Cerebellar involvement in verb generation: an fMRI study

A possible role of the human cerebellum in the generation of verbs corresponding to presented nouns has been suggested. Previous functional brain imaging studies have compared generation of verbs with the reading of nouns as a measure of verb generation. In the present fMRI study involving healthy human subjects, the effects of speech articulation and motor imagery associated with verb production were investigated in greater detail. Generation of verbs to visually presented nouns was compared to a condition in which subjects read those same verbs that had been individually generated by each subject. Activation in lobule HVI/Crus I of the right cerebellar hemisphere was found as a measure of verb generation. In contrast, reading of verbs as a measure of speech articulation evoked cerebellar activations in both left and right paravermal lobule VI. These results suggest an involvement of the right lateral cerebellar hemisphere in linguistic functions during verb generation. Alternatively, effects of inner speech could also possibly explain the results.     

To move or not to move: imperatives modulate action-related verb processing in the motor system

It has been suggested that the processing of action-related words involves activation of the motor circuitry. Using fMRI (functional magnetic resonance imaging), the current study further explored the interaction between action and language by investigating whether the linguistic context, in which an action word occurs, modulates motor circuitry activity related to the processing of action words. To this end, we examined whether the presentation of hand action-related verbs as positive or negative imperatives, for example, “Do grasp” or “Don't write,” modulates neural activity in the hand area of primary motor cortex (M1) or premotor cortex (Pm). Subjects (n = 19) were asked to read silently the imperative phrases, in which both meaningful action verbs and meaningless pseudo-verbs were presented, and to decide whether they made sense (lexical decision task). At the behavioral level, response times in the lexical decision task were significantly longer for negative, compared to positive, imperatives. At the neural level, activity was differentially decreased by action verbs presented as negative imperatives for the premotor and the primary motor cortex of both hemispheres. The data suggest that context (here: positive vs. negative imperatives), in which an action verb is encountered, modulates the neural activity within key areas of the motor system. The finding implies that motor simulation (or motor planning) rather than semantic processing per se may underlie previously observed motor system activation related to action verb processing. Furthermore, the current data suggest that negative imperatives may inhibit motor simulation or motor planning processes.     

Lexical acquisition in progressive aphasia and frontotemporal dementia

We investigated the characteristics of language difficulty in frontotemporal dementia (FTD) by exposing these patients to a new verb in a naturalistic manner and then assessing acquisition of the grammatical, semantic, and thematic matrix information associated with the new word. We found that FTD patients have difficulty relative to healthy seniors in their acquisition of the new verb, but that progressive nonfluent aphasia (PNFA), semantic dementia (SD), and social/dysexecutive variant (SOC/EXEC) subgroups of FTD demonstrate relatively distinct impairment profiles. Specifically, PNFA patients showed relative difficulty assigning the new verb to its correct grammatical form class, reflecting compromised processing of the associated grammatical information. SD patients were impaired at associating the new word with its pictorial representation, suggesting impaired processing of the new verb's semantic attributes. SOC/EXEC patients showed their greatest difficulty judging violations of the new word's associated thematic roles, implying that limited executive resources underlie in part the difficulty in integrating grammatical and semantic information into a coherent thematic matrix. Similar impairment profiles were seen during a follow-up session one week after the initial evaluation. These deficits in lexical acquisition reflect the breakdown of a language-processing system that consists of highly interactive but partially dissociable grammatical, semantic, and resource-based components, leading to relatively distinct language-processing deficits in each subgroup of patients with FTD.     

Lexical acquisition in progressive aphasia and frontotemporal dementia

We investigated the characteristics of language difficulty in frontotemporal dementia (FTD) by exposing these patients to a new verb in a naturalistic manner and then assessing acquisition of the grammatical, semantic, and thematic matrix information associated with the new word. We found that FTD patients have difficulty relative to healthy seniors in their acquisition of the new verb, but that progressive nonfluent aphasia (PNFA), semantic dementia (SD), and social/dysexecutive variant (SOC/EXEC) subgroups of FTD demonstrate relatively distinct impairment profiles. Specifically, PNFA patients showed relative difficulty assigning the new verb to its correct grammatical form class, reflecting compromised processing of the associated grammatical information. SD patients were impaired at associating the new word with its pictorial representation, suggesting impaired processing of the new verb's semantic attributes. SOC/EXEC patients showed their greatest difficulty judging violations of the new word's associated thematic roles, implying that limited executive resources underlie in part the difficulty in integrating grammatical and semantic information into a coherent thematic matrix. Similar impairment profiles were seen during a follow-up session one week after the initial evaluation. These deficits in lexical acquisition reflect the breakdown of a language-processing system that consists of highly interactive but partially dissociable grammatical, semantic, and resource-based components, leading to relatively distinct language-processing deficits in each subgroup of patients with FTD.     

