Attention modulates beta oscillations during prolonged tactile stimulation.

The inter-play between changes in beta-band (14-30-Hz) cortical rhythms and attention during somatosensation informs us about where and when relevant processes occur in the brain. As such, we investigated the effects of attention on somatosensory evoked and induced responses using vibrotactile stimulation and magnetoencephalographic recording. Subjects received trains of vibration at 23 Hz to the right index finger while watching a movie and ignoring the somatosensory stimuli or paying attention to the stimuli to detect a change in the duration of the stimulus. The amplitude of the evoked 23-Hz steady-state response in the contralateral primary somatosensory cortex (SI) was enhanced by attention and the underlying dipole source was located 2 mm more medially, indicating top-down recruitment of additional neuronal populations for the functionally relevant stimulus. Attentional modulation of the somatosensory evoked response indicates facilitation of early processing of the tactile stimulus. Beta-band activity increased after vibration offset in the contralateral primary motor cortex (MI) [event-related synchronization (ERS)] and this increase was larger for attended than ignored stimuli. Beta-band activity decreased in the ipsilateral SI prior to stimulus offset [event-related desynchronization (ERD)] for attended stimuli only. Whereas attention modulation of the evoked response was confined to the contralateral SI, event-related changes of beta-band activity involved contralateral SI-MI and inter-hemispheric SI-SI connections. Modulation of neural activity in such a large sensorimotor network indicates a role for beta activity in higher-order processing.

Correlates of eye blinking as determined by synthetic aperture magnetometry.

OBJECTIVE: To evaluate the spatiotemporal characteristics of ocular and cerebral current sources during voluntary eyeblinking. METHODS: Whole-head magnetoencephalographic (MEG) recordings were acquired during voluntary blinking in eight healthy adults and analysed using synthetic aperture magnetometry (SAM). RESULTS: Fronto-temporal MEG sensors showed a large slow wave lasting approximately 400 ms and a small burst of activity with frequencies above 30 Hz at the initiation of the blink. Group maps of blink-related oscillatory activity at frequencies between 1-18 Hz and 32-64 Hz showed increased activity in and around the orbits during the 400 ms following blink onset. Increased oscillatory activity occurred in occipital regions 200 ms after blink onset at frequencies between 18 and 64 Hz. CONCLUSIONS: Blink-related MEG signals are recorded in the regions of the eyes and in the occipital cortex. The anterior activation is likely a combination of muscle contraction and eyelid currents. Occipital activation likely represents neural processes concerned with re-establishing the visual image after transient ocular occlusion. SIGNIFICANCE: The possibility of eyeblink-related fields should be considered when interpreting frontal and occipital source activities during SAM analyses.

Impaired concentration due to frontal lobe damage from two distinct lesion sites.

BACKGROUND: Investigations of cognitive deficits after frontal lobe damage have commonly relied on multidimensional tests and relatively coarse specification of lesion anatomy. Some form of impairment in attention is often asserted to cause the revealed deficits. OBJECTIVE: To describe a disorder of attention in patients with frontal damage using a theoretical model of the fundamental cognitive processes that underlie attention. METHODS: The ability to perform a task of concentrated responding was studied in 43 patients with well-defined chronic frontal lesions and 38 control subjects using a continuous reaction time (RT) test. Performance measures were mean RT, RT across blocks of the test, and errors. Lesion measures were coarse localization and a hot-spot analysis to detect finer grained lesion effects. RESULTS: Only patients with lesions in the right superomedial (SM) frontal regions had significantly prolonged RT consistently across the entire test. The critical lesion was in Brodmann's areas 24, 32, 9, and 46 days and in the adjacent corpus callosum. Patients with lesions in left lateral frontal (LL) regions made significantly more errors on the 20% of trials in the first block. The critical lesion was in areas 44, 45, and 47/12. CONCLUSION: Concentrating attention to respond is affected by lesions in two different frontal regions for reasons that reflect impairments in different cognitive processes. Right superomedial lesions cause an insufficient energizing of attention to respond. Left lateral lesions cause defective setting of specific stimulus-response contingencies. Constrained tests of attention can demonstrate impairments in specific cognitive operations following lesions to different regions of the frontal lobes.

The effect of attention on the auditory steady-state response.

40-Hz auditory steady state responses to amplitude modulated tones were recorded with magnetoencephalography to investigate the effect of focused attention. A modulation discrimination task and a destructive visual task established the attended and the non-attended experimental conditions. A strong contrast between these conditions was demonstrated by largely enhanced sustained responses under the attention condition. A significant attentional effect on the ASSR amplitude was observed mostly in the left hemisphere between 200 to 500 ms after stimulus onset. In contrast, transient gamma-band and N1 responses were not affected by the different states of attention.

