Early-latency categorical speech sound representations in the left inferior frontal gyrus.

Efficient speech perception requires the mapping of highly variable acoustic signals to distinct phonetic categories. How the brain overcomes this many-to-one mapping problem has remained unresolved. To infer the cortical location, latency, and dependency on attention of categorical speech sound representations in the human brain, we measured stimulus-specific adaptation of neuromagnetic responses to sounds from a phonetic continuum. The participants attended to the sounds while performing a non-phonetic listening task and, in a separate recording condition, ignored the sounds while watching a silent film. Neural adaptation indicative of phoneme category selectivity was found only during the attentive condition in the pars opercularis (POp) of the left inferior frontal gyrus, where the degree of selectivity correlated with the ability of the participants to categorize the phonetic stimuli. Importantly, these category-specific representations were activated at an early latency of 115-140 ms, which is compatible with the speed of perceptual phonetic categorization. Further, concurrent functional connectivity was observed between POp and posterior auditory cortical areas. These novel findings suggest that when humans attend to speech, the left POp mediates phonetic categorization through integration of auditory and motor information via the dorsal auditory stream.

Previous exposure to intact speech increases intelligibility of its digitally degraded counterpart as a function of stimulus complexity.

Recent studies have shown that acoustically distorted sentences can be perceived as either unintelligible or intelligible depending on whether one has previously been exposed to the undistorted, intelligible versions of the sentences. This allows studying processes specifically related to speech intelligibility since any change between the responses to the distorted stimuli before and after the presentation of their undistorted counterparts cannot be attributed to acoustic variability but, rather, to the successful mapping of sensory information onto memory representations. To estimate how the complexity of the message is reflected in speech comprehension, we applied this rapid change in perception to behavioral and magnetoencephalography (MEG) experiments using vowels, words and sentences. In the experiments, stimuli were initially presented to the subject in a distorted form, after which undistorted versions of the stimuli were presented. Finally, the original distorted stimuli were presented once more. The resulting increase in intelligibility observed for the second presentation of the distorted stimuli depended on the complexity of the stimulus: vowels remained unintelligible (behaviorally measured intelligibility 27%) whereas the intelligibility of the words increased from 19% to 45% and that of the sentences from 31% to 65%. This increase in the intelligibility of the degraded stimuli was reflected as an enhancement of activity in the auditory cortex and surrounding areas at early latencies of 130-160ms. In the same regions, increasing stimulus complexity attenuated mean currents at latencies of 130-160ms whereas at latencies of 200-270ms the mean currents increased. These modulations in cortical activity may reflect feedback from top-down mechanisms enhancing the extraction of information from speech. The behavioral results suggest that memory-driven expectancies can have a significant effect on speech comprehension, especially in acoustically adverse conditions where the bottom-up information is decreased.

Memory Stacking in Hierarchical Networks.

Robust representations of sounds with a complex spectrotemporal structure are thought to emerge in hierarchically organized auditory cortex, but the computational advantage of this hierarchy remains unknown. Here, we used computational models to study how such hierarchical structures affect temporal binding in neural networks. We equipped individual units in different types of feedforward networks with local memory mechanisms storing recent inputs and observed how this affected the ability of the networks to process stimuli context dependently. Our findings illustrate that these local memories stack up in hierarchical structures and hence allow network units to exhibit selectivity to spectral sequences longer than the time spans of the local memories. We also illustrate that short-term synaptic plasticity is a potential local memory mechanism within the auditory cortex, and we show that it can bring robustness to context dependence against variation in the temporal rate of stimuli, while introducing nonlinearities to response profiles that are not well captured by standard linear spectrotemporal receptive field models. The results therefore indicate that short-term synaptic plasticity might provide hierarchically structured auditory cortex with computational capabilities important for robust representations of spectrotemporal patterns.

Computational modelling suggests that temporal integration results from synaptic adaptation in auditory cortex.

Incoming sounds are represented in the context of preceding events, and this requires a memory mechanism that integrates information over time. Here, it was demonstrated that response adaptation, the suppression of neural responses due to stimulus repetition, might reflect a computational solution that auditory cortex uses for temporal integration. Adaptation is observed in single-unit measurements as two-tone forward masking effects and as stimulus-specific adaptation (SSA). In non-invasive observations, the amplitude of the auditory N1m response adapts strongly with stimulus repetition, and it is followed by response recovery (the so-called mismatch response) to rare deviant events. The current computational simulations described the serial core-belt-parabelt structure of auditory cortex, and included synaptic adaptation, the short-term, activity-dependent depression of excitatory corticocortical connections. It was found that synaptic adaptation is sufficient for columns to respond selectively to tone pairs and complex tone sequences. These responses were defined as combination sensitive, thus reflecting temporal integration, when a strong response to a stimulus sequence was coupled with weaker responses both to the time-reversed sequence and to the isolated sequence elements. The temporal complexity of the stimulus seemed to be reflected in the proportion of combination-sensitive columns across the different regions of the model. Our results suggest that while synaptic adaptation produces facilitation and suppression effects, including SSA and the modulation of the N1m response, its functional significance may actually be in its contribution to temporal integration. This integration seems to benefit from the serial structure of auditory cortex.