The role of LIFG-based executive control in sentence comprehension

Prior research has shown that patients with damage to the left inferior frontal gyrus (LIFG) can have difficulties in executive control as well as understanding syntactic garden-paths, that is, sentences with a temporary syntactic ambiguity that resolve towards a less preferred interpretation. The present study tested two LIFG patients on object/subject garden path and matched syntactically unambiguous sentences. Besides syntactic ambiguity, support for the preferred but ultimately inappropriate interpretation was manipulated via verb bias, using verbs that were neutral between alternative analyses and verbs that were biased towards the context-inappropriate option. The LIFG patients, a non-LIFG patient, and healthy controls were tested on a sentence interpretation task (Experiment 1) and grammaticality judgement (Experiment 2). In contrast to the non-LIFG patient and controls, the LIFG patients showed impaired thematic role assignment across garden-path as well as unambiguous sentences, which tended to be worse with biased verbs. The results argue for a role of executive control in overcoming verb bias across diverse sentence processing situations, including, but not limited to, garden-path revision.     

Nouns and verbs in the brain: Grammatical class and task specific effects as revealed by fMRI

The wide variety of techniques and tasks used to study the neural correlates of noun and verb processing has resulted in a body of inconsistent evidence. We performed a functional magnetic resonance imaging (fMRI) experiment to detect grammatical class effects that generalize across tasks. A total of 12 participants undertook a grammatical-class switching task (GCST), in which they were presented with a noun (or a verb) and were asked to retrieve the corresponding verb (or noun), and a classical picture naming task (PNT) widely used in the previous aphasiological and imaging literature. The GCST was explicitly designed to ensure control over confounding variables, such as stimulus complexity or imageability. Conjunction analyses of the haemodynamic responses measured in the two tasks indicated a shared verb-related activation of a dorsal premotor and posterior parietal network, pointing to a strong relationship between verb representation and action-oriented (visuo-)spatial knowledge. On the other hand, no brain area was consistently associated with nouns in both tasks. Moreover, there were task-dependent differences between noun and verb retrieval both at behavioural and at physiological level; the grammatical class that elicited the longest reaction times in both tasks (i.e., verbs in the PNT and nouns in the GCST) triggered a greater activation of the left inferior frontal gyrus. Therefore, we suggest that this area reflects a general increase in task demand rather than verb processing per se.     

False belief and verb non-factivity: a common neural basis?

Using fMRI, the present study compares the brain activation underlying false belief thinking induced by pictorial, nonverbal material to that instigated by strong non-factive verbs in a sample of adult Chinese speakers. These verbs obligatorily negate their complements which describe the mind content of the sentence agent, and thus may activate part of the false belief network. Some previous studies have shown a behavioral correlation between verb non-factivity/false complementation and conventional false belief but corresponding neural evidence is lacking. Our results showed that the non-factive grammar and false belief commonly implicated the right temporo-parietal junction (TPJ), which had been shown by past studies to play a role in general mentalizing. Regions that were unique to nonverbal false belief were the left TPJ and right middle frontal gyrus (MFG), whereas the unique regions for the non-factive grammar were the left inferior frontal gyrus (IFG) and right superior temporal gyrus (STG). Hence, conventional nonverbal false belief and verb non-factivity have both shared and unique neural representations.     

