The cortical interactions in short time intervals during the search for verbal associations

Cortical connectivity was studied in tasks of generating the use of words in comparison with reading aloud the same words. These tasks were used earlier in PET and high-density ERP recording studies, which described both the functional anatomy and time course of involvement of cortical areas in word processing. We developed a new method for studying the synchrony of EEG spectral components within the short time intervals compatible with the duration of particular cognitive operations. The wavelet transform of the ERP records and calculation of correlations between the wavelet curves were used to reveal connections between cortical areas. Three stages of intracortical communications developing over the course of task performance were discovered: between the right and left frontal areas (0-200 ms after the stimulus presentation), between the left frontal and left posterior temporo-parietal areas (250-500 ms), and, finally, between the left temporal and right fronto-centro-temporal areas. These findings are in good agreement with the results of the previous PET and ERP studies and supplement them with the circuitry of cortical information transfer. Also, they suggest some differences in information processing during automated reading and performance of more complicated use-generation task.     

Temporal characteristics of the initial stage of the verbal information processing in healthy subjects and in schizophrenia

The study aimed to investigate the early coding of visually presented words and pseudowords using event-related potentials (ERP). We conducted comparative analysis of the characteristics of P100 and N170 in healthy controls and in patients with the first episode of schizophrenia during passive perception of verbal stimuli as well as under conditions of relevant words and pseudowords. The latency of early ERP components P100 and N170 appeared to be shorter in comparison with healthy subjects in the temporal, parietal and occipital areas. The latency of P100 in patients was significantly shorter in the temporal, parietal and occipital areas, whereas the latency of N170 was shorter in the parietal and occipital areas than in controls. The latency of N170 in healthy subjects was significantly longer to words than to pseudowords and in patients - vice versa. The latencies of N170 in all TPO areas were equal in healthy subjects during word processing, and this equality was upset during non-word processing. In patients with schizophrenia the equality was upset, but, opposite to healthy patients, the upset of equality was more expressed during words processing. Thus, the early stage of verbal information processing in schizophrenic patients is insufficient in time. The time deficit of the automatic processes may lead to defective processing of overall information.     

Functional interactions of the inferior frontal cortex during the processing of words and word-like stimuli

The hypothesis that ventral/anterior left inferior frontal gyrus (LIFG) subserves semantic processing and dorsal/posterior LIFG subserves phonological processing was tested by determining the pattern of functional connectivity of these regions with regions in left occipital and temporal cortex during the processing of words and word-like stimuli. In accordance with the hypothesis, we found strong functional connectivity between activity in ventral LIFG and activity in occipital and temporal cortex only for words, and strong functional connectivity between activity in dorsal LIFG and activity in occipital and temporal cortex for words, pseudowords, and letter strings, but not for false font strings. These results demonstrate a task-dependent functional fractionation of the LIFG in terms of its functional links with posterior brain areas.     

Distinct Effects of Memory Retrieval and Articulatory Preparation when Learning and Accessing New Word Forms

Temporal and frontal activations have been implicated in learning of novel word forms, but their specific roles remain poorly understood. The present magnetoencephalography (MEG) study examines the roles of these areas in processing newly-established word form representations. The cortical effects related to acquiring new phonological word forms during incidental learning were localized. Participants listened to and repeated back new word form stimuli that adhered to native phonology (Finnish pseudowords) or were foreign (Korean words), with a subset of the stimuli recurring four times. Subsequently, a modified 1-back task and a recognition task addressed whether the activations modulated by learning were related to planning for overt articulation, while parametrically added noise probed reliance on developing memory representations during effortful perception. Learning resulted in decreased left superior temporal and increased bilateral frontal premotor activation for familiar compared to new items. The left temporal learning effect persisted in all tasks and was strongest when stimuli were embedded in intermediate noise. In the noisy conditions, native phonotactics evoked overall enhanced left temporal activation. In contrast, the frontal learning effects were present only in conditions requiring overt repetition and were more pronounced for the foreign language. The results indicate a functional dissociation between temporal and frontal activations in learning new phonological word forms: the left superior temporal responses reflect activation of newly-established word-form representations, also during degraded sensory input, whereas the frontal premotor effects are related to planning for articulation and are not preserved in noise.     

