Publications

Abstract

Expectancy mechanisms are routinely used by the cognitive system in stimulus processing and in anticipation of appropriate responses. Electrophysiology research has documented negative shifts of brain activity when expectancies are violated within a local stimulus context (e.g., reading an implausible word in a sentence) or more globally between consecutive stimuli (e.g., a narrative of images with an incongruent end). In this EEG study, we examine the interaction between expectancies operating at the level of stimulus plausibility and at more global level of contextual congruency to provide evidence for, or against, a disassociation of the underlying processing mechanisms. We asked participants to verify the congruency of pairs of cross-modal stimuli (a sentence and a scene), which varied in plausibility. ANOVAs on ERP amplitudes in selected windows of interest show that congruency violation has longer-lasting (from 100 to 500 ms) and more widespread effects than plausibility violation (from 200 to 400 ms). We also observed critical interactions between these factors, whereby incongruent and implausible pairs elicited stronger negative shifts than their congruent counterpart, both early on (100–200 ms) and between 400–500 ms. Our results suggest that the integration mechanisms are sensitive to both global and local effects of expectancy in a modality independent manner. Overall, we provide novel insights into the interdependence of expectancy during meaning integration of cross-modal stimuli in a verification task

Abstract

Artificial grammar learning (AGL) has been probed with forced-choice behavioral tests (active tests). Recent attempts to probe the outcomes of learning (implicitly acquired knowledge) with eye-movement responses (passive tests) have shown null results. However, these latter studies have not tested for sensitivity effects, for example, increased eye movements on a printed violation. In this study, we tested for sensitivity effects in AGL tests with (Experiment 1) and without (Experiment 2) concurrent active tests (preference- and grammaticality classification) in an eye-tracking experiment. Eye movements discriminated between sequence types in passive tests and more so in active tests. The eye-movement profile did not differ between preference and grammaticality classification, and it resembled sensitivity effects commonly observed in natural syntax processing. Our findings show that the outcomes of implicit structured sequence learning can be characterized in eye tracking. More specifically, whole trial measures (dwell time, number of fixations) showed robust AGL effects, whereas first-pass measures (first-fixation duration) did not. Furthermore, our findings strengthen the link between artificial and natural syntax processing, and they shed light on the factors that determine performance differences in preference and grammaticality classification tests

Abstract

How can we grasp the temporal structure of events? A few studies have indicated that representations of temporal structure are acquired when there is an intention to learn, but not when learning is incidental. Response-to-stimulus intervals, uncorrelated temporal structures, unpredictable ordinal information, and lack of metrical organization have been pointed out as key obstacles to incidental temporal learning, but the literature includes piecemeal demonstrations of learning under all these circumstances. We suggest that the unacknowledged effects of ordinal load may help reconcile these conflicting findings, ordinal load referring to the cost of identifying the sequence of events (e.g., tones, locations) where a temporal pattern is embedded. In a first experiment, we manipulated ordinal load into simple and complex levels. Participants learned ordinal-simple sequences, despite their uncorrelated temporal structure and lack of metrical organization. They did not learn ordinal-complex sequences, even though there were no response-to-stimulus intervals nor unpredictable ordinal information. In a second experiment, we probed learning of ordinal-complex sequences with strong metrical organization, and again there was no learning. We conclude that ordinal load is a key obstacle to incidental temporal learning. Further analyses showed that the effect of ordinal load is to mask the expression of temporal knowledge, rather than to prevent learning.

Abstract

The suitability of the Artificial Grammar Learning (AGL) paradigm to capture relevant aspects of the acquisition of linguistic structures has been empirically tested in a number of EEG studies. Some have shown a syntax-related P600 component, but it has not been ruled out that the AGL P600 effect is a response to surface features (e.g., subsequence familiarity) rather than the underlying syntax structure. Therefore, in this study, we controlled for the surface characteristics of the test sequences (associative chunk strength) and recorded the EEG before (baseline preference classification) and
after (preference and grammaticality classification) exposure to a grammar. A typical, centroparietal P600 effect was elicited by grammatical violations after exposure, suggesting that the AGL P600 effect signals a response to structural irregularities. Moreover, preference and grammaticality classification showed a qualitatively similar ERP profile, strengthening the idea that the implicit structural mere
exposure paradigm in combination with preference classification is a suitable alternative to the traditional grammaticality classification test.