Differences between nouns and verbs after anterior temporal lobectomy

To test the claim that lesions of left anterior and middle temporal cortical structures specifically impair processing of nouns but not verbs, 56 left-hemisphere-language-dominant patients who had undergone anterior temporal lobectomy (ATL) completed tasks assessing confrontation naming of pictured objects and actions, generation of synonyms for nouns and verbs, and semantic lexical judgments about nouns and verbs. Compared with right ATL patients, left ATL patients were impaired across different tasks that assessed naming and comprehension of high-imageability as well as low-imageability nouns. These groups did not differ, however, in verb naming or comprehension on most tasks. Results are consistent with the hypothesized specialization of left temporal lobe structures for processing nouns and suggest that naming problems commonly seen after left ATL extend beyond difficulties with retrieving object names and may be related to subtle disturbances in comprehension of the meanings underlying nominal word forms.     

Functional magnetic resonance imaging evidence for right-hemisphere involvement in processing unusual semantic relationships

Brain areas active in generating usual (typical) or unusual (atypical) noun-verb relations were examined using functional magnetic resonance imaging. Seven adults generated the 1st verb to come to mind (usual verb) in response to novel and repeated nouns (priming test) and then generated either an unusual verb or the 1st verb to come to mind in response to novel nouns (unusual test). The left inferior prefrontal cortex (LIPC) and right cerebellum were more active when generating usual verbs to novel nouns than to repeated nouns. When participants generated unusual verbs, there was no increased activation in LIPC, but there was increased activity in the right middle and superior frontal gyri, left middle frontal gyrus, and bilateral cerebellum. Results support theories that the right hemisphere is involved in the processing of distant associations that may be useful in creative thought and problem solving.     

Passive reading and motor imagery about hand actions and tool-use actions: an fMRI study

Recent studies have shown that motor activations in action verb comprehension can be modulated by task demands (e.g., motor imagery vs. passive reading) and the specificity of action verb meaning. However, how the two factors work together to influence the involvement of the motor system during action verb comprehension is still unclear. To address the issue, the current study investigated the brain activations in motor imagery and passive reading of verbs about hand actions and tool-use actions. Three types of Chinese verbs were used, including hand-action verbs and two types of tool-use verbs emphasizing either the hand or tools information. Results indicated that all three types of verbs elicited common activations in hand motor areas during passive reading and motor imagery. Contrast analyses showed that in the hand verbs and the tool verbs where the hand information was emphasized, motor imagery elicited stronger effects than passive reading in the superior frontal gyrus, supplemental motor area and cingulate cortex that are related to motor control and regulation. For tool-use verbs emphasizing tools information, the motor imagery task elicited stronger activity than passive reading in occipital regions related to visual imagery. These results suggest that motor activations during action verb comprehension can be modulated by task demands and semantic features of action verbs. The sensorimotor simulation during language comprehension is flexible and determined by the interactions between linguistic and extralinguistic contexts.     

Neural synchronization mediates on-line sentence processing: EEG coherence evidence from filler-gap constructions

This study investigated cognitive and neural processes involved in gap filling during on-line sentence comprehension. Electroencephalogram (EEG) coherences were used to demonstrate that increases in the synchronization of neural activity in different cortical regions occur during gap filling when load in semantic working memory is high due to semantically unrelated words in the filler-gap interval. Sentences could either require gap filling at a verb or not, and the nouns preceding the verb could be either semantically related or unrelated. In the unrelated but not related condition, coherences in the beta band were larger during verb processing for sentences requiring gap filling compared to sentences not requiring gap filling. The coherence changes involved linkages between frontal and posterior temporal-parietal sites in both hemispheres. These results further indicate that semantic working memory is involved in the process of gap filling.     

Implicit/explicit memory dissociation in Alzheimer's disease: the consequence of inappropriate processing?

Dual-process theories of recognition posit that perceptual fluency contributes to both familiarity-based explicit recognition and perceptual priming. However, the priming-without-recognition dissociation, as observed through the intact mere exposure effect and impaired recognition in patients with Alzheimer's disease (AD), might indicate that familiarity and perceptual priming are functionally distinct. This study investigated whether the AD patients' processing strategies at testing may explain this priming-without-recognition dissociation. First, we replicated the priming-without-recognition effect in 16 patients who exhibited intact exposure effects despite null recognition. Second, we showed that, under identical conditions, inducing a holistic processing strategy during recognition testing increased AD patients' recognition--performance was similar for AD patients and healthy control participants. Furthermore, prompting analytic processing during both priming and recognition tasks decreased AD patients' performance in both tasks. These findings suggest that the extent to which AD patients use perceptual fluency in priming and recognition tasks is contingent on their processing approach. The choice of processing strategy may depend on how difficult patients perceive the task to be.     

