Speech listening entails neural encoding of invisible articulatory features.

Speech processing entails a complex interplay between bottom-up and top-down computations. The former is reflected in the neural entrainment to the quasi-rhythmic properties of speech acoustics while the latter is supposed to guide the selection of the most relevant input subspace. Top-down signals are believed to originate mainly from motor regions, yet similar activities have been shown to tune attentional cycles also for simpler, non-speech stimuli. Here we examined whether, during speech listening, the brain reconstructs articulatory patterns associated to speech production. We measured electroencephalographic (EEG) data while participants listened to sentences during the production of which articulatory kinematics of lips, jaws and tongue were also recorded (via Electro-Magnetic Articulography, EMA). We captured the patterns of articulatory coordination through Principal Component Analysis (PCA) and used Partial Information Decomposition (PID) to identify whether the speech envelope and each of the kinematic components provided unique, synergistic and/or redundant information regarding the EEG signals. Interestingly, tongue movements contain both unique as well as synergistic information with the envelope that are encoded in the listener's brain activity. This demonstrates that during speech listening the brain retrieves highly specific and unique motor information that is never accessible through vision, thus leveraging audio-motor maps that arise most likely from the acquisition of speech production during development.

A questionnaire to collect unintended effects of transcranial magnetic stimulation: A consensus based approach.

Transcranial magnetic stimulation (TMS) has been widely used in both clinical and research practice. However, TMS might induce unintended sensations and undesired effects as well as serious adverse effects. To date, no shared forms are available to report such unintended effects. This study aimed at developing a questionnaire enabling reporting of TMS unintended effects. A Delphi procedure was applied which allowed consensus among TMS experts. A steering committee nominated a number of experts to be involved in the Delphi procedure. Three rounds were conducted before reaching a consensus. Afterwards, the questionnaire was publicized on the International Federation of Clinical Neurophysiology website to collect further suggestions by the wider scientific community. A last Delphi round was then conducted to obtain consensus on the suggestions collected during the publicization and integrate them in the questionnaire. The procedure resulted in a questionnaire, that is the TMSens_Q, applicable in clinical and research settings. Routine use of the structured TMS questionnaire and standard reporting of unintended TMS effects will help to monitor the safety of TMS, particularly when applying new protocols. It will also improve the quality of data collection as well as the interpretation of experimental findings.

Motor synergies: Evidence for a novel motor signature in autism spectrum disorder.

In autism spectrum disorder (ASD), socio-communicative impairments and stereotypical behaviours are paralleled by sensorimotor deficits. Individuals with ASD show an altered selection of motor parameters, resulting in clumsy and fragmented actions. Here, we investigated inter-joint coordination and motor synergies as a potential substrate of motor control problems in ASD. Synergies enable co-controlling redundant motor degrees of freedom (DoF, e.g. joint angles, muscles) by mapping behavioural goals into a flexible and low-dimensional set of variables. This mechanism simplifies motor control and helps to find unambiguous solutions for motor tasks. In a reaching-grasping paradigm, children with ASD showed reduced coupling between DoF, which correlated with socio-communicative symptoms severity. Impaired synergies may help to frame well-established motor problems in ASD, including impaired motor sequencing and abnormal trial-to-trial motor variability. On the other hand, synergies also provide an effective and compact coding system of observed actions. Impaired synergies may thus jeopardize motor interaction by initiating bottom-up cascade effects, leading to pervasive impairments of social behaviour. Finally, we trained an automatic classification algorithm to distinguish between ASD and typically developing (TD) participants based on reaching-grasping kinematics. Classification accuracy reached up to 0.947. This result corroborates and expands previous accounts claiming that motor-based early recognition is feasible and effective in ASD.

Motor Recruitment during Action Observation: Effect of Interindividual Differences in Action Strategy.

