A Spatiotemporal Map of Reading Aloud.

Reading words aloud is a fundamental aspect of literacy. The rapid rate at which multiple distributed neural substrates are engaged in this process can only be probed via techniques with high spatiotemporal resolution. We probed this with direct intracranial recordings covering most of the left hemisphere in 46 humans (26 male, 20 female) as they read aloud regular, exception and pseudo-words. We used this to create a spatiotemporal map of word processing and to derive how broadband γ activity varies with multiple word attributes critical to reading speed: lexicality, word frequency, and orthographic neighborhood. We found that lexicality is encoded earliest in mid-fusiform (mFus) cortex, and precentral sulcus, and is represented reliably enough to allow single-trial lexicality decoding. Word frequency is first represented in mFus and later in the inferior frontal gyrus (IFG) and inferior parietal sulcus (IPS), while orthographic neighborhood sensitivity resides solely in IPS. We thus isolate the neural correlates of the distributed reading network involving mFus, IFG, IPS, precentral sulcus, and motor cortex and provide direct evidence for parallel processes via the lexical route from mFus to IFG, and the sublexical route from IPS and precentral sulcus to anterior IFG.SIGNIFICANCE STATEMENT Reading aloud depends on multiple complex cerebral computations: mapping from a written letter string on a page to a sequence of spoken sound representations. Here, we used direct intracranial recordings in a large cohort while they read aloud known and novel words, to track, across space and time, the progression of neural representations of behaviorally relevant factors that govern reading speed. We find, concordant with cognitive models of reading, that known and novel words are differentially processed through a lexical route, sensitive to frequency of occurrence of known words in natural language, and a sublexical route, performing letter-by-letter construction of novel words.

Critical role of the ventral temporal lobe in naming.

OBJECTIVE: Lexical retrieval deficits are characteristic of a variety of different neurological disorders. However, the exact substrates responsible for this are not known. We studied a large cohort of patients undergoing surgery in the dominant temporal lobe for medically intractable epilepsy (n = 95) to localize brain regions that were associated with anomia. METHODS: We performed a multivariate voxel-based lesion-symptom mapping analysis to correlate surgical lesions within the temporal lobe with changes in naming ability. Additionally, we used a surface-based mixed-effects multilevel analysis to estimate group-level broadband gamma activity during naming across a subset of patients with electrocorticographic recordings and integrated these results with lesion-deficit findings. RESULTS: We observed that ventral temporal regions, centered around the middle fusiform gyrus, were significantly associated with a decline in naming. Furthermore, we found that the ventral aspect of temporal lobectomies was linearly correlated to a decline in naming, with a clinically significant decline occurring once the resection extended 6 cm from the anterior tip of the temporal lobe on the ventral surface. On electrocorticography, the majority of these cortical regions were functionally active following visual processing. These loci coincide with the sites of susceptibility artifacts during echoplanar imaging, which may explain why this region has been previously underappreciated as the locus responsible for postoperative naming deficits. SIGNIFICANCE: Taken together, these data highlight the crucial contribution of the ventral temporal cortex in naming and its important role in the pathophysiology of anomia following temporal lobe resections. As such, surgical strategies should attempt to preserve this region to mitigate postoperative language deficits.

Bayesian inference for stationary points in Gaussian process regression models for event-related potentials analysis.

Stationary points embedded in the derivatives are often critical for a model to be interpretable and may be considered as key features of interest in many applications. We propose a semiparametric Bayesian model to efficiently infer the locations of stationary points of a nonparametric function, which also produces an estimate of the function. We use Gaussian processes as a flexible prior for the underlying function and impose derivative constraints to control the function's shape via conditioning. We develop an inferential strategy that intentionally restricts estimation to the case of at least one stationary point, bypassing possible mis-specifications in the number of stationary points and avoiding the varying dimension problem that often brings in computational complexity. We illustrate the proposed methods using simulations and then apply the method to the estimation of event-related potentials derived from electroencephalography (EEG) signals. We show how the proposed method automatically identifies characteristic components and their latencies at the individual level, which avoids the excessive averaging across subjects that is routinely done in the field to obtain smooth curves. By applying this approach to EEG data collected from younger and older adults during a speech perception task, we are able to demonstrate how the time course of speech perception processes changes with age.