Making Memories: A Cross-Sectional Investigation of Episodic Memory Encoding in Childhood Using fMRI

In adults, the neural substrate associated with encoding memories connected to a specific time and place include the prefrontal cortex and medial temporal lobe (MTL). Using functional magnetic resonance imaging, this research studied the developmental trajectory of this frontal-MTL system by comparing 7- and 8-year-old children to those who were 10 or older in conditions that promoted episodic encoding. In 1 condition, participants generated verbs from nouns heard; in another, they listened to short stories for comprehension. Regions in which brain activation predicted subsequent recognition memory performance were identified. These included the left prefrontal cortex, but not MTL, in the verb generation condition for both age groups. In the story comprehension condition, activation in left posterior MTL predicted subsequent memory performance in both age groups, and activation in left anterior MTL (including the hippocampus proper) and left prefrontal cortex predicted subsequent memory only for the older children. These results illustrate both similarities and differences in how brain systems interact in development to mediate the formation of episodic memories.     

Making memories: a cross-sectional investigation of episodic memory encoding in childhood using FMRI

In adults, the neural substrate associated with encoding memories connected to a specific time and place include the prefrontal cortex and medial temporal lobe (MTL). Using functional magnetic resonance imaging, this research studied the developmental trajectory of this frontal-MTL system by comparing 7- and 8-year-old children to those who were 10 or older in conditions that promoted episodic encoding. In 1 condition, participants generated verbs from nouns heard; in another, they listened to short stories for comprehension. Regions in which brain activation predicted subsequent recognition memory performance were identified. These included the left prefrontal cortex, but not MTL, in the verb generation condition for both age groups. In the story comprehension condition, activation in left posterior MTL predicted subsequent memory performance in both age groups, and activation in left anterior MTL (including the hippocampus proper) and left prefrontal cortex predicted subsequent memory only for the older children. These results illustrate both similarities and differences in how brain systems interact in development to mediate the formation of episodic memories.     

The organization of the conceptual system: the case of the "object versus action" dimension

There are very numerous reports in the neuropsychological literature of patients showing, in naming and/or comprehension tasks, a disproportionate deficit for nouns in comparison with verbs or a disproportionate deficit for verbs in comparison with nouns. A number of authors advanced that, in at least some or even in every of these reported cases, the noun/verb dissociation in fact reflected an underlying conceptual deficit disproportionately affecting either object or action concepts. These patterns thus would put an additional constraint on theories of conceptual knowledge organization, which should be able to explain how brain damage could selectively disrupt the concepts of objects or the concepts of actions. We have reviewed 69 papers (published from 1984 to 2009) that reported a pattern of a noun or a verb disproportionate deficit in a single-case, multiple-case, or group study of brain-damaged patients with various aetiologies. From this review, we concluded that none of these studies provided compelling evidence in favour of the interpretation that the observed noun or verb disproportionate deficit arose at the conceptual processing level and, accordingly, that this level may be organized according to the "object/action" dimension. Furthermore, we argue that investigating conceptual impairments in brain-damaged patients according to the "object/action" dichotomy is not empirically fruitful if the purpose is to inform theories of conceptual knowledge organization. In order to provide evidence relevant to these theories, one needs to consider finer grained distinctions within both the object and the action category when investigating the scope of the patients' conceptual impairment.     

