Contextual modulation of hippocampal activity during picture naming.

Picture naming is a standard task used to probe language processes in healthy and impaired speakers. It recruits a broad neural network of language related areas, among which the hippocampus is rarely included. However, the hippocampus could play a role during picture naming, subtending, for example, implicit learning of the links between pictured objects and their names. To test this hypothesis, we recorded hippocampal activity during plain picture naming, without memorization requirement; we further assessed whether this activity was modulated by contextual factors such as repetition priming and semantic interference. Local field potentials recorded from intracerebral electrodes implanted in the healthy hippocampi of epileptic patients revealed a specific and reliable pattern of activity, markedly modulated by repetition priming and semantic context. These results indicate that the hippocampus is recruited during picture naming, presumably in relation to implicit learning, with contextual factors promoting differential hippocampal processes, possibly subtended by different sub-circuitries.

The contribution of frequency-specific activity to hierarchical information processing in the human auditory cortex.

The fact that feed-forward and top-down propagation of sensory information use distinct frequency bands is an appealing assumption for which evidence remains scarce. Here we obtain human depth recordings from two auditory cortical regions in both hemispheres, while subjects listen to sentences, and show that information travels in each direction using separate frequency channels. Bottom-up and top-down propagation dominates in γ- and δ-ß (<40 Hz) bands, respectively. The predominance of low frequencies for top-down information transfer is confirmed by cross-regional frequency coupling, which indicates that the power of γ-activity in A1 is modulated by the phase of δ-ß activity sampled from association auditory cortex (AAC). This cross-regional coupling effect is absent in the opposite direction. Finally, we show that information transfer does not proceed continuously but by time windows where bottom-up or top-down processing alternatively dominates. These findings suggest that the brain uses both frequency- and time-division multiplexing to optimize directional information transfer.

Differential role of visuospatial working memory in the propensity toward uncertainty in patients with obsessive-compulsive disorder and in healthy subjects.

BACKGROUND: Obsessive-compulsive disorder (OCD) is associated with visuospatial working memory deficits. Intolerance of uncertainty is thought to be a core component of OCD symptoms. Recent findings argue for a possible relationship between abilities in visuospatial memory and uncertainty. However, this relationship remains unclear in both OCD patients and healthy subjects. To address this issue, we measured performance in visuospatial working memory and the propensity to express uncertainty during decision making. We assessed their relationship and the temporal direction of this relationship in both OCD patients and healthy subjects. METHOD: Baseline abilities in visuospatial working memory were measured with the Corsi block-tapping test. A delayed matching-to-sample task was used to identify explicit situations of certainty, uncertainty and ignorance and to assess continuous performance in visuospatial working memory. Behavioural variables were recorded over 360 consecutive trials in both groups. RESULTS: Baseline scores of visuospatial working memory did not predict the number of uncertain situations in OCD patients whereas they did in healthy subjects. Uncertain trials led to reduced abilities in visuospatial working memory to 65% of usual performance in OCD patients whereas they remained stable in healthy subjects. CONCLUSIONS: The present findings show an opposite temporal direction in the relationship between abilities in working memory and uncertainty in OCD patients and healthy subjects. Poor working memory performance contributes to the propensity to feel uncertainty in healthy subjects whereas uncertainty contributes to decreased continuous performance in working memory in OCD patients.

Navigated rTMS for the treatment of tinnitus: a pilot study with assessment by fMRI and AEPs.

OBJECTIVE: Repeated transcranial magnetic stimulation (rTMS) of auditory cortex has been proposed to treat refractory chronic tinnitus, but the involved mechanisms of action remain largely unknown. The purpose of this pilot study was to evaluate the impact of rTMS on auditory cortex activity in a series of tinnitus patients, using for the first time both functional magnetic resonance imaging (fMRI) of the brain and auditory evoked potentials (AEPs). METHOD: In six patients with chronic, lateralized refractory tinnitus, we performed five sessions of neuronavigated rTMS delivered at 1Hz over the secondary auditory cortex (defined on morphological MRI), contralateral to tinnitus side. The effects of rTMS were assessed on clinical scales, fMRI, and AEPs (N1 and P2 components). RESULTS: The clinical impact of rTMS on tinnitus was good for three patients (25-50% improvement of tinnitus severity compared to baseline), moderate for two patients (15% improvement), and null for one patient who had the most severe tinnitus at baseline. The changes induced by rTMS on fMRI data varied with the baseline level of auditory cortex activation before rTMS. This baseline level of activation was itself related to the severity of tinnitus. Thus, cortical stimulation increased auditory cortex activation in patients who had less severe tinnitus and low level of activation before rTMS, whereas it decreased auditory cortex activation in patients who had more severe tinnitus and higher level of activation before rTMS. Regarding AEPs, rTMS decreased N1 amplitude in all patients, except in the patient who had the most severe tinnitus at baseline and showed no improvement after rTMS. Conversely, P2 amplitude decreased after rTMS only in patients with severe tinnitus, at least for auditory stimulation contralateral to tinnitus, but increased in patients with less severe tinnitus. CONCLUSIONS: The changes produced by rTMS in auditory cortex activity, as assessed by fMRI and AEPs, appeared to depend on a process of disease-related homeostatic cortical plasticity, regardless of the therapeutic impact of rTMS on tinnitus.

