Distinct Effects of Memory Retrieval and Articulatory Preparation when Learning and Accessing New Word Forms

Temporal and frontal activations have been implicated in learning of novel word forms, but their specific roles remain poorly understood. The present magnetoencephalography (MEG) study examines the roles of these areas in processing newly-established word form representations. The cortical effects related to acquiring new phonological word forms during incidental learning were localized. Participants listened to and repeated back new word form stimuli that adhered to native phonology (Finnish pseudowords) or were foreign (Korean words), with a subset of the stimuli recurring four times. Subsequently, a modified 1-back task and a recognition task addressed whether the activations modulated by learning were related to planning for overt articulation, while parametrically added noise probed reliance on developing memory representations during effortful perception. Learning resulted in decreased left superior temporal and increased bilateral frontal premotor activation for familiar compared to new items. The left temporal learning effect persisted in all tasks and was strongest when stimuli were embedded in intermediate noise. In the noisy conditions, native phonotactics evoked overall enhanced left temporal activation. In contrast, the frontal learning effects were present only in conditions requiring overt repetition and were more pronounced for the foreign language. The results indicate a functional dissociation between temporal and frontal activations in learning new phonological word forms: the left superior temporal responses reflect activation of newly-established word-form representations, also during degraded sensory input, whereas the frontal premotor effects are related to planning for articulation and are not preserved in noise.     

The Role of Phoneme in Mandarin Chinese Production: Evidence from ERPs

Established linguistic theoretical frameworks propose that alphabetic language speakers use phonemes as phonological encoding units during speech production whereas Mandarin Chinese speakers use syllables. This framework was challenged by recent neural evidence of facilitation induced by overlapping initial phonemes, raising the possibility that phonemes also contribute to the phonological encoding process in Chinese. However, there is no evidence of non-initial phoneme involvement in Chinese phonological encoding among representative Chinese speakers, rendering the functional role of phonemes in spoken Chinese controversial. Here, we addressed this issue by systematically investigating the word-initial and non-initial phoneme repetition effect on the electrophysiological signal using a picture-naming priming task in which native Chinese speakers produced disyllabic word pairs. We found that overlapping phonemes in both the initial and non-initial position evoked more positive ERPs in the 180- to 300-ms interval, indicating position-invariant repetition facilitation effect during phonological encoding. Our findings thus revealed the fundamental role of phonemes as independent phonological encoding units in Mandarin Chinese.     

Modelling reading development through phonological decoding and self-teaching: implications for dyslexia

The most influential theory of learning to read is based on the idea that children rely on phonological decoding skills to learn novel words. According to the self-teaching hypothesis, each successful decoding encounter with an unfamiliar word provides an opportunity to acquire word-specific orthographic information that is the foundation of skilled word recognition. Therefore, phonological decoding acts as a self-teaching mechanism or ‘built-in teacher’. However, all previous connectionist models have learned the task of reading aloud through exposure to a very large corpus of spelling–sound pairs, where an ‘external’ teacher supplies the pronunciation of all words that should be learnt. Such a supervised training regimen is highly implausible. Here, we implement and test the developmentally plausible phonological decoding self-teaching hypothesis in the context of the connectionist dual process model. In a series of simulations, we provide a proof of concept that this mechanism works. The model was able to acquire word-specific orthographic representations for more than 25 000 words even though it started with only a small number of grapheme–phoneme correspondences. We then show how visual and phoneme deficits that are present at the outset of reading development can cause dyslexia in the course of reading development.     

Activation of the Left Inferior Frontal Gyrus in the First 200 ms of Reading: Evidence from Magnetoencephalography (MEG)

Background

It is well established that the left inferior frontal gyrus plays a key role in the cerebral cortical network that supports reading and visual word recognition. Less clear is when in time this contribution begins. We used magnetoencephalography (MEG), which has both good spatial and excellent temporal resolution, to address this question.

