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Abstract Understanding language semantically related to actions activates the motor cortex. This activation is sensitive to semantic information such as the body part used to perform the action (e.g. arm-/leg-related action words). Additionally, motor movements of the hands/feet can have a causal effect on memory maintenance of action words, suggesting that the involvement of motor systems extends to working memory. This study examined brain correlates of verbal memory load for action-related words using event-related fMRI. Seventeen participants saw either four identical or four different words from the same category (arm-/leg-related action words) then performed a nonmatching-to-sample task. Results show that verbal memory maintenance in the high-load condition produced greater activation in left premotor and supplementary motor cortex, along with posterior-parietal areas, indicating that verbal memory circuits for action-related words include the cortical action system. Somatotopic memory load effects of arm- and leg-related words were observed, but only at more anterior cortical regions than was found in earlier studies employing passive reading tasks. These findings support a neurocomputational model of distributed action-perception circuits (APCs), according to which language understanding is manifest as full ignition of APCs, whereas working memory is realized as reverberant activity receding to multimodal prefrontal and lateral temporal areas.

Abstract Background Impaired naming is a ubiquitous symptom in all types of aphasia, which often adversely impacts independence, quality of life, and recovery of affected individuals. Previous research has demonstrated that naming can be facilitated by phonological and semantic cueing strategies that are largely incorporated into the treatment of anomic disturbances. Beneficial effects of cueing, whereby naming becomes faster and more accurate, are often attributed to the priming mechanisms occurring within the distributed language network. Objective We proposed and explored two novel cueing techniques: (1) Silent Visuomotor Cues (SVC), which provided articulatory information of target words presented in the form of silent videos, and (2) Semantic Auditory Cues (SAC), which consisted of acoustic information semantically relevant to target words (ringing for “telephone”). Grounded in neurophysiological evidence, we hypothesized that both SVC and SAC might aid communicative effectiveness possibly by triggering activity in perceptual and semantic language regions, respectively. Methods Ten participants with chronic non-fluent aphasia were recruited for a longitudinal clinical intervention. Participants were split into dyads (i.e., five pairs of two participants) and required to engage in a turn-based peer-to-peer language game using the Rehabilitation Gaming System for aphasia (RGSa). The objective of the RGSa sessions was to practice communicative acts, such as making a request. We administered SVCs and SACs in a pseudorandomized manner at the moment when the active player selected the object to be requested from the interlocutor. For the analysis, we compared the times from selection to the reception of the desired object between cued and non-cued trials. Results Naming accuracy, as measured by a standard clinical scale, significantly improved for all stimuli at each evaluation point, including the follow-up. Moreover, the results yielded beneficial effects of both SVC and SAC cues on word naming, especially at the early intervention sessions when the exposure to the target lexicon was infrequent. Conclusions This study supports the efficacy of the proposed cueing strategies which could be integrated into the clinic or mobile technology to aid naming even at the chronic stages of aphasia. These findings are consistent with sensorimotor accounts of language processing, suggesting a coupling between language, motor, and semantic brain regions. Trial registration NCT02928822 . Registered 30 May 2016.

This study seeks to confirm whether lesions in posterior regions of the brain involved in visuo-spatial processing are of functional relevance to the processing of words with spatial meaning. We investigated whether patients with Posterior Cortical Atrophy (PCA), an atypical form of Alzheimer’s Disease which predominantly affects parieto-occipital brain regions, is associated with deficits in working memory for spatial prepositions. Case series of patients with PCA and matched healthy controls performed tests of immediate and delayed serial recall on words from three lexico-semantic word categories: number words (twelve), spatial prepositions (behind) and function words (e.g., shall). The three word categories were closely matched for a number of psycholinguistic and semantic variables including length, bi-/tri-gram frequency, word frequency, valence and arousal. Relative to controls, memory performance of PCA patients on short word lists was significantly impaired on spatial prepositions in the delayed serial recall task. These results suggest that lesions in posterior parieto-occipital regions specifically impair the processing of spatial prepositions. Our findings point to a pertinent role of posterior cortical regions in the semantic processing of words with spatial meaning and provide strong support for modality-specific semantic theories that recognize the necessary contributions of sensorimotor regions to conceptual semantic processing.

