István Winkler

The concept of auditory stimulus representation in cognitive neuroscience

The sequence of neurophysiological processes elicited in the auditory system by a sound is analyzed in search of the stage at which the processes carrying sensory information cross the borderline beyond which they directly underlie sound perception. Neurophysiological data suggest that this transition occurs when the sensory input is mapped onto the physiological basis of sensory memory in the auditory cortex. At this point, the sensory information carried by the stimulus-elicited process corresponds, for the first time, to that contained by the actual sound percept. Before this stage, the sensory stimulus code is fragmentary, lacks the time dimension, cannot enter conscious perception, and is not accessible to top-down processes (voluntary mental operations). On these grounds, 2 distinct stages of auditory sensory processing, prerepresentational and representational, can be distinguished.

Neural Mechanisms of Involuntary Attention to Acoustic Novelty and Change

Behavioral and event-related brain potential (ERP) measures were used to elucidate the neural mechanisms of involuntary engagement of attention by novelty and change in the acoustic environment. The behavioral measures consisted of the reaction time (RT) and performance accuracy (hit rate) in a forced-choice visual RT task where subjects were to discriminate between odd and even numbers. Each visual stimulus was preceded by an irrelevant auditory stimulus, which was randomly either a standard tone (80), a slightly, higher deviant tone (10), or a natural, novel sound (10). Novel sounds prolonged the RT to successive visual stimuli by 17 msec as compared with the RT to visual stimuli that followed standard tones. Deviant tones, in turn, decreased the hit rate but did not significantly affect the RT. In the ERPs to deviant tones, the mismatch negativity (MMN), peaking at 150 msec, and a second negativity, peaking at 400 msec, could be observed. Novel sounds elicited an enhanced N1, with a probable overlap by the MMN, and a large positive P3a response with two different subcomponents: an early centrally dominant P3a, peaking at 230 msec, and a late P3a, peaking at 315 msec with a right-frontal scalp maximum. The present results suggest the involvement of two different neural mechanisms in triggering involuntary attention to acoustic novelty and change: a transient-detector mechanism activated by novel sounds and reflected in the N1 and a stimulus-change detector mechanism activated by deviant tones and novel sounds and reflected in the MMN. The observed differential distracting effects by slightly deviant tones and widely deviant novel sounds support the notion of two separate mechanisms of involuntary attention.

Involuntary attention and distractibility as evaluated with event-related brain potentials

This article reviews recent event-related brain potential (ERP) studies of involuntary attention and distractibility in response to novelty and change in the acoustic environment. These studies show that the mismatch negativity, N1 and P3a ERP components elicited by deviant or novel sounds in an unattended sequence of repetitive stimuli index different processes along the course to involuntary attention switch to distracting stimuli. These studies used new auditory-auditory and auditory-visual distraction paradigms, which enable one to assess objectively abnormal distractibility in several clinical patient groups, such as those suffering from closed-head injuries or chronic alcoholism.

Processing of novel sounds and frequency changes in the human auditory cortex: Magnetoencephalographic recordings

Whole-head magnetoencephalographic (MEG) responses to repeating standard tones and to infrequent slightly higher deviant tones and complex novel sounds were recorded together with event-related brain potentials (ERPs). Deviant tones and novel sounds elicited the mismatch negativity (MMN) component of the ERP and its MEG counterpart (MMNm) both when the auditory stimuli were attended to and when they were ignored. MMNm generators were located bilateral to the superior planes of the temporal lobes where preattentive auditory discrimination appears to occur. A subsequent positive P3a component was elicited by deviant tones and with a larger amplitude by novel sounds even when the sounds were to be ignored. Source localization for the MEG counterpart of P3a (P3am) suggested that the auditory cortex in the superior temporal plane is involved in the neural network of involuntary attention switching to changes in the acoustic environment.

Auditory processing that leads to conscious perception: A unique window to central auditory processing opened by the mismatch negativity and related responses

In this review, we will present a model of brain events leading to conscious perception in audition. This represents an updated version of Näätänens previous model of automatic and attentive central auditory processing. This revised model is mainly based on the mismatch negativity (MMN) and N1 indices of automatic processing, the processing negativity (PN) index of selective attention, and their magnetoencephalographic (MEG) and functional magnetic resonance imaging (fMRI) equivalents. Special attention is paid to determining the neural processes that might underlie conscious perception and the borderline between automatic and attention-dependent processes in audition.

