Somatosensory event-related potentials in the rabbit cerebral and cerebellar cortices: a correspondence with mismatch responses in humans

Somatosensory event-related potentials (ERPs) were recorded from the cerebellar (CerCx), somatosensory (SomCx) and visual (VCx) cortices in rabbits in two stimulus conditions. In the oddball condition, airpuffs to two different locations in the rabbit's muzzle corresponded to infrequently presented deviant stimuli (oddball-deviants) interspersed with frequently presented standard stimuli. In the deviant-alone condition, deviants (alone-deviants) were presented without standards. ERPs to oddball-deviants differed significantly from those to standards in CerCx and SomCx, but not in VCx. Furthermore, some of these differences were not found between ERPs to alone-deviants and those to standards. Thus, ERPs to oddball-deviants contributing these differences were dependent on the presence of preceding standards. The results are in line with the elicitation of mismatch negativity-like ERPs to somatosensory changes in humans.     

Attentional modulation of the mismatch negativity elicited by frequency differences between binaurally presented tone bursts

We examined the attentional sensitivity of the frequency-change mismatch negativity (MMN). Subjects listened to a binaural mixture of a narrative and a series of tone bursts that included 1200-Hz standards and two deviants (1000 and 1400 Hz). In the attend-tones condition, subjects responded to one deviant and ignored the narrative. In the attend-words condition, subjects responded to target words in the narrative and ignored the tones. Event-related potentials (ERPs) were recorded for the tones, and differences waves (deviant ERPs minus standard ERPs) were computed. Two negative peaks in the difference waves, DNI (100–180 ms) and DN2 (200–300 ms), overlapped the known scalp distribution and latency of the MMN. Mean DN1 and DN2 amplitudes were greater in the attend-tones condition than in the attend-words condition. These data suggest that the frequency-change MMN is modulated by nonspatial shifts of auditory attention.     

Event-related potentials to irrelevant deviant motion of visual shapes.

Event-related potentials to visual shapes moving across the visual field were recorded from 10 subjects. The subjects had to respond to the appearance of one of the shapes, while other shapes were irrelevant. On the periphery some of the shapes changed their orientation or their form. Sometimes the direction of movement was different from the standard direction. Subjects did not detect the changes of the pattern on the visual periphery, and the ERPs to these non-detected deviants were identical to the ERPs to the standard stimuli. Six subjects detected the irrelevant direction of movement. In these subjects the deviant direction of motion elicited a fronto-central positive wave (P3s) with 322 ms mean latency. There was no such sharp positive peak in the four subjects who did not detect the deviant direction of movement. Unlike the non-target stimuli, the targets elicited a large positive wave (P3b) with 530 ms mean latency.     

Processing of auditory stimuli during auditory and visual attention as revealed by event-related potentials

Auditory event-related brain potentials (ERPs) were recorded during auditory and visual selective attention tasks. Auditory stimuli consisted of frequent standard tones (1000 Hz) and infrequent deviant tones (1050 Hz and 1300 Hz) delivered randomly to the left and right ears. Visual stimuli were vertical line gratings randomly presented on a video monitor at mean intervals of 6 s. During auditory attention, the subject attended to the stimuli in a designated ear and responded to the 1300-Hz deviants occurring among the attended tones. During visual attention, the subject responded to the occasional visual stimuli. ERPs for tones delivered to the attended ear were negatively displaced relative to ERPs elicited by tones delivered to the unattended ear and to ERPs elicited by auditory stimuli during visual attention. This attention effect consisted of negative difference waves with early and late components. Mismatch negativities (MMNs) were elicited by 1300-Hz and 1050-Hz deviants irrespective of whether they occurred among attended or unattended tones. MMN amplitudes were unaffected by attention, supporting the proposal that the MMN is generated by an automatic cerebral discrimination process.     

