Parietofrontal oscillations show hand-specific interactions with top-down movement plans.

To generate a hand-specific reach plan, the brain must integrate hand-specific signals with the desired movement strategy. Although various neurophysiology/imaging studies have investigated hand-target interactions in simple reach-to-target tasks, the whole brain timing and distribution of this process remain unclear, especially for more complex, instruction-dependent motor strategies. Previously, we showed that a pro/anti pointing instruction influences magnetoencephalographic (MEG) signals in frontal cortex that then propagate recurrently through parietal cortex (Blohm G, Alikhanian H, Gaetz W, Goltz HC, DeSouza JF, Cheyne DO, Crawford JD. NeuroImage 197: 306-319, 2019). Here, we contrasted left versus right hand pointing in the same task to investigate 1) which cortical regions of interest show hand specificity and 2) which of those areas interact with the instructed motor plan. Eight bilateral areas, the parietooccipital junction (POJ), superior parietooccipital cortex (SPOC), supramarginal gyrus (SMG), medial/anterior interparietal sulcus (mIPS/aIPS), primary somatosensory/motor cortex (S1/M1), and dorsal premotor cortex (PMd), showed hand-specific changes in beta band power, with four of these (M1, S1, SMG, aIPS) showing robust activation before movement onset. M1, SMG, SPOC, and aIPS showed significant interactions between contralateral hand specificity and the instructed motor plan but not with bottom-up target signals. Separate hand/motor signals emerged relatively early and lasted through execution, whereas hand-motor interactions only occurred close to movement onset. Taken together with our previous results, these findings show that instruction-dependent motor plans emerge in frontal cortex and interact recurrently with hand-specific parietofrontal signals before movement onset to produce hand-specific motor behaviors.NEW & NOTEWORTHY The brain must generate different motor signals depending on which hand is used. The distribution and timing of hand use/instructed motor plan integration are not understood at the whole brain level. Using MEG we show that different action planning subnetworks code for hand usage and integrating hand use into a hand-specific motor plan. The timing indicates that frontal cortex first creates a general motor plan and then integrates hand specificity to produce a hand-specific motor plan.

Mapping neural dynamics underlying saccade preparation and execution and their relation to reaction time and direction errors.

Our ability to control and inhibit automatic behaviors is crucial for negotiating complex environments, all of which require rapid communication between sensory, motor, and cognitive networks. Here, we measured neuromagnetic brain activity to investigate the neural timing of cortical areas needed for inhibitory control, while 14 healthy young adults performed an interleaved prosaccade (look at a peripheral visual stimulus) and antisaccade (look away from stimulus) task. Analysis of how neural activity relates to saccade reaction time (SRT) and occurrence of direction errors (look at stimulus on antisaccade trials) provides insight into inhibitory control. Neuromagnetic source activity was used to extract stimulus-aligned and saccade-aligned activity to examine temporal differences between prosaccade and antisaccade trials in brain regions associated with saccade control. For stimulus-aligned antisaccade trials, a longer SRT was associated with delayed onset of neural activity within the ipsilateral parietal eye field (PEF) and bilateral frontal eye field (FEF). Saccade-aligned activity demonstrated peak activation 10ms before saccade-onset within the contralateral PEF for prosaccade trials and within the bilateral FEF for antisaccade trials. In addition, failure to inhibit prosaccades on anti-saccade trials was associated with increased activity prior to saccade onset within the FEF contralateral to the peripheral stimulus. This work on dynamic activity adds to our knowledge that direction errors were due, at least in part, to a failure to inhibit automatic prosaccades. These findings provide novel evidence in humans regarding the temporal dynamics within oculomotor areas needed for saccade programming and the role frontal brain regions have on top-down inhibitory control.

Speech Kinematics and Coordination Measured With an MEG-Compatible Speech Tracking System.

