The Wernicke conundrum revisited: evidence from connectome-based lesion-symptom mapping.

Wernicke's area has been assumed since the 1800s to be the primary region supporting word and sentence comprehension. However, in 2015 and 2019, Mesulam and colleagues raised what they termed the 'Wernicke conundrum', noting widespread variability in the anatomical definition of this area and presenting data from primary progressive aphasia that challenged this classical assumption. To resolve the conundrum, they posited a 'double disconnection' hypothesis: that word and sentence comprehension deficits in stroke-based aphasia result from disconnection of anterior temporal and inferior frontal regions from other parts of the brain due to white matter damage, rather than dysfunction of Wernicke's area itself. To test this hypothesis, we performed lesion-deficit correlations, including connectome-based lesion-symptom mapping, in four large, partially overlapping groups of English-speaking chronic left hemisphere stroke survivors. After removing variance due to object recognition and associative semantic processing, the same middle and posterior temporal lobe regions were implicated in both word comprehension deficits and complex non-canonical sentence comprehension deficits. Connectome lesion-symptom mapping revealed similar temporal-occipital white matter disconnections for impaired word and non-canonical sentence comprehension, including the temporal pole. We found an additional significant temporal-parietal disconnection for non-canonical sentence comprehension deficits, which may indicate a role for phonological working memory in processing complex syntax, but no significant frontal disconnections. Moreover, damage to these middle-posterior temporal lobe regions was associated with both word and non-canonical sentence comprehension deficits even when accounting for variance due to the strongest anterior temporal and inferior frontal white matter disconnections, respectively. Our results largely agree with the classical notion that Wernicke's area, defined here as middle superior temporal gyrus and middle-posterior superior temporal sulcus, supports both word and sentence comprehension, suggest a supporting role for temporal pole in both word and sentence comprehension, and speak against the hypothesis that comprehension deficits in Wernicke's aphasia result from double disconnection.

Brain Damage Associated with Impaired Sentence Processing in Acute Aphasia.

Left-hemisphere brain damage commonly affects patients' abilities to produce and comprehend syntactic structures, a condition typically referred to as "agrammatism." The neural correlates of agrammatism remain disputed in the literature, and distributed areas have been implicated as important predictors of performance, for example, Broca's area, anterior temporal areas, and temporo-parietal areas. We examined the association between damage to specific language-related ROIs and impaired syntactic processing in acute aphasia. We hypothesized that damage to the posterior middle temporal gyrus, and not Broca's area, would predict syntactic processing abilities. One hundred four individuals with acute aphasia (<20 days poststroke) were included in the study. Structural MRI scans were obtained, and all participants completed a 45-item sentence-picture matching task. We performed an ROI-based stepwise regression analyses to examine the relation between cortical brain damage and impaired comprehension of canonical and noncanonical sentences. Damage to the posterior middle temporal gyrus was the strongest predictor for overall task performance and performance on noncanonical sentences. Damage to the angular gyrus was the strongest predictor for performance on canonical sentences, and damage to the posterior superior temporal gyrus predicted noncanonical scores when performance on canonical sentences was included as a cofactor. Overall, our models showed that damage to temporo-parietal and posterior temporal areas was associated with impaired syntactic comprehension. Our results indicate that the temporo-parietal area is crucially implicated in complex syntactic processing, whereas the role of Broca's area may be complementary.

Modulation of vocal pitch control through high-definition transcranial direct current stimulation of the left ventral motor cortex.