"Initial-" and "change-orienting reactions": an analysis based on visual single-trial event-related potentials

The present study was concerned with brain potentials elicited by, respectively, the first of a series of stimuli ("initial-orienting reaction", I-OR), and infrequent deviants ("change-orienting reaction", C-OR). Single-trial event-related potentials (ERPs) to visual stimuli were estimated from recordings at Oz, Pz, Cz and Fz. The design included both a habituation series as well as a series of occasional deviant trials against a background of standards. This was done with both task-relevant and neutral stimuli, and in two interstimulus interval (ISI) conditions: 2.45 s and 8.45 s. In the latter ISI condition, skin conductance responses (SCRs) were recorded as well. Decrease (Habituation) in the habituation series was found for a non-specific N1, a posteriorly distributed P3, and the SCR, but not for P2-N2. Deviant stimuli produced an enhancement of the central P2-N2, the P3, the N1 (on the first few deviant trials only, in both ISI conditions), and the SCR (with task-relevant stimuli only). Elongation of ISI delayed both short-term and long-term decrease of P3, but had no effect on enhancement of P2-N2 due to stimulus deviance. It was concluded that, with respect to ERP parameters, the I-OR is marked by the N1, whereas the C-OR coincides with the P2-N2.     

Brain activity from stimuli that are not perceived: Visual mismatch negativity during binocular rivalry suppression

Predictive coding explains visual perception as the result of an interaction between bottom‐up sensory input and top‐down generative models at each level of the visual hierarchy. Evidence for this comes from the visual mismatch negativity (vMMN): a more negative ERP for rare, unpredictable visual stimuli—deviants, than for frequent, predictable visual stimuli—standards. Here, we show that the vMMN does not require conscious experience. We measured the vMMN from monocular luminance‐decrement deviants that were either perceived or not during binocular rivalry dominance or suppression, respectively. We found that both sorts of deviants elicited the vMMN at about 250 ms after stimulus onset, with perceived deviants eliciting a bigger vMMN than not‐perceived deviants. These results show that vMMN occurs in the absence of consciousness, and that consciousness enhances the processing underlying vMMN. We conclude that generative models of visual perception are tested, even when sensory input for those models is not perceived.     

Cortical Activity Elicited by Changes in Auditory Stimuli: Different Sources for the Magnetic N1OOm and Mismatch Responses

Magnetic responses to frequent and infrequent auditory stimuli, all presented in the same stimulus block in randomized order, were recorded. The standard stimuli, comprising 90% of all the stimuli, were 100-ms, 1000 Hz, 90dB sinusoidal tone bursts. There were three deviant tones, each presented at a probability of 3.3%, which differed from the standard tone on one dimension only: frequency deviant (1500 Hz), intensity deviant (67dB SPL), or duration deviant (50 ms). All mismatch fields, i.e., responses elicited by different deviants, as well as N100m to the standards and deviants, could be explained by neural activity in the supratemporal auditory cortex. The source of N100m to standards and deviants was significantly posterior to the sources for the three different mismatch fields. The mean locations of the equivalent dipoles for the different mismatch fields did not differ significantly from each other, but some differences were found for individual subjects.     

Can Echoic Memory Store Two Traces Simultaneously? A Study of Event-Related Brain Potentials

The mismatch negativity, a component of the event-related brain potential elicited by infrequent deviants in sequences of auditory stimuli, is presumably generated by an automatic mismatch process in a mechanism that compares the current stimulus to the trace of the previous one. The present study addressed the possible simultaneous existence of two such traces. Two equiprobable (45% each) frequent stimuli (“standards”), one of 600 Hz and the other of 700 Hz, were presented together with an infrequent (10%), “deviant” stimulus which was of differnet frequency in different blocks. These deviants elicited a mismatch negativity, though a smaller one than that obtained in corresponding blocks with only one standard stimulus. Two aspects of the present results from the blocks with two standard stimuli implicate two parallel stimulus traces in these blocks: 1) deviants elicited a mismatch negativity (MMN) of approximately the same amplitude when preceded by sequences of four identical standards as when preceded by sequences of four stimuli containing both standards; 2) in contrast to the one-standard condition, the magnitude of stimulus deviance did not affect the MMN component elicited by the different deviants.     

The quest for the genuine visual mismatch negativity (vMMN): Event-related potential indications of deviance detection for low-level visual features

Research shows that the visual system monitors the environment for changes. For example, a left-tilted bar, a deviant, that appears after several presentations of a right-tilted bar, standards, elicits a classic visual mismatch negativity (vMMN): greater negativity for deviants than standards in event-related potentials (ERPs) between 100 and 300 ms after onset of the deviant. The classic vMMN is contributed to by adaptation; it can be distinguished from the genuine vMMN that, through use of control conditions, compares standards and deviants that are equally adapted and physically identical. To determine whether the vMMN follows similar principles to the auditory mismatch negativity (MMN), in two experiments we searched for a genuine vMMN from simple, physiologically plausible stimuli that change in fundamental dimensions: orientation, contrast, phase, and spatial frequency. We carefully controlled for attention and eye movements. We found no evidence for the genuine vMMN, despite adequate statistical power. We conclude that either the genuine vMMN is a rather unstable phenomenon that depends on still-to-be-identified experimental parameters, or it is confined to visual stimuli for which monitoring across time is more natural than monitoring over space, such as for high-level features. We also observed an early deviant-related positivity that we propose might reflect earlier predictive processing.     

