Simultaneous simulations of pure, surface and phonological acquired dyslexia within a full computational model of the primary systems hypothesis.

According to the primary systems hypothesis, reading requires interactions of visual-orthographic, phonological and semantic systems. Damage to each primary system generates very different types of acquired dyslexia. Variants of the connectionist 'triangle' models of reading have been developed to investigate individual acquired dyslexia. However, only a few studies have investigated multiple acquired alexia within one framework. Importantly, there are no studies that simultaneously simulate both central dyslexia (e.g. surface and phonological dyslexia) and peripheral dyslexia (e.g. pure alexia). That is largely due to the lack of a visual component in the traditional reading models. To verify the predictions made by the primary systems hypothesis, we developed a connectionist 'deep' multi-layer triangle model of reading including visual, orthographic, phonological and semantic processing layers. We investigated whether damage to the model could produce the general behavioural patterns of impaired performance observed in patients with the corresponding reading deficits. Crucially, damage to the visual-orthographic, phonological or semantic components of the model resulted in the expected reading impairments associated with pure alexia, phonological dyslexia and surface dyslexia, respectively. The simulation results demonstrated for the first time that neurologically-impaired reading including both central and peripheral dyslexia could be addressed within a single triangle model of reading. The findings are consistent with the predictions made by the primary systems hypothesis.

Global versus phonemic similarity: Evidence in support of multi-level representation.

There is long-standing debate about the extent to which children cognitively represent words in terms of global properties or phonological segments, yet few studies have investigated how children's sensitivity to phonemic versus global similarity changes over time. The current study uses a mispronunciation-reconstruction task to measure both types of sensitivity within a cross-sectional (N = 90, aged 3;2 to 5;7) and longitudinal sample (N = 23, aged 3;2 to 5;1). The results show that children's sensitivity to phonemes increases over the first two years of school but does not reach adult levels. The findings indicate that global similarity relations remain important throughout development and support the idea of multi-level representation.

Exploring orthographic neighborhood size effects in a computational model of Chinese character naming.

Orthographic neighborhood (N) size effects have been extensively studied in English consistently producing a facilitatory effect in word naming tasks. In contrast, several recent studies on Chinese character naming have demonstrated an inhibitory effect of neighborhood size. Response latencies tend to be inhibited by inconsistent characters with large neighborhoods relative to small neighborhoods. These differences in neighborhood effects between languages may depend on the characteristics (depth) of the mapping between orthography and phonology. To explore this, we first conducted a behavioral experiment to investigate the relationship between neighborhood size, consistency and reading response. The results showed an inhibitory effect of neighborhood size for inconsistent characters but a facilitatory effect for consistent characters. We then developed two computational models based on parallel distributed processing principles to try and capture the nature of the processing that leads to these results in Chinese character naming. Simulations using models based on the triangle model of reading indicated that consistency and neighborhood size interact with the division of labor between semantics and phonology to produce these effects.

Differential Phonological and Semantic Modulation of Neurophysiological Responses to Visual Word Recognition.

BACKGROUND: Reading words for meaning relies on orthographic, phonological and semantic processing. The triangle model implicates a direct orthography-to-semantics pathway and a phonologically mediated orthography-to-semantics pathway, which interact with each other. The temporal evolution of processing in these routes is not well understood, although theoretical evidence predicts early phonological processing followed by interactive phonological and semantic processing. METHOD: This study used electroencephalography-event-related potential (ERP) analysis and magnetoencephalography (MEG) source localisation to identify temporal markers and the corresponding neural generators of these processes in early (∼200 ms) and late (∼400 ms) neurophysiological responses to visual words, pseudowords and consonant strings. RESULTS: ERP showed an effect of phonology but not semantics in both time windows, although at ∼400 ms there was an effect of stimulus familiarity. Phonological processing at ~200 ms was localised to the left occipitotemporal cortex and the inferior frontal gyrus. At 400 ms, there was continued phonological processing in the inferior frontal gyrus and additional semantic processing in the anterior temporal cortex. There was also an area in the left temporoparietal junction which was implicated in both phonological and semantic processing. In ERP, the semantic response at ∼400 ms appeared to be masked by concurrent processes relating to familiarity, while MEG successfully differentiated these processes. DISCUSSION: The results support the prediction of early phonological processing followed by an interaction of phonological and semantic processing during word recognition. Neuroanatomical loci of these processes are consistent with previous neuropsychological and functional magnetic resonance imaging studies. The results also have implications for the classical interpretation of N400-like responses as markers for semantic processing.

