Tomatoes Are Red: The Perception of Achromatic Objects Elicits Retrieval of Associated Color Knowledge.

When preparing to name an object, semantic knowledge about the object and its attributes is activated, including perceptual properties. It is unclear, however, whether semantic attribute activation contributes to lexical access or is a consequence of activating a concept irrespective of whether that concept is to be named or not. In this study, we measured neural responses using fMRI while participants named objects that are typically green or red, presented in black line drawings. Furthermore, participants underwent two other tasks with the same objects, color naming and semantic judgment, to see if the activation pattern we observe during picture naming is (a) similar to that of a task that requires accessing the color attribute and (b) distinct from that of a task that requires accessing the concept but not its name or color. We used representational similarity analysis to detect brain areas that show similar patterns within the same color category, but show different patterns across the two color categories. In all three tasks, activation in the bilateral fusiform gyri ("Human V4") correlated with a representational model encoding the red-green distinction weighted by the importance of color feature for the different objects. This result suggests that when seeing objects whose color attribute is highly diagnostic, color knowledge about the objects is retrieved irrespective of whether the color or the object itself have to be named.

Parallel or sequential? Decoding conceptual and phonological/phonetic information from MEG signals during language production.

Speaking requires the temporally coordinated planning of core linguistic information, from conceptual meaning to articulation. Recent neurophysiological results suggested that these operations involve a cascade of neural events with subsequent onset times, whilst competing evidence suggests early parallel neural activation. To test these hypotheses, we examined the sources of neuromagnetic activity recorded from 34 participants overtly naming 134 images from 4 object categories (animals, tools, foods and clothes). Within each category, word length and phonological neighbourhood density were co-varied to target phonological/phonetic processes. Multivariate pattern analyses (MVPA) searchlights in source space decoded object categories in occipitotemporal and middle temporal cortex, and phonological/phonetic variables in left inferior frontal (BA 44) and motor cortex early on. The findings suggest early activation of multiple variables due to intercorrelated properties and interactivity of processing, thus raising important questions about the representational properties of target words during the preparatory time enabling overt speaking.

Brain correlates of action word memory revealed by fMRI.

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.

The Time Course of Language Production as Revealed by Pattern Classification of MEG Sensor Data.

Language production involves a complex set of computations, from conceptualization to articulation, which are thought to engage cascading neural events in the language network. However, recent neuromagnetic evidence suggests simultaneous meaning-to-speech mapping in picture naming tasks, as indexed by early parallel activation of frontotemporal regions to lexical semantic, phonological, and articulatory information. Here we investigate the time course of word production, asking to what extent such "earliness" is a distinctive property of the associated spatiotemporal dynamics. Using MEG, we recorded the neural signals of 34 human subjects (26 males) overtly naming 134 images from four semantic object categories (animals, foods, tools, clothes). Within each category, we covaried word length, as quantified by the number of syllables contained in a word, and phonological neighborhood density to target lexical and post-lexical phonological/phonetic processes. Multivariate pattern analyses searchlights in sensor space distinguished the stimulus-locked spatiotemporal responses to object categories early on, from 150 to 250 ms after picture onset, whereas word length was decoded in left frontotemporal sensors at 250-350 ms, followed by the latency of phonological neighborhood density (350-450 ms). Our results suggest a progression of neural activity from posterior to anterior language regions for the semantic and phonological/phonetic computations preparing overt speech, thus supporting serial cascading models of word production.SIGNIFICANCE STATEMENT Current psycholinguistic models make divergent predictions on how a preverbal message is mapped onto articulatory output during the language planning. Serial models predict a cascading sequence of hierarchically organized neural computations from conceptualization to articulation. In contrast, parallel models posit early simultaneous activation of multiple conceptual, phonological, and articulatory information in the language system. Here we asked whether such earliness is a distinctive property of the neural dynamics of word production. The combination of the millisecond precision of MEG with multivariate pattern analyses revealed subsequent onset times for the neural events supporting semantic and phonological/phonetic operations, progressing from posterior occipitotemporal to frontal sensor areas. The findings bring new insights for refining current theories of language production.

Distinct fronto-temporal substrates of distributional and taxonomic similarity among words: evidence from RSA of BOLD signals.

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.

Representational Similarity Mapping of Distributional Semantics in Left Inferior Frontal, Middle Temporal, and Motor Cortex.

Language comprehension engages a distributed network of frontotemporal, parietal, and sensorimotor regions, but it is still unclear how meaning of words and their semantic relationships are represented and processed within these regions and to which degrees lexico-semantic representations differ between regions and semantic types. We used fMRI and representational similarity analysis to relate word-elicited multivoxel patterns to semantic similarity between action and object words. In left inferior frontal (BA 44-45-47), left posterior middle temporal and left precentral cortex, the similarity of brain response patterns reflected semantic similarity among action-related verbs, as well as across lexical classes-between action verbs and tool-related nouns and, to a degree, between action verbs and food nouns, but not between action verbs and animal nouns. Instead, posterior inferior temporal cortex exhibited a reverse response pattern, which reflected the semantic similarity among object-related nouns, but not action-related words. These results show that semantic similarity is encoded by a range of cortical areas, including multimodal association (e.g., anterior inferior frontal, posterior middle temporal) and modality-preferential (premotor) cortex and that the representational geometries in these regions are partly dependent on semantic type, with semantic similarity among action-related words crossing lexical-semantic category boundaries.

