Karl-Magnus Petersson

Publications

Displaying 1 - 100 of 166
  • Araújo, S., Reis, A., Faísca, L., & Petersson, K. M. (in press). Brain sensitivity to words and the “word recognition potential”. In D. Marques, & J. H. Toscano (Eds.), De las neurociencias a la neuropsicologia: el estúdio del cerebro humano. Barranquilla, Colombia: Corporación Universitaria Reformada.
  • Zhu, Z., Bastiaansen, M. C. M., Hakun, J. G., Petersson, K. M., Wang, S., & Hagoort, P. (2019). Semantic unification modulates N400 and BOLD signal change in the brain: A simultaneous EEG-fMRI study. Journal of Neurolinguistics. Advance online publication. doi:10.1016/j.jneuroling.2019.100855.

    Abstract

    Semantic unification during sentence comprehension has been associated with amplitude change of the N400 in event-related potential (ERP) studies, and activation in the left inferior frontal gyrus (IFG) in functional magnetic resonance imaging (fMRI) studies. However, the specificity of this activation to semantic unification remains unknown. To more closely examine the brain processes involved in semantic unification, we employed simultaneous EEG-fMRI to time-lock the semantic unification related N400 change, and integrated trial-by-trial variation in both N400 and BOLD change beyond the condition-level BOLD change difference measured in traditional fMRI analyses. Participants read sentences in which semantic unification load was parametrically manipulated by varying cloze probability. Separately, ERP and fMRI results replicated previous findings, in that semantic unification load parametrically modulated the amplitude of N400 and cortical activation. Integrated EEG-fMRI analyses revealed a different pattern in which functional activity in the left IFG and bilateral supramarginal gyrus (SMG) was associated with N400 amplitude, with the left IFG activation and bilateral SMG activation being selective to the condition-level and trial-level of semantic unification load, respectively. By employing the EEG-fMRI integrated analyses, this study among the first sheds light on how to integrate trial-level variation in language comprehension.
  • Duarte, R., Uhlmann, M., Van den Broek, D., Fitz, H., Petersson, K. M., & Morrison, A. (2018). Encoding symbolic sequences with spiking neural reservoirs. In Proceedings of the 2018 International Joint Conference on Neural Networks (IJCNN). doi:10.1109/IJCNN.2018.8489114.

    Abstract

    Biologically inspired spiking networks are an important tool to study the nature of computation and cognition in neural systems. In this work, we investigate the representational capacity of spiking networks engaged in an identity mapping task. We compare two schemes for encoding symbolic input, one in which input is injected as a direct current and one where input is delivered as a spatio-temporal spike pattern. We test the ability of networks to discriminate their input as a function of the number of distinct input symbols. We also compare performance using either membrane potentials or filtered spike trains as state variable. Furthermore, we investigate how the circuit behavior depends on the balance between excitation and inhibition, and the degree of synchrony and regularity in its internal dynamics. Finally, we compare different linear methods of decoding population activity onto desired target labels. Overall, our results suggest that even this simple mapping task is strongly influenced by design choices on input encoding, state-variables, circuit characteristics and decoding methods, and these factors can interact in complex ways. This work highlights the importance of constraining computational network models of behavior by available neurobiological evidence.
  • Huettig, F., Lachmann, T., Reis, A., & Petersson, K. M. (2018). Distinguishing cause from effect - Many deficits associated with developmental dyslexia may be a consequence of reduced and suboptimal reading experience. Language, Cognition and Neuroscience, 33(3), 333-350. doi:10.1080/23273798.2017.1348528.

    Abstract

    The cause of developmental dyslexia is still unknown despite decades of intense research. Many causal explanations have been proposed, based on the range of impairments displayed by affected individuals. Here we draw attention to the fact that many of these impairments are also shown by illiterate individuals who have not received any or very little reading instruction. We suggest that this fact may not be coincidental and that the performance differences of both illiterates and individuals with dyslexia compared to literate controls are, to a substantial extent, secondary consequences of either reduced or suboptimal reading experience or a combination of both. The search for the primary causes of reading impairments will make progress if the consequences of quantitative and qualitative differences in reading experience are better taken into account and not mistaken for the causes of reading disorders. We close by providing four recommendations for future research.
  • Inacio, F., Faisca, L., Forkstam, C., Araujo, S., Bramao, I., Reis, A., & Petersson, K. M. (2018). Implicit sequence learning is preserved in dyslexic children. Annals of Dyslexia, 68(1), 1-14. doi:10.1007/s11881-018-0158-x.

    Abstract

    This study investigates the implicit sequence learning abilities of dyslexic children using an artificial grammar learning task with an extended exposure period. Twenty children with developmental dyslexia participated in the study and were matched with two control groups—one matched for age and other for reading skills. During 3 days, all participants performed an acquisition task, where they were exposed to colored geometrical forms sequences with an underlying grammatical structure. On the last day, after the acquisition task, participants were tested in a grammaticality classification task. Implicit sequence learning was present in dyslexic children, as well as in both control groups, and no differences between groups were observed. These results suggest that implicit learning deficits per se cannot explain the characteristic reading difficulties of the dyslexics.
  • Silva, S., Folia, V., Inácio, F., Castro, S. L., & Petersson, K. M. (2018). Modality effects in implicit artificial grammar learning: An EEG study. Brain Research, 1687, 50-59. doi:10.1016/j.brainres.2018.02.020.

    Abstract

    Recently, it has been proposed that sequence learning engages a combination of modality-specific operating networks and modality-independent computational principles. In the present study, we compared the behavioural and EEG outcomes of implicit artificial grammar learning in the visual vs. auditory modality. We controlled for the influence of surface characteristics of sequences (Associative Chunk Strength), thus focusing on the strictly structural aspects of sequence learning, and we adapted the paradigms to compensate for known frailties of the visual modality compared to audition (temporal presentation, fast presentation rate). The behavioural outcomes were similar across modalities. Favouring the idea of modality-specificity, ERPs in response to grammar violations differed in topography and latency (earlier and more anterior component in the visual modality), and ERPs in response to surface features emerged only in the auditory modality. In favour of modality-independence, we observed three common functional properties in the late ERPs of the two grammars: both were free of interactions between structural and surface influences, both were more extended in a grammaticality classification test than in a preference classification test, and both correlated positively and strongly with theta event-related-synchronization during baseline testing. Our findings support the idea of modality-specificity combined with modality-independence, and suggest that memory for visual vs. auditory sequences may largely contribute to cross-modal differences.
  • Coco, M. I., Araujo, S., & Petersson, K. M. (2017). Disentangling stimulus plausibility and contextual congruency: Electro-physiological evidence for differential cognitive dynamics. Neuropsychologia, 96, 150-163. doi:10.1016/j.neuropsychologia.2016.12.008.