Concrete symbols (e.g., "sun," "run") can be learned in the context of objects and actions, thereby grounding their meaning in the world. However, it is controversial whether a comparable avenue to semantic learning exists for abstract symbols (e.g., "democracy"). When we simulated the putative brain mechanisms of conceptual/semantic grounding using brain-constrained deep neural networks, the learning of instances of concrete concepts outside of language contexts led to robust neural circuits generating substantial and prolonged activations. In contrast, the learning of instances of abstract concepts yielded much reduced and only short-lived activity. Crucially, when conceptual instances were learned in the context of wordforms, circuit activations became robust and long-lasting for both concrete and abstract meanings. These results indicate that, although the neural correlates of concrete conceptual representations can be built from grounding experiences alone, abstract concept formation at the neurobiological level is enabled by and requires the correlated presence of linguistic forms.

Embodied theories of grounded semantics postulate that, when word meaning is first acquired, a link is established between symbol (word form) and corresponding semantic information present in modality-specific—including primary—sensorimotor cortices of the brain. Direct experimental evidence documenting the emergence of such a link (i.e., showing that presentation of a previously unknown, meaningless word sound induces, after learning, category-specific reactivation of relevant primary sensory or motor brain areas), however, is still missing. Here, we present new neuroimaging results that provide such evidence. We taught participants aspects of the referential meaning of previously unknown, senseless novel spoken words (such as “Shruba” or “Flipe”) by associating them with either a familiar action or a familiar object. After training, we used functional magnetic resonance imaging to analyze the participants’ brain responses to the new speech items. We found that hearing the newly learnt object-related word sounds selectively triggered activity in the primary visual cortex, as well as secondary and higher visual areas.These results for the first time directly document the formation of a link between the novel, previously meaningless spoken items and corresponding semantic information in primary sensory areas in a category-specific manner, providing experimental support for perceptual accounts of word-meaning acquisition in the brain.

A class of semantic theories defines concepts in terms of statistical distributions of lexical items, basing meaning on vectors of word co-occurrence frequencies. A different approach emphasizes abstract hierarchical taxonomic relationships among concepts. However, the functional relevance of these different accounts and how they capture information-encoding of lexical meaning in the brain still remains elusive.We investigated to what extent distributional and taxonomic models explained word-elicited neural responses using cross-validated representational similarity analysis (RSA) of functional magnetic resonance imaging (fMRI) and model comparisons.Our findings show that the brain encodes both types of semantic information, but in distinct cortical regions. Posterior middle temporal regions reflected lexical-semantic similarity based on hierarchical taxonomies, in coherence with the action-relatedness of specific semantic word categories. In contrast, distributional semantics best predicted the representational patterns in left inferior frontal gyrus (LIFG, BA 47). Both representations coexisted in the angular gyrus supporting semantic binding and integration. These results reveal that neuronal networks with distinct cortical distributions across higher-order association cortex encode different representational properties of word meanings. Taxonomy may shape long-term lexical-semantic representations in memory consistently with the sensorimotor details of semantic categories, whilst distributional knowledge in the LIFG (BA 47) may enable semantic combinatorics in the context of language use.Our approach helps to elucidate the nature of semantic representations essential for understanding human language.

Repetitive TMS (rTMS) allows for non-invasive and transient disruption of local neuronal functioning. We used machine learning approaches to assess whether brain tumor patients can be accurately classified into aphasic and non-aphasic groups using their rTMS language mapping results as input features. Given that each tumor affects the subject-specific language networks differently, resulting in heterogenous rTMS functional mappings, we propose the use of machine learning strategies to classify potential patterns of rTMS language mapping results. We retrospectively included 90 patients with left perisylvian world health organization (WHO) grade II-IV gliomas that underwent presurgical navigated rTMS language mapping. Within our cohort, 29 of 90 (32.2%) patients suffered from at least mild aphasia as shown in the Aachen Aphasia Test based Berlin Aphasia Score (BAS). After spatial normalization to MNI 152 of all rTMS spots, we calculated the error rate (ER) in each stimulated cortical area (28 regions of interest, ROI) by automated anatomical labeling parcellation (AAL3) and IIT. We used a support vector machine (SVM) to classify significant areas in relation to aphasia. After feeding the ROIs into the SVM model, it revealed that in addition to age (w = 2.98), the ERs of the left supramarginal gyrus (w = 3.64), left inferior parietal gyrus (w = 2.28) and right pars triangularis (w = 1.34) contributed more than other features to the model. The model’s sensitivity was 86.2%, the specificity was 82.0%, the overall accuracy was 85.5% and the AUC was 89.3%. Our results demonstrate an increased vulnerability of right inferior pars triangularis to rTMS in aphasic patients due to left perisylvian gliomas. This finding points towards a functional relevant involvement of the right pars triangularis in response to aphasia. The tumor location feature, specified by calculating overlaps with white and grey matter atlases, did not affect the SVM model. The left supramarginal gyrus as a feature improved our SVM model the most. Additionally, our results could point towards a decreasing potential for neuroplasticity with age.