Abstract

Non-adjacent dependencies are challenging for the language learning machinery and are acquired later than adjacent dependencies. In this transcranial magnetic stimulation (TMS) study, we show that participants successfully discriminated between grammatical and non-grammatical sequences after having implicitly acquired an artificial language with crossed non-adjacent dependencies. Subsequent to transcranial magnetic stimulation of Broca’s region, discrimination was impaired compared to when a language-irrelevant control region (vertex) was stimulated. These results support the view that Broca’s region is engaged in structured sequence processing and extend previous functional neuroimaging results on artificial grammar learning (AGL) in two directions: first, the results establish that Broca’s region is a causal component in the processing of non-adjacent dependencies, and second, they show that implicit processing of non-adjacent dependencies engages Broca’s region. Since patients with lesions in Broca’s region do not always show grammatical processing difficulties, the result that Broca’s region is causally linked to processing of non-adjacent dependencies is a step towards clarification of the exact nature of syntactic deficits caused by lesions or perturbation to Broca’s region. Our findings are consistent with previous results and support a role for Broca’s region in general structured sequence processing, rather than a specific role for the processing of hierarchically organized sentence structure.

Abstract

We investigated neural mechanisms that support voice recognition in a training paradigm with fMRI. The same listeners were trained on different weeks to categorize the mid-regions of voice-morph continua as an individual's voice. Stimuli implicitly defined a voice-acoustics space, and training explicitly defined a voice-identity space. The predefined centre of the voice category was shifted from the acoustic centre each week in opposite directions, so the same stimuli had different training histories on different tests. Cortical sensitivity to voice similarity appeared over different time-scales and at different representational stages. First, there were short-term adaptation effects: Increasing acoustic similarity to the directly preceding stimulus led to haemodynamic response reduction in the middle/posterior STS and in right ventrolateral prefrontal regions. Second, there were longer-term effects: Response reduction was found in the orbital/insular cortex for stimuli that were most versus least similar to the acoustic mean of all preceding stimuli, and, in the anterior temporal pole, the deep posterior STS and the amygdala, for stimuli that were most versus least similar to the trained voice-identity category mean. These findings are interpreted as effects of neural sharpening of long-term stored typical acoustic and category-internal values. The analyses also reveal anatomically separable voice representations: one in a voice-acoustics space and one in a voice-identity space. Voice-identity representations flexibly followed the trained identity shift, and listeners with a greater identity effect were more accurate at recognizing familiar voices. Voice recognition is thus supported by neural voice spaces that are organized around flexible ‘mean voice’ representations.

Abstract

One implication of the double-deficit hypothesis for dyslexia is that there should be subtypes of dyslexic readers that exhibit rapid naming deficits with or without concomitant phonological processing problems. In the current study, we investigated the validity of this hypothesis for Portuguese orthography, which is more consistent than English orthography, by exploring different cognitive profiles in a sample of dyslexic children. In particular, we were interested in identifying readers characterized by a pure rapid automatized naming deficit. We also examined whether rapid naming and phonological awareness independently account for individual differences in reading performance. We characterized the performance of dyslexic readers and a control group of normal readers matched for age on reading, visual rapid naming and phonological processing tasks. Our results suggest that there is a subgroup of dyslexic readers with intact phonological processing capacity (in terms of both accuracy and speed measures) but poor rapid naming skills. We also provide evidence for an independent association between rapid naming and reading competence in the dyslexic sample, when the effect of phonological skills was controlled. Altogether, the results are more consistent with the view that rapid naming problems in dyslexia represent a second core deficit rather than an exclusive phonological explanation for the rapid naming deficits. Furthermore, additional non-phonological processes, which subserve rapid naming performance, contribute independently to reading development.

Abstract

To clarify whether the neural pathways concerning color processing are the same for natural objects, for artifacts objects and for non-sense objects we examined functional magnetic resonance imaging (FMRI) responses during a covert naming task including the factors color (color vs. black&white (B&W)) and stimulus type (natural vs. artifacts vs. non-sense objects). Our results indicate that the superior parietal lobule and precuneus (BA 7) bilaterally, the right hippocampus and the right fusifom gyrus (V4) make part of a network responsible for color processing both for natural and artifacts objects, but not for non-sense objects. The recognition of non-sense colored objects compared to the recognition of color objects activated the posterior cingulate/precuneus (BA 7/23/31), suggesting that color attribute induces the mental operation of trying to associate a non-sense composition with a familiar objects. When color objects (both natural and artifacts) were contrasted with color nonobjects we observed activations in the right parahippocampal gyrus (BA 35/36), the superior parietal lobule (BA 7) bilaterally, the left inferior middle temporal region (BA 20/21) and the inferior and superior frontal regions (BA 10/11/47). These additional activations suggest that colored objects recruit brain regions that are related to visual semantic information/retrieval and brain regions related to visuo-spatial processing. Overall, the results suggest that color information is an attribute that improve object recognition (based on behavioral results) and activate a specific neural network related to visual semantic information that is more extensive than for B&W objects during object recognition

