Dynamic brain connectivity attuned to the complexity of relative clause sentences revealed by a single-trial analysis.

To explore the issue of how the human brain processes sentences with different levels of complexity, we sought to compare the neural substrates underlying the processing of Chinese subject-extracted relative clause (SRC) and object-extracted relative clause (ORC) sentences in a trial-by-trial fashion. Previous neuroimaging studies have demonstrated that the involvement of the left inferior frontal gyrus (LIFG) and the left superior temporal gyrus (LSTG) is critical for the processing of relative clause (RC) sentences. In this study, we employed independent component analysis (ICA) to decompose brain activity into a set of independent components. Then, the independent component maps were spatially normalized using a surface-based approach in order to further spatially correlate and match the equivalent components from individual participants. The selected equivalent components indicated that the LIFG and the LSTG were consistently engaged in sentence processing among the participants. Subsequently, we observed alterations in the functional coupling between the LIFG and the LSTG in response to SRCs and ORCs using a Granger causality analysis. Specifically, comprehending Chinese ORCs with a canonical word order only involved a unidirectional connection from the LIFG to the LSTG for the integration of lexical-syntactic information. On the other hand, comprehending Chinese SRCs required bi-directional connectivity between the LIFG and the LSTG to fulfill increased integration demands in reconstructing the argument hierarchy due to a non-canonical word order. Furthermore, through a single-trial analysis, the strength of the connectivity from the LIFG to the LSTG was found to be significantly correlated with the complexity of the SRC sentences as quantified by eye-tracking measures. These findings indicated that the effective connectivity from the LIFG to the LSTG played an important role in the comprehension of complex sentences and that enhanced strength of this connectivity might reflect increased integration demands and restructuring attempts during sentence processing. Taken together, the results of the present study reveal that interregional interaction in the brain network for sentence processing can be dynamically engaged in response to different levels of complexity and also shed some light on the interpretation of neuroimaging and behavioral evidence when accounting for the nature of sentence complexity during reading.

Universal brain signature of proficient reading: Evidence from four contrasting languages.

We propose and test a theoretical perspective in which a universal hallmark of successful literacy acquisition is the convergence of the speech and orthographic processing systems onto a common network of neural structures, regardless of how spoken words are represented orthographically in a writing system. During functional MRI, skilled adult readers of four distinct and highly contrasting languages, Spanish, English, Hebrew, and Chinese, performed an identical semantic categorization task to spoken and written words. Results from three complementary analytic approaches demonstrate limited language variation, with speech-print convergence emerging as a common brain signature of reading proficiency across the wide spectrum of selected languages, whether their writing system is alphabetic or logographic, whether it is opaque or transparent, and regardless of the phonological and morphological structure it represents.

Neurolinguistic Studies of Reading in Chinese.

Using the Chinese writing system, which is unique with respect to the composition of each character in terms of its graphic shape, as an example, this chapter addresses the neurobiological underpinnings of reading and writing and how these brain circuits are used in different languages.

Functional and anatomical dissociation between the orthographic lexicon and the orthographic buffer revealed in reading and writing Chinese characters by fMRI.

The contribution of orthographic representations to reading and writing has been intensively investigated in the literature. However, the distinction between neuronal correlates of the orthographic lexicon and the orthographic (graphemic) buffer has rarely been examined in alphabetic languages and never been explored in non-alphabetic languages. To determine whether the neural networks associated with the orthographic lexicon and buffer of logographic materials are comparable to those reported in the literature, the present fMRI experiment manipulated frequency and the stroke number of Chinese characters in the tasks of form judgment and stroke judgment, which emphasized the processing of character recognition and writing, respectively. It was found that the left fusiform gyrus exhibited higher activation when encountering low-frequency than high-frequency characters in both tasks, which suggested this region to be the locus of the orthographic lexicon that represents the knowledge of character forms. On the other hand, the activations in the posterior part of the left middle frontal gyrus and in the left angular gyrus were parametrically modulated by the stroke number of target characters only in the stroke judgment task, which suggested these regions to be the locus of the orthographic buffer that represents the processing of stroke sequence in writing. These results provide the first evidence for the functional and anatomical dissociation between the orthographic lexicon and buffer in reading and writing Chinese characters. They also demonstrate the critical roles of the left fusiform area and the frontoparietal network to the long-term and short-term representations of orthographic knowledge, respectively, across different orthographies.

