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1.  Tactile localization on digits and hand: Structure and development 
Psychological science  2013;24(9):1653-1663.
Localization of tactile stimuli to the hand and digits is fundamental to somatosensory perception. However, little is known about the development or genetic bases of this ability in humans. We examined tactile localization in normally developing children, adolescents, adults, and people with Williams syndrome (WS), a genetic disorder resulting in a wide range of severe visual-spatial deficits. Normally developing four-year-olds made large stimulus localization errors, sometimes across digits, but nevertheless their errors revealed a structured internal representation of the hand. Errors became exponentially smaller over development, reaching adult level by adolescence. In contrast, people with WS showed large localization errors regardless of age and significant cross-digit errors, a profile similar to normal four-year-olds. Thus, tactile localization reflects internal organization of the hand even early in normal development, undergoes substantial development in normal children, and is susceptible to developmental, but not organizational impairment under genetic deficit.
PMCID: PMC4275844  PMID: 23907541
Developmental Disorders; Parietal Lobe; Spatial Perception; Visuospatial Ability; Williams Syndrome
2.  Neuroimaging in aphasia treatment research: Issues of experimental design for relating cognitive to neural changes 
NeuroImage  2012;73:200-207.
The design of functional neuroimaging studies investigating the neural changes that support treatment-based recovery of targeted language functions in acquired aphasia faces a number of challenges. In this paper, we discuss these challenges and focus on experimental tasks and experimental designs that can be used to address the challenges, facilitate the interpretation of results and promote integration of findings across studies.
PMCID: PMC3600065  PMID: 22974976
Aphasia; Language recovery; fMRI
3.  The interface between morphology and phonology: Exploring a morpho-phonological deficit in spoken production 
Cognition  2013;127(2):270-286.
Morphological and phonological processes are tightly interrelated in spoken production. During processing, morphological processes must combine the phonological content of individual morphemes to produce a phonological representation that is suitable for driving phonological processing. Further, morpheme assembly frequently causes changes in a word's phonological well-formedness that must be addressed by the phonology. We report the case of an aphasic individual (WRG) who exhibits an impairment at the morpho-phonological interface. WRG was tested on his ability to produce phonologically complex sequences (specifically, coda clusters of varying sonority) in heteromorphemic and tautomorphemic environments. WRG made phonological errors that reduced coda sonority complexity in multimorphemic words (e.g., passed→[pæstɪd]) but not in monomorphemic words (e.g., past). WRG also made similar insertion errors to repair stress clash in multimorphemic environments, confirming his sensitivity to cross-morpheme well-formedness. We propose that this pattern of performance is the result of an intact phonological grammar acting over the phonological content of morphemic representations that were weakly joined because of brain damage. WRG may constitute the first case of a morpho-phonological impairment—these results suggest that the processes that combine morphemes constitute a crucial component of morpho-phonological processing.
PMCID: PMC3608811  PMID: 23466641
morpho-phonology; aphasia; grammar; production; sonority; stress; well-formedness; markedness
4.  Identifying functional reorganization of spelling networks: an individual peak probability comparison approach 
Previous research has shown that damage to the neural substrates of orthographic processing can lead to functional reorganization during reading (Tsapkini et al., 2011); in this research we ask if the same is true for spelling. To examine the functional reorganization of spelling networks we present a novel three-stage Individual Peak Probability Comparison (IPPC) analysis approach for comparing the activation patterns obtained during fMRI of spelling in a single brain-damaged individual with dysgraphia to those obtained in a set of non-impaired control participants. The first analysis stage characterizes the convergence in activations across non-impaired control participants by applying a technique typically used for characterizing activations across studies: Activation Likelihood Estimate (ALE) (Turkeltaub et al., 2002). This method was used to identify locations that have a high likelihood of yielding activation peaks in the non-impaired participants. The second stage provides a characterization of the degree to which the brain-damaged individual's activations correspond to the group pattern identified in Stage 1. This involves performing a Mahalanobis distance statistics analysis (Tsapkini et al., 2011) that compares each of a control group's peak activation locations to the nearest peak generated by the brain-damaged individual. The third stage evaluates the extent to which the brain-damaged individual's peaks are atypical relative to the range of individual variation among the control participants. This IPPC analysis allows for a quantifiable, statistically sound method for comparing an individual's activation pattern to the patterns observed in a control group and, thus, provides a valuable tool for identifying functional reorganization in a brain-damaged individual with impaired spelling. Furthermore, this approach can be applied more generally to compare any individual's activation pattern with that of a set of other individuals.
