Towards formal models of inhibitory mechanisms involved in motor imagery: a commentary on Bach et al. (2022).

A vast body of research suggests that the primary motor cortex is involved in motor imagery. This raises the issue of inhibition: how is it possible for motor imagery not to lead to motor execution? Bach et al. (Psychol Res Psychol Forschung. 10.1007/s00426-022-01773-w, 2022, this issue) suggest that the motor execution threshold may be "upregulated" during motor imagery to prevent execution. Alternatively, it has been proposed that, in parallel to excitatory mechanisms, inhibitory mechanisms may be actively suppressing motor output during motor imagery. These theories are verbal in nature, with well-known limitations. Here, we describe a toy-model of the inhibitory mechanisms thought to be at play during motor imagery to start disentangling predictions from competing hypotheses.

An open-source toolbox for Multi-patient Intracranial EEG Analysis (MIA).

Intracranial EEG (iEEG) performed during the pre-surgical evaluation of refractory epilepsy provides a great opportunity to investigate the neurophysiology of human cognitive functions with exceptional spatial and temporal precisions. A difficulty of the iEEG approach for cognitive neuroscience, however, is the potential variability across patients in the anatomical location of implantations and in the functional responses therein recorded. In this context, we designed, implemented, and tested a user-friendly and efficient open-source toolbox for Multi-Patient Intracranial data Analysis (MIA), which can be used as standalone program or as a Brainstorm plugin. MIA helps analyzing event related iEEG signals while following good scientific practice recommendations, such as building reproducible analysis pipelines and applying robust statistics. The signals can be analyzed in the temporal and time-frequency domains, and the similarity of time courses across patients or contacts can be assessed within anatomical regions. MIA allows visualizing all these results in a variety of formats at every step of the analysis. Here, we present the toolbox architecture and illustrate the different steps and features of the analysis pipeline using a group dataset collected during a language task.

Inter-individual variability in dorsal stream dynamics during word production.

The current standard model of language production involves a sensorimotor dorsal stream connecting areas in the temporo-parietal junction with those in the inferior frontal gyrus and lateral premotor cortex. These regions have been linked to various aspects of word production such as phonological processing or articulatory programming, primarily through neuropsychological and functional imaging group studies. Most if not all the theoretical descriptions of this model imply that the same network should be identifiable across individual speakers. We tested this hypothesis by quantifying the variability of activation observed across individuals within each dorsal stream anatomical region. This estimate was based on electrical activity recorded directly from the cerebral cortex with millisecond accuracy in awake epileptic patients clinically implanted with intracerebral depth electrodes for pre-surgical diagnosis. Each region's activity was quantified using two different metrics-intra-cerebral evoked related potentials and high gamma activity-at the level of the group, the individual and the recording contact. The two metrics show simultaneous activation of parietal and frontal regions during a picture naming task, in line with models that posit interactive processing during word retrieval. They also reveal different levels of between-patient variability across brain regions, except in core auditory and motor regions. The independence and non-uniformity of cortical activity estimated through the two metrics push the current model towards sub-second and sub-region explorations focused on individualized language speech production. Several hypotheses are considered for this within-region heterogeneity.

Magnetoencephalography can reveal deep brain network activities linked to memory processes.

Recording from deep neural structures such as hippocampus noninvasively and yet with high temporal resolution remains a major challenge for human neuroscience. Although it has been proposed that deep neuronal activity might be recordable during cognitive tasks using magnetoencephalography (MEG), this remains to be demonstrated as the contribution of deep structures to MEG recordings may be too small to be detected or might be eclipsed by the activity of large-scale neocortical networks. In the present study, we disentangled mesial activity and large-scale networks from the MEG signals thanks to blind source separation (BSS). We then validated the MEG BSS components using intracerebral EEG signals recorded simultaneously in patients during their presurgical evaluation of epilepsy. In the MEG signals obtained during a memory task involving the recognition of old and new images, we identified with BSS a putative mesial component, which was present in all patients and all control subjects. The time course of the component selectively correlated with stereo-electroencephalography signals recorded from hippocampus and rhinal cortex, thus confirming its mesial origin. This finding complements previous studies with epileptic activity and opens new possibilities for using MEG to study deep brain structures in cognition and in brain disorders.

