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.

Distinct neural mechanisms support inner speaking and inner hearing.

Humans have the ability to mentally examine speech. This covert form of speech production is often accompanied by sensory (e.g., auditory) percepts. However, the cognitive and neural mechanisms that generate these percepts are still debated. According to a prominent proposal, inner speech has at least two distinct phenomenological components: inner speaking and inner hearing. We used transcranial magnetic stimulation to test whether these two phenomenologically distinct processes are supported by distinct neural mechanisms. We hypothesised that inner speaking relies more strongly on an online motor-to-sensory simulation that constructs a multisensory experience, whereas inner hearing relies more strongly on a memory-retrieval process, where the multisensory experience is reconstructed from stored motor-to-sensory associations. Accordingly, we predicted that the speech motor system will be involved more strongly during inner speaking than inner hearing. This would be revealed by modulations of TMS evoked responses at muscle level following stimulation of the lip primary motor cortex. Overall, data collected from 31 participants corroborated this prediction, showing that inner speaking increases the excitability of the primary motor cortex more than inner hearing. Moreover, this effect was more pronounced during the inner production of a syllable that strongly recruits the lips (vs. a syllable that recruits the lips to a lesser extent). These results are compatible with models assuming that the primary motor cortex is involved during inner speech and contribute to clarify the neural implementation of the fundamental ability of silently speaking in one's mind.

Writing letters in two graphic systems: Behavioral and neural correlates in Latin-Arabic biscripters.

Biscriptuality is the ability to read and write using two scripts. Despite the increasing number of biscripters, this phenomenon remains poorly understood. Here, we focused on investigating graphomotor processing in French-Arabic biscripters. We chose the French and Arabic alphabets because they have comparable visuospatial complexity and linguistic features, but differ dramatically in their graphomotor characteristics. In a first experiment we describe the graphomotor features of the two alphabets and showed that while Arabic and Latin letters are produced with the same velocity and fluency, Arabic letters require more pen lifts, contain more right-to-left strokes and clockwise curves, and take longer to write than Latin letters. These results suggest that Arabic and Latin letters are produced via different motor patterns. In a second experiment we used functional magnetic resonance imaging to ask whether writing the two scripts relies upon partially distinct or fully overlapping neural networks, and whether the elements of the previously described handwriting network are recruited to the same extent by the two scripts. We found that both scripts engaged the so-called "writing network", but that within the network, Arabic letters recruited the left superior parietal lobule (SPL) and the left primary motor cortex (M1) more strongly than Latin letters. Both regions have previously been identified as holding scale-invariant representations of letter trajectories. Arabic and Latin letters also activated distinct regions that do not belong to the writing network. Complementary analyses indicate that the differences observed between scripts at the neural level could be driven by the specific graphomotor features of each script. Overall, our results indicate that particular features of the practiced scripts can lead to different motor organization at both the behavioral and brain levels in biscripters.

Effects of biscriptuality on graphomotor coordination dynamics.

Biscriptuality is the ability to write in two different writing systems. The aim of this study was to examine the effects of biscriptuality on graphomotor coordination dynamics in right-handed adults. Thirty-four French monoscriptuals and 34 French-Arabic biscriptual participants traced series of loops in two writing directions and in two directions of rotation. We found that biscriptuals displayed a general advantage over monoscriptuals in terms of tracing frequency, while both groups displayed a preference for the left-to-right direction. These results provide novel evidence on the effects of writing direction and type of expertise on graphomotor performance by showing that biscriptuality could be an asset. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Typing expertise in a large student population.

Typing has become a pervasive mode of language production worldwide, with keyboards fully integrated in a large part of many daily activities. The bulk of the literature on typing expertise concerns highly trained professional touch-typists, but contemporary typing skills mostly result from unconstrained sustained practice. We measured the typing performance of a large cohort of 1301 university students through an online platform and followed a preregistered plan to analyse performance distributions, practice factors, and cognitive variables. The results suggest that the standard model with a sharp distinction between novice and expert typists may be inaccurate to account for the performance of the current generation of young typists. More generally, this study shows how the mere frequent use of a new tool can lead to the incidental development of high expertise.

The Role of Motor Inhibition During Covert Speech Production.

