Right Inferior Frontal Gyrus Activation Is Associated with Memory Improvement in Patients with Left Frontal Low-Grade Glioma Resection

Patients with low-grade glioma (LGG) have been studied as a model of functional brain reorganization due to their slow-growing nature. However, there is no information regarding which brain areas are involved during verbal memory encoding after extensive left frontal LGG resection. In addition, it remains unknown whether these patients can improve their memory performance after instructions to apply efficient strategies. The neural correlates of verbal memory encoding were investigated in patients who had undergone extensive left frontal lobe (LFL) LGG resections and healthy controls using fMRI both before and after directed instructions were given for semantic organizational strategies. Participants were scanned during the encoding of word lists under three different conditions before and after a brief period of practice. The conditions included semantically unrelated (UR), related-non-structured (RNS), and related-structured words (RS), allowing for different levels of semantic organization. All participants improved on memory recall and semantic strategy application after the instructions for the RNS condition. Healthy subjects showed increased activation in the left inferior frontal gyrus (IFG) and middle frontal gyrus (MFG) during encoding for the RNS condition after the instructions. Patients with LFL excisions demonstrated increased activation in the right IFG for the RNS condition after instructions were given for the semantic strategies. Despite extensive damage in relevant areas that support verbal memory encoding and semantic strategy applications, patients that had undergone resections for LFL tumor could recruit the right-sided contralateral homologous areas after instructions were given and semantic strategies were practiced. These results provide insights into changes in brain activation areas typically implicated in verbal memory encoding and semantic processing.     

Left inferior frontal activations depending on the canonicity determined by the argument structures of ditransitive sentences: an MEG study

To elucidate the relationships between syntactic and semantic processes, one interesting question is how syntactic structures are constructed by the argument structure of a verb, where each argument corresponds to a semantic role of each noun phrase (NP). Here we examined the effects of possessivity [sentences with or without a possessor] and canonicity [canonical or noncanonical word orders] using Japanese ditransitive sentences. During a syntactic decision task, the syntactic structure of each sentence would be constructed in an incremental manner based on the predicted argument structure of the ditransitive verb in a verb-final construction. Using magnetoencephalography, we found a significant canonicity effect on the current density in the left inferior frontal gyrus (IFG) at 530-550 ms after the verb onset. This effect was selective to canonical sentences, and significant even when the precedent NP was physically identical. We suggest that the predictive effects associated with syntactic processing became larger for canonical sentences, where the NPs and verb were merged with a minimum structural distance, leading to the left IFG activations. For monotransitive and intransitive verbs, in which structural computation of the sentences was simpler than that of ditransitive sentences, we observed a significant effect selective to noncanonical sentences in the temporoparietal regions during 480-670 ms. This effect probably reflects difficulty in semantic processing of noncanonical sentences. These results demonstrate that the left IFG plays a predictive role in syntactic processing, which depends on the canonicity determined by argument structures, whereas other temporoparietal regions would subserve more semantic aspects of sentence processing.     

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.     

Altered Regional Homogeneity with Short-term Simulated Microgravity and Its Relationship with Changed Performance in Mental Transformation

In order to further the insight into the explanation of changed performance in mental transformation under microgravity, we discuss the change of performance in mental transformation and its relationship with altered regional homogeneity (ReHo) in resting-state brain by using simulated weightlessness model. Twelve male subjects with age between 24 and 31 received resting-state fMRI scan and mental transformation test both in normal condition and immediately after 72 hours −6° head down tilt (HDT). A paired sample t-test was used to test the difference of behavior performance and brain activity between these two conditions. Compare with normal condition, subjects showed a changed performance in mental transformation with short term simulated microgravity and appeared to be falling. Meanwhile, decreased ReHo were found in right inferior frontal gyrus (IFG) and left inferior parietal lobule (IPL) after 72 hours −6° HDT, while increased ReHo were found in bilateral medial frontal gyrus (MFG) and left superior frontal gyrus (SFG) (P<0.05, corrected). Particularly, there was a significant correlation between ReHo values in left IPL and velocity index of mental transformation. Our findings indicate that gravity change may disrupt the function of right IFG and left IPL in the resting-state, among of which functional change in left IPL may contribute to changed abilities of mental transformation. In addition, the enhanced activity of the bilateral MFG and decreased activity of right IFG found in the current study maybe reflect a complementation effect on inhibitory control process.     