On the Boundaries between Decision and Action: Effector-selective Lateralization of Beta-frequency Power Is Modulated by the Lexical Frequency of Printed Words.

Current computational and neuroscientific models of decision-making posit a discrete, serial processing distinction between upstream decisional stages and downstream processes of motor-response implementation. We investigated this framework in the context of two-alternative forced-choice tasks on linguistic stimuli, words and pseudowords. In two experiments, we assessed the impact of lexical frequency and action semantics on two effector-selective EEG indexes of motor-response activation: the lateralized readiness potential and the lateralization of beta-frequency power. This allowed us to track potentially continuous streams of processing progressively mapping the evaluation of linguistic stimuli onto corresponding response channels. Whereas action semantics showed no influence on EEG indexes of motor-response activation, lexical frequency affected the lateralization of response-locked beta-frequency power. We argue that these observations point toward a continuity between linguistic processing of word input stimuli and implementation of corresponding choice in terms of motor behavior. This interpretation challenges the commonly held assumption of a discrete processing distinction between decisional and motor-response processes in the context of decisions based on symbolic stimuli.

Contributions of electrophysiology for identifying cortical language systems in patients with epilepsy.

A crucial element of the surgical treatment of medically refractory epilepsy is to delineate cortical areas that must be spared in order to avoid clinically relevant neurological and neuropsychological deficits postoperatively. For each patient, this typically necessitates determining the language lateralization between hemispheres and language localization within hemisphere. Understanding cortical language systems is complicated by two primary challenges: the extent of the neural tissue involved and the substantial variability across individuals, especially in pathological populations. We review the contributions made through the study of electrophysiological activity to address these challenges. These contributions are based on the techniques of magnetoencephalography (MEG), intracerebral recordings, electrical-cortical stimulation (ECS), and the electrovideo analyses of seizures and their semiology. We highlight why no single modality alone is adequate to identify cortical language systems and suggest avenues for improving current practice.

A description of verbal and gestural communication during postictal aphasia.

Patients suffering from drug-resistant temporal lobe epilepsy show substantial language deficits (i.e., anomia) during their seizures and in the postictal period (postictal aphasia). Verbal impairments observed during the postictal period may be studied to help localizing the epileptogenic zone. These explorations have been essentially based on simple tasks focused on speech, thus disregarding the multimodal nature of verbal communication, particularly the fact that, when speakers want to communicate, they often produce gestures of various kinds. Here, we propose an innovative procedure for testing postictal language and communication abilities, including the assessment of co-speech gestures. We provide a preliminary description of the changes induced on communication during postictal aphasia. We studied 21 seizures that induced postictal aphasia from 12 patients with drug-refractory epilepsy, including left temporal and left frontal seizures. The experimental task required patients to memorize a highly detailed picture and, briefly after, to describe what they had seen, thus eliciting a communicative meaningful monologue. This allowed comparing verbal communication in postictal and interictal conditions within the same individuals. Co-speech gestures were coded according to two categories: "Rhythmic" gestures, thought to be produced in support of speech building, and "illustrative" gestures, thought to be produced to complement the speech content. When postictal and interictal conditions were compared, there was decreased speech flow along with an increase of rhythmic gesture production at the expense of illustrative gesture production. The communication patterns did not differ significantly after temporal and frontal seizures, yet they were illustrated separately, owing to the clinical importance of the distinction, along with considerations of interindividual variability. A contrast between rhythmic and illustrative gestures production is congruent with previous literature in which rhythmic gestures have been linked to lexical retrieval processes. If confirmed in further studies, such evidence for a facilitative role of co-speech gestures in language difficulties could be put to use in the context of multimodal language therapies.

Tracking Keystroke Sequences at the Cortical Level Reveals the Dynamics of Serial Order Production.

Response selection is often studied by examining single responses, although most actions are performed within an overarching sequence. Understanding processes that order and execute items in a sequence is thus essential to give a complete picture of response selection. In this study, we investigate response selection by comparing single responses and response sequences as well as unimanual and bimanual sequences. We recorded EEG while participants were typing one- or two-keystroke sequences. Irrespective of stimulus modality (visual or auditory), response-locked analysis revealed distinct contralateral and ipsilateral components previously associated with activation and inhibition of alternative responses. Unimanual sequences exhibited a similar activation/inhibition pattern as single responses, but with the activation component of the pattern expressed more strongly, reflecting the fact that the hand will be used for two strokes. In contrast, bimanual sequences were associated with successive activation of each of the corresponding motor cortices controlling each keystroke and no traceable inhibitory component. In short, the activation component of the two-keystroke sequence EEG pattern can be understood from the addition of activation components of single-stroke sequences; the inhibition of the hand not being used is only evidenced when that hand is not planned for the next stroke.

