On the time-course and frequency selectivity of the EEG for different modes of response selection : evidence from speech production and keyboard pressing

Objective To compare brain activity in the alpha and beta bands in relation to different modes of response selection, and to assess the domain generality of the response selection mechanism using verbal and non-verbal tasks. Methods We examined alpha and beta event-related desynchronization (ERD) to analyze brain reactivity during the selection of verbal (word production) and non-verbal motor actions (keyboard pressing) under two different response modes: externally selected and self-selected. Results An alpha and beta ERD was observed for both the verbal and non-verbal tasks in both the externally and the self-selected modes. For both tasks, the beta ERD started earlier and was longer in the self-selected mode than in the externally selected mode. The overall pattern of results between the verbal and non-verbal motor behaviors was similar. Conclusions The pattern of alpha and beta ERD is affected by the mode of response selection suggesting that the activity in both frequency bands contributes to the process of selecting actions. We suggest that activity in the alpha band may reflect attentional processes while activity in the beta band may be more closely related to the execution and selection process. Significance These results suggest that a domain general process contributes to the planning of speech and other motor actions. This finding has potential clinical implications, for the use of diverse motor tasks to treat disorders of motor planning

From language comprehension to action understanding and back again

A controversial question in cognitive neuroscience is whether comprehension of words and sentences engages brain mechanisms specific for decoding linguistic meaning or whether language comprehension occurs through more domain-general sensorimotor processes. Accumulating behavioral and neuroimaging evidence suggests a role for cortical motor and premotor areas in passive action-related language tasks, regions that are known to be involved in action execution and observation. To examine the involvement of these brain regions in language and nonlanguage tasks, we used functional magnetic resonance imaging (fMRI) on a group of 21 healthy adults. During the fMRI session, all participants 1) watched short object-related action movies, 2) looked at pictures of man-made objects, and 3) listened to and produced short sentences describing object-related actions and man-made objects. Our results are among the first to reveal, in the human brain, a functional specialization within the ventral premotor cortex (PMv) for observing actions and for observing objects, and a different organization for processing sentences describing actions and objects. These findings argue against the strongest version of the simulation theory for the processing of action-related language

Music-based interventions for aphasia could act through a motor-speech mechanism : a systematic review and case-control analysis of published individual participant data

Background: Melodic Intonation Therapy, a music-based intervention for the recovery of oral language production in aphasia, has been shown to be particularly effective in patients with Broca’s aphasia compared to other aphasia subtypes. It has been suggested that this therapy might improve language output by acting on motor-speech deficits often associated with Broca’s aphasia. In this article, we examine the relevance of a motor-speech mechanism for music-based interventions designed to improve verbal expression in patients with any type of aphasia. Aim: To test the association between the presence of motor-speech disorders (MSDs) and improvement with music-based protocols targeting verbal expression in participants with aphasia. Methods and procedures: We conducted a systematic review of publications reporting language production outcomes following a music-based intervention in participants with aphasia and performed a case–control analysis on extracted individual participant data (IPD). The databases PubMed, MEDLINE (1800 to 9 March 2018), and PsycINFO (1806 to March 2018) were screened, followed with cross-referencing. We recorded data at the level of study and, when possible, at the IPD level. When not explicitly reported, we applied a series of heuristics to infer the presence/absence of an MSD in participants. Binomial logistic regressions were performed to ascertain the effects of the presence of an MSD, aphasia severity, treatment duration (in weeks), and treatment intensity (hours/week) on the likelihood that participants would show a speech or a language improvement following intervention. Outcomes & Results: Forty original articles were included in this review. Twenty-two reported sufficient details to be included in our IPD analysis, for a total sample of 105 participants. Most interventions included some sort of singing as their primary music-based facilitation technique for language production. For speech improvement, statistically significant predictor variables were the presence of an MSD and treatment intensity. For language improvement, statistically significant predictor variables were the presence of an MSD, treatment intensity, and duration. Severity of aphasia was not associated with the likelihood of speech or language improvement. Conclusion: Music-based interventions for language production in aphasia may act via a motor-speech mechanism. We suggest that music and singing-based therapies might be further investigated as treatment options for patients with MSDs, whether associated with aphasia or not