Visual processing of other's actions is supported by sensorimotor brain activations. Access to sensorimotor representations may, in principle, provide the top-down signal required to bias search and selection of critical visual features. For this to happen, it is necessary that a stable one-to-one mapping exists between observed kinematics and underlying motor commands. However, due to the inherent redundancy of the human musculoskeletal system, this is hardly the case for multijoint actions where everyone has his own moving style (individual motor signature-IMS). Here, we investigated the influence of subject's IMS on subjects' motor excitability during the observation of an actor achieving the same goal by adopting two different IMSs. Despite a clear dissociation in kinematic and electromyographic patterns between the two actions, we found no group-level modulation of corticospinal excitability (CSE) in observers. Rather, we found a negative relationship between CSE and actor-observer IMS distance, already at the single-subject level. Thus, sensorimotor activity during action observation does not slavishly replicate the motor plan implemented by the actor, but rather reflects the distance between what is canonical according to one's own motor template and the observed movements performed by other individuals.

Publisher Correction: Anticipatory postural adjustments during joint action coordination.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Anticipatory postural adjustments during joint action coordination.

There is a current claim that humans are able to effortlessly detect others' hidden mental state by simply observing their movements and transforming the visual input into motor knowledge to predict behaviour. Using a classical paradigm quantifying motor predictions, we tested the role of vision feedback during a reach and load-lifting task performed either alone or with the help of a partner. Wrist flexor and extensor muscle activities were recorded on the supporting hand. Early muscle changes preventing limb instabilities when participants performed the task by themselves revealed the contribution of the visual input in postural anticipation. When the partner performed the unloading, a condition mimicking a split-brain situation, motor prediction followed a pattern evolving along the task course and changing with the integration of successive somatosensory feedback. Our findings demonstrate that during social behaviour, in addition to self-motor representations, individuals cooperate by continuously integrating sensory signals from various sources.

Direction-dependent activation of the insular cortex during vertical and horizontal hand movements.

The planning of any motor action requires a complex multisensory processing by the brain. Gravity - immutable on Earth - has been shown to be a key input to these mechanisms. Seminal fMRI studies performed during visual perception of falling objects and self-motion demonstrated that humans represent the action of gravity in parts of the cortical vestibular system; in particular, the insular cortex and the cerebellum. However, little is known as to whether a specific neural network is engaged when processing non-visual signals relevant to gravity. We asked participants to perform vertical and horizontal hand movements without visual control, while lying in a 3T-MRI scanner. We highlighted brain regions activated in the processing of vertical movements, for which the effects of gravity changed during execution. Precisely, the left insula was activated in vertical movements and not in horizontal movements. Moreover, the network identified by contrasting vertical and horizontal movements overlapped with neural correlates previously associated to the processing of simulated self-motion and visual perception of the vertical direction. Interestingly, we found that the insular cortex activity is direction-dependent which suggests that this brain region processes the effects of gravity on the moving limbs through non-visual signals.

Bimanual non-congruent actions in motor neglect syndrome: a combined behavioral/fMRI study.

In Motor Neglect (MN) syndrome, a specific impairment in non-congruent bimanual movements has been described. In the present case-control study, we investigated the neuro-functional correlates of this behavioral deficit. Two right-brain-damaged (RBD) patients, one with (MN+) and one without (MN-) MN, were evaluated by means of functional Magnetic Resonance Imaging (fMRI) in a bimanual Circles-Lines (CL) paradigm. Patients were requested to perform right-hand movements (lines-drawing) and, simultaneously, congruent (lines-drawing) or non-congruent (circles-drawing) left-hand movements. In the behavioral task, MN- patient showed a bimanual-coupling-effect, while MN+ patient did not. The fMRI study showed that in MN-, a fronto-parietal network, mainly involving the pre-supplementary motor area (pre-SMA) and the posterior parietal cortex (PPC), was significantly more active in non-congruent than in congruent conditions, as previously shown in healthy subjects. On the contrary, MN+ patient showed an opposite pattern of activation both in pre-SMA and in PPC. Within this fronto-parietal network, the pre-SMA is supposed to exert an inhibitory influence on the default coupling of homologous muscles, thus allowing the execution of non-congruent movements. In MN syndrome, the described abnormal pre-SMA activity supports the hypothesis that a failure to inhibit ipsilesional motor programs might determine a specific impairment of non-congruent movements.

Distinct brain signatures of content and structure violation during action observation.

