Publications

Abstract

Naming speed deficits are well documented in developmental dyslexia, expressed by slower naming times and more errors in response to familiar items. Here we used event-related potentials (ERPs) to examine at what processing level the deficits in dyslexia emerge during a discrete-naming task. Dyslexic and skilled adult control readers performed a primed object-naming task, in which the relationship between the prime and the target was manipulated along perceptual, semantic and phonological dimensions. A 3×2 design that crossed Relationship Type (Visual, Phonemic Onset, and Semantic) with Relatedness (Related and Unrelated) was used. An attenuated N/P190 – indexing early visual processing – and N300 – which index late visual processing – was observed to pictures preceded by perceptually related (vs. unrelated) primes in the control but not in the dyslexic group. These findings suggest suboptimal processing in early stages of object processing in dyslexia, when integration and mapping of perceptual information to a more form-specific percept in memory take place. On the other hand, both groups showed an N400 effect associated with semantically related pictures (vs. unrelated), taken to reflect intact integration of semantic similarities in both dyslexic and control readers. We also found an electrophysiological effect of phonological priming in the N400 range – that is, an attenuated N400 to objects preceded by phonemic related primes vs. unrelated – while it showed a more widespread distributed and more pronounced over the right hemisphere in the dyslexics. Topographic differences between groups might have originated from a word form encoding process with different characteristics in dyslexics compared to control readers.

Abstract

his study investigates the interaction between surface and colour knowledge information during object recognition. In two different experiments, participants were instructed to decide whether two presented stimuli belonged to the same object identity. On the non-matching trials, we manipulated the shape and colour knowledge information activated by the two stimuli by creating four different stimulus pairs: (1) similar in shape and colour (e.g. TOMATO–APPLE); (2) similar in shape and dissimilar in colour (e.g. TOMATO–COCONUT); (3) dissimilar in shape and similar in colour (e.g. TOMATO–CHILI PEPPER) and (4) dissimilar in both shape and colour (e.g. TOMATO–PEANUT). The object pictures were presented in typical and atypical colours and also in black-and-white. The interaction between surface and colour knowledge showed to be contingent upon shape information: while colour knowledge is more important for recognising structurally similar shaped objects, surface colour is more prominent for recognising structurally dissimilar shaped objects.

Abstract

During oral reading, the eyes tend to be ahead of the voice (eye-voice span, EVS). It has been hypothesized that the extent to which this happens depends on the automaticity of reading processes, namely on the speed of print-to-sound conversion. We tested whether EVS is affected by another automaticity component – immunity from interference. To that end, we manipulated word familiarity (high-frequency, lowfrequency, and pseudowords, PW) and word length as proxies of immunity from interference, and we used linear mixed effects models to measure the effects of both variables on the time interval at which readers do parallel processing by gazing at word N C 1 while not having articulated word N yet (offset EVS). Parallel processing was enhanced by automaticity, as shown by familiarity length interactions on offset EVS, and it was impeded by lack of automaticity, as shown by the transformation of offset EVS into voice-eye span (voice ahead of the offset of the eyes) in PWs. The relation between parallel processing and automaticity was strengthened by the fact that offset EVS predicted reading velocity. Our findings contribute to understand how the offset EVS, an index that is obtained in oral reading, may tap into different components of automaticity that underlie reading ability, oral or silent. In addition, we compared the duration of the offset EVS with the average reference duration of stages in word production, and we saw that the offset EVS may accommodate for more than the articulatory programming stage of word N.

