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Brain electroencephalography (EEG), the complex, weak, multivariate, nonlinear, and nonstationary time series, has been recently widely applied in neurocognitive disorder diagnoses and brain-machine interface developments. With its specific features, unlabeled EEG is not well addressed by conventional unsupervised time-series learning methods. In this article, we handle the problem of unlabeled EEG time-series clustering and propose a novel EEG clustering algorithm, that we call mwcEEGc. The idea is to map the EEG clustering to the maximum-weight clique (MWC) searching in an improved Fréchet similarity-weighted EEG graph. The mwcEEGc considers the weights of both vertices and edges in the constructed EEG graph and clusters EEG based on their similarity weights instead of calculating the cluster centroids. To the best of our knowledge, it is the first attempt to cluster unlabeled EEG trials using MWC searching. The mwcEEGc achieves high-quality clusters with respect to intracluster compactness as well as intercluster scatter. We demonstrate the superiority of mwcEEGc over ten state-of-the-art unsupervised learning/clustering approaches by conducting detailed experimentations with the standard clustering validity criteria on 14 real-world brain EEG datasets. We also present that mwcEEGc satisfies the theoretical properties of clustering, such as richness, consistency, and order independence.

Human cortical activity measured with magnetoencephalography (MEG) has been shown to track the temporal regularity of linguistic information in connected speech. In the current study, we investigate the underlying neural sources of these responses and test the hypothesis that they can be directly modulated by changes in speech intelligibility. MEG responses were measured to natural and spectrally degraded (noise-vocoded) speech in 19 normal hearing participants. Results showed that cortical coherence to “abstract” linguistic units with no accompanying acoustic cues (phrases and sentences) were lateralized to the left hemisphere and changed parametrically with intelligibility of speech. In contrast, responses coherent to words/syllables accompanied by acoustic onsets were bilateral and insensitive to intelligibility changes. This dissociation suggests that cerebral responses to linguistic information are directly affected by intelligibility but also powerfully shaped by physical cues in speech. This explains why previous studies have reported widely inconsistent effects of speech intelligibility on cortical entrainment and, within a single experiment, provided clear support for conclusions about language lateralization derived from a large number of separately conducted neuroimaging studies. Since noise-vocoded speech resembles the signals provided by a cochlear implant device, the current methodology has potential clinical utility for assessment of cochlear implant performance.

Neural activity has been shown to track hierarchical linguistic units in connected speech and these responses can be directly modulated by changes in speech intelligibility caused by spectral degradation. In the current study, we manipulate prior knowledge to increase the intelligibility of physically identical speech sentences and test the hypothesis that the tracking responses can be enhanced by this intelligibility improvement. Cortical magnetoencephalography (MEG) responses to intelligible speech followed by either the same (matched) or different (unmatched) unintelligible speech were measured in twenty-three normal hearing participants. Driven by prior knowledge, cortical coherence to “abstract” linguistic units with no accompanying acoustic cues (phrases and sentences) was enhanced relative to the unmatched condition, and was lateralized to the left hemisphere. In contrast, cortical responses coherent to word units, aligned with acoustic onsets, were bilateral and insensitive to contextual information changes. No such coherence changes were observed when prior experience was not available (unintelligible speech before intelligible speech). This dissociation suggests that cerebral responses to linguistic information are directly affected by intelligibility, which in turn are powerfully shaped by physical cues in speech. These results provide an objective and sensitive neural index of speech intelligibility, and explain why previous studies have reported no effect of prior knowledge on cortical entrainment.

Currently, electroencephalogram (EEG) is mostly analyzed in a supervised way, which requires EEG labels (e.g., EEG classification). With the ever-increasing amount of unlabeled/mislabeled EEG in neuropsychiatric disorder diagnosis, BCI, and rehabilitation, manually labeling of EEG data is a labor intensive and time-consuming process, and few labs have developed algorithms to analyze EEG in an unsupervised manner (i.e., EEG clustering). In this paper, we propose a cooperative game inspired approach to cluster multi-trial EEG data. The idea is to map multi-trial EEG clustering to the coalition formation in a cooperative game, and then identify cluster center (the EEG trial with highest Shapley value) and assign EEG trials into proper clusters based on their cross correlation-transformed Shapley values. We demonstrate the mapped EEG cooperative game is convex, and it leads to an algorithm for multi-trial EEG clustering named CoGEEGc. The CoGEEGc yields high-quality multi-trial EEG clustering with respect to intra-cluster compactness and inter-cluster scatter. We show that CoGEEGc outperforms 15 state-of-the-art EEG or time series clustering approaches through detailed experimentation on real-world multi-trial EEG datasets. Comparison against 15 methods with four theoretical properties of clustering further illustrates the superiority of CoGEEGc, as it satisfies two properties while other approaches only satisfy one.

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.

This article explores valid brain electroencephalography (EEG) selection for EEG classification with different classifiers, which has been rarely addressed in previous studies and is mostly ignored by existing EEG processing methods and applications. Importantly, traditional selection methods are not able to select valid EEG signals for different classifiers. This article focuses on a source control-based valid EEG selection to reduce the impact of invalid EEG signals and aims to improve EEG-based classification performance for different classifiers. We propose a novel centroid-based EEG selection approach named CenEEGs, which uses a scale-and-shift-invariance similarity metric to measure similarities of EEG signals and then applies a globally optimal centroid strategy to select valid EEG signals with respect to a similarity threshold. A detailed comparison with several state-of-the-art time series selection methods by using standard criteria on 8 EEG datasets demonstrates the efficacy and superiority of CenEEGs for different classifiers.

© 2019 Qing Wu et al. Coupled matrix and tensor factorizations have been successfully used in many data fusion scenarios where datasets are assumed to be exactly coupled. However, in the real world, not all the datasets share the same factor matrices, which makes joint analysis of multiple heterogeneous sources challenging. For this reason, approximate coupling or partial coupling is widely used in real-world data fusion, with exact coupling as a special case of these techniques. However, to fully address the challenge of tensor factorization, in this paper, we propose two improved coupled tensor factorization methods: one for approximately coupled datasets and the other for partially coupled datasets. A series of experiments using both simulated data and three real-world datasets demonstrate the improved accuracy of these approaches over existing baselines. In particular, when experiments on MRI data is conducted, the performance of our method is improved even by 12.47% in terms of accuracy compared with traditional methods.