Hearing and seeing meaning in speech and gesture: insights from brain and behaviour

As we speak, we use not only the arbitrary form–meaning mappings of the speech channel but also motivated form–meaning correspondences, i.e. iconic gestures that accompany speech (e.g. inverted V-shaped hand wiggling across gesture space to demonstrate walking). This article reviews what we know about processing of semantic information from speech and iconic gestures in spoken languages during comprehension of such composite utterances. Several studies have shown that comprehension of iconic gestures involves brain activations known to be involved in semantic processing of speech: i.e. modulation of the electrophysiological recording component N400, which is sensitive to the ease of semantic integration of a word to previous context, and recruitment of the left-lateralized frontal–posterior temporal network (left inferior frontal gyrus (IFG), medial temporal gyrus (MTG) and superior temporal gyrus/sulcus (STG/S)). Furthermore, we integrate the information coming from both channels recruiting brain areas such as left IFG, posterior superior temporal sulcus (STS)/MTG and even motor cortex. Finally, this integration is flexible: the temporal synchrony between the iconic gesture and the speech segment, as well as the perceived communicative intent of the speaker, modulate the integration process. Whether these findings are special to gestures or are shared with actions or other visual accompaniments to speech (e.g. lips) or other visual symbols such as pictures are discussed, as well as the implications for a multimodal view of language.     

Investigating a network model of word generation with positron emission tomography.

By using positron emission tomography (PET) we examined the biological validity of a network model describing changes in cerebral activity associated with intrinsic and extrinsic word generation. The production of words not specified by an extrinsic stimulus constitutes willed or intrinsic generation. Perceiving a heard word is an example of extrinsic generation. The model incorporates three neuronal systems: a pool that stores word representations in a distributed fashion, an afferent system conveying sensory input to the pool and a modulating system that alters the responsivity of neurons in the pool. Simulations based on the model suggested that intrinsic generation would be associated with low activity in the pool, consequent on reduced modulation, and extrinsic generation with high activity. We measured cerebral activity with PET during intrinsic (verbal fluency) and extrinsic (responding to heard words) word generation and found this pattern of changes in the left superior temporal region. We were able to designate this region the site of the distributed word store and implicate the left dorsolateral prefrontal cortex (DLPFC) as the source of modulation. The relation between the superior temporal gyrus and DLPFC was shown by examining the correlation between the two regions in terms of cerebral activity. We conclude that the left DLPFC is responsible for modulating the responsivity of a neural system in the superior temporal gyrus and is the probable mediator of changes in attentional and intentional states that underlies the intrinsic generation of words.     

The temporal dynamics of reading: a PET study.

The temporal dynamics of evoked brain responses are normally characterized using electrophysiological techniques but the positron emission tomography study presented here revealed a temporal aspect of reading by correlating the duration a word remained in the visual field with evoked haemodynamic response. Three distinct types of effects were observed: in visual processing areas, there were linear increases in activity with duration suggesting that visual processing endures throughout the time the stimulus remains in the visual field. In right hemisphere areas, there were monotonic decreases in activity with increased duration which we relate to decreased attention for longer stimulus durations. In left hemisphere word processing areas there were inverted U-shaped dependencies between activity and word duration indicating that, after 400-600 ms, activity in word processing areas is progressively reduced if the word remains in the visual field. We conclude that these inverted U effects in left hemisphere language areas reflect the temporal dynamics of visual word processing and we highlight the implication of these effects for the design of activation studies involving reading.     

The Brain’s Dorsal Route for Speech Represents Word Meaning: Evidence from Gesture

The dual-route model of speech processing includes a dorsal stream that maps auditory to motor features at the sublexical level rather than at the lexico-semantic level. However, the literature on gesture is an invitation to revise this model because it suggests that the premotor cortex of the dorsal route is a major site of lexico-semantic interaction. Here we investigated lexico-semantic mapping using word-gesture pairs that were either congruent or incongruent. Using fMRI-adaptation in 28 subjects, we found that temporo-parietal and premotor activity during auditory processing of single action words was modulated by the prior audiovisual context in which the words had been repeated. The BOLD signal was suppressed following repetition of the auditory word alone, and further suppressed following repetition of the word accompanied by a congruent gesture (e.g. [“grasp” + grasping gesture]). Conversely, repetition suppression was not observed when the same action word was accompanied by an incongruent gesture (e.g. [“grasp” + sprinkle]). We propose a simple model to explain these results: auditory and visual information converge onto premotor cortex where it is represented in a comparable format to determine (in)congruence between speech and gesture. This ability of the dorsal route to detect audiovisual semantic (in)congruence suggests that its function is not restricted to the sublexical level.     

