Neural correlates of semantic associations in patients with schizophrenia.

Patients with schizophrenia have semantic processing disturbances leading to expressive language deficits (formal thought disorder). The underlying pathology has been related to alterations in the semantic network and its neural correlates. Moreover, crossmodal processing, an important aspect of communication, is impaired in schizophrenia. Here we investigated specific processing abnormalities in patients with schizophrenia with regard to modality and semantic distance in a semantic priming paradigm. Fourteen patients with schizophrenia and fourteen demographically matched controls made visual lexical decisions on successively presented word-pairs (SOA = 350 ms) with direct or indirect relations, unrelated word-pairs, and pseudoword-target stimuli during fMRI measurement. Stimuli were presented in a unimodal (visual) or crossmodal (auditory-visual) fashion. On the neural level, the effect of semantic relation indicated differences (patients > controls) within the right angular gyrus and precuneus. The effect of modality revealed differences (controls > patients) within the left superior frontal, middle temporal, inferior occipital, right angular gyri, and anterior cingulate cortex. Semantic distance (direct vs. indirect) induced distinct activations within the left middle temporal, fusiform gyrus, right precuneus, and thalamus with patients showing fewer differences between direct and indirect word-pairs. The results highlight aberrant priming-related brain responses in patients with schizophrenia. Enhanced activation for patients possibly reflects deficits in semantic processes that might be caused by a delayed and enhanced spread of activation within the semantic network. Modality-specific decreases of activation in patients might be related to impaired perceptual integration. Those deficits could induce and increase the prominent symptoms of schizophrenia like impaired speech processing.

Is the motor or the garage more important to the car? The difference between semantic associations in single word and sentence production.

The aim of the present study was to investigate the impact of part-whole (e.g., car-motor) and functional associations (e.g., car-garage) on single word (Experiment 1) and sentence production (Experiment 2). In Experiment 1, a classical picture-word task was used. In Experiment 2, the same stimuli and distractors were embedded into a sentence. The relation between target and distractor was either part-whole, functional or unrelated. At single word level, part-whole and functional relations facilitate naming. Additionally, the facilitation effect was stronger for part-whole in comparison to functional associations. During sentence production, facilitation shifted to interference. The difference between both relations disappeared. The findings of the different effects between functional and part-whole associations depend on the length of utterances and highlight the divergent impact of associations. The differences between part-whole and functional associations in single word production might reflect a differential organization of associative links at the conceptual level. In contrast, during sentence production the syntactic processing at the lexical level seem to be more important than types of semantic associations at the conceptual level.

The influence of emotional associations on the neural correlates of semantic priming.

Emotions influence our everyday life in several ways. With the present study, we wanted to examine the impact of emotional information on neural correlates of semantic priming, a well-established technique to investigate semantic processing. Stimuli were presented with a short SOA of 200 ms as subjects performed a lexical decision task during fMRI measurement. Seven experimental conditions were compared: positive/negative/neutral related, positive/negative/neutral unrelated, nonwords (all words were nouns). Behavioral data revealed a valence specific semantic priming effect (i.e., unrelated > related) only for neutral and positive related word pairs. On a neural level, the comparison of emotional over neutral relations showed activation in left anterior medial frontal cortex, superior frontal gyrus, and posterior cingulate. Interactions for the different relations were located in left anterior part of the medial frontal cortex, cingulate regions, and right hippocampus (positive > neutral + negative) and left posterior part of medial frontal cortex (negative > neutral + positive). The results showed that emotional information have an influence on semantic association processes. While positive and neutral information seem to share a semantic network, negative relations might induce compensatory mechanisms that inhibit the spread of activation between related concepts. The neural correlates highlighted a distributed neural network, primarily involving attention, memory and emotion related processing areas in medial fronto-parietal cortices. The differentiation between anterior (positive) and posterior part (negative) of the medial frontal cortex was linked to the type of affective manipulation with more cognitive demands being involved in the automatic processing of negative information.

Early attention modulates perceptual interpretation of multisensory stimuli.

