Are imaging and lesioning convergent methods for assessing functional specialisation?: investigations using an artificial neural network

This article presents an investigation of the relationship between lesioning and neuroimaging methods of assessing functional specialisation, using synthetic brain imaging (SBI) and lesioning of a connectionist network of past-tense formation. The model comprised two processing ‘routes’: one was a direct route between layers of input and output units, while the other, indirect, route featured an intermediate layer of processing units. Emergent specialisation within the network was assessed (1) by lesioning either the direct or indirect route and measuring past-tense performance for regular and irregular verbs, and (2) by measuring functional activation in each route when processing each verb type (SBI). SBI and lesioning approaches failed to converge when network activation was summed over each route in our SBI approach. Examination of individual network solutions suggested that the verb types might be using the indirect route differently in terms of the pattern of activation across the route, rather than in terms of gross activation. A subsequent SBI analysis compared patterns of activation in the indirect route and confirmed that these patterns were more similar between regular-type verbs than between regular and irregular verbs. As the spatial and temporal resolution of neuroimaging techniques improves, the results of this investigation suggest that the key to finding functional specialisation will be to distinguish local coding differences across behaviours that are the results of developmental processes. Other analyses suggest that lesioning data may be limited because, with increasing damage, they reveal the resting activations of a computational system rather than a computational specialisation per se.     

A neuroconstructivist model of past tense development and processing

We present a neural network model of learning and processing the English past tense that is based on the notion that experience-dependent cortical development is a core aspect of cognitive development. During learning the model adds and removes units and connections to develop a task-specific final architecture. The model provides an integrated account of characteristic errors during learning the past tense, adult generalization to pseudoverbs, and dissociations between verbs observed after brain damage in aphasic patients. We put forward a theory of verb inflection in which a functional processing architecture develops through interactions between experience-dependent brain development and the structure of the environment, in this case, the statistical properties of verbs in the language. The outcome of this process is a structured processing system giving rise to graded dissociations between verbs that are easy and verbs that are hard to learn and process. In contrast to dual-mechanism accounts of inflection, we argue that describing dissociations as a dichotomy between regular and irregular verbs is a post hoc abstraction and is not linked to underlying processing mechanisms. We extend current single-mechanism accounts of inflection by highlighting the role of structural adaptation in development and in the formation of the adult processing system. In contrast to some single-mechanism accounts, we argue that the link between irregular inflection and verb semantics is not causal and that existing data can be explained on the basis of phonological representations alone. This work highlights the benefit of taking brain development seriously in theories of cognitive development.     

German inflection: single route or dual route?

The German plural system has recently become a focal point for conflicting theories of language, both linguistic and cognitive. Marcus et al. (1995) highlight the German plural as support for the dual-route account of inflectional morphology first proposed by Pinker and colleagues (Pinker & Prince, 1988). On the dual-route account, inflectional morphology is universally subserved by a symbolic rule route which deals with regular inflection and an associative memory component which deals with irregular inflection. This contrasts with single-route connectionist systems. We seek to counter supposed evidence for the dual-route account through large-scale simulations as well as through experimental data. We argue that, in its current form, the dual-route account is incapable of generating experimental data provided by Marcus et al. (1995) as support. Finally, we provide direct quantitative comparisons between single-route and dual-route models of German plural inflection and find single-route performance superior on these tests.     

Double dissociations and neurophysiological expectations

Double dissociations are among the most highly regarded and influential evidence for particular modular divisions of cognitive performance. We present here a method for obtaining double dissociations across any task, even within an avowedly indivisible and unmodular system. We argue that the existence of double dissociations is not necessarily evidential and suggest what sort of additional evidence would be necessary to support the desired conclusions. We suggest that the idea of expected behavior after injury is important and should receive more attention.     