The neurophysiological mechanisms underlying motor and language difficulties in autism spectrum disorders (ASD) are still largely unclear. The present work investigates biological indicators of sound processing, (action-) semantic understanding and predictive coding and their correlation with clinical symptoms of ASD. Twenty-two adults with high-functioning ASD and 25 typically developed (TD) participants engaged in an auditory, passive listening, Mismatch Negativity (MMN) task while high-density electroencephalography (EEG) was recorded. Action and non-action words were presented in the context of sounds, which were either semantically congruent with regard to the body part they relate to or semantically incongruent or unrelated. The anticipatory activity before sound onset, the Prediction Potential (PP), was significantly reduced in the ASD group specifically for action, but not for non-action sounds. The early-MMN-like responses to words (latency: 120 ms) were differentially modulated across groups: controls showed larger amplitudes for words in action-sound compared to non-action contexts, whereas ASD participants demonstrated enlarged early-MMN-like responses only in a pure tone context, with no other modulation dependent on action sound context. Late-MMN-like responses around 560 ms post-stimulus onset revealed body-part-congruent action-semantic priming for words in control participants, but not in the ASD group. Importantly, neurophysiological indices of semantic priming in ASD participants correlated with the extent of autistic traits as revealed by the Autism Spectrum Quotient (AQ). The data suggest that high-functioning adults with ASD show a specific deficit in semantic processing and predictive coding of sounds and words related to action, which is absent for neutral, non-action, sounds.

Neuromodulation technologies, such as transcranial magnetic stimulation (TMS), are promising tools for neurorehabilitation, aphasia therapy included, but not yet in common clinical use. Combined with behavioral techniques, in particular treatment-efficient Intensive Language-Action Therapy (ILAT, previously CIAT or CILT), TMS could substantially amplify the beneficial effect of such behavioral therapy alone (Thiel et al., 2013; Martin et al., 2014; Mendoza et al., 2016; Kapoor, 2017). In this randomized study of 17 subjects with post-stroke aphasia in the chronic stage, we studied the combined effect of ILAT and 1-Hz placebo-controlled navigated repetitive TMS (rTMS) to the right-hemispheric inferior frontal cortex—that is, to the anterior part of the non-dominant hemisphere's homolog Broca's area (pars triangularis). Patients were randomized to groups A and B. Patients in group A received a 2-week period of rTMS during naming training where they named pictures displayed on the screen once every 10 s, followed by 2 weeks of rTMS and naming combined with ILAT. Patients in group B received the same behavioral therapy but TMS was replaced by sham stimulation. The primary outcome measures for changes in language performance were the Western Aphasia Battery's aphasia quotient AQ; the secondary outcome measures were the Boston naming test (BNT) and the Action naming test (Action BNT, ANT). All subjects completed the study. At baseline, no statistically significant group differences were discovered for age, post-stroke time or diagnosis. ILAT was associated with significant improvement across groups, as documented by both primary and secondary outcome measures. No significant effect of rTMS could be documented. Our results agree with previous results proving ILAT's ability to improve language in patients with chronic aphasia. In contrast with earlier claims, however, a beneficial effect of rTMS in chronic post-stroke aphasia rehabilitation was not detected in this study.Clinical Trial Registration:www.ClinicalTrials.gov, identifier: NCT03629665

