Does "the motor system" play "a role" in speech perception? If so, where, how, and when? We conducted a systematic review that addresses these questions using both qualitative and quantitative methods. The qualitative review of behavioural, computational modelling, non-human animal, brain damage/disorder, electrical stimulation/recording, and neuroimaging research suggests that distributed brain regions involved in producing speech play specific, dynamic, and contextually determined roles in speech perception. The quantitative review employed region and network based neuroimaging meta-analyses and a novel text mining method to describe relative contributions of nodes in distributed brain networks. Supporting the qualitative review, results show a specific functional correspondence between regions involved in non-linguistic movement of the articulators, covertly and overtly producing speech, and the perception of both nonword and word sounds. This distributed set of cortical and subcortical speech production regions are ubiquitously active and form multiple networks whose topologies dynamically change with listening context. Results are inconsistent with motor and acoustic only models of speech perception and classical and contemporary dual-stream models of the organization of language and the brain. Instead, results are more consistent with complex network models in which multiple speech production related networks and subnetworks dynamically self-organize to constrain interpretation of indeterminant acoustic patterns as listening context requires.

The idea that humans learn and maintain accurate speech by carefully monitoring auditory feedback is widely held. But this view neglects the fact that auditory feedback is highly correlated with somatosensory feedback during speech production. Somatosensory feedback from speech movements could be a primary means by which cortical speech areas monitor the accuracy of produced speech. We tested this idea by placing the somatosensory and auditory systems in competition during speech motor learning. To do this, we combined two speech learning paradigms to simultaneously alter somatosensory and auditory feedback in real-time as subjects spoke. Somatosensory feedback was manipulated by using a robotic device that altered the motion path of the jaw. Auditory feedback was manipulated by changing the frequency of the first formant of the vowel sound and playing back the modified utterance to the subject through headphones. The amount of compensation for each perturbation was used as a measure of sensory reliance. All subjects were observed to correct for at least one of the perturbations, but auditory feedback was not dominant. Indeed, some subjects showed a stable preference for either somatosensory or auditory feedback during speech.

Recent studies of human speech motor learning suggest that learning is accompanied by changes in auditory perception. But what drives the perceptual change? Is it a consequence of changes in the motor system? Or is it a result of sensory inflow during learning? Here, subjects participated in a speech motor-learning task involving adaptation to altered auditory feedback and they were subsequently tested for perceptual change. In two separate experiments, involving two different auditory perceptual continua, we show that changes in the speech motor system that accompany learning drive changes in auditory speech perception. Specifically, we obtained changes in speech perception when adaptation to altered auditory feedback led to speech production that fell into the phonetic range of the speech perceptual tests. However, a similar change in perception was not observed when the auditory feedback that subjects' received during learning fell into the phonetic range of the perceptual tests. This indicates that the central motor outflow associated with vocal sensorimotor adaptation drives changes to the perceptual classification of speech sounds.

The perception of speech is notably malleable in adults, yet alterations in perception seem to have little impact on speech production. We hypothesized that speech perceptual training might immediately influence speech motor learning. To test this, we paired a speech perceptual training task with a speech motor learning task. Subjects performed a series of perceptual tests designed to measure and then manipulate the perceptual distinction between the words “head” and “had”. Subjects then produced “head” with the sound of the vowel altered in real-time so that they heard themselves through headphones producing a word that sounded more like “had”. In support of our hypothesis, the amount of motor learning in response to the voice alterations depended on the perceptual boundary acquired through perceptual training. The studies show that plasticity in adult speech perception can have immediate consequences for speech production in the context of speech learning.

Humans routinely make movements to targets that have different accuracy requirements in different directions. Examples extend from everyday occurrences such as grasping the handle of a coffee cup to the more refined instance of a surgeon positioning a scalpel. The attainment of accuracy in situations such as these might be related to the nervous system's capacity to regulate the limb's resistance to displacement, or impedance. To test this idea, subjects made movements from random starting locations to targets that had shape-dependent accuracy requirements. We used a robotic device to assess both limb impedance and patterns of movement variability just as the subject reached the target. We show that impedance increases in directions where required accuracy is high. Independent of target shape, patterns of limb stiffness are seen to predict spatial patterns of movement variability. The nervous system is thus seen to modulate limb impedance in entirely predictable environments to aid in the attainment of reaching accuracy.

The motor cortex and cerebellum are thought to be critical for learning and maintaining motor behaviors. Here we use transcranial direct current stimulation (tDCS) to test the role of the motor cortex and cerebellum in sensorimotor learning in speech. During productions of "head," "bed," and "dead," the first formant of the vowel sound was altered in real time toward the first formant of the vowel sound in "had," "bad," and "dad." Compensatory changes in first and second formant production were used as a measure of motor adaptation. tDCS to either the motor cortex or the cerebellum improved sensorimotor learning in speech compared with sham stimulation ( n = 20 in each group). However, in the case of cerebellar tDCS, production changes were restricted to the source of the acoustical error (i.e., the first formant). Motor cortex tDCS drove production changes that offset errors in the first formant, but unlike cerebellar tDCS, adaptive changes in the second formant also occurred. The results suggest that motor cortex and cerebellar tDCS have both shared and dissociable effects on motor adaptation. The study provides initial causal evidence in speech production that the motor cortex and the cerebellum support different aspects of sensorimotor learning. We propose that motor cortex tDCS drives sensorimotor learning toward previously learned patterns of movement, whereas cerebellar tDCS focuses sensorimotor learning on error correction.

Highlights d Sensorimotor learning was observed during the fluid production of variable sentences d Sensorimotor learning for sentences transferred robustly to the production of words d The brain predicts the sensory consequences of variable, sentence-level speech d Sensory prediction errors rapidly drive precise changes in fluid speech production

The role of the cerebellum in speech perception remains a mystery. Given its uniform architecture, we tested the hypothesis that it implements a domain-general mechanism whose role in speech is determined by connectivity. We collated all neuroimaging studies reporting cerebellar activity in the Neurosynth database (n = 8206). From this set, we found all studies involving passive speech and sound perception (n = 72, 64% speech, 12.5% sounds, 12.5% music, and 11% tones) and speech production and articulation (n = 175). Standard and coactivation neuroimaging meta-analyses were used to compare cerebellar and associated cortical activations between passive perception and production. We found distinct regions of perception- and production-related activity in the cerebellum and regions of perception–production overlap. Each of these regions had distinct patterns of cortico-cerebellar connectivity. To test for domain-generality versus specificity, we identified all psychological and task-related terms in the Neurosynth database that predicted activity in cerebellar regions associated with passive perception and production. Regions in the cerebellum activated by speech perception were associated with domain-general terms related to prediction. One hallmark of predictive processing is metabolic savings (i.e., decreases in neural activity when events are predicted). To test the hypothesis that the cerebellum plays a predictive role in speech perception, we examined cortical activation between studies reporting cerebellar activation and those without cerebellar activation during speech perception. When the cerebellum was active during speech perception, there was far less cortical activation than when it was inactive. The results suggest that the cerebellum implements a domain-general mechanism related to prediction during speech perception.