Top‐down and bottom‐up influences on the left ventral occipito‐temporal cortex during visual word recognition: An analysis of effective connectivity

The functional role of the left ventral occipito‐temporal cortex (vOT) in visual word processing has been studied extensively. A prominent observation is higher activation for unfamiliar but pronounceable letter strings compared to regular words in this region. Some functional accounts have interpreted this finding as driven by top‐down influences (e.g., Dehaene and Cohen [ 2011 ]: Trends Cogn Sci 15:254–262; Price and Devlin [ 2011 ]: Trends Cogn Sci 15:246–253), while others have suggested a difference in bottom‐up processing (e.g., Glezer et al. [ 2009 ]: Neuron 62:199–204; Kronbichler et al. [ 2007 ]: J Cogn Neurosci 19:1584–1594). We used dynamic causal modeling for fMRI data to test bottom‐up and top‐down influences on the left vOT during visual processing of regular words and unfamiliar letter strings. Regular words (e.g., taxi) and unfamiliar letter strings of pseudohomophones (e.g., taksi) were presented in the context of a phonological lexical decision task (i.e., “Does the item sound like a word?”). We found no differences in top‐down signaling, but a strong increase in bottom‐up signaling from the occipital cortex to the left vOT for pseudohomophones compared to words. This finding can be linked to functional accounts which assume that the left vOT contains neurons tuned to complex orthographic features such as morphemes or words [e.g., Dehaene and Cohen [ 2011 ]: Trends Cogn Sci 15:254‐262; Kronbichler et al. [ 2007 ]: J Cogn Neurosci 19:1584–1594]: For words, bottom‐up signals converge onto a matching orthographic representation in the left vOT. For pseudohomophones, the propagated signals do not converge, but (partially) activate multiple orthographic word representations, reflected in increased effective connectivity. Hum Brain Mapp 35:1668–1680, 2014. © 2013 Wiley Periodicals, Inc.     

Superior Written Over Spoken Picture Naming in a Case of Frontotemporal Dementia

Two main hypotheses have been proposed regarding the role of phonology in written word production. According to the phonological mediation hypothesis, the retrieval of the lexical phonological representation of a word is an obligatory prerequisite to the retrieval of its spelling. Therefore, deficits to the phonological lexicon should affect both spoken and written picture naming. In contrast, the orthographic autonomy hypothesis posits that the lexical orthographic representations of words can be accessed without any necessary phonological mediation. In support of this view, cases of preserved written naming despite impaired lexical phonology have been reported following brain damage. In this report, we replicate this basic pattern of performance in case YP, a 60-year-old woman with a pattern of frontotemporal dementia. As her disease progressed, YP’s ability to write down the names of pictures remained very good despite a severe decline in oral naming. Further testing indicated that this deficit was not primarily due to an articulatory or post-lexical phonological deficit. YP’s case provides strong additional support for the orthographic autonomy hypothesis. The significance of this case with respect to the characterization of dementia syndromes is discussed.     

Superior written over spoken picture naming in a case of frontotemporal dementia

Two main hypotheses have been proposed regarding the role of phonology in written word production. According to the phonological mediation hypothesis, the retrieval of the lexical phonological representation of a word is an obligatory prerequisite to the retrieval of its spelling. Therefore, deficits to the phonological lexicon should affect both spoken and written picture naming. In contrast, the orthographic autonomy hypothesis posits that the lexical orthographic representations of words can be accessed without any necessary phonological mediation. In support of this view, cases of preserved written naming despite impaired lexical phonology have been reported following brain damage. In this report, we replicate this basic pattern of performance in case YP, a 60-year-old woman with a pattern of frontotemporal dementia. As her disease progressed, YP's ability to write down the names of pictures remained very good despite a severe decline in oral naming. Further testing indicated that this deficit was not primarily due to an articulatory or post-lexical phonological deficit. YP's case provides strong additional support for the orthographic autonomy hypothesis. The significance of this case with respect to the characterization of dementia syndromes is discussed.     