The ability to silently hear music in the mind has been argued to be fundamental to musicality. Objective measurements of this subjective imagery experience are needed if this link between imagery ability and musicality is to be investigated. However, previous tests of musical imagery either rely on self-report, rely on melodic memory, or do not cater in range of abilities. The Pitch Imagery Arrow Task (PIAT) was designed to address these shortcomings; however, it is impractically long. In this paper, we shorten the PIAT using adaptive testing and automatic item generation. We interrogate the cognitive processes underlying the PIAT through item response modelling. The result is an efficient online test of auditory mental imagery ability (adaptive Pitch Imagery Arrow Task: aPIAT) that takes 8 min to complete, is adaptive to participant’s individual ability, and so can be used to test participants with a range of musical backgrounds. Performance on the aPIAT showed positive moderate-to-strong correlations with measures of non-musical and musical working memory, self-reported musical training, and general musical sophistication. Ability on the task was best predicted by the ability to maintain and manipulate tones in mental imagery, as well as to resist perceptual biases that can lead to incorrect responses. As such, the aPIAT is the ideal tool in which to investigate the relationship between pitch imagery ability and musicality.

In rodents, hippocampal cell assemblies formed during learning of a navigation task are observed to re-emerge during resting (offline) periods, accompanied by high-frequency oscillations (HFOs). This phenomenon is believed to reflect mechanisms for strengthening newly-formed memory traces. Using magnetoencephalography recordings and a beamforming source location algorithm (synthetic aperture magnetometry), we investigated high-gamma (80–140 Hz) oscillations in the hippocampal region in 18 human participants during inter-trial rest periods in a virtual navigation task. We found right hippocampal gamma oscillations mirrored the pattern of theta power in the same region during navigation, varying as a function of environmental novelty. Gamma power during inter-trial rest periods was positively correlated with theta power during navigation in the first task set when the environment was new and predicted greater performance improvement in the subsequent task set two where the environment became familiar. These findings provide evidence for human hippocampal reactivation accompanied by high-gamma activities immediately after learning and establish a link between hippocampal high-gamma activities and subsequent memory performance.

Highlights • M250 shows amplitude sensitivity to face repetition in adults, but latency sensitivity in children. • Earlier peaks (M100 and M170) show no sensitivity to face repetition in either group. • Face repetition modulates reciprocal connections between OFA and FFA in both groups. • Repetition modulates low-level visual projections to OFA and FFA in adults but only to OFA in children., Electrophysiological studies of adults indicate that brain activity is enhanced during viewing of repeated faces, at a latency of about 250 ms after the onset of the face (M250/N250). The present study aimed to determine if this effect was also present in preschool-aged children, whose brain activity was measured in a custom-sized pediatric MEG system. The results showed that, unlike adults, face repetition did not show any significant modulation of M250 amplitude in children; however children’s M250 latencies were significantly faster for repeated than non-repeated faces. Dynamic causal modelling (DCM) of the M250 in both age groups tested the effects of face repetition within the core face network including the occipital face area (OFA), the fusiform face area (FFA), and the superior temporal sulcus (STS). DCM revealed that repetition of identical faces altered both forward and backward connections in children and adults; however the modulations involved inputs to both FFA and OFA in adults but only to OFA in children. These findings suggest that the amplitude-insensitivity of the immature M250 may be due to a weaker connection between the FFA and lower visual areas.

Articulography and functional neuroimaging are two major tools for studying the neurobiology of speech production. Until now, 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), used for the first time to obtain kinematic profiles of oro-facial movements during speech together with concurrent magnetoencephalographic (MEG) measurements of neuromotor brain activity. MASK was used to characterise speech kinematics in a healthy adult, and the results were compared to measurements from the same participant with a conventional electromagnetic articulography (EMA) setup. We also characterised speech movement kinematics with MASK in a group of ten typically developing children, aged 8-12 years. Analyses targeted the gestural landmarks of the utterances /ida/, /ila/ and reiterated productions of /pataka/. These results demonstrate that the MASK technique can be used to reliably characterise movement profiles and kinematic parameters that reflect development of speech motor control, together with MEG measurements of brain responses from speech sensorimotor cortex. This new capability sets the stage for cross-disciplinary efforts to understand the developmental neurobiology of human speech production.

Providing a plausible neural substrate of speech processing and language comprehension, cortical activity has been shown to track different levels of linguistic structure in connected speech (syllables, phrases and sentences), independent of the physical regularities of the acoustic stimulus. In the current study, we investigated the effect of speech intelligibility on this brain activity as well as the underlying neural sources. Using magnetoencephalography (MEG), brain responses to natural speech and noise-vocoded (spectrally-degraded) speech in nineteen normal hearing participants were measured. Results showed that cortical MEG coherence to linguistic structure changed parametrically with the intelligibility of the speech signal. Cortical responses coherent with phrase and sentence structures were lefthemisphere lateralized, whereas responses coherent to syllable/word structure were bilateral. The enhancement of coherence to intelligible compared to unintelligible speech was also left lateralized and localized to the parasylvian cortex. These results demonstrate that cortical responses to higher level linguistics structures (phrase and sentence level) are sensitive to speech intelligibility. Since the noise-vocoded sentences simulate the auditory input provided by a cochlear implant, such objective neurophysiological measures have potential clinical utility for assessment of cochlear implant performance.