Background Speech Effects on Sentence Processing during Reading: An Eye Movement Study

Effects of background speech on reading were examined by playing aloud different types of background speech, while participants read long, syntactically complex and less complex sentences embedded in text. Readers’ eye movement patterns were used to study online sentence comprehension. Effects of background speech were primarily seen in rereading time. In Experiment 1, foreign-language background speech did not disrupt sentence processing. Experiment 2 demonstrated robust disruption in reading as a result of semantically and syntactically anomalous scrambled background speech preserving normal sentence-like intonation. Scrambled speech that was constructed from the text to-be read did not disrupt reading more than scrambled speech constructed from a different, semantically unrelated text. Experiment 3 showed that scrambled speech exacerbated the syntactic complexity effect more than coherent background speech, which also interfered with reading. Experiment 4 demonstrated that both semantically and syntactically anomalous speech produced no more disruption in reading than semantically anomalous but syntactically correct background speech. The pattern of results is best explained by a semantic account that stresses the importance of similarity in semantic processing, but not similarity in semantic content, between the reading task and background speech.     

Context-dependent EEG spectral characteristics during performance of tasks

Spatial and frequency EEG characteristics of two groups of healthy adult subjects were examined in two series of experiments, which differed in conditions of the second cognitive task in a trial. The first task was the same in the two series: subjects had to evaluate size relationship between two closely spaced circles. The second task successively presented in trials of the first series consisted in the recognition of words/pseudowords, and in the second series, subjects had to localize a target letter in a matrix. It was assumed that the cognitive performance in the first series predominantly involved the ventral visual system, whereas during task performance in the second series, predominant involvement of the ventral and dorsal visual systems alternated. Multichannel EEG fragments recorded prior to the presentation of the task pairs were analyzed. Analysis of variance of the EEG spectral power revealed the generalized significant effect of the factor of the second task in the pair for delta band and lower beta subband, the power being higher in the first series. Factor brain hemisphere had a significant effect for the alpha band in the occipital area, the spectral power being lower in the left hemisphere for both experimental series. The task x hemisphere interaction was significant in the temporal cortical areas for the EEG power in alpha2 band, i.e., the predominant involvement of the ventral visual system was associated with stronger asymmetry of alpha2 rhythm and lower spectral power in this band in the left temporal area. Thus, the character of the forthcoming cognitive activity was shown to be reflected in spatio-frequency characteristics of the preceding EEG.     

Segregation of Lexical and Sub-Lexical Reading Processes in the Left Perisylvian Cortex

A fundamental issue in cognitive neuroscience is the existence of two major, sub-lexical and lexical, reading processes and their possible segregation in the left posterior perisylvian cortex. Using cortical electrostimulation mapping, we identified the cortical areas involved on reading either orthographically irregular words (lexical, “direct” process) or pronounceable pseudowords (sublexical, “indirect” process) in 14 right-handed neurosurgical patients while video-recording behavioral effects. Intraoperative neuronavigation system and Montreal Neurological Institute (MNI) stereotactic coordinates were used to identify the localization of stimulation sites. Fifty-one reading interference areas were found that affected either words (14 areas), or pseudo-words (11 areas), or both (26 areas). Forty-one (80%) corresponded to the impairment of the phonological level of reading processes. Reading processes involved discrete, highly localized perisylvian cortical areas with individual variability. MNI coordinates throughout the group exhibited a clear segregation according to the tested reading route; specific pseudo-word reading interferences were concentrated in a restricted inferior and anterior subpart of the left supramarginal gyrus (barycentre x = −68.1; y = −25.9; z = 30.2; Brodmann’s area 40) while specific word reading areas were located almost exclusively alongside the left superior temporal gyrus. Although half of the reading interferences found were nonspecific, the finding of specific lexical or sublexical interferences is new evidence that lexical and sublexical processes of reading could be partially supported by distinct cortical sub-regions despite their anatomical proximity. These data are in line with many brain activation studies that showed that left superior temporal and inferior parietal regions had a crucial role respectively in word and pseudoword reading and were core regions for dyslexia.     

