Patrick J. C. May

Mismatch negativity (MMN), the deviance-elicited auditory deflection, explained.

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.

A Population Rate Code of Auditory Space in the Human Cortex

Background Previous work on the human auditory cortex has revealed areas specialized in spatial processing but how the neurons in these areas represent the location of a sound source remains unknown. Methodology/Principal Findings Here, we performed a magnetoencephalography (MEG) experiment with the aim of revealing the neural code of auditory space implemented by the human cortex. In a stimulus-specific adaptation paradigm, realistic spatial sound stimuli were presented in pairs of adaptor and probe locations. We found that the attenuation of the N1m response depended strongly on the spatial arrangement of the two sound sources. These location-specific effects showed that sounds originating from locations within the same hemifield activated the same neuronal population regardless of the spatial separation between the sound sources. In contrast, sounds originating from opposite hemifields activated separate groups of neurons. Conclusions/Significance These results are highly consistent with a rate code of spatial location formed by two opponent populations, one tuned to locations in the left and the other to those in the right. This indicates that the neuronal code of sound source location implemented by the human auditory cortex is similar to that previously found in other primates.

Sound localization in the human brain: neuromagnetic observations

&NA; Sound location processing in the human auditory cortex was studied with magnetoencephalography (MEG) by producing spatial stimuli using a modern stimulus generation methodology utilizing head‐related transfer functions (HRTFs). The stimulus set comprised wideband noise bursts filtered through HRTFs in order to produce natural spatial sounds. Neuromagnetic responses for stimuli representing eight equally spaced sound source directions in the azimuthal plane were measured from 10 subjects. The most prominent response, the cortically generated N1m, was investigated above the left and right hemisphere. We found, firstly, that the HRTF‐based stimuli presented from different directions elicited contralaterally prominent N1m responses. Secondly, we found that cortical activity reflecting the processing of spatial sound stimuli was more pronounced in the right than in the left hemisphere.

The auditory transient 40-Hz response is insensitive to changes in stimulus features.

TEN subjects were presented with tone pips occasionally interspersed with deviant tone pips of a higher frequency. The transient 40-Hz response was insensitive to change in qualitative stimulus features. In contrast, stimulus changes elicited a later, and slower event-related potential, the mismatch negativity (MMN). As a response to changes in stimulus features implies the existence of a memory system, and because changes in qualitative stimulus aspects do not activate the generator mechanisms underlying the 40-Hz response, the 40-Hz response can be dissociated from memory mechanisms. Furthermore, the analysis of phase-locked (synchronous), and non-phase-locked (asynchronous) responses revealed that the 40-Hz response might be caused by the synchronization of already active oscillators.

The neural code for interaural time difference in human auditory cortex

A magnetoencephalography study was conducted to reveal the neural code of interaural time difference (ITD) in the human cortex. Widely used crosscorrelator models predict that the code consists of narrow receptive fields distributed to all ITDs. The present findings are, however, more in line with a neural code formed by two opponent neural populations: one tuned to the left and the other to the right hemifield. The results are consistent with models of ITD extraction in the auditory brainstem of small mammals and, therefore, suggest that similar computational principles underlie human sound source localization.

Asymmetrical representation of auditory space in human cortex

Recent single-neuron recordings in monkeys and magnetoencephalography (MEG) data on humans suggest that auditory space is represented in cortex as a population rate code whereby spatial receptive fields are wide and centered at locations to the far left or right of the subject. To explore the details of this code in the human brain, we conducted an MEG study utilizing realistic spatial sound stimuli presented in a stimulus-specific adaptation paradigm. In this paradigm, the spatial selectivity of cortical neurons is measured as the effect the location of a preceding adaptor has on the response to a subsequent probe sound. Two types of stimuli were used: a wideband noise sound and a speech sound. The cortical hemispheres differed in the effects the adaptors had on the response to a probe sound presented in front of the subject. The right-hemispheric responses were attenuated more by an adaptor to the left than by an adaptor to the right of the subject. In contrast, the left-hemispheric responses were similarly affected by adaptors in these two locations. When interpreted in terms of single-neuron spatial receptive fields, these results support a population rate code model where neurons in the right hemisphere are more often tuned to the left than to the right of the perceiver while in the left hemisphere these two neuronal populations are of equal size.