Abstract

Two different forms of parafoveal dysfunction have been hypothesized as core deficits of dyslexic individuals: reduced parafoveal preview benefits (“too little parafovea”) and increased costs of parafoveal load (“too much parafovea”). We tested both hypotheses in a single eye-tracking experiment using a modified serial rapid automatized naming (RAN) task. Comparisons between dyslexic and non-dyslexic adults showed reduced parafoveal preview benefits in dyslexics, without increased costs of parafoveal load. Reduced parafoveal preview benefits were observed in a naming task, but not in a silent letter-finding task, indicating that the parafoveal dysfunction may be consequent to the overload with extracting phonological information from orthographic input. Our results suggest that dyslexics’ parafoveal dysfunction is not based on strict visuo-attentional factors, but nevertheless they stress the importance of extra-phonological processing. Furthermore, evidence of reduced parafoveal preview benefits in dyslexia may help understand why serial RAN is an important reading predictor in adulthood

Abstract

When learning a new language, we build brain networks to process and represent the acquired words and syntax and integrate these with existing language representations. It is an open question whether the same or different neural mechanisms are involved in learning and processing a novel language compared to the native language(s). Here we investigated the neural repetition effects of repeating known and novel word orders while human subjects were in the early stages of learning a new language. Combining a miniature language with a syntactic priming paradigm, we examined the neural correlates of language learning online using functional magnetic resonance imaging (fMRI). In left inferior frontal gyrus (LIFG) and posterior temporal cortex the repetition of novel syntactic structures led to repetition enhancement, while repetition of known structures resulted in repetition suppression. Additional verb repetition led to an
increase in the syntactic repetition enhancement effect in language-related brain regions. Similarly the repetition of verbs led to repetition enhancement effects in areas related to lexical and semantic processing, an effect that continued to increase in a subset of these regions. Repetition enhancement might reflect a mechanism to build and strengthen a neural network to process novel syntactic structures and lexical items. By contrast, the observed repetition suppression points to overlapping neural mechanisms for native and new language constructions when these have sufficient structural similarities.

Abstract

Visual episodic memory encoding was investigated using echoplanar magnetic resonance imaging at 2.0 x 2.0 mm2 resolution and 1.0 mm section thickness, which allows for functional mapping of hippocampal, parahippocampal, and ventral occipital regions with reduced magnetic susceptibility artifact. The memory task was based on 54 image pairs each consisting of a complex visual scene and the face of one of six different photographers. A second group of subjects viewed the same set of images without memory instruction as well as a reversing checkerboard. Apart from visual activation in occipital cortical areas, episodic memory encoding revealed consistent activation in the parahippocampal gyrus but not in the hippocampus proper. This ®nding was most prominently evidenced in sagittal maps covering the right hippocampal formation. Mean activated volumes were 432±293 µl and 259±179 µl for intentional memory encoding and non-instructed viewing, respectively. In contrast, the checkerboard paradigm elicited pure visual activation without parahippocampal involvement.

Abstract

This paper presents a new approach to functional magnetic resonance imaging (FMRI) data analysis. The main difference lies in the view of what comprises an observation. Here we treat the data from one scanning session (comprising t volumes, say) as one observation. This is contrary to the conventional way of looking at the data where each session is treated as t different observations. Thus instead of viewing the v voxels comprising the 3D volume of the brain as the variables, we suggest the usage of the vt hypervoxels comprising the 4D volume of the brain-over-session as the variables. A linear model is fitted to the 4D volumes originating from different sessions. Parameter estimation and hypothesis testing in this model can be performed with standard techniques. The hypothesis testing generates 4D statistical images (SIs) to which any relevant test statistic can be applied. In this paper we describe two test statistics, one voxel based and one cluster based, that can be used to test a range of hypotheses. There are several benefits in treating the data from each session as one observation, two of which are: (i) the temporal characteristics of the signal can be investigated without an explicit model for the blood oxygenation level dependent (BOLD) contrast response function, and (ii) the observations (sessions) can be assumed to be independent and hence inference on the 4D SI can be made by nonparametric or Monte Carlo methods. The suggested 4D approach is applied to FMRI data and is shown to accurately detect the expected signal

Abstract

Recent functional brain imaging studies have shown that sensory-specific brain regions that are activated during perception/encoding of sensory-specific information are reactivated during memory retrieval of the same information. Here we used PET to examine whether verbal retrieval of action phrases is associated with reactivation of motor brain regions if the actions were overtly or covertly performed during encoding. Compared to a verbal condition, encoding by means of overt as well as covert activity was associated with differential activity in regions in contralateral somatosensory and motor cortex. Several of these regions were reactivated during retrieval. Common to both the overt and covert conditions was reactivation of regions in left ventral motor cortex and left inferior parietal cortex. A direct comparison of the overt and covert activity conditions showed that activation and reactivation of left dorsal parietal cortex and right cerebellum was specific to the overt condition. These results support the reactivation hypothesis by showing that verbal-explicit memory of actions involves areas that are engaged during overt and covert motor activity.