In a previous study, we reported the first measurements of pre-movement and sensorimotor cortex activity in preschool age children (ages 3-5 years) using a customized pediatric magnetoencephalographic system. Movement-related activity in the sensorimotor cortex differed from that typically observed in adults, suggesting that maturation of cortical motor networks was still incomplete by late preschool age. Here we compare these earlier results to a group of school age children (ages 6-8 years) including seven children from the original study measured again two years later, and a group of adults (mean age 31.1 years) performing the same task. Differences in movement-related brain activity were observed both longitudinally within children in which repeated measurements were made, and cross-sectionally between preschool age children, school age children, and adults. Movement-related mu (8-12 Hz) and beta (15-30 Hz) oscillations demonstrated linear increases in amplitude and mean frequency with age. In contrast, movement-evoked gamma synchronization demonstrated a step-like transition from low (30-50 Hz) to high (70-90 Hz) narrow-band oscillations, and this occurred at different ages in different children. Notably, pre-movement activity ('readiness fields') observed in adults was absent in even the oldest children. These are the first direct observations of brain activity accompanying motor responses throughout early childhood, confirming that maturation of this activity is still incomplete by mid-childhood. In addition, individual children demonstrated markedly different developmental trajectories in movement-related brain activity, suggesting that individual differences need to be taken into account when studying motor development across age groups.

Hippocampal rhythms are important for spatial navigation. This study examined whether gender differences in human navigation performance are associated with differences in hippocampal rhythms. We measured brain activities in males and females with whole-head magnetoencephalography (MEG), while they performed a virtual Morris water maze task. Behavioural results showed clear gender differences: males were significantly faster than females; unlike males, females did not show improved navigation performance in a familiar vs. new environment. MEG results showed that the magnitudes of right hippocampal/parahippocampal theta rhythm were similar between the two groups during navigation in a new environment; however, unlike males who exhibited a significant decrease in right hippocampal/parahippocampal theta power in the familiar environment shown before, females showed no change. This result may suggest faster environmental learning in males vs. females. After navigating in the new environment during the inter-trial (ITI) rest periods, males showed significantly higher right hippocampal/parahippocampal high-gamma power than females, suggesting greater consolidation in males. Moreover, right hippocampal/parahippocampal theta power during navigation correlated with navigation performance in both genders; high-gamma power during the ITI was correlated with navigation performance only in males. These associations may provide further support for the functional importance of theta and high-gamma rhythms in navigation. Overall, this study provides new insights into the neurophysiological mechanisms underlying gender differences in spatial navigation.

The cause of stuttering has many theoretical explanations. A number of research groups have suggested changes in the volume and/or function of the striatum as a causal agent. Two recent studies in children and one in adults who stutter (AWS) report differences in striatal volume compared that seen in controls; however, the laterality and nature of this anatomical volume difference is not consistent across studies. The current study investigated whether a reduction in striatal grey matter volume, comparable to that seen in children who stutter (CWS), would be found in AWS. Such a finding would support claims that an anatomical striatal anomaly plays a causal role in stuttering. We used voxel-based morphometry to examine the structure of the striatum in a group of AWS and compared it to that in a group of matched adult control subjects. Results showed a statistically significant group difference for the left caudate nucleus, with smaller mean volume in the group of AWS. The caudate nucleus, one of three main structures within the striatum, is thought to be critical for the planning and modulation of movement sequencing. The difference in striatal volume found here aligns with theoretical accounts of stuttering, which suggest it is a motor control disorder that arises from deficient articulatory movement selection and sequencing. Whilst the current study provides further evidence of a striatal volume difference in stuttering at the group level compared to controls, the significant overlap between AWS and controls suggests this difference is unlikely to be diagnostic of stuttering.

Low frequency theta band oscillations (4–8 Hz) are thought to provide a timing mechanism for hippocampal place cell firing and to mediate the formation of spatial memory. In rodents, hippocampal theta has been shown to play an important role in encoding a new environment during spatial navigation, but a similar functional role of hippocampal theta in humans has not been firmly established. To investigate this question, we recorded healthy participants’ brain responses with a 160‐channel whole‐head MEG system as they performed two training sets of a virtual Morris water maze task. Environment layouts (except for platform locations) of the two sets were kept constant to measure theta activity during spatial learning in new and familiar environments. In line with previous findings, left hippocampal/parahippocampal theta showed more activation navigating to a hidden platform relative to random swimming. Consistent with our hypothesis, right hippocampal/parahippocampal theta was stronger during the first training set compared to the second one. Notably, theta in this region during the first training set correlated with spatial navigation performance across individuals in both training sets. These results strongly argue for the functional importance of right hippocampal theta in initial encoding of configural properties of an environment during spatial navigation. Our findings provide important evidence that right hippocampal/parahippocampal theta activity is associated with environmental encoding in the human brain. Hum Brain Mapp 38:1347–1361, 2017. © 2016 Wiley Periodicals, Inc.

While most normal hearing individuals can readily use prosodic information in spoken language to interpret the moods and feelings of conversational partners, people with congenital amusia report that they often rely more on facial expressions and gestures, a strategy that may compensate for deficits in auditory processing. In this investigation, we used EEG to examine the extent to which individuals with congenital amusia draw upon visual information when making auditory or audio-visual judgments. Event-related potentials (ERP) were elicited by a change in pitch (up or down) between two sequential tones paired with a change in spatial position (up or down) between two visually presented dots. The change in dot position was either congruent or incongruent with the change in pitch. Participants were asked to judge (1) the direction of pitch change while ignoring the visual information (AV implicit task), and (2) whether the auditory and visual changes were congruent (AV explicit task). In the AV implicit task, amusic participants performed significantly worse in the incongruent condition than control participants. ERPs showed an enhanced N2-P3 response to incongruent AV pairings for control participants, but not for amusic participants. However when participants were explicitly directed to detect AV congruency, both groups exhibited enhanced N2-P3 responses to incongruent AV pairings. These findings indicate that amusics are capable of extracting information from both modalities in an AV task, but are biased to rely on visual information when it is available, presumably because they have learned that auditory information is unreliable. We conclude that amusic individuals implicitly draw upon visual information when judging auditory information, even though they have the capacity to explicitly recognize conflicts between these two sensory channels.