Dual-echo fMRI can detect activations in inferior temporal lobe during intelligible speech comprehension.

The neural basis of speech comprehension has been investigated intensively during the past few decades. Incoming auditory signals are analysed for speech-like patterns and meaningful information can be extracted by mapping these sounds onto stored semantic representations. Investigation into the neural basis of speech comprehension has largely focused on the temporal lobe, in particular the superior and posterior regions. The ventral anterior temporal lobe (vATL), which includes the inferior temporal gyrus (ITG) and temporal fusiform gyrus (TFG) is consistently omitted in fMRI studies. In contrast, PET studies have shown the involvement of these ventral temporal regions. One crucial factor is the signal loss experienced using conventional echo planar imaging (EPI) for fMRI, at tissue interfaces such as the vATL. One method to overcome this signal loss is to employ a dual-echo EPI technique. The aim of this study was to use intelligible and unintelligible (spectrally rotated) sentences to determine if the vATL could be detected during a passive speech comprehension task using a dual-echo acquisition. A whole brain analysis for an intelligibility contrast showed bilateral superior temporal lobe activations and a cluster of activation within the left vATL. Converging evidence implicates the same ventral temporal regions during semantic processing tasks, which include language processing. The specific role of the ventral temporal region during intelligible speech processing cannot be determined from this data alone, but the converging evidence from PET, MEG, TMS and neuropsychology strongly suggest that it contains the stored semantic representations, which are activated by the speech decoding process.

Mismatch negativity (MMN) reveals inefficient auditory ventral stream function in chronic auditory comprehension impairments.

BACKGROUND: Auditory discrimination is significantly impaired in Wernicke's aphasia (WA) and thought to be causatively related to the language comprehension impairment which characterises the condition. This study used mismatch negativity (MMN) to investigate the neural responses corresponding to successful and impaired auditory discrimination in WA. METHODS: Behavioural auditory discrimination thresholds of consonant-vowel-consonant (CVC) syllables and pure tones (PTs) were measured in WA (n = 7) and control (n = 7) participants. Threshold results were used to develop multiple deviant MMN oddball paradigms containing deviants which were either perceptibly or non-perceptibly different from the standard stimuli. MMN analysis investigated differences associated with group, condition and perceptibility as well as the relationship between MMN responses and comprehension (within which behavioural auditory discrimination profiles were examined). RESULTS: MMN waveforms were observable to both perceptible and non-perceptible auditory changes. Perceptibility was only distinguished by MMN amplitude in the PT condition. The WA group could be distinguished from controls by an increase in MMN response latency to CVC stimuli change. Correlation analyses displayed a relationship between behavioural CVC discrimination and MMN amplitude in the control group, where greater amplitude corresponded to better discrimination. The WA group displayed the inverse effect; both discrimination accuracy and auditory comprehension scores were reduced with increased MMN amplitude. In the WA group, a further correlation was observed between the lateralisation of MMN response and CVC discrimination accuracy; the greater the bilateral involvement the better the discrimination accuracy. CONCLUSIONS: The results from this study provide further evidence for the nature of auditory comprehension impairment in WA and indicate that the auditory discrimination deficit is grounded in a reduced ability to engage in efficient hierarchical processing and the construction of invariant auditory objects. Correlation results suggest that people with chronic WA may rely on an inefficient, noisy right hemisphere auditory stream when attempting to process speech stimuli.

A comparison of dual gradient-echo and spin-echo fMRI of the inferior temporal lobe.

Magnetic susceptibility differences at tissue interfaces lead to signal loss in conventional gradient-echo (GE) EPI. This poses a problem for fMRI in language and memory paradigms, which activate the most affected regions. Two methods proposed to overcome this are spin-echo EPI and dual GE EPI, where two EPI read-outs are serially collected at a short and longer echo time. The spin-echo method applies a refocusing pulse to recover dephased MR signal due to static field inhomogeneities, but is known to have a relatively low blood oxygenation level dependant (BOLD) sensitivity. In comparison, GE has superior BOLD sensitivity, and by employing an additional shorter echo, in a dual GE sequence, it can reduce signal loss due to spin dephasing. We directly compared dual GE and spin-echo fMRI during a semantic categorization task, which has been shown to activate the inferior temporal region-a region known to be affected by magnetic susceptibility. A whole brain analysis showed that the dual GE resulted in significantly higher activation within the left inferior temporal fusiform (ITF) cortex, compared to spin-echo. The inferior frontal gyrus (IFG) was activated for dual GE, but not spin-echo. Regions of interest analysis was carried out on the anterior and posterior ITF, left and right IFG, and part of the cerebellum. Dual GE outperformed spin-echo in the anterior and posterior ITF and bilateral IFG regions, whilst being equal in the cerebellum. Hence, dual GE should be the method of choice for fMRI studies of inferior temporal regions.