Performance in recognition memory is correlated with entorhinal/perirhinal interictal metabolism in temporal lobe epilepsy.

In addition to the hippocampus, the entorhinal/perirhinal cortices are often involved in temporal lobe epilepsy (TLE). It has been proposed that these anterior parahippocampal structures play a key role in recognition memory. We studied the voxel-based PET correlation between number of correctly recognized targets in a new recognition memory paradigm and interictal cerebral metabolic rate for glucose, in 15 patients with TLE with hippocampal sclerosis. In comparison to healthy subjects, patients had decreased recognition of targets (P<0.001) and ipsilateral hypometabolism (relative to side of hippocampal sclerosis) of the hippocampus, entorhinal/perirhinal cortices, medial temporal pole, and middle temporal gyrus (P<0.05, corrected by false discovery rate method). Performance correlated with interictal metabolism of ipsilateral entorhinal/perirhinal cortices (P<0.005, Spearman's rank test), but this relationship was not significant in the hippocampus itself (P>0.18, Spearman's rank test). These findings highlight the preferential involvement of entorhinal/perirhinal cortices in recognition memory in patients with TLE, and suggest that recognition memory paradigms may be useful in assessing anterior parahippocampal functional status in TLE.

Asymmetry of voice onset time-processing in adult developmental dyslexics.

OBJECTIVE: The human auditory cortex codes speech temporally according to sequential acoustico-phonetic cues like the voice onset time (VOT). This coding is predominantly left-lateralized in normal readers. We examined VOT-processing asymmetries in adults with a history of developmental dyslexia (DD-history+). METHODS: Auditory-evoked potentials (AEPs) to voiced (/ba/) and voiceless (/pa/) speech stimuli were recorded from 10 DD-history+ adults and 8 controls. Source modelling of the "release component" (RC: approximately 240 ms; time-locked to voiced consonantal release and considered reflective of VOT-processing) was conducted to explore VOT asymmetries. RESULTS: Controls demonstrated L>R RC source probe amplitude asymmetry in the auditory cortex. DD-history+ subjects with little persistent reading deficit (n=5) demonstrated normal temporal coding but rightward asymmetry. DD-history+ subjects with severe persistent deficits (n=5) exhibited numerous supplemental AEP components (notably left hemispheric) and inconsistent asymmetry (leftward or alternating). CONCLUSIONS: These preliminary findings suggest that DD-history+ adults process auditory speech cues differently than adults without previous DD. The nature of this processing may relate to the severity of persistent reading deficits. SIGNIFICANCE: Previous dyslexics with little persistent deficit can exhibit atypical functional asymmetry with normal auditory temporal coding. Source modelling represents an effective, non-invasive means of exploring processing asymmetries in clinical populations.

Effects of GSM signals on auditory evoked responses.

The article presents a study of the influence of radio frequency (RF) fields emitted by mobile phones on human cerebral activity. Our work was based on the study of Auditory Evoked Potentials (AEPs) recorded on the scalp of healthy humans and epileptic patients. The protocol allowed us to compare AEPs recorded with or without exposure to RFs. To get a reference, a control session was also introduced. In this study, the correlation coefficients computed between AEPs, as well as the correlation coefficients between spectra of AEPs were investigated to detect a possible difference due to RFs. A difference in the correlation coefficients computed in control and experimental sessions was observed, but it was difficult to deduce the effect of RFs on human health.

Auditory evoked potential patterns to voiced and voiceless speech sounds in adult developmental dyslexics with persistent deficits.