Short-Term Habituation and Dishabituation of the Mismatch Negativity of the ERP

Standard tones of 1000 Hz and deviant tones of 1250 Hz were presented in random order, 1 stimulus/second. The probabilities of the standards and deviants were 90% and 10%, respectively. In one condition the subject counted the deviant stimuli and in the other condition he/she read a comic book. ERPs were separately averaged to 1) the standard preceding the deviant, 2) the “first deviant” preceded by at least 4 standards, 3) the “second deviant” (an occasional deviant immediately following the “first deviant”), 4) the first and 5) the second standard following the “first deviant,” 6) the first and 7) the second standard following the “second deviant.” It was found that the mismatch negativity evoked in both conditions by the first deviant was considerably larger than to the second deviant. Also the first standard following the first deviant evoked the mismatch negativity. The results are discussed in terms of parallel neuronal models of the stimuli as reflected by the mismatch negativity.     

Interstimulus interval and the selective-attention effect on auditory ERPs:

The attention effect on the auditory event-related potential (ERP) in dichotic conditions was studied as a function of the interstimulus interval (ISI). Subjects attended to stimuli delivered to a designated ear and responded to infrequent pitch deviants in this input. The mean ISI was either 80, 160, 480, or 800 ms. Negative difference waves (Nds) were computed by subtracting ERPs to unattended standards from ERPs to the same stimuli when attended. The exogenous Nl, as estimated from unattended standard ERPs, was larger contralaterally to the stimulus and inverted in polarity at mastoids. With decreasing ISIs, Nl diminished in amplitude much faster than did the Nd. In addition, Nl latency remained stable, whereas Nd peaked markedly earlier with shorter ISIs, almost perfectly coinciding with the exogenous Nl. However, this temporal coincidence found in grand averages proved to be illusory in single subjects. The early Nd showed no contralateral asymmetry at its peak, but asymmetry was apparent during the ascending slope. These lateral asymmetries resembled those of the exogenous Nl but occurred later. The early Nd peak was, at least mainly, caused by an endogenous attention effect, the processing negativity (PN), even with very short ISIs, but an effect on the exogenous Nl could not be excluded.     

Contra- and ipsilateral auditory stimuli produce different activation patterns at the human auditory cortex

Auditory evoked magnetic fields were recorded over the right hemisphere of healthy humans The stimuli were noise bursts presented either to the contra- (C) or ipsilateral (I) ear in different combinations. The largest deflection of the responses, N100m (magnetic counterpart of electric N100), showed a field pattern which suggests activation of the supratemporal auditory cortex. In an oddball paradigm, where the standards (90%) were 400-ms noise bursts presented to the contralateral ear, and the deviants (10%) similar stimuli to the ipsilateral ear, the deviants elicited on the average 130% stronger equivalent dipoles for N100m than standards. Contralateral standards did not substantially decrease the response amplitude of ipsilateral deviants as compared with the response amplitude to ipsilateral stimuli alone presented at the interstimulus interval of the deviants. When two 50 ms noise bursts, separated by 310 ms, were presented once every 2 s, N100m evoked by the second stimulus of the pair was smaller when the stimuli were presented monaurally (C-C, or I-I) than to different ears (I-C or C-I). The results suggest that contra- and ipsilateral auditory stimuli are analyzed, at least in part, in different neural networks at the human auditory cortex.     