A complementary systems account of word learning: neural and behavioural evidence

In this paper we present a novel theory of the cognitive and neural processes by which adults learn new spoken words. This proposal builds on neurocomputational accounts of lexical processing and spoken word recognition and complementary learning systems (CLS) models of memory. We review evidence from behavioural studies of word learning that, consistent with the CLS account, show two stages of lexical acquisition: rapid initial familiarization followed by slow lexical consolidation. These stages map broadly onto two systems involved in different aspects of word learning: (i) rapid, initial acquisition supported by medial temporal and hippocampal learning, (ii) slower neocortical learning achieved by offline consolidation of previously acquired information. We review behavioural and neuroscientific evidence consistent with this account, including a meta-analysis of PET and functional Magnetic Resonance Imaging (fMRI) studies that contrast responses to spoken words and pseudowords. From this meta-analysis we derive predictions for the location and direction of cortical response changes following familiarization with pseudowords. This allows us to assess evidence for learning-induced changes that convert pseudoword responses into real word responses. Results provide unique support for the CLS account since hippocampal responses change during initial learning, whereas cortical responses to pseudowords only become word-like if overnight consolidation follows initial learning.     

Hemispheric asymmetry of visual evoked potentials during human recognition of facial emotional expressions

Visual evoked potentials (VEP) in standard 16 EEG derivations were recorded in 26 young men and 20 women during recognition of facial emotional expressions and geometric figures. The stimuli were presented on a computer screen in the center of the visual field or randomly in the right or left vision hemifields. Peak VEP latency and mean amplitude in 50-ms epochs were measured; spatiotemporal VEP dynamics was analyzed in a series of topographic maps. The right hemisphere was shown to be more important in processing emotional faces. The character of the asymmetry was dynamic: at earlier stages of emotion processing the electrical activity was higher in the right inferior temporal region compared to the left symmetrical site. Later on the activity was higher in the right frontal and central areas. The dynamic mapping of "face-selective" component N180 of VEPs revealed the onset of activation over the right frontal areas that was followed by the fast activation of symmetrical left zones. Notably, this dynamics didn't correlate with the hemifield of stimuli exposition. The degree of asymmetry was lower during presentation of figures, especially in the inferior temporal and frontal regions. The prominent asymmetry of information processes in the inferior temporal and frontal areas was suggested to be specific for recognition of facial expression.     

Dispersed activation in the left temporal cortex for speech-reading in congenitally deaf people

Does the lateral temporal cortex require acoustic exposure in order to become specialized for speech processing? Six hearing participants and six congenitally deaf participants, all with spoken English as their first langugage, were scanned using functional magnetic resonance imaging while performing a simple speech-reading task. Focal activation of the left lateral temporal cortex was significantly reduced in the deaf group compared with the hearing group. Activation within this region was present in individual deaf participants, but varied in location from person to person. Early acoustic experience may be required for regions within the left temporal cortex in order to develop into a coherent network with subareas devoted to specific speech analysis functions.     

Brain Bases of Reading Fluency in Typical Reading and Impaired Fluency in Dyslexia

Although the neural systems supporting single word reading are well studied, there are limited direct comparisons between typical and dyslexic readers of the neural correlates of reading fluency. Reading fluency deficits are a persistent behavioral marker of dyslexia into adulthood. The current study identified the neural correlates of fluent reading in typical and dyslexic adult readers, using sentences presented in a word-by-word format in which single words were presented sequentially at fixed rates. Sentences were presented at slow, medium, and fast rates, and participants were asked to decide whether each sentence did or did not make sense semantically. As presentation rates increased, participants became less accurate and slower at making judgments, with comprehension accuracy decreasing disproportionately for dyslexic readers. In-scanner performance on the sentence task correlated significantly with standardized clinical measures of both reading fluency and phonological awareness. Both typical readers and readers with dyslexia exhibited widespread, bilateral increases in activation that corresponded to increases in presentation rate. Typical readers exhibited significantly larger gains in activation as a function of faster presentation rates than readers with dyslexia in several areas, including left prefrontal and left superior temporal regions associated with semantic retrieval and semantic and phonological representations. Group differences were more extensive when behavioral differences between conditions were equated across groups. These findings suggest a brain basis for impaired reading fluency in dyslexia, specifically a failure of brain regions involved in semantic retrieval and semantic and phonological representations to become fully engaged for comprehension at rapid reading rates.     

Spatiotemporal dynamics of modality-specific and supramodal word processing

The ability of written and spoken words to access the same semantic meaning provides a test case for the multimodal convergence of information from sensory to associative areas. Using anatomically constrained magnetoencephalography (aMEG), the present study investigated the stages of word comprehension in real time in the auditory and visual modalities, as subjects participated in a semantic judgment task. Activity spread from the primary sensory areas along the respective ventral processing streams and converged in anterior temporal and inferior prefrontal regions, primarily on the left at around 400 ms. Comparison of response patterns during repetition priming between the two modalities suggest that they are initiated by modality-specific memory systems, but that they are eventually elaborated mainly in supramodal areas.     

