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1.  Support for Anterior Temporal Involvement in Semantic Error Production in Aphasia: New Evidence from VLSM 
Brain and language  2010;117(3):110-122.
Semantic errors in aphasia (e.g., naming a horse as “dog”) frequently arise from faulty mapping of concepts onto lexical items. A recent study by our group used voxel-based lesion-symptom mapping (VLSM) methods with 64 patients with chronic aphasia to identify voxels that carry an association with semantic errors. The strongest associations were found in the left anterior temporal lobe (L-ATL), in the mid- to anterior MTG region. The absence of findings in Wernicke’s area was surprising, as were indications that ATL voxels made an essential contribution to the post-semantic stage of lexical access. In this follow-up study, we sought to validate these results by re-defining semantic errors in a manner that was less theory dependent and more consistent with prior lesion studies. As this change also increased the robustness of the dependent variable, it made it possible to perform additional statistical analyses that further refined the interpretation. The results strengthen the evidence for a causal relationship between ATL damage and lexically-based semantic errors in naming and lend confidence to the conclusion that chronic lesions in Wernicke’s area are not causally implicated in semantic error production.
PMCID: PMC3037437  PMID: 20961612
aphasia; voxel-based lesion-symptom mapping; naming; semantic; errors
2.  The dorsal stream contribution to phonological retrieval in object naming 
Brain  2012;135(12):3799-3814.
Meaningful speech, as exemplified in object naming, calls on knowledge of the mappings between word meanings and phonological forms. Phonological errors in naming (e.g. GHOST named as ‘goath’) are commonly seen in persisting post-stroke aphasia and are thought to signal impairment in retrieval of phonological form information. We performed a voxel-based lesion-symptom mapping analysis of 1718 phonological naming errors collected from 106 individuals with diverse profiles of aphasia. Voxels in which lesion status correlated with phonological error rates localized to dorsal stream areas, in keeping with classical and contemporary brain-language models. Within the dorsal stream, the critical voxels were concentrated in premotor cortex, pre- and postcentral gyri and supramarginal gyrus with minimal extension into auditory-related posterior temporal and temporo-parietal cortices. This challenges the popular notion that error-free phonological retrieval requires guidance from sensory traces stored in posterior auditory regions and points instead to sensory-motor processes located further anterior in the dorsal stream. In a separate analysis, we compared the lesion maps for phonological and semantic errors and determined that there was no spatial overlap, demonstrating that the brain segregates phonological and semantic retrieval operations in word production.
PMCID: PMC3525060  PMID: 23171662
aphasia; dual-stream; voxel-based lesion-symptom mapping; naming; phonological errors; semantic errors
3.  Lexical retrieval and semantic knowledge in patients with left inferior temporal lobe lesions 
Aphasiology  2008;22(3):281-304.
It has been proposed that anomia following left inferior temporal lobe lesions may have two different underlying mechanisms with distinct neural substrates. Specifically, naming impairment following damage to more posterior regions (BA 37) has been considered to result from a disconnection between preserved semantic knowledge and phonological word forms (pure anomia), whereas anomia following damage to anterior temporal regions (BAs 38, 20/21) has been attributed to the degradation of semantic representations (semantic anomia). However, the integrity of semantic knowledge in patients with pure anomia has not been demonstrated convincingly, nor were lesions in these cases necessarily confined to BA 37. Furthermore, evidence of semantic anomia often comes from individuals with bilateral temporal lobe damage, so it is unclear whether unilateral temporal lobe lesions are sufficient to produce significant semantic impairment.
The main goals of this study were to determine whether anomia following unilateral left inferior temporal lobe damage reflected a loss of semantic knowledge or a post-semantic deficit in lexical retrieval and to identify the neuroanatomical correlates of the naming impairment.
Methods & Procedures
Eight individuals who underwent left anterior temporal lobectomy (L ATL) and eight individuals who sustained left posterior cerebral artery strokes (L PCA) completed a battery of language measures that assessed lexical retrieval and semantic processing, and 16 age- and education-matched controls also completed this battery. High-resolution structural brain scans were collected to conduct lesion analyses.
Outcomes & Results
Performance of L ATL and L PCA patients was strikingly similar, with both groups demonstrating naming performance ranging from moderately impaired to unimpaired. Anomia in both groups occurred in the context of mild deficits to semantic knowledge, which manifested primarily as greater difficulty in naming living things than nonliving things and greater difficulty in processing visual/perceptual as opposed to functional/associative semantic attributes. Lesion analyses indicated that both patient groups sustained damage to anterior inferior temporal lobe regions implicated in semantic processing.
These results contribute to a better understanding of the cognitive mechanism of naming impairment in patients with temporal lobe damage and support the notion that pure anomia and semantic anomia represent two endpoints along a continuum of semantic impairment. Unilateral left temporal lobe lesions in our patients resulted in relatively mild semantic deficits that were apparent primarily in lexical production tasks, whereas severe semantic impairment likely requires bilateral temporal lobe damage.
PMCID: PMC2743433  PMID: 19756227
4.  Connected speech production in three variants of primary progressive aphasia 
Brain  2010;133(7):2069-2088.