Decompositional Representation of Morphological Complexity: Multivariate fMRI Evidence from Italian.

Derivational morphology is a cross-linguistically dominant mechanism for word formation, combining existing words with derivational affixes to create new word forms. However, the neurocognitive mechanisms underlying the representation and processing of such forms remain unclear. Recent cross-linguistic neuroimaging research suggests that derived words are stored and accessed as whole forms, without engaging the left-hemisphere perisylvian network associated with combinatorial processing of syntactically and inflectionally complex forms. Using fMRI with a "simple listening" no-task procedure, we reexamine these suggestions in the context of the root-based combinatorially rich Italian lexicon to clarify the role of semantic transparency (between the derived form and its stem) and affix productivity in determining whether derived forms are decompositionally represented and which neural systems are involved. Combined univariate and multivariate analyses reveal a key role for semantic transparency, modulated by affix productivity. Opaque forms show strong cohort competition effects, especially for words with nonproductive suffixes (ventura, "destiny"). The bilateral frontotemporal activity associated with these effects indicates that opaque derived words are processed as whole forms in the bihemispheric language system. Semantically transparent words with productive affixes (libreria, "bookshop") showed no effects of lexical competition, suggesting morphologically structured co-representation of these derived forms and their stems, whereas transparent forms with nonproductive affixes (pineta, pine forest) show intermediate effects. Further multivariate analyses of the transparent derived forms revealed affix productivity effects selectively involving left inferior frontal regions, suggesting that the combinatorial and decompositional processes triggered by such forms can vary significantly across languages.

Body-part-specific representations of semantic noun categories.

Word meaning processing in the brain involves ventrolateral temporal cortex, but a semantic contribution of the dorsal stream, especially frontocentral sensorimotor areas, has been controversial. We here examine brain activation during passive reading of object-related nouns from different semantic categories, notably animal, food, and tool words, matched for a range of psycholinguistic features. Results show ventral stream activation in temporal cortex along with category-specific activation patterns in both ventral and dorsal streams, including sensorimotor systems and adjacent pFC. Precentral activation reflected action-related semantic features of the word categories. Cortical regions implicated in mouth and face movements were sparked by food words, and hand area activation was seen for tool words, consistent with the actions implicated by the objects the words are used to speak about. Furthermore, tool words specifically activated the right cerebellum, and food words activated the left orbito-frontal and fusiform areas. We discuss our results in the context of category-specific semantic deficits in the processing of words and concepts, along with previous neuroimaging research, and conclude that specific dorsal and ventral areas in frontocentral and temporal cortex index visual and affective-emotional semantic attributes of object-related nouns and action-related affordances of their referent objects.

A role for the motor system in binding abstract emotional meaning.

Sensorimotor areas activate to action- and object-related words, but their role in abstract meaning processing is still debated. Abstract emotion words denoting body internal states are a critical test case because they lack referential links to objects. If actions expressing emotion are crucial for learning correspondences between word forms and emotions, emotion word-evoked activity should emerge in motor brain systems controlling the face and arms, which typically express emotions. To test this hypothesis, we recruited 18 native speakers and used event-related functional magnetic resonance imaging to compare brain activation evoked by abstract emotion words to that by face- and arm-related action words. In addition to limbic regions, emotion words indeed sparked precentral cortex, including body-part-specific areas activated somatotopically by face words or arm words. Control items, including hash mark strings and animal words, failed to activate precentral areas. We conclude that, similar to their role in action word processing, activation of frontocentral motor systems in the dorsal stream reflects the semantic binding of sign and meaning of abstract words denoting emotions and possibly other body internal states.

Neural dynamics of the intention to speak.

When we talk we communicate our intentions. Although the origin of intentional action is debated in cognitive neuroscience, the question of how the brain generates the intention in speech remains still open. Using magnetoencephalography, we investigated the cortical dynamics engaged when healthy subjects attended to either their intention to speak or their actual speech. We found that activity in the right and left parietal cortex increased before subjects became aware of intending to speak. Within the time window of parietal activation, we also observed a transient left frontal activity in Broca's area, a crucial region for inner speech. During attention to speech, neural activity was detected in left prefrontal and temporal areas and in the temporoparietal junction. In agreement with previous results, our findings suggest that the parietal cortex plays a multimodal role in monitoring intentional mechanisms in both action and language. The coactivation of parietal regions and Broca's area may constitute the cortical circuit specific for controlling intentional processes during speech.