Cortical Dynamics of Semantic Priming and Interference during Word Production: An Intracerebral Study.

Language production requires that semantic representations are mapped to lexical representations on the basis of the ongoing context to select the appropriate words. This mapping is thought to generate two opposing phenomena, "semantic priming," where multiple word candidates are activated, and "interference," where these word activities are differentiated to make a goal-relevant selection. In previous neuroimaging and neurophysiological research, priming and interference have been associated to activity in regions of a left frontotemporal network. Most of such studies relied on recordings that either have high temporal or high spatial resolution, but not both. Here, we employed intracerebral EEG techniques to explore with both high resolutions, the neural activity associated with these phenomena. The data came from nine epileptic patients who were stereotactically implanted for presurgical diagnostics. They performed a cyclic picture-naming task contrasting semantically homogeneous and heterogeneous contexts. Of the 84 brain regions sampled, 39 showed task-evoked activity that was significant and consistent across two patients or more. In nine of these regions, activity was significantly modulated by the semantic manipulation. It was reduced for semantically homogeneous contexts (i.e., priming) in eight of these regions, located in the temporal ventral pathway as well as frontal areas. Conversely, it was increased only in the pre-SMA, notably at an early poststimulus temporal window (200-300 msec) and a preresponse temporal window (700-800 msec). These temporal effects respectively suggest the pre-SMA's role in initial conflict detection (e.g., increased response caution) and in preresponse control. Such roles of the pre-SMA are traditional from a history of neural evidence in simple perceptual tasks, yet are also consistent with recent cognitive lexicosemantic theories that highlight top-down processes in language production. Finally, although no significant semantic modulation was found in the ACC, future intracerebral EEG work should continue to inspect ACC with the pre-SMA.

An intracerebral exploration of functional connectivity during word production.

Language is mediated by pathways connecting distant brain regions that have diverse functional roles. For word production, the network includes a ventral pathway, connecting temporal and inferior frontal regions, and a dorsal pathway, connecting parietal and frontal regions. Despite the importance of word production for scientific and clinical purposes, the functional connectivity underlying this task has received relatively limited attention, and mostly from techniques limited in either spatial or temporal resolution. Here, we exploited data obtained from depth intra-cerebral electrodes stereotactically implanted in eight epileptic patients. The signal was recorded directly from various structures of the neocortex with high spatial and temporal resolution. The neurophysiological activity elicited by a picture naming task was analyzed in the time-frequency domain (10-150 Hz), and functional connectivity between brain areas among ten regions of interest was examined. Task related-activities detected within a network of the regions of interest were consistent with findings in the literature, showing task-evoked desynchronization in the beta band and synchronization in the gamma band. Surprisingly, long-range functional connectivity was not particularly stronger in the beta than in the high-gamma band. The latter revealed meaningful sub-networks involving, notably, the temporal pole and the inferior frontal gyrus (ventral pathway), and parietal regions and inferior frontal gyrus (dorsal pathway). These findings are consistent with the hypothesized network, but were not detected in every patient. Further research will have to explore their robustness with larger samples.

Multi-factor analysis in language production: Sequential sampling models mimic and extend regression results.

For multi-factor analyses of response times, descriptive models (e.g., linear regression) arguably constitute the dominant approach in psycholinguistics. In contrast empirical cognitive models (e.g., sequential sampling models, SSMs) may fit fewer factors simultaneously, but decompose the data into several dependent variables (a multivariate result), offering more information to analyze. While SSMs are notably popular in the behavioural sciences, they are not significantly developed in language production research. To contribute to the development of this modelling in language, we (i) examine SSMs as a measurement modelling approach for spoken word activation dynamics, and (ii) formally compare SSMs to the default method, regression. SSMs model response activation or selection mechanisms in time, and calculate how they are affected by conditions, persons, and items. While regression procedures also model condition effects, it is only in respect to the mean RT, and little work has been previously done to compare these approaches. Through analyses of two language production experiments, we show that SSMs reproduce regression predictors, and further extend these effects through a multivariate decomposition (cognitive parameters). We also examine a combined regression-SSM approach that is hierarchical Bayesian, which can jointly model more conditions than classic SSMs, and importantly, achieve by-item modelling with other conditions. In this analysis, we found that spoken words principally differed from one another by their activation rates and production times, but not their thresholds to be activated.

