Brain Signatures of Embodied Semantics and Language: A Consensus Paper.

According to embodied theories (including embodied, embedded, extended, enacted, situated, and grounded approaches to cognition), language representation is intrinsically linked to our interactions with the world around us, which is reflected in specific brain signatures during language processing and learning. Moving on from the original rivalry of embodied vs. amodal theories, this consensus paper addresses a series of carefully selected questions that aim at determining when and how rather than whether motor and perceptual processes are involved in language processes. We cover a wide range of research areas, from the neurophysiological signatures of embodied semantics, e.g., event-related potentials and fields as well as neural oscillations, to semantic processing and semantic priming effects on concrete and abstract words, to first and second language learning and, finally, the use of virtual reality for examining embodied semantics. Our common aim is to better understand the role of motor and perceptual processes in language representation as indexed by language comprehension and learning. We come to the consensus that, based on seminal research conducted in the field, future directions now call for enhancing the external validity of findings by acknowledging the multimodality, multidimensionality, flexibility and idiosyncrasy of embodied and situated language and semantic processes.

Reduced syntactic recursion in spontaneous speech of Parkinson's disease patients.

Although characterized as a movement disorder, Parkinson's disease (PD) affects more than just the motor system. Within the heterogenous non-motor symptoms, language impairment is frequent but poorly understood beyond semantic processing. This study investigates the impact of PD on syntactic subordination in spontaneous language production. Fifteen PD patients in ON levodopa status narrated a short story guided by a set of pictures. Thirteen PD patients were also assessed in OFF levodopa status. Narrations were digitally recorded, subsequently transcribed and annotated, making the produced speech accessible to systematical quantitative analysis. Compared to a healthy matched control group, PD patients showed a significant reduction of subordinating structures while the number of non-embedding sentences remained unaffected. No significant effect comparing ON versus OFF levodopa status emerged. Our results suggest a contribution of the basal ganglia to language processing, such as syntactic composition, which, however, does not seem to be dopamine dependent.

Altered performance monitoring in Tourette Syndrome: an MEG investigation.

The error-related negativity (ERN) is an event-related potential component indexing processes of performance monitoring during simple stimulus-response tasks: the ERN is typically enhanced for error processing and conflicting response representations. Investigations in healthy participants and different patient groups have linked the ERN to the dopamine system and to prefrontal information processing. As in patients with Tourette Syndrome (TS) both dopamine release and prefrontal information processing are impaired, we hypothesized that performance monitoring would be altered, which was investigated with magnetencephalography (MEG). We examined performance monitoring in TS patients by assessing the magnetic equivalent of the ERN (mERN). The mERN was investigated in tic-free trials of eight adult, unmedicated TS patients without clinically significant comorbidity and ten matched healthy controls while performing a Go/NoGo task in selected frontocentral channels. The analysis of the response-related amplitudes of the event-related magnetic field showed that TS patients, in contrast to controls, did not show earlier amplitude modulation (between 70 and 105 ms after response onset) depending on response type (errors or correct responses). In both groups significant mERN amplitudes in the time-window between 105 and 160 ms after response onset were detected thus pointing at only later error processing in TS patients. In TS patients, early error-related processing might be affected by an enhanced motor control triggered by a conflict between the targeted high task performance and tic suppression. TS patients seem to tend to initially process all responses as erroneous responses.

Induced and Evoked Brain Activation Related to the Processing of Onomatopoetic Verbs.

Grounded cognition theory postulates that cognitive processes related to motor or sensory content are processed by brain networks involved in motor execution and perception, respectively. Processing words with auditory features was shown to activate the auditory cortex. Our study aimed at determining whether onomatopoetic verbs (e.g., "tröpfeln"-to dripple), whose articulation reproduces the sound of respective actions, engage the auditory cortex more than non-onomatopoetic verbs. Alpha and beta brain frequencies as well as evoked-related fields (ERFs) were targeted as potential neurophysiological correlates of this linguistic auditory quality. Twenty participants were measured with magnetoencephalography (MEG) while semantically processing visually presented onomatopoetic and non-onomatopoetic German verbs. While a descriptively stronger left temporal alpha desynchronization for onomatopoetic verbs did not reach statistical significance, a larger ERF for onomatopoetic verbs emerged at about 240 ms in the centro-parietal area. Findings suggest increased cortical activation related to onomatopoeias in linguistically relevant areas.

