FMRI of two measures of phonological processing in visual word recognition: ecological validity matters

Previous functional magnetic resonance imaging (fMRI) studies have investigated the role of phonological processing by utilizing nonword rhyming decision tasks (e.g., Pugh et al., 1996). Although such tasks clearly engage phonological components of visual word recognition, it is clear that decision tasks are more cognitively involved than the simple overt naming tasks, which more closely map onto normal reading behavior. Our research aim for this study was to examine the advantages of overt naming tasks for fMRI studies of word recognition processes. Process models are presented to highlight the similarities and differences between two cognitive tasks that are used in the word recognition literature, pseudohomophone naming (e.g., pronounce BRANE) and rhyming decision (e.g., do LEAT and JEAT rhyme?). An fMRI study identified several differences in cortical activation associated with the differences observed in the process models. Specifically, the results show that the overt naming task involved the insular cortex and inferior frontal gyrus, whereas the rhyming decision task engaged the temporal-parietal regions. It is argued that future fMRI research examining the neuroanatomical components of basic visual word recognition utilize overt naming tasks.     

语言经验对大脑激活的影响：来自第二语言初学者的证据

研究采用功能磁共振成像技术考察语言经验在塑造大脑激活模式中的作用。12名只有很少英语学习经验因而熟练程度很低的小学儿童参与了实验。结果发现,在视觉呈现的押韵判断任务中,英文任务和中文任务共同激活了左侧额下回负责语音加工的脑区。更重要的是,虽然英文任务更难,也更多地激活了双侧的顶叶区域,但是它在额叶诱发的激活强度显著低于中文。这个结果进一步说明第二语言的语音皮层表征是随着学习经验的增加而逐渐发展起来的。     

The functional anatomy of word comprehension and production

This review describes the functional anatomy of word comprehension and production. Data from functional neuroimaging studies of normal subjects are used to determine the distributed set of brain regions that are engaged during particular language tasks and data from studies of patients with neurological damage are used to determine which of these regions are necessary for task performance. This combination of techniques indicates that the left inferior temporal and left posterior inferior parietal cortices are required for accessing semantic knowledge; the left posterior basal temporal lobe and the left frontal operculum are required for translating semantics into phonological output and the left anterior inferior parietal cortex is required for translating orthography to phonology. Further studies are required to establish the specific functions of the different regions and how these functions interact to provide our sophisticated language system.     

自闭症谱系障碍人群词义加工的脑激活模式：基于fMRI研究的元分析

本研究筛选了11项采用功能性磁共振成像技术探究言语自闭症人群词义加工的研究, 探讨了该人群与典型人群脑激活模式的差异是否具有跨研究的稳定性。结果表明, 差异的脑激活模式稳定存在, 且表现为主要涉及左额上回的典型脑区激活不足。该结果为言语ASD人群语言加工的神经机制提供了来自词义加工的跨研究激活证据, 在明确“减弱的额叶激活”这一稳定差异表现的基础上, 强调了针对不同语言加工任务开展元分析研究的必要性。     

Effective brain connectivity in children with reading difficulties during phonological processing

Using Dynamic Causal Modeling (DCM) and functional magnetic resonance imaging (fMRI), we examined effective connectivity between three left hemisphere brain regions (inferior frontal gyrus, inferior parietal lobule, fusiform gyrus) and bilateral medial frontal gyrus in 12 children with reading difficulties (M age=12.4, range: 8.11-14.10) and 12 control children (M age=12.3, range: 8.9-14.11) during rhyming judgments to visually presented words. More difficult conflicting trials either had similar orthography but different phonology (e.g. pint-mint) or similar phonology but different orthography (e.g. jazz-has). Easier non-conflicting trials had similar orthography and phonology (e.g. dime-lime) or different orthography and phonology (e.g. staff-gain). The modulatory effect from left fusiform gyrus to left inferior parietal lobule was stronger in controls than in children with reading difficulties only for conflicting trials. Modulatory effects from left fusiform gyrus and left inferior parietal lobule to left inferior frontal gyrus were stronger for conflicting trials than for non-conflicting trials only in control children but not in children with reading difficulties. Modulatory effects from left inferior frontal gyrus to inferior parietal lobule, from medial frontal gyrus to left inferior parietal lobule, and from left inferior parietal lobule to medial frontal gyrus were positively correlated with reading skill only in control children. These findings suggest that children with reading difficulties have deficits in integrating orthography and phonology utilizing left inferior parietal lobule, and in engaging phonological rehearsal/segmentation utilizing left inferior frontal gyrus possibly through the indirect pathway connecting posterior to anterior language processing regions, especially when the orthographic and phonological information is conflicting.     