Processing of auditory deviants with changes in one versus two stimulus dimensions

Auditory event-related potentials (ERPs) in response to 50-ms tones were recorded from the human scalp. A standard stimulus ( p = .88) and three different deviants were randomly presented via earphones. There were two one-dimensional deviants (one frequency and one location deviant) and one two-dimensional deviant, with changes in both frequency and location. In one condition, subjects read a book and ignored the auditory stimuli, whereas in another condition they tried to discriminate deviants from standards. In the ignore condition, the two-dimensional deviant elicited an enhanced mismatch negativity (MMN) as compared with the MMNs elicited by the one-dimensional deviants. The temporal and the topographic distributions of the two-dimensional MMNs could be modeled by adding the one-dimensional MMNs. This additivity of the MMNs probably results from the independent activity of separate neural populations generating the frequency and the location MMN. In the attend condition, the deviance-related ERP effects were not additive in the N2b and P3 range, implicating that the neural processes involved in the conscious detection of changes in location and frequency were not independent.     

Visual stimulus change and the orienting reaction: event-related potential evidence for a two-stage process.

In a previous study it was found that infrequent deviant visual stimuli, in a series of standards, elicited event-related potentials (ERPs) with enhanced P2-N2s and P3 amplitudes, suggesting that these parameters reflect processes related to the orienting reaction (OR). In the present study a similar oddball series was presented against the background of a second class of stimuli. With respect to the latter stimuli, subjects had to perform either a very involved (hard) or an easy task. EEG was recorded to oddball (probe) stimuli from Oz, Pz, Cz, and Fz. Analysis of average ERPs revealed that, in the easy condition, deviant probes elicited both enhanced P2-N2s and enhanced P3s, relative to the standards. In contrast, in the hard condition P2-N2, but not P3, was enhanced by stimulus change. In addition, overall P3 amplitude to probes was smaller in the hard condition (sequence-independent load effect). Analysis of single-trial ERPs (SERPs) with orthogonal polynomial trend analysis largely replicated these effects. In addition, SERP analysis also revealed a sequence-independent load effect on P2, as well as a decreasing P3 to deviant stimuli in the Easy condition, which was observed at Cz and Fz, but not at Pz or Oz. The results are interpreted as suggesting that P2-N2 and P3 reflect different stages of the OR, one of automatic and one of capacity-limited processing.     

The Effects of Spatial Selective Attention on the Somatosensory Event-Related Potential

Event-related potentials (ERPs) were recorded during a selective attention task involving weak or strong electrical stimuli delivered to the index fingers of the left and right hands. In an attend weak condition, subjects were asked to count the number of weak stimuli (targets) interspersed amongst strong stimuli (standards) delivered to a designated hand, whilst ignoring a similar set of stimuli delivered to the other hand. In an attend strong condition, subjects were asked to count the number of strong targets interspersed amongst weak stimuli. In both conditions, targets and standards occurred with probabilities of .10 and .40 respectively on each hand. Counting weak targets was found to be more difficult than counting strong targets. The latency of the earliest significant effect of selective attention on ERPs to standards was dependent on stimulus intensity: N80 in the case of weak standards, P105 for strong standards. There was no evidence of a later prolonged negative shift in attended standard ERPs. Rather, an enhanced N150 component post-centrally was followed by a prolonged positive shift of attended standard ERPs. This Late positive shift had a similar scalp distribution to the late positive component elicited by attended target stimuli.     

Auditory processing in an inter-modal oddball task: effects of a combined auditory/visual standard on auditory target ERPs.

Previous research examining auditory target ERPs in an inter-modal oddball task has shown that early components are affected by intra-modal processes, whilst components in the later part of the ERP, around 200-400 ms, are affected by both inter-modal and intra-modal processes. These findings led to the conclusion that there are separate stages of auditory target processing--an early modality-specific stage and a later context-dependent stage. The present study investigated this further by simultaneously presenting a visual standard stimulus with an auditory standard stimulus in an oddball task. The aim was to determine whether the inclusion of the visual standard stimulus in this task affected the ERP to targets. The auditory-visual oddball task consisted of a regularly presented combined auditory and visual standard stimulus (80%) and an infrequent auditory target (20%). The ERPs to targets in the auditory-visual oddball task were compared to those in an auditory oddball task, which had identical auditory stimuli and no visual standards. The results showed that the early components N100, P200 and N200 did not differ between tasks. This was in line with earlier results, and confirmed that activity up to 200 ms is unaffected by visual standard stimuli. The later components P250, P300 and P350 were larger and showed topographic differences in the auditory-visual oddball task. This was interpreted as reflecting separate inter-modal and intra-modal processes at later stages. In particular, the P300 and P350 components were argued to represent separate inter-modal and intra-modal components. Overall, this study provides further evidence of auditory processing occurring in two stages, an early modality-dependent stage and a later context-dependent stage.     