Auditory evoked potentials (AEPs) were recorded from eight developmental dyslexic adults with persistent reading, spelling and phonological deficits, and 10 non-dyslexic controls to voiced (/ba/) and voiceless (/pa/) consonant-vowel syllables. Consistent with previous data, non-dyslexics coded these stimuli differentially according to the temporal cues that form the basis of the voiced/voiceless contrast: AEPs had time-locked components with latencies that were determined by the temporal structure of the stimuli. Dyslexics were characterized by one of two electrophysiological patterns: AEP pattern I dyslexics demonstrated a differential coding of stimuli on the basis of some temporal cues, but with an atypically large number of components and a considerable delay in AEP termination time; AEP pattern II dyslexics demonstrated no clear differential coding of stimuli on the basis of temporal cues. These data reveal the presence of anomalies in cortical auditory processing which could underlie persistent perceptual and linguistic impairments in some developmental dyslexics. Furthermore, scalp AEP distribution maps showing the difference observed between /ba/ and /pa/ activity over time suggest that the regions implicated in the processing of crucial time-related acoustic cues were not systematically lateralized to the left hemisphere like they were for non-dyslexics. These findings may be conducive to a better understanding and treatment of perceptual dysfunctions in developmental language disorders.

Analysis of auditory evoked potential parameters in the presence of radiofrequency fields using a support vector machines method.

The paper presents a study of global system for mobile (GSM) phone radiofrequency effects on human cerebral activity. The work was based on the study of auditory evoked potentials (AEPs) recorded from healthy humans and epileptic patients. The protocol allowed the comparison of AEPs recorded with or without exposure to electrical fields. Ten variables measured from AEPs were employed in the design of a supervised support vector machines classifier. The classification performance measured the classifier's ability to discriminate features performed with or without radiofrequency exposure. Most significant features were chosen by a backward sequential selection that ranked the variables according to their pertinence for the discrimination. Finally, the most discriminating features were analysed statistically by a Wilcoxon signed rank test. For both populations, the N100 amplitudes were reduced under the influence of GSM radiofrequency (mean attenuation of -0.36 microV for healthy subjects and -0.60 microV for epileptic patients). Healthy subjects showed a N100 latency decrease (-5.23 ms in mean), which could be consistent with mild, localised heating. The auditory cortical activity in humans was modified by GSM phone radiofrequencies, but an effect on brain functionality has not been proven.

Neuromagnetic source localization of auditory evoked fields and intracerebral evoked potentials: a comparison of data in the same patients.

OBJECTIVE: To compare the localizations of different neural sources (a) obtained from intracerebral evoked responses and (b) calculated from surface auditory evoked field responses recorded in the same subjects. Our aim was to evaluate the resolving power of a source localization method currently used in our laboratory, which is based on a recent spatio-temporal algorithm used in magneto-encephalography (MEG). METHODS: Auditory evoked responses were studied in 4 patients with medically intractable epilepsy. These responses were recorded from depth electrodes implanted in the auditory cortex for pre-surgical evaluation (stereo-electro-encephalography (SEEG)), as well as from surface captors (for MEG) placed on the scalp after removal of the depth electrodes. Auditory stimuli were clicks and short tone bursts with different frequencies. RESULTS: All middle-latency components (from 13 to 70 ms post-stimulus onset) were recorded and localized (via SEEG) along Heschl's gyrus (HG). MEG reliably localized Pam and P1m in the same area of HG that intracerebral recordings localized them in. No significant delay between SEEG and MEG latencies was observed. Both methods suggest that N1 is generated from different sources in the intermediate and lateral parts of the HG and in the planum temporale (PT). The source of P2 (PT and/or Area 22) remains unclear and was in one case, localized in different regions according to the method used. This latter component may therefore also be generated by different sources. CONCLUSIONS: The results suggest that both techniques are useful and may be used together in a complementary fashion. Intracerebral recordings allow the researcher to validate and interpret surface recordings.

Intracerebral evoked potentials in pitch perception reveal a functional asymmetry of the human auditory cortex.