Functional connectivity in an fMRI study of semantic and phonological processes and the effect of L-Dopa

We describe an fMRI experiment examining the functional connectivity (FC) between regions of the brain associated with semantic and phonological processing. We wished to explore whether L-Dopa administration affects the interaction between language network components in semantic and phonological categorization tasks, as revealed by FC. We hypothesized that L-Dopa would decrease FC due to restriction of the semantic network. During two test sessions (placebo and L-Dopa) each participant performed two fMRI runs, involving phonological and semantic processing. A number of brain regions commonly activated by the two tasks were chosen as regions if interest: left inferior frontal, left posterior temporal and left fusiform gyri, and left parietal cortex. FC was calculated and further analyzed for effects of either the drug or task. No main effect for drug was found. A significant main effect for task was found, with a greater average correlation for the phonological task than for the semantic task. These findings suggest that language areas are activated in a more synchronous manner for phonological than for semantic tasks. This may relate to the fact that phonological processes are mediated to a greater extent within language areas, whereas semantic tasks likely require greater interaction outside of the language areas. Alternatively, this may be due to differences in the attentional requirements of the two tasks.     

Reversing spoken items--mind twisting not tongue twisting

Using 12 participants we conducted an fMRI study involving two tasks, word reversal and rhyme judgment, based on pairs of natural speech stimuli, to study the neural correlates of manipulating auditory imagery under taxing conditions. Both tasks engaged the left anterior superior temporal gyrus, reflecting previously established perceptual mechanisms. Engagement of the left inferior frontal gyrus in both tasks relative to baseline could only be revealed by applying small volume corrections to the region of interest, suggesting that phonological segmentation played only a minor role and providing further support for factorial dissociation of rhyming and segmentation in phonological awareness. Most importantly, subtraction of rhyme judgment from word reversal revealed activation of the parietal lobes bilaterally and the right inferior frontal cortex, suggesting that the dynamic manipulation of auditory imagery involved in mental reversal of words seems to engage mechanisms similar to those involved in visuospatial working memory and mental rotation. This suggests that reversing spoken items is a matter of mind twisting rather than tongue twisting and provides support for a link between language processing and manipulation of mental imagery.     

Cognitive control and parsing: Reexamining the role of Broca’s area in sentence comprehension

A century of investigation into the role of the human frontal lobes in complex cognition, including language processing, has revealed several interesting but apparently contradictory findings. In particular, the results of numerous studies suggest that left inferior frontal gyrus (LIFG), which includes Broca’s area, plays a direct role in sentence-level syntactic processing. In contrast, other brain-imaging and neuropsychological data indicate that LIFG is crucial for cognitive control—specifically, for overriding highly regularized, automatic processes, even when a task involves syntactically undemanding material (e.g., single words, a list of letters). We provide a unifying account of these findings, which emphasizes the importance of general cognitive control mechanisms for the syntactic processing of sentences. On the basis of a review of the neurocognitive and sentence-processing literatures, we defend the following three hypotheses: (1) LIFG is part of a network of frontal lobe subsystems that are generally responsible for the detection and resolution of incompatible stimulus representations; (2) the role of LIFG in sentence comprehension is to implement reanalysis in the face of misinterpretation; and (3) individual differences in cognitive control abilities in nonsyntactic tasks predict correlated variation in sentence-processing abilities pertaining to the recovery from misinterpretation.     

