Stimulus variability improves generalization following response inhibition training.

The present study examined the effect of stimulus variability and practice order on generalization to novel stimuli following a single session of response inhibition training. Ninety-six young adults practiced the Go/No-go task online in three training conditions: (1) constant (N = 32)-inhibition practiced on one stimulus; (2) variable-blocked (N = 32)-inhibition practiced on 6 stimuli, each in a separate block; and (3) variable-random (N = 32)-inhibition practiced on 6 stimuli in random order. Generalization was measured by comparing groups on inhibition of novel stimuli and a trained stimulus immediately and 24 h after training. Consistent with our hypothesis, the variable-random and the variable-blocked groups showed better generalization to the novel items than the constant group, demonstrating the benefit of stimulus variability. The variable-random group also showed better generalization than the variable-blocked group, demonstrating the benefit of presenting stimuli in random order. Participants' capacity for working memory maintenance was found to modulate the effect of practice order. While the benefit of variability was retained 24 h after training, the effect of order was not. Results also show generalization to (1) different type of stimuli using the same task and (2) the same stimuli on a different response inhibition task (the Stop-Signal Task), however, the effect of variable practice and order were not evident in these cases. The study findings illustrate the advantage of using variable stimuli presented in random order for generalization and suggest that these principles of motor learning can be applied to learning of cognitive skills.

Inter- and intra- hemispheric interactions in reading ambiguous words.

The present study investigated how the brain processes words with multiple meanings. Specifically, we examined the inter- and intra-hemispheric connectivity of unambiguous words compared to two types of ambiguous words: homophonic homographs, which have multiple meanings mapped to a single phonological representation and orthography, and heterophonic homographs, which have multiple meanings mapped to different phonological representations but the same orthography. Using a semantic relatedness judgment task and effective connectivity analysis via Dynamic Causal Modeling (DCM) on previously published fMRI data (Bitan et al., 2017), we found that the two hemispheres compete in orthographic processing during the reading of unambiguous words. For heterophonic homographs, we observed increased connectivity within the left hemisphere, highlighting the importance of top-down re-activation of orthographic representations by phonological ones for considering alternative meanings. For homophonic homographs, we found a flow of information from the left to the right hemisphere and from the right to the left, indicating that the brain retrieves different meanings using different pathways. These findings provide novel insights into the complex mechanisms involved in language processing and shed light on the different communication patterns within and between hemispheres during the processing of ambiguous and unambiguous words.

Decomposition in early stages of learning novel morphologically derived words: The impact of linear vs. non-linear structure.

We examined whether morphological decomposition takes place in early stages of learning a novel language, and whether morphological structure (linear vs. non-linear) influences decomposition. Across four sessions, 41 native-Hebrew speakers learned morphologically derived words in a novel morpho-lexicon, with two complex conditions: linear and non-linear; and a third simple condition with monomorphemic words. Participants showed faster learning of trained words in the linear condition, and better generalization to untrained words for both complex conditions compared to the simple condition, with better performance for linear than non-linear morphology. Learning the root morpheme, which provides a concrete meaning, was better than learning template/suffix morphemes, which are more abstract. Overall, our results suggest that saliency of discrete units plays an important role in decomposition in early stages of learning derived words, even for speakers highly familiar with the non-linear structure in their L1.

Impaired extraction and consolidation of morphological regularities in developmental dyslexia: A domain general deficit?

The current study examined whether adults with Developmental Dyslexia are impaired in learning linguistic regularities in a novel language, and whether this may be explained by a domain general deficit in the effect of sleep on consolidation. We compared online learning and offline consolidation of morphological regularities in individuals with Developmental Dyslexia (N = 40) and typical readers (N = 38). Participants learned to apply plural inflections to novel words based on morpho-phonological rules embedded in the input and learned to execute a finger motor sequence task. To test the effects of time and sleep on consolidation, participants were assigned into one of two sleep-schedule groups, trained in the evening or in the morning and tested 12 and 24 h later. Unlike typical readers, Dyslexic readers did not extract the morpho-phonological regularities during training and as a group they did not show offline gains in inflecting trained items 24 h after training, suggesting that the deficit in extraction of regularities during training may be related to the deficit in consolidation. The offline gains in dyslexic readers, were correlated with their prior phonological abilities, and were less affected by sleep than those of typical readers. Although no deficit was found in the consolidation of the motor task, dyslexic readers were again less successful in generating an abstract representation of the motor sequence, reflected in a difficulty to generalize the motor sequence knowledge acquired using one hand to the untrained hand. The results suggest that individuals with Developmental Dyslexia have a domain general deficit in extracting statistical regularities from an input. Within the language domain this deficit is reflected in reduced benefits of consolidation, particularly during sleep, perhaps due to reduced prior phonological abilities, which may impede the individual's ability to extract the linguistic regularities during and after training and thus constrain the consolidation process.