Covert speech is accompanied by a subjective multisensory experience with auditory and kinaesthetic components. An influential hypothesis states that these sensory percepts result from a simulation of the corresponding motor action that relies on the same internal models recruited for the control of overt speech. This simulationist view raises the question of how it is possible to imagine speech without executing it. In this perspective, we discuss the possible role(s) played by motor inhibition during covert speech production. We suggest that considering covert speech as an inhibited form of overt speech maps naturally to the purported progressive internalization of overt speech during childhood. We further argue that the role of motor inhibition may differ widely across different forms of covert speech (e.g., condensed vs. expanded covert speech) and that considering this variety helps reconciling seemingly contradictory findings from the neuroimaging literature.

Temporally resolved neural dynamics underlying handwriting.

How do the temporal dynamics of neural activity encode highly coordinated visual-motor behaviour? To capture the millisecond-resolved neural activations associated with fine visual-motor skills, we devised a co-registration system to simultaneously record electroencephalogram and handwriting kinematics while participants were performing four handwriting tasks (writing in Chinese/English scripts with their dominant/non-dominant hand). The neural activation associated with each stroke was clearly identified with a well-structured and reliable pattern. The functional significance of this pattern was validated by its significant associations with language, hand and the cognitive stages and kinematics of handwriting. Furthermore, the handwriting rhythmicity was found to be synchronised to the brain's ongoing theta oscillation, and the synchronisation was associated with the factor of language and hand. These major findings imply an implication between motor skill formation and the interplay between the rhythms in the brain and the peripheral systems.

The Human Basal Ganglia Mediate the Interplay between Reactive and Proactive Control of Response through Both Motor Inhibition and Sensory Modulation.

The basal ganglia (BG) have long been known for contributing to the regulation of motor behaviour by means of a complex interplay between tonic and phasic inhibitory mechanisms. However, after having focused for a long time on phasic reactive mechanisms, it is only recently that psychological research in healthy humans has modelled tonic proactive mechanisms of control. Mutual calibration between anatomo-functional and psychological models is still needed to better understand the unclear role of the BG in the interplay between proactive and reactive mechanisms of control. Here, we implemented an event-related fMRI design allowing proper analysis of both the brain activity preceding the target-stimulus and the brain activity induced by the target-stimulus during a simple go/nogo task, with a particular interest in the ambiguous role of the basal ganglia. Post-stimulus activity was evoked in the left dorsal striatum, the subthalamus nucleus and internal globus pallidus by any stimulus when the situation was unpredictable, pinpointing its involvement in reactive, non-selective inhibitory mechanisms when action restraint is required. Pre-stimulus activity was detected in the ventral, not the dorsal, striatum, when the situation was unpredictable, and was associated with changes in functional connectivity with the early visual, not the motor, cortex. This suggests that the ventral striatum supports modulatory influence over sensory processing during proactive control.

The handwriting brain in middle childhood.

While the brain network supporting handwriting has previously been defined in adults, its organization in children has never been investigated. We compared the handwriting network of 23 adults and 42 children (8- to 11-year-old). Participants were instructed to write the alphabet, the days of the week, and to draw loops while being scanned. The handwriting network previously described in adults (five key regions: left dorsal premotor cortex, superior parietal lobule (SPL), fusiform and inferior frontal gyri, and right cerebellum) was also strongly activated in children. The right precentral gyrus and the right anterior cerebellum were more strongly activated in adults than in children, while the left fusiform gyrus (FuG) was more strongly activated in children than in adults. Finally, we found that, contrary to adults, children recruited prefrontal regions to complete the writing task. This constitutes the first comparative investigation of the neural correlates of writing in children and adults. Our results suggest that the network supporting handwriting is already established in middle childhood. They also highlight the major role of prefrontal regions in learning this complex skill and the importance of right precentral regions and cerebellum in the performance of automated handwriting.

Shared premotor activity in spoken and written communication.

The aim of the present study was to uncover a possible common neural organizing principle in spoken and written communication, through the coupling of perceptual and motor representations. In order to identify possible shared neural substrates for processing the basic units of spoken and written language, a sparse sampling fMRI acquisition protocol was performed on the same subjects in two experimental sessions with similar sets of letters being read and written and of phonemes being heard and orally produced. We found evidence of common premotor regions activated in spoken and written language, both in perception and in production. The location of those brain regions was confined to the left lateral and medial frontal cortices, at locations corresponding to the premotor cortex, inferior frontal cortex and supplementary motor area. Interestingly, the speaking and writing tasks also appeared to be controlled by largely overlapping networks, possibly indicating some domain general cognitive processing. Finally, the spatial distribution of individual activation peaks further showed more dorsal and more left-lateralized premotor activations in written than in spoken language.