Perceptual interpretation of the same multisensory stimuli may rely upon influence of attention on multisensory integration. We tested this hypothesis by recording the brain activity using magnetoencephalography in a passing-bouncing illusion with sound. Early activation of the attention-related brain areas and subsequent involvement of the multisensory areas were associated with the bouncing percept. Early activation of the unimodal sensory areas and later involvement of the attention-related areas were associated with the passing percept. Thus, the bouncing percept occurs when early attentional deployment facilitates multisensory integration modulating visual perception. The passing percept results from hierarchical sequence of the perceptual processes: early activation of the unimodal areas and late attentional deployment. Alternation of the perceptual interpretations may depend on spontaneous fluctuation of attention.

How different types of conceptual relations modulate brain activation during semantic priming.

Semantic priming, a well-established technique to study conceptual representation, has thus far produced variable fMRI results, both regarding the type of priming effects and their correlation with brain activation. The aims of the current study were (a) to investigate two types of semantic relations--categorical versus associative--under controlled processing conditions and (b) to investigate whether categorical and associative relations between words are correlated with response enhancement or response suppression. We used fMRI to examine neural correlates of semantic priming as subjects performed a lexical decision task with a long SOA (800 msec). Four experimental conditions were compared: categorically related trials (couch-bed), associatively related trials (couch-pillow), unrelated trials (couch-bridge), and nonword trials (couch-sibor). We found similar behavioral priming effects for both categorically and associatively related pairs. However, the neural priming effects differed: Categorically related pairs resulted in a neural suppression effect in the right MFG, whereas associatively related pairs resulted in response enhancement in the left IFG. A direct contrast between them revealed activation for categorically related trials in the right insular lobe. We conclude that perceptual and functional similarity of categorically related words may lead to response suppression within right-lateralized frontal regions that represent more retrieval effort and the recruitment of a broader semantic field. Associatively related pairs that require a different processing of the related target compared to the prime may lead to the response enhancement within left inferior frontal regions. Nevertheless, the differences between associative and categorical relations might be parametrical rather than absolutely distinct as both relationships recruit similar regions to a different degree.

Why the leash constrains the dog: the impact of semantic associations on sentence production.

The production of language is one of the most complex and amazing skills in humans. Increasing evidence demonstrated that associative relations (e.g., car - garage) play an important role during concept formation but during speech production the effects and processing of associations are highly debated. Hence, the present study investigated the impact of associations and different SOAs on the production of sentences (Experiment 1) and on naming objects (Experiment 2). In an adapted version of the picture-word interference task, participants were asked to name two pictures using a standardized sentence (e.g., "The car is to the left of the trousers"). Thereby, a simultaneous (SOA is 0 ms) or slightly preceding (SOA is -150 ms) auditory or visual distractor had to be ignored. Distractors were related to the first noun (for example: "The car is to the left to the trousers", distractor: "garage") or to the second noun (distractor: "belt") or unrelated to both nouns (distractor: "bottle") of the sentence. At simultaneous presentation, visual and auditory distractors related to the first noun of the sentence prolonged naming responses (i.e., interference). For slightly preceding distractors, only visual presentation induced interference for the first noun of the sentence. During no condition, longer naming responses were found for the second noun of the sentence. These effects suggest that associatively related concepts are active during speech production and can be competitors, i.e., they lead to semantic interference. In Experiment 2, subjects had to name an object (e.g., car) while ignoring a visually presented distractor (e.g., motor). The stimulus set was the same as in Experiment 1. The results showed a facilitation effect if the distractor and the target were associatively related. Overall, the current results provide new insight in the models of speech production: while during single word production, associations facilitate naming, they interfere during sentence production. Hence, associations have an important influence on producing speech but the impact is varied by the context, i.e., single word or sentential.

Priming words with pictures: neural correlates of semantic associations in a cross-modal priming task using fMRI.