Representations in Distributed Cognitive Tasks

In this article we propose a theoretical framework of distributed representations and a methodology of representational analysis for the study of distributed cognitive tasks—tasks that require the processing of information distributed across the internal mind and the external environment. The basic principle of distributed representations Is that the representational system of a distributed cognitive task is a set of internal and external representations, which together represent the abstract structure of the task. The basic strategy of representational analysis is to decompose the representation of a hierarchical task into its component levels so that the representational properties at each level can be independently examined. The theoretical framework and the methodology are used to analyze the hierarchical structure of the Tower of Hanoi problem. Based on this analysis, four experiments are designed to examine the representational properties of the Tower of Hanoi. Finally, the nature of external representations is discussed.     

Basic Information Processing Abilities at 11 years Account for Deficits in IQ Associated with Preterm Birth

Although it is well established that preterms as a group do poorly relative to their full-term peers on tests of global cognitive functioning, the basis for this relative deficiency is less understood. The present paper examines preterm deficits in core cognitive abilities and determines their role in mediating preterm/full-term differences in IQ. The performance of 11-year-old children born preterm (birth weight < 1750 g) and their full-term controls were compared on a large battery of 15 tasks, covering four basic cognitive domains — memory, attention, speed of processing and representational competence. The validity of these four domains was established using latent variables and confirmatory factor analysis (CFA). Preterms showed pervasive deficits within and across domains. Additionally, preterm deficits in IQ were completely mediated by these four cognitive domains in a structural equation model involving a cascade from elementary abilities (attention and speed), to more complex abilities (memory and representational competence), to IQ. The similarity of findings to those obtained with this cohort in infancy and toddlerhood suggests that preterm deficits persist — across time, across task, and from the non-verbal to the verbal period.     

Is there an advantage for recognizing multi-modal emotional stimuli?

Emotions can be recognized whether conveyed by facial expressions, linguistic cues (semantics), or prosody (voice tone). However, few studies have empirically documented the extent to which multi-modal emotion perception differs from uni-modal emotion perception. Here, we tested whether emotion recognition is more accurate for multi-modal stimuli by presenting stimuli with different combinations of facial, semantic, and prosodic cues. Participants judged the emotion conveyed by short utterances in six channel conditions. Results indicated that emotion recognition is significantly better in response to multi-modal versus uni-modal stimuli. When stimuli contained only one emotional channel, recognition tended to be higher in the visual modality (i.e., facial expressions, semantic information conveyed by text) than in the auditory modality (prosody), although this pattern was not uniform across emotion categories. The advantage for multi-modal recognition may reflect the automatic integration of congruent emotional information across channels which enhances the accessibility of emotion-related knowledge in memory.     

The role of meaning in inflection: why the past tense does not require a rule

How do we produce the past tenses of verbs? For the last 20 years this question has been the focal domain for conflicting theories of language, knowledge representation, and cognitive processing. On one side of the debate have been similarity-based or single-route approaches that propose that all past tenses are formed simply through phonological analogies to existing past tenses stored in memory. On the other side of the debate are rule-based or dual-route approaches which agree that phonological analogy is important for producing irregular past tenses (e.g., think-->thought), but argue that regular past tenses (e.g., walk-->walked) are generated via a +ed rule and that a principled account of regular inflection can only be given by recourse to explicit rules. This debate has become a crucial battleground for arguments concerning the necessity and importance of abstract mental rules, embracing not only language processing, but also the of nature cognition itself. However, in centering on the roles of phonological similarity and rules, the past tense debate has largely ignored the possible role of semantics in determining inflection. This paper presents five studies that demonstrate a striking and decisive role of semantic similarity in inflection. In fact, semantic factors appear to be more important in inflection than the grammatical considerations put forward by the dual-route account. Further, these new findings provide a new way of discriminating between the claims of single-route (similarity-based) and dual-route (rule-based) approaches. It appears that inflection is carried out through analogical reminding based on semantic and phonological similarity and that a rule-based route is not necessary to account for past tense inflection.     