Reading in a regular orthography: an FMRI study investigating the role of visual familiarity

In order to separate the cognitive processes associated with phonological encoding and the use of a visual word form lexicon in reading, it is desirable to compare the processing of words presented in a visually familiar form with words in a visually unfamiliar form. Japanese Kana orthography offers this possibility. Two phonologically equivalent but visually dissimilar syllabaries allow the writing of, for example, foreign loanwords in two ways, only one of which is visually familiar. Familiarly written words, unfamiliarly written words, and pseudowords were presented in both Kana syllabaries (yielding six conditions in total) to participants during an fMRI measurement with a silent articulation task (Experiment 1) and a phonological lexical decision task (Experiment 2) using an event-related design. Consistent over two experimental tasks, the three different stimulus types (familiar, unfamiliar, and pseudoword) were found to activate selectively different brain regions previously associated with phonological encoding and word retrieval or meaning. Compatible with the predictions of the dual-route model for reading, pseudowords and visually unfamiliar words, which have to be read using phonological assembly, caused an increase in brain activity in left inferior frontal regions (BA 44/47), as compared to visually familiar words. Visually familiar and unfamiliar words were found to activate a range of areas associated with lexico-semantic processing more strongly than pseudowords, such as the left and right temporo-parietal region (BA 39/40), a region in the left middle/inferior temporal gyrus (BA 20/21), and the posterior cingulate (BA 31).     

Diagnostic profiles of children with developmental dyslexia in a transparent orthography

In the present study for 108 typical and 122 atypical Dutch readers in second grade, the accuracy and speed of decoding words and pseudowords, as well as the accuracy of spelling words were assessed along with four types of phonological precursor measures: rapid naming, verbal working memory, phoneme awareness and letter knowledge. The data show that the group being diagnosed as poor readers were significantly behind in all reading and spelling measures. It was also found that the criterion measures of reading and spelling explained already two third of the variance associated with the group distinction. Finally, we found word and pseudoword efficiency in the typical group to be explained by phonological awareness (spoonerism) and rapid naming of letters, word and pseudoword accuracy by phonological awareness, and spelling by phonological awareness and letter dictation. In the group of poor readers, a much greater variety of precursor measures was involved in explaining the variance in reading and spelling abilities.     

Analysis of reading strategies in deaf adults as a function of their language and meta-phonological skills

The first aim of this study was to examine the mechanisms used in reading sentences by deaf adults who had completed secondary or higher education. Previous data allowed us to hypothesize that they used the key word strategy, consisting of identifying (some of) the frequent content words, and deriving an overall representation of the sentence's meaning ignoring the function words. The results supported the hypothesis. The second aim was to establish the relationships between this strategy and the linguistic and phonological abilities of deaf participants. The results show that vocabulary increased with reading level, but syntax, evaluated with the use of function words, did not. This suggests that using the key word strategy during long periods of time increases knowledge of content words but not syntax, probably because function words are neglected by this strategy. The results also showed that the deaf participants had a fairly large orthographical lexicon. This implies that the extensive use of the key word strategy allows them to store lexical information. The next question was whether the written word representations of the deaf participants were memorized as mere logograms, or if they had been stored in connection with the phonological representations of the corresponding words. The metaphonological tasks conducted produced evidence indicating that deaf participants used both orthographic and phonological representations. A factor analysis of the metaphonological tasks together with reading and spelling confirmed that both factors were necessary to explain the whole variance in the deaf group.     

Brain activation in teenagers with isolated spelling disorder during tasks involving spelling assessment and comparison of pseudowords. fMRI study

Methods: The present study aimed at defining the specific traits of brain activation in teenagers with isolated spelling disorder in comparison with good spellers. fMRI examination was performed where the subject’s task involved taking a decision 1/whether the visually presented words were spelled correctly or not (the orthographic decision task), and 2/whether the two presented letters strings (pseudowords) were identical or not (the visual decision task). Half of the displays showing meaningful words with an orthographic difficulty contained pairs with both words spelled correctly, and half of them contained one misspelled word. Half of the pseudowords were identical, half of them were not. The participants of the study included 15 individuals with isolated spelling disorder and 14 good spellers, aged 13–15. Results: The results demonstrated that the essential differences in brain activation between teenagers with isolated spelling disorder and good spellers were found in the left inferior frontal gyrus, left medial frontal gyrus and right cerebellum posterior lobe, i.e. structures important for language processes, working memory and automaticity of behaviour. Conclusion: Spelling disorder is not only an effect of language dysfunction, it could be a symptom of difficulties in learning and automaticity of motor and visual shapes of written words, rapid information processing as well as automating use of orthographic lexicon.     