Sentences, musical phrases and goal-directed actions are composed of elements that are linked by specific rules to form meaningful outcomes. In goal-directed actions including a non-canonical element or scrambling the order of the elements alters the action's content and structure, respectively. In the present study we investigated event-related potentials of the electroencephalographic (EEG) activity recorded during observation of both alterations of the action content (obtained by violating the semantic components of an action, e.g. making coffee with cola) and alterations of the action structure (obtained by inverting the order of two temporally adjacent pictures of sequences depicting daily life actions) interfering with the normal flow of the motor acts that compose an action. Action content alterations elicited a bilateral posterior distributed EEG negativity, peaking at around 400 ms after stimulus onset similar to the ERPs evoked by semantic violations in language studies. Alteration of the action structure elicited an early left anterior negativity followed by a late left anterior positivity, which closely resembles the ERP pattern found in language syntax violation studies. Our results suggest a functional dissociation between the processing of action content and structure, reminiscent of a similar dissociation found in the language or music domains. Importantly, this study provides further support to the hypothesis that some basic mechanisms, such as the rule-based structuring of sequential events, are shared between different cognitive domains.

Improving the reliability of single-subject fMRI by weighting intra-run variability.

At present, functional magnetic resonance imaging (fMRI) is one of the most useful methods of studying cognitive processes in the human brain in vivo, both for basic science and clinical goals. Although neuroscience studies often rely on group analysis, clinical applications must investigate single subjects (patients) only. Particularly for the latter, issues regarding the reliability of fMRI readings remain to be resolved. To determine the ability of intra-run variability (IRV) weighting to consistently detect active voxels, we first acquired fMRI data from a sample of healthy subjects, each of whom performed 4 runs (4 blocks each) of self-paced finger-tapping. Each subject's data was analyzed using single-run general linear model (GLM), and each block was then analyzed separately to calculate the IRV weighting. Results show that integrating IRV information into standard single-subject GLM activation maps significantly improved the reliability (p=0.007) of the single-subject fMRI data. This suggests that taking IRV into account can help identify the most constant and relevant neuronal activity at the single-subject level.

Listening to speech recruits specific tongue motor synergies as revealed by transcranial magnetic stimulation and tissue-Doppler ultrasound imaging.

The activation of listener's motor system during speech processing was first demonstrated by the enhancement of electromyographic tongue potentials as evoked by single-pulse transcranial magnetic stimulation (TMS) over tongue motor cortex. This technique is, however, technically challenging and enables only a rather coarse measurement of this motor mirroring. Here, we applied TMS to listeners' tongue motor area in association with ultrasound tissue Doppler imaging to describe fine-grained tongue kinematic synergies evoked by passive listening to speech. Subjects listened to syllables requiring different patterns of dorso-ventral and antero-posterior movements (/ki/, /ko/, /ti/, /to/). Results show that passive listening to speech sounds evokes a pattern of motor synergies mirroring those occurring during speech production. Moreover, mirror motor synergies were more evident in those subjects showing good performances in discriminating speech in noise demonstrating a role of the speech-related mirror system in feed-forward processing the speaker's ongoing motor plan.

Functional effect of short-term immobilization: kinematic changes and recovery on reaching-to-grasp.

Although previous investigations agree in showing significant cortical modifications related to short-term limb immobilization, little is known about the functional changes induced by non-use. To address this issue, we studied the kinematic effect of 10h of hand immobilization. In order to prevent any movement, right handed healthy participants wore on their dominant hand a soft bandage. They were requested to perform the same reaching-to-grasping task immediately after immobilization, 1 day before (baseline 1) and in other two following days without non-use (baseline 2 and baseline 3). While no differences were found among baseline conditions, an increase of the total duration of reaching movement together with an anticipation of the time to peak velocity were observed in the first trial after immobilization. Interestingly, these initial effects decreased quickly trial-by-trial, following an exponential function till reaching values equal to those observed in the control conditions. The present findings show firstly that the transport phase of the reaching-to-grasp task was affected by a temporary reduction of sensory and motor information. Secondly, a trial-by-trial recovery of the immobilization-related changes, likely driven by the sensory inputs and motor outputs associated to the repetition of the movement has been observed. All together these results confirm a fundamental role of a continuous stream of sensorimotor signals in maintaining motor efficiency and in driving recovery process.