Neural interactions between sensorimotor integration mechanisms play critical roles in voice motor control. We investigated how high-definition transcranial direct current stimulation (HD-tDCS) of the left ventral motor cortex modulates neural mechanisms of sensorimotor integration during voice motor control. HD-tDCS was performed during speech vowel production in an altered auditory feedback (AAF) paradigm in response to upward and downward pitch-shift stimuli. In one experiment, two groups received either anodal or cathodal 2 milliamp (mA) HD-tDCS to the left ventral motor cortex while a third group received sham (placebo) stimulation. In a second experiment, two groups received either 1 mA or 2 mA cathodal HD-tDCS to the left ventral motor cortex. Results of the first experiment indicated that the magnitude of vocal compensation was significantly reduced following anodal and cathodal HD-tDCS only in responses to downward pitch-shift AAF stimuli, with stronger effects associated with cathodal HD-tDCS. However, no such effect was observed following sham stimulation. Results of the second experiment indicate that there is not a differential effect of modulation from 1 mA versus 2 mA. Further, these results replicate the directional finding of the first experiment for vocal compensation in response to downward pitch-shift only. These findings suggest that neurostimulation of the left ventral motor cortex modulates sensorimotor mechanisms underlying voice motor control. We speculate that this effect is associated with the increased contribution of feedforward motor mechanisms, leading to reduced compensatory speech responses to AAF.

Cortical and structural-connectivity damage correlated with impaired syntactic processing in aphasia.

Agrammatism in aphasia is not a homogeneous syndrome, but a characterization of a nonuniform set of language behaviors in which grammatical markers and complex syntactic structures are omitted, simplified, or misinterpreted. In a sample of 71 left-hemisphere stroke survivors, syntactic processing was quantified with the Northwestern Assessment of Verbs and Sentences (NAVS). Classification analyses were used to assess the relation between NAVS performance and morphosyntactically reduced speech in picture descriptions. Voxel-based and connectivity-based lesion-symptom mapping were applied to investigate neural correlates of impaired syntactic processing. Despite a nonrandom correspondence between NAVS performance and morphosyntactic production deficits, there was variation in individual patterns of syntactic processing. Morphosyntactically reduced production was predicted by lesions to left-hemisphere inferior frontal cortex. Impaired verb argument structure production was predicted by damage to left-hemisphere posterior superior temporal and angular gyrus, as well as to a ventral pathway between temporal and frontal cortex. Damage to this pathway was also predictive of impaired sentence comprehension and production, particularly of noncanonical sentences. Although agrammatic speech production is primarily predicted by lesions to inferior frontal cortex, other aspects of syntactic processing rely rather on regional integrity in temporoparietal cortex and the ventral stream.

Behavioral and neural correlates of normal aging effects on motor preparatory mechanisms of speech production and limb movement.

Normal aging is associated with decline of the sensorimotor mechanisms that support movement function in the human brain. In this study, we used behavioral and event-related potential (ERP) recordings to investigate the effects of normal aging on the motor preparatory mechanisms of speech production and limb movement. The experiment involved two groups of older and younger adults who performed randomized speech vowel vocalization and button press motor reaction time tasks in response to temporally predictable and unpredictable visual stimuli. Behavioral results revealed age-related slowness of motor reaction time only during speech production in response to temporally unpredictable stimuli, and this effect was accompanied by increased pre-motor ERP activities in older vs. younger adults during the speech task. These results indicate that motor preparatory mechanisms of limb movement during button press are not affected by normal aging, whereas the functional capacity of these mechanisms is reduced in older adults during speech production in response to unpredictable sensory stimuli. These findings suggest that the aging brain selectively compromises the motor timing of speech and recruits additional neural resources for motor planning and execution of speech, as indexed by the increased pre-motor ERP activations in response to temporally unpredictable vs. predictable sensory stimuli.

Anatomy of aphasia revisited.

