Person:

Pascual-Leone, Alvaro

Many empiricist theories hold that concepts are composed of sensory–motor primitives. For example, the meaning of the word “run” is in part a visual image of running. If action concepts are partly visual, then the concepts of congenitally blind individuals should be altered in that they lack these visual features. We compared semantic judgments and neural activity during action verb comprehension in congenitally blind and sighted individuals. Participants made similarity judgments about pairs of nouns and verbs that varied in the visual motion they conveyed. Blind adults showed the same pattern of similarity judgments as sighted adults. We identified the left middle temporal gyrus (lMTG) brain region that putatively stores visual–motion features relevant to action verbs. The functional profile and location of this region was identical in sighted and congenitally blind individuals. Furthermore, the lMTG was more active for all verbs than nouns, irrespective of visual–motion features. We conclude that the lMTG contains abstract representations of verb meanings rather than visual–motion images. Our data suggest that conceptual brain regions are not altered by the sensory modality of learning.

Selective deficits in producing verbs relative to nouns in speech are well documented in neuropsychology and have been associated with left hemisphere frontal cortical lesions resulting from stroke and other neurological disorders. The basis for these impairments is unresolved: Do they arise because of differences in the way grammatical categories of words are organized in the brain, or because of differences in the neural representation of actions and objects? We used repetitive transcranial magnetic stimulation (rTMS) to suppress the excitability of a portion of left prefrontal cortex and to assess its role in producing nouns and verbs. In one experiment subjects generated real words; in a second, they produced pseudowords as nouns or verbs. In both experiments, response latencies increased for verbs but were unaffected for nouns following rTMS. These results demonstrate that grammatical categories have a neuroanatomical basis and that the left prefrontal cortex is selectively engaged in processing verbs as grammatical objects.

A number of researchers have proposed that the premotor and motor areas are critical for the representation of words that refer to actions, but not objects. Recent evidence against this hypothesis indicates that the left premotor cortex is more sensitive to grammatical differences than to conceptual differences between words. However, it may still be the case that other anterior motor regions are engaged in processing a word's sensorimotor features. In the present study, we used singleand paired-pulse transcranial magnetic stimulation to test the hypothesis that left primary motor cortex is activated during the retrieval of words (nouns and verbs) associated with specific actions. We found that activation in the motor cortex increased for action words compared with non-action words, but was not sensitive to the grammatical category of the word being produced. These results complement previous findings and support the notion that producing a word activates some brain regions relevant to the sensorimotor properties associated with that word regardless of its grammatical category.

Neuropsychological and neurophysiological studies suggest that the production of verbs in speech depends on cortical regions in the left frontal lobe. However, the precise topography of these regions, and their functional roles in verb production, remains matters of debate. In an earlier study with repetitive transcranial magnetic stimulation (rTMS), we showed that stimulation to the left anterior midfrontal gyrus disrupted verb production, but not noun production, in a task that required subjects to perform simple morphological alternations. This result raises a number of questions: for example, is the effect of stimulation focal and specific to that brain region? Is the behavioral effect limited to rule-based, regular transformations, or can it be generalized over the grammatical category? In the present study, we used rTMS to suppress the excitability of distinct parts of the left prefrontal cortex to assess their role in producing regular and irregular verbs compared to nouns. We compared rTMS to sham stimulation and to stimulation of homologous areas in the right hemisphere. Response latencies increased for verbs, but were unaffected for nouns, following stimulation to the left anterior midfrontal gyrus. No significant interference specific for verbs resulted after stimulation to two other areas in the left frontal lobe, the posterior midfrontal gyrus and Broca's area. These results therefore reinforce the idea that the left anterior midfrontal cortex is critical for processing verbs. Moreover, none of the regions stimulated was preferentially engaged in the production of regular or irregular inflection, raising questions about the role of the frontal lobes in processing inflectional morphology.

Comprehension of words is an important part of the language faculty, involving the joint activity of frontal and temporo-parietal brain regions. Transcranial Magnetic Stimulation (TMS) enables the controlled perturbation of brain activity, and thus offers a unique tool to test specific predictions about the causal relationship between brain regions and language understanding. This potential has been exploited to better define the role of regions that are classically accepted as part of the language-semantic network. For instance, TMS has contributed to establish the semantic relevance of the left anterior temporal lobe, or to solve the ambiguity between the semantic vs. phonological function assigned to the left inferior frontal gyrus (LIFG). We consider, more closely, the results from studies where the same technique, similar paradigms (lexical-semantic tasks) and materials (words) have been used to assess the relevance of regions outside the classically-defined language-semantic network—i.e., precentral motor regions—for the semantic analysis of words. This research shows that different aspects of the left precentral gyrus (primary motor and premotor sites) are sensitive to the action-non action distinction of words' meanings. However, the behavioral changes due to TMS over these sites are incongruent with what is expected after perturbation of a task-relevant brain region. Thus, the relationship between motor activity and language-semantic behavior remains far from clear. A better understanding of this issue could be guaranteed by investigating functional interactions between motor sites and semantically-relevant regions.