One acoustic feature that plays an important role in pitch perception is frequency. Studies on the processing of frequency in the human and animal brain have shown that the auditory cortex is tonotopically organized: low frequencies are represented laterally whereas high frequencies are represented medially. To date, the study of the functional organization of the human auditory cortex in the processing of frequency has been limited to the use of either scalp-recorded auditory evoked potentials (AEPs), which have relatively poor spatial resolving power, or functional imagery techniques, which have poor temporal resolving power. The present study uses intracerebrally recorded AEPs to explore this topic in the primary and secondary auditory cortices of both hemispheres of the human brain. Recordings were carried out in 45 adult patients with drug-resistant partial seizures. In the right hemisphere, clear spectrally organized tonotopic maps were observed with distinct separations between different frequency-processing regions. AEPs for high frequencies were recorded medially, whereas AEPs for low frequencies were recorded laterally. In the left hemisphere, however, this tonotopic organization was less evident, with different regions involved in the processing of a range of frequencies. The hemisphere-related difference in the processing of tonal frequency is discussed in relation to pitch perception.

Identification reaction times of voiced/voiceless continua: a right-ear advantage for VOT values near the phonetic boundary.

We explored the degree to which the duration of acoustic cues contributes to the respective involvement of the two hemispheres in the perception of speech. To this end, we recorded the reaction time needed to identify monaurally presented natural French plosives with varying VOT values. The results show that a right-ear advantage is significant only when the phonetic boundary is close to the release burst, i.e., when the identification of the two successive acoustical events (the onset of voicing and the release from closure) needed to perceive a phoneme as voiced or voiceless requires rapid information processing. These results are consistent with the recent hypothesis that the left hemisphere is superior in the processing of rapidly changing acoustical information.

[Functional imaging of the auditory cortex: role of magnetoencephalography]. / Imagerie fonctionelle du cortex auditif. Place de la magnétoencéphalographie.

We present the different methods for detecting cerebral activity in the auditory cortex. Positron emission tomography (PET) and functional MRI (fMRI) measure such activity indirectly by calculating the blood flow rate or the consumption of oxygen. The direct methods of detection record the electrical or magnetic activity by eletroencephalography (EEG), or magnetoencephalography (MEG), respectively. The aim of this study was to define the role of MEG amongst these different techniques using the data from recordings of evoked magnetic fields in 5 healthy subjects. The localizations demonstrated the tonotopic organization of the auditory cortex, with high-pitched sounds showing a more medial cortical projection than low-pitched sounds. These various techniques are complementary. PET allows a pharmacological study of the cortex, and could be used in patients with cochlear implants. FMRI is non-invasive, and has a high spatial resolution. EEG has an excellent temporal resolution, and EEG recordings do not require major equipment or infrastructure. MEG has a better spatial resolution, with the same temporal resolution, as EEC. MEG is particularly useful in the localization of the cortical generators of middle-latency auditory evoked responses.

Specialization of left auditory cortex for speech perception in man depends on temporal coding.

Speech perception requires cortical mechanisms capable of analysing and encoding successive spectral (frequency) changes in the acoustic signal. To study temporal speech processing in the human auditory cortex, we recorded intracerebral evoked potentials to syllables in right and left human auditory cortices including Heschl's gyrus (HG), planum temporale (PT) and the posterior part of superior temporal gyrus (area 22). Natural voiced /ba/, /da/, /ga/) and voiceless (/pa/, /ta/, /ka/) syllables, spoken by a native French speaker, were used to study the processing of a specific temporally based acoustico-phonetic feature, the voice onset time (VOT). This acoustic feature is present in nearly all languages, and it is the VOT that provides the basis for the perceptual distinction between voiced and voiceless consonants. The present results show a lateralized processing of acoustic elements of syllables. First, processing of voiced and voiceless syllables is distinct in the left, but not in the right HG and PT. Second, only the evoked potentials in the left HG, and to a lesser extent in PT, reflect a sequential processing of the different components of the syllables. Third, we show that this acoustic temporal processing is not limited to speech sounds but applies also to non-verbal sounds mimicking the temporal structure of the syllable. Fourth, there was no difference between responses to voiced and voiceless syllables in either left or right areas 22. Our data suggest that a single mechanism in the auditory cortex, involved in general (not only speech-specific) temporal processing, may underlie the further processing of verbal (and non-verbal) stimuli. This coding, bilaterally localized in auditory cortex in animals, takes place specifically in the left HG in man. A defect of this mechanism could account for hearing discrimination impairments associated with language disorders.

Contribution of different cortical areas in the temporal lobes to music processing.