Methodology/Principal Findings

MEG data were recorded during a passive viewing paradigm, chosen to emphasize the stimulus-driven component of the cortical response, in which right-handed participants were presented words, consonant strings, and unfamiliar faces to central vision. Time-frequency analyses showed a left-lateralized inferior frontal gyrus (pars opercularis) response to words between 100–250 ms in the beta frequency band that was significantly stronger than the response to consonant strings or faces. The left inferior frontal gyrus response to words peaked at ∼130 ms. This response was significantly later in time than the left middle occipital gyrus, which peaked at ∼115 ms, but not significantly different from the peak response in the left mid fusiform gyrus, which peaked at ∼140 ms, at a location coincident with the fMRI–defined visual word form area (VWFA). Significant responses were also detected to words in other parts of the reading network, including the anterior middle temporal gyrus, the left posterior middle temporal gyrus, the angular and supramarginal gyri, and the left superior temporal gyrus.

Conclusions/Significance

These findings suggest very early interactions between the vision and language domains during visual word recognition, with speech motor areas being activated at the same time as the orthographic word-form is being resolved within the fusiform gyrus. This challenges the conventional view of a temporally serial processing sequence for visual word recognition in which letter forms are initially decoded, interact with their phonological and semantic representations, and only then gain access to a speech code.     

Hierarchical coding of letter strings in the ventral stream: dissecting the inner organization of the visual word-form system

Visual word recognition has been proposed to rely on a hierarchy of increasingly complex neuronal detectors, from individual letters to bigrams and morphemes. We used fMRI to test whether such a hierarchy is present in the left occipitotemporal cortex, at the site of the visual word-form area, and with an anterior-to-posterior progression. We exposed adult readers to (1) false-font strings; (2) strings of infrequent letters; (3) strings of frequent letters but rare bigrams; (4) strings with frequent bigrams but rare quadrigrams; (5) strings with frequent quadrigrams; (6) real words. A gradient of selectivity was observed through the entire span of the occipitotemporal cortex, with activation becoming more selective for higher-level stimuli toward the anterior fusiform region. A similar gradient was also seen in left inferior frontoinsular cortex. Those gradients were asymmetrical in favor of the left hemisphere. We conclude that the left occipitotemporal visual word-form area, far from being a homogeneous structure, presents a high degree of functional and spatial hierarchical organization which must result from a tuning process during reading acquisition.     

Word learning,phonological short-term memory,phonotactic probability and long-term memory: towards an integrated framework

Word learning is studied in a multitude of ways, and it is often not clear what the relationship is between different phenomena. In this article, we begin by outlining a very simple functional framework that despite its simplicity can serve as a useful organizing scheme for thinking about various types of studies of word learning. We then review a number of themes that in recent years have emerged as important topics in the study of word learning, and relate them to the functional framework, noting nevertheless that these topics have tended to be somewhat separate areas of study. In the third part of the article, we describe a recent computational model and discuss how it offers a framework that can integrate and relate these various topics in word learning to each other. We conclude that issues that have typically been studied as separate topics can perhaps more fruitfully be thought of as closely integrated, with the present framework offering several suggestions about the nature of such integration.     

Pseudo-relaxation learning algorithm for complex-valued associative memory

HAM (Hopfield Associative Memory) and BAM (Bidirectinal Associative Memory) are representative associative memories by neural networks. The storage capacity by the Hebb rule, which is often used, is extremely low. In order to improve it, some learning methods, for example, pseudo-inverse matrix learning and gradient descent learning, have been introduced. Oh introduced pseudo-relaxation learning algorithm to HAM and BAM. In order to accelerate it, Hattori proposed quick learning. Noest proposed CAM (Complex-valued Associative Memory), which is complex-valued HAM. The storage capacity of CAM by the Hebb rule is also extremely low. Pseudo-inverse matrix learning and gradient descent learning have already been generalized to CAM. In this paper, we apply pseudo-relaxation learning algorithm to CAM in order to improve the capacity.     

Word learning and the cerebral hemispheres: from serial to parallel processing of written words

Reading familiar words differs from reading unfamiliar non-words in two ways. First, word reading is faster and more accurate than reading of unfamiliar non-words. Second, effects of letter length are reduced for words, particularly when they are presented in the right visual field in familiar formats. Two experiments are reported in which right-handed participants read aloud non-words presented briefly in their left and right visual fields before and after training on those items. The non-words were interleaved with familiar words in the naming tests. Before training, naming was slow and error prone, with marked effects of length in both visual fields. After training, fewer errors were made, naming was faster, and the effect of length was much reduced in the right visual field compared with the left. We propose that word learning creates orthographic word forms in the mid-fusiform gyrus of the left cerebral hemisphere. Those word forms allow words to access their phonological and semantic representations on a lexical basis. But orthographic word forms also interact with more posterior letter recognition systems in the middle/inferior occipital gyri, inducing more parallel processing of right visual field words than is possible for any left visual field stimulus, or for unfamiliar non-words presented in the right visual field.     

