Recruitment of anterior and posterior structures in lexical-semantic processing: an fMRI study comparing implicit and explicit tasks

Previous studies examining explicit semantic processing have consistently shown activation of the left inferior frontal gyrus (IFG). In contrast, implicit semantic processing tasks have shown activation in posterior areas including the superior temporal gyrus (STG) and the middle temporal gyrus (MTG) with less consistent activation in the IFG. These results raise the question whether the functional role of the IFG is related to those processes needed to make a semantic decision or to processes involved in the extraction and analysis of meaning. This study examined neural activation patterns during a semantic judgment task requiring overt semantic analysis, and then compared these activation patterns to previously obtained results using the same semantically related and unrelated word pairs in a lexical decision task which required only implicit semantic processing (Rissman, J., Eliassen, J. C., & Blumstein, S. E. (2003). An event-related fMRI investigation of implicit semantic priming. Journal of Cognitive Neuroscience, 15, 1160-1175). The behavioral results demonstrated that the tasks were equivalent in difficulty. fMRI results indicated that the IFG and STG bilaterally showed greater activation for semantically unrelated than related word pairs across the two tasks. Comparison of the two task types across conditions revealed greater activation for the semantic judgment task only in the STG bilaterally and not in the IFG. These results suggest that the pre-frontal cortex is recruited similarly in the service of both the lexical decision and semantic judgment tasks. The increased activation in the STG in the semantic judgment task reflects the greater depth of semantic processing required in this task and indicates that the STG is not simply a passive store of lexical-semantic information but is involved in the active retrieval of this information.     

Prominence vs. aboutness in sequencing: A functional distinction within the left inferior frontal gyrus

Prior research on the neural bases of syntactic comprehension suggests that activation in the left inferior frontal gyrus (lIFG) correlates with the processing of word order variations. However, there are inconsistencies with respect to the specific subregion within the IFG that is implicated by these findings: the pars opercularis or the pars triangularis. Here, we examined the hypothesis that the dissociation between pars opercularis and pars triangularis activation may reflect functional differences between clause-medial and clause-initial word order permutations, respectively. To this end, we directly compared clause-medial and clause-initial object-before-subject orders in German in a within-participants, event-related fMRI design. Our results showed increased activation for object-initial sentences in a bilateral network of frontal, temporal and subcortical regions. Within the lIFG, posterior and inferior subregions showed only a main effect of word order, whereas more anterior and superior subregions showed effects of word order and sentence type, with higher activation for sentences with an argument in the clause-initial position. These findings are interpreted as evidence for a functional gradation of sequence processing within the left IFG: posterior subportions correlate with argument prominence-based (local) aspects of sequencing, while anterior subportions correlate with aboutness-based aspects of sequencing, which are crucial in linking the current sentence to the wider discourse. This proposal appears compatible with more general hypotheses about information processing gradients in prefrontal cortex (Koechlin & Summerfield, 2007).     

Aging affects both perceptual and lexical/semantic components of word stem priming: An event-related fMRI study

In this event-related fMRI study, brain activity patterns were compared in extensive groups of young (N=25) and older (N=38) adults, while they were performing a word stem completion priming task. Based on behavioral findings, we tested the hypothesis that aging affects only the lexical/semantic, but not the perceptual component of word stem priming. To this end, we distinguished between priming-related activity reductions in posterior regions involved in visual processing, and regions associated with lexical/semantic retrieval processes, i.e., left lateral temporal and left prefrontal regions. Both groups revealed significant priming-related response time reductions. However, in accordance with earlier findings, a larger priming effect was found in the group of young participants. In line with previous imaging studies, the groups showed common priming-related activity reductions in the anterior cingulate, and the left inferior prefrontal cortex extending into the anterior portion of the left superior temporal gyrus, and at lower thresholds also in the right occipital lobe. However, when directly comparing the groups, greater priming-related reductions were found for the young group in the left anterior superior temporal gyrus and the right posterior occipital lobe. These findings suggest that, converse to current psychological views, aging affects both perceptual and lexical/semantic components of repetition priming.     

