The current review constitutes the first comprehensive look at the possibility that the mismatch negativity (MMN, the deflection of the auditory ERP/ERF elicited by stimulus change) might be generated by so-called fresh-afferent neuronal activity. This possibility has been repeatedly ruled out for the past 30 years, with the prevailing theoretical accounts relying on a memory-based explanation instead. We propose that the MMN is, in essence, a latency-and amplitude-modulated expression of the auditory N1 response, generated by fresh-afferent activity of cortical neurons that are under nonuniform levels of adaptation.

Being able to detect unusual, possibly dangerous events in the environment is a fundamental ability that helps ensure the survival of biological organisms. Novelty detection requires a memory system that models (builds neural representations of) events in the environment, so that changes are detected because they violate the predictions of the model. The earliest physiologically measurable brain response to novel auditory stimuli is the mismatch negativity, MMN, a component of the event-related potential. It is elicited when a predictable series of unvarying stimuli is unexpectedly followed by a deviating stimulus. As the occurrence of MMN is not usually affected by the direction of attention, MMN reflects the operation of automatic sensory (echoic) memory, the earliest memory system that builds traces of the acoustic environment against which new stimuli can be compared. The dependence of attentive novelty detection on earlier, pre-attentive processes, however, has remained elusive. Previous, related studies seem to suggest a relationship between MMN and attentive processes, although no conclusive evidence has so far been shown. Here we address novelty detection in humans both on a physiological and behavioural level, and show how attentive novelty detection is governed by a pre-attentive sensory memory mechanism.

Life or death in hostile environments depends crucially on one's ability to detect and gate novel sounds to awareness, such as that of a twig cracking under the paw of a stalking predator in a noisy jungle. Two distinct auditory cortex processes have been thought to underlie this phenomenon: (i) attenuation of the so-called N1 response with repeated stimulation and (ii) elicitation of a mismatch negativity response (MMN) by changes in repetitive aspects of auditory stimulation. This division has been based on previous studies suggesting that, unlike for the N1, repetitive ''standard'' stimuli preceding a physically different ''novel'' stimulus constitute a prerequisite to MMN elicitation, and that the source loci of MMN and N1 are different. Contradicting these findings, our combined electromagnetic, hemodynamic, and psychophysical data indicate that the MMN is generated as a result of differential adaptation of anterior and posterior auditory cortex N1 sources by preceding auditory stimulation. Early (Ϸ85 ms) neural activity within posterior auditory cortex is adapted as sound novelty decreases. This alters the center of gravity of electromagnetic N1 source activity, creating an illusory difference between N1 and MMN source loci when estimated by using equivalent current dipole fits. Further, our electroencephalography data show a robust MMN after a single standard event when the interval between two consecutive novel sounds is kept invariant. Our converging findings suggest that transient adaptation of feature-specific neurons within human posterior auditory cortex filters superfluous sounds from entering one's awareness.

Studies of human auditory and somatosensory modalities have shown that there is an oscillatory response in the gamma-band (at about 40 Hz) frequency which is elicited by either steady state or transient stimulation. The auditory 40-Hz response is generated at least partially in the auditory cortex as a result of thalamocortical interaction and may serve perceptual integration and conscious perception. A connection to selective attention has been implied in human and animal studies, although the evidence is inconclusive. Moreover, fundamental differences between the human and animal 40-Hz responses prohibit generalization. Furthermore, most experiments have used steady-state stimulation during which the brain does not regain its resting state between stimuli as it does when transient stimulation is used. Here we study the effect of selective attention on the auditory gamma-band (40-Hz) transient response using subjects listening to tone pips presented in one ear while ignoring a concurrent sequence of tone pips in the other ear. The 40-Hz response was larger when subjects paid attention to stimuli rather than ignored them. This attention effect was most pronounced over the frontal and central scalp areas. Our results demonstrate a physiological correlate of selective attention in the 40-Hz transient response in humans.

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The mismatch negativity (MMN) component of the auditory event-related potential (ERP) is elicited by infrequent, physically deviant stimuli in a sequence of frequent homogeneous stimuli (standards). It has been suggested that the MMN is generated by an automatic (attention-independent) neural mismatch process with a memory trace that encodes the physical features of the standard stimulus. The proposed MMN independence of attention was addressed in the present study. Standard stimuli and two types of deviant stimuli, differing from standards either in frequency or intensity, were dichotically presented in random order and at a rapid rate. The subject attended either to left- or right-ear stimuli, counting the number of a designated type of deviants in that ear. In the present conditions of very strongly focused attention, the MMN was elicited even by frequency change in the ignored input stream, and its amplitude was very similar to that of the MMN elicited by equivalent deviant stimuli (targets) in the attended input stream. In contrast, the MMN to intensity deviation was clearly attenuated in the absence of attention. This effect is, however, probably due to the attention effect on the MMN generator itself rather than the antecedent sensory-analysis and -storing functions.

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Here, the perception of auditory spatial information as indexed by behavioral measures is linked to brain dynamics as reflected by the N1m response recorded with whole-head magnetoencephalography (MEG). Broadband noise stimuli with realistic spatial cues corresponding to eight direction angles in the horizontal plane were constructed via custom-made, individualized binaural recordings (BAR) and generic head-related transfer functions (HRTF). For comparison purposes, stimuli with impoverished acoustical cues were created via interaural time and level differences (ITDs and ILDs) and their combinations. MEG recordings in ten subjects revealed that the amplitude and the latency of the N1m exhibits directional tuning to sound location, with the amplitude of the right-hemispheric N1m being particularly sensitive to the amount of spatial cues in the stimuli. The BAR, HRTF, and combined ITD + ILD stimuli resulted both in a larger dynamic range and in a more systematic distribution of the N1m amplitude across stimulus angle than did the ITD or ILD stimuli alone. Further, the righthemispheric source loci of the N1m responses for the BAR and HRTF stimuli were anterior to those for the ITD and ILD stimuli. In behavioral tests, we measured the ability of the subjects to localize BAR and HRTF stimuli in terms of azimuthal error and front-back confusions. We found that behavioral performance correlated positively with the amplitude of the N1m. Thus, the activity taking place already in the auditory cortex predicts behavioral sound detection of spatial stimuli, and the amount of spatial cues embedded in the signal are reflected in the activity of this brain area. D