Auditory spatial processing in the human cortex

The auditory system codes spatial locations in a way that deviates from the spatial representations found in other modalities. This difference is especially striking in the cortex, where neurons form topographical maps of visual and tactile space but where auditory space is represented through a population rate code. In this hemifield code, sound source location is represented in the activity of two widely tuned opponent populations, one tuned to the right and the other to the left side of auditory space. Scientists are only beginning to uncover how this coding strategy adapts to various spatial processing demands. This review presents the current understanding of auditory spatial processing in the cortex. To this end, the authors consider how various implementations of the hemifield code may exist within the auditory cortex and how these may be modulated by the stimulation and task context. As a result, a coherent set of neural strategies for auditory spatial processing emerges.

Long-term stimulation attenuates the transient 40-Hz response

The effect of long-term stimulation on the transient 40-Hz response was studied in 5-h long experimental sessions. Ten human subjects were presented with series of 1000 Hz tone pips occasionally replaced by 1200 Hz tone pips. The transient 40-Hz response was compared with two event-related potential (ERP) components, the N1 and the mismatch negativity (MMN) during the different phases of the long-lasting sessions. The 40-Hz and N1 onset responses attenuated while the MMN did not. Thus, the 40-Hz response might indicate vigilance.

Cortical sensitivity to periodicity of speech sounds

Previous non-invasive brain research has reported auditory cortical sensitivity to periodicity as reflected by larger and more anterior responses to periodic than to aperiodic vowels. The current study investigated whether there is a lower fundamental frequency (F0) limit for this effect. Auditory evoked fields (AEFs) elicited by natural-sounding 400 ms periodic and aperiodic vowel stimuli were measured with magnetoencephalography. Vowel F0 ranged from normal male speech (113 Hz) to exceptionally low values (9 Hz). Both the auditory N1m and sustained fields were larger in amplitude for periodic than for aperiodic vowels. The AEF sources for periodic vowels were also anterior to those for the aperiodic vowels. Importantly, the AEF amplitudes and locations were unaffected by the F0 decrement of the periodic vowels. However, the N1m latency increased monotonically as F0 was decreased down to 19 Hz, below which this trend broke down. Also, a cascade of transient N1m-like responses was observed in the lowest F0 condition. Thus, the auditory system seems capable of extracting the periodicity even from very low F0 vowels. The behavior of the N1m latency and the emergence of a response cascade at very low F0 values may reflect the lower limit of pitch perception.

Sensitivity of the human auditory cortex to acoustic degradation of speech and non-speech sounds

BackgroundRecent studies have shown that the human right-hemispheric auditory cortex is particularly sensitive to reduction in sound quality, with an increase in distortion resulting in an amplification of the auditory N1m response measured in the magnetoencephalography (MEG). Here, we examined whether this sensitivity is specific to the processing of acoustic properties of speech or whether it can be observed also in the processing of sounds with a simple spectral structure. We degraded speech stimuli (vowel /a/), complex non-speech stimuli (a composite of five sinusoidals), and sinusoidal tones by decreasing the amplitude resolution of the signal waveform. The amplitude resolution was impoverished by reducing the number of bits to represent the signal samples. Auditory evoked magnetic fields (AEFs) were measured in the left and right hemisphere of sixteen healthy subjects.ResultsWe found that the AEF amplitudes increased significantly with stimulus distortion for all stimulus types, which indicates that the right-hemispheric N1m sensitivity is not related exclusively to degradation of acoustic properties of speech. In addition, the P1m and P2m responses were amplified with increasing distortion similarly in both hemispheres. The AEF latencies were not systematically affected by the distortion.ConclusionsWe propose that the increased activity of AEFs reflects cortical processing of acoustic properties common to both speech and non-speech stimuli. More specifically, the enhancement is most likely caused by spectral changes brought about by the decrease of amplitude resolution, in particular the introduction of periodic, signal-dependent distortion to the original sound. Converging evidence suggests that the observed AEF amplification could reflect cortical sensitivity to periodic sounds.