The domain-specificity of serial order working memory.

Making a turn while driving is simple: turn on the indicator, check for cars, then turn. Two types of information are required to perform this sequence of events: information about the items (e.g., the correct indicator), and the serial order of those items (e.g., checking before turning rather than vice-versa). Previous research has found distinct working memory capacities (WMCs) for item and serial order information in both verbal and nonverbal domains. The current study investigates whether the serial order WMC is shared for sequences from different content domains. One hundred and fifty-three participants performed sequence matching tasks with verbal (letters and words) and nonverbal (locations and arrows) stimuli. The accuracy of detecting mismatched item-identity and serial order information in sequences was used to operationalize item and order WMC. Using structural equation modeling analyses, we directly compared models that included either domain-specific or domain-general serial order WMC latent variables, finding that models with domain-specific serial order WMC latent variables for verbal and nonverbal materials fit the data better than models with domain-general latent variables. The findings support the hypothesis that there are separate capacities for serial order working memory depending on the type of material being ordered.

Word deafness with preserved number word perception.

We describe the performance of an aphasic individual, K.A., who showed a selective impairment affecting his ability to perceive spoken language, while largely sparing his ability to perceive written language and to produce spoken language. His spoken perception impairment left him unable to distinguish words or nonwords that differed on a single phoneme and he was no better than chance at auditory lexical decision or single spoken word and single picture matching with phonological foils. Strikingly, despite this profound impairment, K.A. showed a selective sparing in his ability to perceive number words, which he was able to repeat and comprehend largely without error. This case adds to a growing literature demonstrating modality-specific dissociations between number word and non-number word processing. Because of the locus of K.A.'s speech perception deficit for non-number words, we argue that this distinction between number word and non-number word processing arises at a sublexical level of representations in speech perception, in a parallel fashion to what has previously been argued for in the organization of the sublexical level of representation for speech production.

How modality-specific is morphology?

Writing has long been considered to be dependent on speaking. However, modality-specific dissociations between written and spoken word production imply that word production is supported by distinct neural mechanisms in writing, which can be impaired or spared regardless of the intactness of spoken word production. Rapp et al. (2015). Modality and morphology: What we write may not be what we say. Psychological Science, 26, 892-902 documented a double dissociation where problems with regular inflections were selectively restricted to writing or speaking. We report on two English-speaking aphasic individuals who exhibit this same modality-specific dissociation of inflectional processing, replicating the original findings. We expand on Rapp et al.'s study by examining whether the dissociations observed with regular inflections extend to other morphological forms, such as derivation and irregular inflection. Results showed that the dissociation holds for derivation; however, both participants were impaired with irregular inflections in both output modalities. Implications of these findings for morphological processing and the independence of the orthographic system are discussed.

Brain Modularity Mediates the Relation between Task Complexity and Performance.

Recent work in cognitive neuroscience has focused on analyzing the brain as a network, rather than as a collection of independent regions. Prior studies taking this approach have found that individual differences in the degree of modularity of the brain network relate to performance on cognitive tasks. However, inconsistent results concerning the direction of this relationship have been obtained, with some tasks showing better performance as modularity increases and other tasks showing worse performance. A recent theoretical model [Chen, M., & Deem, M. W. 2015. Development of modularity in the neural activity of children's brains. Physical Biology, 12, 016009] suggests that these inconsistencies may be explained on the grounds that high-modularity networks favor performance on simple tasks whereas low-modularity networks favor performance on more complex tasks. The current study tests these predictions by relating modularity from resting-state fMRI to performance on a set of simple and complex behavioral tasks. Complex and simple tasks were defined on the basis of whether they did or did not draw on executive attention. Consistent with predictions, we found a negative correlation between individuals' modularity and their performance on a composite measure combining scores from the complex tasks but a positive correlation with performance on a composite measure combining scores from the simple tasks. These results and theory presented here provide a framework for linking measures of whole-brain organization from network neuroscience to cognitive processing.

Single-case cognitive neuropsychology in the age of big data.