Cortical processing of degraded speech sounds: effects of distortion type and continuity.

Human speech perception is highly resilient to acoustic distortions. In addition to distortions from external sound sources, degradation of the acoustic structure of the sound itself can substantially reduce the intelligibility of speech. The degradation of the internal structure of speech happens, for example, when the digital representation of the signal is impoverished by reducing its amplitude resolution. Further, the perception of speech is also influenced by whether the distortion is transient, coinciding with speech, or is heard continuously in the background. However, the complex effects of the acoustic structure and continuity of the distortion on the cortical processing of degraded speech are unclear. In the present magnetoencephalography study, we investigated how the cortical processing of degraded speech sounds as measured through the auditory N1m response is affected by variation of both the distortion type (internal, external) and the continuity of distortion (transient, continuous). We found that when the distortion was continuous, the N1m was significantly delayed, regardless of the type of distortion. The N1m amplitude, in turn, was affected only when speech sounds were degraded with transient internal distortion, which resulted in larger response amplitudes. The results suggest that external and internal distortions of speech result in divergent patterns of activity in the auditory cortex, and that the effects are modulated by the temporal continuity of the distortion.

Enhanced early-latency electromagnetic activity in the left premotor cortex is associated with successful phonetic categorization.

Sensory-motor interactions between auditory and articulatory representations in the dorsal auditory processing stream are suggested to contribute to speech perception, especially when bottom-up information alone is insufficient for purely auditory perceptual mechanisms to succeed. Here, we hypothesized that the dorsal stream responds more vigorously to auditory syllables when one is engaged in a phonetic identification/repetition task subsequent to perception compared to passive listening, and that this effect is further augmented when the syllables are embedded in noise. To this end, we recorded magnetoencephalography while twenty subjects listened to speech syllables, with and without noise masking, in four conditions: passive perception; overt repetition; covert repetition; and overt imitation. Compared to passive listening, left-hemispheric N100m equivalent current dipole responses were amplified and shifted posteriorly when perception was followed by covert repetition task. Cortically constrained minimum-norm estimates showed amplified left supramarginal and angylar gyri responses in the covert repetition condition at ~100ms from stimulus onset. Longer-latency responses at ~200ms were amplified in the covert repetition condition in the left angular gyrus and in all three active conditions in the left premotor cortex, with further enhancements when the syllables were embedded in noise. Phonetic categorization accuracy and magnitude of voice pitch change between overt repetition and imitation conditions correlated with left premotor cortex responses at ~100 and ~200ms, respectively. Together, these results suggest that the dorsal stream involvement in speech perception is dependent on perceptual task demands and that phonetic categorization performance is influenced by the left premotor cortex.

Transient and sustained cortical activity elicited by connected speech of varying intelligibility.

BACKGROUND: The robustness of speech perception in the face of acoustic variation is founded on the ability of the auditory system to integrate the acoustic features of speech and to segregate them from background noise. This auditory scene analysis process is facilitated by top-down mechanisms, such as recognition memory for speech content. However, the cortical processes underlying these facilitatory mechanisms remain unclear. The present magnetoencephalography (MEG) study examined how the activity of auditory cortical areas is modulated by acoustic degradation and intelligibility of connected speech. The experimental design allowed for the comparison of cortical activity patterns elicited by acoustically identical stimuli which were perceived as either intelligible or unintelligible. RESULTS: In the experiment, a set of sentences was presented to the subject in distorted, undistorted, and again in distorted form. The intervening exposure to undistorted versions of sentences rendered the initially unintelligible, distorted sentences intelligible, as evidenced by an increase from 30% to 80% in the proportion of sentences reported as intelligible. These perceptual changes were reflected in the activity of the auditory cortex, with the auditory N1m response (~100 ms) being more prominent for the distorted stimuli than for the intact ones. In the time range of auditory P2m response (>200 ms), auditory cortex as well as regions anterior and posterior to this area generated a stronger response to sentences which were intelligible than unintelligible. During the sustained field (>300 ms), stronger activity was elicited by degraded stimuli in auditory cortex and by intelligible sentences in areas posterior to auditory cortex. CONCLUSIONS: The current findings suggest that the auditory system comprises bottom-up and top-down processes which are reflected in transient and sustained brain activity. It appears that analysis of acoustic features occurs during the first 100 ms, and sensitivity to speech intelligibility emerges in auditory cortex and surrounding areas from 200 ms onwards. The two processes are intertwined, with the activity of auditory cortical areas being modulated by top-down processes related to memory traces of speech and supporting speech intelligibility.