Neurophysiological mechanisms of perspective-taking: An MEG investigation of agency.

According to the embodied cognition framework, sensory and motor areas are recruited during language understanding through simulation processes. Behavioral and imaging findings point to a dependence of the latter on perspective-taking (e.g., first person "I" versus third person "s/he"). The current study aims at identifying possible neurophysiological correlates of perspective in a linguistic context. Twenty healthy participants were measured with magnetoencephalography (MEG) while semantically processing visually presented inflected German verbs in the first- and third-person perspective, simple present tense. Results show that the first-person perspective induces stronger beta (15-25 Hz) desynchronization in the right-hemispheric posterior superior temporal sulcus, ventral posterior cingulate gyrus, and V5/MT+ area; no modulation of sensorimotor cortex emerged. Moreover, a stronger event-related field (ERF) was observed for the first-person perspective at about 150 ms after pronoun-verb onset, originating in occipital and moving to central and left temporal cortical sites. No effect of perspective on sensory gating was found when targeting the N1 component related to tones following the linguistic stimuli. Results indicate an effect of linguistic perspective-taking on brain activation patterns. The contribution of the single brain areas and their role in self-other distinction is further discussed.

Auditory cortex sensitivity to the loudness attribute of verbs.

The auditory cortex was shown to be activated during the processing of words describing actions with acoustic features. The present study further examines whether processing visually presented action words characterized by different levels of loudness, i.e. "loud" (to shout) and "quiet" actions (to whisper), differentially engage the auditory cortex. Twenty healthy participants were measured with magnetoencephalography (MEG) while reading inflected verbs followed by a short tone and semantic tasks. Based on the results of a localizer task, loudness sensitive temporal Brodmann areas A22, A41/42, and pSTS were inspected in the word paradigm. "Loud" actions induced significantly stronger beta power suppression compared to "quiet" actions in the left hemisphere. Smaller N100m amplitude related to tones following "loud" compared to "quiet" actions confirmed that auditory cortex sensitivity was modulated by action words. Results point to possible selective auditory simulation mechanisms involved in verb processing and support embodiment theories.

Action verb processing specifically modulates motor behaviour and sensorimotor neuronal oscillations.

Understanding action-related language recruits the brain's motor system and can interact with motor behaviour. The current study shows MEG oscillatory patterns during verb-motor priming. Hand and foot verbs were followed by hand or foot responses, with faster reaction times for congruent conditions. In ROIs placed in the hand/arm and foot/leg portions of the sensorimotor cortex, this behavioural priming effect was accompanied by modulations in MEG oscillatory patterns preceding the responses. Power suppression in the alpha/beta frequency bands was reduced in congruent conditions in the body-part-specific ROIs. These results imply that the verb-motor priming effect may be a direct consequence of motor cortex contributions to action word processing.

A peek into premonitory urges in Tourette syndrome: Temporal evolution of neurophysiological oscillatory signatures.

BACKGROUND: Tics are the core symptom of patients with Gilles de la Tourette syndrome, yet the spatial-temporal dynamics of neural activity causing a tic remains to be determined. OBJECTIVE: Identification of cortical events preceding tic onset. METHODS: In twelve patients with Tourette syndrome we performed magnetoencephalography to trace the time course of beta oscillations (15-30 Hz) in motor cortical areas before tic onset. RESULTS: Patients showed a biphasic modulation of cortical beta activity during the second before tic onset. We observed an initial increase of beta power over the left-hemispheric channels overlying the motor cortex. This increase was subsequently replaced by a decrease in beta power. The beta decrease close to tic onset resembled the typical pattern accompanying preparation of voluntary movements. Only the initial increase in beta power positively correlated with the intensity of motor urges preceding tics. CONCLUSIONS: The spatial-temporal dynamics of cortical activity suggests a voluntary component of tics that might be triggered by a failure of compensatory motor inhibitory mechanisms.

Semantic discrimination impacts tDCS modulation of verb processing.