A multi-channel low-power system-on-chip for single-unit recording and narrowband wireless transmission of neural signal.

This paper reports a multi-channel neural recording system-on-chip (SoC) with digital data compression and wireless telemetry. The circuit consists of a 16 amplifiers, an analog time division multiplexer, an 8-bit SAR AD converter, a digital signal processor (DSP) and a wireless narrowband 400-MHz binary FSK transmitter. Even though only 16 amplifiers are present in our current die version, the whole system is designed to work with 64 channels demonstrating the feasibility of a digital processing and narrowband wireless transmission of 64 neural recording channels. A digital data compression, based on the detection of action potentials and storage of correspondent waveforms, allows the use of a 1.25-Mbit/s binary FSK wireless transmission. This moderate bit-rate and a low frequency deviation, Manchester-coded modulation are crucial for exploiting a narrowband wireless link and an efficient embeddable antenna. The chip is realized in a 0.35- εm CMOS process with a power consumption of 105 εW per channel (269 εW per channel with an extended transmission range of 4 m) and an area of 3.1 × 2.7 mm(2). The transmitted signal is captured by a digital TV tuner and demodulated by a wideband phase-locked loop (PLL), and then sent to a PC via an FPGA module. The system has been tested for electrical specifications and its functionality verified in in-vivo neural recording experiments.

Lexicality drives audio-motor transformations in Broca's area.

Broca's area is classically associated with speech production. Recently, Broca's area has also been implicated in speech perception and non-linguistic information processing. With respect to the latter function, Broca's area is considered to be a central area in a network constituting the human mirror system, which maps observed or heard actions onto motor programs to execute analogous actions. These mechanisms share some similarities with Liberman's motor theory, where objects of speech perception correspond to listener's intended articulatory gestures. The aim of the current series of behavioral, TMS and fMRI studies was to test if Broca's area is indeed implicated in such audio-motor transformations. More specifically, using a classical phonological rhyme priming paradigm, we investigated whether the role of Broca's area could be purely phonological or rather, is lexical in nature. In the behavioral baseline study, we found a large priming effect in word prime/target pairs (W-W) and no effect for pseudo-words (PW-PW). Online TMS interference of Broca's area canceled the priming difference between W-W and PW-PW by enhancing the effects for PW-PW. Finally, the fMRI study showed activation of Broca's area for W-W pairs, but not for PW-PW pairs. Our data show that Broca's area plays a significant role in speech perception strongly linked to the lexicality of a stimulus.

The influence of hand posture on corticospinal excitability during motor imagery: a transcranial magnetic stimulation study.

In order to study the interaction between proprioceptive information and motor imagery, we herein investigate how compatible and incompatible postural signals influence corticospinal excitability during the mental simulation of hand movements. Subjects were asked to imagine themselves joining the tips of the thumb and the little finger while they maintained one of the two following hand postures: posture A (PA, compatible), little finger, index and thumb extended, the remaining fingers flexed; or posture B (PB, incompatible), index and thumb extended, other fingers flexed. All subjects rated the imagined finger opposition movements as easier to perform when the hand was kept in PA than in PB (P < 0.01) and the correlation between the duration of motor imagery and movement execution was also higher for PA than PB (P < 0.01). For each posture, motor evoked potentials (MEPs) elicited by focal transcranial magnetic stimulation (TMS) of the left motor cortex were recorded from the right opponens pollicis muscle during both motor imagery (MI) and rest (R) conditions. MEP area varied according to the hand posture: PA induced a higher increase in corticospinal excitability, when compared with PB. These results indicate that the actual limb posture affects the process of motor imagery. The source of this postural modulation effect is discussed.

Long-latency motor evoked potentials in congenital hemiplegia.

