Interaction among the components of multiple auditory steady-state responses: enhancement in tinnitus patients, inhibition in controls.

Amplitude and phase of steady-state signals recorded in response to amplitude-modulated (AM) sine tones vary over time, suggesting that the steady-state response (SSR) reflects not only stimulus input but also its interaction with other input streams or internally generated signals. Alterations of the interaction between simultaneous SSRs associated with tinnitus were studied by recording the magnetic field evoked by AM-tones with one of three carrier and one of three modulation frequencies. Single AM-tones were presented in single presentation mode and superpositions of three AM-tones differing in carrier and modulation frequency in multiple presentation mode. Modulation frequency-specific SSR components were recovered by bandpass filtering. Compared with single mode, in multiple mode SSR amplitude was reduced in healthy controls, but increased in tinnitus patients. Thus, while in controls multiple response components seem to reciprocally inhibit one another, in tinnitus reciprocal facilitation seems to predominate. Reciprocal inhibition was unrelated to the phase coherence among SSR components, but was correlated with the frequency of phase slips, indicating that the lateral interaction among SSR components acts in a quasi-paroxysmal manner and manifests itself in terms of a random train of phase reset events. Phase slips were more frequent in patients than controls both in single and multiple mode. Together, these findings indicate that lateral or surround inhibition of single units in auditory cortex is reduced and suggest that in-field inhibition is increased in tinnitus.

Silent echo-planar imaging for auditory FMRI.

INTRODUCTION: The effect of the acoustic scanner noise produced by gradient coil switching on the auditory evoked BOLD signal represents a well-known problem in auditory functional MRI (FMRI). In this paper, a new low-noise echo-planar imaging (EPI) sequence is presented that is optimized for auditory FMRI measurements. METHODS: The sequence produces a narrow-band acoustic frequency spectrum by using a sinusoidal readout echo train and a constant phase encoding gradient. This narrow band is adapted to the frequency response function of the MR scanner by varying the switching frequency of the sinusoidal readout gradient. RESULTS: Compared to a manufacturer-provided standard EPI sequence, the acoustic noise reduction amounts to up to 20 dBA. Using a simple block design paradigm contrasting presentation of a pure tone during ON blocks and "silence" (absence of the tone) during OFF blocks, the new low-noise sequence was evaluated and compared to the standard EPI sequence. Statistical parametric mapping (SPM) resulted in higher levels of significance of auditory activation for the low-noise sequence. DISCUSSION: These findings strongly suggest that the low-noise sequence may generate enhanced BOLD contrasts compared to the standard EPI sequences commonly used in FMRI.

Extinction training for tinnitus.

Recent accounts of tinnitus development and maintenance assign an important role to central mechanisms. Residual inhibition is a frequent phenomenon in individuals with tinnitus, and refers to the fact that tinnitus can temporarily be reduced by presenting sounds or noises that inhibit tinnitus for a limited time even after termination of the sound. This kind of stimulation-induced inhibition of tinnitus could potentially be used for treatment by combining it with additional interventions to enhance the extinction of tinnitus. Here we propose a training program aimed at the amplification and the extension in time of residual inhibition as well as the extinction of negative emotional responses to the tinnitus. The program is tested alone or in combination with a pharmacological intervention that is aimed at decreasing central hyperactivity. Treatment effects are assessed by tinnitus questionnaires, electroencephalographic measures (reduction in the amplitude of the N(100) component of the event-related potential as an indicator of habituation) as well as skin conductance responses to 1000 Hz tones or tinnitus-like tones. This training is an example of the use of centrally acting and mechanism-oriented tinnitus treatments.

Multiple frequency steady-state evoked magnetic field mapping of digit representation in primary somatosensory cortex.