Parallel processing across neural systems: implications for a multiple memory system hypothesis

A common conceptualization of the organization of memory systems in brain is that different types of memory are mediated by distinct neural systems. Strong support for this view comes from studies that show double (or triple) dissociations between spatial, response, and emotional memories following selective lesions of hippocampus, striatum, and the amygdala. Here, we examine the extent to which hippocampal and striatal neural activity patterns support the multiple memory systems view. A comparison is made between hippocampal and striatal neural correlates with behavior during asymptotic performance of spatial and response maze tasks. Location- (or place), movement, and reward-specific firing patterns were found in both structures regardless of the task demands. Many, but not all, place fields of hippocampal and striatal neurons were similarly affected by changes in the visual and reward context regardless of the cognitive demands. Also, many, but not all, hippocampal and striatal movement-sensitive neurons showed significant changes in their behavioral correlates after a change in visual context, irrespective of cognitive strategy. Similar partial reorganization was observed following manipulations of the reward condition for cells recorded from both structures, again regardless of task. Assuming that representations that persist across context changes reflect learned information, we make the following conclusions. First, the consistent pattern of partial reorganization supports a view that the analysis of spatial, response, and reinforcement information is accomplished via an error-driven, or match-mismatch, algorithm across neural systems. Second, task-relevant processing occurs continuously within hippocampus and striatum regardless of the cognitive demands of the task. Third, given the high degree of parallel processing across allegedly different memory systems, we propose that different neural systems may effectively compete for control of a behavioral expression system. The strength of the influence of any one neural system on behavioral output is likely modulated by factors such as motivation, experience, or hormone status.     

Grasping objects by their handles: a necessary interaction between cognition and action

Research has illustrated dissociations between "cognitive" and "action" systems, suggesting that different representations may underlie phenomenal experience and visuomotor behavior. However, these systems also interact. The present studies show a necessary interaction when semantic processing of an object is required for an appropriate action. Experiment 1 demonstrated that a semantic task interfered with grasping objects appropriately by their handles, but a visuospatial task did not. Experiment 2 assessed performance on a visuomotor task that had no semantic component and showed a reversal of the effects of the concurrent tasks. In Experiment 3, variations on concurrent word tasks suggested that retrieval of semantic information was necessary for appropriate grasping. In all, without semantic processing, the visuomotor system can direct the effective grasp of an object, but not in a manner that is appropriate for its use.     

Cortical integration of audio–visual speech and non-speech stimuli

Using fMRI we investigated the neural basis of audio–visual processing of speech and non-speech stimuli using physically similar auditory stimuli (speech and sinusoidal tones) and visual stimuli (animated circles and ellipses). Relative to uni-modal stimuli, the different multi-modal stimuli showed increased activation in largely non-overlapping areas. Ellipse-Speech, which most resembles naturalistic audio–visual speech, showed higher activation in the right inferior frontal gyrus, fusiform gyri, left posterior superior temporal sulcus, and lateral occipital cortex. Circle-Tone, an arbitrary audio–visual pairing with no speech association, activated middle temporal gyri and lateral occipital cortex. Circle-Speech showed activation in lateral occipital cortex, and Ellipse-Tone did not show increased activation relative to uni-modal stimuli. Further analysis revealed that middle temporal regions, although identified as multi-modal only in the Circle-Tone condition, were more strongly active to Ellipse-Speech or Circle-Speech, but regions that were identified as multi-modal for Ellipse-Speech were always strongest for Ellipse-Speech. Our results suggest that combinations of auditory and visual stimuli may together be processed by different cortical networks, depending on the extent to which multi-modal speech or non-speech percepts are evoked.     