Detection of power changes between conditions using split-half resampling of synthetic aperture magnetometry data.

Synthetic Aperture Magnetometry (SAM) measures changes in task-related power using pseudo-t values which are affected by changes in both signal and noise. Detecting significant signal power changes between two separate experimental conditions should not be done directly due to possible fluctuation in the noise as well as the response. This study proposes a method to estimate the noise within a single condition, which is then used to test the null hypothesis of no difference between the conditions. The noise estimation is based on a split-half resampling technique. For each resampling, the data of a given condition is divided into two halves. The difference of the pseudo-t volumes between the pair of the datasets is calculated. After multiple resamplings, the confidence limits of the differences within this single condition are computed for a given p-value so that one can test the null hypotheses that the second condition is within the same distribution as the first. The limits are calculated using a bootstrap technique to correct for any bias in the estimated threshold. Power changes between the two conditions are considered significantly different if the difference of the pseudo-t value is larger than expected within conditions. To demonstrate the effectiveness of the technique, the proposed method was applied to MEG responses to two distinct visual stimuli recorded from a single subject. Major differences of brain activity between the two conditions were found in the occipital region. These results were validated using four pairs of split-half datasets, generated from either the odd or even trials in each condition. The method of split-half resampling should therefore be useful for localizing significant differences in brain activity between conditions within individual subjects.

Cortical oscillations modulated by congruent and incongruent audiovisual stimuli.

Congruent or incongruent grapheme-phoneme stimuli are easily perceived as one or two linguistic objects. The main objective of this study was to investigate the changes in cortical oscillations that reflect the processing of congruent and incongruent audiovisual stimuli. Graphemes were Japanese Hiragana characters for four different vowels (/a/, /o/, /u/, and /i/). They were presented simultaneously with their corresponding phonemes (congruent) or non-corresponding phonemes (incongruent) to native-speaking Japanese participants. Participants' reaction times to the congruent audiovisual stimuli were significantly faster by 57 ms as compared to reaction times to incongruent stimuli. We recorded the brain responses for each condition using a whole-head magnetoencephalograph (MEG). A novel approach to analysing MEG data, called synthetic aperture magnetometry (SAM), was used to identify event-related changes in cortical oscillations involved in audiovisual processing. The SAM contrast between congruent and incongruent responses revealed greater event-related desynchonization (8-16 Hz) bilaterally in the occipital lobes and greater event-related synchronization (4-8 Hz) in the left transverse temporal gyrus. Results from this study further support the concept of interactions between the auditory and visual sensory cortices in multi-sensory processing of audiovisual objects.

Efficient stimuli for evoking auditory steady-state responses.

OBJECTIVE: To compare the magnitudes of the steady-state responses evoked by several types of stimuli, and the times required to recognize these responses as significant. DESIGN: In the first two experiments, we examined auditory steady-state responses to pure tones, broadband noise and band-limited noise. The stimuli were amplitude modulated in the 75 to 100 Hz range with sinusoidal or exponential envelopes. A third experiment investigated the effects of exponential envelopes on the responses to broadband noise. The final experiment examined auditory steady-state responses evoked by rapidly presented transient stimuli, such as clicks, brief tones and brief noise-bursts. All stimuli were presented dichotically at intensities 30 to 50 dB above behavioral thresholds. The subjects were adults, who drowsed or slept during the recording sessions. RESULTS: The responses to the noise were larger than the responses to the tones. At an intensity of 32 dB nHL, the average amount of time needed to obtain significant responses for the amplitude-modulated noise was 43 sec and the maximum time was 2 minutes. The average time for pure tone stimuli was approximately 2 minutes but 25% of the responses remained undetected after 5 minutes. Combining the responses to all the frequency-specific stimuli showed results similar to using noise stimuli. Using exponential envelopes did not increase response amplitudes for noise stimuli. At 45 dB nHL, the steady-state responses to clicks and other transient stimuli were larger than responses to the broadband noise. The average time to detect steady-state responses to transient stimuli was approximately 20 sec, which was a little faster than for amplitude modulated noise. CONCLUSIONS: Auditory steady-state potentials evoked by amplitude modulated noise or transient stimuli might be useful in providing rapid and objective tests of hearing during screening procedures. Another approach might be to record responses to multiple frequency-specific stimuli and to evaluate the combined responses for a rapid indication that some hearing is present.

Bottom-up and top-down influences on auditory scene analysis: evidence from event-related brain potentials.

