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.

Phoneme Representation and Articulatory Impairment: Insights from Adults with Comorbid Motor Coordination Disorder and Dyslexia.

Phonemic processing skills are impaired both in children and adults with dyslexia. Since phoneme representation development is based on articulatory gestures, it is likely that these gestures influence oral reading-related skills as assessed through phonemic awareness tasks. In our study, fifty-two young dyslexic adults, with and without motor impairment, and fifty-nine skilled readers performed reading, phonemic awareness, and articulatory tasks. The two dyslexic groups exhibited slower articulatory rates than skilled readers and the comorbid dyslexic group presenting with an additional difficulty in respiratory control (reduced speech proportion and increased pause duration). Two versions of the phoneme awareness task (PAT) with pseudoword strings were administered: a classical version under time pressure and a delayed version in which access to phonemic representations and articulatory programs was facilitated. The two groups with dyslexia were outperformed by the control group in both versions. Although the two groups with dyslexia performed equally well on the classical PAT, the comorbid group performed significantly less efficiently on the delayed PAT, suggesting an additional contribution of articulatory impairment in the task for this group. Overall, our results suggest that impaired phoneme representations in dyslexia may be explained, at least partially, by articulatory deficits affecting access to them.

Stronger impulse capture and impaired inhibition of prepotent action in children with ADHD performing a Simon task: An electromyographic study.

OBJECTIVE: A deficit in interference control is commonly reported in children with attention deficit hyperactivity disorder (ADHD). This has mainly been interpreted as a difficulty in inhibiting inappropriate responses. However, it could be due to at least two distinct and independent processes, which are often confounded: The activation or suppression of impulsive responses. The aim of the present study was to separate the contribution of these two processes. METHOD: We compared performance of 26 children with ADHD to that of 26 nonADHD children using a novel approach based on electromyographic activity (EMG) analysis. EMG allows two distinct indices to be computed: Incorrect activation rate, which is an index of the intensity of impulse capture and correction rate, which provides a direct measure of the ability to suppress automatic responses. RESULTS: Children with ADHD were slower, committed more errors, and had a larger interference effect than nonADHD children. Moreover, we observed a greater incorrect activation rate and a lower correction rate in the ADHD group. CONCLUSIONS: Our data suggest that the difficulties in interference control found in children with ADHD are explained by both impaired inhibitory processes and a greater propensity to activate automatic responses. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Centroparietal activity mirrors the decision variable when tracking biased and time-varying sensory evidence.

Decision-making is a fundamental human activity requiring explanation at the neurocognitive level. Current theoretical frameworks assume that, during sensory-based decision-making, the stimulus is sampled sequentially. The resulting evidence is accumulated over time as a decision variable until a threshold is reached and a response is initiated. Several neural signals, including the centroparietal positivity (CPP) measured from the human electroencephalogram (EEG), appear to display the accumulation-to-bound profile associated with the decision variable. Here, we evaluate the putative computational role of the CPP as a model-derived accumulation-to-bound signal, focussing on point-by-point correspondence between model predictions and data in order to go beyond simple summary measures like average slope. In two experiments, we explored the CPP under two manipulations (namely non-stationary evidence and probabilistic decision biases) that complement one another by targeting the shape and amplitude of accumulation respectively. We fit sequential sampling models to the behavioural data, and used the resulting parameters to simulate the decision variable, before directly comparing the simulated profile to the CPP waveform. In both experiments, model predictions deviated from our naïve expectations, yet showed similarities with the neurodynamic data, illustrating the importance of a formal modelling approach. The CPP appears to arise from brain processes that implement a decision variable (as formalised in sequential-sampling models) and may therefore inform our understanding of decision-making at both the representational and implementational levels of analysis, but at this point it is uncertain whether a single model can explain how the CPP varies across different kinds of task manipulation.

The Neurodynamic Decision Variable in Human Multi-alternative Perceptual Choice.

The neural dynamics underpinning binary perceptual decisions and their transformation into actions are well studied, but real-world decisions typically offer more than two response alternatives. How does decision-related evidence accumulation dynamically influence multiple action representations in humans? The heightened conservatism required in multiple compared with binary choice scenarios suggests a mechanism that compensates for increased uncertainty when multiple choices are present by suppressing baseline activity. Here, we tracked action representations using corticospinal excitability during four- and two-choice perceptual decisions and modeled them using a sequential sampling framework. We found that the predictions made by leaky competing accumulator models to accommodate multiple choices (i.e., reduced baseline activity to compensate increased uncertainty) were borne out by dynamic changes in human action representations. This suggests a direct and continuous influence of interacting evidence accumulators, each favoring a different decision alternative, on downstream corticospinal excitability during complex choice.

Neurodynamic Evidence Supports a Forced-Excursion Model of Decision-Making under Speed/Accuracy Instructions.