On the selection of words and oral motor responses : evidence of a response-independent fronto-parietal network

Several brain areas including the medial and lateral premotor areas, and the prefrontal cortex, are thought to be involved in response selection. It is unclear, however, what the specific contribution of each of these areas is. It is also unclear whether the response selection process operates independent of response modality or whether a number of specialized processes are recruited depending on the behaviour of interest. In the present study, the neural substrates for different response selection modes (volitional and stim- ulus-driven) were compared, using sparse-sampling functional magnetic resonance imaging, for two different response modalities: words and comparable oral motor gestures. Results demonstrate that response selection relies on a network of prefrontal, premotor and parietal areas, with the pre-supplementary motor area (pre-SMA) at the core of the process. Overall, this network is sensitive to the manner in which responses are selected, despite the absence of a medio-lateral axis, as was suggested by Goldberg (1985). In contrast, this network shows little sensitivity to the modality of the response, suggesting of a domain-general selection process. Theoretical implications of these results are discussed

Motor Response Selection in Overt Sentence Production: A Functional MRI Study

Many different cortical areas are thought to be involved in the process of selecting motor responses, from the inferior frontal gyrus, to the lateral and medial parts of the premotor cortex. The objective of the present study was to examine the neural underpinnings of motor response selection in a set of overt language production tasks. To this aim, we compared a sentence repetition task (externally constrained selection task) with a sentence generation task (volitional selection task) in a group of healthy adults. In general, the results clarify the contribution of the pre-SMA, cingulate areas, PMv, and pars triangularis to the process of selecting motor responses in the context of sentence production, and shed light on the manner in which this network is modulated by selection mode. Further, the present study suggests that response selection in sentence production engages neural resources similar to those engaged in the production of isolated words and oral motor gestures

Contribution of the frontal lobe to externally and internally specified verbal responses : fMRI evidence

It has been suggested that within the frontal cortex there is a lateral to medial shift in the control of action, with the lateral premotor area (PMA) involved in externally specified actions and the medial supplementary motor areas (SMA) involved in internally specified actions. Recent brain imaging studies demonstrate, however, that the control of externally and internally specified actions may involve more complex and overlapping networks involving not only the PMA and the SMA, but also the pre-SMA and the lateral prefrontal cortex (PFC). The aim of the present study was to determine whether these frontal regions are differentially involved in the production of verbal responses, when they are externally specified and when they are internally specified. Participants engaged in three overt speaking tasks in which the degree of response specification differed. The tasks involved reading aloud words (externally specified), or generating words aloud from narrow or broad semantic categories (internally specified). Using fMRI, the location and magnitude of the BOLD activity for these tasks was measured in a group of ten participants. Compared with rest, all tasks activated the primary motor area and the SMA-proper, reflecting their common role in speech production. The magnitude of the activity in the PFC (Brodmann area 45), the left PMAv and the pre-SMA increased for word generation, suggesting that each of these three regions plays a role in internally specified action selection. This confirms previous reports concerning the participation of the pre-SMA in verbal response selection. The pattern of activity in PMAv suggests participation in both externally and internally specified verbal actions

Beyond the arcuate fasciculus : consensus and controversy in the connectional anatomy of language

The growing consensus that language is distributed into large-scale cortical and subcortical networks has brought with it an increasing focus on the connectional anatomy of language, or how particular fibre pathways connect regions within the language network. Understanding connectivity of the language network could provide critical insights into function, but recent investigations using a variety of methodologies in both humans and non-human primates have provided conflicting accounts of pathways central to language. Some of the pathways classically considered language pathways, such as the arcuate fasciculus, are now argued to be domain-general rather than specialized, which represents a radical shift in perspective. Other pathways described in the non-human primate remain to be verified in humans. In this review, we examine the consensus and controversy in the study of fibre pathway connectivity for language. We focus on seven fibre pathways—the superior longitudinal fasciculus and arcuate fasciculus, the uncinate fasciculus, extreme capsule, middle longitudinal fasciculus, inferior longitudinal fasciculus and inferior fronto-occipital fasciculus—that have been proposed to support language in the human. We examine the methods in humans and non-human primate used to investigate the connectivity of these pathways, the historical context leading to the most current understanding of their anatomy, and the functional and clinical correlates of each pathway with reference to language. We conclude with a challenge for researchers and clinicians to establish a coherent framework within which fibre pathway connectivity can be systematically incorporated to the study of language