    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
  • Silva, S., Inácio, F., Folia, V., & Petersson, K. M. (2017). Eye movements in implicit artificial grammar learning. Journal of Experimental Psychology: Learning, Memory, and Cognition, 43(9), 1387-1402. doi:10.1037/xlm0000350.

    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
  • Silva, S., Petersson, K. M., & Castro, S. L. (2017). The effects of ordinal load on incidental temporal learning. Quarterly Journal of Experimental Psychology, 70(4), 664-674. doi:10.1080/17470218.2016.1146909.

    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.
  • Silva, S., Folia, V., Hagoort, P., & Petersson, K. M. (2017). The P600 in Implicit Artificial Grammar Learning. Cognitive Science, 41(1), 137-157. doi:10.1111/cogs.12343.

    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.
  • Udden, J., Ingvar, M., Hagoort, P., & Petersson, K. M. (2017). Broca’s region: A causal role in implicit processing of grammars with crossed non-adjacent dependencies. Cognition, 164, 188-198. doi:10.1016/j.cognition.2017.03.010.

    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.
  • Araújo, S., Faísca, L., Reis, A., Marques, J. F., & Petersson, K. M. (2016). Visual naming deficits in dyslexia: An ERP investigation of different processing domains. Neuropsychologia, 91, 61-76. doi:10.1016/j.neuropsychologia.2016.07.007.

    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.
  • Bramão, I., Reis, A., Petersson, K. M., & Faísca, L. (2016). Knowing that strawberries are red and seeing red strawberries: The interaction between surface colour and colour knowledge information. Journal of Cognitive Psychology, 28(6), 641-657. doi:10.1080/20445911.2016.1182171.

    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.
  • Silva, S., Reis, A., Casaca, L., Petersson, K. M., & Faísca, L. (2016). When the eyes no longer lead: Familiarity and length effects eye-voice span. Frontiers in Psychology, 7: 1720. doi:10.3389/fpsyg.2016.01720.

    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.
  • Silva, S., Petersson, K. M., & Castro, S. (2016). Rhythm in the brain: Is music special? In D. Da Silva Marques, & J. Avila-Toscano (Eds.), Neuroscience to neuropsychology: The study of the human brain (pp. 29-54). Barranquilla, Colombia: Ediciones CUR.
  • Silva, S., Faísca, L., Araújo, S., Casaca, L., Carvalho, L., Petersson, K. M., & Reis, A. (2016). Too little or too much? Parafoveal preview benefits and parafoveal load costs in dyslexic adults. Annals of Dyslexia, 66(2), 187-201. doi:10.1007/s11881-015-0113-z.

    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
  • Weber, K., Christiansen, M., Petersson, K. M., Indefrey, P., & Hagoort, P. (2016). fMRI syntactic and lexical repetition effects reveal the initial stages of learning a new language. The Journal of Neuroscience, 36, 6872-6880. doi:10.1523/JNEUROSCI.3180-15.2016.

    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.
  • Araújo, S., Faísca, L., Bramão, I., Reis, A., & Petersson, K. M. (2015). Lexical and sublexical orthographic processing: An ERP study with skilled and dyslexic adult readers. Brain and Language, 141, 16-27. doi:10.1016/j.bandl.2014.11.007.

    Abstract

    This ERP study investigated the cognitive nature of the P1–N1 components during orthographic processing. We used an implicit reading task with various types of stimuli involving different amounts of sublexical or lexical orthographic processing (words, pseudohomophones, pseudowords, nonwords, and symbols), and tested average and dyslexic readers. An orthographic regularity effect (pseudowords– nonwords contrast) was observed in the average but not in the dyslexic group. This suggests an early sensitivity to the dependencies among letters in word-forms that reflect orthographic structure, while the dyslexic brain apparently fails to be appropriately sensitive to these complex features. Moreover, in the adults the N1-response may already reflect lexical access: (i) the N1 was sensitive to the familiar vs. less familiar orthographic sequence contrast; (ii) and early effects of the phonological form (words-pseudohomophones contrast) were also found. Finally, the later N320 component was attenuated in the dyslexics, suggesting suboptimal processing in later stages of phonological analysis.
  • Araújo, S., Reis, A., Petersson, K. M., & Faísca, L. (2015). Rapid automatized naming and reading performance: A meta-analysis. Journal of Educational Psychology, 107(3), 868-883. doi:10.1037/edu0000006.

    Abstract

    Evidence that rapid naming skill is associated with reading ability has become increasingly prevalent in recent years. However, there is considerable variation in the literature concerning the magnitude of this relationship. The objective of the present study was to provide a comprehensive analysis of the evidence on the relationship between rapid automatized naming (RAN) and reading performance. To this end, we conducted a meta-analysis of the correlational relationship between these 2 constructs to (a) determine the overall strength of the RAN–reading association and (b) identify variables that systematically moderate this relationship. A random-effects model analysis of data from 137 studies (857 effect sizes; 28,826 participants) indicated a moderate-to-strong relationship between RAN and reading performance (r = .43, I2 = 68.40). Further analyses revealed that RAN contributes to the 4 measures of reading (word reading, text reading, non-word reading, and reading comprehension), but higher coefficients emerged in favor of real word reading and text reading. RAN stimulus type and type of reading score were the factors with the greatest moderator effect on the magnitude of the RAN–reading relationship. The consistency of orthography and the subjects’ grade level were also found to impact this relationship, although the effect was contingent on reading outcome. It was less evident whether the subjects’ reading proficiency played a role in the relationship. Implications for future studies are discussed.
  • Araújo, S., Faísca, L., Bramão, I., Petersson, K. M., & Reis, A. (2014). Lexical and phonological processes in dyslexic readers: Evidences from a visual lexical decision task. Dyslexia, 20, 38-53. doi:10.1002/dys.1461.

    Abstract

    The aim of the present study was to investigate whether reading failure in the context of an orthography of intermediate consistency is linked to inefficient use of the lexical orthographic reading procedure. The performance of typically developing and dyslexic Portuguese-speaking children was examined in a lexical decision task, where the stimulus lexicality, word frequency and length were manipulated. Both lexicality and length effects were larger in the dyslexic group than in controls, although the interaction between group and frequency disappeared when the data were transformed to control for general performance factors. Children with dyslexia were influenced in lexical decision making by the stimulus length of words and pseudowords, whereas age-matched controls were influenced by the length of pseudowords only. These findings suggest that non-impaired readers rely mainly on lexical orthographic information, but children with dyslexia preferentially use the phonological decoding procedure—albeit poorly—most likely because they struggle to process orthographic inputs as a whole such as controls do. Accordingly, dyslexic children showed significantly poorer performance than controls for all types of stimuli, including words that could be considered over-learned, such as high-frequency words. This suggests that their orthographic lexical entries are less established in the orthographic lexicon
  • Basnakova, J., Weber, K., Petersson, K. M., Van Berkum, J. J. A., & Hagoort, P. (2014). Beyond the language given: The neural correlates of inferring speaker meaning. Cerebral Cortex, 24(10), 2572-2578. doi:10.1093/cercor/bht112.