Evolutionary pressures suggest that choices should be optimized to maximize rewards, by appropriately trading speed for accuracy. This speed-accuracy tradeoff (SAT) is commonly explained by variation in just the baseline-to-boundary distance, i.e., the excursion, of accumulation-to-bound models of perceptual decision-making. However, neural evidence is not consistent with this explanation. A compelling account of speeded choice should explain both overt behavior and the full range of associated brain signatures. Here, we reconcile seemingly contradictory behavioral and neural findings. In two variants of the same experiment, we triangulated upon the neural underpinnings of the SAT in the human brain using both EEG and transcranial magnetic stimulation (TMS). We found that distinct neural signals, namely the event-related potential (ERP) centroparietal positivity (CPP) and a smoothed motor-evoked potential (MEP) signal, which have both previously been shown to relate to decision-related accumulation, revealed qualitatively similar average neurodynamic profiles with only subtle differences between SAT conditions. These signals were then modelled from behavior by either incorporating traditional boundary variation or utilizing a forced excursion. These model variants are mathematically equivalent, in terms of their behavioral predictions, hence providing identical fits to correct and erroneous reaction time distributions. However, the forced-excursion version instantiates SAT via a more global change in parameters and implied neural activity, a process conceptually akin to, but mathematically distinct from, urgency. This variant better captured both ERP and MEP neural profiles, suggesting that the SAT may be implemented via neural gain modulation, and reconciling standard modelling approaches with human neural data.

Beyond decision! Motor contribution to speed-accuracy trade-off in decision-making.

Both in real life and experimental settings, increasing response speed typically leads to more error-prone actions. Processes underlying such a "speed-accuracy trade-off" (SAT) are usually assumed to be purely decisional: cautiousness would be determined only by the amount of sensory evidence required to select a response. The present data challenges this largely accepted view, by directly showing that motor processes are speeded up under time pressure. In a choice reaction time task where emphasis was put either on response speed or accuracy, motor processes were investigated through the analysis of muscular activity related to response execution. When response speed was emphasized, the time between electromyographic onset and behavioral response (motor time) was also speeded up (contributing to more than 20 % of the total effect on global reaction time). This speeded execution (likely due to a more efficient motor command) may also explain why participants are less able to interrupt incorrect response execution once started (Burle et al., Psychonomic Bulletin & Review, 21(4), 1003-1010, 2014), leading to more overt errors. Pointing to a speed-accuracy exchange within motor processes themselves, the present results call for a re-evaluation of widely accepted assumptions about SAT, and more generally, decision-making processes. They are discussed in the context of recent extensions of the drift diffusion model framework, questioning the strict separation between decisional and motor processes.

An independent, landmark-dominated head-direction signal in dysgranular retrosplenial cortex.

We investigated how landmarks influence the brain's computation of head direction and found that in a bidirectionally symmetrical environment, some neurons in dysgranular retrosplenial cortex showed bidirectional firing patterns. This indicates dominance of neural activity by local environmental cues even when these conflicted with the global head direction signal. It suggests a mechanism for associating landmarks to or dissociating them from the head direction signal, according to their directional stability and/or utility.

Preventing (impulsive) errors: Electrophysiological evidence for online inhibitory control over incorrect responses.

In a rich environment, with multiple action affordances, selective action inhibition is critical in preventing the execution of inappropriate responses. Here, we studied the origin and the dynamics of incorrect response inhibition and how it can be modulated by task demands. We used EEG in a conflict task where the probability of compatible and incompatible trials was varied. This allowed us to modulate the strength of the prepotent response, and hence to increase the risk of errors, while keeping the probability of the two responses equal. The correct response activation and execution was not affected by compatibility or by probability. In contrast, incorrect response inhibition in the primary motor cortex ipsilateral to the correct response was more pronounced on incompatible trials, especially in the condition where most of the trials were compatible, indicating a modulation of inhibitory strength within the course of the action. Two prefrontal activities, one medial and one lateral, were also observed before the response, and their potential links with the observed inhibitory pattern observed are discussed.

Linking EEG signals, brain functions and mental operations: Advantages of the Laplacian transformation.

Electroencephalography (EEG) is a very popular technique for investigating brain functions and/or mental processes. To this aim, EEG activities must be interpreted in terms of brain and/or mental processes. EEG signals being a direct manifestation of neuronal activity it is often assumed that such interpretations are quite obvious or, at least, straightforward. However, they often rely on (explicit or even implicit) assumptions regarding the structures supposed to generate the EEG activities of interest. For these assumptions to be used appropriately, reliable links between EEG activities and the underlying brain structures must be established. Because of volume conduction effects and the mixture of activities they induce, these links are difficult to establish with scalp potential recordings. We present different examples showing how the Laplacian transformation, acting as an efficient source separation method, allowed to establish more reliable links between EEG activities and brain generators and, ultimately, with mental operations. The nature of those links depends on the depth of inferences that can vary from weak to strong. Along this continuum, we show that 1) while the effects of experimental manipulation can appear widely distributed with scalp potentials, Laplacian transformation allows to reveal several generators contributing (in different manners) to these modulations, 2) amplitude variations within the same set of generators can generate spurious differences in scalp potential topographies, often interpreted as reflecting different source configurations. In such a case, Laplacian transformation provides much more similar topographies, evidencing the same generator(s) set, and 3) using the LRP as an index of response activation most often produces ambiguous results, Laplacian-transformed response-locked ERPs obtained over motor areas allow resolving these ambiguities.