In most cases, aphasia is caused by strokes involving the left hemisphere, with more extensive damage typically being associated with more severe aphasia. The classical model of aphasia commonly adhered to in the Western world is the Wernicke-Lichtheim model. The model has been in existence for over a century, and classification of aphasic symptomatology continues to rely on it. However, far more detailed models of speech and language localization in the brain have been formulated. In this regard, the dual stream model of cortical brain organization proposed by Hickok and Poeppel is particularly influential. Their model describes two processing routes, a dorsal stream and a ventral stream, that roughly support speech production and speech comprehension, respectively, in normal subjects. Despite the strong influence of the dual stream model in current neuropsychological research, there has been relatively limited focus on explaining aphasic symptoms in the context of this model. Given that the dual stream model represents a more nuanced picture of cortical speech and language organization, cortical damage that causes aphasic impairment should map clearly onto the dual processing streams. Here, we present a follow-up study to our previous work that used lesion data to reveal the anatomical boundaries of the dorsal and ventral streams supporting speech and language processing. Specifically, by emphasizing clinical measures, we examine the effect of cortical damage and disconnection involving the dorsal and ventral streams on aphasic impairment. The results reveal that measures of motor speech impairment mostly involve damage to the dorsal stream, whereas measures of impaired speech comprehension are more strongly associated with ventral stream involvement. Equally important, many clinical tests that target behaviours such as naming, speech repetition, or grammatical processing rely on interactions between the two streams. This latter finding explains why patients with seemingly disparate lesion locations often experience similar impairments on given subtests. Namely, these individuals' cortical damage, although dissimilar, affects a broad cortical network that plays a role in carrying out a given speech or language task. The current data suggest this is a more accurate characterization than ascribing specific lesion locations as responsible for specific language deficits.5705668782001awx363media15705668782001.

Revealing the dual streams of speech processing.

Several dual route models of human speech processing have been proposed suggesting a large-scale anatomical division between cortical regions that support motor-phonological aspects vs. lexical-semantic aspects of speech processing. However, to date, there is no complete agreement on what areas subserve each route or the nature of interactions across these routes that enables human speech processing. Relying on an extensive behavioral and neuroimaging assessment of a large sample of stroke survivors, we used a data-driven approach using principal components analysis of lesion-symptom mapping to identify brain regions crucial for performance on clusters of behavioral tasks without a priori separation into task types. Distinct anatomical boundaries were revealed between a dorsal frontoparietal stream and a ventral temporal-frontal stream associated with separate components. Collapsing over the tasks primarily supported by these streams, we characterize the dorsal stream as a form-to-articulation pathway and the ventral stream as a form-to-meaning pathway. This characterization of the division in the data reflects both the overlap between tasks supported by the two streams as well as the observation that there is a bias for phonological production tasks supported by the dorsal stream and lexical-semantic comprehension tasks supported by the ventral stream. As such, our findings show a division between two processing routes that underlie human speech processing and provide an empirical foundation for studying potential computational differences that distinguish between the two routes.

Effects of aging on temporal predictive mechanisms of speech and hand motor reaction time.

Evidence from previous studies has suggested that movement execution in younger adults is accelerated in response to temporally predictable vs. unpredictable sensory stimuli. This effect indicates that external temporal information can modulate motor behavior; however, how aging can influence temporal predictive mechanisms in motor system has yet to be understood. The objective of the present study was to investigate aging effects on the initiation and inhibition of speech and hand movement reaction times in response to temporally predictable and unpredictable sensory stimuli. Fifteen younger (mean age 22.6) and fifteen older (mean age 63.8) adults performed a randomized speech vowel vocalization or button press initiation and inhibition tasks in two counterbalanced blocks in response to temporally predictable and unpredictable visual cue stimuli. Results showed that motor reaction time was accelerated in both younger and older adults for predictable vs. unpredictable stimuli during initiation and inhibition of speech and hand movement. However, older adults were significantly slower than younger adults in motor execution of speech and hand movement when stimulus timing was unpredictable. Moreover, we found that overall, motor inhibition of speech and hand was executed faster than their initiation. Our findings suggest that older adults can compensate age-related decline in motor reaction times by incorporating external temporal information and execute faster movement in response to predictable stimuli, whereas unpredictable temporal information cannot counteract aging effects efficiently and lead to less accurate motor timing predictive codes for speech production and hand movement.

