Previous research indicates that, under explicit instructions to listen to spoken stimuli or in speech-oriented behavioural tasks, the brain's responses to senseless pseudowords are larger than those to meaningful words; the reverse is true in non-attended conditions. These differential responses could be used as a tool to trace linguistic processes in the brain and their interaction with attention. However, as previous studies relied on explicit instructions to attend or ignore the stimuli, a technique for automatic attention modulation (i.e., not dependent on explicit instruction) would be more advantageous, especially when cooperation with instructions may not be guaranteed (e.g., neurological patients, children etc). Here we present a novel paradigm in which the stimulus context automatically draws attention to speech. In a non-attend passive auditory oddball sequence, rare words and pseudowords were presented among frequent non-speech tones of variable frequency and length. The low percentage of spoken stimuli guarantees an involuntary attention switch to them. The speech stimuli, in turn, could be disambiguated as words or pseudowords only in their end, at the last phoneme, after the attention switch would have already occurred. Our results confirmed that this paradigm can indeed be used to induce automatic shifts of attention to spoken input. At ~250ms after the stimulus onset, a P3a-like neuromagnetic deflection was registered to spoken (but not tone) stimuli indicating an involuntary attention shift. Later, after the word-pseudoword divergence point, we found a larger oddball response to pseudowords than words, best explained by neural processes of lexical search facilitated through increased attention. Furthermore, we demonstrate a breakdown of this orderly pattern of neurocognitive processes as a result of sleep deprivation. The new paradigm may thus be an efficient way to assess language comprehension processes and their dynamic interaction with those of attention allocation. It does it in an automatic and task-free fashion, indicating its potential benefit for assessing uncooperative clinical populations.
Object identification is enabled through a distributed neural network but the relative contribution of the single components of this network is largely unknown. In the present study, we used online interference by repetitive transcranial magnetic stimulation (rTMS) to investigate the role of the dorso-lateral prefrontal cortex (DLPFC) in identifying semantically different stimuli presented as to make the decision process easy or difficult, according to the amount of sensory information available. Nineteen healthy volunteers performed an object identification task. Stimuli belonging to living and non-living categories were presented at different levels of spatial filtering following a coarse-to-fine order that gradually integrated spatial information. Six-pulse trains of 10-Hz rTMS were delivered at an intensity of 90% resting motor threshold simultaneously to the picture presentation. rTMS of either the left or right DLPFC produced a significant lengthening in the identification process of spatially filtered living stimuli, as shown by the increase in the reaction time, but not of non-filtered living stimuli or of non-living objects. rTMS over the vertex did not interfere with the identification task. These data indicate that DLPFC role in the network underlying object recognition is more crucial when this neural process is challenged by the level of sensory information available to the observer. Specificity of this effect for living objects is discussed taking into account the crucial role of DLPFC in recruitment of cognitive resources for accomplishing perceptual decision-making.
Recent actions can benefit or disrupt our current actions and the prefrontal cortex (PFC) is thought to play a major role in the regulation of these actions before they occur. The left PFC has been associated with overcoming interference from past events in the context of language production and working memory. The right PFC, and especially the right IFG, has been associated with preparatory inhibition processes. But damage to the right PFC has also been associated with impairment in sustaining actions in motor intentional disorders. Moreover, bilateral dorsolateral PFC has been associated with the ability to maintain task-sets, and improve the performance of current actions based on previous experience. However, potential hemispheric asymmetries in anticipatory regulation of action have not yet been delineated. In the present study, patients with left (n=7) vs. right (n=6) PFC damage due to stroke and 14 aged- and education-matched controls performed a picture naming and a verbal Simon task (participants had to say "right" or "left" depending on the color of the picture while ignoring its position). In both tasks, performance depended on the nature of the preceding trial, but in different ways. In the naming task, performance decreased if previous pictures were from the same rather than from different semantic categories (i.e., semantic interference effect). In the Simon task, performance was better for both compatible (i.e., response matching the position of the stimulus) and incompatible trials when preceded by a trial of the same compatibility (i.e. Gratton effect) relative to sequential trials of different compatibility. Left PFC patients were selectively impaired in picture naming; they had an increased semantic interference effect compared to both right PFC patients and aged-matched controls. Conversely, right PFC patients were selectively impaired in the Simon task compared to controls or left PFC patients; they showed no benefit when sequential trials were compatible (cC vs. iC trials) or a decreased Gratton effect. These results provide evidence for a double dissociation between left and right PFC in the anticipatory regulation of action. Our results are in agreement with a preponderant role of the left PFC in overcoming proactive interference from competing memory representations and provide evidence that the right PFC, plays a role in sustaining goal-directed actions consistent with clinical data in right PFC patients with motor intentional disorders.