Effects of TMS on Different Stages of Motor and Non-Motor Verb Processing in the Primary Motor Cortex

The embodied cognition hypothesis suggests that motor and premotor areas are automatically and necessarily involved in understanding action language, as word conceptual representations are embodied. This transcranial magnetic stimulation (TMS) study explores the role of the left primary motor cortex in action-verb processing. TMS-induced motor-evoked potentials from right-hand muscles were recorded as a measure of M1 activity, while participants were asked either to judge explicitly whether a verb was action-related (semantic task) or to decide on the number of syllables in a verb (syllabic task). TMS was applied in three different experiments at 170, 350 and 500 ms post-stimulus during both tasks to identify when the enhancement of M1 activity occurred during word processing. The delays between stimulus onset and magnetic stimulation were consistent with electrophysiological studies, suggesting that word recognition can be differentiated into early (within 200 ms) and late (within 400 ms) lexical-semantic stages, and post-conceptual stages. Reaction times and accuracy were recorded to measure the extent to which the participants'' linguistic performance was affected by the interference of TMS with M1 activity. No enhancement of M1 activity specific for action verbs was found at 170 and 350 ms post-stimulus, when lexical-semantic processes are presumed to occur (Experiments 1–2). When TMS was applied at 500 ms post-stimulus (Experiment 3), processing action verbs, compared with non-action verbs, increased the M1-activity in the semantic task and decreased it in the syllabic task. This effect was specific for hand-action verbs and was not observed for action-verbs related to other body parts. Neither accuracy nor RTs were affected by TMS. These findings suggest that the lexical-semantic processing of action verbs does not automatically activate the M1. This area seems to be rather involved in post-conceptual processing that follows the retrieval of motor representations, its activity being modulated (facilitated or inhibited), in a top-down manner, by the specific demand of the task.     

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.     

Dissociation between the activity of the right middle frontal gyrus and the middle temporal gyrus in processing semantic priming

The aim of this event-related functional magnetic resonance imaging (fMRI) study was to test whether the right middle frontal gyrus (MFG) and middle temporal gyrus (MTG) would show differential sensitivity to the effect of prime-target association strength on repetition priming. In the experimental condition (RP), the target occurred after repetitive presentation of the prime within an oddball design. In the control condition (CTR), the target followed a single presentation of the prime with equal probability of the target as in RP. To manipulate semantic overlap between the prime and the target both conditions (RP and CTR) employed either the onomatopoeia "oink" as the prime and the referent "pig" as the target (OP) or vice-versa (PO) since semantic overlap was previously shown to be greater in OP. The results showed that the left MTG was sensitive to release of adaptation while both the right MTG and MFG were sensitive to sequence regularity extraction and its verification. However, dissociated activity between OP and PO was revealed in RP only in the right MFG. Specifically, target "pig" (OP) and the physically equivalent target in CTR elicited comparable deactivations whereas target "oink" (PO) elicited less inhibited response in RP than in CTR. This interaction in the right MFG was explained by integrating these effects into a competition model between perceptual and conceptual effects in priming processing.     

Children with reading disability show brain differences in effective connectivity for visual, but not auditory word comprehension

Background

Previous literature suggests that those with reading disability (RD) have more pronounced deficits during semantic processing in reading as compared to listening comprehension. This discrepancy has been supported by recent neuroimaging studies showing abnormal activity in RD during semantic processing in the visual but not in the auditory modality. Whether effective connectivity between brain regions in RD could also show this pattern of discrepancy has not been investigated.

Methodology/Principal Findings

Children (8- to 14-year-olds) were given a semantic task in the visual and auditory modality that required an association judgment as to whether two sequentially presented words were associated. Effective connectivity was investigated using Dynamic Causal Modeling (DCM) on functional magnetic resonance imaging (fMRI) data. Bayesian Model Selection (BMS) was used separately for each modality to find a winning family of DCM models separately for typically developing (TD) and RD children. BMS yielded the same winning family with modulatory effects on bottom-up connections from the input regions to middle temporal gyrus (MTG) and inferior frontal gyrus(IFG) with inconclusive evidence regarding top-down modulations. Bayesian Model Averaging (BMA) was thus conducted across models in this winning family and compared across groups. The bottom-up effect from the fusiform gyrus (FG) to MTG rather than the top-down effect from IFG to MTG was stronger in TD compared to RD for the visual modality. The stronger bottom-up influence in TD was only evident for related word pairs but not for unrelated pairs. No group differences were noted in the auditory modality.