Motor cortex activation observed during body-related verb processing hints at simulation accompanying linguistic understanding. By exploiting the up- and down-regulation that anodal and cathodal transcranial direct current stimulation (tDCS) exert on motor cortical excitability, we aimed at further characterizing the functional contribution of the motor system to linguistic processing. In a double-blind sham-controlled within-subjects design, online stimulation was applied to the left hemispheric hand-related motor cortex of 20 healthy subjects. A dual, double-dissociation task required participants to semantically discriminate concrete (hand/foot) from abstract verb primes as well as to respond with the hand or with the foot to verb-unrelated geometric targets. Analyses were conducted with linear mixed models. Semantic priming was confirmed by faster and more accurate reactions when the response effector was congruent with the verb's body part. Cathodal stimulation induced faster responses for hand verb primes thus indicating a somatotopical distribution of cortical activation as induced by body-related verbs. Importantly, this effect depended on performance in semantic discrimination. The current results point to verb processing being selectively modifiable by neuromodulation and at the same time to a dependence of tDCS effects on enhanced simulation. We discuss putative mechanisms operating in this reciprocal dependence of neuromodulation and motor resonance.

Body-part specific interactions of action verb processing with motor behaviour.

The interaction of action-related language processing with actual movement is an indicator of the functional role of motor cortical involvement in language understanding. This paper describes two experiments using single action verb stimuli. Motor responses were performed with the hand or the foot. To test the double dissociation of language-motor facilitation effects within subjects, Experiments 1 and 2 used a priming procedure where both hand and foot reactions had to be performed in response to different geometrical shapes, which were preceded by action verbs. In Experiment 1, the semantics of the verbs could be ignored whereas Experiment 2 included semantic decisions. Only Experiment 2 revealed a clear double dissociation in reaction times: reactions were facilitated when preceded by verbs describing actions with the matching effector. In Experiment 1, by contrast, there was an interaction between verb-response congruence and a semantic variable related to motor features of the verbs. Thus, the double dissociation paradigm of semantic motor priming was effective, corroborating the role of the motor system in action-related language processing. Importantly, this effect was body part specific.

Correction: Grasping Hand Verbs: Oscillatory Beta and Alpha Correlates of Action-Word Processing.

[This corrects the article DOI: 10.1371/journal.pone.0108059.].

Increased beta rhythm as an indicator of inhibitory mechanisms in tourette syndrome.

BACKGROUND: Inhibitory oscillatory mechanisms subserving tic compensation have been put forward in Tourette syndrome. Modulation of the beta rhythm (15-25 Hz) as the well-established oscillatory movement execution-inhibition indicator was tested during a cognitive-motor task in patients with Tourette syndrome. METHODS: Performing a Go/NoGo task, 12 patients with Tourette syndrome and 12 matched controls were recorded using whole-head magnetoencephalography. RESULTS: Compared to healthy participants, patients showed less beta suppression in the sensorimotor area and enhanced beta power in parieto-occipital brain regions contralaterally to the response hand. Average beta power and power gain correlated negatively with tic severity. CONCLUSIONS: Increased motor inhibitory as well as visuomotor attentional processes are likely to subserve tic compensation. Correlational results suggest that stronger inhibitory compensation accompanies less tic severity.

Language-motor interference reflected in MEG beta oscillations.

The involvement of the brain's motor system in action-related language processing can lead to overt interference with simultaneous action execution. The aim of the current study was to find evidence for this behavioural interference effect and to investigate its neurophysiological correlates using oscillatory MEG analysis. Subjects performed a semantic decision task on single action verbs, describing actions executed with the hands or the feet, and abstract verbs. Right hand button press responses were given for concrete verbs only. Therefore, longer response latencies for hand compared to foot verbs should reflect interference. We found interference effects to depend on verb imageability: overall response latencies for hand verbs did not differ significantly from foot verbs. However, imageability interacted with effector: while response latencies to hand and foot verbs with low imageability were equally fast, those for highly imageable hand verbs were longer than for highly imageable foot verbs. The difference is reflected in motor-related MEG beta band power suppression, which was weaker for highly imageable hand verbs compared with highly imageable foot verbs. This provides a putative neuronal mechanism for language-motor interference where the involvement of cortical hand motor areas in hand verb processing interacts with the typical beta suppression seen before movements. We found that the facilitatory effect of higher imageability on action verb processing time is perturbed when verb and motor response relate to the same body part. Importantly, this effect is accompanied by neurophysiological effects in beta band oscillations. The attenuated power suppression around the time of movement, reflecting decreased cortical excitability, seems to result from motor simulation during action-related language processing. This is in line with embodied cognition theories.