Humans show variable degrees of success in acquiring a second language (L2). In many cases, morphological and syntactic knowledge remain deficient, although some learners succeed in reaching nativelike levels, even if they begin acquiring their L2 relatively late. In this study, we use psycholinguistic, online language proficiency tests and a neurophysiological index of syntactic processing, the syntactic mismatch negativity (sMMN) to local agreement violations, to compare behavioural and neurophysiological markers of grammar processing between native speakers (NS) of English and non-native speakers (NNS). Variable grammar proficiency was measured by psycholinguistic tests. When NS heard ungrammatical word sequences lacking agreement between subject and verb (e.g. * we kicks), the MMN was enhanced compared with syntactically legal sentences (e.g. he kicks). More proficient NNS also showed this difference, but less proficient NNS did not. The main cortical sources of the MMN responses were localised in bilateral superior temporal areas, where, crucially, source strength of grammar-related neuronal activity correlated significantly with grammatical proficiency of individual L2 speakers as revealed by the psycholinguistic tests. As our results show similar, early MMN indices to morpho-syntactic agreement violations among both native speakers and non-native speakers with high grammar proficiency, they appear consistent with the use of similar brain mechanisms for at least certain aspects of L1 and L2 grammars. © 2016 The Authors.
Despite a persistent interest in verb processing, data on the neural underpinnings of verb retrieval are frag- mentary. The present study is the first to analyze the contributions of both grey and white matter damage affecting verb retrieval through action naming in stroke. We used voxel-based lesion-symptom mapping (VLSM) with an action naming task in 40 left-hemisphere stroke patients. Within the grey matter, we revealed the critical involvement of the left precentral and inferior frontal gyri, insula, and parts of basal ganglia. An overlay of white matter tract probability masks on the VLSM lesion map revealed involvement of left-hemisphere long and short association tracts with terminations in the frontal areas; and several projection tracts. The involvement of these structures is interpreted in the light of existing picture naming models, semantic control processes, and the embodiment cognition framework. Our results stress the importance of both cortico-cortical and cortico-sub- cortical networks of language processing.
The extent to which action and perception systems of the brain are involved in semantic comprehension remains controversial. Whether figurative language, such as metaphors and idioms, is grounded in sensory-motor systems is especially contentious. Here, we used high-definition transcranial direct current stimulation (HD-tDCS) in healthy adults to examine the role of the left-hemisphere motor cortex during the comprehension of action sentences, relative to comprehension of sentences with visual verbs. Action sentences were divided into three types: literal, metaphoric, or idiomatic. This allowed us to ask whether processing of action verbs used in figurative contexts relies on motor cortex. The results revealed that action sentence comprehension response times were facilitated relative to the visual sentence control. Significant interaction relative to visual sentences was observed for literal, metaphoric, and idiomatic action sentences with HD-tDCS of the motor cortex. These results suggest that the left motor cortex is functionally involved in action sentence comprehension. Furthermore, this involvement exists when the action content of the sentences is figurative, for both idiomatic and metaphoric cases. The results provide evidence for functional links between conceptual and action systems of the brain.
Previous studies have demonstrated that efficient neurorehabilitation in post stroke aphasia leads to clinical language improvements and promotes neuroplasticity. Brain areas frequently implicated in functional restitution of language after stroke comprise perilesional sites in the left hemisphere and homotopic regions in the right hemisphere. However, the neuronal mechanisms underlying therapy-induced language restitution are still largely unclear. In this study, magnetoencephalography was used to investigate neurophysiological changes in a group of chronic aphasia patients who underwent intensive language action therapy (ILAT), also known as constraint-induced aphasia therapy (CIAT). Before and immediately after ILAT, patients’ language and communication skills were assessed and their brain responses were recorded during a lexical magnetic mismatch negativity (MMNm) paradigm, presenting familiar spoken words and meaningless pseudowords. After the two-week therapy interval, patients showed significant clinical improvements of language and communication skills. Spatio-temporal dynamics of neuronal changes revealed a significant increase in word-specific neuro-magnetic MMNm activation around 200 ms after stimulus identification points. This enhanced brain response occurred specifically for words and was most pronounced over perilesional areas in the left hemisphere. Therapy-related changes in neuromagnetic activation for words in both hemispheres significantly correlated with performance on a clinical language test. The findings indicate that functional recovery of language in chronic post stroke aphasia is associated with neuroplastic changes in both cerebral hemispheres, with stronger left-hemispheric contribution during automatic stages of language processing.