Grammatical number agreement processing using the visual half-field paradigm: An event-related brain potential study

Despite indications in the split-brain and lesion literatures that the right hemisphere is capable of some syntactic analysis, few studies have investigated right hemisphere contributions to syntactic processing in people with intact brains. Here we used the visual half-field paradigm in healthy adults to examine each hemisphere's processing of correct and incorrect grammatical number agreement marked either lexically, e.g., antecedent/reflexive pronoun (“The grateful niece asked herself/*themselves…”) or morphologically, e.g., subject/verb (“Industrial scientists develop/*develops…”). For reflexives, response times and accuracy of grammaticality decisions suggested similar processing regardless of visual field of presentation. In the subject/verb condition, we observed similar response times and accuracies for central and right visual field (RVF) presentations. For left visual field (LVF) presentation, response times were longer and accuracy rates were reduced relative to RVF presentation. An event-related brain potential (ERP) study using the same materials revealed similar ERP responses to the reflexive pronouns in the two visual fields, but very different ERP effects to the subject/verb violations. For lexically marked violations on reflexives, P600 was elicited by stimuli in both the LVF and RVF; for morphologically marked violations on verbs, P600 was elicited only by RVF stimuli. These data suggest that both hemispheres can process lexically marked pronoun agreement violations, and do so in a similar fashion. Morphologically marked subject/verb agreement errors, however, showed a distinct LH advantage.     

The organization of the conceptual system: The case of the “object versus action” dimension

There are very numerous reports in the neuropsychological literature of patients showing, in naming and/or comprehension tasks, a disproportionate deficit for nouns in comparison with verbs or a disproportionate deficit for verbs in comparison with nouns. A number of authors advanced that, in at least some or even in every of these reported cases, the noun/verb dissociation in fact reflected an underlying conceptual deficit disproportionately affecting either object or action concepts. These patterns thus would put an additional constraint on theories of conceptual knowledge organization, which should be able to explain how brain damage could selectively disrupt the concepts of objects or the concepts of actions. We have reviewed 69 papers (published from 1984 to 2009) that reported a pattern of a noun or a verb disproportionate deficit in a single-case, multiple-case, or group study of brain-damaged patients with various aetiologies. From this review, we concluded that none of these studies provided compelling evidence in favour of the interpretation that the observed noun or verb disproportionate deficit arose at the conceptual processing level and, accordingly, that this level may be organized according to the “object/action” dimension. Furthermore, we argue that investigating conceptual impairments in brain-damaged patients according to the “object/action” dichotomy is not empirically fruitful if the purpose is to inform theories of conceptual knowledge organization. In order to provide evidence relevant to these theories, one needs to consider finer grained distinctions within both the object and the action category when investigating the scope of the patients' conceptual impairment.     

Functional neuroimaging of grammatical class: Ambiguous and unambiguous nouns and verbs

Neuropsychological and neuroimaging studies of grammatical-class differences suggest that nouns and verbs may be associated with different brain regions. The current functional magnetic resonance imaging (fMRI) study focused on auditory grammaticality judgements of two-word stimuli consisting of noun/verb ambiguous and unambiguous nouns and verbs preceded by either “the” or “to”. The fMRI results showed effects of class ambiguity in the left inferior frontal gyrus, possibly due to greater selection demands, and effects of grammaticality (“yes” vs. “no” response) in left superior temporal gyrus, consistent with greater activation for trials with greater conflict. In addition, posterior left temporal lobe regions showed increased activity on unambiguous trials for verbs than for nouns, consistent with prior imaging studies. Thus, in a task specifically focused on the grammatical (rather than semantic) aspects of words and that used morphologically simple nouns and verbs controlled for imageability, verbs preferentially activated posterior temporal but not frontal lobe regions. However, for ambiguous trials in this same region, nouns showed greater activation than verbs, suggesting that these effects can be modulated by class ambiguity.     

Task-dependent neural correlates of the processing of verbal threat-related stimuli in social phobia

The neural basis of abnormal processing of phobia-related linguistic cues in individuals suffering from social phobia is unknown, particularly in respect to different task conditions. Using event-related functional magnetic resonance imaging, this study investigated brain activation to phobia-related and phobia-unrelated words in 19 socially phobic patients and 18 healthy control subjects (HC) while subjects had to attend either to social meaning or to grammatical category of words (direct or indirect task). During the indirect task, patients, compared to HC, showed an increased activation of the amygdala and orbitofrontal cortex (OFC) in response to phobia-related vs. phobia-unrelated words. Activation of the insula was positively correlated with patients’ symptom severity during the direct task. The results suggest a specific role of the amygdala and OFC during the processing of verbal phobia-relevant distracting information. In contrast, insula activation seems to be more important for direct processing of disorder-related words, especially in more severe cases of social phobia.