Adaptation of the human visual system to the statistics of letters and line configurations.

By adulthood, literate humans have been exposed to millions of visual scenes and pages of text. Does the human visual system become attuned to the statistics of its inputs? Using functional magnetic resonance imaging, we examined whether the brain responses to line configurations are proportional to their natural-scene frequency. To further distinguish prior cortical competence from adaptation induced by learning to read, we manipulated whether the selected configurations formed letters and whether they were presented on the horizontal meridian, the familiar location where words usually appear, or on the vertical meridian. While no natural-scene frequency effect was observed, we observed letter-status and letter frequency effects on bilateral occipital activation, mainly for horizontal stimuli. The findings suggest a reorganization of the visual pathway resulting from reading acquisition under genetic and connectional constraints. Even early retinotopic areas showed a stronger response to letters than to rotated versions of the same shapes, suggesting an early visual tuning to large visual features such as letters.

Neural correlates of merging number words.

Complex number words (e.g., "twenty two") are formed by merging together several simple number words (e.g., "twenty" and "two"). In the present study, we explored the neural correlates of this operation and investigated to what extent it engages brain areas involved processing numerical quantity and linguistic syntactic structure. Participants speaking two typologically distinct languages, French and Chinese, were required to read aloud sequences of simple number words while their cerebral activity was recorded by functional magnetic resonance imaging. Each number word could either be merged with the previous ones (e.g., 'twenty three') or not (e.g., 'three twenty'), thus forming four levels ranging from lists of number words to complex numerals. When a number word could be merged with the preceding ones, it was named faster than when it could not. Neuroimaging results showed that the number of merges correlated with activation in the left inferior frontal gyrus and in the left inferior parietal lobule. Consistent findings across Chinese and French participants suggest that these regions serve as the neural bases for forming complex number words in different languages.

Phonological similarity and orthographic similarity affect probed serial recall of Chinese characters.

The previous literature on working memory (WM) has indicated that verbal materials are dominantly retained in phonological representations, whereas other linguistic information (e.g., orthography, semantics) only contributes to verbal WM minimally, if not negligibly. Although accumulating evidence has suggested that multiple linguistic components jointly support verbal WM, the visual/orthographic contribution has rarely been addressed in alphabetic languages, possibly due to the difficulty of dissociating the effects of word forms from the effects of their pronunciations in relatively shallow orthography. In the present study, we examined whether the orthographic representations of Chinese characters support the retention of verbal materials in this language of deep orthography. In Experiments 1a and 2, we independently manipulated the phonological and orthographic similarity of horizontal and vertical characters, respectively, and found that participants' accuracy of probed serial recall was reduced by both similar pronunciations and shared phonetic radicals in the to-be-remembered stimuli. Moreover, Experiment 1b showed that only the effect of phonological, but not that of orthographic, similarity was affected by concurrent articulatory suppression. Taken together, the present results indicate the indispensable contribution of orthographic representations to verbal WM of Chinese characters, and suggest that the linguistic characteristics of a specific language not only determine long-term linguistic-processing mechanisms, but also delineate the organization of verbal WM for that language.

Big time is not always long: numerical magnitude automatically affects time reproduction.

To reproduce the duration of an event precisely, one needs to represent the temporal information without being influenced by other magnitude attributes (e.g., size) of the event. In the present study, however, task-irrelevant numerical magnitude automatically affected participants' reproduction of the duration of a stimulus. In Experiment 1, participants made key-press responses to reproduce the duration of numbers. Reproduced durations were shorter for small numbers (e.g., 1) than for large numbers (e.g., 9). In contrast, in Experiment 2, participants' reproductions of a standard duration were longer when their key-press response was accompanied by visual presentation of a small number than when it was accompanied by presentation of a large number. These results clearly demonstrate that number-time interference extends beyond simple mapping between stimulus categories and response alternatives. The findings support the notion that either a common magnitude representation or closely connected magnitude representations underlie numerical and temporal processing.

Dynamic allocation of attention to metrical and grouping accents in rhythmic sequences.