Implicit-statistical learning (ISL) research investigates whether domain-general mechanisms are recruited in the linguistic processes that require manipulation of patterned regularities (e.g. syntax). Aphasia is a language disorder caused by focal brain damage in the left fronto-temporal-parietal network. Research shows that people with aphasia (PWA) with frontal lobe lesions manifest convergent deficits in syntax and ISL mechanisms. So far, ISL mechanisms in PWA with temporal or parietal lobe lesions have not been systematically investigated.
We investigated two complementary hypotheses: 1) the anatomical hypothesis, that PWA with frontal lesions display more severely impaired ISL abilities than PWA with posterior lesions and 2) the behavioural hypothesis, that the magnitude of impairment in ISL mechanisms correlates to syntactic deficits in aphasia.
We tested 13 PWA, 5 with frontal lesions and 8 with posterior lesions, and 11 non-brain-damaged controls on a visual statistical learning (VSL) task. In addition, all PWA completed several linguistic tasks. Reaction times, obtained in the VSL task, were analyzed using linear mixed-effects model. Correlational statistics were used to assess the relationship between VSL task performance and linguistic measures.
Results and Discussion
We did not find support for the anatomical hypothesis as patients with spared frontal regions also manifested impaired ISL mechanisms. This is attributed to a) ISL mechanisms being vulnerable to other cognitive dysfunctions and/or b) ISL mechanisms anatomically extending to the posterior brain regions. Notably, ISL mechanisms were impaired, but not absent in aphasia. With regards to the behavioural hypothesis, we provide empirical evidence of correlation between ISL mechanisms and syntactic, but not lexical impairment in aphasia. We discuss both the theoretical contributions to the debate of domain-independence of ISL mechanisms and clinical implications for implicit language therapy.
The neural mechanisms underlying perceptual learning are still under investigation. Eureka effect is a form of rapid, long-lasting perceptual learning by which a degraded image, which appears meaningless when first seen, becomes recognizable after a single exposure to its undegraded version. We used online interference by focal 10-Hz repetitive transcranial magnetic stimulation (rTMS) to evaluate whether the parietal cortex (PC) is involved in Eureka effect, as suggested by neuroimaging data. RTMS of the PC did not affect recognition of degraded pictures when displayed 2s after the presentation of their undegraded version (learning phase). However, rTMS delivered over either right or left intraparietal sulcus simultaneously to the undegraded image presentation, disrupted identification of the degraded version of the same pictures when displayed 30 min after the learning phase. In contrast, recognition of degraded images was unaffected by rTMS over the vertex or by sham rTMS, or when rTMS of either PC was delivered 2s after the presentation of the undegraded image. Findings strongly support the hypothesis that both PC at the level of the intraparietal sulcus play a pivotal role in the Eureka effect particularly in consolidation processes, and contribute to elucidate the neural network underlying rapid perceptual learning
The frontal aslant tract (FAT) is a white-matter tract connecting the inferior frontal gyrus (IFG) and the supplementary motor complex (SMC). Damage to either component of the network causes spontaneous speech dysfluency, indicating its critical role in language production. However, spontaneous speech dysfluency may stem from various lower-level linguistic deficits, precluding inferences about the nature of linguistic processing subserved by the IFG-SMC network. Since the IFG and the SMC are attributed a role in conceptual and lexical selection during language production, we hypothesized that these processes rely on the IFG-SMC connectivity via the FAT. We analysed the effects of FAT volume on conceptual and lexical selection measures following frontal lobe stroke. The measures were obtained from the sentence completion (SC) task, tapping into conceptual and lexical selection, and the picture-word interference (PWI) task, providing a more specific measure of lexical selection. Lower FAT volume was not associated with lower conceptual or lexical selection abilities in our patient cohort. Current findings stand in marked discrepancy with previous lesion and neuroimaging evidence for the joint contribution of the IFG and the SMC to lexical and conceptual selection. A plausible explanation reconciling this discrepancy is that the IFG-SMC connectivity via the FAT does contribute to conceptual and/or lexical selection but its disrupted function undergoes reorganisation over the course of post-stroke recovery. Thus, our negative findings stress the importance of testing the causal role of the FAT in lexical and conceptual selection in patients with more acute frontal lobe lesions.
