Effects of attention on the processing of physical and abstract auditory regularities: An exploratory MMN study.

The mismatch-negativity (MMN) component of the auditory event-related potential (ERP) reflects preattentive memory mechanisms encoding various types of regularities in the auditory environment. In an oddball paradigm, two types of deviant stimuli (in separate blocks) were presented among frequent standard stimuli: physical deviants (higher in pitch than the standards) and more complex, "abstract" deviants (tone pairs descending in pitch, presented among ascending standard tone pairs). The attentional load of the participant was manipulated under three conditions, where the participants either (1) watched a silent video, (2) played a computer game (Tetris) or (3) attended to the auditory stimuli and tried to detect infrequent target stimuli which were of lower intensity than the standard stimuli. The goal was to find out, whether the possible attention effects (suppression/enhancement) on the MMN are similar or different to stimuli requiring the extraction of either physical or abstract invariances. Both the physical and abstract deviants elicited in all conditions MMNs but no statistically significant amplitude differences between the conditions were found. The N2b and P3b components were elicited only in the attend condition and only by the soft target tones. The results further confirm that the MMN is a robust response to various types of regularity violations, showing no major effects of attentional manipulations. The results also suggest that the most commonly used primary task in MMN experiments, watching a silent video, usually keeps the participants' attention well enough directed away from the auditory stimuli. However, in cases where a cognitively more demanding but still participant-friendly primary task is needed, a simple computer game such as Tetris can be used, enabling better control of the participants' attention and vigilance.

Benefits of choir singing on complex auditory encoding in the aging brain: An ERP study.

Aging is accompanied by difficulties in auditory information processing, especially in more complex sound environments. Choir singing requires efficient processing of multiple sound features and could, therefore, mitigate the detrimental effects of aging on complex auditory encoding. We recorded auditory event-related potentials during passive listening of sounds in healthy older adult (≥ 60 years) choir singers and nonsinger controls. We conducted a complex oddball condition involving encoding of abstract regularities in combinations of pitch and location features, as well as in two simple oddball conditions, in which only either the pitch or spatial location of the sounds was varied. We analyzed change-related mismatch negativity (MMN) and obligatory P1 and N1 responses in each condition. In the complex condition, the choir singers showed a larger MMN than the controls, which also correlated with better performance in a verbal fluency test. In the simple pitch and location conditions, the choir singers had smaller N1 responses compared to the control subjects, whereas the MMN responses did not differ between groups. These results suggest that regular choir singing is associated both with more enhanced encoding of complex auditory regularities and more effective adaptation to simple sound features.

No evidence for theta power as a marker of hypnotic state in highly hypnotizable subjects.

EEG spectral-power density was analyzed in a group of nine highly hypnotizable subjects via ten frontal, central, parietal, and occipital electrodes under four conditions: 1) wake state, 2) neutral hypnosis, 3) hypnotic suggestion for altering perception of tones, and 4) post-hypnosis. Results indicate no theta-power changes between conditions, challenging previous findings that increased theta power is a marker of hypnosis. A decrease in gamma power under hypnotic suggestion and an almost significant decrease under neutral hypnosis were observed, compared to post-hypnosis. Anteroposterior power distribution remained stable over all conditions. The results are discussed and compared to earlier studies, which report heterogenous findings.

THE EFFECTS OF HYPNOSIS AND HYPNOTIC SUGGESTIONS ON THE MISMATCH NEGATIVITY IN HIGHLY HYPNOTIZABLE SUBJECTS.

The neural mechanisms associated with hypnosis were investigated in a group of 9 high hypnotizable subjects by measuring the mismatch negativity (MMN) component of the auditory event-related potential (ERP). ERPs were recorded using a passive oddball paradigm to sinusoidal standard and deviant tone stimuli of 500 and 520 Hz, respectively, in four conditions: prehypnosis, neutral hypnosis, hypnotic suggestion for altering the tone perception, and posthypnotic conditions. Earlier studies have indicated that hypnosis and hypnotic suggestions might have an effect on MMN, but the results of our study contradict these results: No statistically significant differences were found between the conditions in the MMN amplitudes.

Mismatch negativity (MMN) elicited by abstract regularity violations in two concurrent auditory streams.