Non-fluent speech following stroke is caused by impaired efference copy.

Efference copy is a cognitive mechanism argued to be critical for initiating and monitoring speech: however, the extent to which breakdown of efference copy mechanisms impact speech production is unclear. This study examined the best mechanistic predictors of non-fluent speech among 88 stroke survivors. Objective speech fluency measures were subjected to a principal component analysis (PCA). The primary PCA factor was then entered into a multiple stepwise linear regression analysis as the dependent variable, with a set of independent mechanistic variables. Participants' ability to mimic audio-visual speech ("speech entrainment response") was the best independent predictor of non-fluent speech. We suggest that this "speech entrainment" factor reflects integrity of internal monitoring (i.e., efference copy) of speech production, which affects speech initiation and maintenance. Results support models of normal speech production and suggest that therapy focused on speech initiation and maintenance may improve speech fluency for individuals with chronic non-fluent aphasia post stroke.

The Role of Fundamental Frequency and Temporal Envelope in Processing Sentences with Temporary Syntactic Ambiguities.

Previous experiments have demonstrated the impact of speech prosody on syntactic processing. The present study was designed to examine how listeners use specific acoustic properties of prosody for grammatical interpretation. We investigated the independent contributions of two acoustic properties associated with the pitch and rhythmic properties of speech; the fundamental frequency and temporal envelope, respectively. The effect of degrading these prosodic components was examined by testing listeners' ability to parse early-closure garden-path sentences. A second aim was to investigate how effects of prosody interact with semantic effects of sentence plausibility. Using a task that required both a comprehension and a production response, we were able to determine that degradation of the speech envelope more consistently affects syntactic processing than degradation of the fundamental frequency. These effects are exacerbated in sentences with plausible misinterpretations, showing that prosodic degradation interacts with contextual cues to sentence interpretation.

On the Use of the Terms Anodal and Cathodal in High-Definition Transcranial Direct Current Stimulation: A Technical Note.

BACKGROUND: The terms "anodal" and "cathodal" are widely used to describe transcranial direct current stimulation (tDCS) of opposing polarities, often interpreted as excitatory and inhibitory, respectively. However, high-definition tDCS allows for complex electrode configurations that may not be characterized accurately as "anodal" and "cathodal." METHOD: To illustrate challenges to data interpretation that may result from unclarity about the neuromodulatory effects of different field orientations, we present two high-definition tDCS experiments in the language domain, with different electrode configurations. We also present the modeled electric fields for a traditional tDCS setup, showing how brain stimulation may far exceed target regions. CONCLUSIONS: More research is warranted on the hypothesized inhibitory or excitatory effects of different electrode configurations. Moreover, conventional bicephalic 1 × 1 configurations using sponges or HD electrodes may not be accurately described by the terms "anodal" and "cathodal" either, as these terms only pertain to the desired effects over an area of interest, but not any other areas affected. Therefore, design and interpretation of (HD-)tDCS and conventional tDCS research studies should not be constrained by the anodal/cathodal dichotomy.

Simulating the neural correlates of stuttering.

For functional neuroimaging studies of stuttering, two challenges are (1) the elicitation of naturally stuttered versus fluent speech and (2) the separation of activation associated with abnormal motor execution from activation that reflects the cognitive substrates of stuttering. This paper reports on a proof-of-concept study, in which a single-subject approach was applied to address these two issues. A stuttering speaker used his insight into his own stuttering behavior to create a list of stutter-prone words versus a list of "fluent" words. He was then matched to a non-stuttering speaker, who imitated the specific articulatory and orofacial motor pattern of the stuttering speaker. Both study participants performed a functional MRI experiment of single word reading, each being presented with the same lexical items. Results suggest that the generally observed right-hemisphere lateralization appears to reflect a true neural correlate of stuttering. Some of the classically reported activation associated with stuttering appears to be driven more by nonspecific motor patterns than by cognitive substrates of stuttering, while anterior cingulate activation may reflect awareness of (upcoming) dysfluencies. This study shows that it is feasible to match stuttering speakers' utterances more closely to simulated stutters for the investigation of neural correlates of real stuttering. Significant main effects and contrast effects were obtained for the differences between fluent and stuttered speech, and right-hemisphere lateralization associated with real stuttered speech was shown in a single subject.