Recent evidence shows that hippocampal theta oscillations, usually linked to memory and navigation, are also observed during online language processing, suggesting a shared neurophysiological mechanism between language and memory. However, it remains to be established what specific roles hippocampal theta oscillations may play in language, and whether and how theta mediates the communication between the hippocampus and the perisylvian cortical areas, generally thought to support language processing. With whole-head magnetoencephalographic (MEG) recordings, the present study investigated these questions with two experiments. Using a violation paradigm, extensively used for studying neural underpinnings of different aspects of linguistic processing, we found increased theta power (4–8 ​Hz) in the hippocampal formation, when participants read a semantically incorrect vs. correct sentence ending. Such a pattern of results was replicated using different sentence stimuli in another cohort of participants. Importantly, no significant hippocampal theta power increase was found when participants read a semantically correct but syntactically incorrect sentence ending vs. a correct sentence ending. These findings may suggest that hippocampal theta oscillations are specifically linked to lexical-semantic related processing, and not general information processing in sentence reading. Furthermore, we found significantly transient theta phase coupling between the hippocampus and the left superior temporal gyrus, a hub area of the cortical network for language comprehension. This transient theta phase coupling may provide an important channel that links the memory and language systems for the generation of sentence meaning. Overall, these findings help specify the role of hippocampal theta in language, and provide a novel neurophysiological mechanism at the network level that may support the interface between memory and language.

In a previous MEG study of movement-related brain activity in preschool age children, we reported that pre-movement fields and sensorimotor cortex oscillations differed from those typically observed in adults, suggesting that maturation of cortical motor networks is still incomplete by late preschool age (Cheyne et al., 2014). Here we describe the same measurements in an older group of school-aged children (6 to 8 years old) and an adult control group, in addition to repeated recordings in seven children from the original study approximately two years later. Differences were observed both longitudinally within children and between age groups. Pre-movement (readiness) fields were still not present in the oldest children, however both frequency and magnitude of movement-related mu (8-12Hz) and beta (15-30Hz) oscillations demonstrated linear increases with age. In contrast, movement-evoked gamma synchronization demonstrated a step-like transition from low (30-50 Hz) to high (70-90 Hz) narrow-band oscillations, and this occurred at different ages in different children. These data provide novel evidence of linear and non-linear changes in motor cortex oscillations and delayed development of the readiness field throughout early childhood. Individual children showed large differences in maturation of movement-related brain activity, possibly reflecting differing rates of motor development.

Purpose Hearing loss is associated with changes in brain volume in regions supporting auditory and cognitive processing. The purpose of this study was to determine whether there is a systematic association between hearing ability and brain volume in cross-sectional data from a large nonclinical cohort of middle-aged adults available from the UK Biobank Resource ( http://www.ukbiobank.ac.uk ). Method We performed a set of regression analyses to determine the association between speech reception threshold in noise (SRTn) and global brain volume as well as predefined regions of interest (ROIs) based on T1-weighted structural images, controlling for hearing-related comorbidities and cognition as well as demographic factors. In a 2nd set of analyses, we additionally controlled for hearing aid (HA) use. We predicted statistically significant associations globally and in ROIs including auditory and cognitive processing regions, possibly modulated by HA use. Results Whole-brain gray matter volume was significantly lower for individuals with poorer SRTn. Furthermore, the volume of 9 predicted ROIs including both auditory and cognitive processing regions was lower for individuals with poorer SRTn. The greatest percentage difference (−0.57%) in ROI volume relating to a 1 SD worsening of SRTn was found in the left superior temporal gyrus. HA use did not substantially modulate the pattern of association between brain volume and SRTn. Conclusions In a large middle-aged nonclinical population, poorer hearing ability is associated with lower brain volume globally as well as in cortical and subcortical regions involved in auditory and cognitive processing, but there was no conclusive evidence that this effect is moderated by HA use. This pattern of results supports the notion that poor hearing leads to reduced volume in brain regions recruited during speech understanding under challenging conditions. These findings should be tested in future longitudinal, experimental studies. Supplemental Material https://doi.org/10.23641/asha.7949357

Evidence is accumulating that similar cognitive resources are engaged to process syntactic structure in music and language. Congenital amusia – a neurodevelopmental disorder that primarily affects music perception, including musical syntax – provides a special opportunity to understand the nature of this overlap. Using electroencephalography (EEG), we investigated whether individuals with congenital amusia have parallel deficits in processing language syntax in comparison to control participants. Twelve amusic participants (eight females) and 12 control participants (eight females) were presented melodies in one session, and spoken sentences in another session, both of which had syntactic-congruent and -incongruent stimuli. They were asked to complete a music-related and a language-related task that were irrelevant to the syntactic incongruities. Our results show that amusic participants exhibit impairments in the early stages of both music- and language-syntactic processing. Specifically, we found that two event-related potential (ERP) components – namely Early Right Anterior Negativity (ERAN) and Left Anterior Negativity (LAN), associated with music- and language-syntactic processing respectively, were absent in the amusia group. However, at later processing stages, amusics showed similar brain responses as controls to syntactic incongruities in both music and language. This was reflected in a normal N5 in response to melodies and a normal P600 to spoken sentences. Notably, amusics' parallel music- and language-syntactic impairments were not accompanied by deficits in semantic processing (indexed by normal N400 in response to semantic incongruities). Together, our findings provide further evidence for shared music and language syntactic processing, particularly at early stages of processing., Highlights • Amusics displayed abnormal brain responses to music-syntactic irregularities. • They also exhibited abnormal brain responses to language-syntactic irregularities. • These impairments affect an early stage of syntactic processing not a later stage. • Music and language involve similar cognitive mechanisms for processing syntax.

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.

The current study examined event-related fields (ERFs) evoked by vocal response inhibition in a stimulus-selective stop-signal task. We compared inhibition-related ERFs across a younger and an older group of adults. Behavioural results revealed that stop-signal reaction times (RTs), go-RTs, ignore-stop RTs and failed stop RTs were longer in the older, relative to the younger group by 38, 123, 149 and 116 ms, respectively. The amplitude of the ERF M2 peak (approximately 200 ms after the stop signal) evoked on successful stop trials was larger compared to that evoked on both failed stop and ignore-stop trials. The M4 peak (approximately 450 ms after stop signal) was of larger amplitude in both successful and failed stops compared to ignore-stop trials. In the older group, the M2, M3 and M4 peaks were smaller in amplitude and peaked later in time (by 24, 50 and 76 ms, respectively). We demonstrate that vocal response inhibition-related ERFs exhibit a similar temporal evolution to those previously described for manual response inhibition: an early peak at 200 ms (i.e. M2) that differentiates successful from failed stopping, and a later peak (i.e. M4) that is consistent with a neural marker of response checking and error processing. Across groups, our data support a more general decline of stimulus processing speed with age.