Conclusions/Significance

This study revealed a modality-specific deficit for children with RD in bottom-up effective connectivity from orthographic to semantic processing regions. There were no group differences in connectivity from frontal regions, suggesting that the core deficit in RD is not in top-down modulation.     

Relating Inter-Individual Differences in Verbal Creative Thinking to Cerebral Structures: An Optimal Voxel-Based Morphometry Study

Creativity can be defined the capacity of an individual to produce something original and useful. An important measurable component of creativity is divergent thinking. Despite existing studies on creativity-related cerebral structural basis, no study has used a large sample to investigate the relationship between individual verbal creativity and regional gray matter volumes (GMVs) and white matter volumes (WMVs). In the present work, optimal voxel-based morphometry (VBM) was employed to identify the structure that correlates verbal creativity (measured by the verbal form of Torrance Tests of Creative Thinking) across the brain in young healthy subjects. Verbal creativity was found to be significantly positively correlated with regional GMV in the left inferior frontal gyrus (IFG), which is believed to be responsible for language production and comprehension, new semantic representation, and memory retrieval, and in the right IFG, which may involve inhibitory control and attention switching. A relationship between verbal creativity and regional WMV in the left and right IFG was also observed. Overall, a highly verbal creative individual with superior verbal skills may demonstrate a greater computational efficiency in the brain areas involved in high-level cognitive processes including language production, semantic representation and cognitive control.     

De fonologische variant van primaire progressieve afasie, een gevalsstudie

Primary progressive aphasia (PPA) is a neurodegenerative syndrome characterized by an insidious onset and gradual progression of deficits that can involve any aspect of language, including word finding, object naming, fluency, syntax, phonology and word comprehension. The initial symptoms occur in the absence of major deficits in other cognitive domains, including episodic memory, visuospatial abilities and visuoconstruction. According to recent diagnostic guidelines, PPA is typically divided into three variants: nonfluent variant PPA (also termed progressive nonfluent aphasia), semantic variant PPA (also termed semantic dementia) and logopenic/phonological variant PPA (also termed logopenic progressive aphasia). The paper describes a 79-yr old man, who presented with normal motor speech and production rate, impaired single word retrieval and phonemic errors in spontaneous speech and confrontational naming. Confrontation naming was strongly affected by lexical frequency. He was impaired on repetition of sentences and phrases. Reading was intact for regularly spelled words but not for irregular words (surface dyslexia). Comprehension was spared at the single word level, but impaired for complex sentences. He performed within the normal range on the Dutch equivalent of the Pyramids and Palm Trees (PPT) Pictures Test, indicating that semantic processing was preserved. There was, however, a slight deficiency on the PPT Words Test, which appeals to semantic knowledge of verbal associations. His core deficit was interpreted as an inability to retrieve stored lexical-phonological information for spoken word production in spontaneous speech, confrontation naming, repetition and reading aloud.     

Brain mechanisms of the semantic analysis of words-homonyms

The brain mechanisms of semantic comprehension of a word were comparatively studied in three experimental conditions: simple perception of the nuclear value of a word-homonym determined by the preceding context, perception of its circumferential value, and during active semantic analysis of these values. It was shown that the amplitude of the evoked potential component P200 was correlated with complexity of the semantic analysis. A decrease in this amplitude under conditions of complication of semantic problem was associated with an increase in the activities of the caudate nucleus and hippocampus and a parallel slight decrease in the activity of cortical areas.     

Word learning and the cerebral hemispheres: from serial to parallel processing of written words

Reading familiar words differs from reading unfamiliar non-words in two ways. First, word reading is faster and more accurate than reading of unfamiliar non-words. Second, effects of letter length are reduced for words, particularly when they are presented in the right visual field in familiar formats. Two experiments are reported in which right-handed participants read aloud non-words presented briefly in their left and right visual fields before and after training on those items. The non-words were interleaved with familiar words in the naming tests. Before training, naming was slow and error prone, with marked effects of length in both visual fields. After training, fewer errors were made, naming was faster, and the effect of length was much reduced in the right visual field compared with the left. We propose that word learning creates orthographic word forms in the mid-fusiform gyrus of the left cerebral hemisphere. Those word forms allow words to access their phonological and semantic representations on a lexical basis. But orthographic word forms also interact with more posterior letter recognition systems in the middle/inferior occipital gyri, inducing more parallel processing of right visual field words than is possible for any left visual field stimulus, or for unfamiliar non-words presented in the right visual field.     

Facilitating memory for novel characters by reducing neural repetition suppression in the left fusiform cortex

Background

The left midfusiform and adjacent regions have been implicated in processing and memorizing familiar words, yet its role in memorizing novel characters has not been well understood.