Explicit benefits: Motor sequence acquisition and short-term retention in adults who do and do not stutter.

Motor sequencing skills have been found to distinguish individuals who experience developmental stuttering from those who do not stutter, with these differences extending to non-verbal sequencing behaviour. Previous research has focused on measures of reaction time and practice under externally cued conditions to decipher the motor learning abilities of persons who stutter. Without the confounds of extraneous demands and sensorimotor processing, we investigated motor sequence learning under conditions of explicit awareness and focused practice among adults with persistent development stuttering. Across two consecutive practice sessions, 18 adults who stutter (AWS) and 18 adults who do not stutter (ANS) performed the finger-to-thumb opposition sequencing (FOS) task. Both groups demonstrated significant within-session performance improvements, as evidenced by fast on-line learning of finger sequences on day one. Additionally, neither participant group showed deterioration of their learning gains the following day, indicating a relative stabilization of finger sequencing performance during the off-line period. These findings suggest that under explicit and focused conditions, early motor learning gains and their short-term retention do not differ between AWS and ANS. Additional factors influencing motor sequencing performance, such as task complexity and saturation of learning, are also considered. Further research into explicit motor learning and its generalization following extended practice and follow-up in persons who stutter is warranted. The potential benefits of motor practice generalizability among individuals who stutter and its relevance to supporting treatment outcomes are suggested as future areas of investigation.

Start shallow and grow deep: The development of a Hebrew reading brain.

Brain plasticity implies that readers of different orthographies can have different reading networks. Theoretical models suggest that reading acquisition in transparent orthographies relies on mapping smaller orthographic units to phonology, than reading opaque orthographies; but what are the neural mechanisms underlying this difference? Hebrew has a transparent (pointed) script used for beginners, and a non-transparent script used for skilled readers. The current study examined the developmental changes in brain regions associated with phonological and orthographic processes during reading pointed and un-pointed words. Our results highlight some changes that are universal in reading development, such as a developmental increase in frontal involvement (in bilateral inferior frontal gyrus (IFG) pars opercularis), and increase in left asymmetry (in IFG pars opercularis and superior temporal gyrus, STG) of the reading network. Our results also showed a developmental increase in activation in STG, which stands in contrast to previous studies in other orthographies. We further found an interaction of word length and diacritics in bilateral STG and the visual word form area (VWFA) across both groups. These findings suggest that children slightly adjust their reading depending on orthographic transparency, relying on smaller units when reading a transparent script and on larger units when reading an opaque script. Our results also showed that phonological abilities across groups correlated with activation in the VWFA, regardless of transparency, supporting the continued role of phonology at all levels of orthographic transparency. Our findings are consistent with multiple route reading models, in which both phonological and orthographic processing of multiple size units continue to play a role in children's reading of transparent and opaque scripts during reading development. The results further demonstrate the importance of taking into account differences between orthographies when constructing neural models of reading acquisition.

Differences in implicit motor learning between adults who do and do not stutter.

Implicit learning allows us to acquire complex motor skills through repeated exposure to sensory cues and repetition of motor behaviours, without awareness or effort. Implicit learning is also critical to the incremental fine-tuning of the perceptual-motor system. To understand how implicit learning and associated domain-general learning processes may contribute to motor learning differences in people who stutter, we investigated implicit finger-sequencing skills in adults who do (AWS) and do not stutter (ANS) on an Alternating Serial Reaction Time task. Our results demonstrated that, while all participants showed evidence of significant sequence-specific learning in their speed of performance, male AWS were slower and made fewer sequence-specific learning gains than their ANS counterparts. Although there were no learning gains evident in accuracy of performance, AWS performed the implicit learning task more accurately than ANS, overall. These findings may have implications for sex-based differences in the experience of developmental stuttering, for the successful acquisition of complex motor skills during development by individuals who stutter, and for the updating and automatization of speech motor plans during the therapeutic process.

Neural Processing of Morphology During Reading in Children.