Spatial and temporal resolutions of EEG: Is it really black and white? A scalp current density view.

Among the different brain imaging techniques, electroencephalography (EEG) is classically considered as having an excellent temporal resolution, but a poor spatial one. Here, we argue that the actual temporal resolution of conventional (scalp potentials) EEG is overestimated, and that volume conduction, the main cause of the poor spatial resolution of EEG, also distorts the recovered time course of the underlying sources at scalp level, and hence degrades the actual temporal resolution of EEG. While Current Source Density (CSD) estimates, through the Surface Laplacian (SL) computation, are well known to dramatically reduce volume conduction effects and hence improve EEG spatial resolution, its positive impact on EEG temporal resolution is much less recognized. In two simulation studies, we first show how volume conduction and reference electrodes distort the scalp potential time course, and how SL transform provides a much better spatio-temporal description. We then exemplify similar effects on two empirical datasets. We show how the time courses of the scalp potentials mis-estimate the latencies of the relevant brain events and that CSD provides a much richer, and much more accurate, view of the spatio-temporal dynamics of brain activity.

Controlling your impulses: electrical stimulation of the human supplementary motor complex prevents impulsive errors.

To err is human. However, an inappropriate urge does not always result in error. Impulsive errors thus entail both a motor system capture by an urge to act and a failed inhibition of that impulse. Here we show that neuromodulatory electrical stimulation of the supplementary motor complex in healthy humans leaves action urges unchanged but prevents them from turning into overt errors. Subjects performed a choice reaction-time task known to trigger impulsive responses, leading to fast errors that can be revealed by analyzing accuracy as a function of poststimulus time. Yet, such fast errors are only the tip of the iceberg: electromyography (EMG) revealed fast subthreshold muscle activation in the incorrect response hand in an even larger proportion of overtly correct trials, revealing covert response impulses not discernible in overt behavior. Analyzing both overt and covert response tendencies enables to gauge the ability to prevent these incorrect impulses from turning into overt action errors. Hyperpolarizing the supplementary motor complex using transcranial direct current stimulation (tDCS) preserves action impulses but prevents their behavioral expression. This new combination of detailed behavioral, EMG, and tDCS techniques clarifies the neurophysiology of impulse control, and may point to avenues for improving impulse control deficits in various neurologic and psychiatric disorders.

Distributional reaction time properties in the Eriksen task: marked differences or hidden similarities with the Simon task?

In conflict tasks, the irrelevant stimulus attribute needs to be suppressed for the correct response to be produced. In the Simon task, earlier researchers have proposed that this suppression is the reason that, after an initial increase, the interference effect decreases for longer RTs, as reflected by late, negative-going delta plots. This view has been challenged by observations of positive-going delta plots, even for long RTs, in other conflict tasks, despite a similar necessity for suppression. For late negative-going delta plots to be interpreted as reflecting suppression, a necessary, although maybe not sufficient, condition is that similar patterns should be observed for other conflict tasks. We reasoned that a similar suppression could be present, but hidden, in the Eriksen flanker task. By recording and analyzing electromyograms of the muscles involved in response execution, we could compute delta plots separately for trials that elicited a subthreshold incorrect response activation (partial error). Late negative-going delta plots were observable on partial-error trials, although they were weaker than for the Simon task, reducing the impact of this inversion on the overall distribution. We further showed that this pattern is modulated by time pressure. Those results indicate that mechanisms leading to negative-going delta plots, similar to those observed in the Simon task, are also at play in the Eriksen task. The link between negative-going delta plots and executive online control is discussed.

Corticospinal control of wrist muscles during expectation of a motor perturbation: a transcranial magnetic stimulation study.

Voluntary movement is often perturbed by the external forces in the environment. Because corticospinal (CS) control of wrist muscles during preparation of voluntary movement has been extensively studied without variation in the external forces, very little is known about the way CS control adapts when subjects expect motor perturbations. Here, we studied the CS control of wrist muscles during expectation of an imposed wrist extension. Subjects were instructed either to compensate (COMP) the perturbation (applied at variable delays) or not to intervene (NINT). In a quarter of all trials at random, in the time window when perturbation might occur, TMS was applied over contralateral M1. Motor evoked potentials (MEPs) were measured in the FCR (flexor carpi radialis) and ECR (extensor carpi radialis) muscles, as well as the silent period (SP) in the FCR. Following the perturbation, we found a larger long-latency stretch reflex in COMP than in NINT. During the expectation of the perturbation, MEP amplitudes did not differ across conditions in FCR. However, those evoked in ECR were greater in COMP than in NINT condition. Moreover in the FCR, the silent period lasted longer in NINT. Thus, we showed a selective effect of the prepared reaction on the anticipatory tuning of CS excitability and cortical inhibition in the agonist/antagonist muscles. This tuning clearly differed from the tuning during voluntary movement preparation without variation in the external forces. This shows that the tuning of the CS system during motor preparation depends on the dynamical context of movement production.