Person:

Shapiro, Kevin Alfred

The ability to generate novel sentences depends on cognitive operations that specify the syntactic function of nouns, verbs, and other words retrieved from the mental lexicon. Although neuropsychological studies suggest that such operations rely on neural circuits distinct from those encoding word form and meaning, it has not been possible to characterize this distinction definitively with neuroimaging. We used functional magnetic resonance imaging (fMRI) to show that a brain area engaged in a given grammatical operation can be identified uniquely by a monotonic decrease in activation as that operation is repeated. We applied this methodology to identify areas involved selectively in the operation of inflection of nouns or verbs. By contrast, areas involved in processing word meaning do not show this monotonic adaptation across stimuli. These results are the first to demonstrate adaptation in the fMR signal evoked not by specific stimuli, but by well-defined cognitive linguistic operations.

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.