Neural correlates of lexical and sublexical processes in reading

The purpose of the present study was to compare the brain regions and systems that subserve lexical and sublexical processes in reading. In order to do so, three types of tasks were used: (i). silent reading of very high frequency regular words (lexical task); (ii). silent reading of nonwords (sublexical task); and, (iii). silent reading of very low frequency regular words (sublexical task). All three conditions were contrasted with a visual/phonological baseline condition. The lexical condition engaged primarily an area at the border of the left angular and supramarginal gyri. Activation found in this region suggests that this area may be involved in mapping orthographic-to-phonological whole word representations. Both sublexical conditions elicited significantly greater activation in the left inferior prefrontal gyrus. This region is thought to be associated with sublexical processes in reading such as grapheme-to-phoneme conversion, phoneme assembly and underlying verbal working memory processes. Activation in the left IFG was also associated with left superior and middle temporal activation. These areas are thought to be functionally correlated with the left IFG and to contribute to a phonologically based form of reading. The results as a whole demonstrate that lexical and sublexical processes in reading activate different regions within a complex network of brain structures.     

Adult age differences in visual word identification: functional neuroanatomy by positron emission tomography

Adult age differences in the neural systems mediating semantic (context-independent) memory were investigated using positron emission tomography (PET). Younger (20-29 years) and older (62-70 years) participants performed lexical decision (word/nonword discrimination) and nonsemantic (simple visual search) baseline tasks during PET scanning. Within the lexical decision task, display duration and presentation rate were varied across scans. The behavioral data suggested that although an age-related slowing was evident in visual feature and response processing, the retrieval of semantic/lexical information was similar for younger and older adults. For both age groups, lexical-related activation occurred in inferior prefrontal and occipitotemporal regions of the left hemisphere. Differential activation, as a function of age group, was observed in the left occipitotemporal pathway as a result of older adults' maintaining higher levels of neural activity in striate cortex (during visual search) and in inferior temporal cortex (during lexical decision). The prefrontal activation was similar for the two age groups. Thus, although this form of semantic memory retrieval does not undergo significant age-related decline, an age-related change in the associated pattern of neural activation is evident. These findings differ from previous neuroimaging studies of episodic (context-dependent) memory retrieval, which have suggested that age-related compensatory mechanisms are expressed primarily by greater activation of prefrontal regions for older adults than for younger adults.     

GRAMMATICAL GENDER IS ON THE TIP OF ITALIAN TONGUES

Abstract— To correctly produce words, speakers must have access to three broad classes of information lexical semantics, syntax, and sound structure. The relevant information must be organized in ways that permit rapid and accurate retrieval of specific lexical targets. Current models of language production do this by a two-stage process. The first stage incorporates lexical meanings and syntax, and the second, sound structure. We used studies of the tip-of-the-tongue phenomenon (i e, the condition in which a speaker cannot produce a well-known word) to evaluate this organization and in so doing, we provide the first clear experimental evidence for a lexical stage that includes syntax and is distinct from both sound structure and the conceptual correlates of syntactic features     

Reading and lexicalization in opaque and transparent orthographies: Word naming and word learning in English and Spanish

Do skilled readers of opaque and transparent orthographies make differential use of lexical and sublexical processes when converting words from print to sound? Two experiments are reported, which address that question, using effects of letter length on naming latencies as an index of the involvement of sublexical letter–sound conversion. Adult native speakers of English (Experiment 1) and Spanish (Experiment 2) read aloud four- and seven-letter high-frequency words, low-frequency words, and nonwords in their native language. The stimuli were interleaved and presented 10 times in a first testing session and 10 more times in a second session 28 days later. Effects of lexicality were observed in both languages, indicating the deployment of lexical representations in word naming. Naming latencies to both words and nonwords reduced across repetitions on Day 1, with those savings being retained to Day 28. Length effects were, however, greater for Spanish than English word naming. Reaction times to long and short nonwords converged with repeated presentations in both languages, but less in Spanish than in English. The results support the hypothesis that reading in opaque orthographies favours the rapid creation and use of lexical representations, while reading in transparent orthographies makes more use of a combination of lexical and sublexical processing.     