Memory prerequisites of mismatch negativity in the auditory event-related potential (ERP)

The mismatch negativity (MMN) is a component of the auditory event-related brain potential that occurs in response to infrequent changes in the physical properties of homogeneous series of sounds, even when subjects are instructed to ignore the auditory channel of stimulation. It has been proposed (e.g., Näätänen, 1990) that the MMN is generated by an automatic process in which a difference between the deviant sound and the previous, standard sound is detected by the brain. However, it is unclear how the form of memory involved is related to the rest of the memory system. The present study indicates that, for an MMN to be elicited in response to a change in tone frequency, the representation of the standard tone must be both (a) well-established as a standard in memory, and (b) in a currently active state. The relation between physiological and psychological aspects of memory representation is discussed.

Adaptive modeling of the unattended acoustic environment reflected in the mismatch negativity event-related potential

The mismatch negativity (MMN) event-related potential is elicited by changes in repetitive auditory stimuli. The present paper suggests that: (1) an acoustic model of the auditory environment is maintained even in the absence of attention focussed on auditory stimuli, preattentively detecting repetitive features of the acoustic stimulation; and (2) the MMN reflects modifications to existing parts of this model during incorporation of a new stimulus into the model. MMN responses were investigated during the period when a repetitive stimulus (standard) was replaced by a new standard sound. It was found that whereas the new standard stimulus stopped eliciting an MMN after its third presentation with respect to the old standard, a probe stimulus, differing from both standards, elicited an MMN with respect to the old standard, even when following four presentations of the new standard. The probe stimulus also elicited an MMN with respect to the new standard after four or more presentations of this new standard stimulus, thus eliciting two consecutive MMNs. The comparison (conducted on the basis of the present and some previous findings) of the present hypothesis with alternative explanations of MMN based on the presence and strength of auditory transient memory traces supported the model adjustment hypothesis.

Newborn infants detect the beat in music

To shed light on how humans can learn to understand music, we need to discover what the perceptual capabilities with which infants are born. Beat induction, the detection of a regular pulse in an auditory signal, is considered a fundamental human trait that, arguably, played a decisive role in the origin of music. Theorists are divided on the issue whether this ability is innate or learned. We show that newborn infants develop expectation for the onset of rhythmic cycles (the downbeat), even when it is not marked by stress or other distinguishing spectral features. Omitting the downbeat elicits brain activity associated with violating sensory expectations. Thus, our results strongly support the view that beat perception is innate.

Phase Entrainment of Human Delta Oscillations Can Mediate the Effects of Expectation on Reaction Speed

The more we anticipate a response to a predictable stimulus, the faster we react. This empirical observation has been confirmed and quantified by many investigators suggesting that the processing of behaviorally relevant stimuli is facilitated by probability-based confidence of anticipation. However, the exact neural mechanisms underlying this phenomenon are largely unknown. Here we show that performance changes related to different levels of expectancy originate in dynamic modulation of delta oscillation phase. Our results obtained in rhythmic auditory target detection tasks indicated significant entrainment of the EEG delta rhythm to the onset of the target tones with increasing phase synchronization at higher levels of predictability. Reaction times correlated with the phase of the delta band oscillation at target onset. The fastest reactions occurred during the delta phase that most commonly coincided with the target event in the high expectancy conditions. These results suggest that low-frequency oscillations play a functional role in human anticipatory mechanisms, presumably by modulating synchronized rhythmic fluctuations in the excitability of large neuronal populations and by facilitating efficient task-related neuronal communication among brain areas responsible for sensory processing and response execution.

Memory-based detection of task-irrelevant visual changes.

Colored grating patterns were presented to 8 participants in a passive oddball condition (standard, 87.5% and deviant, 12.5%, differing in their color). In the corresponding multicolor condition, grating patterns of eight different colors were presented, their probabilities set equal both to each other and to that of the deviant in the oddball task. Compared with the ERP response elicited by the standard stimulus, the deviant response was negatively displaced over posterior areas, the difference wave peaking at 136 ms. A similar negative wave was obtained when the ERP response to the deviant was compared with the ERP elicited by the same stimulus in the multicolor condition. This result rules out stimulus- (color-) specific refractoriness as a major factor in the generation of the deviance-related posterior negativity. The observed posterior negativity can therefore be regarded as a visual analog of the mismatch negativity (vMMN).