Music processing ability was studied in 65 right-handed patients who had undergone unilateral temporal cortectomy for the relief of intractable epilepsy, and 24 matched normal controls. The ability to recognize changes in note intervals and to distinguish between different rhythms and metres was tested by presentation of sequences of simple musical phrases with variations in either pitch or temporal dimensions. The responses (right or wrong) enabled us to determine in which component of the music processing mechanism the patients had deficits and hence, knowing the positions of the surgical lesions, to identify their separate cerebral locations. The results showed that a right temporal cortectomy impaired the use of both contour and interval information in the discrimination of melodies and a left temporal cortectomy impaired only the use of interval information. Moreover, they underlined the importance of the superior temporal gyrus in melody processing. The excision of a part of the auditory areas (posterior part of the superior temporal gyrus) was found to be most detrimental for pitch and temporal variation processing. In the temporal dimension, we observed a dissociation between metre and rhythm and the critical involvement of the anterior part of the superior temporal gyrus in metric processing. This study highlights the relevance of dissociating musical abilities into their most significant cognitive components in order to identify their separate cerebral locations.

Pitch perception: a difference between right- and left-handed listeners.

Various results indicate that the perception of a complex tone's "virtual" pitch is generally lateralized in the right cerebral hemisphere. The primary aim of this work was to test the hypothesis that this is not the case for the "spectral" pitch percepts induced by complex tones. Forty right-handed and 18 left-handed listeners were monaurally presented with pairs of successive tones made up of n consecutive equal-amplitude harmonics of a missing fundamental (F0). n varied from two to four across subjects. In "test" conditions, the paired tones differed in F0 but the spectral components of the tone with the lower F0 were higher in frequency than the corresponding components of the other tone (except for one component, which was identical). The subjects had to say if, from one tone to the other, pitch rose or fell. From such judgements, one could infer that the pitch dominantly perceived in each tone was a virtual pitch (corresponding to F0) or a spectral pitch (i.e., the pitch of a single spectral component, or a perceptual quality corresponding to the centroid of the power spectrum). For n = 2, the results indicated that virtual pitch was less salient than spectral pitch; the opposite occurred for n = 3 and n = 4. The ear (left or right) to which the stimuli were presented had some influence on the judgements, in the expected direction. However, this influence was not a robust one. Unexpectedly, a reliable effect of the listeners' handedness was observed: for each value of n, the judgements indicating virtual pitch perception were less frequent in the left-handers than in the right-handers. Discrimination performances measured in "control" conditions showed that the handedness factor was not confounded with a factor of frequency discrimination ability.

[Cortical mechanisms of auditive perception in man: contribution of cerebral potentials and evoked magnetic fields by auditive stimulations]. / Etude des mécanismes corticaux de la perception auditive chez l'homme: apport des potentiels intracérébraux et des champs magnétiques évoqués par les stimulations auditives.

The goal of this study is to determine and localize the generators of different components of middle latency auditory evoked potentials (MLAEPs) through intracerebral recordings in auditory cortex in Human (Heschl's gyrus and Planum Temporale). The intracerebral data show that the generators of components at 30, 50, 60 and 75 msec latency are distributed medio-laterally along the Heschl's gyrus. The 30 msec component is generated in the dorso-postero-medial part of the Heschl's gyrus (primary area) and the 50 msec component is generated laterally in the primary area. The generators of the later components (60-75 msec) are localized in the lateral part of the Heschl's gyrus that are the secondary areas. The comparison with the generators of the components of the magnetic auditory evoked field and the tonotopic organization of the auditory cortex are discussed.

Evoked potentials recorded from the auditory cortex in man: evaluation and topography of the middle latency components.

The goal of this study is to determine and localize the generators of different components of middle latency auditory evoked potentials (MLAEPs) through intracerebral recording in auditory cortex in man (Heschl's gyrus and planum temporale). The present results show that the generators of components at 30, 50, 60 and 75 msec latency are distributed medio-laterally along Heschl's gyrus. The 30 msec component is generated in the dorso-postero-medial part of Heschl's gyrus (primary area) and the 50 msec component is generated laterally in the primary area. The generators of the later components (60-75 msec) are localized in the lateral part of Heschl's gyrus that forms the secondary areas. The localization of N100 generators is discussed.

Localization of the primary auditory area in man.

The localization of the primary auditory cortex in man was studied by direct recordings in 150 different sites in the superior transverse gyrus, especially in Heschl's gyrus and the planum temporale. The distribution of the primary evoked responses (N13/P17/N26) was studied in 15 epileptic patients who were candidates for surgical treatment. Precise topography of recording sites was determined stereotactically. Our results provide evidence for considering only a restricted portion of Heschl's gyrus (its posteromedial part) as the primary auditory area.