Network-based statistics distinguish anomic and Broca aphasia.

Aphasia is a speech-language impairment commonly caused by damage to the left hemisphere. Due to the complexity of speech-language processing, the neural mechanisms that underpin various symptoms between different types of aphasia are still not fully understood. We used the network-based statistic method to identify distinct subnetwork(s) of connections differentiating the resting-state functional networks of the anomic and Broca groups. We identified one such subnetwork that mainly involved the brain regions in the premotor, primary motor, primary auditory, and primary sensory cortices in both hemispheres. The majority of connections in the subnetwork were weaker in the Broca group than the anomic group. The network properties of the subnetwork were examined through complex network measures, which indicated that the regions in the superior temporal gyrus and auditory cortex bilaterally exhibit intensive interaction, and primary motor, premotor and primary sensory cortices in the left hemisphere play an important role in information flow and overall communication efficiency. These findings underlied articulatory difficulties and reduced repetition performance in Broca aphasia, which are rarely observed in anomic aphasia. This research provides novel findings into the resting-state brain network differences between groups of individuals with anomic and Broca aphasia. We identified a subnetwork of, rather than isolated, connections that statistically differentiate the resting-state brain networks of the two groups, in comparison with standard lesion symptom mapping results that yield isolated connections.

Grammatical Parallelism in Aphasia: A Lesion-Symptom Mapping Study.

Sentence structure, or syntax, is potentially a uniquely creative aspect of the human mind. Neuropsychological experiments in the 1970s suggested parallel syntactic production and comprehension deficits in agrammatic Broca's aphasia, thought to result from damage to syntactic mechanisms in Broca's area in the left frontal lobe. This hypothesis was sometimes termed overarching agrammatism, converging with developments in linguistic theory concerning central syntactic mechanisms supporting language production and comprehension. However, the evidence supporting an association among receptive syntactic deficits, expressive agrammatism, and damage to frontal cortex is equivocal. In addition, the relationship among a distinct grammatical production deficit in aphasia, paragrammatism, and receptive syntax has not been assessed. We used lesion-symptom mapping in three partially overlapping groups of left-hemisphere stroke patients to investigate these issues: grammatical production deficits in a primary group of 53 subjects and syntactic comprehension in larger sample sizes (N = 130, 218) that overlapped with the primary group. Paragrammatic production deficits were significantly associated with multiple analyses of syntactic comprehension, particularly when incorporating lesion volume as a covariate, but agrammatic production deficits were not. The lesion correlates of impaired performance of syntactic comprehension were significantly associated with damage to temporal lobe regions, which were also implicated in paragrammatism, but not with the inferior and middle frontal regions implicated in expressive agrammatism. Our results provide strong evidence against the overarching agrammatism hypothesis. By contrast, our results suggest the possibility of an alternative grammatical parallelism hypothesis rooted in paragrammatism and a central syntactic system in the posterior temporal lobe.

Network-based statistics distinguish anomic and Broca's aphasia.