Grasping hand verbs: oscillatory beta and alpha correlates of action-word processing.

The grounded cognition framework proposes that sensorimotor brain areas, which are typically involved in perception and action, also play a role in linguistic processing. We assessed oscillatory modulation during visual presentation of single verbs and localized cortical motor regions by means of isometric contraction of hand and foot muscles. Analogously to oscillatory activation patterns accompanying voluntary movements, we expected a somatotopically distributed suppression of beta and alpha frequencies in the motor cortex during processing of body-related action verbs. Magnetoencephalographic data were collected during presentation of verbs that express actions performed using the hands (H) or feet (F). Verbs denoting no bodily movement (N) were used as a control. Between 150 and 500 msec after visual word onset, beta rhythms were suppressed in H and F in comparison with N in the left hemisphere. Similarly, alpha oscillations showed left-lateralized power suppression in the H-N contrast, although at a later stage. The cortical oscillatory activity that typically occurs during voluntary movements is therefore found to somatotopically accompany the processing of body-related verbs. The combination of a localizer task with the oscillatory investigation applied to verb reading as in the present study provides further methodological possibilities of tracking language processing in the brain.

Neuromagnetic hand and foot motor sources recruited during action verb processing.

The current study investigated sensorimotor involvement in the processing of verbs describing actions performed with the hands, feet, or no body part. Actual movements were used to identify neuromagnetic sources for hand and foot actions. These sources constrained the analysis of verb processing. While hand and foot sources picked up activation in all three verb conditions, peak amplitudes showed an interaction of source and verb condition at 200 ms after word onset, thereby reflecting effector-specificity. Specifically, hand verbs elicited significantly higher peak amplitudes than foot verbs in hand sources. Our results are in line with theories of embodied cognition that assume an involvement of sensorimotor areas in early stages of lexico-semantic processing, even for single words without a semantic or motor task.

Distinct neural processes in grapheme-colour synaesthetes and semantic controls.

In synaesthetes, stimulation of one sensory pathway provokes a sensory experience (e.g. a colour concurrent) in a different sensory modality or sub-modality. Results of synaesthetic Stroop and priming tests indicate that the perception of a colour concurrent interferes with the processing of a veridical colour in synaesthetes. We here examined the congruency between a stimulus' colour and the colour concurrent both in grapheme-colour synaesthetes and in non-synaesthetes trained on grapheme-colour associations. Electrophysiological (electroencephalogram) and behavioural measurements were collected during a priming task that included grapheme-grapheme and grapheme-colour patch pairs. To investigate covert bidirectional synaesthesia, an additional inverted colour patch-grapheme condition was included. Both groups of participants showed longer reaction time and more negative-going N300 and N400 event-related potential (ERP) components on incongruent trials. Whereas ERP effects in the non-synaesthetes were largely confined to the late cognitive components N300, P300 and N400, the synaesthetes also showed congruency-dependent modulation of the early sensory component N170. Our results suggest that early cognitive processes distinguish cross-modal synaesthetic perceptions from acquired associations. The involvement of both early- and late-stage cognitive components in bidirectional synaesthesia possibly indicates similar feature-binding mechanisms during processing of opposite flow directions of information, namely grapheme-colour and colour-grapheme.

Synaesthetic perception of colour and visual space in a blind subject: an fMRI case study.

In spatial sequence synaesthesia (SSS) ordinal stimuli are perceived as arranged in peripersonal space. Using fMRI, we examined the neural bases of SSS and colour synaesthesia for spoken words in a late-blind synaesthete, JF. He reported days of the week and months of the year as both coloured and spatially ordered in peripersonal space; parts of the days and festivities of the year were spatially ordered but uncoloured. Words that denote time-units and triggered no concurrents were used in a control condition. Both conditions inducing SSS activated the occipito-parietal, infero-frontal and insular cortex. The colour area hOC4v was engaged when the synaesthetic experience included colour. These results confirm the continued recruitment of visual colour cortex in this late-blind synaesthetes. Synaesthesia also involved activation in inferior frontal cortex, which may be related to spatial memory and detection, and in the insula, which might contribute to audiovisual integration related to the processing of inducers and concurrents.