The study investigated whether violations of abstract regularities in two parallel auditory stimulus streams can elicit the MMN (mismatch negativity) event-related potential. Tone pairs from a low (220-392 Hz) and a high (1319-2349 Hz) stream were delivered in an alternating order either at a fast or a slow pace. With the slow pace, the pairs were perceptually heard as a single stream obeying an alternating low pair-high pair pattern, whereas with the fast pace, an experience of two separate auditory streams, low and high, emerged. Both streams contained standard and deviant pairs. The standard pairs were either in both streams ascending in the direction of the within-pair pitch change or in the one stream ascending and in the other stream descending. The direction of the deviant pairs was opposite to that of the same-stream standard pairs. The participant's task was either to ignore the auditory stimuli or to detect the deviant pairs in the designated stream. The deviant pairs elicited an MMN both when the directions of the standard pairs in the two streams were the same or when they were opposite. The MMN was present irrespective of the pace of stimulation. The results indicate that the preattentive brain mechanisms, reflected by the MMN, can extract abstract regularities from two concurrent streams even when the regularities are opposite in the two streams, and independently of whether there perceptually exists only one stimulus stream or two segregated streams. These results demonstrate the brain's remarkable ability to model various regularities embedded in the auditory environment and update the models when the regularities are violated. The observed phenomena can be related to several aspects of auditory information processing, e.g., music and speech perception and different forms of attention.

Mismatch negativity (MMN) to speech sounds is modulated systematically by manual grip execution.

Manual actions and speech are connected: for example, grip execution can influence simultaneous vocalizations and vice versa. Our previous studies show that the consonant [k] is associated with the power grip and the consonant [t] with the precision grip. Here we studied whether the interaction between speech sounds and grips could operate already at a pre-attentive stage of auditory processing, reflected by the mismatch-negativity (MMN) component of the event-related potential (ERP). Participants executed power and precision grips according to visual cues while listening to syllable sequences consisting of [ke] and [te] utterances. The grips modulated the MMN amplitudes to these syllables in a systematic manner so that when the deviant was [ke], the MMN response was larger with a precision grip than with a power grip. There was a converse trend when the deviant was [te]. These results suggest that manual gestures and speech can interact already at a pre-attentive processing level of auditory perception, and show, for the first time that manual actions can systematically modulate the MMN.

Predictive coding of phonological rules in auditory cortex: A mismatch negativity study.

The brain is constantly generating predictions of future sensory input to enable efficient adaptation. In the auditory domain, this applies also to the processing of speech. Here we aimed to determine whether the brain predicts the following segments of speech input on the basis of language-specific phonological rules that concern non-adjacent phonemes. Auditory event-related potentials (ERP) were recorded in a mismatch negativity (MMN) paradigm, where the Finnish vowel harmony, determined by the first syllables of pseudowords, either constrained or did not constrain the phonological composition of pseudoword endings. The phonological rule of vowel harmony was expected to create predictions about phonologically legal pseudoword endings. Results showed that MMN responses were larger for phonologically illegal than legal pseudowords, and P3a was elicited only for illegal pseudowords. This supports the hypothesis that speech input is evaluated against context-dependent phonological predictions that facilitate speech processing.

Attention-shift vs. response-priming explanations for the spatial cueing effect in cross-modal tasks.

The task-irrelevant spatial location of a cue stimulus affects the processing of a subsequent target. This "Posner effect" has been explained by an exogenous attention shift to the spatial location of the cue, improving perceptual processing of the target. We studied whether the left/right location of task-irrelevant and uninformative tones produces cueing effects on the processing of visual targets. Tones were presented randomly from left or right. In the first condition, the subsequent visual target, requiring response either with the left or right hand, was presented peripherally to left or right. In the second condition, the target was a centrally presented left/right-pointing arrow, indicating the response hand. In the third condition, the tone and the central arrow were presented simultaneously. Data were recorded on compatible (the tone location and the response hand were the same) and incompatible trials. Reaction times were longer on incompatible than on compatible trials. The results of the second and third conditions are difficult to explain with the attention-shift model emphasizing improved perceptual processing in the cued location, as the central target did not require any location-based processing. Consequently, as an alternative explanation they suggest response priming in the hand corresponding to the spatial location of the tone. Simultaneous lateralized readiness potential (LRP) recordings were consistent with the behavioral data, the tone cues eliciting on incompatible trials a fast preparation for the incorrect response and on compatible trials preparation for the correct response.