Abstract

In order to clarify whether the influence of color knowledge information in object recognition depends on the presence of the appropriate surface color, we designed a name—object verification task. The relationship between color and shape information provided by the name and by the object photo was manipulated in order to assess color interference independently of shape interference. We tested three different versions for each object: typically colored, black and white, and nontypically colored. The response times on the nonmatching trials were used to measure the interference between the name and the photo. We predicted that the more similar the name and the photo are, the longer it would take to respond. Overall, the color similarity effect disappeared in the black-and-white and nontypical color conditions, suggesting that the influence of color knowledge on object recognition depends on the presence of the appropriate surface color information.

Abstract

This article briefly reviews some recent work on artificial language learning in children and adults. The final part of the article is devoted to a theoretical formulation of the language learning problem from a mechanistic neurobiological viewpoint and we show that it is logically possible to combine the notion of innate language constraints with, for example, the notion of domain general learning mechanisms. A growing body of empirical evidence suggests that the mechanisms involved in artificial language learning and in structured sequence processing are shared with those of natural language acquisition and natural language processing. Finally, by theoretically analyzing a formal learning model, we highlight Fodor’s insight that it is logically possible to combine innate, domain-specific constraints with domain-general learning mechanisms.

Abstract

Language in high-functioning autism is characterized by pragmatic and semantic deficits, and people with autism have a reduced tendency to integrate information. Because the left and right inferior frontal (LIF and RIF) regions are implicated with integration of speaker information, world knowledge, and semantic knowledge, we hypothesized that abnormal functioning of the LIF and RIF regions might contribute to pragmatic and semantic language deficits in autism. Brain activation of sixteen 12- to 18-year-old, high-functioning autistic participants was measured with functional magnetic resonance imaging during sentence comprehension and compared with that of twenty-six matched controls. The content of the pragmatic sentence was congruent or incongruent with respect to the speaker characteristics (male/female, child/adult, and upper class/lower class). The semantic- and world-knowledge sentences were congruent or incongruent with respect to semantic expectancies and factual expectancies about the world, respectively. In the semanticknowledge and world-knowledge condition, activation of the LIF region did not differ between groups. In sentences that required integration of speaker information, the autism group showed abnormally reduced activation of the LIF region. The results suggest that people with autism may recruit the LIF region in a different manner in tasks that demand integration of social information.

Abstract

Aging is accompanied by an impairment of associative memory. The medial temporal lobe and fronto-striatal network, both involved in associative memory, are known to decline functionally and structurally with age, leading to the so-called associative binding deficit and the resource deficit. Because the MTL and fronto-striatal network interact, they might also be able to support each other. We therefore employed an episodic memory task probing memory for sequences of object–location associations, where the demand on self-initiated processing was manipulated during encoding: either all the objects were visible simultaneously (rich environmental support) or every object became visible transiently (poor environmental support). Following the concept of resource deficit, we hypothesised that the elderly probably have difficulty using their declarative memory system when demands on self-initiated processing are high (poor environmental support). Our behavioural study showed that only the young use the rich environmental support in a systematic way, by placing the objects next to each other. With the task adapted for fMRI, we found that elderly showed stronger activity than young subjects during retrieval of environmentally richly encoded information in the basal ganglia, thalamus, left middle temporal/fusiform gyrus and right medial temporal lobe (MTL). These results indicate that rich environmental support leads to recruitment of the declarative memory system in addition to the fronto-striatal network in elderly, while the young use more posterior brain regions likely related to imagery. We propose that elderly try to solve the task by additional recruitment of stimulus-response associations, which might partly compensate their limited attentional resources.