PMCID: PMC3872307  PMID: 24399981
spelling; dysgraphia; ALE; IPPC; fMRI; orthography; mahalanobis
5.  Lexical neighborhood effects in pseudoword spelling 
The general aim of this study is to contribute to a better understanding of the cognitive processes that underpin skilled adult spelling. More specifically, it investigates the influence of lexical neighbors on pseudo-word spelling with the goal of providing a more detailed account of the interaction between lexical and sublexical sources of knowledge in spelling. In prior research examining this topic, adult participants typically heard lists composed of both words and pseudo-words and had to make a lexical decision to each stimulus before writing the pseudo-words. However, these priming paradigms are susceptible to strategic influence and may therefore not give a clear picture of the processes normally engaged in spelling unfamiliar words. In our two Experiments involving 71 French-speaking literate adults, only pseudo-words were presented which participants were simply requested to write to dictation using the first spelling that came to mind. Unbeknownst to participants, pseudo-words varied according to whether they did or did not have a phonological word neighbor. Results revealed that low-probability phoneme/grapheme mappings (e.g., /o/ -> aud in French) were used significantly more often in spelling pseudo-words with a close phonological lexical neighbor with that spelling (e.g., /krepo/ derived from “crapaud,” /krapo/) than in spelling pseudo-words with no close neighbors (e.g., /frøpo/). In addition, the strength of this lexical influence increased with the lexical frequency of the word neighbors as well as with their degree of phonetic overlap with the pseudo-word targets. These results indicate that information from lexical and sublexical processes is integrated in the course of spelling, and a specific theoretical account as to how such integration may occur is introduced.
PMCID: PMC3842689  PMID: 24348436
pseudoword spelling; skilled spelling; literacy; spelling models; neighborhood activation
6.  Letter representations in writing: an fMRI adaptation approach 
Behavioral and neuropsychological research in reading and spelling has provided evidence for the role of the following types of orthographic representations in letter writing: letter shapes, letter case, and abstract letter identities. We report on the results of an fMRI investigation designed to identify the neural substrates of these different representational types. Using an fMRI adaptation paradigm we examined the neural distribution of inhibition and release from inhibition in a letter-writing task in which, on every trial, participants produced three repetitions of the same letter and a fourth letter that was either identical to (no-change trial) or different from the previous three (change trial). Change trials involved a change in the shape, case, and/or identity of the letter. After delineating the general letter writing network by identifying areas that exhibited significant neural adaptation effects on no-change trials, we used deconvolution analysis to examine this network for effects of release from inhibition on change trials. In this way we identified regions specifically associated with the representation of letter shape (in the left SFS and SFG/pre-CG) and letter identity [in the left fusiform gyrus (FG)] or both [right cerebellum, left post-central gyrus (post-CG), and left middle frontal gyrus (MFG)]. No regions were associated with the representation of letter case. This study showcases an investigational approach that allows for the differentiation of the neurotopography of the representational types that are key to our ability to produce written language.
PMCID: PMC3809555  PMID: 24194724
letter; writing; fMRI; fMRI adaptation; neural habituation; letter shape; letter identity; letter case
7.  Temporal stability and representational distinctiveness: Key functions of orthographic working memory 
Cognitive neuropsychology  2012;28(5):338-362.
A primary goal of working memory research has been to understand the mechanisms that permit working memory systems to effectively maintain the identity and order of the elements held in memory for sufficient time as to allow for their selection and transfer to subsequent processing stages. Based on the performance of two individuals with acquired dysgraphia affecting orthographic WM (the graphemic buffer) we present evidence of two distinct and dissociable functions of orthographic WM. One function is responsible for maintaining the temporal stability of letters held in orthographic WM, while the other is responsible for maintaining their representational distinctiveness. The failure to maintain temporal stability and representational distinctiveness give rise, respectively, to decay and interference effects that manifest themselves in distinctive error patterns, including distinct serial position effects. The findings we report have implications beyond our understanding of orthographic WM, as the need to maintain temporal stability and representational distinctiveness in WM is common across cognitive domains.