Magnetic source imaging of multiple frequency steady-state somatosensory evoked responses was examined using a 151-channel magnetoencephalography (MEG) system and a dual-channel electrical stimulator. Somatotopy of digit representation was studied in healthy subjects and effects of injury-related cortical plasticity in patients with unilateral transections of the median or the ulnar nerve. Dipole source locations exhibited somatotopic order with overlap between neighboring digits. In two of three nerve injury patients evidence for cortical reorganization was found. The location of sources related to digits neighboring deafferented digits was changed and their dipole moments were enlarged by comparsion with the sources related to contralateral homologue control digits. As a basis for magnetic source imaging, the recording of multiple frequency somatosensory steady-state evoked responses may be a viable and time saving alternative to the recording of transient evoked responses.

Topographic and temporal indices of vowel spectral envelope extraction in the human auditory cortex.

The auditory-evoked neuromagnetic field elicited by single vowel formants and two-formant vowels was recorded under active listening conditions using a 37-channel magnetometer. There were three single formants with formant frequencies of 200, 400, and 800 Hz, another single formant with a formant frequency of 2600 Hz, and three vowels that were constructed by linear superimposition of the high- onto one of the low-frequency formants. P50 m and N100 m latency values were inversely correlated with the formant frequency of single formants. A strong effect of formant frequency on source location was obtained along the postero-anterior axis, which is orthogonal to the well-established latero-medial tonotopic gradient. Regardless of whether single formants or first formants of vowels were considered, N100 m sources were more anterior and sustained field sources were more posterior for higher-frequency than for lower-frequency formants. The velocity of the apparent posterior-to-anterior movement across cortical surface of N100 m sources first reported by Rogers et al. [Rogers, R. L., Papanicolaou, A. C., Baumann, S. B., Saydjari, C., & Eisenberg, H. M. (1990). Neuromagnetic evidence of a dynamic excitation pattern generating the N100 auditory response. Electroencephalography and Clinical Neurophysiology,77, 237-240] decreased as a function of latency. The amount of deceleration was positively correlated with formant frequency. Responses to the vowels were superadditive, indicating that the processes elicited by the constituents of composite stimuli interact at one or more stages of the afferent auditory pathway. Such interaction may account for the absence of a lateral-to-medial tonotopic mapping of first formant frequency. The source topography found may reflect activity in auditory fields adjacent to AI with the strength of the contribution varying with formant frequency. Alternatively, it may reflect sharpness-of-tuning and inhibitory response-area asymmetry gradients along isofrequency stripes within AI. Either alternative may be interpreted in terms of a spectral blurring mechanism that abstracts spectral envelope information from the details of spectral composition, an important step towards the formation of invariant phonetic percepts.

Involvement of the human frontal eye field and multiple parietal areas in covert visual selection during conjunction search.

Searching for a target object in a cluttered visual scene requires active visual attention if the target differs from distractors not by elementary visual features but rather by a feature conjunction. We used functional magnetic resonance imaging (fMRI) in human subjects to investigate the functional neuroanatomy of attentional mechanisms employed during conjunction search. In the experimental condition, subjects searched for a target defined by a conjunction of colour and orientation. In the baseline condition, subjects searched for a uniquely coloured target, regardless of its orientation. Eye movement recordings outside the scanner verified subjects' ability to maintain fixation during search. Reaction times indicated that the experimental condition was attentionally more demanding than the baseline condition. Differential activations between conditions were therefore ascribed to top-down modulation of neural activity. The frontal eye field, the ventral precentral sulcus and the following posterior parietal regions were consistently activated: (i) the postcentral sulcus; (ii) the posterior; and (iii) the anterior part of the intraparietal sulcus; and (iv) the junction of the intraparietal with the transverse occipital sulcus. Parietal regions were spatially distinct and displayed differential amplitudes of signal increase with a maximal amplitude in the posterior intraparietal sulcus. Less consistent activation was found in the lateral fusiform gyrus. These results suggest an involvement of the human frontal eye field in covert visual selection of potential targets during search. These results also provide evidence for a subdivision of posterior parietal cortex in multiple areas participating in covert visual selection, with a major contribution of the posterior intraparietal sulcus.