    Abstract

    Even though language allows us to say exactly what we mean, we often use language to say things indirectly, in a way that depends on the specific communicative context. For example, we can use an apparently straightforward sentence like "It is hard to give a good presentation" to convey deeper meanings, like "Your talk was a mess!" One of the big puzzles in language science is how listeners work out what speakers really mean, which is a skill absolutely central to communication. However, most neuroimaging studies of language comprehension have focused on the arguably much simpler, context-independent process of understanding direct utterances. To examine the neural systems involved in getting at contextually constrained indirect meaning, we used functional magnetic resonance imaging as people listened to indirect replies in spoken dialog. Relative to direct control utterances, indirect replies engaged dorsomedial prefrontal cortex, right temporo-parietal junction and insula, as well as bilateral inferior frontal gyrus and right medial temporal gyrus. This suggests that listeners take the speaker's perspective on both cognitive (theory of mind) and affective (empathy-like) levels. In line with classic pragmatic theories, our results also indicate that currently popular "simulationist" accounts of language comprehension fail to explain how listeners understand the speaker's intended message.
  • Folia, V., & Petersson, K. M. (2014). Implicit structured sequence learning: An fMRI study of the structural mere-exposure effect. Frontiers in Psychology, 5: 41. doi:10.3389/fpsyg.2014.00041.

    Abstract

    In this event-related FMRI study we investigated the effect of five days of implicit acquisition on preference classification by means of an artificial grammar learning (AGL) paradigm based on the structural mere-exposure effect and preference classification using a simple right-linear unification grammar. This allowed us to investigate implicit AGL in a proper learning design by including baseline measurements prior to grammar exposure. After 5 days of implicit acquisition, the FMRI results showed activations in a network of brain regions including the inferior frontal (centered on BA 44/45) and the medial prefrontal regions (centered on BA 8/32). Importantly, and central to this study, the inclusion of a naive preference FMRI baseline measurement allowed us to conclude that these FMRI findings were the intrinsic outcomes of the learning process itself and not a reflection of a preexisting functionality recruited during classification, independent of acquisition. Support for the implicit nature of the knowledge utilized during preference classification on day 5 come from the fact that the basal ganglia, associated with implicit procedural learning, were activated during classification, while the medial temporal lobe system, associated with explicit declarative memory, was consistently deactivated. Thus, preference classification in combination with structural mere-exposure can be used to investigate structural sequence processing (syntax) in unsupervised AGL paradigms with proper learning designs.
  • Pacheco, A., Araújo, S., Faísca, L., de Castro, S. L., Petersson, K. M., & Reis, A. (2014). Dyslexia's heterogeneity: Cognitive profiling of Portuguese children with dyslexia. Reading and Writing, 27(9), 1529-1545. doi:10.1007/s11145-014-9504-5.

    Abstract

    Recent studies have emphasized that developmental dyslexia is a multiple-deficit disorder, in contrast to the traditional single-deficit view. In this context, cognitive profiling of children with dyslexia may be a relevant contribution to this unresolved discussion. The aim of this study was to profile 36 Portuguese children with dyslexia from the 2nd to 5th grade. Hierarchical cluster analysis was used to group participants according to their phonological awareness, rapid automatized naming, verbal short-term memory, vocabulary, and nonverbal intelligence abilities. The results suggested a two-cluster solution: a group with poorer performance on phoneme deletion and rapid automatized naming compared with the remaining variables (Cluster 1) and a group characterized by underperforming on the variables most related to phonological processing (phoneme deletion and digit span), but not on rapid automatized naming (Cluster 2). Overall, the results seem more consistent with a hybrid perspective, such as that proposed by Pennington and colleagues (2012), for understanding the heterogeneity of dyslexia. The importance of characterizing the profiles of individuals with dyslexia becomes clear within the context of constructing remediation programs that are specifically targeted and are more effective in terms of intervention outcome.

    Supplementary material

    11145_2014_9504_MOESM1_ESM.doc
  • Silva, S., Branco, P., Barbosa, F., Marques-Teixeira, J., Petersson, K. M., & Castro, S. L. (2014). Musical phrase boundaries, wrap-up and the closure positive shift. Brain Research, 1585, 99-107. doi:10.1016/j.brainres.2014.08.025.

    Abstract

    We investigated global integration (wrap-up) processes at the boundaries of musical phrases by comparing the effects of well and non-well formed phrases on event-related potentials time-locked to two boundary points: the onset and the offset of the boundary pause. The Closure Positive Shift, which is elicited at the boundary offset, was not modulated by the quality of phrase structure (well vs. non-well formed). In contrast, the boundary onset potentials showed different patterns for well and non-well formed phrases. Our results contribute to specify the functional meaning of the Closure Positive Shift in music, shed light on the large-scale structural integration of musical input, and raise new hypotheses concerning shared resources between music and language.
  • Silva, S., Barbosa, F., Marques-Teixeira, J., Petersson, K. M., & Castro, S. L. (2014). You know when: Event-related potentials and theta/beat power indicate boundary prediction in music. Journal of Integrative Neuroscience, 13(1), 19-34. doi:10.1142/S0219635214500022.

    Abstract

    Neuroscientific and musicological approaches to music cognition indicate that listeners familiarized in the Western tonal tradition expect a musical phrase boundary at predictable time intervals. However, phrase boundary prediction processes in music remain untested. We analyzed event-related potentials (ERPs) and event-related induced power changes at the onset and offset of a boundary pause. We made comparisons with modified melodies, where the pause was omitted and filled by tones. The offset of the pause elicited a closure positive shift (CPS), indexing phrase boundary detection. The onset of the filling tones elicited significant increases in theta and beta powers. In addition, the P2 component was larger when the filling tones started than when they ended. The responses to boundary omission suggest that listeners expected to hear a boundary pause. Therefore, boundary prediction seems to coexist with boundary detection in music segmentation.
  • Van Leeuwen, T. M., Petersson, K. M., Langner, O., Rijpkema, M., & Hagoort, P. (2014). Color specificity in the human V4 complex: An fMRI repetition suppression study. In T. D. Papageorgiou, G. I. Cristopoulous, & S. M. Smirnakis (Eds.), Advanced Brain Neuroimaging Topics in Health and Disease - Methods and Applications (pp. 275-295). Rijeka, Croatia: Intech. doi:10.5772/58278.
  • Van Leeuwen, T. M., Lamers, M. J. A., Petersson, K. M., Gussenhoven, C., Poser, B., & Hagoort, P. (2014). Phonological markers of information structure: An fMRI study. Neuropsychologia, 58(1), 64-74. doi:10.1016/j.neuropsychologia.2014.03.017.