Primary progressive aphasia is a clinical syndrome defined by progressive deficits isolated to speech and/or language, and can be classified into non-fluent, semantic and logopenic variants based on motor speech, linguistic and cognitive features. The connected speech of patients with primary progressive aphasia has often been dichotomized simply as ‘fluent’ or ‘non-fluent’, however fluency is a multidimensional construct that encompasses features such as speech rate, phrase length, articulatory agility and syntactic structure, which are not always impacted in parallel. In this study, our first objective was to improve the characterization of connected speech production in each variant of primary progressive aphasia, by quantifying speech output along a number of motor speech and linguistic dimensions simultaneously. Secondly, we aimed to determine the neuroanatomical correlates of changes along these different dimensions. We recorded, transcribed and analysed speech samples for 50 patients with primary progressive aphasia, along with neurodegenerative and normal control groups. Patients were scanned with magnetic resonance imaging, and voxel-based morphometry was used to identify regions where atrophy correlated significantly with motor speech and linguistic features. Speech samples in patients with the non-fluent variant were characterized by slow rate, distortions, syntactic errors and reduced complexity. In contrast, patients with the semantic variant exhibited normal rate and very few speech or syntactic errors, but showed increased proportions of closed class words, pronouns and verbs, and higher frequency nouns, reflecting lexical retrieval deficits. In patients with the logopenic variant, speech rate (a common proxy for fluency) was intermediate between the other two variants, but distortions and syntactic errors were less common than in the non-fluent variant, while lexical access was less impaired than in the semantic variant. Reduced speech rate was linked with atrophy to a wide range of both anterior and posterior language regions, but specific deficits had more circumscribed anatomical correlates. Frontal regions were associated with motor speech and syntactic processes, anterior and inferior temporal regions with lexical retrieval, and posterior temporal regions with phonological errors and several other types of disruptions to fluency. These findings demonstrate that a multidimensional quantification of connected speech production is necessary to characterize the differences between the speech patterns of each primary progressive aphasic variant adequately, and to reveal associations between particular aspects of connected speech and specific components of the neural network for speech production.
PMCID: PMC2892940  PMID: 20542982
primary progressive aphasia; progressive non-fluent aphasia; semantic dementia; logopenic progressive aphasia; speech production
5.  Capturing multidimensionality in stroke aphasia: mapping principal behavioural components to neural structures 
Brain  2014;137(12):3248-3266.
Butler et al. relate behavioural deficits in 31 patients with chronic stroke aphasia to underlying neural structures. Using principal components analysis, they reduce a neuropsychological battery to three independent dimensions: phonological, semantic and executive-cognition. Phonological and semantic processing are linked to dorsal and ventral pathway integrity, respectively
Stroke aphasia is a multidimensional disorder in which patient profiles reflect variation along multiple behavioural continua. We present a novel approach to separating the principal aspects of chronic aphasic performance and isolating their neural bases. Principal components analysis was used to extract core factors underlying performance of 31 participants with chronic stroke aphasia on a large, detailed battery of behavioural assessments. The rotated principle components analysis revealed three key factors, which we labelled as phonology, semantic and executive/cognition on the basis of the common elements in the tests that loaded most strongly on each component. The phonology factor explained the most variance, followed by the semantic factor and then the executive-cognition factor. The use of principle components analysis rendered participants’ scores on these three factors orthogonal and therefore ideal for use as simultaneous continuous predictors in a voxel-based correlational methodology analysis of high resolution structural scans. Phonological processing ability was uniquely related to left posterior perisylvian regions including Heschl’s gyrus, posterior middle and superior temporal gyri and superior temporal sulcus, as well as the white matter underlying the posterior superior temporal gyrus. The semantic factor was uniquely related to left anterior middle temporal gyrus and the underlying temporal stem. The executive-cognition factor was not correlated selectively with the structural integrity of any particular region, as might be expected in light of the widely-distributed and multi-functional nature of the regions that support executive functions. The identified phonological and semantic areas align well with those highlighted by other methodologies such as functional neuroimaging and neurostimulation. The use of principle components analysis allowed us to characterize the neural bases of participants’ behavioural performance more robustly and selectively than the use of raw assessment scores or diagnostic classifications because principle components analysis extracts statistically unique, orthogonal behavioural components of interest. As such, in addition to improving our understanding of lesion–symptom mapping in stroke aphasia, the same approach could be used to clarify brain–behaviour relationships in other neurological disorders.
PMCID: PMC4240295  PMID: 25348632
lesion symptom mapping; aphasia; language processing; phonology; semantics
6.  Voxel-based lesion-parameter mapping: Identifying the neural correlates of a computational model of word production 
Cognition  2013;128(3):380-396.
The dual-route interactive two-step model explains the variation in the error patterns of aphasic speakers in picture naming, and word and nonword repetition tasks. The model has three parameters that can vary across individuals: the efficiency of the connections between semantic and lexical representations (s-weight), between lexical and phonological representations (p-weight), and between representations of auditory input and phonological representations (nl-weight). We determined these parameter values in 103 participants with chronic aphasia from left hemisphere stroke whose lesion locations had been determined. Then, using voxel-based lesion-parameter mapping, we mapped the parameters onto the brain, thus determining the neural correlates of the model’s mechanisms. The maps and the behavioral findings supported the model’s central claim that word repetition is affected by both the p and nl parameters. We propose that these two parameters constitute the model’s analogue of the “dorsal stream” component of neurocognitive models of language processing.