Measuring the perceptual magnet effect in the perception of /i/ by German listeners.

In a study of the internal category structure of the vowel /i/, Kuhl found a "perceptual magnet effect": Discrimination sensitivity was poorer for category instances that were acoustically similar to the category prototype than it was for category instances that were not. The typicality of category exemplars was determined by goodness judgments and was found to correlate with the acoustics of average production. Analysis and interpretation of discrimination performance relied on two important assumptions: that listeners perceived all stimuli presented as exemplars of the same vowel category and that, apart from the influence of phonetic coding, discrimination sensitivity was the same across the investigated part of the vowel space. In the present study, it is shown that production and perception estimates of the category prototype may diverge, possibly because listeners seem to prefer hyperarticulated variants of vowel categories. An approach towards measurement of intra-category discrimination minima is put forward and tested that protects against intercategory confounds and avoids the isosensitivity assumption.

The magnetic mismatch field elicited by words and phonological non-words.

The speech-evoked magnetic mismatch field was measured using a 49-channel gradiometer. The standard stimuli were words in one condition and phonological non-words in another condition. The deviants were non-words throughout. The equivalent current dipole fitted to the mismatch field was deeper inside the brain and its dipole moment was stronger for non-word than word standards. The factor structure of field amplitude, source dipole moment, and depth suggested that the lexicality conditions differed in source surface area and depth, but not in source current density. This lexicality effect is compatible with a modular rather than an interactive view of the relationship between lexical and phonetic representation.

Magnetic fields elicited by tones and vowel formants reveal tonotopy and nonlinear summation of cortical activation.

A long-latency response component (N1m) and the sustained field (SF) of the auditory evoked magnetic field elicited by two composite stimuli (a two-tone combination and a two-formant vowel) and their individually presented components (a 600-Hz and a 2100-Hz pure tone and two single-vowel formants with formant frequencies matched to the tone frequencies) were recorded using a 37-channel magnetometer. The response to the composite stimuli differed from the linear sum of the responses to the respective components in latency, equivalent dipole moment, and equivalent dipole location, suggesting an interaction among the processes elicited by the constituents of composite stimuli. N1m and SF source locations were more medial for the response to the high tone than to the low tone and more medial for the response to the high vowel formant than to the low vowel formant. The N1m formant sources were more lateral than the N1m tone sources. These findings suggest that, at the level of the auditory cortex, vowels are represented in terms of both the spectral pitches determined by their most prominent harmonics and, within the latency range of the N1m, the virtual pitch determined by the spacing of the harmonics.

Magnetic mismatch fields elicited by vowels and consonants.

Auditory-evoked mismatch fields (MMFs) elicited by vowel contrasts and plosive stop consonant place-of-articulation contrasts were recorded over the left hemisphere of neurologically and audiologically normal subjects. Two experiments were conducted: vowels were presented in isolation in experiment 1 and embedded in consonant-vowel syllables in experiment 2. Bestfit equivalent MMF sources were obtained using the model of a single, spatiotemporal current dipole in a sphere. In both experiments, MMF sources activated by place-of-articulation contrasts were later in latency and smaller in dipole moment amplitude than MMF sources excited by vowel contrasts. There was evidence, albeit not unambiguous, for the vowel-contrast MMF sources being located more posteriorly than the consonant-contrast MMF sources in experiment 1 and more laterally in experiment 2. In both experiments, the MMF source excited by the contrast between /da/ and /ga/ was more anterior than the MMF source excited by the contrast between /da/ and /ba/. The effects on latency and dipole moment may be interpreted to mirror differences in perceptual discriminability and auditory memory decay between consonantal place-of-articulation contrasts and vowel contrasts. Similarly, the effects on location may be interpreted to reflect featural specificity of the mismatch response. Interestingly, the dipole source analysis results show a correspondence to the pattern of preservation and loss of the mismatch response to vowel and consonant place-of-articulation contrasts recently observed in Wernicke's aphasia.