The mismatch-negativity (MMN) component of the auditory event-related potential to violations of abstract regularities: a review.

The mismatch-negativity (MMN) component of the event-related potential (ERP) has been extensively used to study the preattentive processing and storage of regularities in basic physical stimulus features (e.g., frequency, intensity, spatial location). However, studies reviewed in the present article reveal that the auditory analysis reflected by MMN also includes the detection and use of more complex, "abstract", regularities based, for example, on relationships between various physical features of the stimuli or in patterns present in the auditory stream. When these regularities are violated, then MMN is elicited. Thus, the central auditory system performs even at the pre-attentive, auditory-cortex level surprisingly "cognitive" operations, such as generalization leading to simple concept formation, rule extraction and prediction of future stimuli. The information extracted often seems to be in an implicit form, not directly available to conscious processes and difficult to express verbally. It can nevertheless influence the behavior of the subject, for example, the regularity violations can temporarily impair performance in the primary task. Neural, behavioral and cognitive events associated with the development of the regularity representations are discussed.

Processing of audiovisual associations in the human brain: dependency on expectations and rule complexity.

In order to respond to environmental changes appropriately, the human brain must not only be able to detect environmental changes but also to form expectations of forthcoming events. The events in the external environment often have a number of multisensory features such as pitch and form. For integrated percepts of objects and events, crossmodal processing, and crossmodally induced expectations of forthcoming events are needed. The aim of the present study was to determine whether the expectations created by visual stimuli can modulate the deviance detection in the auditory modality, as reflected by auditory event-related potentials (ERPs). Additionally, it was studied whether the complexity of the rules linking auditory and visual stimuli together affects this process. The N2 deflection of the ERP was observed in response to violations in the subjects' expectation of a forthcoming tone. Both temporal aspects and cognitive demands during the audiovisual deviance detection task modulated the brain processes involved.

The preattentive processing of major vs. minor chords in the human brain: An event-related potential study.

Western music has two classifications that are highly familiar to all Western listeners: the dichotomy between the major and minor modalities and consonance vs. dissonance. We aimed at determining whether these classifications already take place at the level of the elicitation of the change-related mismatch negativity (MMN) component of the event-related potential (ERP). To this end, we constructed an oddball-paradigm with root minor, dissonant and inverted major chords in a context of root major chords. These stimuli were composed so that the standard and deviant chords did not include a physically deviant frequency which could cause the MMN. The standard chords were transposed into 12 different keys (=pitch levels) and delivered to the participants while they were watching a silent movie (ignore condition) or detecting softer target sounds (detection condition). In the ignore condition, the MMN was significant for all but inverted major chords. In the detection condition, the MMN was significant for dissonant chords and soft target chords. Our results indicate that the processes underlying MMN are able to make discriminations which are qualitative by nature. Whether the classifications between major and minor modalities and consonance vs. dissonance are innate or based on implicit learning remains a question for the future.

Preattentive detection of nonsalient contingencies between auditory features.

Extraction of contingencies between features of successive auditory stimuli was studied by exploiting the mismatch negativity component of the event-related potential. According to the rules hidden in the stimulus sequences, one stimulus feature (duration) predicted another feature (pitch) of the next stimulus. Occasional deviant stimuli, violating the rules, elicited a mismatch negativity although the participants were ignoring the stimuli. Mismatch negativity was also elicited when the participants tried to detect the deviant stimuli. Their detection performance, however, was poor and on the basis of subsequent interviews, they were not consciously aware of the rules. The results suggest that contingencies across-features of successive stimuli are extracted already at the early preattentive level in the auditory sensory memory.

Implicit, intuitive, and explicit knowledge of abstract regularities in a sound sequence: an event-related brain potential study.