Abstract

ehavioral studies have suggested that placebo analgesia is partly mediated by the endogenous opioid system. Expanding on these results we have shown that the opioid-receptor-rich rostral anterior cingulate cortex (rACC) is activated in both placebo and opioid analgesia. However, there are also differences between the two treatments. While opioids have direct pharmacological effects, acting on the descending pain inhibitory system, placebo analgesia depends on neocortical top-down mechanisms. An important difference may be that expectations are met to a lesser extent in placebo treatment as compared with a specific treatment, yielding a larger error signal. As these processes previously have been shown to influence other types of perceptual experiences, we hypothesized that they also may drive placebo analgesia. Imaging studies suggest that lateral orbitofrontal cortex (lObfc) and ventrolateral prefrontal cortex (vlPFC) are involved in processing expectation and error signals. We re-analyzed two independent functional imaging experiments related to placebo analgesia and emotional placebo to probe for a differential processing in these regions during placebo treatment vs. opioid treatment and to test if this activity is associated with the placebo response. In the first dataset lObfc and vlPFC showed an enhanced activation in placebo analgesia vs. opioid analgesia. Furthermore, the rACC activity co-varied with the prefrontal regions in the placebo condition specifically. A similar correlation between rACC and vlPFC was reproduced in another dataset involving emotional placebo and correlated with the degree of the placebo effect. Our results thus support that placebo is different from specific treatment with a prefrontal top-down influence on rACC.

Abstract

A aquisição da leitura decorre ao longo de diversas etapas, desde o momento em que a criança inicia o contacto com o alfabeto até ao momento em que se torna um leitor competente, apto a ler correcta e fluentemente. Compreender a evolução desta competência através de uma análise da diferenciação do peso de variáveis preditoras da leitura possibilita teorizar sobre os mecanismos cognitivos envolvidos nas diferentes fases de desenvolvimento da leitura. Realizámos um estudo transversal com 568 alunos do segundo ao quarto ano do primeiro ciclo do Ensino Básico, em que se avaliou o impacto de capacidades de processamento fonológico, nomeação rápida, conhecimento letra-som e vocabulário, bem como de capacidades cognitivas mais gerais (inteligência não-verbal e memória de trabalho), na exactidão e velocidade da leitura. De uma forma geral, os resultados mostraram que, apesar da consciência fonológica permanecer como o preditor mais importante da exactidão e fluência da leitura, o seu peso decresce à medida que a escolaridade aumenta. Observou-se também que, à medida que o contributo da consciência fonológica para a explicação da velocidade de leitura diminuía, aumentava o contributo de outras variáveis mais associadas ao automatismo e reconhecimento lexical, tais como a nomeação rápida e o vocabulário. Em suma, podemos dizer que ao longo da escolaridade se observa uma alteração dinâmica dos processos cognitivos subjacentes à leitura, o que sugere que a criança evolui de uma estratégia de leitura ancorada em processamentos sub-lexicais, e como tal mais dependente de processamentos fonológicos, para uma estratégia baseada no reconhecimento ortográfico das palavras.

Abstract

In a recent fMRI study we showed that left posterior middle temporal gyrus (LpMTG) subserves the retrieval of a word's lexical-syntactic properties from the mental lexicon (long-term memory), while left posterior inferior frontal gyrus (LpIFG) is involved in unifying (on-line integration of) this information into a sentence structure (Snijders et al., 2009). In addition, the right IFG, right MTG, and the right striatum were involved in the unification process. Here we report results from a psychophysical interactions (PPI) analysis in which we investigated the effective connectivity between LpIFG and LpMTG during unification, and how the right hemisphere areas and the striatum are functionally connected to the unification network. LpIFG and LpMTG both showed enhanced connectivity during the unification process with a region slightly superior to our previously reported LpMTG. Right IFG better predicted right temporal activity when unification processes were more strongly engaged, just as LpIFG better predicted left temporal activity. Furthermore, the striatum showed enhanced coupling to LpIFG and LpMTG during unification. We conclude that bilateral inferior frontal and posterior temporal regions are functionally connected during sentence-level unification. Cortico-subcortical connectivity patterns suggest cooperation between inferior frontal and striatal regions in performing unification operations on lexical-syntactic representations retrieved from LpMTG.

Abstract

Two decades of pharmacologic research on the human capacity to implicitly acquire knowledge as well as cognitive skills and procedures have yielded surprisingly few conclusive insights. We review the empirical literature of the neuropharmacology of implicit learning. We evaluate the findings in the context of relevant computational models related to neurotransmittors such as dopamine, serotonin, acetylcholine and noradrenalin. These include models for reinforcement learning, sequence production, and categorization. We conclude, based on the reviewed literature, that one can predict improved implicit acquisition by moderately elevated dopamine levels and impaired implicit acquisition by moderately decreased dopamine levels. These effects are most prominent in the dorsal striatum. This is supported by a range of behavioral tasks in the empirical literature. Similar predictions can be made for serotonin, although there is yet a lack of support in the literature for serotonin involvement in classical implicit learning tasks. There is currently a lack of evidence for a role of the noradrenergic and cholinergic systems in implicit and related forms of learning. GABA modulators, including benzodiazepines, seem to affect implicit learning in a complex manner and further research is needed. Finally, we identify allosteric AMPA receptors modulators as a potentially interesting target for future investigation of the neuropharmacology of procedural and implicit learning.