    Abstract

    In this fMRI study we investigate the neural correlates of information structure integration during sentence comprehension in Dutch. We looked into how prosodic cues (pitch accents) that signal the information status of constituents to the listener (new information) are combined with other types of information during the unification process. The difficulty of unifying the prosodic cues into overall sentence meaning was manipulated by constructing sentences in which the pitch accent did (focus-accent agreement), and sentences in which the pitch accent did not (focus-accent disagreement) match the expectations for focus constituents of the sentence. In case of a mismatch, the load on unification processes increases. Our results show two anatomically distinct effects of focus-accent disagreement, one located in the posterior left inferior frontal gyrus (LIFG, BA6/44), and one in the more anterior-ventral LIFG (BA 47/45). Our results confirm that information structure is taken into account during unification, and imply an important role for the LIFG in unification processes, in line with previous fMRI studies.

    Supplementary material

    mmc1.doc
  • Andics, A., McQueen, J. M., & Petersson, K. M. (2013). Mean-based neural coding of voices. NeuroImage, 79, 351-360. doi:10.1016/j.neuroimage.2013.05.002.

    Abstract

    The social significance of recognizing the person who talks to us is obvious, but the neural mechanisms that mediate talker identification are unclear. Regions along the bilateral superior temporal sulcus (STS) and the inferior frontal cortex (IFC) of the human brain are selective for voices, and they are sensitive to rapid voice changes. Although it has been proposed that voice recognition is supported by prototype-centered voice representations, the involvement of these category-selective cortical regions in the neural coding of such "mean voices" has not previously been demonstrated. Using fMRI in combination with a voice identity learning paradigm, we show that voice-selective regions are involved in the mean-based coding of voice identities. Voice typicality is encoded on a supra-individual level in the right STS along a stimulus-dependent, identity-independent (i.e., voice-acoustic) dimension, and on an intra-individual level in the right IFC along a stimulus-independent, identity-dependent (i.e., voice identity) dimension. Voice recognition therefore entails at least two anatomically separable stages, each characterized by neural mechanisms that reference the central tendencies of voice categories.
  • Kristensen, L. B., Wang, L., Petersson, K. M., & Hagoort, P. (2013). The interface between language and attention: Prosodic focus marking recruits a general attention network in spoken language comprehension. Cerebral Cortex, 23, 1836-1848. doi:10.1093/cercor/bhs164.

    Abstract

    In spoken language, pitch accent can mark certain information as focus, whereby more attentional resources are allocated to the focused information. Using functional magnetic resonance imaging, this study examined whether pitch accent, used for marking focus, recruited general attention networks during sentence comprehension. In a language task, we independently manipulated the prosody and semantic/pragmatic congruence of sentences. We found that semantic/pragmatic processing affected bilateral inferior and middle frontal gyrus. The prosody manipulation showed bilateral involvement of the superior/inferior parietal cortex, superior and middle temporal cortex, as well as inferior, middle, and posterior parts of the frontal cortex. We compared these regions with attention networks localized in an auditory spatial attention task. Both tasks activated bilateral superior/inferior parietal cortex, superior temporal cortex, and left precentral cortex. Furthermore, an interaction between prosody and congruence was observed in bilateral inferior parietal regions: for incongruent sentences, but not for congruent ones, there was a larger activation if the incongruent word carried a pitch accent, than if it did not. The common activations between the language task and the spatial attention task demonstrate that pitch accent activates a domain general attention network, which is sensitive to semantic/pragmatic aspects of language. Therefore, attention and language comprehension are highly interactive.

    Supplementary material

    Kirstensen_Cer_Cor_Suppl_Mat.doc
  • Nieuwenhuis, I. L., Folia, V., Forkstam, C., Jensen, O., & Petersson, K. M. (2013). Sleep promotes the extraction of grammatical rules. PLoS One, 8(6): e65046. doi:10.1371/journal.pone.0065046.

    Abstract

    Grammar acquisition is a high level cognitive function that requires the extraction of complex rules. While it has been proposed that offline time might benefit this type of rule extraction, this remains to be tested. Here, we addressed this question using an artificial grammar learning paradigm. During a short-term memory cover task, eighty-one human participants were exposed to letter sequences generated according to an unknown artificial grammar. Following a time delay of 15 min, 12 h (wake or sleep) or 24 h, participants classified novel test sequences as Grammatical or Non-Grammatical. Previous behavioral and functional neuroimaging work has shown that classification can be guided by two distinct underlying processes: (1) the holistic abstraction of the underlying grammar rules and (2) the detection of sequence chunks that appear at varying frequencies during exposure. Here, we show that classification performance improved after sleep. Moreover, this improvement was due to an enhancement of rule abstraction, while the effect of chunk frequency was unaltered by sleep. These findings suggest that sleep plays a critical role in extracting complex structure from separate but related items during integrative memory processing. Our findings stress the importance of alternating periods of learning with sleep in settings in which complex information must be acquired.
  • Segaert, K., Kempen, G., Petersson, K. M., & Hagoort, P. (2013). Syntactic priming and the lexical boost effect during sentence production and sentence comprehension: An fMRI study. Brain and Language, 124, 174-183. doi:10.1016/j.bandl.2012.12.003.

    Abstract

    Behavioral syntactic priming effects during sentence comprehension are typically observed only if both the syntactic structure and lexical head are repeated. In contrast, during production syntactic priming occurs with structure repetition alone, but the effect is boosted by repetition of the lexical head. We used fMRI to investigate the neuronal correlates of syntactic priming and lexical boost effects during sentence production and comprehension. The critical measure was the magnitude of fMRI adaptation to repetition of sentences in active or passive voice, with or without verb repetition. In conditions with repeated verbs, we observed adaptation to structure repetition in the left IFG and MTG, for active and passive voice. However, in the absence of repeated verbs, adaptation occurred only for passive sentences. None of the fMRI adaptation effects yielded differential effects for production versus comprehension, suggesting that sentence comprehension and production are subserved by the same neuronal infrastructure for syntactic processing.

    Supplementary material

    Segaert_Supplementary_data_2013.docx
  • Segaert, K., Weber, K., De Lange, F., Petersson, K. M., & Hagoort, P. (2013). The suppression of repetition enhancement: A review of fMRI studies. Neuropsychologia, 51, 59-66. doi:10.1016/j.neuropsychologia.2012.11.006.

    Abstract

    Repetition suppression in fMRI studies is generally thought to underlie behavioural facilitation effects (i.e., priming) and it is often used to identify the neuronal representations associated with a stimulus. However, this pays little heed to the large number of repetition enhancement effects observed under similar conditions. In this review, we identify several cognitive variables biasing repetition effects in the BOLD response towards enhancement instead of suppression. These variables are stimulus recognition, learning, attention, expectation and explicit memory. We also evaluate which models can account for these repetition effects and come to the conclusion that there is no one single model that is able to embrace all repetition enhancement effects. Accumulation, novel network formation as well as predictive coding models can all explain subsets of repetition enhancement effects.
  • Whitmarsh, S., Udden, J., Barendregt, H., & Petersson, K. M. (2013). Mindfulness reduces habitual responding based on implicit knowledge: Evidence from artificial grammar learning. Consciousness and Cognition, (3), 833-845. doi:10.1016/j.concog.2013.05.007.