The influence of speech rate and accent on access and use of semantic information

Circumstances in which the speech input is presented in sub-optimal conditions generally lead to processing costs affecting spoken word recognition. The current study indicates that some processing demands imposed by listening to difficult speech can be mitigated by feedback from semantic knowledge. A set of lexical decision experiments examined how foreign accented speech and word duration impact access to semantic knowledge in spoken word recognition. Results indicate that when listeners process accented speech, the reliance on semantic information increases. Speech rate was not observed to influence semantic access, except in the setting in which unusually slow accented speech was presented. These findings support interactive activation models of spoken word recognition in which attention is modulated based on speech demands.     

Co-occurring anxiety influences patterns of brain activity in depression

Brain activation associated with anhedonic depression and co-occurring anxious arousal and anxious apprehension was measured by fMRI during performance of an emotion word Stroop task. Consistent with EEG findings, depression was associated with rightward frontal lateralization in the dorsolateral prefrontal cortex (DLPFC), but only when anxious arousal was elevated and anxious apprehension was low. Activity in the right inferior frontal gyrus (IFG) was also reduced for depression under the same conditions. In contrast, depression was associated with more activity in the anterior cingulate cortex (dorsal ACC and rostral ACC) and the bilateral amygdala. Results imply that depression, particularly when accompanied by anxious arousal, may result in a failure to implement top-down processing by appropriate brain regions (left DLPFC, right IFG) due to increased activation in regions associated with responding to emotionally salient information (right DLPFC, amygdala).     

Quantifying the Interplay of Semantics and Phonology During Failures of Word Retrieval by People With Aphasia Using a Multiplex Lexical Network

Investigating instances where lexical selection fails can lead to deeper insights into the cognitive machinery and architecture supporting successful word retrieval and speech production. In this paper, we used a multiplex lexical network approach that combines semantic and phonological similarities among words to model the structure of the mental lexicon. Network measures at different levels of analysis (degree, network distance, and closeness centrality) were used to investigate the influence of network structure on picture naming accuracy and errors by people with Anomic, Broca's, Conduction, and Wernicke's aphasia. Our results reveal that word retrieval is influenced by the multiplex lexical network structure in at least two ways—(a) the accuracy of production and error type on incorrect productions were influenced by the degree and closeness centrality of the target word, and (b) error type also varied in terms of network distance between the target word and produced error word. Taken together, the analyses demonstrate that network science techniques, particularly the use of the multiplex lexical network to simultaneously represent semantic and phonological relationships among words, reveal how the structure of the mental lexicon influences language processes beyond traditionally examined psycholinguistic variables. We propose a framework for how the multiplex lexical network approach allows for understanding the influence of mental lexicon structure on word retrieval processes, with an eye toward a better understanding of the nature of clinical impairments, like aphasia.     

The role of lexical competition and acoustic-phonetic structure in lexical processing: evidence from normal subjects and aphasic patients

This study examined the effects that the acoustic-phonetic structure of a stimulus exerts on the processes by which lexical candidates compete for activation. An auditory lexical decision paradigm was used to investigate whether shortening the VOT of an initial voiceless stop consonant in a real word results in the activation of the lexical-semantic network of its voiced competitor, i.e., does acoustically modified time prime penny via dime. Results for normal subjects showed semantic priming for related pairs and mediated priming for voiced competitors, consistent with cascade models of language processing allowing for interaction between phonological and semantic levels of processing. Although Broca's aphasics showed semantic priming (dime primed penny), they failed to show priming in the context of a lexical competitor. These results are consistent with the hypothesis that these patients have a lexical processing deficit characterized by an overall reduction in lexical activation.     