A complementary systems account of word learning: neural and behavioural evidence

In this paper we present a novel theory of the cognitive and neural processes by which adults learn new spoken words. This proposal builds on neurocomputational accounts of lexical processing and spoken word recognition and complementary learning systems (CLS) models of memory. We review evidence from behavioural studies of word learning that, consistent with the CLS account, show two stages of lexical acquisition: rapid initial familiarization followed by slow lexical consolidation. These stages map broadly onto two systems involved in different aspects of word learning: (i) rapid, initial acquisition supported by medial temporal and hippocampal learning, (ii) slower neocortical learning achieved by offline consolidation of previously acquired information. We review behavioural and neuroscientific evidence consistent with this account, including a meta-analysis of PET and functional Magnetic Resonance Imaging (fMRI) studies that contrast responses to spoken words and pseudowords. From this meta-analysis we derive predictions for the location and direction of cortical response changes following familiarization with pseudowords. This allows us to assess evidence for learning-induced changes that convert pseudoword responses into real word responses. Results provide unique support for the CLS account since hippocampal responses change during initial learning, whereas cortical responses to pseudowords only become word-like if overnight consolidation follows initial learning.     

Rapid Extraction of Lexical Tone Phonology in Chinese Characters: A Visual Mismatch Negativity Study

Background

In alphabetic languages, emerging evidence from behavioral and neuroimaging studies shows the rapid and automatic activation of phonological information in visual word recognition. In the mapping from orthography to phonology, unlike most alphabetic languages in which there is a natural correspondence between the visual and phonological forms, in logographic Chinese, the mapping between visual and phonological forms is rather arbitrary and depends on learning and experience. The issue of whether the phonological information is rapidly and automatically extracted in Chinese characters by the brain has not yet been thoroughly addressed.

Methodology/Principal Findings

We continuously presented Chinese characters differing in orthography and meaning to adult native Mandarin Chinese speakers to construct a constant varying visual stream. In the stream, most stimuli were homophones of Chinese characters: The phonological features embedded in these visual characters were the same, including consonants, vowels and the lexical tone. Occasionally, the rule of phonology was randomly violated by characters whose phonological features differed in the lexical tone.

Conclusions/Significance

We showed that the violation of the lexical tone phonology evoked an early, robust visual response, as revealed by whole-head electrical recordings of the visual mismatch negativity (vMMN), indicating the rapid extraction of phonological information embedded in Chinese characters. Source analysis revealed that the vMMN was involved in neural activations of the visual cortex, suggesting that the visual sensory memory is sensitive to phonological information embedded in visual words at an early processing stage.     

Task-guided selection of the dual neural pathways for reading

The visual perception of words is known to activate the auditory representation of their spoken forms automatically. We examined the neural mechanism for this phonological activation using transcranial magnetic stimulation (TMS) with a masked priming paradigm. The stimulation sites (left superior temporal gyrus [L-STG] and inferior parietal lobe [L-IPL]), modality of targets (visual and auditory), and task (pronunciation and lexical decision) were manipulated independently. For both within- and cross-modal conditions, the repetition priming during pronunciation was eliminated when TMS was applied to the L-IPL, but not when applied to the L-STG, whereas the priming during lexical decision was eliminated when the L-STG, but not the L-IPL, was stimulated. The observed double dissociation suggests that the conscious task instruction modulates the stimulus-driven activation of the lateral temporal cortex for lexico-phonological activation and the inferior parietal cortex for spoken word production, and thereby engages a different neural network for generating the appropriate behavioral response.     