In our everyday life we process information from different modalities simultaneously with great ease. With the current study we had the following goals: to detect the neural correlates of (1) automatic semantic processing of associates and (2) to investigate the influence of different visual modalities on semantic processing. Stimuli were presented with a short SOA (350 ms) as subjects performed a lexical decision task. To minimize the variance and increase homogeneity within our sample, only male subjects were measured. Three experimental conditions were compared while brain activation was measured with a 3 T fMRI scanner: related word-pairs (e.g., frame-picture), unrelated word-pairs (e.g., frame-car) as well as word-nonword pairs (e.g., frame-fubber). They were presented uni- (word-word) and cross-modally (picture-word). Behavioral data revealed a priming effect for cross-modal and unimodal word-pairs. On a neural level, the unimodal condition revealed response suppression in bilateral fronto-parietal regions. Cross-modal priming led to response suppression within the right inferior frontal gyrus. Common areas of deactivation for both modalities were found in bilateral fronto-tempo-parietal regions. These results suggest that the processing of semantic associations presented in different modalities lead to modality-specific activation caused by early access routes. However, common activation for both modalities refers to a common neural network for semantic processing suggesting amodal processing.

Spatially congruent visual motion modulates activity of the primary auditory cortex.

We investigated the brain responses to the transitions from the static to moving audiovisual stimuli using magnetoencephalography. The spatially congruent auditory and visual stimuli moved in the same direction whereas the incongruent stimuli moved in the opposite directions. Using dipole modeling we found that the static-to-moving transitions evoked a neural response in the primary auditory cortex bilaterally. The response started about 100 ms after the motion onset from a negative component (mvN1) and lasted during the entire interval of the stimulus motion. The mvN1 component was similar to the classical auditory N1 response to the static sound, but had smaller amplitude and later latency. The coordinates of the mvN1 and N1 dipoles in the primary auditory cortex were also similar. The amplitude of the auditory response to the moving stimuli appears to be sensitive to spatial congruency of the audiovisual motion; it was larger in the incongruent than congruent condition. This is evidence that the moving visual stimuli modulate the early sensory activity in the primary auditory cortex. Such early audiovisual integration may be specific for motion processing.

Taxonomic and thematic categories: Neural correlates of categorization in an auditory-to-visual priming task using fMRI.

Categorization is a basic principle of knowledge organization in the brain. The goal of the current fMRI study was to compare the neural correlates of thematic (e.g., car - garage) and taxonomic (e.g., couch - bed) categories under automatic processing conditions using auditory-to-visual semantic priming. Behavioral data revealed a priming effect for thematically but not for taxonomically related word pairs. On a neural level, thematically related words led to a left-lateralized temporal activation (superior temporal sulcus), whereas taxonomically related word pairs evoked a right-lateralized frontal activation and within the hippocampus. A direct comparison between both categories revealed enhanced activation for thematically related and response suppression for taxonomically related trials in the left superior temporal sulcus. These results suggest that processing of thematic and taxonomic categories leads to activation of distinct brain areas. The mainly right-lateralized fronto-temporal activation for taxonomic relations suggests increased attention and effort for processing this category. The interaction within the left superior temporal sulcus reflects the processing and retrieval of semantic relations whereby specific memory contents seem to influence the direction of activation.

Lion - tiger - stripes: Neural correlates of indirect semantic priming across processing modalities.

"Lions" do not have "stripes". However, via the word "tiger" both words are closely connected within the semantic network. With the present study we pursued two goals: to detect neural correlates of (1) directly and indirectly related word pairs by means of priming, and (2) to assess the effect of presentation modality. Stimuli were presented with a short SOA of 350 ms as subjects performed a lexical decision task during fMRI measurement. Four experimental conditions were compared: directly related (picture-frame), indirectly related (anvil-nail), unrelated (steamboat-needle) and nonword trials (chalk-edan), presented in a uni- (word-word) and cross-modal (auditory-word) version. Behavioral data revealed a modality-independent priming effect only for direct semantic priming. On a neural level, directly linked words led to left-lateralized activations in fronto-temporo-parietal areas. Indirect priming led to right-hemispheric fronto-parietal signal changes. Common areas of activation for uni- and cross-modal priming were found within the left middle temporal gyrus and right precuneus for direct priming and within the right insula for indirect priming. The comparison of the semantic distances (direct>indirect) showed one region activated modality-independent: the precuneus. Direct priming is associated with activation clusters corresponding to a large left-lateralized network. Indirect priming recruits right-hemispheric regions, reflecting widespread semantic fields and attentional components. The modality-independent comparison of direct and indirect priming revealed common areas of activation supporting an amodal rather than multiple semantic systems. The activation related to semantic distances underpins the special role of the precuneus. This region is involved in semantic priming and association processing whereas episodic memory contents might be addressed.