Methodology/Principal Findings

Using functional MRI, the present study examined the hypothesis that the left midfusiform is also involved in memorizing novel characters and spaced learning could enhance the memory by enhancing the left midfusiform activity during learning. Nineteen native Chinese readers were scanned while memorizing the visual form of 120 Korean characters that were novel to the subjects. Each character was repeated four times during learning. Repetition suppression was manipulated by using two different repetition schedules: massed learning and spaced learning, pseudo-randomly mixed within the same scanning session. Under the massed learning condition, the four repetitions were consecutive (with a jittered inter-repetition interval to improve the design efficiency). Under the spaced learning condition, the four repetitions were interleaved with a minimal inter-repetition lag of 6 stimuli. Spaced learning significantly improved participants'' performance during the recognition memory test administered one hour after the scan. Stronger left midfusiform and inferior temporal gyrus activities during learning (summed across four repetitions) were associated with better memory of the characters, based on both within- and cross-subjects analyses. Compared to massed learning, spaced learning significantly reduced neural repetition suppression and increased the overall activities in these regions, which were associated with better memory for novel characters.

Conclusions/Significance

These results demonstrated a strong link between cortical activity in the left midfusiform and memory for novel characters, and thus challenge the visual word form area (VWFA) hypothesis. Our results also shed light on the neural mechanisms of the spacing effect in memorizing novel characters.     

Concurrent word generation and motor performance: further evidence for language-motor interaction

Embodied/modality-specific theories of semantic memory propose that sensorimotor representations play an important role in perception and action. A large body of evidence supports the notion that concepts involving human motor action (i.e., semantic-motor representations) are processed in both language and motor regions of the brain. However, most studies have focused on perceptual tasks, leaving unanswered questions about language-motor interaction during production tasks. Thus, we investigated the effects of shared semantic-motor representations on concurrent language and motor production tasks in healthy young adults, manipulating the semantic task (motor-related vs. nonmotor-related words) and the motor task (i.e., standing still and finger-tapping). In Experiment 1 (n = 20), we demonstrated that motor-related word generation was sufficient to affect postural control. In Experiment 2 (n = 40), we demonstrated that motor-related word generation was sufficient to facilitate word generation and finger tapping. We conclude that engaging semantic-motor representations can have a reciprocal influence on motor and language production. Our study provides additional support for functional language-motor interaction, as well as embodied/modality-specific theories.     

Anorexia Nervosa during Adolescence Is Associated with Decreased Gray Matter Volume in the Inferior Frontal Gyrus

Anorexia nervosa (AN) is an eating disorder characterized by the relentless pursuit to lose weight, mostly through self-starvation, and a distorted body image. AN tends to begin during adolescence among women. However, the underlying neural mechanisms related to AN remain unclear. Using voxel-based morphometry based on magnetic resonance imaging scans, we investigated whether the presence of AN was associated with discernible changes in brain morphology. Participants were 20 un-medicated, right-handed patients with early-onset AN and 14 healthy control subjects. Group differences in gray matter volume (GMV) were assessed using high-resolution, T1-weighted, volumetric magnetic resonance imaging datasets (3T Trio scanner; Siemens AG) and analyzed after controlling for age and total GMV, which was decreased in the bilateral inferior frontal gyrus (IFG) (left IFG: FWE corrected, p < 0.05; right IFG: uncorrected, p < 0.05) of patients with AN. The GMV in the bilateral IFG correlated significantly with current age (left IFG: r = -.481, p < .05; right IFG: r = -.601, p < .01) and was limited to the AN group. We speculate that decreased IFG volume might lead to deficits in executive functioning or inhibitory control within neural reward systems. Precocious or unbalanced neurological trimming within this particular region might be an important factor for the pathogenesis of AN onset.     

Written production of English compounds: effects of morphology and semantic transparency

The results of research on the processing of morphologically complex words are consistent with a lexical system that activates both whole-word and constituent representations during word recognition. In this study, we focus on written production and examine whether semantically priming the first constituent of a compound influences the ease of producing a compound (as measured by typing latencies), and whether any such priming effect depends on the semantic transparency of the compound’s constituents. We found that semantic transparency of the constituents affects whether semantic priming results in changes to processing. However, it is not only the semantic transparency of the primed constituent that exerts an influence—for example, the semantic transparency of the head affects whether semantically priming the modifier results in a change in typing times. We discuss these effects in terms of competition among the various representations as the compound is output, such that overall performance is a combination of facilitation and inhibition that changes over the course of the output.