OBJECTIVE: To investigate long-latency motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation in congenital hemiplegia (CH) and to seek for correlation with paretic hand movement deficits. METHODS: MEPs were recorded from the first dorsal interosseous of both hands in 12 CH patients and 12 age-matched controls; dexterity and upper limb function were quantitatively assessed in both groups. RESULTS: In CH patients, long-latency MEPs, occurring much later than the commonly reported MEPs, were frequently observed in the paretic and non-paretic hands. Four distinct groups of long-latency MEPs were found, each cluster being identified by its mean latency, namely 35, 85, 160 and 225 ms. The residual dexterity of the paretic hand was correlated with the presence of contralateral MEPs with a 20 and 225 ms latency and was negatively correlated with ipsilateral MEPs, irrespective of their latency. In controls, only few MEPs with a latency of 225 ms were found in 4 out of 12 subjects. CONCLUSIONS: The pattern of MEPs found in CH patients differs dramatically from that reported in adult stroke patients, suggesting that long-latency MEPs are a rather distinctive consequence of early corticospinal lesions. The hypothesis that a given cluster of long-latency MEPs is mediated by a particular pathway appears very unlikely. Rather, we suggest that an exacerbation of cortical and/or spinal excitability is at the origin of these long-latency MEPs.

Peripheral oculomotor palsy affects orienting of visuospatial attention.

Patients affected by VI cranial nerve palsy were required to orient their attention in monocular vision and to detect a stimulus appearing either in attended or in unattended locations. Results showed that while during non-paretic eye vision stimulus detection in the attended location was faster than that in the unattended one, during paretic eye vision no difference in detection speed was present. However, in this latter condition, detection speed in both attended and un attended locations were as fast as that measured during non-paretic eye vision in attended location. Demonstration that peripheral oculomotor impairment influences monocular covert orienting of visuospatial attention strongly support the idea that visuospatial attention and oculomotor mechanisms share similar cortical networks.

I know what you are doing. a neurophysiological study.

In the ventral premotor cortex of the macaque monkey, there are neurons that discharge both during the execution of hand actions and during the observation of the same actions made by others (mirror neurons). In the present study, we show that a subset of mirror neurons becomes active during action presentation and also when the final part of the action, crucial in triggering the response in full vision, is hidden and can therefore only be inferred. This implies that the motor representation of an action performed by others can be internally generated in the observer's premotor cortex, even when a visual description of the action is lacking. The present findings support the hypothesis that mirror neuron activation could be at the basis of action recognition.

An integrated environment for stereoscopic acquisition, off-line 3D elaboration, and visual presentation of biological actions.

We present an integrated environment for stereoscopic acquisition, off-line 3D elaboration, and visual presentation of biological hand actions. The system is used in neurophysiological experiments aimed at the investigation of the parameters of the external stimuli that mirror neurons visually extract and match on their movement related activity.

Cortical mechanism for the visual guidance of hand grasping movements in the monkey: A reversible inactivation study.

Picking up an object requires two basic motor operations: reaching and grasping. Neurophysiological studies in monkeys have suggested that the visuomotor transformations necessary for these two operations are carried out by separate parietofrontal circuits and that, for grasping, a key role is played by a specific sector of the ventral premotor cortex: area F5. The aim of the present study was to test the validity of this hypothesis by reversibly inactivating area F5 in monkeys trained to grasp objects of different shape, size and orientation. In separate sessions, the hand field of the primary motor cortex (area F1 or area 4) was also reversibly inactivated. The results showed that after inactivation of area F5 buried in the bank of the arcuate sulcus (the F5 sector where visuomotor neurones responding to object presentation are located), the hand shaping preceding grasping was markedly impaired and the hand posture was not appropriate for the object size and shape. The monkeys were eventually able to grasp the objects, but only after a series of corrections made under tactile control. With small inactivations the deficits concerned the contralesional hand, with larger inactivations the ipsilateral hand as well. In addition, there were signs of peripersonal neglect in the hemispace contralateral to the inactivation site. Following inactivation of area F5 lying on the cortical convexity (the F5 sector where visuomotor neurones responding to action observation, 'mirror neurones', are found) only a motor slowing was observed, the hand shaping being preserved. The inactivation of the hand field of area F1 produced a severe paralysis of contralateral finger movements with hypotonia. The results of this study indicate the crucial role of the ventral premotor cortex in visuomotor transformations for grasping movements. More generally, they provide strong support for the notion that distal and proximal movement organization relies upon distinct cortical circuits. Clinical data on distal movement deficits in humans are re-examined in the light of the present findings.