PMCID: PMC3427759  PMID: 22248210
working memory; spelling; dysgraphia; orthographic representations
8.  Underlying Cause(s) of Letter Perseveration Errors 
Neuropsychologia  2011;50(2):305-318.
Perseverations, the inappropriate intrusion of elements from a previous response into a current response, are commonly observed in individuals with acquired deficits. This study specifically investigates the contribution of failure-to activate and failure-to-inhibit deficit(s) in the generation of letter perseveration errors in acquired dysgraphia. We provide evidence from the performance 12 dysgraphic individuals indicating that a failure to activate graphemes for a target word gives rise to letter perseveration errors. In addition, we also provide evidence that, in some individuals, a failure-to-inhibit deficit may also contribute to the production of perseveration errors.
PMCID: PMC3259193  PMID: 22178232
letter perseveration errors; dysgraphia; spelling; inhibition
9.  Case Series in Cognitive Neuropsychology: Promise, Perils and Proper Perspective 
Cognitive neuropsychology  2011;28(7):435-444.
Schwartz & Dell (2010) advocated for a major role for case series investigations in cognitive neuropsychology. They defined the key features of this approach and presented a number of arguments and examples illustrating the benefits of case series studies and their contribution to computational cognitive neuropsychology. In the Special Issue on “Case Series in Cognitive Neuropsychology” there are six commentaries on Schwartz and Dell (2010) as well as a response to the six commentaries by Dell and Schwartz. In this paper, I provide a brief summary of the key points made in Schwartz and Dell (2010) and I review the promise and perils of case series design as revealed by the six commentaries. I conclude by placing the set of papers within a broader perspective, providing some clarification of the historical record on case series and single case approaches, raising some cautionary notes for case series studies and situating both case series and single case approaches within the larger context of theory development in the cognitive sciences.
PMCID: PMC3427758  PMID: 22746685
10.  When do combinatorial mechanisms apply in the production of inflected words? 
Cognitive neuropsychology  2010;10.1080/02643294.2010.524467.
A central question for theories of inflected word processing is to determine under what circumstances compositional procedures apply. Some accounts (e.g., the Dual Mechanism Model; Clahsen, 1999) propose that compositional processes only apply to verbs that take productive affixes. For all other verbs, inflected forms are assumed to be stored in the lexicon in a non-decomposed manner. This account makes clear predictions about the consequences of disruption to the lexical access mechanisms involved in the spoken production of inflected forms. Briefly, it predicts that non-productive forms (which require lexical access) should be more affected than productive forms (which, depending on the language task, may not). We tested these predictions through the detailed analysis of the spoken production of a German-speaking individual with an acquired lexical impairment resulting from a stroke. Analyses of response accuracy, error types, and frequency effects revealed that combinatorial processes are not restricted to verbs that take productive inflections. On this basis, we propose an alternative account, the Stem-based Assembly Model (SAM) that posits that combinatorial processes may be available to all stems, and not only those that combine with productive affixes.
PMCID: PMC3117036  PMID: 21104479
Morphology; inflections; productivity; affixation; speech production; neuropsychology
11.  Patterns of brain reorganization subsequent to left fusiform damage: fMRI evidence from visual processing of words and pseudowords, faces and objects 
NeuroImage  2010;55(3):1357-1372.
Little is known about the neural reorganization that takes place subsequent to lesions that affect orthographic processing (reading and/or spelling). We report on an fMRI investigation of an individual with a left mid-fusiform resection that affected both reading and spelling (Tsapkini & Rapp, 2010). To investigate possible patterns of functional reorganization, we compared the behavioral and neural activation patterns of this individual with those of a group of control participants for the tasks of silent reading of words and pseudowords and the passive viewing of faces and objects, all tasks that typically recruit the inferior temporal lobes. This comparison was carried out with methods that included a novel application of Mahalanobis distance statistics, and revealed: (1) normal behavioral and neural responses for face and object processing, (2) evidence of neural reorganization bilaterally in the posterior fusiform that supported normal performance in pseudoword reading and which contributed to word reading (3) evidence of abnormal recruitment of the bilateral anterior temporal lobes indicating compensatory (albeit insufficient) recruitment of mechanisms for circumventing the word reading deficit.