文盲与非文盲汉字字形和语音加工的脑机制

利用功能性磁共振成像(fMRI)技术探讨文盲和非文盲汉字字形和语音加工脑机制的差异。实验1使用汉字字形和图形比较了中国人文盲和非文盲字形加工过程脑机制的左侧差异。实验2使用汉字语音和纯音比较了文盲和非文盲语音加工过程脑机制的双侧差异。结果表明文盲与非文盲汉字字形和语音加工脑机制不同,且非文盲的脑活动强。     

The pars triangularis in dyslexia and ADHD: A comprehensive approach

Limited research has been conducted on the structure of the pars triangularis (PT) in dyslexia despite functional neuroimaging research finding it may play a role in phonological processing. Furthermore, research to date has not examined PT size in ADHD even though the right inferior frontal region has been implicated in the disorder. Hence, one of the purposes of this study was to examine the structure of the PT in dyslexia and ADHD. The other purposes included examining the PT in relation to overall expressive language ability and in relation to several specific linguistic functions given language functioning often is affected in both dyslexia and ADHD. Participants included 50 children: 10 with dyslexia, 15 with comorbid dyslexia/ADHD, 15 with ADHD, and 10 controls. Using a 2 (dyslexia or not) × 2 (ADHD or not) MANCOVA, findings revealed PT length and shape were comparable between those with and without dyslexia. However, children with ADHD had smaller right PT lengths than those without ADHD, and right anterior ascending ramus length was related to attention problems in the total sample. In terms of linguistic functioning, presence of an extra sulcus in the left PT was related to poor expressive language ability. In those with adequate expressive language functioning, left PT length was related to phonological awareness, phonological short-term memory and rapid automatic naming (RAN). Right PT length was related to RAN and semantic processing. Further work on PT morphology in relation to ADHD and linguistic functioning is warranted.     

The phonological-similarity effect differentiates between two working memory tasks

Working memory is a set of interactive cognitive processes that maintain information on–line and available for analysis. Part of the system is specialized for maintaining verbal information, a core component of which is thought to be a phonological store. On the basis of the study of patients with acquired brain lesions, this store has been localized to the supramarginal and angular gyri of the speech–dominant hemisphere, and some functional neuroimaging studies support this localization. However, other imaging studies localize the phonological store in a more dorsal region of the parietal lobe. To reconcile these findings, we examined the phonological–similarity effect in two different tasks. A phonological–similarity effect was observed only in the task that involved sequential presentation and explicit verbal rehearsal. We conclude that at least one possible source of the differences in brain activation between different working memory tasks may be differences in phonological processing.     

Sex differences in cerebral laterality of language and visuospatial processing

Sex differences on language and visuospatial tasks are of great interest, with differences in hemispheric laterality hypothesized to exist between males and females. Some functional imaging studies examining sex differences have shown that males are more left lateralized on language tasks and females are more right lateralized on visuospatial tasks; however, findings are inconsistent. Here we used functional magnetic resonance imaging to study thirty participants, matched on task performance, during phonological and visuospatial tasks. For each task, region-of-interest analyses were used to test differences in cerebral laterality. Results indicate that lateralization differences exist, with males more left lateralized during the phonological task and showing greater bilateral activity during the visuospatial task, whereas females showed greater bilateral activity during the phonological task and were more right lateralized during the visuospatial task. Our data provide clear evidence for differences in laterality between males and females when processing language versus visuospatial information.     

Effects of frontal lobe damage on interference effects in working memory

Working memory is hypothesized to comprise a collection of distinct components or processes, each of which may have a unique neural substrate. Recent neuroimaging studies have isolated a region of the left inferior frontal gyrus that appears to be related specifically to one such component: resolving interference from previous items in working memory. In the present study, we examined working memory in patients with unilateral frontal lobe lesions by using a modified version of an item recognition task in which interference from previous trials was manipulated. In particular, we focused on patient R.C., whose lesion uniquely impinged on the region identified in the neuroimaging studies of interference effects. We measured baseline working memory performance and interference effects in R.C. and other frontal patients and in age-matched control subjects and young control subjects. Comparisons of each of these groups supported the following conclusions. Normal aging is associated with changes to both working memory and interference effects. Patients with frontal damage exhibited further declines in working memory but normal interference effects, with the exception of R.C., who exhibited a pronounced interference effect on both response time and accuracy. We propose that the left inferior frontal gyrus subserves a general, nonmnemonic function of selecting relevant information in the face of competing alternatives and that this function may be required by some working memory tasks.     