Two Systems of Maintenance in Verbal Working Memory: Evidence from the Word Length Effect

The extended time-based resource-sharing (TBRS) model suggested a working memory architecture in which an executive loop and a phonological loop could both support the maintenance of verbal information. The consequence of such a framework is that phonological effects known to impact the maintenance of verbal information, like the word length effect (WLE), should depend on the use of the phonological loop, but should disappear under the maintenance by the executive loop. In two previous studies, introducing concurrent articulation in complex span tasks barely affected WLE, contradicting the prediction from the TBRS model. The present study re-evaluated the WLE in a complex span task while controlling for time parameters and the amount of concurrent articulation. Specifically, we used a computer-paced span task in which participants remembered lists of either short or long words while concurrently either articulating or making a location judgment. Whereas the WLE appeared when participants remained silent, concurrent articulation eliminated the effect. Introducing a concurrent attention demand reduced recall, but did not affect WLE, and did not interact with concurrent articulation. These results support the existence of two systems of maintenance for verbal information.     

Quantitative-trait locus for specific language and reading deficits on chromosome 6p.

Reading disability (RD), or dyslexia, is a complex cognitive disorder manifested by difficulties in learning to read, in otherwise normal individuals. Individuals with RD manifest deficits in several reading and language skills. Previous research has suggested the existence of a quantitative-trait locus (QTL) for RD on the short arm of chromosome 6. In the present study, RD subjects' performance in several measures of word recognition and component skills of orthographic coding, phonological decoding, and phoneme awareness were individually subjected to QTL analysis, with a new sample of 126 sib pairs, by means of a multipoint mapping method and eight informative DNA markers on chromosome 6 (D6S461, D6S276, D6S105, D6S306, D6S258, D6S439, D6S291, and D6S1019). The results indicate significant linkage across a distance of at least 5 cM for deficits in orthographic (LOD = 3.10) and phonological (LOD = 2.42) skills, confirming previous findings.     

Mismatch Response to Polysyllabic Nonwords: A Neurophysiological Signature of Language Learning Capacity

Background

The ability to repeat polysyllabic nonwords such as “blonterstaping” has frequently been shown to correlate with language learning ability but it is not clear why such a correlation should exist. Three alternative explanations have been offered, stated in terms of differences in: (a) perceptual ability; (b) efficiency of phonological loop functioning; (c) pre-existing vocabulary knowledge and/or articulatory skills. In the present study, we used event-related potentials to assess the contributions from these three factors to explaining individual variation in nonword repetition ability.

Methodology/Principal Findings

59 adults who were subdivided according to whether they were good or poor nonword-repeaters participated. Electrophysiologically measured mismatch responses were recorded to changes in consonants as participants passively listened to a repeating four syllable CV-string. The consonant change could occur in one of four positions along the CV-string and we predicted that: (a) if nonword repetition depended purely on auditory discrimination ability, then reduced mismatch responses to all four consonant changes would be observed in the poor nonword-repeaters, (b) if it depended on encoding or decay of information in a capacity-limited phonological store, then a position specific decrease in mismatch response would be observed, (c) if neither cognitive capacity was involved, then the two groups of participants would provide equivalent mismatch responses. Consistent with our second hypothesis, a position specific difference located on the third syllable was observed in the late discriminative negativity (LDN) window (230–630 ms post-syllable onset).

Conclusions/Significance

Our data thus confirm that people who are poorer at nonword repetition are less efficient in early processing of polysyllabic speech materials, but this impairment is not attributable to deficits in low level auditory discrimination. We conclude by discussing the significance of the observed relationship between LDN amplitude and nonword repetition ability and describe how this relatively little understood ERP component provides a biological window onto processes required for successful language learning.     

The legacy of Donald O. Hebb: more than the Hebb synapse

Neuroscientists associate the name of Donald O. Hebb with the Hebbian synapse and the Hebbian learning rule, which underlie connectionist theories and synaptic plasticity, but Hebb's work has also influenced developmental psychology, neuropsychology, perception and the study of emotions, as well as learning and memory. Here, we review the work of Hebb and its lasting influence on neuroscience in honour of the 2004 centenary of his birth.     

