ways. In these subjects, fewer significantly elicited MMNs, diminished MMN amplitudes, as well as prolonged latencies were found. In addition, the MMN generator loci differed between the subjects with AS and control subjects. These findings were predominant over the right cerebral hemisphere. These results show impaired neurobiological basis for speech-prosody processing at an early, pre-attentive auditory discrimination stage in AS.     

Event-related potentials and item recognition in depressed, schizophrenic and alcoholic patients

Auditory event-related potentials (ERPs) to pure tones and performance on a measure of item recognition were compared in 20 controls, 14 alcoholics, 20 depressed and 21 schizophrenic patients. Compared with normal controls, P2 and N2 were delayed and of diminished amplitude in the psychopathological groups. Increased amplitude of P1 in alcoholics, diminished N1 in depressed patients, increased latencies of N1 in schizophrenics and N2 in alcoholics were pathology-specific. Unusual patterns of response in the item recognition test (elevated intercept and flattened slope) and its relationship with ERPs distinguished the diagnostic groups from the controls. Support for the preferential involvement of the left hemisphere in schizophrenia and of the right hemisphere in depression was found. Disinhibition of CNS activity in the response of alcoholics (increased P1 and delayed P3) was indicated. The findings suggested that discriminant analysis of auditory ERPs to simple, pure tones, in conjunction with psychometric data significantly differentiated pathologic groups from each other and from controls.     

Maturation processes in automatic change detection as revealed by event-related brain potentials and dipole source localization: significance for adult AD/HD.

Mismatch negativity (MMN) is an event-related potential reflecting automatic attention-related information processing marking the detection of auditory change. The bilateral scalp distribution develops by 14 years of age, and is elicited with adult latencies by 17 years. But consistent with reports of continued brain maturation after adolescence, we show here that features of the temporal and frontal lobe dipole sources also continue to develop in the third decade of life. This has consequences for studies of the developmental course of MMN anomalies, from childhood into adulthood, in attention-deficit/hyperactivity disorder. Two groups of healthy subjects with mean ages of 17 and 30 years were presented with a 3-tone auditory oddball. The duration-deviant MMN was recorded during attention to a visual discrimination (auditory-passive condition) and an active auditory discrimination. MMN amplitudes were smaller in the older subjects and the MMN lasted longer over the right hemisphere. Latencies and moments of the four dipoles in the temporal and frontal lobes did not distinguish the two subject-groups. But both temporal lobe sources were located significantly more ventrally and further left in the young adult than in the adolescent subjects. The left cingular source moved posteriorly and the right inferior frontal source moved antero-medially in the older subjects. Brain development in the third decade may cause the two frontal sources to move apart on the rostro-caudal axis but the temporal lobe sources to move left on the lateral and down on the dorsoventral axes. Thus special care is necessary in interpreting putative dysfunctional neurobiological changes in developmental attention-deficit disorders where as-yet-unspecified sub-groups may show a late developmental lag, partial lag, or no lag at all, associated with other impairments.     

Electroencephalographic and reaction time asymmetries to musical chord stimuli

EEG was recorded over left and right hemispheres at temporal leads (T3, T4) referred to the vertex (Cz) in 16 right-handed male subjects. Musical chords were presented randomly in monaural sequence, during a task which required a selective motor response to stimuli presented to one ear. The integrated amplitude of the 8-13 Hz alpha rhythm was measured when subjects listened passively. Under all conditions, a lower mean alpha amplitude was recorded over the right hemisphere than the left, regardless of which ear was stimulated. Alpha suppression over the right temporal area was accentuated when the performance task directed attention to the stimulus. Reaction time to left ear stimulation was shorter than that for the right ear. With monaural stimulus presentation behavioral asymmetry, and various EEG asymmetries can be observed. There is hemispheric asymmetry associated with attention to task relevant stimuli indicated by reduction in the alpha rhythm over the right temporal area and asymmetry in reaction time with greater efficiency of the left ear to muscial chords.     

Investigation of functional hemispheric asymmetry of language in tinnitus sufferers

The authors tested functional hemispheric asymmetry through word dichotic listening and lateralized lexical decision tasks in tinnitus patients and controls stimulated by a continuous tinnitus-like noise to test the influence of a tinnitus-like external stimulation. A classic right-ear advantage was shown in the auditory task for all but right-ear tinnitus patients, who performed as equally badly when the stimuli were presented to the right and left ears. Concerning the visual task, all participants demonstrated the expected right visual field advantage for word stimuli. Moreover, those who submitted to external stimulation demonstrated normal asymmetric patterns. These data suggest a specific effect of tinnitus on central processing and provide evidence for a functional reorganization induced by this auditory phantom perception.