The importance of morphological segmentation for reading has been shown in numerous behavioral studies in children and adults. However, little is known about developmental changes in the neural basis of morphological processing. In addition to effects of age and reading skill, morphological processing during reading may be affected by the morphological structure of the language and the transparency of its orthography. Hebrew provides a unique opportunity to study these factors, with its rich morphological structure, and two versions of script that differ in orthographic transparency. Two groups of children (2nd-3rd and 5th-6th graders) were scanned using fMRI while reading aloud Hebrew nouns. Half of the words were composed of roots and templates (bi-morphemic) and half were mono-morphemic. The words were presented at two levels of transparency: with or without diacritics. ROI analyses showed greater activation for mono over bi-morphemic words across groups in the anterior portions of bilateral middle and superior temporal gyri, especially for the transparent script. These results diverge from a previous finding in adults, showing left frontal activation in the non-transparent script with the same stimuli. These results support the early sensitivity of young Hebrew readers to the rich morphological structure of their language but suggest a developmental change in the role of morphological processes during reading. While in adults morpho-phonological segmentation during reading may compensate for orthographic opacity, morphological processes in children may rely more on semantic aspects, and are enhanced by orthographic transparency.

Effects of Sleep on Language and Motor Consolidation: Evidence of Domain General and Specific Mechanisms.

The current study explores the effects of time and sleep on the consolidation of a novel language learning task containing both item-specific knowledge and the extraction of grammatical regularities. We also compare consolidation effects in language and motor sequence learning tasks, to ask whether consolidation mechanisms are domain general. Young adults learned to apply plural inflections to novel words based on morphophonological rules embedded in the input, and learned to type a motor sequence using a keyboard. Participants were randomly assigned into one of two groups, practicing each task during either the morning or evening hours. Both groups were retested 12 and 24 hours post-training. Performance on frequent trained items in the language task stabilized only following sleep, consistent with a hippocampal mechanism for item-specific learning. However, regularity extraction, indicated by generalization to untrained items in the linguistic task, as well as performance on motor sequence learning, improved 24 hours post-training, irrespective of the timing of sleep. This consolidation process is consistent with a frontostriatal skill-learning mechanism, common across the language and motor domains. This conclusion is further reinforced by cross-domain correlations at the individual level between improvement across 24 hours in the motor task and in the low-frequency trained items in the linguistic task, which involve regularity extraction. Taken together, our results at the group and individual levels suggest that some aspects of consolidation are shared across the motor and language domains, and more specifically, between motor sequence learning and grammar learning.

Simultaneous Normalization and Compensatory Changes in Right Hemisphere Connectivity during Aphasia Therapy.

Changes in brain connectivity during language therapy were examined among participants with aphasia (PWA), aiming to shed light on neural reorganization in the language network. Four PWA with anomia following left hemisphere stroke and eight healthy controls (HC) participated in the study. Two fMRI scans were administered to all participants with a 3.5-month interval. The fMRI scans included phonological and semantic tasks, each consisting of linguistic and perceptual matching conditions. Between the two fMRI scans, PWA underwent Phonological Components Analysis treatment. Changes in effective connectivity during the treatment were examined within right hemisphere (RH) architecture. The results illustrate that following the treatment, the averaged connectivity of PWA across all perceptual and linguistic conditions in both tasks increased resemblance to HC, reflecting the normalization of neural processes associated with silent object name retrieval. In contrast, connections that were specifically enhanced by the phonological condition in PWA decreased in their resemblance to HC, reflecting emerging compensatory reorganization in RH connectivity to support phonological processing. These findings suggest that both normalization and compensation play a role in neural language reorganization at the chronic stage, occurring simultaneously in the same brain.

Mechanisms underlying anomia treatment outcomes.

Treatments for anomia have demonstrated short- and long-term efficacy. However, individual outcomes can be variable, and evidence for treatment generalization is limited. We investigated whether treatment-related measures of access to- and learning of language, namely, a) responsiveness to cues, and b) during-treatment improvements in naming, are good predictors of treatment outcomes. In addition, we investigated mechanisms underlying treatment generalization. Ten adults with chronic, post-stroke aphasia received a phonological treatment for anomia three times a week for five weeks. Naming accuracy of treated and untreated words was assessed pre- and post-treatment and at four- and eight-week follow-ups. Generalization to an untrained naming task, which involved analyses of naming accuracy and speech errors, was also assessed; speech errors were analyzed according to the Interactive Activation (IA) model of word retrieval. Group analyses indicate significant improvements in naming treated compared to untreated words, at all timepoints after therapy. Additional analyses showed significant long-term improvements in naming untreated words. Initial responsiveness to cueing and early improvement emerged as significant predictors of overall pre- to post-treatment improvements in naming treated words; naming improvements made early-on in treatment were also predictive of improvements in naming of the untreated words at follow-up. Furthermore, our study is the first to demonstrate that generalization after a phonological treatment for anomia may be driven by a strengthening of lexical-phonological connections. This study provides novel insights regarding mechanisms driving anomia treatment outcomes. Understanding such mechanisms is critical to improving existing assessment practices, optimizing treatment selection and building treatment protocols that are more likely to generalize.