Stopping outright (reactive inhibition) and slowing down (proactive inhibition) are types of response inhibition which have mainly been investigated in the manual effector system. This study compared reactive inhibition across manual and vocal effector systems, examined the effects of excitatory anodal transcranial direct current stimulation (anodal tDCS) over the right prefrontal cortex (right-PFC) and looked at the relationship between reactive and proactive inhibition. We hypothesised (1) that vocal reactive inhibition would be less effective than manual reactive inhibition as evidenced by longer stop signal reaction times; (2) that anodal tDCS would enhance both vocal and manual reactive inhibitions and (3) that proactive and reactive inhibitions would be positively related. We tested 14 participants over two sessions (one session with anodal tDCS and one session with sham stimulation) and applied stimulation protocol in the middle of the session, i.e. only during the second of three phases. We used a stop signal task across two stop conditions: relevant and irrelevant stop conditions in which stopping was required or ignored, respectively. We found that reactive inhibition was faster during and immediately after anodal tDCS relative to sham. We also found that greater level of proactive inhibition enhanced reactive inhibition (indexed by shorter stop signal reaction times). These results support the hypothesis that the right-PFC is part of a core network for reactive inhibition and supports previous contention that proactive inhibition is possibly modulated via preactivating the reactive inhibition network.

In rodents, hippocampal cell assemblies formed during learning of a navigation task are observed to re-emerge during resting (offline) periods, accompanied by high-frequency oscillations (HFOs). This phenomenon is believed to reflect mechanisms for strengthening newly-formed memory traces. Using magnetoencephalography recordings and a beamforming source location algorithm (synthetic aperture magnetometry), we investigated high-gamma (80 – 140 Hz) oscillations in the hippocampal region in 18 human participants during inter-trial rest periods in a virtual navigation task. We found right hippocampal gamma oscillations mirrored the pattern of theta power in the same region during navigation, varying as a function of environmental novelty. Gamma power during inter-trial rest periods was positively correlated with theta power during navigation in the first training set when the environment was new and predicted faster learning in the subsequent training set two where the environment became familiar. These findings provide evidence for human hippocampal reactivation accompanied by high-gamma activities immediately after learning and establish a link between hippocampal high-gamma activities and memory consolidation.

This study measured effective connectivity within the core face network in young children using a paediatric magnetoencephalograph (MEG). Dynamic casual modeling (DCM) of brain responses was performed in a group of adults (N = 14) and a group of young children aged from 3 to 6 years (N = 15). Three candidate DCM models were tested, and the fits of the MEG data to the three models were compared at both individual and group levels. The results show that the connectivity structure of the core face network differs significantly between adults and children. Further, the relative strengths of face network connections were differentially modulated by experimental conditions in the two groups. These results support the interpretation that the core face network undergoes significant structural configuration and functional specialization between four years of age and adulthood.

The auditory processing atypicalities experienced by many individuals on the autism spectrum disorder might be understood in terms of difficulties parsing the sound energy arriving at the ears into discrete auditory 'objects'. Here, we asked whether autistic adults are able to make use of two important spatial cues to auditory object formation - the relative timing and amplitude of sound energy at the left and right ears. Using electroencephalography, we measured the brain responses of 15 autistic adults and 15 age- and verbal-IQ-matched control participants as they listened to dichotic pitch stimuli - white noise stimuli in which interaural timing or amplitude differences applied to a narrow frequency band of noise typically lead to the perception of a pitch sound that is spatially segregated from the noise. Responses were contrasted with those to stimuli in which timing and amplitude cues were removed. Consistent with our previous studies, autistic adults failed to show a significant object-related negativity (ORN) for timing-based pitch, although their ORN was not significantly smaller than that of the control group. Autistic participants did show an ORN to amplitude cues, indicating that they do not experience a general impairment in auditory object formation. However, their P400 response - thought to indicate the later attention-dependent aspects of auditory object formation - was missing. These findings provide further evidence of atypical auditory object processing in autism with potential implications for understanding the perceptual and communication difficulties associated with the condition.

We examined sensorimotor brain activity associated with voluntary movements in preschool children using a customized pediatric magnetoencephalographic system. A videogame-like task was used to generate self-initiated right or left index finger movements in 17 healthy right-handed subjects (8 females, ages 3.2–4.8 years). We successfully identified spatiotemporal patterns of movement-related brain activity in 15/17 children using beamformer source analysis and surrogate MRI spatial normalization. Readiness fields in the contralateral sensorimotor cortex began ∼0.5 s prior to movement onset (motor field, MF), followed by transient movement-evoked fields (MEFs), similar to that observed during self-paced movements in adults, but slightly delayed and with inverted source polarities. We also observed modulation of mu (8–12 Hz) and beta (15–30 Hz) oscillations in sensorimotor cortex with movement, but with different timing and a stronger frequency band coupling compared to that observed in adults. Adult-like high-frequency (70–80 Hz) gamma bursts were detected at movement onset. All children showed activation of the right superior temporal gyrus that was independent of the side of movement, a response that has not been reported in adults. These results provide new insights into the development of movement-related brain function, for an age group in which no previous data exist. The results show that children under 5 years of age have markedly different patterns of movement-related brain activity in comparison to older children and adults, and indicate that significant maturational changes occur in the sensorimotor system between the preschool years and later childhood. Hum Brain Mapp 35:4858–4875, 2014. © 2014 Wiley Periodicals, Inc.