Most people find it easy to perform rhythmic movements in synchrony with music, which reflects their ability to perceive the temporal periodicity and to allocate attention in time accordingly. Musicians and non-musicians were tested in a click localization paradigm in order to investigate how grouping and metrical accents in metrical rhythms influence attention allocation, and to reveal the effect of musical expertise on such processing. We performed two experiments in which the participants were required to listen to isochronous metrical rhythms containing superimposed clicks and then to localize the click on graphical and ruler-like representations with and without grouping structure information, respectively. Both experiments revealed metrical and grouping influences on click localization. Musical expertise improved the precision of click localization, especially when the click coincided with a metrically strong beat. Critically, although all participants located the click accurately at the beginning of an intensity group, only musicians located it precisely when it coincided with a strong beat at the end of the group. Removal of the visual cue of grouping structures enhanced these effects in musicians and reduced them in non-musicians. These results indicate that musical expertise not only enhances attention to metrical accents but also heightens sensitivity to perceptual grouping.

Enhanced left inferior frontal to left superior temporal effective connectivity for complex sentence comprehension: fMRI evidence from Chinese relative clause processing.

Previous studies investigating the processing of complex sentences have demonstrated the involvement of the left inferior frontal gyrus (LIFG) and left superior temporal gyrus (LSTG), which might subserve ordering and storage of linguistic components, respectively, for sentence comprehension. However, how these brain regions are interconnected, especially during the processing of Chinese sentences, need to be further explored. In this study, the neural network supporting the comprehension of Chinese relative clause was identified. Both the LIFG and LSTG exhibited higher activation in processing subject-extracted relative clauses (SRCs) than object-extracted relative clauses (ORCs). Moreover, a Granger causality analysis revealed that the effective connectivity from the LIFG to LSTG was significant only when participants read Chinese SRCs, which were argued to be more difficult than ORCs. Contrary to the observations of an SRC advantage in most other languages, the present results provide clear neuroimaging evidence for an ORC advantage in Chinese.

A Minimum Temporal Window for Direction Detection of Frequency-Modulated Sweeps: A Magnetoencephalography Study.

The ability to rapidly encode the direction of frequency contour contained in frequency-modulated (FM) sweeps is essential for speech processing, music appreciation, and conspecific communications. Psychophysical evidence points to a common temporal window threshold for human listeners in processing rapid changes in frequency glides. No neural evidence has been provided for the existence of a cortical temporal window threshold underlying the encoding of rapid transitions in frequency glides. The present magnetoencephalography study used the cortical mismatch negativity activity (MMNm) to investigate the minimum temporal window required for detecting different magnitudes of directional changes in frequency-modulated sweeps. A deviant oddball paradigm was used in which directional upward or downward frequency sweep serves as the standard and the same type of sweep with the opposite direction serves as its deviant. Stimuli consisted of unidirectional linear frequency-sweep complexes that swept across speech-relevant frequency bands in durations of 10, 20, 40, 80, 160, and 320 ms (with corresponding rates of 50, 25, 12.5, 6.2, 3.1, 1.5 oct/s). The data revealed significant magnetic mismatch field responses across all sweep durations, with slower-rate sweeps eliciting larger MMNm responses. A greater temporally related enhancement in MMNm response was obtained for rising but not falling frequency sweep contours. A hemispheric asymmetry in the MMNm response pattern was observed corresponding to the directionality of frequency sweeps. Contrary to psychophysical findings, we report a temporal window as short as 10 ms sufficient to elicit a robust MMNm response to a directional change in speech-relevant frequency contours. The results suggest that auditory cortex requires extremely brief temporal window to implicitly differentiate a dynamic change in frequency of linguistically relevant pitch contours. That the brain is extremely sensitive to fine spectral changes contained in speech-relevant glides provides cortical evidence for the ecological importance of FM sweeps in speech processing.

Cortical encoding of linguistic constituent with and without morphosyntactic cues.

This study examined the brain areas involved in combining words into larger units when there are few or no morphosyntactic cues. We manipulated constituent length in word strings of the same length under two conditions: Mandarin sentence, which had sparse morphosyntactic cues, and nominal phrase that had no morphosyntactic cues [e.g., ((honey mustard) (chicken burger))]. Contrasting sentences to word lists revealed a network that largely overlapped with the one reported in languages with rich morphosyntactic cues, including left IFGorb/IFGtri and areas along left STG/STS. Both conditions showed increased activation in left IFGtri/IFGorb in functional ROIs defined based on previous study in sentence processing, while the nominal phrases additionally revealed a constituent length effect in bilateral dorsal IFGtri, left IFGoper, left pMTG/pSTG, left IPL, and several subcortical areas, which might reflect an increased reliance on semantic and pragmatic information. Moreover, in upper left IFGtri/IFGoper and left thalamus/caudate, this effect increased with the participants' tendency to combine nouns into phrases. The absence of syntactic constraints on linguistic composition might highlight individual differences in cognitive control, which helps to integrate non-syntactic information.