PMCID: PMC3709005  PMID: 23765000
Aphasia; Language production; Computational models; Voxel-based Lesion Parameter Mapping
7.  Repeating with the right hemisphere: reduced interactions between phonological and lexical-semantic systems in crossed aphasia? 
Knowledge on the patterns of repetition amongst individuals who develop language deficits in association with right hemisphere lesions (crossed aphasia) is very limited. Available data indicate that repetition in some crossed aphasics experiencing phonological processing deficits is not heavily influenced by lexical-semantic variables (lexicality, imageability, and frequency) as is regularly reported in phonologically-impaired cases with left hemisphere damage. Moreover, in view of the fact that crossed aphasia is rare, information on the role of right cortical areas and white matter tracts underpinning language repetition deficits is scarce. In this study, repetition performance was assessed in two patients with crossed conduction aphasia and striatal/capsular vascular lesions encompassing the right arcuate fasciculus (AF) and inferior frontal-occipital fasciculus (IFOF), the temporal stem and the white matter underneath the supramarginal gyrus. Both patients showed lexicality effects repeating better words than non-words, but manipulation of other lexical-semantic variables exerted less influence on repetition performance. Imageability and frequency effects, production of meaning-based paraphrases during sentence repetition, or better performance on repeating novel sentences than overlearned clichés were hardly ever observed in these two patients. In one patient, diffusion tensor imaging disclosed damage to the right long direct segment of the AF and IFOF with relative sparing of the anterior indirect and posterior segments of the AF, together with fully developed left perisylvian white matter pathways. These findings suggest that striatal/capsular lesions extending into the right AF and IFOF in some individuals with right hemisphere language dominance are associated with atypical repetition patterns which might reflect reduced interactions between phonological and lexical-semantic processes.
PMCID: PMC3798981  PMID: 24151460
right hemisphere; language; crossed aphasia; conduction aphasia; language network; structural connectivity
8.  Neurocognitive contributions to verbal fluency deficits in frontotemporal lobar degeneration 
Neurology  2009;73(7):535-542.
To test the hypothesis that different neurocognitive networks underlie verbal fluency deficits in frontotemporal lobar degeneration (FTLD).
Letter (“FAS”) and semantic (“animal”) fluency tests were administered to patients with a behavioral/dysexecutive disorder (bvFTLD; n = 71), semantic dementia (SemD; n = 21), and progressive nonfluent aphasia (PNFA; n = 26). Tests measuring working memory, naming/lexical retrieval, and semantic knowledge were also obtained. MRI voxel-based morphometry (VBM) studies were obtained on a subset of these patients (bvFTLD, n = 51; PNFA, n = 11; SemD, n = 10).
Patients with SemD were disproportionately impaired on the semantic fluency measure. Reduced output on this test was correlated with impaired performance on naming/lexical retrieval tests. VBM analyses related reduced letter and semantic fluency to anterior and inferior left temporal lobe atrophy. Patients with bvFTLD were equally impaired on both fluency tests. Poor performance on both fluency tests was correlated with low scores on working memory and naming/lexical retrieval measures. In this group, MRI-VBM analyses related letter fluency to bilateral frontal atrophy and semantic fluency to left frontal/temporal atrophy. Patients with PNFA were also equally impaired on fluency tests. Reduced semantic fluency output was correlated with reduced performance on naming/lexical retrieval tests. MRI-VBM analyses related semantic fluency to the right frontal lobe and letter fluency to left temporal atrophy.
Distinct neurocognitive networks underlie impaired performance on letter and semantic fluency tests in frontotemporal lobar degeneration subgroups.
= Alzheimer disease;
= analysis of variance;
= behavioral/dysexecutive subgroup;
= frontotemporal lobar degeneration;
= Mini-Mental State Examination;
= Montreal Neurological Institute;
= magnetic resonance;
= progressive nonfluent aphasia;
= semantic dementia;
= echo time;
= repetition time;
= voxel-based morphometry.
PMCID: PMC2730797  PMID: 19687454
9.  The anterior temporal lobes support residual comprehension in Wernicke’s aphasia 
Brain  2014;137(3):931-943.
Robson et al. use fMRI to investigate preserved written word and picture comprehension in Wernicke’s aphasia (impaired verbal comprehension following left temporoparietal damage). Bilaterally enhanced activation in the ventral and anterior temporal lobes as patients semantically process visually presented material emphasizes the importance of these regions for multimodal comprehension.
Wernicke’s aphasia occurs after a stroke to classical language comprehension regions in the left temporoparietal cortex. Consequently, auditory–verbal comprehension is significantly impaired in Wernicke’s aphasia but the capacity to comprehend visually presented materials (written words and pictures) is partially spared. This study used functional magnetic resonance imaging to investigate the neural basis of written word and picture semantic processing in Wernicke’s aphasia, with the wider aim of examining how the semantic system is altered after damage to the classical comprehension regions. Twelve participants with chronic Wernicke’s aphasia and 12 control participants performed semantic animate–inanimate judgements and a visual height judgement baseline task. Whole brain and region of interest analysis in Wernicke’s aphasia and control participants found that semantic judgements were underpinned by activation in the ventral and anterior temporal lobes bilaterally. The Wernicke’s aphasia group displayed an ‘over-activation’ in comparison with control participants, indicating that anterior temporal lobe regions become increasingly influential following reduction in posterior semantic resources. Semantic processing of written words in Wernicke’s aphasia was additionally supported by recruitment of the right anterior superior temporal lobe, a region previously associated with recovery from auditory-verbal comprehension impairments. Overall, the results provide support for models in which the anterior temporal lobes are crucial for multimodal semantic processing and that these regions may be accessed without support from classic posterior comprehension regions.