    Abstract

    Participants were unknowingly exposed to complex regularities in a working memory task. The existence of implicit knowledge was subsequently inferred from a preference for stimuli with similar grammatical regularities. Several affective traits have been shown to influence AGL performance positively, many of which are related to a tendency for automatic responding. We therefore tested whether the mindfulness trait predicted a reduction of grammatically congruent preferences, and used emotional primes to explore the influence of affect. Mindfulness was shown to correlate negatively with grammatically congruent responses. Negative primes were shown to result in faster and more negative evaluations. We conclude that grammatically congruent preference ratings rely on habitual responses, and that our findings provide empirical evidence for the non-reactive disposition of the mindfulness trait.
  • Araújo, S., Bramão, I., Faísca, L., Petersson, K. M., & Reis, A. (2012). Electrophysiological correlates of impaired reading in dyslexic pre-adolescent children. Brain and Cognition, 79, 79-88. doi:10.1016/j.bandc.2012.02.010.

    Abstract

    In this study, event related potentials (ERPs) were used to investigate the extent to which dyslexics (aged 9–13 years) differ from normally reading controls in early ERPs, which reflect prelexical orthographic processing, and in late ERPs, which reflect implicit phonological processing. The participants performed an implicit reading task, which was manipulated in terms of letter-specific processing, orthographic familiarity, and phonological structure. Comparing consonant- and symbol sequences, the results showed significant differences in the P1 and N1 waveforms in the control but not in the dyslexic group. The reduced P1 and N1 effects in pre-adolescent children with dyslexia suggest a lack of visual specialization for letter-processing. The P1 and N1 components were not sensitive to the familiar vs. less familiar orthographic sequence contrast. The amplitude of the later N320 component was larger for phonologically legal (pseudowords) compared to illegal (consonant sequences) items in both controls and dyslexics. However, the topographic differences showed that the controls were more left-lateralized than the dyslexics. We suggest that the development of the mechanisms that support literacy skills in dyslexics is both delayed and follows a non-normal developmental path. This contributes to the hemispheric differences observed and might reflect a compensatory mechanism in dyslexics.
  • Bramão, I., Francisco, A., Inácio, F., Faísca, L., Reis, A., & Petersson, K. M. (2012). Electrophysiological evidence for colour effects on the naming of colour diagnostic and noncolour diagnostic objects. Visual Cognition, 20, 1164-1185. doi:10.1080/13506285.2012.739215.

    Abstract

    In this study, we investigated the level of visual processing at which surface colour information improves the naming of colour diagnostic and noncolour diagnostic objects. Continuous electroencephalograms were recorded while participants performed a visual object naming task in which coloured and black-and-white versions of both types of objects were presented. The black-and-white and the colour presentations were compared in two groups of event-related potentials (ERPs): (1) The P1 and N1 components, indexing early visual processing; and (2) the N300 and N400 components, which index late visual processing. A colour effect was observed in the P1 and N1 components, for both colour and noncolour diagnostic objects. In addition, for colour diagnostic objects, a colour effect was observed in the N400 component. These results suggest that colour information is important for the naming of colour and noncolour diagnostic objects at different levels of visual processing. It thus appears that the visual system uses colour information, during naming of both object types, at early visual stages; however, for the colour diagnostic objects naming, colour information is also recruited during the late visual processing stages.
  • Bramão, I., Faísca, L., Forkstam, C., Inácio, F., Araújo, S., Petersson, K. M., & Reis, A. (2012). The interaction between surface color and color knowledge: Behavioral and electrophysiological evidence. Brain and Cognition, 78, 28-37. doi:10.1016/j.bandc.2011.10.004.

    Abstract

    In this study, we used event-related potentials (ERPs) to evaluate the contribution of surface color and color knowledge information in object identification. We constructed two color-object verification tasks – a surface and a knowledge verification task – using high color diagnostic objects; both typical and atypical color versions of the same object were presented. Continuous electroencephalogram was recorded from 26 subjects. A cluster randomization procedure was used to explore the differences between typical and atypical color objects in each task. In the color knowledge task, we found two significant clusters that were consistent with the N350 and late positive complex (LPC) effects. Atypical color objects elicited more negative ERPs compared to typical color objects. The color effect found in the N350 time window suggests that surface color is an important cue that facilitates the selection of a stored object representation from long-term memory. Moreover, the observed LPC effect suggests that surface color activates associated semantic knowledge about the object, including color knowledge representations. We did not find any significant differences between typical and atypical color objects in the surface color verification task, which indicates that there is little contribution of color knowledge to resolve the surface color verification. Our main results suggest that surface color is an important visual cue that triggers color knowledge, thereby facilitating object identification.
  • Bramão, I., Faísca, L., Petersson, K. M., & Reis, A. (2012). The contribution of color to object recognition. In I. Kypraios (Ed.), Advances in object recognition systems (pp. 73-88). Rijeka, Croatia: InTech. Retrieved from http://www.intechopen.com/books/advances-in-object-recognition-systems/the-contribution-of-color-in-object-recognition.

    Abstract

    The cognitive processes involved in object recognition remain a mystery to the cognitive sciences. We know that the visual system recognizes objects via multiple features, including shape, color, texture, and motion characteristics. However, the way these features are combined to recognize objects is still an open question. The purpose of this contribution is to review the research about the specific role of color information in object recognition. Given that the human brain incorporates specialized mechanisms to handle color perception in the visual environment, it is a fair question to ask what functional role color might play in everyday vision.
  • Menenti, L., Petersson, K. M., & Hagoort, P. (2012). From reference to sense: How the brain encodes meaning for speaking. Frontiers in Psychology, 2, 384. doi:10.3389/fpsyg.2011.00384.

    Abstract

    In speaking, semantic encoding is the conversion of a non-verbal mental representation (the reference) into a semantic structure suitable for expression (the sense). In this fMRI study on sentence production we investigate how the speaking brain accomplishes this transition from non-verbal to verbal representations. In an overt picture description task, we manipulated repetition of sense (the semantic structure of the sentence) and reference (the described situation) separately. By investigating brain areas showing response adaptation to repetition of each of these sentence properties, we disentangle the neuronal infrastructure for these two components of semantic encoding. We also performed a control experiment with the same stimuli and design but without any linguistic task to identify areas involved in perception of the stimuli per se. The bilateral inferior parietal lobes were selectively sensitive to repetition of reference, while left inferior frontal gyrus showed selective suppression to repetition of sense. Strikingly, a widespread network of areas associated with language processing (left middle frontal gyrus, bilateral superior parietal lobes and bilateral posterior temporal gyri) all showed repetition suppression to both sense and reference processing. These areas are probably involved in mapping reference onto sense, the crucial step in semantic encoding. These results enable us to track the transition from non-verbal to verbal representations in our brains.
  • Petersson, K. M., & Hagoort, P. (2012). The neurobiology of syntax: Beyond string-sets [Review article]. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 367, 1971-1883. doi:10.1098/rstb.2012.0101.