The physiological processes underlying the segregation of concurrent sounds were investigated through the use of event-related brain potentials. The stimuli were complex sounds containing multiple harmonics, one of which could be mistuned so that it was no longer an integer multiple of the fundamental. Perception of concurrent auditory objects increased with degree of mistuning and was accompanied by negative and positive waves that peaked at 180 and 400 ms poststimulus, respectively. The negative wave, referred to as object-related negativity, was present during passive listening, but the positive wave was not. These findings indicate bottom-up and top-down influences during auditory scene analysis. Brain electrical source analyses showed that distinguishing simultaneous auditory objects involved a widely distributed neural network that included auditory cortices, the medial temporal lobe, and posterior association cortices.

The use of phase in the detection of auditory steady-state responses.

OBJECTIVE: To investigate how phase measurements might facilitate the detection of auditory steady-state responses. METHODS: Multiple steady-state responses were evoked by auditory stimuli modulated at rates between 78 and 95 Hz and with intensities between 50 and 0 dB SPL. The responses were evaluated in 20 subjects after 1, 2, 4, and 6 min. The responses were analyzed in the frequency domain using 4 different detection protocols: (1) phase-coherence, (2) phase-weighted coherence, (3) F test for hidden periodicity, and (4) phase-weighted t test. The phase-weighted measurements were either based on the mean phase of a group of normal subjects or derived for each subject from the phase of the response at higher intensities. RESULTS: Detection protocols based on both phase and amplitude (F test and phase-weighted t test) were more effective than those based on phase alone (phase coherence and phase-weighted coherence) although the difference was small. Protocols using phase-weighting were more effective than those without phase-weighting. The lowest thresholds for the steady-state responses were obtained using the phase-weighted t test. CONCLUSION: Threshold detection can be improved by weighting the detection protocols toward an expected phase, provided that the expected phase can be reliably predicted.

Comparative electrophysiological and hemodynamic measures of neural activation during memory-retrieval.

The spatial and temporal characteristics of the brain processes underlying memory retrieval were studied with both event-related potentials (ERP) and positron emission tomography (PET) techniques. Subjects studied lists of 20 words and then performed episodic (old/new judgment) or semantic (living/nonliving decision) retrieval tasks on multiple four-item test lists, each lasting 10 sec. The PET and ERP measurements at test were assessed in relation to both the task (episodic vs. semantic) and the item (old vs. new or living vs. nonliving). Episodic retrieval was associated with increased blood flow in the right frontal lobe (Brodmann Area 10) and a sustained, slowly developing positive ERP shift recorded from the right frontopolar scalp. Semantic retrieval was associated with increased blood flow in the left frontal (Area 45) and temporal (Area 21) lobes but no clear ERP concomitant. The two retrieval tasks also differed from each other in the ERPs to single items in an early (300-500 ms) time window. Item-related comparisons yielded convergent results mainly if the retrieved information was relevant to the given task (e.g., old/new items during episodic retrieval and living/nonliving items during semantic retrieval). Episodically retrieved old items were associated with increased blood flow in the left medial temporal lobe and a transient increase in the amplitude of the late positive component (500-700 ms) of the ERP. Semantically retrieved living items were associated with increased blood flow in the left frontal cortex and anterior cingulate and a transient late frontal slow wave (700-1,500 ms) in the ERPs. These results indicate that the brain regions engaged in memory retrieval are active in either a sustained or transient manner. They map task-related processes to sustained and item-related processes to transient neural activity. But they also suggest that task-related factors can transiently affect early stages of item processing.

Human auditory steady-state responses to tones independently modulated in both frequency and amplitude.

OBJECTIVE: Independent amplitude and frequency modulation (IAFM) of a carrier tone uses two different modulating frequencies, one for amplitude modulation (AM) and one for frequency modulation (FM). This study measured the human steady-state responses to multiple IAFM tones. The first question was whether the IAFM responses could be recorded without attenuation of the AM and FM components. The second question was whether IAFM stimuli would provide a more effective demonstration of responses at intensities near threshold than the responses to AM tones. The third question was whether the responses to multiple IAFM stimuli would relate to the discrimination of words at different intensities. DESIGN: Multiple AM, FM, or IAFM stimuli were presented simultaneously. Responses were recorded between the vertex and the neck and analysed in the frequency domain. The first experiment compared IAFM responses with AM and FM responses. The second experiment compared IAFM responses with AM responses between intensities 20 to 50 dB SPL. The third experiment related the IAFM responses to the discrimination of monosyllabic words at intensities between 20 and 70 dB SPL. RESULTS: Steady-state responses to the individual component of the IAFM stimuli were clearly recognizable although attenuated a little (14%) from the responses to AM or FM alone. Using IAFM stimuli was not different than simply using AM stimuli when trying to recognize responses at low intensities. The number of responses detected during multiple IAFM stimulation and the amplitudes of these responses correlated significantly with word discrimination. CONCLUSIONS: IAFM of a carrier using two different modulating frequencies (one for AM and one for FM) elicits separate AM and FM responses that are relatively independent of each other. These separate responses can be used to detect whether a particular carrier has been processed in the cochlea, but they are not as effective as measuring responses to carriers that have been modulated in both amplitude and frequency at the same modulation frequency (mixed modulation). The detectability of eight different responses (four AM and four FM) to an IAFM stimuli relates well to the ability of subjects to discriminate words. IAFM stimuli therefore show promise as an objective test for assessing suprathreshold hearing.