Argument Structure and Morphological Factors in Noun and Verb Processing: An fMRI Study

In a functional MRI (fMRI) study, we have investigated the grammatical categories of object noun, event noun and verb in order to assess the cortical regions of activation supporting their processing. Twelve Italian healthy participants performed a lexical decision task. They had to decide whether a string was an Italian word or not. Words could be objects like medaglia (medal), or events like the noun pianto (cry); or the verb dormire (to sleep). Noun and verb comparison shows differences in regions of activation in the left Inferior Frontal cortex and in the extent of the same areas. We have found specific areas of activation for object noun, and similarities in the pattern of activation for event noun and verb. The activations induced by pseudowords highly resembled the areas activated by the corresponding word category. The implications of the results are discussed in light of the recent debate on the role of grammatical category in the brain.     

Lexical context affects mismatch negativity caused by pseudowords

The aim of this study was to investigate the effect of lexical context on the latency and the amplitude of the mismatch negativity (MMN) brain potential caused by perception of pseudowords. The eventrelated potentials were recorded according to the multideviant passive odd-ball paradigm by using only pseudowords (control condition) or pseudowords with Russian words with different lexical frequencies (lexical context). It was found that different MMN patterns were generated when the same pseudoword was presented in different contexts. Pseudoword presentation in a context with other pseudowords resulted in a relatively small amplitude and large latency of MMN. If the same pseudoword was presented in a context with words, it induced significantly increased amplitude and reduced latency of MMN varying in the range of 100–200 ms. It is supposed that the pseudoword presented in a context with words is perceived as conceptually different stimulus, which leads to a significant increase in MMN. Moreover, our findings support the hypothesis that MMN is affected by lexical frequency. In particular, presentation of a high-frequency word induced a significantly more pronounced MMN response than a low-frequency one. High-frequency words also evoked earlier response, which indicates more rapid access to a frequently used lexical entry. More frequent use of certain words results in stronger internal connections in the corresponding memory circuit, which in turn is determined by the lexical context. We hypothesize that different intensities of activation depends on the strength of lexical representation.     

Interaction between the Frontal and Left Parietotemporal Areas in Verbal Thinking

The synchronization of the rhythmical components of evoked potentials (EP) was studied during verbal-task solving. A novel method of the calculation of Wavelet curve correlation was used to reveal synchronization between the evoked rhythmical components in short time intervals. This method was applied to earlier EP records, which were conducted during the search for verbal associations and revealed the successive activation of the frontal and left parietal cortical areas. Two stages of task solving were identified. Independently of the task type, the first stage was characterized by a diffuse synchronization in a broad frequency band below 22 Hz immediately after the stimulus presentation. This stage results in a realization of the verbal stimulus. The second stage was manifested in a localized synchronization between the frontal and left temporal (Wernicke's) areas in the narrow frequency band about 17 Hz only during search for associations. This specific and local synchronization took place earlier than the diffuse activation of the left temporal cortex. This stage appears to reflect the information transmission from the frontal cortex to the left parietotemporal area.     

Suggestion-Induced Modulation of Semantic Priming during Functional Magnetic Resonance Imaging

Using functional magnetic resonance imaging during a primed visual lexical decision task, we investigated the neural and functional mechanisms underlying modulations of semantic word processing through hypnotic suggestions aimed at altering lexical processing of primes. The priming task was to discriminate between target words and pseudowords presented 200 ms after the prime word which was semantically related or unrelated to the target. In a counterbalanced study design, each participant performed the task once at normal wakefulness and once after the administration of hypnotic suggestions to perceive the prime as a meaningless symbol of a foreign language. Neural correlates of priming were defined as significantly lower activations upon semantically related compared to unrelated trials. We found significant suggestive treatment-induced reductions in neural priming, albeit irrespective of the degree of suggestibility. Neural priming was attenuated upon suggestive treatment compared with normal wakefulness in brain regions supporting automatic (fusiform gyrus) and controlled semantic processing (superior and middle temporal gyri, pre- and postcentral gyri, and supplementary motor area). Hence, suggestions reduced semantic word processing by conjointly dampening both automatic and strategic semantic processes.     