Cortical encoding of aperiodic and periodic speech sounds: evidence for distinct neural populations.

Most speech sounds are periodic due to the vibration of the vocal folds. Non-invasive studies of the human brain have revealed a periodicity-sensitive population in the auditory cortex which might contribute to the encoding of speech periodicity. Since the periodicity of natural speech varies from (almost) periodic to aperiodic, one may argue that speech aperiodicity could similarly be represented by a dedicated neuron population. In the current magnetoencephalography study, cortical sensitivity to periodicity was probed with natural periodic vowels and their aperiodic counterparts in a stimulus-specific adaptation paradigm. The effects of intervening adaptor stimuli on the N1m elicited by the probe stimuli (the actual effective stimuli) were studied under interstimulus intervals (ISIs) of 800 and 200 ms. The results indicated a periodicity-dependent release from adaptation which was observed for aperiodic probes alternating with periodic adaptors under both ISIs. Such release from adaptation can be attributed to the activation of a distinct neural population responsive to aperiodic (probe) but not to periodic (adaptor) stimuli. Thus, the current results suggest that the aperiodicity of speech sounds may be represented not only by decreased activation of the periodicity-sensitive population but, additionally, by the activation of a distinct cortical population responsive to speech aperiodicity.

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

BACKGROUND: Recent 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. RESULTS: We 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. CONCLUSIONS: We 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.

Temporal integration of vowel periodicity in the auditory cortex.

Cortical sensitivity to the periodicity of speech sounds has been evidenced by larger, more anterior responses to periodic than to aperiodic vowels in several non-invasive studies of the human brain. The current study investigated the temporal integration underlying the cortical sensitivity to speech periodicity by studying the increase in periodicity-specific cortical activation with growing stimulus duration. Periodicity-specific activation was estimated from magnetoencephalography as the differences between the N1m responses elicited by periodic and aperiodic vowel stimuli. The duration of the vowel stimuli with a fundamental frequency (F0=106 Hz) representative of typical male speech was varied in units corresponding to the vowel fundamental period (9.4 ms) and ranged from one to ten units. Cortical sensitivity to speech periodicity, as reflected by larger and more anterior responses to periodic than to aperiodic stimuli, was observed when stimulus duration was 3 cycles or more. Further, for stimulus durations of 5 cycles and above, response latency was shorter for the periodic than for the aperiodic stimuli. Together the current results define a temporal window of integration for the periodicity of speech sounds in the F0 range of typical male speech. The length of this window is 3-5 cycles, or 30-50 ms.

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.

Modeling the categorical perception of speech sounds: a step toward biological plausibility.

Our native language has a lifelong effect on how we perceive speech sounds. Behaviorally, this is manifested as categorical perception, but the neural mechanisms underlying this phenomenon are still unknown. Here, we constructed a computational model of categorical perception, following principles consistent with infant speech learning. A self-organizing network was exposed to a statistical distribution of speech input presented as neural activity patterns of the auditory periphery, resembling the way sound arrives to the human brain. In the resulting neural map, categorical perception emerges from most single neurons of the model being maximally activated by prototypical speech sounds, while the largest variability in activity is produced at category boundaries. Consequently, regions in the vicinity of prototypes become perceptually compressed, and regions at category boundaries become expanded. Thus, the present study offers a unifying framework for explaining the neural basis of the warping of perceptual space associated with categorical perception.

Representation of the vocal roughness of aperiodic speech sounds in the auditory cortex.

Aperiodicity of speech alters voice quality. The current study investigated the relationship between vowel aperiodicity and human auditory cortical N1m and sustained field (SF) responses with magnetoencephalography. Behavioral estimates of vocal roughness perception were also collected. Stimulus aperiodicity was experimentally varied by increasing vocal jitter with techniques that model the mechanisms of natural speech production. N1m and SF responses for vowels with high vocal jitter were reduced in amplitude as compared to those elicited by vowels of normal vocal periodicity. Behavioral results indicated that the ratings of vocal roughness increased up to the highest jitter values. Based on these findings, the representation of vocal jitter in the auditory cortex is suggested to be formed on the basis of reduced activity in periodicity-sensitive neural populations.

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.