The interaction between orthographic and phonological information in children: an fMRI study

We examined the neural representations of orthographic and phonological processing in children, while manipulating the consistency between orthographic and phonological information. Participants, aged 9-15, were scanned while performing rhyming and spelling judgments on pairs of visually presented words. The orthographic and phonological similarity between words in the pair was independently manipulated, resulting in four conditions. In the nonconflicting conditions, both orthography and phonology of the words were either (1) similar (lime-dime) or (2) different (staff-gain); in conflicting conditions, words had (3) similar phonology and different orthography (jazz-has) or (4) different phonology and similar orthography (pint-mint). The comparison between tasks resulted in greater activation for the rhyming task in bilateral inferior frontal gyri (BA 45/47), and greater activation for the spelling task in bilateral inferior/superior parietal lobules (BA 40/7), suggesting greater involvement of phonological and semantic processing in the rhyming task, and nonlinguistic spatial processing in the spelling task. Conflicting conditions were more difficult in both tasks and resulted in greater activation in the above regions. The results suggest that when children encounter inconsistency between orthographic and phonological information they show greater engagement of both orthographic and phonological processing.     

Taxi vs. taksi: on orthographic word recognition in the left ventral occipitotemporal cortex

The importance of the left occipitotemporal cortex for visual word processing is highlighted by numerous functional neuroimaging studies, but the precise function of the visual word form area (VWFA) in this brain region is still under debate. The present functional magnetic resonance imaging study varied orthographic familiarity independent from phonological-semantic familiarity by presenting orthographically familiar and orthographically unfamiliar forms (pseudohomophones) of the same words in a phonological lexical decision task. Consistent with orthographic word recognition in the VWFA, we found lower activation for familiar compared with unfamiliar forms, but no difference between pseudohomophones and pseudowords. This orthographic familiarity effect in the VWFA differed from the phonological familiarity effect in left frontal regions, where phonologically unfamiliar pseudowords led to higher activation than phonologically familiar pseudohomophones. We suggest that the VWFA not only computes letter string representations but also hosts word-specific orthographic representations. These representations function as recognition units with the effect that letter strings that readily match with stored representations lead to less activation that letter strings that do not.     

An investigation of twenty/20 vision in reading

One functional anatomical model of reading, drawing on human neuropsychological and neuroimaging data, proposes that a region in left ventral occipitotemporal cortex (vOT) becomes, through experience, specialized for written word perception. We tested this hypothesis by presenting numbers in orthographical and digital form with two task demands, phonological and numerical. We observed a main effect of task on left vOT activity but not stimulus type, with increased activity during the phonological task that was also associated with increased activity in the left inferior frontal gyrus, a region implicated in speech production. Region-of-interest analysis confirmed that there was equal activity for orthographical and digital written forms in the left vOT during the phonological task, despite greater visual complexity of the orthographical forms. This evidence is incompatible with a predominantly feedforward model of written word recognition that proposes that the left vOT is a specialized cortical module for word recognition in literate subjects. Rather, the physiological data presented here fits better with interactive computational models of reading that propose that written word recognition emerges from bidirectional interactions between three processes: visual, phonological, and semantic. Further, the present study is in accord with others that indicate that the left vOT is a route through which nonlinguistic stimuli, perhaps high contrast two-dimensional objects in particular, gain access to a predominantly left-lateralized language and semantic system.     

Letter fluency in 7-8-year-old children is related to the anterior,but not posterior,ventral occipito-temporal cortex during an auditory phonological task

Previous studies have shown that reading skill in 3- to 6-year-old children is related to the automatic activation of the posterior left ventral occipitotemporal cortex (vOT) during spoken language processing, whereas 8- to 15-year-old children and adult readers activate the anterior vOT. However, it is unknown how children who are between these two age groups automatically activate orthographic representations in vOT for spoken language. In the current study, we recruited 153 7- to 8-year-old children to fill the age gap from previous studies. Using functional magnetic resonance imaging (fMRI), we measured children’s reading-related skills and brain activity during an auditory phonological task with both a small (i.e. onset) and a large (i.e. rhyme) grain size condition. We found that letter fluency, but not reading accuracy, was correlated with activation in the anterior vOT for the rhyme condition. There were no reading-related skill correlations for the posterior vOT or for activation during the onset condition in this age group. Our findings reveal that automatic activation in the anterior vOT during spoken language processing already occurs in higher skilled 7- to 8-year-old children. In addition, increases in naming automaticity is the primary determinant of the engagement of vOT during phonological awareness tasks.     