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.     

Reduction of left visual field lexical decision accuracy as a result of concurrent nonverbal auditory stimulation

To investigate whether concurrent nonverbal sound sequences would affect visual-hemifield lexical processing, lexical-decision performance of 24 strongly right-handed students (12 men, 12 women) was measured in three conditions: baseline, concurrent neutral sound sequence, and concurrent emotional sound sequence. With the neutral sequence, Naveteru, Roy, Ovelac, and Steinling (1992) had observed a right greater than left cerebral blood flow, and an opposite pattern with the emotional sequence. In the present study, the neutral sound sequence induced a significant accuracy reduction for lexical decisions to stimuli presented in the left visual field. It is hypothesized that RH activation in response to neutral sounds interferes with the limited lexical processing resources of that hemisphere.     

Sex differences in semantic language processing: a functional MRI study

Predictions based on two models of sex differences in cerebral organization of language were compared by examining fMRI patterns of 10 females and 9 males during a semantic processing task. Both groups displayed activation of left inferior frontal gyrus (IFG), left superior temporal gyrus (STG), and cingulate. Females, but not males, showed bilateral IFG and STG activation. Further analyses revealed females had less diffuse left activation and greater right posterior temporal and insula region activation than males. Results support both an interhemispheric and an intrahemispheric model of sex differences in language, suggesting that the models may not be mutually exclusive.     

Individual differences in skilled adult readers reveal dissociable patterns of neural activity associated with component processes of reading

We used fMRI to examine patterns of brain activity associated with component processes of visual word recognition and their relationships to individual differences in reading skill. We manipulated both the judgments adults made on written stimuli and the characteristics of the stimuli. Phonological processing led to activation in left inferior frontal and temporal regions whereas semantic processing was associated with bilateral middle frontal activation. Individual differences in reading subskills were reflected in differences in the degree to which cortical regions were engaged during reading. Variation in sight word reading efficiency was associated with degree of activation in visual cortex. Increased phonological decoding skill was associated with greater activation in left temporo-parietal cortex. Greater reading comprehension ability was associated with decreased activation in anterior cingulate and temporal regions. Notably, associations between reading ability and neural activation indicate that brain/behavior relationships among skilled readers differ from patterns associated with dyslexia and reading development.     

Syntactic structure building in the anterior temporal lobe during natural story listening

The neural basis of syntax is a matter of substantial debate. In particular, the inferior frontal gyrus (IFG), or Broca’s area, has been prominently linked to syntactic processing, but the anterior temporal lobe has been reported to be activated instead of IFG when manipulating the presence of syntactic structure. These findings are difficult to reconcile because they rely on different laboratory tasks which tap into distinct computations, and may only indirectly relate to natural sentence processing. Here we assessed neural correlates of syntactic structure building in natural language comprehension, free from artificial task demands. Subjects passively listened to Alice in Wonderland during functional magnetic resonance imaging and we correlated brain activity with a word-by-word measure of the amount syntactic structure analyzed. Syntactic structure building correlated with activity in the left anterior temporal lobe, but there was no evidence for a correlation between syntactic structure building and activity in inferior frontal areas. Our results suggest that the anterior temporal lobe computes syntactic structure under natural conditions.     

Word and letter string processing networks in schizophrenia: evidence for anomalies and compensation

Imaging studies show that in normal language correlated activity between anterior and posterior brain regions increases as the linguistic and semantic content (i.e., from false fonts, letter strings, pseudo words, to words) of stimuli increase. In schizophrenia however, disrupted functional connectivity between frontal and posterior brain regions has been frequently reported and these disruptions may change the nature of language organization. We characterized basic linguistic operations in word and letter string processing in a region-of-interest network using structural equation modeling (SEM). Healthy volunteers and volunteers with schizophrenia performed an fMRI one-back matching task with real words and consonant letter strings. We hypothesized that left hemisphere network dysfunction in schizophrenia would be present during processes dealing with linguistic/semantic content. The modeling results suggest aberrant left hemisphere function in schizophrenia, even in tasks requiring minimal access to language. Alternative mechanisms included increases in right hemisphere involvement and increased top-down influence from frontal to posterior regions.     