Domain-dependent activation during spatial and nonspatial auditory working memory

Visual system has been proposed to be divided into two, the ventral and dorsal, processing streams. The ventral pathway is thought to be involved in object identification whereas the dorsal pathway processes information regarding the spatial locations of objects and the spatial relationships among objects. Several studies on working memory (WM) processing have further suggested that there is a dissociable domain-dependent functional organization within the prefrontal cortex for processing of spatial and nonspatial visual information. Also the auditory system is proposed to be organized into two domain-specific processing streams, similar to that seen in the visual system. Recent studies on auditory WM have further suggested that maintenance of nonspatial and spatial auditory information activates a distributed neural network including temporal, parietal, and frontal regions but the magnitude of activation within these activated areas shows a different functional topography depending on the type of information being maintained. The dorsal prefrontal cortex, specifically an area of the superior frontal sulcus (SFS), has been shown to exhibit greater activity for spatial than for nonspatial auditory tasks. Conversely, ventral frontal regions have been shown to be more recruited by nonspatial than by spatial auditory tasks. It has also been shown that the magnitude of this dissociation is dependent on the cognitive operations required during WM processing. Moreover, there is evidence that within the nonspatial domain in the ventral prefrontal cortex, there is an across-modality dissociation during maintenance of visual and auditory information. Taken together, human neuroimaging results on both visual and auditory sensory systems support the idea that the prefrontal cortex is organized according to the type of information being maintained in WM.     

押韵加工的认知神经机制

押韵是指一对词语中从最后一个发音的元音到词尾的语音结构均相同的现象。现有押韵加工主要分为押韵识别与押韵产生两个研究领域,两者的认知加工过程相似,包括字形编码、形音转换、语音表征与语音分段等阶段。从语音加工与字形加工两方面对押韵过程的神经基础进行探讨,发现左半球颞上回与额下回分别负责语音表征与语音分段,左半球梭状回参与着字形编码,而有效的形音转换依赖于左半球顶下小叶与额下回组成的神经网络。今后应进一步整合不同研究方法与任务下的研究结果,并对押韵产生加工进行更深入的探讨。     

On Broca, brain, and binding: a new framework

In speaking and comprehending language, word information is retrieved from memory and combined into larger units (unification). Unification operations take place in parallel at the semantic, syntactic and phonological levels of processing. This article proposes a new framework that connects psycholinguistic models to a neurobiological account of language. According to this proposal the left inferior frontal gyrus (LIFG) plays an important role in unification. Research in other domains of cognition indicates that left prefrontal cortex has the necessary neurobiological characteristics for its involvement in the unification for language. I offer here a psycholinguistic perspective on the nature of language unification and the role of LIFG.     

Short-term memory and sentence processing: Evidence from neuropsychology

Traditional models of memory assume that short-term memory, as measured by memory span, plays an important role in linguistic processing and the learning ofverbal information. Contradicting this view are findings from a brain-damaged patient, E.A., who, despite a verbal memory span of about two items, demonstrated normal sentence comprehension in a variety oftasks. She was, however, impaired whenever verbatim phonological information had to be maintained or learned. These results and those from other patients with reduced span suggest that the phonological storage capacity that is critical to memory span plays only a limited role in language processing, specifically in the maintenance and learning of phonological forms. Implications for models of short-term memory are discussed. It is argued that short-termmemory should be seen as deriving from the processing and retentive capacities of language processing modules, with span tasks drawing on only a subset of these modules.     