INTRODUCTION: Aphasia is a speech-language impairment commonly caused by damage to the left hemisphere. The neural mechanisms that underpin different types of aphasia and their symptoms are still not fully understood. This study aims to identify differences in resting-state functional connectivity between anomic and Broca's aphasia measured through resting-state functional magnetic resonance imaging (rs-fMRI). METHODS: We used the network-based statistic (NBS) method, as well as voxel- and connectome-based lesion symptom mapping (V-, CLSM), to identify distinct neural correlates of the anomic and Broca's groups. To control for lesion effect, we included lesion volume as a covariate in both the NBS method and LSM. RESULTS: NBS identified a subnetwork located in the dorsal language stream bilaterally, including supramarginal gyrus, primary sensory, motor, and auditory cortices, and insula. The connections in the subnetwork were weaker in the Broca's group than the anomic group. The properties of the subnetwork were examined through complex network measures, which indicated that regions in right inferior frontal sulcus, right paracentral lobule, and bilateral superior temporal gyrus exhibit intensive interaction. Left superior temporal gyrus, right postcentral gyrus, and left supramarginal gyrus play an important role in information flow and overall communication efficiency. Disruption of this network underlies the constellation of symptoms associated with Broca's aphasia. Whole-brain CLSM did not detect any significant connections, suggesting an advantage of NBS when thousands of connections are considered. However, CLSM identified connections that differentiated Broca's from anomic aphasia when analysis was restricted to a hypothesized network of interest. DISCUSSION: We identified novel signatures of resting-state brain network differences between groups of individuals with anomic and Broca's aphasia. We identified a subnetwork of connections that statistically differentiated the resting-state brain networks of the two groups, in comparison with standard CLSM results that yielded isolated connections. Network-level analyses are useful tools for the investigation of the neural correlates of language deficits post-stroke.

Network modulation during complex syntactic processing.

Complex sentence processing is supported by a left-lateralized neural network including inferior frontal cortex and posterior superior temporal cortex. This study investigates the pattern of connectivity and information flow within this network. We used fMRI BOLD data derived from 12 healthy participants reported in an earlier study (Thompson, C. K., Den Ouden, D. B., Bonakdarpour, B., Garibaldi, K., & Parrish, T. B. (2010b). Neural plasticity and treatment-induced recovery of sentence processing in agrammatism. Neuropsychologia, 48(11), 3211-3227) to identify activation peaks associated with object-cleft over syntactically less complex subject-cleft processing. Directed Partial Correlation Analysis was conducted on time series extracted from participant-specific activation peaks and showed evidence of functional connectivity between four regions, linearly between premotor cortex, inferior frontal gyrus, posterior superior temporal sulcus and anterior middle temporal gyrus. This pattern served as the basis for Dynamic Causal Modeling of networks with a driving input to posterior superior temporal cortex, which likely supports thematic role assignment, and networks with a driving input to inferior frontal cortex, a core region associated with syntactic computation. The optimal model was determined through both frequentist and Bayesian Model Selection and turned out to reflect a network with a primary drive from inferior frontal cortex and modulation of the connection between inferior frontal cortex and posterior superior temporal cortex by complex sentence processing. The winning model also showed a substantive role for a feedback mechanism from posterior superior temporal cortex back to inferior frontal cortex. We suggest that complex syntactic processing is driven by word-order analysis, supported by inferior frontal cortex, in an interactive relation with posterior superior temporal cortex, which supports verb argument structure processing.

Automatic Subtyping of Individuals with Primary Progressive Aphasia.

BACKGROUND: The classification of patients with primary progressive aphasia (PPA) into variants is time-consuming, costly, and requires combined expertise by clinical neurologists, neuropsychologists, speech pathologists, and radiologists. OBJECTIVE: The aim of the present study is to determine whether acoustic and linguistic variables provide accurate classification of PPA patients into one of three variants: nonfluent PPA, semantic PPA, and logopenic PPA. METHODS: In this paper, we present a machine learning model based on deep neural networks (DNN) for the subtyping of patients with PPA into three main variants, using combined acoustic and linguistic information elicited automatically via acoustic and linguistic analysis. The performance of the DNN was compared to the classification accuracy of Random Forests, Support Vector Machines, and Decision Trees, as well as to expert clinicians' classifications. RESULTS: The DNN model outperformed the other machine learning models as well as expert clinicians' classifications with 80% classification accuracy. Importantly, 90% of patients with nfvPPA and 95% of patients with lvPPA was identified correctly, providing reliable subtyping of these patients into their corresponding PPA variants. CONCLUSION: We show that the combined speech and language markers from connected speech productions can inform variant subtyping in patients with PPA. The end-to-end automated machine learning approach we present can enable clinicians and researchers to provide an easy, quick, and inexpensive classification of patients with PPA.