Abstract

The study of illiterate subjects, which for specific socio-cultural reasons did not have the opportunity to acquire basic reading and writing skills, represents one approach to study the interaction between neurobiological and cultural factors in cognitive development and the functional organization of the human brain. In addition the naturally occurring illiteracy may serve as a model for studying the influence of alphabetic orthography on auditory-verbal language. In this paper we have reviewed some recent behavioral and functional neuroimaging data indicating that learning an alphabetic written language modulates the auditory-verbal language system in a non-trivial way and provided support for the hypothesis that the functional architecture of the brain is modulated by literacy. We have also indicated that the effects of literacy and formal schooling is not limited to language related skills but appears to affect also other cognitive domains. In particular, we indicate that formal schooling influences 2D but not 3D visual naming skills. We have also pointed to the importance of using ecologically relevant tasks when comparing literate and illiterate subjects. We also demonstrate the applicability of a network approach in elucidating differences in the functional organization of the brain between groups. The strength of such an approach is the ability to study patterns of interactions between functionally specialized brain regions and the possibility to compare such patterns of brain interactions between groups or functional states. This complements the more commonly used activation approach to functional neuroimaging data, which characterize functionally specialized regions, and provides important data characterizing the functional interactions between these regions.

Abstract

The modulatory influence of literacy on the cognitive system of the human brain has been indicated in behavioral, neuroanatomic, and functional neuroimaging studies. In this study we explored the functional consequences of formal education and the acquisition of an alphabetic written language on two- and three-dimensional visual naming. The results show that illiterate subjects perform significantly worse on immediate naming of two-dimensional representations of common everyday objects compared to literate subjects, both in terms of accuracy and reaction times. In contrast, there was no significant difference when the subjects named the corresponding real objects. The results suggest that formal education and learning to read and to write modulate the cognitive process involved in processing two- but not three-dimensional representations of common everyday objects. Both the results of the reaction time and the error pattern analyses can be interpreted as indicating that the major influence of literacy affects the visual system or the interaction between the visual and the language systems. We suggest that the visual system in a wide sense and/or the interface between the visual and the language system are differently formatted in literate and illiterate subjects. In other words, we hypothesize that the pattern of interactions in the functional–anatomical networks subserving visual naming, that is, the interactions within and between the visual and language processing networks, differ in literate and illiterate subjects

Abstract

The strength of a memory trace is modulated by a variety of factors such as arousal, attention, context, type of processing during encoding, salience and novelty of the experience. Some of these factors can be modeled as a variable plasticity level in the memory system, controlled by arousal or relevance-estimating systems. We demonstrate that a Bayesian confidence propagation neural network with learning time constant modulated in this way exhibits enhanced recall of an item tagged as salient. Proactive and retroactive inhibition of other items is also demonstrated as well as an inverted U-shape response to overall plasticity

Abstract

In the present PET study we explore some functional aspects of the interaction between attentional/control processes and learning/memory processes. The network of brain regions supporting recall of abstract designs were studied in a less practiced and in a well practiced state. The results indicate that automaticity, i.e., a decreased dependence on attentional and working memory resources, develops as a consequence of practice. This corresponds to the practice related decreases of activity in the prefrontal, anterior cingulate, and posterior parietal regions. In addition, the activity of the medial temporal regions decreased as a function of practice. This indicates an inverse relation between the strength of encoding and the activation of the MTL during retrieval. Furthermore, the pattern of practice related increases in the auditory, posterior insular-opercular extending into perisylvian supra marginal region, and the right mid occipito-temporal region, may reflect a lower degree of inhibitory attentional modulation of task irrelevant processing and more fully developed representations of the abstract designs, respectively. We also suggest that free recall is dependent on bilateral prefrontal processing, in particular non-automatic free recall. The present results cofirm previous functional neuroimaging studies of memory retrieval indicating that recall is subserved by a network of interacting brain regions. Furthermore, the results indicate that some components of the neural network subserving free recall may have a dynamic role and that there is a functional restructuring of the information processing networks during the learning process.