PMCID: PMC3927705  PMID: 24519979
Wernicke’s aphasia; semantic processing; language comprehension; anterior temporal lobe; Wernicke’s area
10.  Theoretical analysis of word production deficits in adult aphasia 
The cognitive analysis of adult language disorders continues to draw heavily on linguistic theory, but increasingly it reflects the influence of connectionist, spreading activation models of cognition. In the area of spoken word production, ‘localist’ connectionist models represent a natural evolution from the psycholingistic theories of earlier decades. By contrast, the parallel distributed processing framework forces more radical rethinking of aphasic impairments. This paper exemplifies these multiple influences in contemporary cognitive aphasiology. Topics include (i) what aphasia reveals about semantic-phonological interaction in lexical access; (ii) controversies surrounding the interpretation of semantic errors and (iii) a computational account of the relationship between naming and word repetition in aphasia. Several of these topics have been addressed using case series methods, including computational simulation of the individual, quantitative error patterns of diverse groups of patients and analysis of brain lesions that correlate with error rates and patterns. Efforts to map the lesion correlates of nonword errors in naming and repetition highlight the involvement of sensorimotor areas in the brain and suggest the need to better integrate models of word production with models of speech and action.
PMCID: PMC3866420  PMID: 24324234
aphasia; naming; repetition; errors; semantic; phonological
11.  Lexical-semantic Activation in Broca’s and Wernicke’s Aphasia: Evidence from Eye Movements 
Journal of cognitive neuroscience  2008;20(4):592-612.
Lexical processing requires both activating stored representations, and selecting among active candidates. The current work uses an eye-tracking paradigm to conduct a detailed temporal investigation of lexical processing. Patients with Broca's and Wernicke's aphasia are studied to shed light on the roles of anterior and posterior brain regions in lexical processing as well as the effects of lexical competition on such processing. Experiment 1 investigates whether objects semantically related to an uttered word are preferentially fixated, e.g., given the auditory target 'hammer', do participants fixate a picture of a nail? Results show that, like normals, both groups of patients are more likely to fixate on an object semantically related to the target than an unrelated object. Experiment 2 explores whether Broca's and Wernicke's aphasics show competition effects when words share onsets with the uttered word, e.g., given the auditory target 'hammer', do participants fixate a picture of a hammock? Experiment 3 investigates whether these patients activate words semantically related to onset competitors of the uttered word, e.g., given the auditory target 'hammock' do participants fixate a nail due to partial activation of the onset competitor hammer? Results of Experiments 2 and 3 show pathological patterns of performance for both Broca's and Wernicke's aphasics under conditions of lexical onset competition. However, the patterns of deficit differed, suggesting different functional and computational roles for anterior and posterior areas in lexical processing. Implications of the findings for the functional architecture of the lexical processing system and its potential neural substrates are considered.
PMCID: PMC3474198  PMID: 18052783
Broca’s aphasia; Wernicke’s aphasia; spoken word recognition; eye movements; cohort competition; semantic activation; lexical access
12.  Mechanisms of recovery from aphasia: evidence from positron emission tomography studies 
OBJECTIVES—Language functions comprise a distributed neural system, largely lateralised to the left cerebral hemisphere. Late recovery from aphasia after a focal lesion, other than by behavioural strategies, has been attributed to one of two changes at a systems level: a laterality shift, with mirror region cortex in the contralateral cortex assuming the function(s) of the damaged region; or a partial lesion effect, with recovery of perilesional tissue to support impaired language functions. Functional neuroimaging with PET allows direct observations of brain functions at systems level. This study used PET to compare regional brain activations in response to a word retrieval task in normal subjects and in aphasic patients who had shown at least some recovery and were able to attempt the task. Emphasis has been placed on single subject analysis of the results as there is no reason to assume that the mechanisms of recovery are necessarily uniform among aphasic patients.
METHODS—Six right handed aphasic patients, each with a left cerebral hemispheric lesion (five strokes and one glioma), were studied. Criteria for inclusion were symptomatic or formal test evidence of at least some recovery and an ability to attempt word retrieval in response to heard word cues. Each patient underwent 12 PET scans using oxygen-15 labelled water (H215O) as tracer to index regional cerebral blood flow (rCBF). The task, repeated six times, required the patient to think of verbs appropriate to different lists of heard noun cues. The six scans obtained during word retrieval were contrasted with six made while the subject was "at rest". The patients' individual results were compared with those of nine right handed normal volunteers undergoing the same activation study. The data were analysed using statistical parametric mapping (SPM96, Wellcome Department of Cognitive Neurology, London, UK).
RESULTS—Perception of the noun cues would be expected to result in bilateral dorsolateral temporal cortical activations, but as the rate of presentation was only four per minute the auditory perceptual activations were not evident in all people. Anterior cingulate, medial premotor (supplementary speech area) and dorsolateral frontal activations were evident in all normal subjects and patients. There were limited right dorsolateral frontal activations in three of the six patients, but a similar pattern was also found in four of the nine normal subjects. In the left inferolateral temporal cortex, activation was found for the normal subjects and five of the six patients, including two of the three subjects with lesions involving the left temporal lobe. The only patient who showed subthreshold activation in the left inferolateral temporal activation had a very high error rate when performing the verb retrieval task.