Mismatch negativity for item rather than serial-order information in a 150-ms tone series that is not repeated as a melodic pattern

Speech, for example, consists of fast-paced (>5/s) sounds in specific spectrotemporal patterns. Humans are generally held to be able to represent not only such sounds themselves (item information) but also their serial order (order-information) as a repeated melody with ease, as suggested by studies on the mismatch negativity (MMN) of event-related potentials (ERPs). The present study tested whether this ability tolerates the absence of the support of melodic repetitiveness. ERPs were recorded from adult humans presented with rare 150-ms series of three 50-ms tones ('deviants') interspersed with frequently repeated ones ('standards'). The frequency of each tone was pseudorandomly one of four alternative frequencies. The series were of type 'AAB' (two tones of one frequency followed by a tone of another frequency), 'ABB' (two tones of one frequency preceded by a tone of another frequency), or 'AAA' (three tones of one frequency). The MMN was robustly elicited by AAA deviants against AAB standards. It was, however, less distinct for ABB deviants against AAB standards and even statistically non-significant for AAB deviants against ABB standards. MMN generation in the human brain thus seems to be based on item rather than serial-order information in a rapid spectro-temporal pattern of acoustic signals that is not repeated frequently in the short term.     

Neurodynamic Studies on Emotional and Inverted Faces in an Oddball Paradigm

The detection of a change in a face stimulus was studied in an oddball paradigm. Event-related potentials (ERPs) and MEG responses to face stimuli were recorded in four conditions: 1) happy standard, neutral deviant; 2) neutral standard, neutral deviant; 3) inverted happy standard, inverted neutral deviant; 4) inverted neutral standard, inverted neutral deviant. In all conditions, the target was a face with glasses. Neutral deviants elicited a negative deflection (with a maximum around 280 ms) in ERP and MEG responses, an effect similar to auditory mismatch negativity. Face inversion diminished deviance-related negativity, implying an important role of face recognition in the observed effect. Emotional content and larger physical differences between stimuli in conditions 1 and 3 compared to conditions 2 and 4 did not show statistically significant effect on the neutral-deviant-related negativity.     

Backward masking and visual mismatch negativity: electrophysiological evidence for memory-based detection of deviant stimuli

Sequences composed of two different colored checkerboard patterns (standard and deviant) were presented to adults. Each pattern was followed by a mask with stimulus onset asynchronies (SOAs) varying between 14 and 174 ms. ERPs were recorded to the deviant and standard stimuli while the participants detected changes of a cross, which was continuously present at the center of the screen. In further experiments, the participants performed a Go-NoGo task detecting the deviant checkerboards. Deviant stimuli elicited an occipital negative component with 124-132 ms mean latency (the visual mismatch negativity, vMMN) at test (standard or deviant)-to-mask SOAs longer than 27 ms. No vMMN amplitude increase was observed beyond 40 ms test-to-mask intervals, whereas detection of deviant checkerboard patterns improved up to 174-ms SOA. Therefore the processes underlying vMMN elicitation cannot fully explain the overt detection of visual deviance.     

P300 and slow wave from oddball and single-stimulus visual tasks: inter-stimulus interval effects.

The effects of inter-stimulus intervals on P300 from an oddball task (target and standard stimuli) and a single-stimulus task (targets only) employing simple visual stimuli were assessed in order to determine how a relatively long ISI affects event-related brain potentials (ERPs). Young adult subjects (n=16) responded by pressing a button to a visual target stimulus of each task condition. ISI was either 2.5 or 30 s and paradigm type was either the oddball or single-stimulus task. ERPs were recorded from the midline electrodes, with amplitude, mean area, and latency of the P300 and other components assessed. The results showed that P300 morphology was dramatically affected by task and ISI such that under the 2.5 s condition, the oddball paradigm produced typical ERP components, whereas the single-stimulus condition demonstrated minimal P300 amplitude. When ISI was 30 s, both the oddball and single-stimulus tasks produced robust P300 components but also evinced strong slow wave (SW) potentials, which contributed to the ERP measurement outcomes. It is concluded that P300 from visual stimuli can be elicited with both oddball and single-stimulus tasks when ISI is relatively long. ERPs from both paradigms produced appreciable SW activity, which needs to be considered when long ISI procedures are employed.