Late acquisition of literacy in a native language

With event-related functional MRI (fMRI) and with behavioral measures we studied the brain processes underlying the acquisition of native language literacy. Adult dialect speakers were scanned while reading words belonging to three different conditions: dialect words, i.e., the native language in which subjects are illiterate (dialect), German words, i.e., the second language in which subjects are literate, and pseudo-words. Investigating literacy acquisition of a dialect may reveal how novel readers of a language build an orthographic lexicon, i.e., establish a link between already available semantic and phonological representations and new orthographic word forms. The main results of the study indicate that a set of regions, including the left anterior hippocampal formation and subcortical nuclei, is involved in the buildup of orthographic representations. The repeated exposure to written dialect words resulted in a convergence of the neural substrate to that of the language in which these subjects were already proficient readers. The latter result is compatible with a "fast" brain plasticity process that may be related to a shift of reading strategies.     

Assembling and encoding word representations: fMRI subsequent memory effects implicate a role for phonological control

Novel word learning is central to the flexibility inherent in the human language capacity. Word learning may partially depend on long-term memory formation during the assembly of phonological representations from orthographic inputs. In the present study, event-related functional magnetic resonance imaging (fMRI) examined the contributions of phonological control-a component of the verbal working memory system-to phonological assembly and word learning. Subjects were scanned while making syllable decisions about visually presented familiar (English) and novel (pseudo-English and Foreign) words, a task that required retrieval and analysis of existing phonological codes or the assembly and analysis of novel representations. Results revealed that left inferior prefrontal cortex (LIPC) and bilateral parietal cortices were differentially engaged during the processing of novel words, suggesting that this circuit is recruited during phonological assembly. A subsequent memory analysis that examined the relation between fMRI signal and the subject's ability to later remember the words (a measure of effective memory formation) revealed that the magnitude of activation in LIPC, bilateral superior parietal, and left inferior parietal cortices was positively correlated with later memory. Moreover, although the magnitude of the subsequent memory effect in parietal cortex was not significantly affected by word type, this effect was greater in posterior LIPC for novel (pseudo-English) than for familiar (English) words. In the course of subserving the assembly of novel word representations, the phonological (articulatory) control component of the phonological system appears to play a central role in the encoding of novel words into long-term memory.     

From orthography to phonetics: ERP measures of grapheme-to-phoneme conversion mechanisms in reading

Neuroimaging has provided evidence that the first stages of visual word recognition activate a visual word-form center localized in the left extrastriate cortex (fusiform gyrus). Accordingly, neurological cases of patients suffering from pure alexia reported the left posterior occipital lobe as the possible locus of orthographic analysis. There is less agreement in the literature about which brain structures are involved in the subsequent stages of word processing and, in particular, their time course of activation. Functional magnetic resonance imaging and magnetic source imaging studies recently reported data that could indicate a dual route model of reading. These findings are particularly relevant to studies on the functional deficits associated with phonological and surface dyslexia. There is evidence for the existence of two different brain mechanisms supporting phonological processing in visual word recognition: one mechanism subserving "assembled phonology" for reading letter strings and another one subserving "addressed phonology" for reading meaningful words. However, available knowledge on the time course and neural locus of grapheme-to-phoneme conversion mechanisms in reading is still inadequate. In this study, we compared processing of meaningful and meaningless Italian words in a task requiring a phonemic/phonetic decision task. Stimuli were 1152 different orthographic stimuli presented in the central visual field. Half the stimuli were Italian words (with a high or low frequency of occurrence), the other half were meaningless strings of letters (legal pseudowords and letter strings). Event-related potentials were recorded from 28 scalp sites in 10 Italian university students. The task consisted of deciding about the presence/absence of a given "phone" in the hypothetical enunciation of word read: for example, "Is there a /k/ in cheese?". Results showed that lexical frequency and orthographical regularity affected linguistic processing within 150 msec poststimulus. Indeed, the amplitude of a centroparietal P150 varied as a function of stimulus type, being larger in response to high-frequency words than to low-frequency ones and to words and pseudowords than to letter strings. This component might index visual categorization processes and recognition of familiar objects, being highly sensitive to orthographic regularity and "ill-formedness" of words. The amplitude of the P150 was the same in response to well-formed meaningless and to meaningful words, when these latter had a low lexical frequency. This might indicate that highly familiar words are recognized as meaningful unitary visual objects at very early stages of processing, through a visual route to an orthographic input lexicon. Moreover, the amplitude of the negativity recorded between 250 and 350 msec showed an anteroposterior topographic dissociation for access to the phonemic representation of well- or ill-formed strings of characters. Brain responses were larger over the left occipito-temporal regions during reading of words and pseudowords and over the left frontal regions during reading of letter strings.     