The contribution of high frequencies to human brain activity underlying horizontal localization of natural spatial sounds.

BACKGROUND: In the field of auditory neuroscience, much research has focused on the neural processes underlying human sound localization. A recent magnetoencephalography (MEG) study investigated localization-related brain activity by measuring the N1m event-related response originating in the auditory cortex. It was found that the dynamic range of the right-hemispheric N1m response, defined as the mean difference in response magnitude between contralateral and ipsilateral stimulation, reflects cortical activity related to the discrimination of horizontal sound direction. Interestingly, the results also suggested that the presence of realistic spectral information within horizontally located spatial sounds resulted in a larger right-hemispheric N1m dynamic range. Spectral cues being predominant at high frequencies, the present study further investigated the issue by removing frequencies from the spatial stimuli with low-pass filtering. This resulted in a stepwise elimination of direction-specific spectral information. Interaural time and level differences were kept constant. The original, unfiltered stimuli were broadband noise signals presented from five frontal horizontal directions and binaurally recorded for eight human subjects with miniature microphones placed in each subject's ear canals. Stimuli were presented to the subjects during MEG registration and in a behavioral listening experiment. RESULTS: The dynamic range of the right-hemispheric N1m amplitude was not significantly affected even when all frequencies above 600 Hz were removed. The dynamic range of the left-hemispheric N1m response was significantly diminished by the removal of frequencies over 7.5 kHz. The subjects' behavioral sound direction discrimination was only affected by the removal of frequencies over 600 Hz. CONCLUSION: In accord with previous psychophysical findings, the current results indicate that frontal horizontal sound localization and related right-hemispheric cortical processes are insensitive to the presence of high-frequency spectral information. The previously described changes in localization-related brain activity, reflected in the enlarged N1m dynamic range elicited by natural spatial stimuli, can most likely be attributed to the processing of individualized spatial cues present already at relatively low frequencies. The left-hemispheric effect could be an indication of left-hemispheric processing of high-frequency sound information unrelated to sound localization. Taken together, these results provide converging evidence for a hemispheric asymmetry in sound localization.

The right-hemispheric auditory cortex in humans is sensitive to degraded speech sounds.

We investigated how degraded speech sounds activate the auditory cortices of the left and right hemisphere. To degrade the stimuli, we introduce uniform scalar quantization, a controlled and replicable manipulation, not used before, in cognitive neuroscience. Three Finnish vowels (/a/, /e/ and /u/) were used as stimuli for 10 participants in magnetoencephalography registrations. Compared with the original vowel sounds, the degraded sounds increased the amplitude of the right-hemispheric N1m without affecting the latency whereas the amplitude and latency of the N1m in the left hemisphere remained unaffected. Although the participants were able to identify the stimuli correctly, the increased degradation led to increased reaction times which correlated positively with the N1m amplitude. Thus, the auditory cortex of right hemisphere might be particularly involved in processing degraded speech and possibly compensates for the poor signal quality by increasing its activity.

Predictive processing increases intelligibility of acoustically distorted speech: Behavioral and neural correlates.

INTRODUCTION: We examined which brain areas are involved in the comprehension of acoustically distorted speech using an experimental paradigm where the same distorted sentence can be perceived at different levels of intelligibility. This change in intelligibility occurs via a single intervening presentation of the intact version of the sentence, and the effect lasts at least on the order of minutes. Since the acoustic structure of the distorted stimulus is kept fixed and only intelligibility is varied, this allows one to study brain activity related to speech comprehension specifically. METHODS: In a functional magnetic resonance imaging (fMRI) experiment, a stimulus set contained a block of six distorted sentences. This was followed by the intact counterparts of the sentences, after which the sentences were presented in distorted form again. A total of 18 such sets were presented to 20 human subjects. RESULTS: The blood oxygenation level dependent (BOLD)-responses elicited by the distorted sentences which came after the disambiguating, intact sentences were contrasted with the responses to the sentences presented before disambiguation. This revealed increased activity in the bilateral frontal pole, the dorsal anterior cingulate/paracingulate cortex, and the right frontal operculum. Decreased BOLD responses were observed in the posterior insula, Heschl's gyrus, and the posterior superior temporal sulcus. CONCLUSIONS: The brain areas that showed BOLD-enhancement for increased sentence comprehension have been associated with executive functions and with the mapping of incoming sensory information to representations stored in episodic memory. Thus, the comprehension of acoustically distorted speech may be associated with the engagement of memory-related subsystems. Further, activity in the primary auditory cortex was modulated by prior experience, possibly in a predictive coding framework. Our results suggest that memory biases the perception of ambiguous sensory information toward interpretations that have the highest probability to be correct based on previous experience.