Historically, single-case studies of brain-damaged individuals have contributed substantially to our understanding of cognitive processes. However, the role of single-case cognitive neuropsychology has diminished with the proliferation of techniques that measure neural activity in humans. Instead, large-scale informatics approaches in which data are gathered from hundreds of neuroimaging studies have become popular. It has been claimed that utilizing these informatics approaches can address problems found in single imaging studies. We first discuss reasons for why cognitive neuropsychology is thought to be in decline. Next, we note how these informatics approaches, while having benefits, are not particularly suited for understanding functional architectures. We propose that the single-case cognitive neuropsychological approach, which is focused on developing models of cognitive processing, addresses several of the weaknesses inherent in informatics approaches. Furthermore, we discuss how using neural data from brain-damaged individuals provides data that can inform both cognitive and neural models of cognitive processing.

Neuroplasticity and the logic of cognitive neuropsychology.

More than thirty years ago, Alfonso Caramazza laid out assumptions for drawing inferences about the undamaged cognitive system from individuals with brain damage. Since then, these assumptions have been challenged including the transparency or subtractivity assumption, that the cognitive system does not reorganize following brain damage. It has been repeatedly demonstrated that brains are highly plastic. However, there is no clear connection between brain plasticity and cognitive reorganization. Brain plasticity research does not require a rethinking of the core logic of cognitive neuropsychology. Differences in task-based activation between damaged and undamaged brains provide little insight into the cognitive architectures of brain-damaged patients. Theory and methods are needed to understand cognitive neuroplasticity, or how neural reorganization that follows brain damage relates to reorganization of functions. We discuss alternative types of cognitive neuroplasticity that may occur in damaged brains and consider how they impact the basic logic of cognitive neuropsychology.

The Cognitive Neuroplasticity of Reading Recovery following Chronic Stroke: A Representational Similarity Analysis Approach.

Damage to certain left hemisphere regions leads to reading impairments, at least acutely, though some individuals eventually recover reading. Previous neuroimaging studies have shown a relationship between reading recovery and increases in contralesional and perilesional activation during word reading tasks, relative to controls. Questions remain about how to interpret these changes in activation. Do these changes reflect functional take-over, a reorganization of functions in the damaged brain? Or do they reveal compensatory masquerade or the use of alternative neural pathways to reading that are available in both patients and controls? We address these questions by studying a single individual, CH, who has made a partial recovery of reading familiar words following stroke. We use an fMRI analysis technique, representational similarity analysis (RSA), which allows us to decode cognitive function from distributed patterns of neural activity. Relative to controls, we find that CH shows a shift from visual to orthographic processing in contralesional regions, with a marginally significant result in perilesional regions as well. This pattern supports a contralesional reorganization of orthographic processing following stroke. More generally, these analyses demonstrate how powerful RSA can be for mapping the neural plasticity of language function.

Modality and morphology: what we write may not be what we say.

Written language is an evolutionarily recent human invention; consequently, its neural substrates cannot be determined by the genetic code. How, then, does the brain incorporate skills of this type? One possibility is that written language is dependent on evolutionarily older skills, such as spoken language; another is that dedicated substrates develop with expertise. If written language does depend on spoken language, then acquired deficits of spoken and written language should necessarily co-occur. Alternatively, if at least some substrates are dedicated to written language, such deficits may doubly dissociate. We report on 5 individuals with aphasia, documenting a double dissociation in which the production of affixes (e.g., the -ing in jumping) is disrupted in writing but not speaking or vice versa. The findings reveal that written- and spoken-language systems are considerably independent from the standpoint of morpho-orthographic operations. Understanding this independence of the orthographic system in adults has implications for the education and rehabilitation of people with written-language deficits.

The analysis of perseverations in acquired dysgraphia reveals the internal structure of orthographic representations.