Neurofunctionally dissecting the reading system in children

The reading system can be broken down into four basic subcomponents in charge of prelexical, orthographic, phonological, and lexico-semantic processes. These processes need to jointly work together to become a fluent and efficient reader. Using functional magnetic resonance imaging (fMRI), we systematically analyzed differences in neural activation patterns of these four basic subcomponents in children (N = 41, 9–13 years) using tasks specifically tapping each component (letter identification, orthographic decision, phonological decision, and semantic categorization). Regions of interest (ROI) were selected based on a meta-analysis of child reading and included the left ventral occipito-temporal cortex (vOT), left posterior parietal cortex (PPC), left inferior frontal gyrus (IFG), and bilateral supplementary motor area (SMA). Compared to a visual baseline task, enhanced activation in vOT and IFG was observed for all tasks with very little differences between tasks. Activity in the dorsal PPC system was confined to prelexical and phonological processing. Activity in the SMA was found in orthographic, phonological, and lexico-semantic tasks. Our results are consistent with the idea of an early engagement of the vOT accompanied by executive control functions in the frontal system, including the bilateral SMA.     

Right hemisphere reading mechanisms in a global alexic patient

We investigated the implicit, or covert, reading ability of a global alexic patient (EA) to help determine the contribution of the right hemisphere to reading. Previous studies of alexic patients with left hemisphere damage have suggested that the ability to derive meaning from printed words that cannot be read out loud may reflect right hemisphere reading mechanisms. Other investigators have argued that residual left hemisphere abilities are sufficient to account for implicit reading and moreover do not require the postulation of a right hemisphere system that has no role in normal reading processes. However, few studies have assessed covert reading in patients with lesions as extensive as the one in EA, which affected left medial, inferior temporal-occipital cortex, hippocampus, splenium, and dorsal white matter. EA was presented with lexical decision, semantic categorization, phonemic categorization, and letter matching tasks. Although EA was unable to access phonology and could not overtly name words or letters, she was nevertheless capable of making lexical and semantic decisions at above chance levels, with an advantage for concrete versus abstract words. Her oral and written spelling were relatively intact, suggesting that orthographic knowledge is retained, although inaccessible through the visual modality. Based on her ability to access lexical and semantic information without contacting phonological representations, we propose that EA's implicit reading emerges from, and is supported, by the right hemisphere. Finally, we conclude that her spelling and writing abilities are supported by left hemisphere mechanisms.     

Age of acquisition effects on spelling in surface dysgraphia

Background: It is not known whether the age of acquisition (AoA) of a word has any significant effect on the spelling performance of normal or impaired speakers. One way to explore this question is to examine the effects of AoA on surface dysgraphia. Aims: We report MK, a patient who suffered herpes simplex viral encephalitis (HSVE) resulting in semantic anomia and surface dysgraphia that we characterise as a tendency to produce a legitimate alternative spelling of a component defined as a LASC error. Methods & Procedures: We describe MK's neuropsychological profile, giving details of impaired naming, spoken and written word comprehension, written picture naming, oral reading, and spelling to dictation, but preserved repetition of whole words. We report experiments designed (1) to test the effect of AoA on spelling by controlling for word frequency as well as other correlated variables using logistic regression and ANOVA, and (2) to test the effects of AoA and sound to spelling predictability on MK's spelling. Outcomes and Results: The results show an effect of AoA but no independent effect of frequency on spelling and an interaction between AoA and predictability, i.e., an effect of AoA on unpredictable word spelling but no effect of AoA on predictable word spelling. Conclusions: We discuss these data with reference to accounts of AoA that locate the effect at the level of mappings between input (phonological) and output (orthographic) representations. We argue that the effect of AoA on spelling is not the result of lexeme activation per se but instead reflects the largely unpredictable mappings between sound and spelling that characterise the majority of English word spellings.     