Articulography and functional neuroimaging are two major tools for studying the neurobiology of speech production. Until recently, however, it has generally not been possible to use both in the same experimental setup because of technical incompatibilities between the two methodologies. Here we describe results from a novel articulography system dubbed Magneto-articulography for the Assessment of Speech Kinematics (MASK), which we used to derive kinematic profiles of oro-facial movements during speech. MASK was used to characterize speech kinematics in two healthy adults, and the results were compared to measurements from a separate participant with a conventional Electromagnetic Articulography (EMA) system. Analyses targeted the gestural landmarks of reiterated utterances /ipa/, /api/ and /pataka/. The results demonstrate that MASK reliably characterizes key kinematic and movement coordination parameters of speech motor control. Since these parameters are intrinsically registered in time with concurrent magnetoencephalographic (MEG) measurements of neuromotor brain activity, this methodology paves the way for innovative cross-disciplinary studies of the neuromotor control of human speech production, speech development, and speech motor disorders.

Speech-induced suppression of evoked auditory fields in children who stutter.

Auditory responses to speech sounds that are self-initiated are suppressed compared to responses to the same speech sounds during passive listening. This phenomenon is referred to as speech-induced suppression, a potentially important feedback-mediated speech-motor control process. In an earlier study, we found that both adults who do and do not stutter demonstrated a reduced amplitude of the auditory M50 and M100 responses to speech during active production relative to passive listening. It is unknown if auditory responses to self-initiated speech-motor acts are suppressed in children or if the phenomenon differs between children who do and do not stutter. As stuttering is a developmental speech disorder, examining speech-induced suppression in children may identify possible neural differences underlying stuttering close to its time of onset. We used magnetoencephalography to determine the presence of speech-induced suppression in children and to characterize the properties of speech-induced suppression in children who stutter. We examined the auditory M50 as this was the earliest robust response reproducible across our child participants and the most likely to reflect a motor-to-auditory relation. Both children who do and do not stutter demonstrated speech-induced suppression of the auditory M50. However, children who stutter had a delayed auditory M50 peak latency to vowel sounds compared to children who do not stutter indicating a possible deficiency in their ability to efficiently integrate auditory speech information for the purpose of establishing neural representations of speech sounds.

Auditory evoked fields to vocalization during passive listening and active generation in adults who stutter.

We used magnetoencephalography to investigate auditory evoked responses to speech vocalizations and non-speech tones in adults who do and do not stutter. Neuromagnetic field patterns were recorded as participants listened to a 1 kHz tone, playback of their own productions of the vowel /i/ and vowel-initial words, and actively generated the vowel /i/ and vowel-initial words. Activation of the auditory cortex at approximately 50 and 100 ms was observed during all tasks. A reduction in the peak amplitudes of the M50 and M100 components was observed during the active generation versus passive listening tasks dependent on the stimuli. Adults who stutter did not differ in the amount of speech-induced auditory suppression relative to fluent speakers. Adults who stutter had shorter M100 latencies for the actively generated speaking tasks in the right hemisphere relative to the left hemisphere but the fluent speakers showed similar latencies across hemispheres. During passive listening tasks, adults who stutter had longer M50 and M100 latencies than fluent speakers. The results suggest that there are timing, rather than amplitude, differences in auditory processing during speech in adults who stutter and are discussed in relation to hypotheses of auditory-motor integration breakdown in stuttering.

Non-invasive Investigation of Human Hippocampal Rhythms Using Magnetoencephalography: A Review.

Hippocampal rhythms are believed to support crucial cognitive processes including memory, navigation, and language. Due to the location of the hippocampus deep in the brain, studying hippocampal rhythms using non-invasive magnetoencephalography (MEG) recordings has generally been assumed to be methodologically challenging. However, with the advent of whole-head MEG systems in the 1990s and development of advanced source localization techniques, simulation and empirical studies have provided evidence that human hippocampal signals can be sensed by MEG and reliably reconstructed by source localization algorithms. This paper systematically reviews simulation studies and empirical evidence of the current capacities and limitations of MEG "deep source imaging" of the human hippocampus. Overall, these studies confirm that MEG provides a unique avenue to investigate human hippocampal rhythms in cognition, and can bridge the gap between animal studies and human hippocampal research, as well as elucidate the functional role and the behavioral correlates of human hippocampal oscillations.

Cortical reorganization after modified constraint-induced movement therapy in pediatric hemiplegic cerebral palsy.