    Abstract

    The human capacity to acquire language is an outstanding scientific challenge to understand. Somehow our language capacities arise from the way the human brain processes, develops and learns in interaction with its environment. To set the stage, we begin with a summary of what is known about the neural organization of language and what our artificial grammar learning (AGL) studies have revealed. We then review the Chomsky hierarchy in the context of the theory of computation and formal learning theory. Finally, we outline a neurobiological model of language acquisition and processing based on an adaptive, recurrent, spiking network architecture. This architecture implements an asynchronous, event-driven, parallel system for recursive processing. We conclude that the brain represents grammars (or more precisely, the parser/generator) in its connectivity, and its ability for syntax is based on neurobiological infrastructure for structured sequence processing. The acquisition of this ability is accounted for in an adaptive dynamical systems framework. Artificial language learning (ALL) paradigms might be used to study the acquisition process within such a framework, as well as the processing properties of the underlying neurobiological infrastructure. However, it is necessary to combine and constrain the interpretation of ALL results by theoretical models and empirical studies on natural language processing. Given that the faculty of language is captured by classical computational models to a significant extent, and that these can be embedded in dynamic network architectures, there is hope that significant progress can be made in understanding the neurobiology of the language faculty.
  • Petersson, K. M., Folia, V., & Hagoort, P. (2012). What artificial grammar learning reveals about the neurobiology of syntax. Brain and Language, 120, 83-95. doi:10.1016/j.bandl.2010.08.003.

    Abstract

    In this paper we examine the neurobiological correlates of syntax, the processing of structured sequences, by comparing FMRI results on artificial and natural language syntax. We discuss these and similar findings in the context of formal language and computability theory. We used a simple right-linear unification grammar in an implicit artificial grammar learning paradigm in 32 healthy Dutch university students (natural language FMRI data were already acquired for these participants). We predicted that artificial syntax processing would engage the left inferior frontal region (BA 44/45) and that this activation would overlap with syntax-related variability observed in the natural language experiment. The main findings of this study show that the left inferior frontal region centered on BA 44/45 is active during artificial syntax processing of well-formed (grammatical) sequence independent of local subsequence familiarity. The same region is engaged to a greater extent when a syntactic violation is present and structural unification becomes difficult or impossible. The effects related to artificial syntax in the left inferior frontal region (BA 44/45) were essentially identical when we masked these with activity related to natural syntax in the same subjects. Finally, the medial temporal lobe was deactivated during this operation, consistent with the view that implicit processing does not rely on declarative memory mechanisms that engage the medial temporal lobe. In the context of recent FMRI findings, we raise the question whether Broca’s region (or subregions) is specifically related to syntactic movement operations or the processing of hierarchically nested non-adjacent dependencies in the discussion section. We conclude that this is not the case. Instead, we argue that the left inferior frontal region is a generic on-line sequence processor that unifies information from various sources in an incremental and recursive manner, independent of whether there are any processing requirements related to syntactic movement or hierarchically nested structures. In addition, we argue that the Chomsky hierarchy is not directly relevant for neurobiological systems.
  • Scheeringa, R., Petersson, K. M., Kleinschmidt, A., Jensen, O., & Bastiaansen, M. C. M. (2012). EEG alpha power modulation of fMRI resting state connectivity. Brain Connectivity, 2, 254-264. doi:10.1089/brain.2012.0088.

    Abstract

    In the past decade, the fast and transient coupling and uncoupling of functionally related brain regions into networks has received much attention in cognitive neuroscience. Empirical tools to study network coupling include fMRI-based functional and/or effective connectivity, and EEG/MEG-based measures of neuronal synchronization. Here we use simultaneously recorded EEG and fMRI to assess whether fMRI-based BOLD connectivity and frequency-specific EEG power are related. Using data collected during resting state, we studied whether posterior EEG alpha power fluctuations are correlated with connectivity within the visual network and between visual cortex and the rest of the brain. The results show that when alpha power increases BOLD connectivity between primary visual cortex and occipital brain regions decreases and that the negative relation of the visual cortex with anterior/medial thalamus decreases and ventral-medial prefrontal cortex is reduced in strength. These effects were specific for the alpha band, and not observed in other frequency bands. Decreased connectivity within the visual system may indicate enhanced functional inhibition during higher alpha activity. This higher inhibition level also attenuates long-range intrinsic functional antagonism between visual cortex and other thalamic and cortical regions. Together, these results illustrate that power fluctuations in posterior alpha oscillations result in local and long range neural connectivity changes.
  • Segaert, K., Menenti, L., Weber, K., Petersson, K. M., & Hagoort, P. (2012). Shared syntax in language production and language comprehension — An fMRI study. Cerebral Cortex, 22, 1662-1670. doi:10.1093/cercor/bhr249.

    Abstract

    During speaking and listening syntactic processing is a crucial step. It involves specifying syntactic relations between words in a sentence. If the production and comprehension modality share the neuronal substrate for syntactic processing then processing syntax in one modality should lead to adaptation effects in the other modality. In the present functional magnetic resonance imaging experiment, participants either overtly produced or heard descriptions of pictures. We looked for brain regions showing adaptation effects to the repetition of syntactic structures. In order to ensure that not just the same brain regions but also the same neuronal populations within these regions are involved in syntactic processing in speaking and listening, we compared syntactic adaptation effects within processing modalities (syntactic production-to-production and comprehension-to-comprehension priming) with syntactic adaptation effects between processing modalities (syntactic comprehension-to-production and production-to-comprehension priming). We found syntactic adaptation effects in left inferior frontal gyrus (Brodmann's area [BA] 45), left middle temporal gyrus (BA 21), and bilateral supplementary motor area (BA 6) which were equally strong within and between processing modalities. Thus, syntactic repetition facilitates syntactic processing in the brain within and across processing modalities to the same extent. We conclude that that the same neurobiological system seems to subserve syntactic processing in speaking and listening.
  • Silva, C., Faísca, L., Ingvar, M., Petersson, K. M., & Reis, A. (2012). Literacy: Exploring working memory systems. Journal of Clinical and Experimental Neuropsychology, 34(4), 369-377. doi:10.1080/13803395.2011.645017.

    Abstract

    Previous research showed an important association between reading and writing skills (literacy) and the phonological loop. However, the effects of literacy on other working memory components remain unclear. In this study, we investigated performance of illiterate subjects and their matched literate controls on verbal and nonverbal working memory tasks. Results revealed that the phonological loop is significantly influenced by literacy, while the visuospatial sketchpad appears to be less affected or not at all. Results also suggest that the central executive might be influenced by literacy, possibly as an expression of cognitive reserve.