Semantic priming is affected by real-time phonological competition: Evidence for continuous cascading systems

Lexical-semantic access is affected by the phonological structure of the lexicon. What is less clear is whether such effects are the result of continuous activation between lexical form and semantic processing or whether they arise from a more modular system in which the timing of accessing lexical form determines the timing of semantic activation. This study examined this issue using the visual world paradigm by investigating the time course of semantic priming as a function of the number of phonological competitors. Critical trials consisted of high or low density auditory targets (e.g., horse) and a visual display containing a target, a semantically related object (e.g., saddle), and two phonologically and semantically unrelated objects (e.g., chimney, bikini). Results showed greater magnitude of priming for semantically related objects of low than of high density words, and no differences for high and low density word targets in the time course of looks to the word semantically related to the target. This pattern of results is consistent with models of cascading activation, which predict that lexical activation has continuous effects on the level of semantic activation, with no delays in the onset of semantic activation for phonologically competing words.     

The influence of syllable onset complexity and syllable frequency on speech motor control

Functional imaging studies have delineated a "minimal network for overt speech production", encompassing mesiofrontal structures (supplementary motor area, anterior cingulate gyrus), bilateral pre- and postcentral convolutions, extending rostrally into posterior parts of the inferior frontal gyrus (IFG) of the language-dominant hemisphere, left anterior insula as well as bilateral components of the basal ganglia, the cerebellum, and the thalamus. In order to further elucidate the specific contribution of these cerebral regions to speech motor planning, subjects were asked to read aloud visually presented bisyllabic pseudowords during functional magnetic resonance imaging (fMRI). The test stimuli systematically varied in onset complexity (CCV versus CV) and frequency of occurrence (high-frequency, HF versus low-frequency, LF) of the initial syllable. A cognitive subtraction approach revealed a significant main effect of syllable onset complexity (CCV versus CV) at the level of left posterior IFG, left anterior insula, and both cerebellar hemispheres. Conceivably, these areas closely cooperate in the sequencing of subsyllabic aspects of the sound structure of verbal utterances. A significant main effect of syllable frequency (LF versus HF), by contrast, did not emerge. However, calculation of the time series of hemodynamic activation within the various cerebral structures engaged in speech motor control revealed this factor to enhance functional connectivity between Broca's area and ipsilateral anterior insula.     

Age-Specific Effects of Lexical–Semantic Networks on Word Production

The lexical–semantic organization of the mental lexicon is bound to change across the lifespan. Nevertheless, the effects of lexical–semantic factors on word processing are usually based on studies enrolling young adult cohorts. The current study aims to investigate to what extent age-specific semantic organization predicts performance in referential word production over the lifespan, from school-age children to older adults. In Study 1, we conducted a free semantic association task with participants from six age-groups (ranging from 10 to 80 years old) to compute measures that capture age-specific properties of the mental lexicon across the lifespan. These measures relate to lifespan changes in the Available Richness of the mental lexicon and in the lexical–semantic Network Prototypicality of concrete words. In Study 2, we used the collected data to predict performance in a picture-naming task on a new group of participants within the same age-groups as for Study 1. The results show that age-specific semantic Available Richness and Network Prototypicality affect word production speed while the semantic variables collected only in young adults do not. A richer and more prototypical semantic network across subjects from a given age-group is associated with faster word production speed. The current results indicate that age-specific semantic organization is crucial to predict lexical–semantic behaviors across the lifespan. Similarly, these results also provide cues to the understanding of the lexical–semantic properties of the mental lexicon and to lexical selection in referential tasks.