Morphological decomposition compensates for imperfections in phonological decoding. Neural evidence from typical and dyslexic readers of an opaque orthography.

The current study examined the widely held, but un-tested, assumption that morphological decomposition can compensate for missing phonological information in reading opaque orthographies. In addition, we tested whether morphological decomposition can compensate for the phonological decoding deficits in readers with dyslexia. Hebrew provides a unique opportunity to test these questions as it has a rich Semitic morphology, and two versions of script: a transparent orthography (with diacritic marks, 'pointed') and an opaque orthography (without diacritic marks, 'un-pointed'). In two experiments, one behavioral and one fMRI, skilled and dyslexic readers read aloud Hebrew nouns: half bi-morphemic (root + pattern) and half mono-morphemic (non-decomposable). Each word was presented both in the transparent orthography (pointed), and in the opaque orthography (un-pointed). While skilled readers were faster, and showed no effects of diacritics or morphology, dyslexic readers read pointed words more slowly than un-pointed words and bi-morphemic words faster than mono-morphemic words. The imaging results showed: 1) In both groups a morphological effect was found in un-pointed words, in left inferior and middle frontal gyri, associated with morpho-phonological decomposition. 2) Only readers with dyslexia showed a morphological effect in pointed words in the left occipito-temporal cortex, associated with orthographic processing. 3) Dyslexic readers also showed a positive association between morphological awareness and activation in the left occipito-temporal cortex during reading of all words, and activation in inferior frontal cortex during reading of un-pointed bi-morphemic words. Altogether, these findings suggest that in both typical and dyslexic readers morphological decomposition can compensate for the missing phonological information in an opaque orthography. The results also show that readers with dyslexia can rely on morphological decomposition to compensate for their deficits in phonological decoding. Finally, these results highlight the way in which unique language specific properties shape the neural mechanisms underlying typical and atypical reading.

A neuroimaging dataset on orthographic, phonological and semantic word processing in school-aged children.

Here we describe the public neuroimaging and behavioral dataset entitled "Cross-Sectional Multidomain Lexical Processing" available on the OpenNeuro project (https://openneuro.org). This dataset explores the neural mechanisms and development of lexical processing through task based functional magnetic resonance imaging (fMRI) of rhyming, spelling, and semantic judgement tasks in both the auditory and visual modalities. Each task employed varying degrees of trial difficulty, including conflicting versus non-conflicting orthography-phonology pairs (e.g. harm - warm, wall - tall) in the rhyming and spelling tasks as well as high versus low word pair association in the semantic tasks (e.g. dog - cat, dish - plate). In addition, this dataset contains scores from a battery of standardized psychoeducational assessments allowing for future analyses of brain-behavior relations. Data were collected from a cross-sectional sample of 91 typically developing children aged 8.7- to 15.5- years old. The cross-sectional design employed in this dataset as well as the inclusion of multiple measures of lexical processing in varying difficulties and modalities allows for multiple avenues of future research on reading development.

The role of executive control in post-stroke aphasia treatment.

Executive control (EC) ability is increasingly emerging as an important predictor of post-stroke aphasia recovery. This study examined whether EC predicted immediate treatment gains, treatment maintenance and generalization after naming therapy in ten adults with mild to severe chronic post-stroke aphasia. Performance on multiple EC tasks allowed for the creation of composite scores for common EC, and the EC processes of shifting, inhibition and working memory (WM) updating. Participants were treated three times a week for five weeks with a phonological naming therapy; difference scores in naming accuracy of treated and untreated words (assessed pre, post, four- and eight-weeks after therapy) served as the primary outcome measures. Results from simple and multiple linear regressions indicate that individuals with better shifting and WM updating abilities demonstrated better maintenance of treated words at four-week follow-up, and those with better common EC demonstrated better maintenance of treated words at both four- and eight-week follow-ups. Better shifting ability also predicted better generalization to untreated words post-therapy. Measures of EC were not indicative of improvements on treated words immediately post-treatment, nor of generalization to untreated words at follow-up. Findings suggest that immediate treatment gains, maintenance and generalization may be supported by different underlying mechanisms.