Reconstructing coherent networks from electroencephalography and magnetoencephalography with reduced contamination from volume conduction or magnetic field spread.

Volume conduction (VC) and magnetic field spread (MFS) induce spurious correlations between EEG/MEG sensors, such that the estimation of functional networks from scalp recordings is inaccurate. Imaginary coherency [1] reduces VC/MFS artefacts between sensors by assuming that instantaneous interactions are caused predominantly by VC/MFS and do not contribute to the imaginary part of the cross-spectral densities (CSDs). We propose an adaptation of the dynamic imaging of coherent sources (DICS) [2] - a method for reconstructing the CSDs between sources, and subsequently inferring functional connectivity based on coherences between those sources. Firstly, we reformulate the principle of imaginary coherency by performing an eigenvector decomposition of the imaginary part of the CSD to estimate the power that only contributes to the non-zero phase-lagged (NZPL) interactions. Secondly, we construct an NZPL-optimised spatial filter with two a priori assumptions: (1) that only NZPL interactions exist at the source level and (2) the NZPL CSD at the sensor level is a good approximation of the projected source NZPL CSDs. We compare the performance of the NZPL method to the standard method by reconstructing a coherent network from simulated EEG/MEG recordings. We demonstrate that, as long as there are phase differences between the sources, the NZPL method reliably detects the underlying networks from EEG and MEG. We show that the method is also robust to very small phase lags, noise from phase jitter, and is less sensitive to regularisation parameters. The method is applied to a human dataset to infer parts of a coherent network underpinning face recognition.

Why bilateral damage is worse than unilateral damage to the brain.

Human and animal lesion studies have shown that behavior can be catastrophically impaired after bilateral lesions but that unilateral damage often produces little or no effect, even controlling for lesion extent. This pattern is found across many different sensory, motor, and memory domains. Despite these findings, there has been no systematic, computational explanation. We found that the same striking difference between unilateral and bilateral damage emerged in a distributed, recurrent attractor neural network. The difference persists in simple feedforward networks, where it can be understood in explicit quantitative terms. In essence, damage both distorts and reduces the magnitude of relevant activity in each hemisphere. Unilateral damage reduces the relative magnitude of the contribution to performance of the damaged side, allowing the intact side to dominate performance. In contrast, balanced bilateral damage distorts representations on both sides, which contribute equally, resulting in degraded performance. The model's ability to account for relevant patient data suggests that mechanisms similar to those in the model may operate in the brain.

Modelling normal and impaired letter recognition: implications for understanding pure alexic reading.

Letter recognition is the foundation of the human reading system. Despite this, it tends to receive little attention in computational modelling of single word reading. Here we present a model that can be trained to recognise letters in various spatial transformations. When presented with degraded stimuli the model makes letter confusion errors that correlate with human confusability data. Analyses of the internal representations of the model suggest that a small set of learned visual feature detectors support the recognition of both upper case and lower case letters in various fonts and transformations. We postulated that a damaged version of the model might be expected to act in a similar manner to patients suffering from pure alexia. Summed error score generated from the model was found to be a very good predictor of the reading times of pure alexic patients, outperforming simple word length, and accounting for 47% of the variance. These findings are consistent with a hypothesis suggesting that impaired visual processing is a key to understanding the strong word-length effects found in pure alexic patients.

"Serial" effects in parallel models of reading.

There is now considerable evidence showing that the time to read a word out loud is influenced by an interaction between orthographic length and lexicality. Given that length effects are interpreted by advocates of dual-route models as evidence of serial processing this would seem to pose a serious challenge to models of single word reading which postulate a common parallel processing mechanism for reading both words and nonwords (Coltheart, Rastle, Perry, Langdon, & Ziegler, 2001; Rastle, Havelka, Wydell, Coltheart, & Besner, 2009). However, an alternative explanation of these data is that visual processes outside the scope of existing parallel models are responsible for generating the word-length related phenomena (Seidenberg & Plaut, 1998). Here we demonstrate that a parallel model of single word reading can account for the differential word-length effects found in the naming latencies of words and nonwords, provided that it includes a mapping from visual to orthographic representations, and that the nature of those orthographic representations are not preconstrained. The model can also simulate other supposedly "serial" effects. The overall findings were consistent with the view that visual processing contributes substantially to the word-length effects in normal reading and provided evidence to support the single-route theory which assumes words and nonwords are processed in parallel by a common mechanism.