Representational and executive selection resources in 'theory of mind': evidence from compromised belief-desire reasoning in old age

Effective belief-desire reasoning requires both specialized representational capacities-the capacity to represent the mental states as such-as well as executive selection processes for accurate performance on tasks requiring the prediction and explanation of the actions of social agents. Compromised belief-desire reasoning in a given population may reflect failures in either or both of these systems. We report evidence supporting this two-process model from belief-desire reasoning tasks conducted with younger and older adult populations. When task inferential complexity is held constant, neither group showed specific difficulty with reasoning about mental state content as compared with non-mental state content. However, manipulations that systematically increase executive performance demands within belief-desire reasoning caused systematic decreases in task performance in both older and younger adult groups. Moreover, the effect of increasing executive demands was disproportionately greater in the older group. Regression analysis indicated that measures of processing speed and inhibition contributed most to explaining variance in accuracy and response times in the belief-desire reasoning tasks. These results are consistent with the idea that compromised belief-desire reasoning in old age is likely the result of age-related decline in executive selection skills that supplement core mental state representational abilities, rather than as a result of failures in the representational system itself.     

Insight without awareness: On the interaction of verbalization, instruction and practice in a simulated process control task

Four experiments in which subjects learned to control two versions of a complex simulated process control task show that verbalizable knowledge of procedures used to perform these tasks is very limited and is acquired late in learning. Individual learning curves associated with these tasks showed sudden improvements in performance, which were not accompanied by a similar increase in verbalizable knowledge. It was also found that verbal instructions consisting of exemplar memorization, strategies for rule induction, simple heuristics, and experts' instructions were all effective in enhancing novice subjects' performance. A theoretical framework is proposed in which subjects draw on two separate but interacting knowledge structures to perform these tasks. One knowledge structure is based on memory for past experiences (close analogies), and the other is based on one's current mental model of the task. Implicit sets of competing rules that control response selection are derived from both sources of knowledge. It is suggested that dissociations between task performance and verbalizing occur because memory-based processing tends to have more control over response selection because of its greater specificity, whereas a mental model tends to be the preferred mode for verbal reporting because of its greater accessibility.     

Why do children learn to say "Broke"? A model of learning the past tense without feedback

Learning the English past tense is characterized by a U-shaped learning function for the irregular verbs. Existing cognitive models often rely on a sudden increase in vocabulary, a high token-frequency of regular verbs, and complicated schemes of feedback in order to model this phenomenon. All these assumptions are at odds with empirical data. In this paper a hybrid ACT-R model is presented that shows U-shaped learning without direct feedback, changes in vocabulary, or unrealistically high rates of regular verbs. The model is capable of learning the default rule, even if regular forms are infrequent. It can also help explore the question of why there is a distinction between regular and irregular verbs in the first place, by examining the costs and benefits of both types of verbs.     

On the processing of regular and irregular forms of verbs and nouns: evidence from neuropsychology

Following acquired brain damage, a native English speaking patient (AW) encountered problems accessing phonology in speech production, while her ability to access word meaning appeared to be intact. In a series of tasks, AW was presented either with a verb, and was asked to produce its past tense or past participle (walk --> "walked"), or with a noun, and was asked to produce its plural (glove --> "gloves"). A stark dissociation was found: while AW responded accurately with regular forms of verbs (walked) and nouns (gloves), performance was significantly less accurate with irregular forms (found; children). The appearance of a selective deficit for irregular forms in conditions of impaired lexical access is in line with dual-mechanism accounts, which proposes that irregular forms are specified in the lexicon whereas regular forms are computed via rule-based mechanisms. In contrast, AW's data are problematic for connectionist accounts that do not posit separate mechanisms for processing regular and irregular forms, including the connectionist model recently proposed by Joanisse and Seidenberg (Proceedings of the National Academy of Sciences USA 96 (1999) 7592) which successfully simulated a variety of earlier neuropsychological findings. Analyses of AW's responses shed light on further details of the representation and processing of regular and irregular inflected forms.     

Is categorization performance really intact in amnesia? A meta-analysis

Most published studies of category learning in amnesia have reported intact categorization performance. These results have been used to challenge single-system accounts of categorization and recognition, in which a single representational system mediates performance in these two tasks. Many of the published studies, however, have shown a numerical advantage for controls over amnesics and often have had low statistical power. A meta-analysis was conducted to assess whether this numerical advantage is significant when the data are pooled across studies. This analysis indicates that amnesic subjects do, in fact, show deficits in categorization tasks, which is consistent with single-system exemplar model predictions.     