Neuromagnetic recordings reveal the temporal dynamics of auditory spatial processing in the human cortex.

In an attempt to delineate the assumed 'what' and 'where' processing streams, we studied the processing of spatial sound in the human cortex by using magnetoencephalography in the passive and active recording conditions and two kinds of spatial stimuli: individually constructed, highly realistic spatial (3D) stimuli and stimuli containing interaural time difference (ITD) cues only. The auditory P1m, N1m, and P2m responses of the event-related field were found to be sensitive to the direction of sound source in the azimuthal plane. In general, the right-hemispheric responses to spatial sounds were more prominent than the left-hemispheric ones. The right-hemispheric P1m and N1m responses peaked earlier for sound sources in the contralateral than for sources in the ipsilateral hemifield and the peak amplitudes of all responses reached their maxima for contralateral sound sources. The amplitude of the right-hemispheric P2m response reflected the degree of spatiality of sound, being twice as large for the 3D than ITD stimuli. The results indicate that the right hemisphere is specialized in the processing of spatial cues in the passive recording condition. Minimum current estimate (MCE) localization revealed that temporal areas were activated both in the active and passive condition. This initial activation, taking place at around 100 ms, was followed by parietal and frontal activity at 180 and 200 ms, respectively. The latter activations, however, were specific to attentional engagement and motor responding. This suggests that parietal activation reflects active responding to a spatial sound rather than auditory spatial processing as such.

From the repression of contents to the rules of the (narrative) self: a present-day cognitive view of the Freudian phenomenon of repressed contents.

In psychoanalysis, it is commonly thought that ideas (desires, fears, etc.) may be repressed, and that they can be made conscious. In this article, we shall apply cognitive viewpoints and assert that ideas do not exist in the unconscious as 'ready made', and thus repressed ideas cannot be 'brought' into consciousness. We suggest that the contents of consciousness are formed by processes on four levels: (1) unconscious brain processes, (2) the level of consciousness, (3) the level of self-consciousness, and (4) the level of narrative self-consciousness. From this point of view, the absence (or repression) of certain contents appears to be due to the missing of processes on Levels 1-4. Consequently, repressed contents appear in consciousness when appropriate processes take place. When studied in terms of our four-level model, repression may be treated as part of the study of the self. By applying the viewpoint of the self to the phenomenon of repression, the danger of the homunculus problem can be avoided. It also becomes apparent that certain fundamental problems met in the study of the self are the ones that Freud tried to solve in his meta-psychological writings.

Disentangling the effects of phonation and articulation: hemispheric asymmetries in the auditory N1m response of the human brain.

BACKGROUND: The cortical activity underlying the perception of vowel identity has typically been addressed by manipulating the first and second formant frequency (F1 & F2) of the speech stimuli. These two values, originating from articulation, are already sufficient for the phonetic characterization of vowel category. In the present study, we investigated how the spectral cues caused by articulation are reflected in cortical speech processing when combined with phonation, the other major part of speech production manifested as the fundamental frequency (F0) and its harmonic integer multiples. To study the combined effects of articulation and phonation we presented vowels with either high (/a/) or low (/u/) formant frequencies which were driven by three different types of excitation: a natural periodic pulseform reflecting the vibration of the vocal folds, an aperiodic noise excitation, or a tonal waveform. The auditory N1m response was recorded with whole-head magnetoencephalography (MEG) from ten human subjects in order to resolve whether brain events reflecting articulation and phonation are specific to the left or right hemisphere of the human brain. RESULTS: The N1m responses for the six stimulus types displayed a considerable dynamic range of 115-135 ms, and were elicited faster (approximately 10 ms) by the high-formant /a/ than by the low-formant /u/, indicating an effect of articulation. While excitation type had no effect on the latency of the right-hemispheric N1m, the left-hemispheric N1m elicited by the tonally excited /a/ was some 10 ms earlier than that elicited by the periodic and the aperiodic excitation. The amplitude of the N1m in both hemispheres was systematically stronger to stimulation with natural periodic excitation. Also, stimulus type had a marked (up to 7 mm) effect on the source location of the N1m, with periodic excitation resulting in more anterior sources than aperiodic and tonal excitation. CONCLUSION: The auditory brain areas of the two hemispheres exhibit differential tuning to natural speech signals, observable already in the passive recording condition. The variations in the latency and strength of the auditory N1m response can be traced back to the spectral structure of the stimuli. More specifically, the combined effects of the harmonic comb structure originating from the natural voice excitation caused by the fluctuating vocal folds and the location of the formant frequencies originating from the vocal tract leads to asymmetric behaviour of the left and right hemisphere.