The role of plasticity-related functional reorganization in the explanation of central dyslexias.

This investigation explored the hypothesis that patterns of acquired dyslexia may reflect, in part, plasticity-driven relearning that dynamically alters the division of labour (DOL) between the direct, orthography → phonology (O → P) pathway and the semantically mediated, orthography → semantics → phonology (O → S → P) pathway. Three simulations were conducted using a variant of the triangle model of reading. The model demonstrated core characteristics of normal reading behaviour in its undamaged state. When damage was followed by reoptimization (mimicking spontaneous recovery), the model reproduced the deficits observed in the central dyslexias-acute phonological damage combined with recovery matched data taken from a series of 12 phonological dyslexic patients-whilst progressive semantic damage interspersed with recovery reproduced data taken from 100 observations of semantic dementia patients. The severely phonologically damaged model also produced symptoms of deep dyslexia (imageability effects, production of semantic and mixed semantic/visual errors). In all cases, the DOL changed significantly in the recovery period, suggesting that postmorbid functional reorganization is important in understanding behaviour in chronic-stage patients.

Solving the paradox of the equipotential and modular brain: a neurocomputational model of stroke vs. slow-growing glioma.

In acute brain damage (e.g., stroke), patients can be left with specific deficits while other domains are unaffected, consistent with the classical 'modular' view of cortical organization. On this view, relearning of impaired function is limited because the remaining brain regions, tuned to other domains, have minimal capacity to assimilate an alternative activity. A clear paradox arises in low-grade glioma where an even greater amount of cortex may be affected and resected without impairment. Using a neurocomputational model we account for the modular nature of normal function as well as the contrasting types of brain insult through the interaction of three computational principles: patterns of connectivity; experience-dependent plasticity; and the time course of damage. This work provides support for a neo-Lashleyan view of cortical organization.

Using parallel distributed processing models to simulate phonological dyslexia: the key role of plasticity-related recovery.

PMSP96 [Plaut, D. C., McClelland, J. L., Seidenberg, M. S., & Patterson, K. Understanding normal and impaired word reading: Computational principles in quasi-regular domains. Psychological Review, 103, 56-115, 1996, Simulation 4] is an implementation of the triangle model of reading, which was able to simulate effects found in normal and surface dyslexic readers. This study replicated the original findings and explored the possibility that damage to the phonological portion of the model might produce symptoms of phonological dyslexia. The first simulation demonstrated that this implementation of PMSP96 was able to reproduce the standard effects of reading, and that when damaged by removal of the semantic input to phonology, it produced the kind of frequency/consistency interactions and regularization errors typical of surface dyslexia. The second simulation explored the effect of phonological damage. Phonological damage alone did not result in a convincing simulation of phonological dyslexia. However, when the damage was followed by a period of recovery, the network was able to simulate large lexicality and imageability effects characteristic of phonological dyslexia-the first time that both surface and phonological dyslexia have been simulated in the same parallel distributed processing network. This result supports the view that plasticity-related changes should be a significant factor in our understanding of chronic behavioral dissociations.

Exploring the impact of plasticity-related recovery after brain damage in a connectionist model of single-word reading.

The effect of retraining a damaged connectionist model of single-word reading was investigated with the aim of establishing whether plasticity-related changes occurring during the recovery process can contribute to our understanding of the pattern of dissociations found in brain-damaged patients. In particular, we sought to reproduce the strong frequency x consistency interactions found in surface dyslexia. A replication of Plaut, McClelland, Seidenberg, and Patterson's (1996) model of word reading was damaged and then retrained, using a standard backpropagation algorithm. Immediately after damage, there was only a small frequency x consistency interaction. Retraining the damaged model crystallized out these small differences into a strong dissociation, very similar to the pattern found in surface dyslexic patients. What is more, the percentage of regularization errors, always high in surface dyslexics, increased greatly over the retraining period, moving from under 10% to over 80% in some simulations. These results suggest that the performance patterns of brain-damaged patients can owe as much to the substantial changes in the pattern of connectivity occurring during recovery as to the original premorbid structure. This finding is discussed in relation to the traditional cognitive neuropsychological assumptions of subtractivity and transparency.