CONCLUSIONS—The normal subjects showed a left lateralised inferolateral temporal activation, reflecting retrieval of words appropriate in meaning to the cue from the semantic system. Lateralisation of frontal activations to the left was only relative, with right prefrontal involvement in half of the normal subjects. Frontal activations are associated with parallel psychological processes involved in word retrieval, including task initiation, short term (working) memory for the cue and responses, and prearticulatory processes (even though no overt articulation was required). There was little evidence of a laterality shift of word retrieval functions to the right temporal lobe after a left hemispheric lesion. In particular, left inferolateral temporal activation was seen in all patients except one, and he proved to be very inefficient at the task. The results provide indirect evidence that even limited salvage of peri-infarct tissue with acute stroke treatments will have an important impact on the rehabilitation of cognitive functions.

PMCID: PMC1736204  PMID: 10071093
13.  Where (in the brain) do semantic errors come from? 
Semantic errors result from the disruption of access either to semantics or to lexical representations. One way to determine the origins of these errors is to evaluate comprehension of words that elicit semantic errors in naming. We hypothesized that in acute stroke there are different brain regions where dysfunction results in semantic errors in both naming and comprehension versus those with semantic errors in oral naming alone.
A consecutive series of 196 patients with acute left hemispheric stroke who met inclusion criteria were evaluated with oral naming and spoken word/picture verification tasks and magnetic resonance imaging within 48 hours of stroke onset. We evaluated the relationship between tissue dysfunction in 10 pre-specified Brodmann's areas (BA) and the production of coordinate semantic errors resulting from (1) semantic deficits or (2) lexical access deficits.
Semantic errors arising from semantic deficits were most associated with tissue dysfunction/infarct of left BA 22. Semantic errors resulting from lexical access deficits were associated with hypoperfusion/infarct of left BA 37.
Our study shows that semantic errors arising from damage to distinct cognitive processes reflect dysfunction of different brain regions.
PMCID: PMC2659726  PMID: 19084219
aphasia; perfusion-weighted magnetic resonance imaging; semantics; acute ischemic stroke
14.  The impact of semantic impairment on verbal short-term memory in stroke aphasia and semantic dementia: A comparative study 
Journal of memory and language  2008;58(1):66-87.
This study presents the first direct comparison of immediate serial recall in semantic dementia (SD) and transcortical sensory aphasia (TSA). Previous studies of the effect of semantic impairment on verbal short-term memory (STM) have led to important theoretical advances. However, different conclusions have been drawn from these two groups. This research aimed to explain these inconsistencies. We observed (a) qualitative differences between SD and TSA in the nature of the verbal STM impairment and (b) considerable variation within the TSA group. The SD and TSA patients all had poor semantic processing and good phonology. Reflecting this, both groups remained sensitive to phonological similarity and showed a reduced effect of lexicality in immediate serial recall. The SD patients showed normal serial position effects; in contrast, the TSA patients had poor recall of the initial list items and exhibited large recency effects on longer lists. The error patterns of the two groups differed: the SD patients made numerous phoneme migration errors whereas the TSA group were more likely to produce entire words in the wrong order, often initiating recall with terminal list items. The SD cases also showed somewhat larger effects of word frequency and imageability. We propose that these contrasting performance patterns are explicable in terms of the nature of the underlying semantic impairment. SD is associated with anterior lobe atrophy and produces degradation of semantic knowledge – this is more striking for less frequent/imageable items, accentuating the effects of these lexical/semantic variables in STM. SD patients frequently recombine the phonemes of different list items due to the reduced semantic constraint upon phonology (semantic binding: Patterson et al., 1994). In contrast, the semantic impairment in TSA follows frontal or temporoparietal lesions and is associated with poor executive control of semantic processing (deregulated semantic cognition: Jefferies and Lambon Ralph, 2006), explaining why these patients are liable to recall entire words out of serial order.
PMCID: PMC2344152  PMID: 18438454
15.  Grey and white matter correlates of picture naming: Evidence from a voxel-based lesion analysis of the Boston Naming Test 
A number of recent studies utilizing both functional neuroimaging and lesion analysis techniques in neurologic patients have produced conflicting results with respect to the neural correlates of picture naming. Picture naming involves a number of cognitive processes, from visual perception/recognition to lexical-semantic retrieval to articulation. This middle process, the ability to retrieve a name associated with an object, has been attributed in some cases to posterior portions of the left lateral temporal lobe and in other cases, to anterior temporal cortex. In the current study, we used voxel-based lesion symptom mapping (VLSM) to identify neural correlates of picture naming in a large sample of well-characterized left hemisphere (LH) patients suffering from a range of naming deficits. We tested patients on the Boston Naming Test (BNT), a clinical, standardized measure of picture naming that is widely used in both clinical and research settings. We found that overall performance on the BNT was associated with a network of LH regions that included significant portions of the left anterior to posterior middle temporal gyrus (MTG) and superior temporal gyrus (STG) and underlying white matter, and extended into left inferior parietal cortex. However, when we added covariates to this analysis that controlled for deficits in visual recognition and motor speech in order to isolate brain regions specific to lexical-semantic retrieval, the significant regions that remained were confined almost exclusively to the left mid-posterior MTG and underlying white matter. These findings support the notion that a large network in left peri-Sylvian cortex supports picture naming, but that the left mid-posterior MTG and underlying white matter play a critical role in the core ability to retrieve a name associated with an object or picture.