The Scope of Planning Serial Actions during Typing.

Human activities consisting of multiple component actions require the generation of ordered sequences. This study investigated the scope of response planning in highly serial task, typing, by means of ERPs indexing motor response preparation. Specifically, we compared motor-related ERPs yielded by words typed using a single hand against words that had all keystrokes typed with a single hand, except for a deviant one, typed with the opposite hand. The deviant keystroke occurred either early in the typed sequence, corresponding to the second or third letters, or late, corresponding to the penultimate or last letter. Motor-related ERPs detected before response onset were affected only by deviant keystrokes located at the beginning of the sequence, whereas deviant keystrokes located at the end yielded ERPs that were undistinguishable from unimanual responses. These results impose some constraints on the notion of parallel processing of component actions.

Estimating Parallel Processing in a Language Task Using Single-Trial Intracerebral Electroencephalography.

We provide a quantitative assessment of the parallel-processing hypothesis included in various language-processing models. First, we highlight the importance of reasoning about cognitive processing at the level of single trials rather than using averages. Then, we report the results of an experiment in which the hypothesis was tested at an unprecedented level of granularity with intracerebral data recorded during a picture-naming task. We extracted patterns of significant high-gamma activity from multiple patients and combined them into a single analysis framework that identified consistent patterns. Average signals from different brain regions, presumably indexing distinct cognitive processes, revealed a large degree of concurrent activity. In comparison, at the level of single trials, the temporal overlap of detected significant activity was unexpectedly low, with the exception of activity in sensory cortices. Our novel methodology reveals some limits on the degree to which word production involves parallel processing.

Lesions to the left lateral prefrontal cortex impair decision threshold adjustment for lexical selection.

Patients with lesions in the left prefrontal cortex (PFC) have been shown to be impaired in lexical selection, especially when interference between semantically related alternatives is increased. To more deeply investigate which computational mechanisms may be impaired following left PFC damage due to stroke, a psychometric modelling approach is employed in which we assess the cognitive parameters of the patients from an evidence accumulation (sequential information sampling) modelling of their response data. We also compare the results to healthy speakers. Analysis of the cognitive parameters indicates an impairment of the PFC patients to appropriately adjust their decision threshold, in order to handle the increased item difficulty that is introduced by semantic interference. Also, the modelling contributes to other topics in psycholinguistic theory, in which specific effects are observed on the cognitive parameters according to item familiarization, and the opposing effects of priming (lower threshold) and semantic interference (lower drift) which are found to depend on repetition. These results are developed for the blocked-cyclic picture naming paradigm, in which pictures are presented within semantically homogeneous (HOM) or heterogeneous (HET) blocks, and are repeated several times per block. Overall, the results are in agreement with a role of the left PFC in adjusting the decision threshold for lexical selection in language production.

The time course of visual influences in letter recognition.

This study builds on a specific characteristic of letters of the Roman alphabet-namely, that each letter name is associated with two visual formats, corresponding to their uppercase and lowercase versions. Participants had to read aloud the names of single letters, and event-related potentials (ERPs) for six pairs of visually dissimilar upper- and lowercase letters were recorded. Assuming that the end product of processing is the same for upper- and lowercase letters sharing the same vocal response, ERPs were compared backward, starting from the onset of articulatory responses, and the first significant divergence was observed 120 ms before response onset. Given that naming responses were produced at around 414 ms, on average, these results suggest that letter processing is influenced by visual information until 294 ms after stimulus onset. This therefore provides new empirical evidence regarding the time course and interactive nature of visual letter perception processes.

On the resolution of phonological constraints in spoken production: Acoustic and response time evidence.