PMCID: PMC3107009  PMID: 21168516
functional reorganization; reading; fusiform gyrus; Mahalanobis; anterior temporal lobe
12.  Examining the Central and Peripheral Processes of Written Word Production Through Meta-Analysis 
Producing written words requires “central” cognitive processes (such as orthographic long-term and working memory) as well as more peripheral processes responsible for generating the motor actions needed for producing written words in a variety of formats (handwriting, typing, etc.). In recent years, various functional neuroimaging studies have examined the neural substrates underlying the central and peripheral processes of written word production. This study provides the first quantitative meta-analysis of these studies by applying activation likelihood estimation (ALE) methods (Turkeltaub et al., 2002). For alphabet languages, we identified 11 studies (with a total of 17 experimental contrasts) that had been designed to isolate central and/or peripheral processes of word spelling (total number of participants = 146). Three ALE meta-analyses were carried out. One involved the complete set of 17 contrasts; two others were applied to subsets of contrasts to distinguish the neural substrates of central from peripheral processes. These analyses identified a network of brain regions reliably associated with the central and peripheral processes of word spelling. Among the many significant results, is the finding that the regions with the greatest correspondence across studies were in the left inferior temporal/fusiform gyri and left inferior frontal gyrus. Furthermore, although the angular gyrus (AG) has traditionally been identified as a key site within the written word production network, none of the meta-analyses found it to be a consistent site of activation, identifying instead a region just superior/medial to the left AG in the left posterior intraparietal sulcus. These meta-analyses and the discussion of results provide a valuable foundation upon which future studies that examine the neural basis of written word production can build.
PMCID: PMC3190188  PMID: 22013427
spelling; writing; meta-analysis; fMRI; dysgraphia; fusiform gyrus; intraparietal sulcus; angular
13.  Distinctions between orthographic long-term memory and working memory 
Cognitive neuropsychology  2009;26(8):724-751.
Research in the cognitive and neural sciences has long posited a distinction between the long-term memory (LTM) storage of information and the short-term buffering of information that is being actively manipulated in working memory (WM). This basic type of distinction has been posited in a variety of domains, including written language production—spelling. In the domain of spelling, the primary source of empirical evidence regarding this distinction has been cognitive neuropsychological studies reporting deficits selectively affecting what the cognitive neuropsychological literature has referred to as the orthographic lexicon (LTM) or the graphemic buffer (WM). Recent papers have reexamined several of the hallmark characteristics of impairment affecting the graphemic buffer, with implications for our understanding of the nature of the orthographic LTM and WM systems. In this paper, we present a detailed case series study of 4 individuals with acquired spelling deficits and report evidence from both error types and factors influencing error rates that support the traditional distinction between these cognitive systems involved in spelling. In addition, we report evidence indicating possible interaction between these systems, which is consistent with a variety of recent findings in research on spelling.
PMCID: PMC3145833  PMID: 20425660
Orthography; Spelling; Long-term memory; Working memory
14.  The Literate Brain: The Relationship between Spelling and Reading 
Journal of cognitive neuroscience  2010;23(5):1180-1197.
We report the results of an fMRI investigation of the neural bases of written language comprehension (reading) and production (spelling). Both tasks were examined in the same individuals, allowing greater precision in establishing the relationship between the neural underpinnings of these two cognitive functions. Also examined was the relationship between written language substrates and those involved in face and object (house) processing. The results reveal that reading and spelling share specific left hemisphere substrates in the mid-fusiform gyrus and in the inferior frontal gyrus/junction. Furthermore, the results indicate that the left mid-fusiform substrates are specifically involved in lexical orthographic processing. We also find that written language and face processing exhibit largely complementary activation patterns in both the fusiform and the inferior frontal/junction areas, with left and right lateralization, respectively. In sum, these results provide perhaps the strongest evidence to date of components that are shared by written language comprehension (reading) and production (spelling), and they further our understanding of the role of literacy within the larger repertoire of cognitive operations and their neural substrates.