PMCID: PMC3613759  PMID: 22482693
Picture naming; Word production; Temporal cortex; Wernicke’s aphasia; Aphasia; Middle temporal gyrus; Lexical-semantics
16.  Critical brain regions for action recognition: lesion symptom mapping in left hemisphere stroke 
Brain  2010;133(11):3269-3280.
A number of conflicting claims have been advanced regarding the role of the left inferior frontal gyrus, inferior parietal lobe and posterior middle temporal gyrus in action recognition, driven in part by an ongoing debate about the capacities of putative mirror systems that match observed and planned actions. We report data from 43 left hemisphere stroke patients in two action recognition tasks in which they heard and saw an action word (‘hammering’) and selected from two videoclips the one corresponding to the word. In the spatial recognition task, foils contained errors of body posture or movement amplitude/timing. In the semantic recognition task, foils were semantically related (sawing). Participants also performed a comprehension control task requiring matching of the same verbs to objects (hammer). Using regression analyses controlling for both the comprehension control task and lesion volume, we demonstrated that performance in the semantic gesture recognition task was predicted by per cent damage to the posterior temporal lobe, whereas the spatial gesture recognition task was predicted by per cent damage to the inferior parietal lobule. A whole-brain voxel-based lesion symptom-mapping analysis suggested that the semantic and spatial gesture recognition tasks were associated with lesioned voxels in the posterior middle temporal gyrus and inferior parietal lobule, respectively. The posterior middle temporal gyrus appears to serve as a central node in the association of actions and meanings. The inferior parietal lobule, held to be a homologue of the monkey parietal mirror neuron system, is critical for encoding object-related postures and movements, a relatively circumscribed aspect of gesture recognition. The inferior frontal gyrus, on the other hand, was not predictive of performance in any task, suggesting that previous claims regarding its role in action recognition may require refinement.
PMCID: PMC2965423  PMID: 20805101
action; recognition; apraxia; stroke; voxel-based lesion symptom mapping
17.  Bilateral Capacity for Speech Sound Processing in Auditory Comprehension: Evidence from Wada Procedures 
Brain and language  2008;107(3):179-184.
Data from lesion studies suggests that the ability to perceive speech sounds, as measured by auditory comprehension tasks, is supported by temporal lobe systems in both the left and right hemisphere. For example, patients with left temporal lobe damage and auditory comprehension deficits (i.e., Wernicke’s aphasics), nonetheless comprehend isolated words better than one would expect if their speech perception system had been largely destroyed (70–80% accuracy). Further, when comprehension fails in such patients their errors are more often semantically-based, than-phonemically based. The question addressed by the present study is whether this ability of the right hemisphere to process speech sounds is a result of plastic reorganization following chronic left hemisphere damage, or whether the ability exists in undamaged language systems. We sought to test these possibilities by studying auditory comprehension in acute left versus right hemisphere deactivation during Wada procedures. A series of 20 patients undergoing clinically indicated Wada procedures were asked to listen to an auditorily presented stimulus word, and then point to its matching picture on a card that contained the target picture, a semantic foil, a phonemic foil, and an unrelated foil. This task was performed under three conditions, baseline, during left carotid injection of sodium amytal, and during right carotid injection of sodium amytal. Overall, left hemisphere injection led to a significantly higher error rate than right hemisphere injection. However, consistent with lesion work, the majority (75%) of these errors were semantic in nature. These findings suggest that auditory comprehension deficits are predominantly semantic in nature, even following acute left hemisphere disruption. This, in turn, supports the hypothesis that the right hemisphere is capable of speech sound processing in the intact brain.
PMCID: PMC2644214  PMID: 18976806
18.  Neurology of anomia in the semantic variant of primary progressive aphasia 
Brain  2009;132(9):2553-2565.
The semantic variant of primary progressive aphasia (PPA) is characterized by the combination of word comprehension deficits, fluent aphasia and a particularly severe anomia. In this study, two novel tasks were used to explore the factors contributing to the anomia. The single most common factor was a blurring of distinctions among members of a semantic category, leading to errors of overgeneralization in word–object matching tasks as well as in word definitions and object descriptions. This factor was more pronounced for natural kinds than artifacts. In patients with the more severe anomias, conceptual maps were more extensively disrupted so that inter-category distinctions were as impaired as intra-category distinctions. Many objects that could not be named aloud could be matched to the correct word in patients with mild but not severe anomia, reflecting a gradual intensification of the semantic factor as the naming disorder becomes more severe. Accurate object descriptions were more frequent than accurate word definitions and all patients experienced prominent word comprehension deficits that interfered with everyday activities but no consequential impairment of object usage or face recognition. Magnetic resonance imaging revealed three characteristics: greater atrophy of the left hemisphere; atrophy of anterior components of the perisylvian language network in the superior and middle temporal gyri; and atrophy of anterior components of the face and object recognition network in the inferior and medial temporal lobes. The left sided asymmetry and perisylvian extension of the atrophy explains the more profound impairment of word than object usage and provides the anatomical basis for distinguishing the semantic variant of primary progressive aphasia from the partially overlapping group of patients that fulfil the widely accepted diagnostic criteria for semantic dementia.