Attention and successful episodic encoding: an event-related potential study.

Event-related potentials (ERPs) were used to delineate the cerebral processes occurring when information is encoded into episodic memory and to determine how these processes are affected by divided attention. ERPs were recorded during encoding under focused or divided attention, and were selectively averaged on the basis of their retrieval during later free recall and recognition tests (with remember-know judgments). Items retrieved with conscious recollection of the encoding episode (remembered, recalled) were distinguished at encoding from later missed items by an enhanced left fronto-temporal negative wave (N340), a negative posterior sustained potential and a positive frontal sustained potential. These effects occurred independently of the level of attention. Items later retrieved on the basis of familiarity (known) elicited a larger N340 than missed items, but did not demonstrate the increased sustained potentials. We suggest that item-specific conceptual processing (N340) is sufficient to produce familiarity-based recognition, but additional elaborative processing (sustained interaction of frontal and posterior regions) is necessary for conscious recollection. The effect of divided attention on these processes was related to the difficulty of the secondary task, with the more difficult task causing greater and earlier interference.

Multiple auditory steady-state responses to AM and FM stimuli.

Multiple auditory steady-state responses were recorded using tonal stimuli that were amplitude-modulated (AM), frequency-modulated (FM) or modulated simultaneously in both amplitude and frequency (mixed modulation or MM). When MM stimuli combined 100% AM and 25% FM (12.5% above and below the carrier frequency) and the maximum frequency occurred simultaneously with maximum amplitude, the MM response was one third larger than the simple AM response. This enhancement occurred at intensities between 50 and 30 dB SPL and at carrier frequencies between 500 and 4000 Hz. The AM and FM components of a MM stimulus generate independent responses that add together to give the MM response. Since AM responses generally occur with a slightly later phase delay than FM responses, the largest MM response is recorded when the maximum frequency of the MM stimulus occurs just after the maximum amplitude.

Weighted averaging of steady-state responses.

OBJECTIVE: To compare weighted averaging and artifact-rejection to normal averaging in the detection of steady-state responses. METHODS: Multiple steady-state responses were evoked by auditory stimuli modulated at rates between 78 and 95 Hz. The responses were evaluated after recording periods of 3, 6 and 10 min, using 5 averaging protocols: (1) normal averaging; (2) sample-weighted averaging; (3) noise-weighted averaging; (4) amplitude-based artifact-rejection; and (5) percentage-based artifact rejection. The responses were analyzed in the frequency domain and the signal-to-noise ratio was estimated by comparing the signals at the modulation-frequencies to the noise at adjacent frequencies. RESULTS: Weighted averaging gave the best signal-to-noise ratios. Artifact-rejection was better than normal averaging but not as good as weighted averaging. Responses that were not significant with normal averaging became significant with weighted averaging much more frequently than vice versa. False alarm rates did not significantly differ among the protocols. The advantage of weighted averaging was especially evident when stimuli were presented at lower intensities or when smaller amounts (e.g. only 3 or 6 min) of data were evaluated. Weighted averaging was most effective when the background noise levels were variable. Weighted averaging underestimated the amplitude of the responses by about 2%. CONCLUSION: Weighted averaging should be used instead of normal averaging for detecting steady-state responses.

Fast temporal interactions in human auditory cortex.

The temporal resolution of the human primary auditory cortex (AC) was studied using middle-latency evoked fields. Paired sounds with either the same or different spectral characteristics were presented with gaps between the sounds of 1, 4, 8 and 14 ms. Spatio-temporal modelling showed (1) that the response to the second sound was recognizable with gaps of 1 ms and rapidly increased in amplitude with increasing gap durations, (2) an enhanced N40m amplitude at gaps > 4 ms, (3) delayed N19m-P30m latencies when the stimuli were different. The median psychoacoustical thresholds were 1.6 ms for the same stimuli and 2.5 ms for different stimuli, confirming the electrophysiological evidence for rapid pattern-specific temporal processing in human primary auditory cortex.