The common magnitude code underlying numerical and size processing for action but not for perception.

The interaction between numbers and action-related process has received increasing attention in the literature of numerical cognition. In the current study, two dual-task experiments were conducted to explore the interaction among numerical, prehension, and perceptual color/size judgments. The results revealed the commonality and distinctness of the magnitude representations that are involved in these tasks. Specifically, a photograph of a graspable object with a superimposed Arabic digit was presented in each trial. Participants were required to first judge the parity of the digit with a manual response while simultaneously planning a subsequent vocal response pertaining to the depicted object. When parity and action judgments were performed close in time, the compatibility effect between the numerical magnitude of the digit and the appropriate action (pinch vs. clutch) for the object was demonstrated in both manual and vocal responses. In contrast, such compatibility effect was absent when parity judgment was coupled with color-related or perceptual size judgment. The findings of the current study support the existence of a common magnitude code underlying numerical and non-numerical dimensions for action-related purposes, as proposed by the ATOM model (Walsh in Trends Cogn Sci 7:483-488, 2003). Furthermore, based on the selective presence of the compatibility effect, we argue that the interaction among different quantity dimensions conforms to the "dorsal-action and ventral-perception" organizational principle of the human brain.

Preference for Object Relative Clauses in Chinese Sentence Comprehension: Evidence From Online Self-Paced Reading Time.

Most prior studies have reported that subject-extracted relative clauses (SRCs) are easier to process than object-extracted relative clauses (ORCs). However, whether such an SRC preference is universal across different languages remains an open question. Several reports from Chinese have provided conflicting results; thus, in the present study, we conducted two self-paced reading experiments to examine the comprehension of Chinese relative clauses. The results demonstrated a clear ORC preference that Chinese ORCs were easier to comprehend than Chinese SRCs. These findings were most compatible with the prediction of the integration cost account, which claims that the processing difference between SRCs and ORCs arises at the point of dependency formation. The ORC preference in Chinese poses a challenge to the universality of the SRC preference assumed by the structural distance hypothesis and highlights the values of cross-linguistic research.

Neural substrates of processing path and manner information of a moving event.

Languages consistently distinguish the path and the manner of a moving event in different constituents, even if the specific constituents themselves vary across languages. Children also learn to categorize moving events according to their path and manner at different ages. Motivated by these linguistic and developmental observations, we employed fMRI to test the hypothesis that perception of and attention to path and manner of motion is segregated neurally. Moreover, we hypothesize that such segregation respects the "dorsal-where and ventral-what" organizational principle of vision. Consistent with this proposal, we found that attention to the path of a moving event was associated with greater activity within bilateral inferior/superior parietal lobules and the frontal eye-field, while attention to manner was associated with greater activity within bilateral postero-lateral inferior/middle temporal regions. Our data provide evidence that motion perception, traditionally considered as a dorsal "where" visual attribute, further segregates into dorsal path and ventral manner attributes. This neural segregation of the components of motion, which are linguistically tagged, points to a perceptual counterpart of the functional organization of concepts and language.

Flexible spatial mapping of different notations of numbers in Chinese readers.

The spatial component of numerical and ordinal information has been explored in previous research. However, how such mapping emerges and how it is affected by the learning experience are issues still under debate. In the current study, we examined the orientation of the mental number line for different numerical notations (e.g., "1", "---", "[symbol: see text]") in Chinese readers. Our data demonstrated that Arabic numbers are mentally aligned horizontally with a left-to-right directionality, while Chinese number words are aligned vertically with a top-to-bottom directionality. These findings indicate that different notations of the same concept have flexible mappings within space, which is plausibly shaped by the dominant context in which the numerical notations appear.

Is There a Processing Preference for Object Relative Clauses in Chinese? Evidence From ERPs.