Holistic processing of words modulated by reading experience

Perceptual expertise has been studied intensively with faces and object categories involving detailed individuation. A common finding is that experience in fulfilling the task demand of fine, subordinate-level discrimination between highly similar instances is associated with the development of holistic processing. This study examines whether holistic processing is also engaged by expert word recognition, which is thought to involve coarser, basic-level processing that is more part-based. We adopted a paradigm widely used for faces--the composite task, and found clear evidence of holistic processing for English words. A second experiment further showed that holistic processing for words was sensitive to the amount of experience with the language concerned (native vs. second-language readers) and with the specific stimuli (words vs. pseudowords). The adoption of a paradigm from the face perception literature to the study of expert word perception is important for further comparison between perceptual expertise with words and face-like expertise.     

Characteristics of the perception and imagination of emotions in patients with focal brain pathology

Voluntary components of the emotion perception were shown to be related to the left hemisphere temporal area in patients with focal lesions of postero-frontal and temporal areas of both hemispheres, whereas the involuntary components required involvement of the right hemisphere temporal area. The voluntary components of the emotion reproduction are associated with involvement of the left hemisphere postero-frontal area, whereas involuntary components of perception are related to work of the right hemisphere postero-frontal area. The data obtained suggest that the voluntary (conscious) recognition and reproduction of emotions are mainly related to the sensory and motor speech centres of the left hemisphere, whereas involuntary those involve symmetrical areas of the opposite hemisphere.     

Organization of Regional Interactions of the Brain Cortical Activity during the Common-root Word Derivation Task

This study analyzed specificities in the activity of the neurophysiological mechanisms underlying the organization of active word-derivation processes. The regularities in the reorganization of the spatial structure for the systemic interaction of bioelectrical activity between different cortical areas of the cerebral hemispheres were studied in adult subjects during the test for mental derivation of common root words (i.е., using the modern methods of the so-called “functional connectome” investigations). Сross-correlation and coherent analysis of EEG has shown that the ipsilateral statistical EEG interactions in the left hemisphere, including Broca’s and Wernicke’s areas, were significantly increased in adults during mental derivation of common root words and, simultaneously, the interhemispheric connectivity and the EEG interactions in the right hemisphere were reduced. Comparison of our results with the previous data of verbal activity associated with speech perception and production has revealed significant differences in the degree of involvement of the left and right hemisphere cortical activity in verbal processing. For example, unlike the data of current study, an equal involvement of both hemispheres cortical activity was recorded during the phoneme recognition in auditory perceived words, grammatical and semantic errors in sentences, as well as during mental formation of words from a set of phonemes and sentences from a set of words, which was particularly manifested in the increased of hemispheric interactions, predominantly, in the inferior frontal and temporal areas and the overlapped areas of the temporal, parietal, and occipital cortical zones (TPO) of both hemispheres. Thus, the data obtained in this study indicate the presence of expressed specificities in the lateralization of activity in the neurophysiological mechanisms underlying the processes of active word derivation and inflexion.     

Neural Substrates for Semantic Memory of Familiar Songs: Is There an Interface between Lyrics and Melodies?

Findings on song perception and song production have increasingly suggested that common but partially distinct neural networks exist for processing lyrics and melody. However, the neural substrates of song recognition remain to be investigated. The purpose of this study was to examine the neural substrates involved in the accessing “song lexicon” as corresponding to a representational system that might provide links between the musical and phonological lexicons using positron emission tomography (PET). We exposed participants to auditory stimuli consisting of familiar and unfamiliar songs presented in three ways: sung lyrics (song), sung lyrics on a single pitch (lyrics), and the sung syllable ‘la’ on original pitches (melody). The auditory stimuli were designed to have equivalent familiarity to participants, and they were recorded at exactly the same tempo. Eleven right-handed nonmusicians participated in four conditions: three familiarity decision tasks using song, lyrics, and melody and a sound type decision task (control) that was designed to engage perceptual and prelexical processing but not lexical processing. The contrasts (familiarity decision tasks versus control) showed no common areas of activation between lyrics and melody. This result indicates that essentially separate neural networks exist in semantic memory for the verbal and melodic processing of familiar songs. Verbal lexical processing recruited the left fusiform gyrus and the left inferior occipital gyrus, whereas melodic lexical processing engaged the right middle temporal sulcus and the bilateral temporo-occipital cortices. Moreover, we found that song specifically activated the left posterior inferior temporal cortex, which may serve as an interface between verbal and musical representations in order to facilitate song recognition.