Constraint-induced movement therapy improves motor function in the affected hand of children with hemiplegic cerebral palsy and results in cortical changes in adults with stroke. This study measured clinical improvement and cortical reorganization in a child with hemiplegia who underwent modified constraint-induced movement therapy for 3 weeks. Clinical, functional magnetic resonance imaging and magnetoencephalography measurements were done at baseline, after therapy, and 6 months after therapy. Modified constraint-induced movement therapy resulted in clinical improvement as measured by the Pediatric Motor Activity Log. Functional magnetic resonance imaging showed bilateral sensorimotor activation before and after therapy and a shift in the laterality index from ipsilateral to contralateral hemisphere after therapy. Magnetoencephalography showed increased cortical activation in the ipsilateral motor field and contralateral movement evoked field after therapy. Cortical reorganization was maintained at the 6-month follow-up. This is the first study to demonstrate cortical reorganization after any version of constraint-induced movement therapy in a child with hemiplegia.

Neural processing of observed oro-facial movements reflects multiple action encoding strategies in the human brain.

In this experiment, the oscillatory responses of the MEG were characterized during the observation of four viewing conditions: (a) observation of mouth movements, (b) observation of a non-biological motion stimulus (a mechanical aperture opening and shutting), (c) observation of object-directed mouth movements and (d) observation of speech-like mouth movements. Data were analyzed using synthetic aperture magnetometry (SAM) in three frequency bands, beta (15-35 Hz), gamma (35-70 Hz) and alpha/mu (8-15 Hz). Results showed that observations of biological motion resulted in beta desynchronization over lateral sensorimotor areas, while observations of non-biological motion resulted in a more medial desynchronization, an effect that may be related to the processing of a structured event sequence. Observation of linguistic movements resulted in less alpha/beta desynchronization in posterior brain regions in comparison to biological motion stimuli, suggesting that linguistically-relevant stimuli are processed with different neuronal systems than those recruited by normal action observation. We suggest that non-linguistic actions recruit dorsal systems while linguistic actions engage ventral processing systems. Object-directed movements showed the largest sensorimotor activations, suggesting that, as is the case for observations of hand movements, motoric processing is particularly sensitive to the viewing of goal-directed actions. Taken together, the results indicate that the brain utilizes multiple action encoding strategies, tailored to the function of the observed movement.

Spatiotemporal analysis of feedback processing during a card sorting task using spatially filtered MEG.

A card sorting paradigm was used to observe the neural correlates of feedback processing in adult participants. Visually presented feedback was used to indicate response accuracy and the requirement to shift response set in a 2-category card sorting task. Magnetoencephalography (MEG) responses to feedback cues were analysed using a beamformer-based spatial filtering algorithm (event-related Synthetic Aperture Magnetometry, erSAM). Analysis of source power revealed activity in rostral anterior cingulate cortex (ACC) only to negative feedback processing, which peaked at 260 ms after stimulus onset. The results are in agreement with both evidence from fMRI on spatial characteristics of negative feedback processing, and evidence from event-related potentials (ERPs) on the temporal profile of this ACC response. The superior temporal gyrus was activated only with positive feedback, reflecting integration of actions with successful outcomes. The present MEG erSAM findings are the first to provide both accurate spatial localization as well as temporal specificity for the neural correlates of feedback processing.

Sensorimotor Oscillations Prior to Speech Onset Reflect Altered Motor Networks in Adults Who Stutter.

Adults who stutter (AWS) have demonstrated atypical coordination of motor and sensory regions during speech production. Yet little is known of the speech-motor network in AWS in the brief time window preceding audible speech onset. The purpose of the current study was to characterize neural oscillations in the speech-motor network during preparation for and execution of overt speech production in AWS using magnetoencephalography (MEG). Twelve AWS and 12 age-matched controls were presented with 220 words, each word embedded in a carrier phrase. Controls were presented with the same word list as their matched AWS participant. Neural oscillatory activity was localized using minimum-variance beamforming during two time periods of interest: speech preparation (prior to speech onset) and speech execution (following speech onset). Compared to controls, AWS showed stronger beta (15-25 Hz) suppression in the speech preparation stage, followed by stronger beta synchronization in the bilateral mouth motor cortex. AWS also recruited the right mouth motor cortex significantly earlier in the speech preparation stage compared to controls. Exaggerated motor preparation is discussed in the context of reduced coordination in the speech-motor network of AWS. It is further proposed that exaggerated beta synchronization may reflect a more strongly inhibited motor system that requires a stronger beta suppression to disengage prior to speech initiation. These novel findings highlight critical differences in the speech-motor network of AWS that occur prior to speech onset and emphasize the need to investigate further the speech-motor assembly in the stuttering population.