The impact of spelling regularity on handwriting production: A coupled fMRI and kinematics study.

Current models of writing assume that the orthographic processes involved in spelling retrieval and the motor processes involved in the control of the hand are independent. This view has been challenged by behavioral studies, which showed that the linguistic features of words impact motor execution during handwriting. We designed an experiment coupling functional magnetic resonance imaging and kinematic recordings during a writing to dictation task. Participants wrote orthographically regular and irregular words. The presence of an irregularity impacts both the initiation of the movement and its fine motor execution. At the brain level, the left inferior frontal and fusiform gyri, two regions belonging to the core of the written language system, were found to be sensitive to the presence of an irregularity and to its position in the word during writing execution. Moreover, the left superior parietal lobule, the left superior frontal gyrus and the right cerebellum, three motor-related regions, displayed a stronger response to irregular than regular words. These results constitute direct evidence that orthographic and motor processes occur in a continuous and interactive fashion during writing.

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.

The serial order of response units in word production: The case of typing.

The selection and ordering of response units (phonemes, letters, keystrokes) represents a transversal issue across different modalities of language production. Here, the issue of serial order was investigated with respect to typewriting. Following seminal investigations in the spoken modality, we conducted an experiment where participants typed as many times as possible a pair of words during a fixed time-window. The 2 words shared either their first 2 keystrokes, the last 2 ones, all the keystrokes, or were unrelated. Fine-grained performance measures were recorded at the level of individual keystrokes. In contrast with previous results from the spoken modality, we observed an overall facilitation for words sharing the initial keystrokes. In addition, the initial overlap briefly delayed the execution of the following keystroke. The results are discussed with reference to different theoretical perspectives on serial order, with a particular attention to the competing accounts offered by position coding models and chaining models. Our findings point to potential major differences between the speaking and typing modalities in terms of interactive activation between lexical and response units processing levels. (PsycINFO Database Record

Motor control of handwriting in the developing brain: A review.

This review focuses on the acquisition of writing motor aspects in adults, and in 5-to 12-year-old children without learning disabilities. We first describe the behavioural aspects of adult writing and dominant models based on the notion of motor programs. We show that handwriting acquisition is characterized by the transition from reactive movements programmed stroke-by-stroke in younger children, to an automatic control of the whole trajectory when the motor programs are memorized at about 10 years old. Then, we describe the neural correlates of adult writing, and the changes that could occur with learning during childhood. The acquisition of a new skill is characterized by the involvement of a network more restricted in space and where neural specificity is increased in key regions. The cerebellum and the left dorsal premotor cortex are of fundamental importance in motor learning, and could be at the core of the acquisition of handwriting.

Effect of training status on beta-range corticomuscular coherence in agonist vs. antagonist muscles during isometric knee contractions.

Antagonist muscle co-activation is thought to be partially regulated by cortical influences, but direct motor cortex involvement is not fully understood. Corticomuscular coherence (CMC) measures direct functional coupling of the motor cortex and muscles. As antagonist co-activation differs according to training status, comparison of CMC in agonist and antagonist muscles and in strength-trained and endurance-trained individuals may provide in-depth knowledge of cortical implication in antagonist muscle co-activation. Electroencephalographic and electromyographic signals were recorded, while 10 strength-trained and 11 endurance-trained participants performed isometric knee contractions in flexion and extension at various torque levels. CMC magnitude in 13-21 and 21-31 Hz frequency bands was quantified by CMC analysis between Cz electroencephalographic electrode activity and all electromyographic signals. CMC was significant in both 13-21 and 21-31 Hz frequency bands in flexor and extensor muscles regardless of participant group, torque level, and direction of contraction. CMC magnitude decreased more in antagonist than in agonist muscles as torque level increased. Finally, CMC magnitude was higher in strength-trained than in endurance-trained participants. These findings provide experimental evidence that the motor cortex directly regulates both agonist and antagonist muscles. Nevertheless, the mechanisms underlying muscle activation may be specific to their function. Between-group modulation of corticomuscular coherence may result from training-induced adaptation, re-emphasizing that corticomuscular coherence analysis may be efficient in characterizing corticospinal adaptations after long-term muscle specialization.