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  • Udden, J., Ingvar, M., Hagoort, P., & Petersson, K. M. (2012). Implicit acquisition of grammars with crossed and nested non-adjacent dependencies: Investigating the push-down stack model. Cognitive Science, 36, 1078-1101. doi:10.1111/j.1551-6709.2012.01235.x.

    Abstract

    A recent hypothesis in empirical brain research on language is that the fundamental difference between animal and human communication systems is captured by the distinction between finite-state and more complex phrase-structure grammars, such as context-free and context-sensitive grammars. However, the relevance of this distinction for the study of language as a neurobiological system has been questioned and it has been suggested that a more relevant and partly analogous distinction is that between non-adjacent and adjacent dependencies. Online memory resources are central to the processing of non-adjacent dependencies as information has to be maintained across intervening material. One proposal is that an external memory device in the form of a limited push-down stack is used to process non-adjacent dependencies. We tested this hypothesis in an artificial grammar learning paradigm where subjects acquired non-adjacent dependencies implicitly. Generally, we found no qualitative differences between the acquisition of non-adjacent dependencies and adjacent dependencies. This suggests that although the acquisition of non-adjacent dependencies requires more exposure to the acquisition material, it utilizes the same mechanisms used for acquiring adjacent dependencies. We challenge the push-down stack model further by testing its processing predictions for nested and crossed multiple non-adjacent dependencies. The push-down stack model is partly supported by the results, and we suggest that stack-like properties are some among many natural properties characterizing the underlying neurophysiological mechanisms that implement the online memory resources used in language and structured sequence processing.
  • De Vries, M. H., Petersson, K. M., Geukes, S., Zwitserlood, P., & Christiansen, M. H. (2012). Processing multiple non-adjacent dependencies: Evidence from sequence learning. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 367, 2065-2076. doi:10.1098/rstb.2011.0414.

    Abstract

    Processing non-adjacent dependencies is considered to be one of the hallmarks of human language. Assuming that sequence-learning tasks provide a useful way to tap natural-language-processing mechanisms, we cross-modally combined serial reaction time and artificial-grammar learning paradigms to investigate the processing of multiple nested (A1A2A3B3B2B1) and crossed dependencies (A1A2A3B1B2B3), containing either three or two dependencies. Both reaction times and prediction errors highlighted problems with processing the middle dependency in nested structures (A1A2A3B3_B1), reminiscent of the ‘missing-verb effect’ observed in English and French, but not with crossed structures (A1A2A3B1_B3). Prior linguistic experience did not play a major role: native speakers of German and Dutch—which permit nested and crossed dependencies, respectively—showed a similar pattern of results for sequences with three dependencies. As for sequences with two dependencies, reaction times and prediction errors were similar for both nested and crossed dependencies. The results suggest that constraints on the processing of multiple non-adjacent dependencies are determined by the specific ordering of the non-adjacent dependencies (i.e. nested or crossed), as well as the number of non-adjacent dependencies to be resolved (i.e. two or three). Furthermore, these constraints may not be specific to language but instead derive from limitations on structured sequence learning.
  • Araújo, S., Inácio, F., Francisco, A., Faísca, L., Petersson, K. M., & Reis, A. (2011). Component processes subserving rapid automatized naming in dyslexic and non-dyslexic readers. Dyslexia, 17, 242-255. doi:10.1002/dys.433.

    Abstract

    The current study investigated which time components of rapid automatized naming (RAN) predict group differences between dyslexic and non-dyslexic readers (matched for age and reading level), and how these components relate to different reading measures. Subjects performed two RAN tasks (letters and objects), and data were analyzed through a response time analysis. Our results demonstrated that impaired RAN performance in dyslexic readers mainly stem from enhanced inter-item pause times and not from difficulties at the level of post-access motor production (expressed as articulation rates). Moreover, inter-item pause times account for a significant proportion of variance in reading ability in addition to the effect of phonological awareness in the dyslexic group. This suggests that non-phonological factors may lie at the root of the association between RAN inter-item pauses and reading ability. In normal readers, RAN performance was associated with reading ability only at early ages (i.e. in the reading-matched controls), and again it was the RAN inter-item pause times that explain the association.
  • Araújo, S., Faísca, L., Bramão, I., Inácio, F., Petersson, K. M., & Reis, A. (2011). Object naming in dyslexic children: More than a phonological deficit. The Journal of General Psychology, 138, 215-228. doi:10.1080/00221309.2011.582525.

    Abstract

    In the present study, the authors investigate how some visual factors related to early stages of visual-object naming modulate naming performance in dyslexia. The performance of dyslexic children was compared with 2 control groups—normal readers matched for age and normal readers matched for reading level—while performing a discrete naming task in which color and dimensionality of the visually presented objects were manipulated. The results showed that 2-dimensional naming performance improved for color representations in control readers but not in dyslexics. In contrast to control readers, dyslexics were also insensitive to the stimulus's dimensionality. These findings are unlikely to be explained by a phonological processing problem related to phonological access or retrieval but suggest that dyslexics have a lower capacity for coding and decoding visual surface features of 2-dimensional representations or problems with the integration of visual information stored in long-term memory.
  • Araújo, S., Faísca, L., Petersson, K. M., & Reis, A. (2011). What does rapid naming tell us about dyslexia? Avances en Psicología Latinoamericana, 29, 199-213.

    Abstract

    This article summarizes some of the important findings from research evaluating the relationship between poor rapid naming and impaired reading performance. Substantial evidence shows that dyslexic readers have problems with rapid naming of visual items. Early research assumed that this was a consequence of phonological processing deficits, but recent findings suggest that non-phonological processes may lie at the root of the association between slow naming speed and poor reading. The hypothesis that rapid naming reflects an independent core deficit in dyslexia is supported by the main findings: (1) some dyslexics are characterized by rapid naming difficulties but intact phonological skills; (2) evidence for an independent association between rapid naming and reading competence in the dyslexic readers, when the effect of phonological skills was controlled; (3) rapid naming and phonological processing measures are not reliably correlated. Recent research also reveals greater predictive power of rapid naming, in particular the inter-item pause time, for high-frequency word reading compared to pseudoword reading in developmental dyslexia. Altogether, the results are more consistent with the view that a phonological component alone cannot account for the rapid naming performance in dyslexia. Rather, rapid naming problems may emerge from the inefficiencies in visual-orthographic processing as well as in phonological processing.
  • Bramão, I., Inácio, F., Faísca, L., Reis, A., & Petersson, K. M. (2011). The influence of color information on the recognition of color diagnostic and noncolor diagnostic objects. The Journal of General Psychology, 138(1), 49-65. doi:10.1080/00221309.2010.533718.