Modality and variability of synesthetic experience.

In synesthesia, stimulation of one sensory or cognitive pathway leads to additional, involuntary experiences in a second sensory or cognitive pathway. We here review previous surveys on this neurologically based phenomenon and report the results of 63 synesthetes who completed our Internet and paper questionnaire on synesthesia. In addition to asking for personal data and information on the participant's synesthesia, the questionnaire focused on the components of the inducer that elicit or modulate synesthesia. Synesthesia was most often developmental (92%) and of the grapheme-color type (86%). Sixty-two percent of the participants perceived time-related words in a spatial configuration. Music-color synesthesia was common (41%), and synesthesia for natural and artificial sounds (33%) was higher than in previous estimates. Eighty-one percent of participants experienced more than one form of synesthesia. Multimodal synesthesia, in which inducer and concurrent belong to 2 different sensory modalities, occurred in 92% of the participants. Overall, auditory stimuli were most often reported as inducers, and visual concurrents were most common. Modulations of the synesthetic experiences such as changes of the concurrent color, expansion within the same or to a different sensory modality, or reduction of the number of inducers over time were reported by 17% of participants. This challenges the presumed consistency of synesthesia and the adequacy of the test-retest consistency score still most commonly used to assess the veracity of reported synesthesia. Implications of the high prevalence of cross-modal synesthesia and the variability of synesthesia are discussed.

Objective and subjective measures in recovery from a 35% carbon dioxide challenge.

OBJECTIVES: Because hyperventilation, dyspnea, and a feeling of choking are often core features of a panic attack, respiration has been one of the most widely studied physiological parameters in panic disorder (PD) patients. A respiratory subgroup of PD, with distinct etiological pathways, has also been suggested. Investigation of the recovery phase following a respiratory challenge may be a reliable way to establish respiratory impairment in PD patients. The objective of the present study was to investigate the recovery phase from a 35% carbon dioxide challenge in PD patients and in healthy controls, and to test the hypothesis of a different respiratory pattern in patients, compared to control subjects. METHODS: Eleven nonmedicated PD patients with or without agoraphobia, 11 medicated PD patients, and 11 control subjects took part in a 35% carbon dioxide and 65% oxygen inhalation challenge. Respiratory rate, partial pressure of carbon dioxide, heart rate, and blood pressure were recorded during the baseline phase (10 minutes) and the recovery phase (10 minutes). Visual Analogue Scale of Anxiety and Panic Symptom List scores were collected pre- and post-challenge. RESULTS: Nonmedicated patients had increased variability in respiratory rate and partial pressure of carbon dioxide during recovery, compared with control subjects and medicated PD patients. Also, PD patients tended to have higher heart rates and to need more time to recover from the challenge than control subjects. CONCLUSIONS: Results suggest that PD patients have less effective homeostatic control after their physiological equilibrium has been disrupted by a respiratory stressor.

Cognitive-behavioral group treatment for panic disorder with agoraphobia.

Cognitive-behavioral therapy (CBT) is well documented in the treatment of panic disorder with or without agoraphobia; however, little is known about the efficacy of group treatment. The purpose of this open study is to investigate the benefits of a combination of the major cognitive and behavioral techniques used in the several specific versions of CBT thus far developed, in a psychotherapeutic group approach for panic and agoraphobia. Seventy-six outpatients meeting the Diagnostic and Statistical Manual of Mental Disorders, third edition, revised (DSM-III-R; American Psychiatric Association, 1987) criteria for panic disorder with or without agoraphobia were included in the study. The treatment consisted of 14 weekly 2-hr group sessions and included: (a) an educational component, (b) interoceptive exposure, (c) cognitive restructuring, (d) problem solving, and (e) in vivo exposure. Patients achieved significant treatment gains on all dimensions assessed with a high rate of panic remission and significant improvement in the associated symptoms. Furthermore, these gains were maintained at 6-months' follow-up. Our results suggest the feasibility of this combination of cognitive and behavioral techniques. The findings raise questions about the specificity and the impact of each technique.