Implicit knowledge has been proposed to be the substrate of intuition because intuitive judgments resemble implicit processes. We investigated whether the automatically elicited mismatch negativity (MMN) component of the auditory event-related potentials (ERPs) can reflect implicit knowledge and whether this knowledge can be utilized for intuitive sound discrimination. We also determined the sensitivity of the attention-and task-dependent P3 component to intuitive versus explicit knowledge. We recorded the ERPs elicited in an "abstract" oddball paradigm. Tone pairs roving over different frequencies but with a constant ascending inter-pair interval, were presented as frequent standard events. The standards were occasionally replaced by deviating, descending tone pairs. The ERPs were recorded under both ignore and attend conditions. Subjects were interviewed and classified on the basis of whether or not they could datect the deviants. The deviants elicited an MMN even in subjects who subsequent to the MMN recording did not express awareness of the deviants. This suggests that these subjects possessed implicit knowledge of the sound-sequence structure. Some of these subjects learned, in an associative training session, to detect the deviants intuitively, that is, they could detect the deviants but did not give a correct description of how the deviants differed from the standards. Intuitive deviant detection was not accompanied by P3 elicitation whereas subjects who developed explicit knowledge of the sound sequence during the training did show a P3 to the detected deviants.

Event-related potentials suggest early interaction between syntax and semantics during on-line sentence comprehension.

Event-related potentials (ERPs) were used to investigate interaction between syntactic parsing and semantic integration processes during a visual sentence comprehension task. The linguistic stimuli were Finnish five-word sentences containing morphosyntactic and/or semantic violations. Single morphosyntactic violations evoked left anterior negativity (LAN) and P600 components. Single semantic violations elicited a robust N400 effect over the left hemisphere. A later and weaker N400-like response was also observed in the right hemisphere, left-right hemispheric latency difference being 40 ms. Combined morphosyntactic and semantic violations elicited a P600 component and a negative ERP component within the latency range of the LAN and N400 components. Further analysis of these ERP effects provided evidence for early processual interaction between syntax and semantics during on-line sentence comprehension. The hemispheric distribution of the LAN and N400 components was taken to suggest lateralization of initial morphosyntactic parsing and semantic integration processes to the left hemisphere. In contrast, the later syntax-related P600 component was observed as being more pronounced over the posterior areas of the right hemisphere.

Electric brain responses indicate preattentive processing of abstract acoustic regularities in children.

This study investigated the preattentive processing of abstract acoustic regularities in children aged 8-14 years. Event-related brain potentials (ERPs) were elicited by frequent (standard) pairs ascending in pitch (the second tone having a higher frequency than the first tone) and by infrequent (deviant) pairs descending in pitch. In the easy condition, the second tone of the pair was always one step higher (standard) or lower (deviant) than the first tone, while in the hard condition, the second tone was randomly 1-10 steps higher or lower than the first tone. In the easy condition we found the mismatch negativity (MMN) and a subsequent positive P3a-like deflection. In the hard condition, the amplitude of MMN was lower over frontal sites than in the easy condition, while the temporal component of MMN was not impaired by complexity of abstract regularities. These results suggest that the complexity of the auditory stimulation affects preattentive auditory change detection in children.

Evidence for the different additivity of the temporal and frontal generators of mismatch negativity: a human auditory event-related potential study.

The auditory sensory-memory mechanisms in the human brain were investigated using the mismatch negativity (MMN) component of the event-related potential. MMNs were recorded to stimuli deviating from the repetitive standard stimuli simultaneously either in one or two features (frequency, intensity). If the processing of these two features is independent of each other, the MMN to the double deviant should equal the sum of the MMNs elicited by the corresponding single deviants. The double-deviant MMNs were found to be additive at the electrode sites below the Sylvian fissure but not at the frontal scalp areas. The results suggest that the temporal subcomponent of MMN is additive whereas the frontal is non-additive. The pattern of results was similar in ignore and attend conditions, suggesting that the components were not attentionally modulated.

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

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

Simultaneous storage of two complex temporal sound patterns in auditory sensory memory.

We wished to determine whether multiple sound patterns can be simultaneously represented in the temporary auditory buffer (auditory sensory memory), when subjects have no task related to the sounds. To this end we used the mismatch negativity (MMN) event-related potential, an electric brain response elicited when a frequent sound is infrequently replaced by a different sound. The MMN response is based on the presence of the auditory sensory memory trace of the frequent sounds, which exists whether or not these sounds are in the focus of the subject's attention. Subjects watching a movie were presented with sound sequences consisting of two frequent sound patterns, each formed of four different tones and an infrequent pattern consisting of the first two tones of one of the frequent sound pattern and the last two tones of the other frequent sound pattern. The infrequent sound pattern elicited an MMN, indicating that multiple sound patterns are formed at an early, largely automatic stage of auditory processing.