Currently, a distributed bilateral network of frontal-parietal areas is regarded as the neural substrate of working memory (WM), with the verbal WM network being more left-lateralized. This conclusion is based primarily on functional magnetic resonance imaging (fMRI) data that provides correlational evidence for brain regions involved in a task. However, fMRI cannot differentiate the areas that are fundamentally required for performing a task. These data can only come from brain-injured individuals who fail the task after the loss of specific brain areas. In addition to the lack of complimentary data, is the issue of the variety in the WM tasks used to assess verbal WM. When different tasks are assumed to measure the same behavior, this may mask the contributions of different brain regions. Here, we investigated the neural substrate of WM by using voxel-based lesion symptom mapping (VLSM) in 49 individuals with stroke-induced left hemisphere brain injuries. These participants completed two different verbal WM tasks: complex listening span and a word 2-back task. Behavioral results indicated that the two tasks were only slightly related, while the VLSM analysis revealed different critical regions associated with each task. Specifically, significant detriments in performance on the complex span task were found with lesions in the inferior frontal gyrus, while for the 2-back task, significant deficits were seen after injury to the superior and middle temporal gyri. Thus, the two tasks depend on the structural integrity of different, non-overlapping frontal and temporal brain regions, suggesting distinct neural and cognitive mechanisms triggered by the two tasks: rehearsal and cue-dependent selection in the complex span task, versus updating/auditory recognition in the 2-back task. These findings call into question the common practice of using these two tasks interchangeably in verbal WM research and undermine the legitimacy of aggregating data from studies with different WM tasks. Thus, the present study points out the importance of lesion studies in complementing functional neuroimaging findings and highlights the need to consider task demands in neuroimaging and neuropsychological investigations of WM.
Among various questions pertinent to grounding human cognitive functions in a neurobiological substrate, the association between language and motor brain structures is a particularly debated one in neuroscience and psychology. While many studies support a broadly distributed model of language and semantics grounded, among other things, in the general modality-specific systems, theories disagree as to whether motor and sensory cortex activity observed during language processing is functional or epiphenomenal. Here, we assessed the role of motor areas in linguistic processing by investigating the responses of 28 healthy volunteers to different word types in semantic and lexical decision tasks, following repetitive transcranial magnetic stimulation (rTMS) of primary motor cortex. We found that early rTMS (delivered within 200ms of word onset) produces a left-lateralised and meaning-specific change in reaction speed, slowing down behavioural responses to action-related words, and facilitating abstract words - an effect present only during semantic, but not lexical, decision. We interpret these data in light of action-perception theory of language, bolstering the claim that motor cortical areas play a functional role in language comprehension.
The purpose of the present study was to identify general and syndrome-specific deficits in the lexical processing of individuals with non-fluent and fluent aphasia compared to individuals without cognitive, neurological or language impairments. The time course of lexical access, as well as lexical selection and integration was studied using a visual-world paradigm in three groups of Russian speakers: 36 individuals in the control group, 15 individuals with non-fluent aphasia and eight individuals with fluent aphasia. Participants listened to temporarily ambiguous sentences wherein the context biased the interpretation of an ambiguous word toward one of its two meanings. In half of the experimental sentences, a reanalysis was needed upon encountering the disambiguating phrase. The effect of the length of the intervening material between the ambiguous word and the disambiguation point was additionally monitored. All groups of participants showed intact lexical access under slowed speech rate, but non-fluent participants experienced difficulties with timely activation of multiple referents. At later stages of lexical processing, they additionally demonstrated a specific impairment of reanalysis. The deficit in participants with fluent aphasia was not focalized at any specific stage of lexical processing. Rather, the breakdown of lexical processes in fluent aphasia was likely related to difficulties with the inhibition of irrelevant lexical activation, which is further supported by the finding that increased phonological distance between the ambiguous word and ambiguity resolution was influential to the offline performance in this group.