Prior Knowledge Predicts Early Consolidation in Second Language Learning.

Language learning occurs in distinct phases. Whereas some improvement is evident during training, offline memory consolidation processes that take place after the end of training play an important role in learning of linguistic information. The timing of offline consolidation is thought to depend on the type of task, with generalization of implicit knowledge suggested to take more time and sleep to consolidate. The current study aims to investigate individual differences in the timing of consolidation following learning of morphological inflections in a novel language in typical adults. Participants learned to make plural inflections in an artificial language, where inflection was based on morpho-phonological regularities. Participants were trained in the evening, and consolidation was measured after two intervals: 12 h (one night) and 36 h (two nights) post training. We measured both inflection of trained items, which may rely on item-specific learning, and generalization to new untrained items, which requires extraction of morpho-phonological regularities. The results for both trained and un-trained items showed two patterns of consolidation: early versus late, that is while some participants improved during the first night, others, who deteriorated in performance during the first night, improved in the later consolidation interval. Importantly, phonological awareness in L1 predicted early consolidation for trained items. Furthermore, there was no association between participants' consolidation trajectory in trained and untrained items. Our results suggest that consolidation timing depends on the interaction between task characteristics and individual abilities. Moreover, the results show that prior meta-linguistic knowledge predicts the quality of early consolidation processes. These results are consistent with studies in rodents and humans, showing that prior knowledge accelerates consolidation of newly learnt episodic memory. Finally, the rate of consolidation across exposures to the language might explain some of the variability found in the attained level of second language proficiency.

Interhemispheric interactions during sentence comprehension in patients with aphasia.

Right-hemisphere involvement in language processing following left-hemisphere damage may reflect either compensatory processes, or a release from homotopic transcallosal inhibition, resulting in excessive right-to-left suppression that is maladaptive for language performance. Using fMRI, we assessed inter-hemispheric effective connectivity in fifteen patients with post-stroke aphasia, along with age-matched and younger controls during a sentence comprehension task. Dynamic Causal Modeling was used with four bilateral regions including inferior frontal gyri (IFG) and primary auditory cortices (A1). Despite the presence of lesions, satisfactory model fit was obtained in 9/15 patients. In young controls, the only significant homotopic connection (RA1-LA1), was excitatory, while inhibitory connections emanated from LIFG to both left and right A1's. Interestingly, these connections were also correlated with language comprehension scores in patients. The results for homotopic connections show that excitatory connectivity from RA1-to-LA1 and inhibitory connectivity from LA1-to-RA1 are associated with general auditory verbal comprehension. Moreover, negative correlations were found between sentence comprehension and top-down coupling for both heterotopic (LIFG-to-RA1) and intra-hemispheric (LIFG-to-LA1) connections. These results do not show an emergence of a new compensatory right to left excitation in patients nor do they support the existence of left to right transcallosal suppression in controls. Nevertheless, the correlations with performance in patients are consistent with some aspects of both the compensation model, and the transcallosal suppression account for the role of the RH. Altogether our results suggest that changes to both excitatory and inhibitory homotopic and heterotopic connections due to LH damage may be maladaptive, as they disrupt the normal inter-hemispheric coordination and communication.

Changes in Resting-State Connectivity following Melody-Based Therapy in a Patient with Aphasia.

Melody-based treatments for patients with aphasia rely on the notion of preserved musical abilities in the RH, following left hemisphere damage. However, despite evidence for their effectiveness, the role of the RH is still an open question. We measured changes in resting-state functional connectivity following melody-based intervention, to identify lateralization of treatment-related changes. A patient with aphasia due to left frontal and temporal hemorrhages following traumatic brain injuries (TBI) more than three years earlier received 48 sessions of melody-based intervention. Behavioral measures improved and were maintained at the 8-week posttreatment follow-up. Resting-state fMRI data collected before and after treatment showed an increase in connectivity between motor speech control areas (bilateral supplementary motor areas and insulae) and RH language areas (inferior frontal gyrus pars triangularis and pars opercularis). This change, which was specific for the RH, was greater than changes in a baseline interval measured before treatment. No changes in RH connectivity were found in a matched control TBI patient scanned at the same intervals. These results are compatible with a compensatory role for RH language areas following melody-based intervention. They further suggest that this therapy intervenes at the level of the interface between language areas and speech motor control areas necessary for language production.