PMCID: PMC2766179  PMID: 19506067
aphasia; frontotemporal lobar degeneration; language processing; progressive aphasia; semantic categorization
Brain and language  2011;117(1):28-33.
Few studies have directly compared the clinical and anatomical characteristics of patients with progressive aphasia to those of patients with aphasia caused by stroke. In the current study we examined fluent forms of aphasia in these two groups, specifically the semantic dementia (SD) and persisting Wernicke's aphasia (WA) due to stroke. We compared 10 patients with SD to 10 age- and education-matched patients with WA in three language domains: language comprehension (single words and sentences), spontaneous speech and visual semantics. Neuroanatomical involvement was analyzed using disease-specific image analysis techniques: voxel-based morphometry (VBM) for patients with SD and overlays of lesion masks in patients with WA. Patients with SD and WA were both impaired on tasks that involved visual semantics, but patients with SD were less impaired in spontaneous speech and sentence comprehension. The anatomical findings showed that different regions were most affected in the two disorders: the left anterior temporal lobe in SD and the left posterior middle temporal gyrus in chronic WA. This study highlights that the two syndromes classically associated with language comprehension deficits in aphasia due to stroke and neurodegenerative disease are clinically distinct, most likely due to distinct distributions of damage in the temporal lobe.
PMCID: PMC3160783  PMID: 21315437
semantic dementia; Wernicke’s aphasia; voxel-based morphometry; comprehension impairments; stroke; primary progressive aphasia
20.  The Neural Correlates of Abstract and Concrete Words: Evidence from Brain-Damaged Patients 
Brain Sciences  2013;3(3):1229-1243.
Neuropsychological and activation studies on the neural correlates of abstract and concrete words have produced contrasting results. The present study explores the anatomical substrates of abstract/concrete words in 22 brain-damaged patients with a single vascular lesion either in the right or left hemisphere. One hundred and twenty (60 concrete and 60 abstract) noun triplets were used for a semantic similarity judgment task. We found a significant interaction in word type × group since left temporal brain-damaged patients performed significantly better with concrete than abstract words. Lesion mapping of patients with predominant temporal damage showed that the left superior and middle temporal gyri and the insula were the areas of major overlapping, while the anterior portion of the left temporal lobe was generally spared. Errors on abstract words mainly concerned (although at a non-significant level) semantically associate targets, while in the case of concrete words, coordinate targets were significantly more impaired than associate ones. Our results suggest that the left superior and middle temporal gyri and the insula are crucial regions in processing abstract words. They also confirm the hypothesis of a semantic similarity vs. associative organization of concrete and abstract concepts.
PMCID: PMC4061881  PMID: 24961527
concreteness; temporal lobe; insula; associative vs. categorical architecture
21.  Reading Disorders in Primary Progressive Aphasia: a behavioral and neuroimaging study 
Neuropsychologia  2009;47(8-9):1893-1900.
Previous neuropsychological studies on acquired dyslexia revealed a double dissociation in reading impairments. Patients with phonological dyslexia have selective difficulty reading pseudo-words, while those with surface dyslexia misread exception words. This double dissociation in reading abilities has often been reported in brain damaged patients, but it has not been consistently shown in patients with neurodegenerative diseases.
In this study, we investigated reading impairments and their anatomical correlates in various neurodegenerative diseases. First, we performed a behavioral analysis to characterize the reading of different word types in primary progressive aphasia (PPA). Then, we conducted a voxel-based morphometry neuroimaging study to map the brain areas in which gray matter volume correlated with accurate reading of exception and pseudo-words.
The results showed a differential pattern of exception and pseudo-word reading abilities in different clinical variants of PPA. Patients with semantic dementia, a disorder characterized by selective loss of semantic memory, revealed a pattern of surface dyslexia, while patients with logopenic/phonological progressive aphasia, defined by phonological loop deficits, showed phonological dyslexia. Neuroimaging results showed that exception word reading accuracy correlated with gray matter volume in the left anterior temporal structures, including the temporal pole, the anterior superior and middle temporal and fusiform gyri, while pseudo-word reading accuracy correlated with left temporoparietal regions, including the posterior superior and middle temporal and fusiform gyri, and the inferior parietal lobule.
These results suggest that exception and pseudo-word reading not only rely upon different language mechanisms selectively damaged in PPA, but also that these processes are sustained by separate brain structures.
PMCID: PMC2734967  PMID: 19428421
exception words; pseudo-words; neurodegenerative diseases; voxel-based morphometry
22.  The neural basis of inhibitory effects of semantic and phonological neighbors in spoken word production 
Journal of cognitive neuroscience  2013;25(9):1504-1516.
Theories of word production and word recognition generally agree that multiple word candidates are activated during processing. The facilitative and inhibitory effects of these “lexical neighbors” have been studied extensively using behavioral methods and have spurred theoretical development in psycholinguistics, but relatively little is known about the neural basis of these effects and how lesions may affect them. The present study used voxel-wise lesion overlap subtraction to examine semantic and phonological neighbor effects in spoken word production following left hemisphere stroke. Increased inhibitory effects of near semantic neighbors were associated with inferior frontal lobe lesions, suggesting impaired selection among strongly activated semantically-related candidates. Increased inhibitory effects of phonological neighbors were associated with posterior superior temporal and inferior parietal lobe lesions. In combination with previous studies, these results suggest that such lesions cause phonological-to-lexical feedback to more strongly activate phonologically-related lexical candidates. The comparison of semantic and phonological neighbor effects and how they are affected by left hemisphere lesions provides new insights into the cognitive dynamics and neural basis of phonological, semantic, and cognitive control processes in spoken word production.