Abstract

Background: In synaesthesia, sensations in a particular modality cause additional experiences in a second, unstimulated modality (e.g., letters elicit colour). Understanding how synaesthesia is mediated in the brain can help to understand normal processes of perceptual awareness and multisensory integration. In several neuroimaging studies, enhanced brain activity for grapheme-colour synaesthesia has been found in ventral-occipital areas that are also involved in real colour processing. Our question was whether the neural correlates of synaesthetically induced colour and real colour experience are truly shared. Methodology/Principal Findings: First, in a free viewing functional magnetic resonance imaging (fMRI) experiment, we located main effects of synaesthesia in left superior parietal lobule and in colour related areas. In the left superior parietal lobe, individual differences between synaesthetes (projector-associator distinction) also influenced brain activity, confirming the importance of the left superior parietal lobe for synaesthesia. Next, we applied a repetition suppression paradigm in fMRI, in which a decrease in the BOLD (blood-oxygenated-level-dependent) response is generally observed for repeated stimuli. We hypothesized that synaesthetically induced colours would lead to a reduction in BOLD response for subsequently presented real colours, if the neural correlates were overlapping. We did find BOLD suppression effects induced by synaesthesia, but not within the colour areas. Conclusions/Significance: Because synaesthetically induced colours were not able to suppress BOLD effects for real colour, we conclude that the neural correlates of synaesthetic colour experience and real colour experience are not fully shared. We propose that synaesthetic colour experiences are mediated by higher-order visual pathways that lie beyond the scope of classical, ventral-occipital visual areas. Feedback from these areas, in which the left parietal cortex is likely to play an important role, may induce V4 activation and the percept of synaesthetic colour.

Abstract

A recurrently connected attractor neural network with a Hebbian learning rule is currently our best ANN analogy for a piece cortex. Functionally biological memory operates on a spectrum of time scales with regard to induction and retention, and it is modulated in complex ways by sub-cortical neuromodulatory systems. Moreover, biological memory networks are commonly believed to be highly distributed and engage many co-operating cortical areas. Here we focus on the temporal aspects of induction and retention of memory in a connectionist type attractor memory model of a piece of cortex. A continuous time, forgetful Bayesian-Hebbian learning rule is described and compared to the characteristics of LTP and LTD seen experimentally. More generally, an attractor network implementing this learning rule can operate as a long-term, intermediate-term, or short-term memory. Modulation of the print-now signal of the learning rule replicates some experimental memory phenomena, like e.g. the von Restorff effect.

Abstract

It has previously been suggested that the activity in sensory regions of the brain can be modulated by attentional mechanisms during parallel cognitive processing. To investigate whether such attention-related modulations are present in the processing of pain, the regional cerebral blood ¯ow was measured using [15O]butanol and positron emission tomography in conditions involving both pain and parallel cognitive demands. The painful stimulus consisted of the standard cold pressor test and the cognitive task was a computerised perceptual maze test. The activations during the maze test reproduced findings in previous studies of the same cognitive task. The cold pressor test evoked signi®cant activity in the contralateral S1, and bilaterally in the somatosensory association areas (including S2), the ACC and the mid-insula. The activity in the somatosensory association areas and periaqueductal gray/midbrain were significantly modified, i.e. relatively decreased, when the subjects also were performing the maze task. The altered activity was accompanied with significantly lower ratings of pain during the cognitive task. In contrast, lateral orbitofrontal regions showed a relative increase of activity during pain combined with the maze task as compared to only pain, which suggests the possibility of the involvement of frontal cortex in modulation of regions processing pain

Abstract

Capacity limited memory systems need to gradually forget old information in order to avoid catastrophic forgetting where all stored information is lost. This can be achieved by allowing new information to overwrite old, as in the so-called palimpsest memory. This paper describes a new such learning rule employed in an attractor neural network. The network does not exhibit catastrophic forgetting, has a capacity dependent on the learning time constant and exhibits recency e!ects in retrieval