Therapy-Induced Neuroplasticity in Chronic Aphasia After Phonological Component Analysis: A Matter of Intensity.

Despite the growing evidence regarding the importance of intensity and dose in aphasia therapy, few well-controlled studies contrasting the effects of intensive and non-intensive treatment have been conducted to date. Phonological components analysis (PCA) treatment for anomia has been associated with improvements in some patients with chronic aphasia; however, the effect of treatment intensity has not yet been studied with PCA. Thus, the aim of the present study was to identify the effect of intensity on neural processing associated with word retrieval abilities after PCA treatment. We used functional magnetic resonance imaging to examine therapy-induced changes in activation during an overt naming task in two patients who suffered from a stroke in the left middle cerebral artery territory. P1 received intensive PCA treatment whereas P2 received the standard, non-intensive, PCA treatment. Behavioral results indicate that both standard and intensive conditions yielded improved naming performance with treated nouns, but the changes were only significant for the patient who received the intensive treatment. The improvements were found to be long lasting as both patients maintained improved naming at 2-months follow-ups. The associated neuroimaging data indicate that the two treatment conditions were associated with different neural activation changes. The patient who received the standard PCA showed significant increase in activation with treatment in the right anterior cingulate, as well as extensive areas in bilateral posterior and lateral cortices. By contrast, the patient who received intensive PCA showed more decreases in activation following the treatment. Unexpectedly, this patient showed subcortical increase in activation, specifically in the right caudate nucleus. We speculate that the recruitment of the caudate nucleus and the anterior cingulate in these patients reflects the need to suppress errors to improve naming. Thus, both short-term intensive and standard, non-intensive, PCA treatment can improve word retrieval in chronic aphasia, but neuroimaging data suggest that improved naming is associated with different neural activation patterns in the two treatment conditions.

The Effect of Stimulus Variability on Learning and Generalization of Reading in a Novel Script.

Purpose: The benefit of stimulus variability for generalization of acquired skills and knowledge has been shown in motor, perceptual, and language learning but has rarely been studied in reading. We studied the effect of variable training in a novel language on reading trained and untrained words. Method: Sixty typical adults received 2 sessions of training in reading an artificial script. Participants were assigned to 1 of 3 groups: a variable training group practicing a large set of 24 words, and 2 nonvariable training groups practicing a smaller set of 12 words, with twice the number of repetitions per word. Results: Variable training resulted in higher accuracy for both trained and untrained items composed of the same graphemes, compared to the nonvariable training. Moreover, performance on untrained items was correlated with phonemic awareness only for the nonvariable training groups. Conclusions: High stimulus variability increases the reliance on small unit decoding in adults reading in a novel script, which is beneficial for both familiar and novel words. These results show that the statistical properties of the input during reading acquisition influence the type of acquired knowledge and have theoretical and practical implications for planning efficient reading instruction methods. Supplemental Material: https://doi.org/10.23641/asha.5302195.

Phonological ambiguity modulates resolution of semantic ambiguity during reading: An fMRI study of Hebrew.

OBJECTIVES: The current fMRI study examined the role of phonology in the extraction of meaning from print in each hemisphere by comparing homophonic and heterophonic homographs (ambiguous words in which both meanings have the same or different sounds respectively, e.g., bank or tear). The analysis distinguished between the first phase, in which participants read ambiguous words without context, and the second phase in which the context resolves the ambiguity. METHOD: Native Hebrew readers were scanned during semantic relatedness judgments on pairs of words in which the first word was either a homophone or a heterophone and the second word was related to its dominant or subordinate meaning. RESULTS: In Phase 1 there was greater activation for heterophones in left inferior frontal gyrus (IFG), pars opercularis, and more activation for homophones in bilateral IFG pars orbitalis, suggesting that resolution of the conflict at the phonological level has abolished the semantic ambiguity for heterophones. Reduced activation for all ambiguous words in temporo-parietal regions suggests that although ambiguity enhances controlled lexical selection processes in frontal regions it reduces reliance on bottom-up mapping processes. After presentation of the context, a larger difference between the dominant and subordinate meaning was found for heterophones in all reading-related regions, suggesting a greater engagement for heterophones with the dominant meaning. CONCLUSIONS: Altogether these results are consistent with the prominent role of phonological processing in visual word recognition. Finally, despite differences in hemispheric asymmetry between homophones and heterophones, ambiguity resolution, even toward the subordinate meaning, is largely left lateralized. (PsycINFO Database Record