Agrammatism and Paragrammatism: A Cortical Double Dissociation Revealed by Lesion-Symptom Mapping.

The fundamental distinction of grammatical deficits in aphasia, agrammatism and paragrammatism, was made over a century ago. However, the extent to which the agrammatism/paragrammatism distinction exists independently of differences in speech fluency has not clearly been investigated. Despite much research on agrammatism, the lesion correlates of paragrammatism are essentially unknown. Lesion-symptom mapping was used to investigate the degree to which the lesion correlates of agrammatism and paragrammatism overlap or dissociate. Four expert raters assessed videos of 53 right-handed patients with aphasia following chronic left-hemisphere stroke retelling the Cinderella story. Consensus discussion determined each subject's classification with respect to grammatical deficits as Agrammatic, Paragrammatic, Both, or No Grammatical Deficit. Each subject's lesion was manually drawn on a high-resolution MRI and warped to standard space for group analyses. Lesion-symptom mapping analyses were performed in NiiStat including lesion volume as a covariate. Secondary analyses included speech rate (words per minute) as an additional covariate. Region of interest analyses identified a double dissociation between these syndromes: damage to Broca's area was significantly associated with agrammatism, p = 0.001 (but not paragrammatism, p = 0.930), while damage to the left posterior superior and middle temporal gyri was significantly associated with paragrammatism, p < 0.001 (but not agrammatism, p = 0.873). The same results obtained when regressing out the effect of speech rate, and nonoverlapping lesion distributions between the syndromes were confirmed by uncorrected whole brain analyses. Our results support a fundamental distinction between agrammatism and paragrammatism.

Argument structure effects in action verb naming in static and dynamic conditions.

Argument structure, as in the participant roles entailed within the lexical representation of verbs, affects verb processing. Recent neuroimaging studies show that when verbs are heard or read, the posterior temporoparietal region shows increased activation for verbs with greater versus lesser argument structure complexity, usually bilaterally. In addition, patients with agrammatic aphasia show verb production deficits, graded based on argument structure complexity. In the present study, we used fMRI to examine the neural correlates of verb production in overt action naming conditions. In addition, we tested the differential effects of naming when verbs were presented dynamically in video segments versus statically in line drawings. Results showed increased neuronal activity associated with production of transitive as compared to intransitive verbs not only in posterior regions, but also in left inferior frontal cortex. We also found significantly greater activation for transitive versus intransitive action naming for videos compared to pictures in the right inferior and superior parietal cortices, areas associated with object manipulation. These findings indicate that verbs with greater argument structure density engender graded activation of both anterior and posterior portions of the language network and support verb naming deficit patterns reported in lesion studies. In addition, the similar findings derived under video and static picture naming conditions provide validity for using videos in neuroimaging studies, which are more naturalistic and perhaps ecologically valid than using static pictures to investigate action naming.

The electrophysiological manifestation of Dutch Verb Second violations.

We investigated the processing of violations of the verb position in Dutch, in a group of healthy subjects, by measuring event-related potentials (ERPs) through electroencephalography (EEG). In Dutch, the base position of the verb is clause final, but in matrix clauses, the finite verb is in second position, a construction known as Verb Second. In embedded clauses, the finite verb remains in its clause-final base position. The results show that ungrammatical placement of finite verbs in second position in embedded clauses yields a P600 response, which suggests that the parser treats this type of violation as a clear syntactic anomaly. This is in contrast to accounts by which a general preference for subject-verb-object word order in languages like Dutch is reflected by an absence of P600 effects in response to violations of Verb Second.