Language lateralization in phonological, semantic and orthographic tasks: a slow evoked potential study

Most of literature on language has shown how different word-classes activate distinct neural networks within linguistic cortical areas. The present investigation aimed to demonstrate that, by means of slow evoked potentials and using the same set of words in different tasks, it is possible to activate cortical networks that are spatially and temporally distinguished. Twenty healthy subjects had to evaluate, in a word pair matching session, whether two words rhymed (phonological task), were semantically related (semantic task) or were written in the same letter case (orthographic task). Slow wave amplitude was computed in three relevant time windows: the last 0.5 s of first word presentation (W1), the initial contingent negative variation (iCNV) and the terminal CNV (tCNV). During W1 and iCNV intervals, both the orthographic and the phonological tasks were left lateralized. Furthermore, the phonological task was more lateralized than the orthographic because of a greater inhibition of the right hemisphere, whereas the orthographic task was characterized by a greater bilateral posterior activation. During the tCNV, only the phonological task remained left lateralized while orthographic and semantic were bilaterally distributed. Although the use of the same set of words tends to activate widely overlapped networks, in the present research task manipulation was effective in demonstrating task dependent differences in brain lateralization. Thus, the present paradigm and the adopted tasks are especially suited for studying deficit and recovery of language in patients affected by linguistic disorders such as developmental dyslexia and aphasia.     

Explaining left lateralization for words in the ventral occipitotemporal cortex

Reading is a uniquely human task and therefore any sign of neuronal activation that is specific to reading is of considerable interest. One intriguing observation is that ventral occipitotemporal (vOT) activation is more strongly left lateralized for written words than other visual stimuli. This has contributed to claims that left vOT plays a special role in reading. Here, we investigated whether left lateralized vOT responses for words were the consequence of visual feature processing, visual word form selectivity, or higher level language processing. Using fMRI in 82 skilled readers, our paradigm compared activation and lateralization for words and nonlinguistic stimuli during different tasks. We found that increased left lateralization for words relative to pictures was the consequence of reduced activation in right vOT rather than increased activation in left vOT. We also found that the determinants of lateralization varied with the subregion of vOT tested. In posterior vOT, lateralization depended on the spatial frequency of the visual inputs. In anterior vOT, lateralization depended on the semantic demands of the task. In middle vOT, lateralization depended on a combination of visual expertise in the right hemisphere and semantics in the left hemisphere. These results have implications for interpreting left lateralized vOT activation during reading. Specifically, left lateralized activation in vOT does not necessarily indicate an increase in left vOT processing but is instead a consequence of decreased right vOT function. Moreover, the determinants of lateralization include both visual and semantic factors depending on the subregion tested.     

Using orthographic neighbours to treat a case of graphemic buffer disorder

Background: The characteristics of graphemic buffer disorder have been described by Miceli, Silveri, and Caramazza (1985) and Caramazza and Miceli (1990) and include a length effect in spelling words and nonwords, in both written and oral format. Error patterns typically consist of omissions, substitutions, additions, and movement errors. Recently, lexical effects on spelling accuracy in many buffer cases have been shown (Sage & Ellis, 2004) and, in patient BH in particular, these included an influence of orthographic neighbourhood size (the number of words that can be generated by changing one letter in the target word; Coltheart, Davelaar, Jonassen, & Besner, 1977). Aims: This paper aims to show that orthographic neighbours used in a therapy programme can bring about improvements in the spelling of targets that have not directly received therapy. Methods & procedures: This was a single case treatment study of BH, who showed classic features of graphemic buffer disorder in her spelling. Two priming studies contrasting no prime, control primes, and orthographic neighbour primes (both word and nonword) established that positive effects on spelling accuracy and error pattern could be achieved. The use of orthographic neighbours was then extended into a therapy programme. Three word sets contrasted direct therapy to the words themselves (Set1), no therapy at all (Set 2), and therapy to neighbours of the words in a set (Set 3). An errorless learning paradigm was used throughout the therapy programme. Outcome & results: Improvement was made both to the treated words in Set 1 and to the words in Set 3, even though these words had received no direct therapy. There was no change in accuracy in Set 2, the control set that had received no therapy. The paper also explores changes in error patterns due to therapy, showing that error patterns changed following therapy. Conclusions: The priming studies showed that changes in accuracy could be achieved when orthographic neighbours were used. Following on from this, a therapy programme based on neighbours was effective in assisting the graphemic buffer. Specifically, the interaction between lexicon and buffer was used therapeutically to improve not only the spelling of words that had received direct treatment but also to a word set that did not directly receive treatment. These changes were brought about using an errorless learning paradigm.