Background Previous magnetoencephalography (MEG) studies have failed to find a facesensitive, brain response-M170 in children. If this is the case, this suggests that the developmental trajectory of the M170 is different from that of its electrical equivalent, the N170. We investigated the alternative possibility that the child M170 may not be detectable in conventional adult-sized MEG systems. New method Brain responses to pictures of faces and well controlled stimuli were measured from the same four-year-old child with a custom child MEG system and an adult-sized MEG system. Results The goodness of fit of the child's head was about the same over the occipital head surface in both systems, but was much worse over all other parts of the head surface in the adult MEG system compared to the child MEG system. The face-sensitive M170 was measured from the child in both MEG systems, but was larger in amplitude, clearer in morphology, and had a more accurate source localization when measured in the child MEG system. Comparison with existing method The custom-sized child MEG system is superior for measuring the face-sensitive M170 brain response in children than the conventional adult MEG system. Conclusions The present results show that the face-sensitive M170 brain response can be elicited in a four-year-old child. This provides new evidence for early maturation of face processing brain mechanisms in humans, and offers new opportunities for the study of neurodevelopmental disorders that show atypical face processing capabilities, such as autism spectrum disorder.

During bimanual load lifting, the brain not only contends with muscle activations in the load-lifting arm, but also has to pre-emptively modulate muscle activations in the load-bearing arm with temporal precision in order to minimize upward arm deflection. Premature or late inhibition in the load-bearing arm activity would result in augmented arm deflection. Little is currently known about the timing operation of motor systems subserving coordinated, bimanual actions. In this study, we measured neuromagnetic brain activity with whole-head magnetoencephalography while 15 participants performed a bimanual load-lifting task. To investigate neural processes prior to load lifting, a beamformer was applied to 6 contiguous 200 ms time epochs spanning the entire premovement phase of the motor task. The sequence of neural activations, following a signal to lift the weight, was chronologically ordered: firstly, the primary motor cortex contralateral to the load-lifting arm was activated, then the cerebellum, and lastly, the basal ganglia, thalamus and primary-/pre- motor areas contralateral to the load-bearing arm. The current data extend our understanding of the neural underpinnings of bimanual coordination. A model is proposed to account for the central organization of volitional and anticipatory motor control in bimanual load lifting.

The ability to imagine objects undergoing rotation (mental rotation) improves markedly with practice, but an explanation of this plasticity remains controversial. Some researchers propose that practice speeds up the rate of a general-purpose rotation algorithm. Others maintain that performance improvements arise through the adoption of a new cognitive strategy-repeated exposure leads to rapid retrieval from memory of the required response to familiar mental rotation stimuli. In two experiments we provide support for an integrated explanation of practice effects in mental rotation by combining behavioral and EEG measures in a way that provides more rigorous inference than is available from either measure alone. Before practice, participants displayed two well-established signatures of mental rotation: Both response time and EEG negativity increased linearly with rotation angle. After extensive practice with a small set of stimuli, both signatures of mental rotation had all but disappeared. In contrast, after the same amount of practice with a much larger set both signatures remained, even though performance improved markedly. Taken together, these results constitute a reversed association, which cannot arise from variation in a single cause, and so they provide compelling evidence for the existence of two routes to expertise in mental rotation. We also found novel evidence that practice with the large but not the small stimulus set increased the magnitude of an early visual evoked potential, suggesting increased rotation speed is enabled by improved efficiency in extracting three-dimensional information from two-dimensional stimuli.

Neuroimaging studies have shown that the neural mechanisms of motor imagery (MI) overlap substantially with the mechanisms of motor execution (ME). Surprisingly, however, the role of several regions of the motor circuitry in MI remains controversial, a variability that may be due to differences in neuroimaging techniques, MI training, instruction types, or tasks used to evoke MI. The objectives of this study were twofold: (i) to design a novel task that reliably invokes MI, provides a reliable behavioral measure of MI performance, and is transferable across imaging modalities; and (ii) to measure the common and differential activations for MI and ME with functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG). We present a task in which it is difficult to give accurate responses without the use of either motor execution or motor imagery. The behavioral results demonstrate that participants performed similarly on the task when they imagined vs. executed movements and this performance did not change over time. The fMRI results show a spatial overlap of MI and ME in a number of motor and premotor areas, sensory cortices, cerebellum, inferior frontal gyrus, and ventrolateral thalamus. MI uniquely engaged bilateral occipital areas, left parahippocampus, and other temporal and frontal areas, whereas ME yielded unique activity in motor and sensory areas, cerebellum, precuneus, and putamen. The MEG results show a robust event-related beta band desynchronization in the proximity of primary motor and premotor cortices during both ME and MI. Together, these results further elucidate the neural circuitry of MI and show that our task robustly and reliably invokes motor imagery, and thus may prove useful for interrogating the functional status of the motor circuitry in patients with motor disorders.

We examined central auditory processing in typically- and atypically-developing readers. Concurrent EEG and MEG brain measurements were obtained from a group of 16 children with dyslexia aged 8–12 years, and a group of 16 age-matched children with normal reading ability. Auditory responses were elicited using 500ms duration broadband noise. These responses were strongly lateralized in control children. Children with dyslexia showed significantly less lateralisation of auditory cortical functioning, and a different pattern of development of auditory lateralization with age. These results provide further evidence that the core neurophysiological deficit of dyslexia is a problem in the balance of auditory function between the two hemispheres.

The consequences of losing the ability to move a limb are traumatic. One approach that examines the impact of pathological limb nonuse on the brain involves temporary immobilization of a healthy limb. Here, we investigated immobilization-induced plasticity in the motor imagery (MI) circuitry during hand immobilization. We assessed these changes with a multimodal paradigm, using functional magnetic resonance imaging (fMRI) to measure neural activation, magnetoencephalography (MEG) to track neuronal oscillatory dynamics, and transcranial magnetic stimulation (TMS) to assess corticospinal excitability. fMRI results show a significant decrease in neural activation for MI of the constrained hand, localized to sensorimotor areas contralateral to the immobilized hand. MEG results show a significant decrease in beta desynchronization and faster resynchronization in sensorimotor areas contralateral to the immobilized hand. TMS results show a significant increase in resting motor threshold in motor cortex contralateral to the constrained hand, suggesting a decrease in corticospinal excitability in the projections to the constrained hand. These results demonstrate a direct and rapid effect of immobilization on MI processes of the constrained hand, suggesting that limb nonuse may not only affect motor execution, as evidenced by previous studies, but also MI. These findings have important implications for the effectiveness of therapeutic approaches that use MI as a rehabilitation tool to ameliorate the negative effects of limb nonuse.