Neural interaction of speech and gesture: differential activations of metaphoric co-verbal gestures.

Gestures are an important part of human communication. However, little is known about the neural correlates of gestures accompanying speech comprehension. The goal of this study is to investigate the neural basis of speech-gesture interaction as reflected in activation increase and decrease during observation of natural communication. Fourteen German participants watched video clips of 5 s duration depicting an actor who performed metaphoric gestures to illustrate the abstract content of spoken sentences. Furthermore, video clips of isolated gestures (without speech), isolated spoken sentences (without gestures) and gestures in the context of an unknown language (Russian) were additionally presented while functional magnetic resonance imaging (fMRI) data were acquired. Bimodal speech and gesture processing led to left hemispheric activation increases of the posterior middle temporal gyrus, the premotor cortex, the inferior frontal gyrus, and the right superior temporal sulcus. Activation reductions during the bimodal condition were located in the left superior temporal gyrus and the left posterior insula. Gesture related activation increases and decreases were dependent on language semantics and were not found in the unknown-language condition. Our results suggest that semantic integration processes for bimodal speech plus gesture comprehension are reflected in activation increases in the classical left hemispheric language areas. Speech related gestures seem to enhance language comprehension during the face-to-face communication.

Automatic processing of semantic relations in fMRI: neural activation during semantic priming of taxonomic and thematic categories.

Most current models of knowledge organization are based on hierarchical or taxonomic categories (animals, tools). Another important organizational pattern is thematic categorization, i.e. categories held together by external relations, a unifying scene or event (car and garage). The goal of this study was to compare the neural correlates of these categories under automatic processing conditions that minimize strategic influences. We used fMRI to examine neural correlates of semantic priming for category members with a short stimulus onset asynchrony (SOA) of 200 ms as subjects performed a lexical decision task. Four experimental conditions were compared: thematically related words (car-garage); taxonomically related (car-bus); unrelated (car-spoon); non-word trials (car-derf). We found faster reaction times for related than for unrelated prime-target pairs for both thematic and taxonomic categories. However, the size of the thematic priming effect was greater than that of the taxonomic. The imaging data showed signal changes for the taxonomic priming effects in the right precuneus, postcentral gyrus, middle frontal and superior frontal gyri and thematic priming effects in the right middle frontal gyrus and anterior cingulate. The contrast of neural priming effects showed larger signal changes in the right precuneus associated with the taxonomic but not with thematic priming response. We suggest that the greater involvement of precuneus in the processing of taxonomic relations indicates their reduced salience in the knowledge structure compared to more prominent thematic relations.

Categorical and thematic knowledge representation in the brain: neural correlates of taxonomic and thematic conceptual relations.

Most current models of knowledge organization in the brain are based on hierarchical or taxonomic categories (animals, tools). Another important organizational pattern is thematic categorization, i.e. categories held together by external relations, a unifying scene or event (car and garage). We used fMRI to examine neural activation patterns as subjects performed a category construction task where these two category types were contrasted. Subjects were visually presented with a target word followed by the presentation of two match words and had to choose by button press one match that goes best with the target word. In the balanced or cross-categorization condition (Car/Garage Bus) both match words fit the target; in the biased conditions only one match word fit the target either thematically (Car/Garage Brush) or taxonomically (Car/Bus Eraser). We found that in the biased conditions, thematic and taxonomic categories recruited very similar cortical regions: left inferior frontal, middle temporal and occipital regions. In the balanced condition subjects showed no behavioral preference for either thematic or taxonomic matches. However, contrasting signal changes during a subjective taxonomic choice in the presence of a thematic alternative vs. a subjective thematic choice in the presence of a taxonomic alternative required the additional recruitment of right middle frontal gyrus, left precuneus and left thalamus. Our results suggest that thematic relations between objects are processed similarly to taxonomic relations, but require less cerebral processing demand, providing validation for thematic categories as an alternative principle of conceptual organization.