PMCID: PMC3106999  PMID: 20433242
15.  Representation of Letter Position in Spelling: Evidence from Acquired Dysgraphia 
Cognition  2010;115(3):466-490.
The graphemic representations that underlie spelling performance must encode not only the identities of the letters in a word, but also the positions of the letters. This study investigates how letter position information is represented. We present evidence from two dysgraphic individuals, CM and LSS, who perseverate letters when spelling: that is, letters from previous spelling responses intrude into subsequent responses. The perseverated letters appear more often than expected by chance in the same position in the previous and subsequent responses. We used these errors to address the question of how letter position is represented in spelling. In a series of analyses we determined how often the perseveration errors produced maintain position as defined by a number of alternative theories of letter position encoding proposed in the literature. The analyses provide strong evidence that the grapheme representations used in spelling encode letter position such that position is represented in a graded manner based on distance from both edges of the word.
PMCID: PMC2953246  PMID: 20378104
letter position coding; dysgraphia; spelling; letter perseveration errors; orthographic processing
16.  The benefits and protective effects of behavioural treatment for dysgraphia in a case of primary progressive aphasia 
Aphasiology  2009;23(2):236-265.
Spoken and written language difficulties are the predominant symptoms in the progressive neurodegenerative disease referred to as primary progressive aphasia (PPA). There has been very little research on the effectiveness of intervention on spoken language impairments in this context and none directed specifically at progressive written language impairment.
To examine the effectiveness of behavioural intervention for dysgraphia in a case of primary progressive aphasia.
Methods & Procedures:
We carried out a longitudinal single-case study that allowed us to examine the effectiveness of a non-intensive spell-study-spell intervention procedure. We did so by comparing performance on four sets of words: trained, repeated, homework, and control words at five evaluations: baseline, during intervention, after the intervention, and at 6- and 12-month follow-up.
Outcomes & Results:
We find that: (1) at the end of the intervention, Trained words show a small but statistically significant improvement relative to baseline and an advantage in accuracy over Control, Homework, and Repeated word sets. (2) All word sets exhibited a decline in accuracy from the end of treatment to the 6-month follow-up evaluation, consistent with the degenerative nature of the illness. Nonetheless, accuracy on Trained words continued to be superior to that of Control words and not statistically different from pre-intervention baseline levels. (3) Repeated testing and practice at home yielded modest numerical advantages relative to Control words; but these differences were, for many comparisons, not statistically significant. (4) At 12 months post-intervention, all words sets had significantly declined relative to pre-intervention baselines and performance on the four sets was comparable.
This investigation documents—for the first time—that behavioural intervention can provide both immediate and short-term benefits for dysgraphia in the context of primary progressive aphasia.
PMCID: PMC3096931  PMID: 21603153
17.  Allographic agraphia: A case study 
We report the case of patient MN, diagnosed with frontotemporal dementia, who exhibited a severe impairment in writing letters and words in upper-case print in the face of accurate production of the same stimuli in lower-case cursive. In contrast to her written production difficulties, MN was unimpaired in recognizing visually presented letters and words in upper-case print. We find a modest benefit of visual form cueing in the written production of upper-case letters, despite an inability to describe or report visual features of letters in any case or font. This case increases our understanding of the allographic level of letter-shape representation in written language production. It provides strong support for previous reports indicating the neural independence of different types of case and font-specific letter-shape information; it provides evidence that letter-shape production does not require explicit access to information about the visual attributes of letter shapes and, finally, it reveals the possibility of interaction between processes involved in letter-shape production and perception.