Contribution of the pre-SMA to the production of words and non-speech oral motor gestures, as revealed by repetitive transcranial magnetic stimulation (rTMS)

An emerging theoretical perspective, largely based on neuroimaging studies, suggests that the pre-SMA is involved in planning cognitive aspects of motor behavior and language, such as linguistic and non-linguistic response selection. Neuroimaging studies, however, cannot indicate whether a brain region is equally important to all tasks in which it is activated. In the present study, we tested the hypothesis that the pre-SMA is an important component of response selection, using an interference technique. High frequency repetitive TMS (10 Hz) was used to interfere with the functioning of the pre-SMA during tasks requiring selection of words and oral gestures under different selection modes (forced, volitional) and attention levels (high attention, low attention). Results show that TMS applied to the pre-SMA interferes selectively with the volitional selection condition, resulting in longer RTs. The low- and high-attention forced selection conditions were unaffected by TMS, demonstrating that the pre-SMA is sensitive to selection mode but not attentional demands. TMS similarly affected the volitional selection of words and oral gestures, reflecting the response- independent nature of the pre-SMA contribution to response selection. The implications of these results are discussed

Broca and wernicke are dead, or moving past the classic model of language neurobiology

With the advancement of cognitive neuroscience and neuropsychological research, the field of language neurobiology is at a cross-roads with respect to its framing theories. The central thesis of this article is that the major historical framing model, the Classic “Wernicke-Lichtheim-Geschwind” model, and associated terminology, is no longer adequate for contemporary investigations into the neurobiology of language. We argue that the Classic model (1) is based on an outdated brain anatomy; (2) does not adequately represent the distributed connectivity relevant for language, (3) offers a modular and “language centric” perspective, and (4) focuses on cortical structures, for the most part leaving out subcortical regions and relevant connections. To make our case, we discuss the issue of anatomical specificity with a focus on the contemporary usage of the terms “Broca’s and Wernicke’s area”, including results of a survey that was conducted within the language neurobiology community. We demonstrate that there is no consistent anatomical definition of “Broca’s and Wernicke’s Areas”, and propose to replace these terms with more precise anatomical definitions. We illustrate the distributed nature of the language connectome, which extends far beyond the single-pathway notion of arcuate fasciculus connectivity established in the Geschwind’s version of the Classic Model. By illustrating the definitional confusion surrounding “Broca’s and Wernicke’s areas”, and the by illustrating the difficulty integrating the emerging literature on perisylvian white matter connectivity into this model, we hope to expose the limits of the model, argue for its obsolescence, and suggest a path forward in defining a replacement

On the context-dependent nature of the contribution of the ventral premotor cortex to speech perception

What is the nature of the interface between speech perception and production, where auditory and motor representations converge? One set of explanations suggests that during perception, the motor circuits involved in producing a perceived action are in some way enacting the action without actually causing movement (covert simulation) or sending along the motor information to be used to predict its sensory consequences (i.e., efference copy). Other accounts either reject entirely the involvement of motor representations in perception, or explain their role as being more supportive than integral, and not employing the identical circuits used in production. Using fMRI, we investigated whether there are brain regions that are conjointly active for both speech perception and production, and whether these regions are sensitive to articulatory (syllabic) complexity during both processes, which is predicted by a covert simulation account. A group of healthy young adults (1) observed a female speaker produce a set of familiar words (perception), and (2) observed and then repeated the words (production). There were two types of words, varying in articulatory complexity, as measured by the presence or absence of consonant clusters. The simple words contained no consonant cluster (e.g. “palace”), while the complex words contained one to three consonant clusters (e.g. “planet”). Results indicate that the left ventral premotor cortex (PMv) was significantly active during speech perception and speech production but that activation in this region was scaled to articulatory complexity only during speech production, revealing an incompletely specified efferent motor signal during speech perception. The right planum temporal (PT) was also active during speech perception and speech production, and activation in this region was scaled to articulatory complexity during both production and perception. These findings are discussed in the context of current theories of speech perception, with particular attention to accounts that include an explanatory role for mirror neurons