This study examines the production of words the pronunciation of which depends on the phonological context. Participants produced adjective-noun phrases starting with the French determiner un. The pronunciation of this determiner requires a liaison consonant before vowels. Naming latencies and determiner acoustic durations were shorter when the adjective and the noun both started with vowels or both with consonants, than when they had different onsets. These results suggest that the liaison process is not governed by the application of a local contextual phonological rule; they rather favor the hypothesis that pronunciation variants with and without the liaison consonant are stored in memory.

Simultaneous recording of MEG, EEG and intracerebral EEG during visual stimulation: from feasibility to single-trial analysis.

Electroencephalography (EEG), magnetoencephalography (MEG), and intracerebral stereotaxic EEG (SEEG) are the three neurophysiological recording techniques, which are thought to capture the same type of brain activity. Still, the relationships between non-invasive (EEG, MEG) and invasive (SEEG) signals remain to be further investigated. In early attempts at comparing SEEG with either EEG or MEG, the recordings were performed separately for each modality. However such an approach presents substantial limitations in terms of signal analysis. The goal of this technical note is to investigate the feasibility of simultaneously recording these three signal modalities (EEG, MEG and SEEG), and to provide strategies for analyzing this new kind of data. Intracerebral electrodes were implanted in a patient with intractable epilepsy for presurgical evaluation purposes. This patient was presented with a visual stimulation paradigm while the three types of signals were simultaneously recorded. The analysis started with a characterization of the MEG artifact caused by the SEEG equipment. Next, the average evoked activities were computed at the sensor level, and cortical source activations were estimated for both the EEG and MEG recordings; these were shown to be compatible with the spatiotemporal dynamics of the SEEG signals. In the average time-frequency domain, concordant patterns between the MEG/EEG and SEEG recordings were found below the 40 Hz level. Finally, a fine-grained coupling between the amplitudes of the three recording modalities was detected in the time domain, at the level of single evoked responses. Importantly, these correlations have shown a high level of spatial and temporal specificity. These findings provide a case for the ability of trimodal recordings (EEG, MEG, and SEEG) to reach a greater level of specificity in the investigation of brain signals and functions.

Reconciling phonological neighborhood effects in speech production through single trial analysis.

A crucial step for understanding how lexical knowledge is represented is to describe the relative similarity of lexical items, and how it influences language processing. Previous studies of the effects of form similarity on word production have reported conflicting results, notably within and across languages. The aim of the present study was to clarify this empirical issue to provide specific constraints for theoretical models of language production. We investigated the role of phonological neighborhood density in a large-scale picture naming experiment using fine-grained statistical models. The results showed that increasing phonological neighborhood density has a detrimental effect on naming latencies, and re-analyses of independently obtained data sets provide supplementary evidence for this effect. Finally, we reviewed a large body of evidence concerning phonological neighborhood density effects in word production, and discussed the occurrence of facilitatory and inhibitory effects in accuracy measures. The overall pattern shows that phonological neighborhood generates two opposite forces, one facilitatory and one inhibitory. In cases where speech production is disrupted (e.g. certain aphasic symptoms), the facilitatory component may emerge, but inhibitory processes dominate in efficient naming by healthy speakers. These findings are difficult to accommodate in terms of monitoring processes, but can be explained within interactive activation accounts combining phonological facilitation and lexical competition.

An exploration of anomia rehabilitation in drug-resistant temporal lobe epilepsy.

Around 40% of patients who undergo a left temporal lobe epilepsy (LTLE) surgery suffer from anomia (word-finding difficulties), a condition that negatively impacts quality of life. Despite these observations, language rehabilitation is still understudied in LTLE. We assessed the effect of a four-week rehabilitation on four drug-resistant LTLE patients after their surgery. The anomia rehabilitation was based on cognitive descriptions of word finding deficits in LTLE. Its primary ingredients were psycholinguistic tasks and a psychoeducation approach to help patients cope with daily communication issues. We repeatedly assessed naming skills for trained and untrained words, before and during the therapy using an A-B design with follow-up and replication. Subjective anomia complaint and standardized language assessments were also collected. We demonstrated the effectiveness of the rehabilitation program for trained words despite the persistence of seizures. Furthermore, encouraging results were observed for untrained items. Variable changes in anomia complaint were observed. One patient who conducted the protocol as self-rehabilitation responded similarly to the others, despite the different manner of intervention. These results open promising avenues for helping epileptic patients suffering from anomia. For example, this post-operative program could easily be adapted to be conducted preoperatively.