The neural effect of stimulus-response modality compatibility on dual-task performance: an fMRI study

Recent fMRI studies suggest that the inferior frontal sulcus (IFS) is involved in the coordination of interfering processes in dual-task situations. The present study aims to further specify this assumption by investigating whether the compatibility between stimulus and response modalities modulates dual-task-related activity along the IFS. It has been shown behaviorally that the degree of interference, as measured by dual-task costs, increases in modality-incompatible conditions (e.g. visual–vocal tasks combined with auditory–manual tasks) as compared to modality-compatible conditions (e.g. visual–manual tasks combined with auditory–vocal tasks). Using fMRI, we measured IFS activity when participants performed modality-compatible and modality-incompatible single and dual tasks. Behaviorally, we replicated the finding of higher dual-task costs for modality-incompatible tasks compared to modality-compatible tasks. The fMRI data revealed higher activity along the IFS in modality-incompatible dual tasks compared with modality-compatible dual tasks when inter-individual variability in functional brain organization is taken into account. We argue that in addition to temporal order coordination (Szameitat et al., 2002), the IFS is involved in the coordination of cognitive processes associated with the concurrent mapping of sensory information onto corresponding motor responses in dual-task situations.     

The Parieto-Frontal Integration Theory (P-FIT) of intelligence: converging neuroimaging evidence

"Is there a biology of intelligence which is characteristic of the normal human nervous system?" Here we review 37 modern neuroimaging studies in an attempt to address this question posed by Halstead (1947) as he and other icons of the last century endeavored to understand how brain and behavior are linked through the expression of intelligence and reason. Reviewing studies from functional (i.e., functional magnetic resonance imaging, positron emission tomography) and structural (i.e., magnetic resonance spectroscopy, diffusion tensor imaging, voxel-based morphometry) neuroimaging paradigms, we report a striking consensus suggesting that variations in a distributed network predict individual differences found on intelligence and reasoning tasks. We describe this network as the Parieto-Frontal Integration Theory (P-FIT). The P-FIT model includes, by Brodmann areas (BAs): the dorsolateral prefrontal cortex (BAs 6, 9, 10, 45, 46, 47), the inferior (BAs 39, 40) and superior (BA 7) parietal lobule, the anterior cingulate (BA 32), and regions within the temporal (BAs 21, 37) and occipital (BAs 18, 19) lobes. White matter regions (i.e., arcuate fasciculus) are also implicated. The P-FIT is examined in light of findings from human lesion studies, including missile wounds, frontal lobotomy/leukotomy, temporal lobectomy, and lesions resulting in damage to the language network (e.g., aphasia), as well as findings from imaging research identifying brain regions under significant genetic control. Overall, we conclude that modern neuroimaging techniques are beginning to articulate a biology of intelligence. We propose that the P-FIT provides a parsimonious account for many of the empirical observations, to date, which relate individual differences in intelligence test scores to variations in brain structure and function. Moreover, the model provides a framework for testing new hypotheses in future experimental designs.     

Neural correlates of mapping from phonology to orthography in children performing an auditory spelling task

Age-related differences (9- to 15-year-olds) in the neural correlates of mapping from phonology to orthography were examined with functional magnetic resonance imaging (fMRI). Participants were asked to determine if two spoken words had the same spelling for the rime (corresponding letters after the first consonant or consonant cluster). Some of the word pairs had conflicting orthography and phonology (e.g. jazz-has, pint-mint) whereas other pairs had non-conflicting information (e.g. press-list, gate-hate) (see Table 1). There were age-related increases in activation for lexical processing (across conflicting and non-conflicting conditions) in left inferior parietal lobule, suggesting that older children have a more elaborated system for mapping between phonology and orthography that includes connections at different grain sizes (e.g. phonemes, onset-rimes, syllables). In addition, we found that the conflicting conditions had lower accuracy, slower reaction time and greater activation in left inferior frontal gyrus as compared to non-conflicting conditions. Higher accuracy was also correlated with greater activation in left inferior frontal gyrus for the most difficult conflicting condition (e.g. jazz-has). The finding of both a conflict effect and a correlation with accuracy in left inferior frontal gyrus suggests that this region may be involved in resolving the conflict between orthographic and phonological representations.