While stuttering is known to be characterized by anomalous brain activations during speech, very little data is available describing brain activations during stuttering. To our knowledge there are no reports describing brain activations that precede blocking. In this case report we present magnetoencephalographic data from a person who stutters who had significant instances of blocking whilst performing a vowel production task. This unique data set has allowed us to compare the brain activations leading up to a block with those leading up to successful production. Surprisingly, the results are very consistent with data comparing fluent production in stutterers to controls. We show here that preceding a block there is significantly less activation of the left orbitofrontal and inferiorfrontal cortices. Furthermore, there is significant extra activation in the right orbitofrontal and inferiorfrontal cortices, and the sensorimotor and auditory areas bilaterally. This data adds weight to the argument forwarded by Kell et al. (2009) that the best functional sign of optimal repair in stutterering is activation of the left BA 47/12 in the orbitofrontal cortex. Educational objectives: At the end of this activity the reader will be able to (a) identify brain regions associated with blocked vocalization, (b) discuss the functions of the orbitofrontal and inferior frontal cortices in regard to speech production and (c) describe the usefulness and limitations of magnetoencephalography (MEG) in stuttering research.

Event-related potentials (ERPs) to tones that are self-initiated are reduced in their magnitude in comparison with ERPs to tones that are externally generated. This phenomenon has been taken as evidence for an efference copy of the motor command acting to suppress the sensory response. However, self-initiation provides a strong temporal cue for the stimulus which might also contribute to the ERP suppression for self-initiated tones. The current experiment sought to investigate the suppression of monaural tones by temporal cueing and also whether the addition of self-initiation enhanced this suppression. Lastly, the experiment sought to investigate the lateralisation of the ERP suppression via presenting these monaural tones to each ear respectively. We examined source waveforms extracted from the lateralised auditory cortices and measured the modulation of the N1 and P2 components by cueing and self-initiation. Self-initiation significantly reduced the amplitude of the N1 component. Temporal cueing without self-initiation significantly reduced the P2 component. There were no significant differences in the amplitude of either the N1 or the P2 between self-initiation and temporal cuing. There was a significant lateralisation effect on the N1—it being significantly larger contralateral to the ear of stimulation. There was no interaction between lateralisation and side of the temporal cue or side of self-initiation suggesting that the effects of self-initiation and temporal cuing are equal bilaterally. We conclude that a significant proportion of ERP suppression by self-initiation is a result of inherent temporal cueing.

This study was designed to test two hypotheses about binaural hearing: (1) that binaural cues are primarily processed in the hemisphere contralateral to the perceived location of a sound; and (2) that the two main binaural cues, interaural timing differences and interaural level differences, are processed in separate channels in the auditory cortex. Magnetoencephalography was used to measure brain responses to dichotic pitches – a perception of pitch created by segregating a narrow band of noise from a wider band of noise – derived from interaural timing or level disparities. Our results show a strong modulation of interhemispheric M100 amplitudes by ITD cues. When these cues simulated source presentation unilaterally from the right hemispace, M100 amplitude changed from a predominant right hemisphere pattern to a bilateral pattern. In contrast, ILD cues lacked any capacity to alter the right hemispheric distribution. These data indicate that intrinsic hemispheric biases are large in comparison to any contralaterality biases in the auditory system. Importantly, both types of binaural cue elicited a circa 200 ms latency object-related negativity component, believed to reflect automatic cortical processes involved in distinguishing concurrent auditory objects. These results support the conclusion that ITDs and ILDs are processed by distinct neuronal populations to relatively late stages of cortical processing indexed by the M100. However information common to the two cues seems to be extracted for use in a subsequent stage of auditory scene segregation indexed by the object related negativity. This may place a new bound on the extent to which sound location cues are processed in separate channels of the auditory cortex.

Objective: Conventional whole-head MEG systems have fixed sensor arrays designed to accommodate most adult heads. However arrays optimised for adult brain measurements are suboptimal for research with the significantly smaller heads of young children. We wished to measure brain activity in children using a novel whole-head MEG system custom sized to fit the heads of pre-school-aged children. Methods: Auditory evoked fields were measured from seven 4-year-old children in a 64-channel KIT whole-head gradiometer MEG system. Results: The fit of heads in the MEG helmet dewars, defined as the mean of sensor-to-head centre distances, were substantially better for children in the child helmet dewar than in the adult helmet dewar, and were similar to head fits obtained for adults in a conventional adult MEG system. Auditory evoked fields were successfully measured from all seven children and dipole source locations were computed. Conclusions: These results demonstrate the feasibility of routinely measuring neuromagnetic brain function in healthy, awake pre-school-aged children. Significance: The advent of child-sized whole-head MEG systems opens new opportunities for the study of cognitive brain development in young children.

We investigated event-related oscillatory responses to acoustic stimulation with EEG and MEG measurements of brain function. EEG and MEG measurements were obtained from healthy adults presented with 500 ms segments of broadband noise presented to both ears via insert earphones. The results showed that auditory stimulation resulted in a reduction in beta band activity within 200 to 400 ms after stimulus onset. The magnitude of this effect was markedly larger when listeners actively attended to sounds than when they ignored them. These results support the conclusion that beta-band oscillations are directly influenced by activation of auditory cortex with acoustic stimulation. However these responses are also strongly modulated by attentional and/or motoric factors related to the auditory task.

In this paper a theoretical framework is proposed for how the brain processes the information necessary for us to achieve the understanding of others that we experience in our social worlds. Our framework attempts to expand several previous approaches to more fully account for the various data on interpersonal understanding and to respond to theoretical critiques in this area. Specifically, we propose that social understanding must be achieved by at least two mechanisms in the brain that are capable of parallel information processing. The first mechanism, based on research into mirror matching systems in the brain, suggests that representations of others are mapped onto an observer's representations of these same schemas in order to understand them. The second mechanism requires semantic analysis of a given social situation in order to understand the actions of others and most likely involves conscious processes. We suggest that experimental correlates of these systems should be dissociable using both beh...