One of the most controversial debates in cognitive neuroscience concerns the cortical locus of semantic knowledge and processing in the human brain. Experimental data revealed the existence of various cortical regions relevant for meaning processing, ranging from semantic hubs generally involved in semantic processing to modality-preferential sensorimotor areas involved in the processing of specific conceptual categories. Why and how the brain uses such complex organization for conceptualization can be investigated using biologically constrained neurocomputational models. Here, we improve pre-existing neurocomputational models of semantics by incorporating spiking neurons and a rich connectivity structure between the model ‘areas’ to mimic important features of the underlying neural substrate. Semantic learning and symbol grounding in action and perception were simulated by associative learning between co-activated neuron populations in frontal, temporal and occipital areas. As a result of Hebbian learning of the correlation structure of symbol, perception and action information, distributed cell assembly circuits emerged across various cortices of the network. These semantic circuits showed category-specific topographical distributions, reaching into motor and visual areas for action- and visually-related words, respectively. All types of semantic circuits included large numbers of neurons in multimodal connector hub areas, which is explained by cortical connectivity structure and the resultant convergence of phonological and semantic information on these zones. Importantly, these semantic hub areas exhibited some category-specificity, which was less pronounced than that observed in primary and secondary modality-preferential cortices. The present neurocomputational model integrates seemingly divergent experimental results about conceptualization and explains both semantic hubs and category-specific areas as an emergent process causally determined by two major factors: neuroanatomical connectivity structure and correlated neuronal activation during language learning.

Recent neuropsychological studies in neurological patients and healthy subjects suggest a close functional relationship between the brain systems for language and action. Facilitation and inhibition effects of motor system activity on language processing have been demonstrated as well as causal effects in the reverse direction, from language processes on motor excitability or performance. However, as the documented effects between motor and language systems were sometimes facilitatory and sometimes inhibitory, the “sign” of these effects still remains to be explained. In a previous study, we reported a word-category-specific differential impairment of verbal working memory for concordant arm- and leg-related action words brought about by complex sequential movements of the hands and feet. In this article, we seek to determine whether the sign of the functional interaction between language and action systems of the human brain can be changed in a predictable manner by changing movement type. We here report that the sign of the effect of motor movement on action word memory can be reversed from interference to facilitation if, instead of complex movement sequences, simple repetitive movements are performed. Specifically, when engaged in finger tapping, subjects were able to remember relatively more arm-related action words (as compared to control conditions), thus documenting an enhancement of working memory brought about by simple hand movements. In contrast, when performing complex sequences of finger movements, an effector-specific degradation of action word memory was found. By manipulating the sign of the effect in accord with theory-driven predictions, these findings provide support for shared neural bases for motor movement and verbal working memory for action-related words and strengthen the argument that motor systems play a causal and functionally relevant role in language processing semantically related to action.

Separable affix verbs consist of a stem and a derivational affix, which, in some languages can appear together or in discontinuous, distributed form, e.g., German “aufgreifen” and “greifen … auf” [“up-pick(ing)” and “pick … up”]. Certain stems can combine with only certain affixes. However, many such combinations are evaluated not as clearly correct or incorrect, but frequently take an intermediate status with participants rating them ambiguously. Here, we mapped brain responses to combinations of verb stems and affixes realized in short sentences, including more and less common particle verbs, borderline acceptable combinations and clear violations. Event-related potential responses to discontinuous particle verbs were obtained for five affixes re-combined with 10 verb stems, situated within short, German sentences, i.e., “sie en es ,” English: “they it .” The congruity of combinations was assessed both with behavioral ratings of the stimuli and corpus-derived probability measures. The size of a frontal N400 correlated with the degree of incongruency between stem and affix, as assessed by both measures. Behavioral ratings performed better than corpus-derived measures in predicting N400 magnitudes, and a combined model performed best of all. No evidence for a discrete, right/wrong effect was found. We discuss methodological implications and integrate the results into past research on the N400 and neurophysiological studies on separable-affix verbs, generally.

A range of methods in clinical research aim to assess treatment-induced progress in aphasia therapy. Here, we used a crossover randomized controlled design to compare the suitability of utterance-centered and dialogue-sensitive outcome measures in speech-language testing. Fourteen individuals with post-stroke chronic non-fluent aphasia each received two types of intensive training in counterbalanced order: conventional confrontation naming, and communicative-pragmatic speech-language therapy (Intensive Language-Action Therapy, an expanded version of Constraint-Induced Aphasia Therapy). Motivated by linguistic-pragmatic theory and neuroscience data, our dependent variables included a newly created diagnostic instrument, the Action Communication Test (ACT). This diagnostic instrument requires patients to produce target words in two conditions: (i) utterance-centered object naming, and (ii) communicative-pragmatic social interaction based on verbal requests. In addition, we administered a standardized aphasia test battery, the Aachen Aphasia Test (AAT). Composite scores on the ACT and the AAT revealed similar patterns of changes in language performance over time, irrespective of the treatment applied. Changes in language performance were relatively consistent with the AAT results also when considering both ACT subscales separately from each other. However, only the ACT subscale evaluating verbal requests proved to be successful in distinguishing between different types of training in our patient sample. Critically, testing duration was substantially shorter for the entire ACT (10–20 min) than for the AAT (60–90 min). Taken together, the current findings suggest that communicative-pragmatic methods in speech-language testing provide a sensitive and time-effective measure to determine the outcome of aphasia therapy.