A consistent finding across head-initial languages, such as English, is that subject relative clauses (SRCs) are easier to comprehend than object relative clauses (ORCs). However, several studies in Mandarin Chinese, a head-final language, revealed the opposite pattern, which might be modulated by working memory (WM) as suggested by recent results from self-paced reading performance. In the present study, event-related potentials (ERPs) were recorded when participants with high and low WM spans (measured by forward digit span and operation span tests) read Chinese ORCs and SRCs. The results revealed an N400-P600 complex elicited by ORCs on the relativizer, whose magnitude was modulated by the WM span. On the other hand, a P600 effect was elicited by SRCs on the head noun, whose magnitude was not affected by the WM span. These findings paint a complex picture of relative clause processing in Chinese such that opposing factors involving structural ambiguities and integration of filler-gap dependencies influence processing dynamics in Chinese relative clauses.

Neural correlates of processing sentences and compound words in Chinese.

Sentence reading involves multiple linguistic operations including processing of lexical and compositional semantics, and determining structural and grammatical relationships among words. Previous studies on Indo-European languages have associated left anterior temporal lobe (aTL) and left interior frontal gyrus (IFG) with reading sentences compared to reading unstructured word lists. To examine whether these brain regions are also involved in reading a typologically distinct language with limited morphosyntax and lack of agreement between sentential arguments, an FMRI study was conducted to compare passive reading of Chinese sentences, unstructured word lists and disconnected character lists that are created by only changing the order of an identical set of characters. Similar to previous findings from other languages, stronger activation was found in mainly left-lateralized anterior temporal regions (including aTL) for reading sentences compared to unstructured word and character lists. On the other hand, stronger activation was identified in left posterior temporal sulcus for reading unstructured words compared to unstructured characters. Furthermore, reading unstructured word lists compared to sentences evoked stronger activation in left IFG and left inferior parietal lobule. Consistent with the literature on Indo-European languages, the present results suggest that left anterior temporal regions subserve sentence-level integration, while left IFG supports restoration of sentence structure. In addition, left posterior temporal sulcus is associated with morphological compounding. Taken together, reading Chinese sentences engages a common network as reading other languages, with particular reliance on integration of semantic constituents.

When "He" Can Also Be "She": An ERP Study of Reflexive Pronoun Resolution in Written Mandarin Chinese.

The gender information in written Chinese third person pronouns is not symmetrically encoded: the character for "he" (, with semantic radical , meaning human) is used as a default referring to every individual, while the character for "she" (, with semantic radical , meaning woman) indicates females only. This critical feature could result in different patterns of processing of gender information in text, but this is an issue that has seldom been addressed in psycholinguistics. In Chinese, the written forms of the reflexive pronouns are composed of a pronoun plus the reflexive "/self" (/himself and /herself). The present study focuses on how such gender specificity interacts with the gender type of an antecedent, whether definitional (proper name) or stereotypical (stereotypical role noun) during reflexive pronoun resolution. In this event-related potential (ERP) study, gender congruity between a reflexive pronoun and its antecedent was studied by manipulating the gender type of antecedents and the gender specificity of reflexive pronouns (default: /himself vs. specific: /herself). Results included a P200 "attention related" congruity effect for /himself and a P600 "integration difficulty" congruity effect for /herself. Reflexive pronoun specificity independently affected the P200 and N400 components. These results highlight the role of /himself as a default applicable to both genders and indicate that only the processing of /herself supports a two-stage model for anaphor resolution. While both reflexive pronouns are evaluated at the bonding stage, the processing of the gender-specific reflexive pronoun is completed in the resolution stage.

The dimensional arrow: agreement in directional mapping of dimensions among Mandarin Chinese- and English-speakers.

We present evidence that English- and Mandarin-speakers agree about how to map dimensions (e.g., size and clarity) to vertical space and that they do so in a directional way. We first developed visual stimuli for four dimensions-size, clarity, complexity, and darkness-and in each case we varied the stimuli to express a range of the dimension (e.g., there were five total items expressing the range covering big, medium, and small). In our study, English- and Mandarin-speakers mapped these stimuli to an unlabelled vertical scale. Most people mapped dimensional endpoints in similar ways; using size as a standard, we found that the majority of participants mapped the clearest, most complex, and darkest items to the same end of the vertical scale as they mapped the biggest items. This indicates that all four dimensions have a weighted or unmarked end (i.e., all are directional or polar). The strong similarities in polarity across language groups contrasted with group differences on a lexical task, for which there was little cross-linguistic agreement about which comparative words to use to describe stimulus pairs (e.g., "bigger" vs. "smaller"). Thus, we found no evidence in this study that the perception of these dimensions is influenced by language.