A dynamic over games drives selfish agents to win–win outcomes

Understanding human institutions, animal cultures and other social systems requires flexible formalisms that describe how their members change them from within. We introduce a framework for modelling how agents change the games they participate in. We contrast this between-game ‘institutional evolution’ with the more familiar within-game ‘behavioural evolution’. We model institutional change by following small numbers of persistent agents as they select and play a changing series of games. Starting from an initial game, a group of agents trace trajectories through game space by navigating to increasingly preferable games until they converge on ‘attractor’ games. Agents use their ‘institutional preferences'' for game features (such as stability, fairness and efficiency) to choose between neighbouring games. We use this framework to pose a pressing question: what kinds of games does institutional evolution select for; what is in the attractors? After computing institutional change trajectories over the two-player space, we find that attractors have disproportionately fair outcomes, even though the agents who produce them are strictly self-interested and indifferent to fairness. This seems to occur because game fairness co-occurs with the self-serving features these agents do actually prefer. We thus present institutional evolution as a mechanism for encouraging the spontaneous emergence of cooperation among small groups of inherently selfish agents, without space, reputation, repetition, or other more familiar mechanisms. Game space trajectories provide a flexible, testable formalism for modelling the interdependencies of behavioural and institutional evolutionary processes, as well as a mechanism for the evolution of cooperation.     

Word learning and lexical development across the lifespan

Word learning is one of the core components of language acquisition. In this article, we provide an overview of the theme issue on word learning, describing some of the ways in which research in the area has progressed and diverged. In recent years, word learning has become central in a wider range of research areas, and is important to research on adult, as well as child and infant language. We introduce 10 papers that cover the recent developments from a wide range of perspectives, focusing on developmental research, the influence of reading skills, neuroimaging and the relationship between word learning and general models of memory.     

Dosage suppression of the Kluyveromyces lactis zymocin by Saccharomyces cerevisiae ISR1 and UGP1

The Kluyveromyces lactis zymocin complex kills Saccharomyces cerevisiae cells in a process that involves tRNA cleavage by its tRNAse gamma-toxin subunit. In contrast to the gamma-toxin mode of action, the early steps of the zymocin response are less well characterized. Here, we present high-dosage suppressors of zymocin that encode a putative Pkc1-related kinase (ISR1) and UDP-glucose pyrophosphorylase (UGPase) (UGP1). Anti-UGPase Western blots and GAL10 - ISR1 overexpression suggest that zymocin suppression correlates with overproduction of UGPase or Isr1. As judged from protection against exo-zymocin and unaltered sensitivity to endogenous gamma-toxin, high-copy ISR1 and UGP1 operate in early, nontarget steps of the zymocin pathway. Consistent with a recent report on in vitro phosphorylation of Isr1 and UGPase by the CDK Pho85, high-copy ISR1 and UGP1 suppression of zymocin is abolished in a pho85 null mutant lacking CDK activity of Pho85. Moreover, suppression requires UGPase enzyme activity, and ISR1 overexpression also protects against CFW, a chitin-interfering poison. Our data agree with roles for UGPase in cell wall biosynthetic processes and for Isr1 in Pkc1-related cell wall integrity. In sum, high-copy ISR1 and UGP1 cells affect early steps of the zymocin response and potentially prevent the lethal K. lactis killer complex from establishing cell surface recognition and/or contact.     

Deep generative modeling for volume reconstruction in cryo-electron microscopy

Advances in cryo-electron microscopy (cryo-EM) for high-resolution imaging of biomolecules in solution have provided new challenges and opportunities for algorithm development for 3D reconstruction. Next-generation volume reconstruction algorithms that combine generative modelling with end-to-end unsupervised deep learning techniques have shown promise, but many technical and theoretical hurdles remain, especially when applied to experimental cryo-EM images. In light of the proliferation of such methods, we propose here a critical review of recent advances in the field of deep generative modelling for cryo-EM reconstruction. The present review aims to (i) provide a unified statistical framework using terminology familiar to machine learning researchers with no specific background in cryo-EM, (ii) review the current methods in this framework, and (iii) outline outstanding bottlenecks and avenues for improvements in the field.     

Phonological,grammatical, and written words in Wichi

This article aims at investigating the linguistic criteria to determine what a word is in Wichi (Matacoan), a polysynthetic and agglutinative language spoken in the Gran Chaco Region, in South America. The main phonological criteria proposed are phonological rules and stress. We also apply some grammatical criteria that have been proposed cross linguistically, some of which are useful to determine the boundaries of grammatical words in Wichi. Finally, we explore the relationship between the phonological and grammatical word with the written word. We base our analysis of written words on a textbook (Tsalanawu) used in many bilingual schools in Northeastern Argentina.