Localization of human somatosensory cortex using spatially filtered magnetoencephalography.

A spatial filter algorithm based on minimum-variance beamforming (synthetic aperture magnetometry (SAM)) was applied to single trial neuromagnetic recordings in order to localize primary somatosensory cortex. Magnetoencephalography (MEG) responses to electrical stimulation of the right and left median nerve were recorded using a whole-head MEG system and localized using both SAM spatial filtering and dipole analysis. Spatial filtering was applied to single trial neuromagnetic recordings to produce 3-dimensional difference images of source power between active (0-50 ms) and control states (-50-0 ms) in the range of 15-300 Hz. Average difference between N20m dipole location and location of maximal increase in power in the SAM images was 3.7 mm (1.5 mm SD) and localized to primary somatosensory cortex. Time-frequency analysis of spatially filtered output for the peak SAM locations showed a brief (10 ms) increase in the 60-100 Hz band coincident with the N20m response and a longer duration (approx. 80 ms) increase in power in the 10-40 Hz band following N20m onset. These results indicate that beamformer based spatial filter methods such as SAM can be used to localize temporally discrete cortical activity produced by median nerve stimulation.

Adaptive cluster analysis approach for functional localization using magnetoencephalography.

In this paper we propose an agglomerative hierarchical clustering Ward's algorithm in tandem with the Affinity Propagation algorithm to reliably localize active brain regions from magnetoencephalography (MEG) brain signals. Reliable localization of brain areas with MEG has been difficult due to variations in signal strength, and the spatial extent of the reconstructed activity. The proposed approach to resolve this difficulty is based on adaptive clustering on reconstructed beamformer images to find locations that are consistently active across different participants and experimental conditions with high spatial resolution. Using data from a human reaching task, we show that the method allows more accurate and reliable localization from MEG data alone without using functional magnetic resonance imaging (fMRI) or any other imaging techniques.

Intended actions and unexpected outcomes: automatic and controlled processing in a rapid motor task.

Human action involves a combination of controlled and automatic behavior. These processes may interact in tasks requiring rapid response selection or inhibition, where temporal constraints preclude timely intervention by conscious, controlled processes over automatized prepotent responses. Such contexts tend to produce frequent errors, but also rapidly executed correct responses, both of which may sometimes be perceived as surprising, unintended, or "automatic". In order to identify neural processes underlying these two aspects of cognitive control, we measured neuromagnetic brain activity in 12 right-handed subjects during manual responses to rapidly presented digits, with an infrequent target digit that required switching response hand (bimanual task) or response finger (unimanual task). Automaticity of responding was evidenced by response speeding (shorter response times) prior to both failed and fast correct switches. Consistent with this automaticity interpretation of fast correct switches, we observed bilateral motor preparation, as indexed by suppression of beta band (15-30 Hz) oscillations in motor cortex, prior to processing of the switch cue in the bimanual task. In contrast, right frontal theta activity (4-8 Hz) accompanying correct switch responses began after cue onset, suggesting that it reflected controlled inhibition of the default response. Further, this activity was reduced on fast correct switch trials suggesting a more automatic mode of inhibitory control. We also observed post-movement (event-related negativity) ERN-like responses and theta band increases in medial and anterior frontal regions that were significantly larger on error trials, and may reflect a combination of error and delayed inhibitory signals. We conclude that both automatic and controlled processes are engaged in parallel during rapid motor tasks, and that the relative strength and timing of these processes may underlie both optimal task performance and subjective experiences of automaticity or control.