In the neuroscience of language, phonemes are frequently described as multimodal units whose neuronal representations are distributed across perisylvian cortical regions, including auditory and sensorimotor areas. A different position views phonemes primarily as acoustic entities with posterior temporal localization, which are functionally independent from frontoparietal articulatory programs. To address this current controversy, we here discuss experimental results from neuroimaging (fMRI) as well as transcranial magnetic stimulation (TMS) studies. On first glance, a mixed picture emerges, with earlier research documenting neurofunctional distinctions between phonemes in both temporal and frontoparietal sensorimotor systems, but some recent work seemingly failing to replicate the latter. Detailed analysis of methodological differences between studies reveals that the way experiments are set up explains whether sensorimotor cortex maps phonological information during speech perception or not. In particular, acoustic noise during the experiment and ‘motor noise’ caused by button press tasks work against the frontoparietal manifestation of phonemes. We highlight recent studies using sparse imaging and passive speech perception tasks along with multivariate pattern analysis (MVPA) and especially representational similarity analysis (RSA), which succeeded in separating acoustic-phonological from general-acoustic processes and in mapping specific phonological information on temporal and frontoparietal regions. The question about a causal role of sensorimotor cortex on speech perception and understanding is addressed by reviewing recent TMS studies. We conclude that frontoparietal cortices, including ventral motor and somatosensory areas, reflect phonological information during speech perception and exert a causal influence on understanding.

Recent cognitive neuroscience research improved our understanding of where, when, how, and why language circuits emerge and activate in the human brain. Where: Regions crucial for very specific linguistic processes were delineated; phonetic features and fine semantic categories could be mapped onto specific sets of cortical areas. When: Brain correlates of phonological, syntactic and semantic processes were documented early-on, suggesting language understanding in an instant (within 250ms). How: New mechanistic network models mimicking structure and function of left-perisylvian language areas suggest that multimodal action-perception circuits — rather than separate modules for action and perception — carry the processing resources for language use and understanding. Why language circuits emerge in specific areas, become active at specific early time points and are connected in specific ways is best addressed in light of neuroscience principles governing neuronal activation, correlation learning, and, critical-ly, partly predetermined structural information wired into connections between cortical neurons and areas.

Memory traces for words are frequently conceptualized neurobiologically as networks of neurons interconnected via reciprocal links developed through associative learning in the process of language acquisition. Neurophysiological reflection of activation of such memory traces has been reported using the mismatch negativity brain potential (MMN), which demonstrates an enhanced response to meaningful words over meaningless items. This enhancement is believed to be generated by the activation of strongly intraconnected long-term memory circuits for words that can be automatically triggered by spoken linguistic input and that are absent for unfamiliar phonological stimuli. This conceptual framework critically predicts different amounts of activation depending on the strength of the word's lexical representation in the brain. The frequent use of words should lead to more strongly connected representations, whereas less frequent items would be associated with more weakly linked circuits. A word with higher frequency of occurrence in the subject's language should therefore lead to a more pronounced lexical MMN response than its low-frequency counterpart. We tested this prediction by comparing the event-related potentials elicited by low- and high-frequency words in a passive oddball paradigm; physical stimulus contrasts were kept identical. We found that, consistent with our prediction, presenting the high-frequency stimulus led to a significantly more pronounced MMN response relative to the low-frequency one, a finding that is highly similar to previously reported MMN enhancement to words over meaningless pseudowords. Furthermore, activation elicited by the higher-frequency word peaked earlier relative to low-frequency one, suggesting more rapid access to frequently used lexical entries. These results lend further support to the above view on word memory traces as strongly connected assemblies of neurons. The speed and magnitude of their activation appears to be linked to the strength of internal connections in a memory circuit, which is in turn determined by the everyday use of language elements.