Dissociations in performance on novel versus irregular items: single-route demonstrations with input gain in localist and distributed models

Four pairs of connectionist simulations are presented in which quasi-regular mappings are computed using localist and distributed representations. In each simulation, a control parameter termed input gain was modulated over the only level of representation that mapped inputs to outputs. Input gain caused both localist and distributed models to shift between regularity-based and item-based modes of processing. Performance on irregular items was selectively impaired in the regularity-based modes, whereas performance on novel items was selectively impaired in the item-based modes. Thus, the models exhibited double dissociations without separable processing components. These results are discussed in the context of analogous dissociations found in language domains such as word reading and inflectional morphology.     

An encoding-complex approach to numerical cognition in Chinese-English bilinguals.

We present a model of the cognitive architecture of basic numerical skills in adult Chinese-English bilinguals. The model is based on data reported by Campbell, Kanz, and Xue (1999) and combines Dehaene and Cohen's triple-code theory with Campbell and Clark's encoding-complex approach to modeling number processing. Participants were required to name, add or multiply Arabic or Mandarin numerals and to respond in English or Chinese. They also performed magnitude comparisons on pairs of Arabic or Mandarin numerals. The proposed model of their performance on this set of tasks assumes 1) that number processing is modular with respect to representational code (e.g., visual, visuo-spatial, verbal) rather than with respect to numerical function, 2) task-specific communication between representational codes is interactive rather than additive, and 3) memory for arithmetic facts is at least partially language-based and our Chinese-English bilinguals possessed both Chinese and English-language number-fact representations. We provide new analyses of the arithmetic data and a review of research on the role of language in simple arithmetic to substantiate our claims about linguistic codes for number-fact memory.     

Cognitive Load in Cross‐Modal Dual‐Task Processing

Unitary‐resource models posit a collective attentional resource responsible for sustaining information processing across all modalities. In contrast, multiple‐resource models presume a system of modality‐independent resources. The two models predict similar within‐modal performance, but seemingly contradictory cross‐modal performance. Discordant outcomes in past research are here hypothesized to result from inconsistent levels of cognitive load imposed within experimental tasks. Participants monitored two simultaneously presented alphabetic letter arrays consisting of the serial presentation of a target and multiple distractors. The dual‐task design required the identification of targets in within‐modal and cross‐modal trials. Cognitive load was manipulated by altering the presentation rate of the letters. While cross‐modal performance was generally superior and cognitive load was associated with decreasing performance, the cross‐modal advantage prominent at low levels of cognitive load was absent under high cognitive load. Therefore, neither resource model can alone account for dual‐task processing, suggesting the role of a hierarchical resource model. Copyright © 2015 John Wiley & Sons, Ltd.     

A model‐based approach to limb apraxia in Alzheimer's disease

Limb apraxia is a neurological deficit characterized by an inability to pantomime and/or imitate gestures, which can result from neurodegenerative disorders such as Alzheimer's disease (AD). The major goal of the study was to describe comprehensively the apraxia deficits observed in AD patients and to relate those deficits to general cognitive status, measures of daily activity, and other neuropsychological measures. Limb apraxia was assessed on a variety of conceptual and gesture production tasks in 30 AD patients. As a group, AD patients were impaired across gesture production tasks: of note was the greater impairment in imitation, as opposed to pantomime, which was especially pronounced when patients were imitating with a delay. Imitation performance was best predicted by measures of visuospatial processing, while imitation with delay was best predicted by measures of working memory. In addition, pantomime in response to pictures of tools was less accurate than Pantomime to Verbal Command and holding the tool during performance did not decrease the participants' impairment, while introducing a verbal cue during imitation increased the severity of deficits. Furthermore, investigation into patterns of deficits clearly demonstrated that the nature of limb apraxia deficits observed in AD can be quite heterogeneous and that dissociations between the conceptual and the production system exist. Finally, we also report on significant correlations between general cognitive status and limb apraxia.