This study measured proactive and reactive response inhibition and their relationships with self-reported impulsivity. We examined the domains of both vocal and manual responding using a stop signal task (SST) with two stop probabilities: high and low probability stop (1/3 and 1/6 stops respectively). Our aim was to evaluate the effect stop probability would have on reactive and proactive inhibition. We tested 44 subjects and found that for the high compared to low probability stop signal condition, more proactive inhibition was evident and this was correlated with a reduction in the stop signal reaction time (SSRT). We found that reactive inhibition had a positive relationship with dysfunctional but not functional impulsivity in both vocal and manual domains of responding. These findings support the hypothesis that proactive inhibition may pre-activate the network for reactive inhibition.

Stuttering is a disorder of speech affecting millions of people around the world. Whilst the exact aetiology of stuttering remains unknown, it has been hypothesised that it is a disorder of the neural mechanisms that support speech timing. In this article, we used magnetoencephalography (MEG) to examine activity from auditory regions of the brain in stuttering and non-stuttering children aged 3-9years. For typically developing children, we found that MEG oscillations in the beta band responded to rhythmic sounds with a peak near the time of stimulus onset. In contrast, stuttering children showed an opposite phase of beta band envelope, with a trough of activity at stimulus onset. These results suggest that stuttering may result from abnormalities in predictive brain responses which are reflected in abnormal entrainment of the beta band envelope to rhythmic sounds.

Background Cortical oscillatory activity in the beta frequency band (13–30 Hz) is associated with voluntary movement and may be altered in motor disorders such as stroke. Methods We used a multimodal case-series approach to investigate movement-related beta oscillations, cortical excitability and upper-limb motor-function in 10 chronic stroke-patients across a broad range of motor-impairment. Assessments included: (i) whole-head magnetoencephalography (MEG) during a voluntary motor task; (ii) resting and active motor-thresholds to transcranial magnetic stimulation (TMS); and (iii) assessments of upper-limb motor-function. Results MEG revealed distinct patterns of movement-related beta oscillations according to motor-function. An elongated event-related desynchronisation (ERD) was correlated with poorer motor-function (p=0.004) whereas an elongated event-related resynchronisation (ERS) was correlated with higher motor function and lower TMS motor thresholds on the more-affected side (p=0.014 and p

Objective This study investigated auditory cortical processing of linguistically-relevant temporal modulations in the developing brains of young children. Methods Auditory envelope following responses to white noise amplitude modulated at rates of 1–80Hz in healthy children (aged 3–5years) and adults were recorded using a paediatric magnetoencephalography (MEG) system and a conventional MEG system, respectively. Results For children, there were envelope following responses to slow modulations but no significant responses to rates higher than about 25Hz, whereas adults showed significant envelope following responses to almost the entire range of stimulus rates. Conclusion Our results show that the auditory cortex of preschool-aged children has a sharply limited capacity to process rapid amplitude modulations in sounds, as compared to the auditory cortex of adults. Significance These neurophysiological results are consistent with previous psychophysical evidence for a protracted maturational time course for auditory temporal processing. The findings are also in good agreement with current linguistic theories that posit a perceptual bias for low frequency temporal information in speech during language acquisition. These insights also have clinical relevance for our understanding of language disorders that are associated with difficulties in processing temporal information in speech.

Musical imagery is a relatively unexplored area, partly because of deficiencies in existing experimental paradigms, which are often difficult, unreliable, or do not provide objective measures of performance. Here we describe a novel protocol, the Pitch Imagery Arrow Task (PIAT), which induces and trains pitch imagery in both musicians and non-musicians. Given a tonal context and an initial pitch sequence, arrows are displayed to elicit a scale-step sequence of imagined pitches, and participants indicate whether the final imagined tone matches an audible probe. It is a staircase design that accommodates individual differences in musical experience and imagery ability. This new protocol was used to investigate the roles that musical expertise, self-reported auditory vividness and mental control play in imagery performance. Performance on the task was significantly better for participants who employed a musical imagery strategy compared to participants who used an alternative cognitive strategy and positively correlated with scores on the Control subscale from the Bucknell Auditory Imagery Scale (BAIS). Multiple regression analysis revealed that Imagery performance accuracy was best predicted by a combination of strategy use and scores on the Vividness subscale of BAIS. These results confirm that competent performance on the PIAT requires active musical imagery and is very difficult to achieve using alternative cognitive strategies. Auditory vividness and mental control were more important than musical experience in the ability to perform manipulation of pitch imagery.

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.

Recent electrophysiological studies have shown that the human electroencephalographic mu rhythm is suppressed during the observation of actions performed by other persons, an effect that may be functionally related to the behaviour of so-called "mirror neurons" observed in area F5 of nonhuman primates. Because mirror neuron activity has been reported to be functionally specific to object-oriented actions, the present study was designed to determine if the human mu rhythm also exhibits this property. EEG measurements were obtained from 12 normal subjects while they observed either a precision grip of a manipulandum or an empty grip using the same hand position. Our results showed that the magnitude of the mu rhythm was significantly lower for the object grip condition than for the empty grip condition. These data support the notion that the human mu rhythm indexes a brain system that is functionally comparable to the monkey mirror neuron system. We propose that nonobject-directed actions may result in representational schemas that are either different or less salient than motorically equivalent actions that are directed toward objects.

The time to decide if a letter is normal or backwards (mirror-reversed) increases as the letter is rotated away from the upright. It is widely accepted that this increase in time reflects the mental rotation of the stimulus to the upright orientation in order to determine the mirror-normal status of the stimulus. Although response times tend to be longer for mirrored stimuli than for normal stimuli, the difference is constant across orientation. Little work has been focused on why mirror-image stimuli produce longer response times than normal stimuli. This study examines the question of whether or not mirrored stimuli are rotated in the picture plane at the same time as normal stimuli, and if so, why response times to mirrored stimuli are longer than that for normal stimuli. Both the behavioural and electrophysiological findings suggest that the mirrored stimuli are not only rotated in the picture plane, but that they are subsequently rotated to the normal view. It is this additional rotation that produces, at least in part, the delayed response times for mirror-image stimuli.