The goal of the present study was to investigate event-related potential (ERP) responses to Dutch negative and positive polarity adverbs of degree presented in licensed and unlicensed contexts with negative and affirmative particles directly preceding the polarity item. To control for effects of the processing of negation as such, neutral adverbs were also presented in negative and affirmative contexts. The results did not show any significant effect of negation for the non-polar adverbs, allowing context effects for polarity items to be interpreted as being due to the appropriateness of the context. Negative polarity violations elicited an N400 response that might reflect the lack of semantic congruity of the negative polarity item in an affirmative context. In contrast, processing positive polarity items in context of negation resulted in a positive effect resembling the P600, which may be considered as a marker of a different sort of integration difficulty caused by violation of licensing conditions and/or a search for a licensor in the wider discourse context. The study presented here is the first to show an unambiguous dissociation between responses to negative and positive polarity violations. This dissociation argues for different mechanisms underlying the processing of these two types of polarity; we propose that positive polarity items are sensitive to wider discourse context, while negative polarity items are more sensitive to local lexical context.
especially impaired on regular past-tense forms like played, whether the task requires production, comprehension or even the judgement that "play" and "played" sound different. Within a dual-mechanism account of inflectional morphology, these deficits reflect disruption to the rule-based process that adds (or strips) the suffix -ed to regular verb stems; but the fact that the patients are also impaired at detecting the difference between word pairs like "tray" and "trade" (the latter being a phonological but not a morphological twin to "played") suggests an important role for phonological characteristics of the regular past tense. The present study examined MEG brain responses in healthy participants evoked by spoken regular past-tense forms and phonological twin words (plus twin pseudowords and a non-speech control) presented in a passive oddball paradigm. Deviant forms (played, trade, kwade/kwayed) relative to their standards (play, tray, kway) elicited a pronounced neuromagnetic response at approximately 130 ms after the onset of the affix; this response was maximal at sensors over temporal areas of both hemispheres but stronger on the left, especially for played and kwayed. Relative to the same standards, a different set of deviants ending in /t/--plate, trait and kwate--produced stronger difference responses especially over the right hemisphere. Results are discussed with regard to dual- and single-mechanism theories of past tense processing and the need to consider neurobiological evidence in attempts to understand inflectional morphology.
Background: Agrammatic speakers have problems with grammatical encoding and decoding. However, not all syntactic processes are equally problematic: present time reference, who questions, and reflexives can be processed by narrow syntax alone and are relatively spared compared to past time reference, which questions, and personal pronouns, respectively. The latter need additional access to discourse and information structures to link to their referent outside the clause (Avrutin, 2006). Linguistic processing that requires discourse linking is difficult for agrammatic individuals: verb morphology with reference to the past is more difficult than with reference to the present (Bastiaanse et al., 2011). The same holds for which questions compared to who questions and for pronouns compared to reflexives (Avrutin, 2006). These results have been reported independently for different populations in different languages. The current study, for the first time, tested all conditions within the same population.
Aims: We had two aims with the current study. Firstly, we wanted to investigate whether discourse linking is the common denominator of the deficits in time reference, wh-questions, and object pronouns. Secondly, we aimed to compare the comprehension of discourse-linked elements in people with agrammatic and fluent aphasia, to define the degree of the discourse linking deficit specificity.
Methods & Procedures: Three sentence-picture-matching tasks were administered to 10 agrammatic, 10 fluent aphasic, and 10 non-brain-damaged Russian speakers (NBDs): (1) the Test for Assessing Reference of Time (TART) for present imperfect (reference to present) and past perfect (reference to past), (2) the Wh-Extraction Assessment Tool (WHEAT) for which and who subject questions, and (3) the Reflexive-Pronoun Test (RePro) for reflexive and pronominal reference.
Outcomes & Results: NBDs scored at ceiling and significantly higher than the aphasic participants. We found an overall effect of discourse linking in the TART and WHEAT for the agrammatic speakers, and in all three tests for the fluent speakers. Scores on the RePro were at ceiling.
Conclusions: The problems that individuals with agrammatic and fluent aphasia experience when comprehending sentences that contain verbs with past time reference, which question words and pronouns are caused by the fact that these elements need to involve operations at the discourse level. The effect is not specific to agrammatism, although might result from different underlying disorders in agrammatic and fluent aphasia.