At a minimum, our long-term memory representations of word spellings consist of ordered strings of single letter identities. While letter identity and position must certainly be represented, it is by no means obvious that this is the only information that is included in orthographic representations, nor that representations necessarily have a one-dimensional "flat" structure. Evidence favours the alternative hypothesis that orthographic representations, much like phonological ones, are internally rich, complex multidimensional structures, though many questions remain regarding the precise nature of the internal complexity of orthographic representations. In this investigation, we test competing accounts of the internal structure of orthographic representations by analysing the perseveration errors produced by an individual with acquired dysgraphia, L.S.S. The analysis of perseveration errors provides a novel and powerful method for investigating the question of the independence of different representational components. The results provide clear support for the hypothesis that letter quantity and syllabic role information are associated with, but separable from, letter identity information. Furthermore, the results indicate that digraphs-letter pairs associated with a single phoneme (e.g., the SH in FISH)-are units of orthographic representation. These results contribute substantially to the further development of the multidimensional hypothesis, providing both new and converging evidence regarding the nature of the internal complexity of orthographic representations.

The primacy of taxonomic semantic organization over thematic semantic organization during picture naming.

Different organizational structures have been argued to underlie semantic knowledge about concepts; taxonomic organization, based on shared features, and thematic organization based on co-occurrence in common scenes and scenarios. The goal of the current study is to examine which of the two organizational systems are more engaged in the semantic context of a picture naming task. To address this question, we examined the representational structure underlying the semantic space in different picture naming tasks by applying representational similarity analysis (RSA) to electroencephalography (EEG) datasets. In a series of experiments, EEG signals were collected while participants named pictures under different semantic contexts. Study 1 reanalyzes existing data from semantic contexts directing attention to taxonomic organization and semantic contexts that are not biased towards either taxonomic or thematic organization. In Study 2 we keep the stimuli the same and vary semantic contexts to draw attention to either taxonomic or thematic organization. The RSA approach allows us to examine the pairwise similarity in scalp-recorded amplitude patterns at each time point following the onset of the picture and relate it to theoretical taxonomic and thematic measures derived from computational models of semantics. Across all tasks, the similarity structure of scalp-recorded neural activity correlated better with taxonomic than thematic measures, in time windows associated with semantic processing. Most strikingly, we found that the scalp-recorded patterns of neural activity between taxonomically related items were more similar to each other than the scalp-recorded patterns of neural activity for thematically related or unrelated items, even in tasks that makes thematic information more salient. These results suggest that the principle semantic organization of these concepts during picture naming is taxonomic, at least in the context of picture naming.

Stuck in the past: neural events that predict intrusions from prior trials.

Neurologically intact adults perseverate in immediate serial recall, intruding items from a previous trial into the current response. We applied the electroencephalogram/event-related-potential subsequent-memory paradigm to immediate serial recall to investigate the causes of these errors. In line with previous studies using this paradigm, results revealed that words that were correctly recalled elicited a greater frontal positivity during encoding when compared with words that were either perseverated over or not produced for some other reason. More surprisingly, differences were also found at encoding between the words perseverated into the subsequent response and words that were not perseverated. These findings support a theory stating that abnormalities in both how the current target and the previous trial are processed can contribute to perseveration errors. These results inform existing theories of immediate serial recall and theories of the control of irrelevant information.

Making sense of deviance: identifying dissociating cases within the case series approach.

The case series approach in cognitive neuropsychology provides a means to test theories that make quantitative predictions about associations between different components of the cognitive system [Schwartz, M. F., & Dell, G. S. (2010). Case series investigations in cognitive neuropsychology. Cognitive Neuropsychology, 27, 477-494]. However, even when the predicted association is borne out the study may include outliers-observations that deviate significantly from the rest of the data. These outliers may reveal individual cases whose cognitive impairments dissociate from other cases included in the study. These dissociating cases can pose a significant challenge to the theory being tested. Using a recent case series that investigated the underlying causes of letter perseveration in spelling [Fischer-Baum, S., & Rapp, B. (2012). Underlying cause(s) of letter perseveration errors. Neuropsychologia, 50, 305-318], I discuss statistical and theoretical issues that arise when using outlier detection techniques to identify dissociating cases in a case series study.

Representation of letter position in single-word reading: evidence from acquired dyslexia.