PMCID: PMC3095950  PMID: 18489965
Allographic agraphia; Allograph; Letter font; Letter case; Graphemic representations
18.  The orthography-specific functions of the left fusiform gyrus: Evidence of modality and category specificity 
We report on an investigation of the cognitive functions of an individual with a resection of the left fusiform gyrus. This individual and a group of control participants underwent testing to examine the question of whether or not there are neural substrates within the left fusiform gyrus that are dedicated to orthographic processing. We evaluated the modality specificity (written vs. spoken language) and the category specificity (written language vs. other visual categories) of this individual’s impairments. The results clearly reveal deficits affecting lexical processes in both reading and spelling. Specifically, we find disruption of normal, rapid access to meaning from print in reading and of accurate retrieval of the spellings of words from their meaning in writing. These deficits stand in striking contrast with intact processing of spoken language and categories of visual stimuli such as line drawings of objects and faces. The modality and category specificity of the deficits provide clear evidence of neural substrates within the left mid-fusiform gyrus that are specialized and necessary for normal orthographic processing.
PMCID: PMC2844337  PMID: 19428003
fusiform gyrus; VWFA; reading; spelling; orthographic processing
19.  Mrs. Malaprop’s Neighborhood: Using Word Errors to Reveal Neighborhood Structure 
Journal of memory and language  2010;62(2):113-134.
Many theories of language production and perception assume that in the normal course of processing a word, additional non-target words (lexical neighbors) become active. The properties of these neighbors can provide insight into the structure of representations and processing mechanisms in the language processing system. To infer the properties of neighbors, we examined the non-semantic errors produced in both spoken and written word production by four individuals who suffered neurological injury. Using converging evidence from multiple language tasks, we first demonstrate that the errors originate in disruption to the processes involved in the retrieval of word form representations from long-term memory. The targets and errors produced were then examined for their similarity along a number of dimensions. A novel statistical simulation procedure was developed to determine the significance of the observed similarities between targets and errors relative to multiple chance baselines. The results reveal that in addition to position-specific form overlap (the only consistent claim of traditional definitions of neighborhood structure) the dimensions of lexical frequency, grammatical category, target length and initial segment independently contribute to the activation of non-target words in both spoken and written production. Additional analyses confirm the relevance of these dimensions for word production showing that, in both written and spoken modalities, the retrieval of a target word is facilitated by increasing neighborhood density, as defined by the results of the target-error analyses.
PMCID: PMC2808630  PMID: 20161591
20.  Phantom Tactile Sensations Modulated by Body Position 
Current biology : CB  2008;18(24):1937-1942.
Various lines of evidence reveal bilateral activation of somatosensory areas after unilateral stimulation [1-6] assumed to be mediated by cross-hemispheric connections [7-11]. Despite evidence of bilateral activity in response to unilateral stimulation, neurologically intact humans do not experience bilateral percepts when stimulated on one side of the body. This may be due to active suppression of ipsilateral neural activity [12, 13] by inhibitory mechanisms whose functioning is poorly understood. We describe an individual with left fronto-parietal damage who experiences bilateral sensations in response to unilateral tactile stimulation—a rarely reported condition known as synchiria (previously described in visual [14], auditory [15] and somatosensory modalities [16-19]). Presumably the phantom sensations result from normal bilateral cross-hemispheric activation, combined with a failure of inhibitory mechanisms to prevent bilateral perceptual experiences. The disruption of these mechanisms provides a valuable opportunity to examine their internal functioning. We find that the synchiria rate is affected by hand position relative to multiple reference frames. Specifically, synchiria decreases as the hands move from right (contralesional) to left (ipsilesional) space in trunk- and head-centered reference frames and disappears when the hands are crossed. These findings provide, for the first time, evidence that the mechanisms that inhibit bilateral percepts operate in multiple reference frames [20-27].
PMCID: PMC2701404  PMID: 19062276
21.  Deafness for the meanings of number words 
Neuropsychologia  2007;46(1):63-81.
We describe the performance of an aphasic individual who showed a selective impairment affecting his comprehension of auditorily presented number words and not other word categories. His difficulty in number word comprehension was restricted to the auditory modality, given that with visual stimuli (written words, Arabic numerals and pictures) his comprehension of number and non-number words was intact. While there have been previous reports of selective difficulty or sparing of number words at the semantic and post-semantic levels, this is the first reported case of a pre-semantic deficit that is specific to the category of number words. This constitutes evidence that lexical semantic distinctions are respected by modality-specific neural mechanisms responsible for providing access to the meanings of words.
PMCID: PMC2274996  PMID: 17915265
aphasia; numerals; category-specific deficits

Results 1-21 (21)