PMCID: PMC4038097  PMID: 23647518
lexical neighborhood; phonological neighborhood; semantic neighborhood; spoken word production; aphasia; lesion overlap
23.  “Pre-semantic” cognition revisited: Critical differences between semantic aphasia and semantic dementia 
Neuropsychologia  2010;48(1):248-261.
Patients with semantic dementia show a specific pattern of impairment on both verbal and non-verbal “pre-semantic” tasks: e.g., reading aloud, past tense generation, spelling to dictation, lexical decision, object decision, colour decision and delayed picture copying. All seven tasks are characterised by poorer performance for items that are atypical of the domain and “regularisation errors” (irregular/atypical items are produced as if they were domain-typical). The emergence of this pattern across diverse tasks in the same patients indicates that semantic memory plays a key role in all of these types of “pre-semantic” processing. However, this claim remains controversial because semantically-impaired patients sometimes fail to show an influence of regularity. This study demonstrates that (a) the location of brain damage and (b) the underlying nature of the semantic deficit affect the likelihood of observing the expected relationship between poor comprehension and regularity effects. We compared the effect of multimodal semantic impairment in the context of semantic dementia and stroke aphasia on the seven “pre-semantic” tasks listed above. In all of these tasks, the semantic aphasia patients were less sensitive to typicality than the semantic dementia patients, even though the two groups obtained comparable scores on semantic tests. The semantic aphasia group also made fewer regularisation errors and many more unrelated and perseverative responses. We propose that these group differences reflect the different locus for the semantic impairment in the two conditions: patients with semantic dementia have degraded semantic representations, whereas semantic aphasia patients show deregulated semantic cognition with concomitant executive deficits. These findings suggest a reinterpretation of single case studies of comprehension-impaired aphasic patients who fail to show the expected effect of regularity on “pre-semantic” tasks. Consequently, such cases do not demonstrate the independence of these tasks from semantic memory.
PMCID: PMC2805958  PMID: 19766662
24.  Anatomic, clinical, and neuropsychological correlates of spelling errors in Primary Progressive Aphasia 
Neuropsychologia  2012;50(8):1929-1935.
This study evaluates spelling errors in the three subtypes of primary progressive aphasia (PPA): agrammatic (PPA-G), logopenic (PPA-L), and semantic (PPA-S). Forty one PPA-patients and 36 age-matched healthy controls were administered a test of spelling. The total number of errors and types of errors in spelling to dictation of regular words, exception words and nonwords, were recorded. Error types were classified based on phonetic plausibility. In the first analysis, scores were evaluated by clinical diagnosis. Errors in spelling exception words and phonetically plausible errors were seen in PPA-S. Conversely, PPA-G was associated with errors in nonword spelling and phonetically implausible errors. In the next analysis, spelling scores were correlated to other neuropsychological language test scores. Significant correlations were found between exception word spelling and measures of naming and single word comprehension. Nonword spelling correlated with tests of grammar and repetition. Global language measures did not correlate significantly with spelling scores, however. Cortical thickness analysis based on MRI showed that atrophy in several language regions of interest were correlated with spelling errors. Atrophy in the left supramarginal gyrus and inferior frontal gyrus (IFG) pars orbitalis correlated with errors in nonword spelling, while thinning in the left temporal pole and fusiform gyrus correlated with errors in exception word spelling. Additionally, phonetically implausible errors in regular word spelling correlated with thinning in the left IFG pars triangularis and pars opercularis. Together, these findings suggest two independent systems for spelling to dictation, one phonetic (phoneme to grapheme conversion), and one lexical (whole word retrieval).
PMCID: PMC3527105  PMID: 22579708
Dementia; Primary Progressive Aphasia; Agraphia; Cortical atrophy; Frontotemporal dementia
25.  Density Pervades: An Analysis of Phonological Neighborhood Density Effects in Aphasic Speakers with Different Types of Naming Impairment 
Cognitive neuropsychology  2010;27(5):401-427.
We investigated the influence of phonological neighborhood density (PND) on the performance of aphasic speakers whose naming impairments differentially implicate phonological or semantic stages of lexical access. A word comes from a dense phonological neighborhood if many words sound like it. Limited evidence suggests that higher density facilitates naming in aphasic speakers, as it does in healthy speakers. Using well controlled stimuli, Experiment 1 confirmed the influence of PND on accuracy and phonological error rates in two aphasic speakers with phonological processing deficits. In Experiments 2 and 3, we extended the investigation to an aphasic speaker who is prone to semantic errors, indicating a semantic deficit and/or a deficit in the mapping from semantics to words. This individual had higher accuracy, and fewer semantic errors, in naming targets from high versus low density neighborhoods. It is argued that the results provide strong support for interactive approaches to lexical access, where reverberatory feedback between word- and phoneme-level lexical representations not only facilitates phonological level processes but also privileges the selection of a target word over its semantic competitors.
PMCID: PMC3132149  PMID: 21718214
phonology; neighborhood density; aphasia; interactivity; lexical access

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