Visual word recognition requires information about the positions as well as the identities of the letters in a word. This study addresses representation of letter position at prelexical levels of the word recognition process. We present evidence from an acquired dyslexic patient, L.H.D., who perseverates letters in single-word reading tasks: Far more often than expected by chance, L.H.D.'s reading responses include letters from preceding responses (e.g., SAILOR read as SAILOG immediately after FLAG was read correctly). Analyses carried out over two large data sets compared the positions of perseverated letters (e.g., the G in SAILOG) with the positions of the corresponding "source" letters (e.g., the G in FLAG). The analyses assessed the extent to which the perseverations preserved source position as defined by various theories of letter position representation. The results provided strong evidence for graded both-edges position representations, in which the position of each letter is encoded coarsely relative to both the beginning and the end of the word. Alternative position representation schemes, including letter-context and orthosyllabic schemes, were not supported.

Representational Similarity Analysis for Tracking Neural Correlates of Haptic Learning on a Multimodal Device.

A goal of wearable haptic devices has been to enable haptic communication, where individuals learn to map information typically processed visually or aurally to haptic cues via a process of cross-modal associative learning. Neural correlates have been used to evaluate haptic perception and may provide a more objective approach to assess association performance than more commonly used behavioral measures of performance. In this article, we examine Representational Similarity Analysis (RSA) of electroencephalography (EEG) as a framework to evaluate how the neural representation of multifeatured haptic cues changes with association training. We focus on the first phase of cross-modal associative learning, perception of multimodal cues. A participant learned to map phonemes to multimodal haptic cues, and EEG data were acquired before and after training to create neural representational spaces that were compared to theoretical models. Our perceptual model showed better correlations to the neural representational space before training, while the feature-based model showed better correlations with the post-training data. These results suggest that training may lead to a sharpening of the sensory response to haptic cues. Our results show promise that an EEG-RSA approach can capture a shift in the representational space of cues, as a means to track haptic learning.

Cerebellar contributions to orthographic working memory: A single case cognitive neuropsychological investigation.

Single case cognitive neuropsychological investigations involve the precise characterization of cognitive impairment at the level of an individual participant. This deep data precision affords a more fine-grained understanding of the cognitive and neural underpinnings of complex tasks, and continues to provide unique insights that inform theory in cognitive neuroscience. Here, we present a single case study of an individual, F.R., who suffered a stroke that led to chronic reading and writing problems that include an impairment to the orthographic working memory system proposed to be involved in both written language production and comprehension. Individuals who have been previously reported with a similar cognitive impairment commonly have left parietal lesions. However, F.R.'s orthographic working memory deficit resulted from damage to the right cerebellum, specifically to a region that is both structurally and functionally connected to the left parietal lobe and has been identified as part of the spelling network in previous meta-analyses of writing fMRI studies. From this lesion-symptom association, we argue that orthographic working memory is subserved by a cortical-cerebellar circuit, with damage at any point in the circuit resulting in an impairment to this function. Such a conclusion is warranted by observations from this single case approach, and we argue that these observations would likely have been missed if F.R. had been included in a larger, shallower group study. In addition to elucidating our understanding of the neural basis of spelling, this case study demonstrates the value that single case neuropsychology can continue to bring to cognitive neuroscience.

The psychological reality of picture name agreement.

Picture name agreement is commonly used as both a control variable and independent variable in studies of language production. It describes the proportion of participants who volunteer a picture's modal name in a norming study-a population-level descriptor-but researchers often assume that name agreement also indexes cognitive processes that occur within individuals. For instance, if norms show that 50% of speakers name a picture as couch, then each time a person tries to name the picture, they might have a 50% chance of selecting couch. An alternative, however, is that name agreement may simply reflect population-level sampling of more stable individual preferences (e.g., 50% of speakers prefer the name couch), continually developed through experience. One way to distinguish between these possibilities - and assess the psychological reality of name agreement - is simply to re-norm pictures with the same individuals. In Experiment 1, we therefore collected timed naming norms for a large set of line drawings from the same 25 native British English speakers twice, 1-2 weeks apart. Results show participants' name choices in Session 2 are jointly predicted by population-level name agreement, from our previous norms, and individuals' own productions in Session 1. Experiment 2 replicated this result and further showed that prior selections predicted Session 3 outcomes better than those in Session 2, in line with an incremental learning account. This is the first direct demonstration that picture name agreement has some psychological validity, but also reveals that it does not directly index within-participant lexical competition as previously assumed.

Temporal stability and representational distinctiveness: key functions of orthographic working memory.

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 working memory (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 gives 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.