The neurotopography of vowels as mirrored by evoked magnetic field measurements.

The auditory evoked neuromagnetic field elicited by synthetic specimens of the vowels [a], [ae], [u], and [i] was recorded over the left and the right hemisphere of 11 subjects. The N100m and the SF deflection of the recorded signal was submitted to equivalent current source analysis using the model of a single dipole in a spherical volume conductor. Vowel processing was hypothesized to occur in a multistage process rendering a sequence of representations of the acoustic input. Vowel representations were considered to differ among each other in the features they make salient, thus, in the kind of dissimilarity relationship they establish, and, by implication, in terms of the vowel space defined by the respective set of dissimilarities. It was investigated whether a mapping exists between at least one of a number of hypothetical vowel spaces and the cortical response space spanned by the spatial distribution of vowel evoked equivalent current dipoles. Although the spatial configuration of vowel evoked sources proved to be highly variable across subjects, the ordering of distances between N100m and SF equivalent current dipole locations turned out to correspond to the ordering of distances between the corners of a vowel trapezium. There were some, albeit weak, indications of hemispheric differences in vowel processing. The results suggest that the spatial distribution of the equivalent current dipole sources of both the N100m and the SF deflection of the neuromagnetic field elicited by vowels reflect a processing stage transitional between auditory and phonetic representation.

Left and right hemifield advantages of fusions and combinations in audiovisual speech perception.

If a place-of-articulation contrast is created between the auditory and the visual component syllables of videotaped speech, frequently the syllable that listeners report they have heard differs phonetically from the auditory component. These "McGurk effects", as they have come to be called, show that speech perception may involve some kind of intermodal process. There are two classes of these phenomena: fusions and combinations. Perception of the syllable /da/ when auditory /ba/ and visual /ga/ are presented provides a clear example of the former, and perception of the string /bga/ after presentation of auditory /ga/ and visual /ba/ an unambiguous instance of the latter. Besides perceptual fusions and combinations, hearing visually presented component syllables also shows an influence of vision on audition. It is argued that these "visual" responses arise from basically the same underlying processes that yield fusions and combinations, respectively. In the present study, the visual component of audiovisually incongruous CV-syllables was presented in the left and the right visual hemifield, respectively. Audiovisual fusion responses showed a left hemifield advantage, and audiovisual combination responses a right hemifield advantage. This finding suggests that the process of audiovisual integration differs between audiovisual fusions and combinations and, furthermore, that the two cerebral hemispheres contribute differentially to the two classes of response.

Magnetic and electric brain activity evoked by the processing of tone and vowel stimuli.

Sustained magnetic and electric brain waves may reflect linguistic processing when elicited by auditory speech stimuli. In the present study, only in the latency interval subsequent to the N1m/N1 has a sensitivity of brain responses to features of speech been demonstrated. We conclude this from studying the auditory-evoked magnetic field (AEF) and the corresponding evoked potential (AEP) in response to vowels and a tone. Brain activity was recorded from the left and the right hemisphere of 11 subjects. Three aspects of transient activity were examined: (1) the amplitudes and source characteristics of the N1m component of the AEF; (2) the amplitudes and source characteristics of the sustained field (SF), and (3) the corresponding amplitude characteristics of the AEP. Sustained potential amplitudes and SF root mean square amplitudes, as well as the dipole strength of the SF source, were found to be larger for vowel-evoked signals than for signals elicited by the tone stimulus. The amplitude and dipole strength effects had an interaction with hemisphere, with larger interhemispheric differences for the vowel condition, as well as larger tone-vowel differences of these parameters in the speech-dominant left hemisphere. No statistically significant hemisphere-by-stimulus-type interactions were found in N1/N1m amplitudes and N1m source parameters.