Event-related potentials during conscious and automatic memory retrieval.

The effects of study-test lags of between 0 and 32 items on conscious (C) and automatic (A) memory processes in a running word-completion task were investigated with event-related potentials (ERPs). The process dissociation procedure (PDP) can distinguish between C and A contributions to memory by comparing performance when subjects respond with either an old item (inclusion) or a new item (exclusion). C can be estimated by subtracting the probability of an intrusion of an old item during the exclusion task (due to A without C) from the probability of correctly producing an old item during the inclusion task (due to C and/or A). The behavioral results showed that C was stronger when the test item followed the studied word in the next trial or after a lag of one stimulus. The strength of A did not vary with lag. The ERP waveforms contained a broad parietal positive wave between 300 and 800 ms. This parietal wave distinguished between correctly recalled old and new words. The early portion of this old-new effect was significantly affected by lag. Subtracting waveforms to obtain a measure of C revealed an effect in the later portion of this wave, lateralized over the left hemisphere. A sustained frontal negativity occurred during all recordings and was larger during conscious retrieval. There was no consistent ERP effect related to automatic memory retrieval.

Mismatch negativity: different water in the same river.

The mismatch negativity (MMN) is a frontal negative deflection in the human event-related potential that typically occurs when a repeating auditory stimulus changes in some manner. The MMN can be elicited by many kinds of stimulus change, varying from simple changes in a single stimulus feature to abstract changes in the relationship between stimuli. The main intracerebral sources for the MMN are located in the auditory cortices of the temporal lobe. Since it occurs whether or not stimuli are being attended, the MMN represents an automatic cerebral process for detecting change. The MMN is clinically helpful in terms of demonstrating disordered sensory processing or disordered memory in groups of patients. Improvements in the techniques for measuring the MMN and in the paradigms for eliciting it will be needed before the MMN can become clinically useful as an objective measurement of such disorders in individual patients.

Effects of visual attentional load on auditory processing.

Auditory evoked potentials were recorded while participants attended to visually presented digits. The difficulty of the visual task was manipulated by requiring participants to process only the current digit (0-back) or both the current and the preceding digit (1-back). Tones deviating in frequency from standard tones elicited a frontal mismatch negativity peaking around 200 ms which did not vary with visual task. However, decreasing the visual task load enhanced a right-temporal positive wave peaking around 200 ms when tones were presented slowly, and a frontocentral negative wave peaking around 450 ms when tones were presented more rapidly. The degree to which task-irrelevant sounds are processed therefore depends on the degree to which a visual task engages attentional resources.

Guidelines for using human event-related potentials to study cognition: recording standards and publication criteria.

Event-related potentials (ERPs) recorded from the human scalp can provide important information about how the human brain normally processes information and about how this processing may go awry in neurological or psychiatric disorders. Scientists using or studying ERPs must strive to overcome the many technical problems that can occur in the recording and analysis of these potentials. The methods and the results of these ERP studies must be published in a way that allows other scientists to understand exactly what was done so that they can, if necessary, replicate the experiments. The data must then be analyzed and presented in a way that allows different studies to be compared readily. This paper presents guidelines for recording ERPs and criteria for publishing the results.

Human auditory steady-state responses to amplitude-modulated tones: phase and latency measurements.

Human auditory steady-state responses were recorded to four stimuli, with carrier frequencies (f(c)) of 750, 1500, 3000 and 6000 Hz, presented simultaneously at 60 dB SPL. Each carrier frequency was modulated by a specific modulation frequency (f(m)) of 80.6, 85.5, 90.3 or 95.2 Hz. By using four different recording conditions we obtained responses for all permutations of f(m) and f(c). The phase delays (P) of the responses were unwrapped and converted to latency (L) using the equation: L=P/(360xf(m)). The number of cycles of the stimulus that occurred prior to the recorded response was estimated by analyzing the effect of modulation frequency on the responses. These calculations provided latencies of 20.7, 17.7, 16.1 and 16.1 ms for carrier frequencies 750, 1500, 3000 and 6000 Hz. This latency difference of about 4.5 ms between low and high carrier frequencies remained constant over many different manipulations of the stimuli: faster modulation rates (150-190 Hz), binaural rather than monaural presentation, different intensities, stimuli presented alone or in conjunction with other stimuli, and modulation frequencies that were separated by as little as 0.24 Hz. This frequency-related delay is greater than that measured using transient evoked potentials, most likely because of differences in how transient and steady-state responses are generated and how their latencies are determined.