Abstract

Recent event-related FMRI studies indicate that the prefrontal (PFC) and the medial temporal lobe (MTL) regions are more active during effective encoding than during ineffective encoding. The within-subject design and the use of well-educated young college students in these studies makes it important to replicate these results in other study populations. In this PET study, we used an auditory word-pair association cued-recall paradigm and investigated a group of healthy upper middle-aged/older illiterate women. We observed a positive correlation between cued-recall success and the regional cerebral blood flow of the left inferior PFC (BA 47) and the MTLs. Specifically, we used the cuedrecall success as a covariate in a general linear model and the results confirmed that the left inferior PFC and the MTLare more active during effective encoding than during ineffective encoding. These effects were observed during encoding of both semantically and phonologically related word pairs, indicating that these effects are robust in the studied population, that is, reproducible within group. These results generalize the results of Brewer et al. (1998, Science 281, 1185– 1187) and Wagner et al. (1998, Science 281, 1188–1191) to an upper middle aged/older illiterate population. In addition, the present study indicates that effective relational encoding correlates positively with the activity of the anterior medial temporal lobe regions.

Abstract

A fundamental problem in the study of learning is that learning-related changes may be confounded by nonspecific time effects. There are several strategies for handling this problem. This problem may be of greater significance in functional magnetic resonance imaging (fMRI) compared to positron emission tomography (PET). Using the general linear model, we describe, compare, and discuss two approaches for separating learning-related from nonspecific time effects. The first approach makes assumptions on the general behavior of nonspecific effects and explicitly models these effects, i.e., nonspecific time effects are incorporated as a linear or nonlinear confounding covariate in the statistical model. The second strategy makes no a priori assumption concerning the form of nonspecific time effects, but implicitly controls for nonspecific effects using an interaction approach, i.e., learning effects are assessed with an interaction contrast. The two approaches depend on specific assumptions and have specific limitations. With certain experimental designs, both approaches may be used and the results compared, lending particular support to effects that are independent of the method used. A third and perhaps better approach that sometimes may be practically unfeasible is to use a completely temporally balanced experimental design. The choice of approach may be of particular importance when learning related effects are studied with fMRI.

Abstract

The objective of this study was to investigate the central processing of dynamic mechanical allodynia in patients with mononeuropathy. Regional cerebral bloodflow, as an indicator of neuronal activity, was measured with positron emission tomography. Paired comparisons were made between three different states; rest, allodynia during brushing the painful skin area, and brushing of the homologous contralateral area. Bilateral activations were observed in the primary somatosensory cortex (S1) and the secondary somatosensory cortex (S2) during allodynia compared to rest. The S1 activation contralateral to the site of the stimulus was more expressed during allodynia than during innocuous touch. Significant activations of the contralateral posterior parietal cortex, the periaqueductal gray (PAG), the thalamus bilaterally and motor areas were also observed in the allodynic state compared to both non-allodynic states. In the anterior cingulate cortex (ACC) there was only a suggested activation when the allodynic state was compared with the non-allodynic states. In order to account for the individual variability in the intensity of allodynia and ongoing spontaneous pain, rCBF was regressed on the individually reported pain intensity, and significant covariations were observed in the ACC and the right anterior insula. Significantly decreased regional blood flow was observed bilaterally in the medial and lateral temporal lobe as well as in the occipital and posterior cingulate cortices when the allodynic state was compared to the non-painful conditions. This finding is consistent with previous studies suggesting attentional modulation and a central coping strategy for known and expected painful stimuli. Involvement of the medial pain system has previously been reported in patients with mononeuropathy during ongoing spontaneous pain. This study reveals a bilateral activation of the lateral pain system as well as involvement of the medial pain system during dynamic mechanical allodynia in patients with mononeuropathy.