Testing the physiological plausibility of conflicting psychological models of response inhibition: A forward inference fMRI study.

The neural mechanisms underlying response inhibition and related disorders are unclear and controversial for several reasons. First, it is a major challenge to assess the psychological bases of behaviour, and ultimately brain-behaviour relationships, of a function which is precisely intended to suppress overt measurable behaviours. Second, response inhibition is difficult to disentangle from other parallel processes involved in more general aspects of cognitive control. Consequently, different psychological and anatomo-functional models coexist, which often appear in conflict with each other even though they are not necessarily mutually exclusive. The standard model of response inhibition in go/no-go tasks assumes that inhibitory processes are reactively and selectively triggered by the stimulus that participants must refrain from reacting to. Recent alternative models suggest that action restraint could instead rely on reactive but non-selective mechanisms (all automatic responses are automatically inhibited in uncertain contexts) or on proactive and non-selective mechanisms (a gating function by which reaction to any stimulus is prevented in anticipation of stimulation when the situation is unpredictable). Here, we assessed the physiological plausibility of these different models by testing their respective predictions regarding event-related BOLD modulations (forward inference using fMRI). We set up a single fMRI design which allowed for us to record simultaneously the different possible forms of inhibition while limiting confounds between response inhibition and parallel cognitive processes. We found BOLD dynamics consistent with non-selective models. These results provide new theoretical and methodological lines of inquiry for the study of basic functions involved in behavioural control and related disorders.

How specialized are writing-specific brain regions? An fMRI study of writing, drawing and oral spelling.

Several brain imaging studies identified brain regions that are consistently involved in writing tasks; the left premotor and superior parietal cortices have been associated with the peripheral components of writing performance as opposed to other regions that support the central, orthographic components. Based on a meta-analysis by Planton, Jucla, Roux, and Demonet (2013), we focused on five such writing areas and questioned the task-specificity and hemispheric lateralization profile of the brain response in an functional magnetic resonance imaging (fMRI) experiment where 16 right-handed participants wrote down, spelled out orally object names, and drew shapes from object pictures. All writing-related areas were activated by drawing, and some of them by oral spelling, thus questioning their specialization for written production. The graphemic/motor frontal area (GMFA), a subpart of the superior premotor cortex close to Exner's area (Roux et al., 2009), was the only area with a writing-specific lateralization profile, that is, clear left lateralization during handwriting, and bilateral activity during drawing. Furthermore, the relative lateralization and levels of activation in the superior parietal cortex, ventral premotor cortex, ventral occipitotemporal cortex and right cerebellum across the three tasks brought out new evidence regarding their respective contributions to the writing processes.

Activation of writing-specific brain regions when reading Chinese as a second language. Effects of training modality and transfer to novel characters.

We examined the implication of training modality on the cortical representation of Chinese words in adult second language learners of Chinese. In particular, we tested the implication of the neural substrates of writing in a reading task. The brain network sustaining finger writing was defined neuroanatomically based on an independent functional localizer. We examined the brain activations elicited by Chinese words learned via writing vs. pronunciation, and by novel untrained words, within regions of interest (ROIs) defined according to the position of the activation peaks in the localizer, and at the whole brain level. We revealed activations in the reading task that overlapped with several parts of the finger writing network. In addition, our results provide evidence that the neural substrates of writing are differentially involved in reading depending on the stored knowledge for words, as revealed by the fine-grained response of several regions including the left superior parietal lobule and left precentral gyrus / superior frontal sulcus to the experimental manipulations. Training modality and the linguistic properties of the characters also impacted the response of the left mid-fusiform gyrus, confirming its involvement as the brain region where linguistic, visual and sensorimotor information converge during orthographic processing. At the behavioral level, global handwriting quality during the training sessions was positively correlated to the final translation performance. Our results demonstrate substantial overlap in the neural substrates of reading and writing, and indicate that some regions sustaining handwriting are differentially involved in reading depending on the type of knowledge associated with words.