    Abstract

    In the present study, the authors explore in detail the level of visual object recognition at which perceptual color information improves the recognition of color diagnostic and noncolor diagnostic objects. To address this issue, 3 object recognition tasks, with different cognitive demands, were designed: (a) an object verification task; (b) a category verification task; and (c) a name verification task. They found that perceptual color information improved color diagnostic object recognition mainly in tasks for which access to the semantic knowledge about the object was necessary to perform the task; that is, in category and name verification. In contrast, the authors found that perceptual color information facilitates noncolor diagnostic object recognition when access to the object’s structural description from long-term memory was necessary—that is, object verification. In summary, the present study shows that the role of perceptual color information in object recognition is dependent on color diagnosticity
  • Bramão, B., Reis, A., Petersson, K. M., & Faísca, L. (2011). The role of color in object recognition: A review and meta-analysis. Acta Psychologica, 138, 244-253. doi:10.1016/j.actpsy.2011.06.010.

    Abstract

    In this study, we systematically review the scientific literature on the effect of color on object recognition. Thirty-five independent experiments, comprising 1535 participants, were included in a meta-analysis. We found a moderate effect of color on object recognition (d = 0.28). Specific effects of moderator variables were analyzed and we found that color diagnosticity is the factor with the greatest moderator effect on the influence of color in object recognition; studies using color diagnostic objects showed a significant color effect (d = 0.43), whereas a marginal color effect was found in studies that used non-color diagnostic objects (d = 0.18). The present study did not permit the drawing of specific conclusions about the moderator effect of the object recognition task; while the meta-analytic review showed that color information improves object recognition mainly in studies using naming tasks (d = 0.36), the literature review revealed a large body of evidence showing positive effects of color information on object recognition in studies using a large variety of visual recognition tasks. We also found that color is important for the ability to recognize artifacts and natural objects, to recognize objects presented as types (line-drawings) or as tokens (photographs), and to recognize objects that are presented without surface details, such as texture or shadow. Taken together, the results of the meta-analysis strongly support the contention that color plays a role in object recognition. This suggests that the role of color should be taken into account in models of visual object recognition.

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  • Folia, V., Forkstam, C., Ingvar, M., Hagoort, P., & Petersson, K. M. (2011). Implicit artificial syntax processing: Genes, preference, and bounded recursion. Biolinguistics, 5(1/2), 105-132.

    Abstract

    The first objective of this study was to compare the brain network engaged by preference classification and the standard grammaticality classification after implicit artificial syntax acquisition by re-analyzing previously reported event-related fMRI data. The results show that preference and grammaticality classification engage virtually identical brain networks, including Broca’s region, consistent with previous behavioral findings. Moreover, the results showed that the effects related to artificial syntax in Broca’s region were essentially the same when masked with variability related to natural syntax processing in the same participants. The second objective was to explore CNTNAP2-related effects in implicit artificial syntax learning by analyzing behavioral and event-related fMRI data from a subsample. The CNTNAP2 gene has been linked to specific language impairment and is controlled by the FOXP2 transcription factor. CNTNAP2 is expressed in language related brain networks in the developing human brain and the FOXP2–CNTNAP2 pathway provides a mechanistic link between clinically distinct syndromes involving disrupted language. Finally, we discuss the implication of taking natural language to be a neurobiological system in terms of bounded recursion and suggest that the left inferior frontal region is a generic on-line sequence processor that unifies information from various sources in an incremental and recursive manner.
  • Petersson, K. M., Forkstam, C., Inácio, F., Bramão, I., Araújo, S., Souza, A. C., Silva, S., & Castro, S. L. (2011). Artificial language learning. In A. Trevisan, & V. Wannmacher Pereira (Eds.), Alfabeltização e cognição (pp. 71-90). Porto Alegre, Brasil: Edipucrs.

    Abstract

    Neste artigo fazemos uma revisão breve de investigações actuais com técnicas comportamentais e de neuroimagem funcional sobre a aprendizagem de uma linguagem artificial em crianças e adultos. Na secção final, discutimos uma possível associação entre dislexia e aprendizagem implícita. Resultados recentes sugerem que a presença de um défice ao nível da aprendizagem implícita pode contribuir para as dificuldades de leitura e escrita observadas em indivíduos disléxicos.
  • Reis, A., Faísca, L., & Petersson, K. M. (2011). Literacia: Modelo para o estudo dos efeitos de uma aprendizagem específica na cognição e nas suas bases cerebrais. In A. Trevisan, J. J. Mouriño Mosquera, & V. Wannmacher Pereira (Eds.), Alfabeltização e cognição (pp. 23-36). Porto Alegro, Brasil: Edipucrs.

    Abstract

    A aquisição de competências de leitura e de escrita pode ser vista como um processo formal de transmissão cultural, onde interagem factores neurobiológicos e culturais. O treino sistemático exigido pela aprendizagem da leitura e da escrita poderá produzir mudanças quantitativas e qualitativas tanto a nível cognitivo como ao nível da organização do cérebro. Estudar sujeitos iletrados e letrados representa, assim, uma oportunidade para investigar efeitos de uma aprendizagem específica no desenvolvimento cognitivo e suas bases cerebrais. Neste trabalho, revemos um conjunto de investigações comportamentais e com métodos de imagem cerebral que indicam que a literacia tem um impacto nas nossas funções cognitivas e na organização cerebral. Mais especificamente, discutiremos diferenças entre letrados e iletrados para domínios cognitivos verbais e não-verbais, sugestivas de que a arquitectura cognitiva é formatada, em parte, pela aprendizagem da leitura e da escrita. Os dados de neuroimagem funcionais e estruturais são também indicadores que a aquisição de uma ortografia alfabética interfere nos processos de organização e lateralização das funções cognitivas.
  • Scheeringa, R., Fries, P., Petersson, K. M., Oostenveld, R., Grothe, I., Norris, D. G., Hagoort, P., & Bastiaansen, M. C. M. (2011). Neuronal dynamics underlying high- and low- frequency EEG oscillations contribute independently to the human BOLD signal. Neuron, 69, 572-583. doi:10.1016/j.neuron.2010.11.044.

    Abstract

    Work on animals indicates that BOLD is preferentially sensitive to local field potentials, and that it correlates most strongly with gamma band neuronal synchronization. Here we investigate how the BOLD signal in humans performing a cognitive task is related to neuronal synchronization across different frequency bands. We simultaneously recorded EEG and BOLD while subjects engaged in a visual attention task known to induce sustained changes in neuronal synchronization across a wide range of frequencies. Trial-by-trial BOLD luctuations correlated positively with trial-by-trial fluctuations in high-EEG gamma power (60–80 Hz) and negatively with alpha and beta power. Gamma power on the one hand, and alpha and beta power on the other hand, independently contributed to explaining BOLD variance. These results indicate that the BOLD-gamma coupling observed in animals can be extrapolated to humans performing a task and that neuronal dynamics underlying high- and low-frequency synchronization contribute independently to the BOLD signal.

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  • Tesink, C. M. J. Y., Buitelaar, J. K., Petersson, K. M., Van der Gaag, R. J., Teunisse, J.-P., & Hagoort, P. (2011). Neural correlates of language comprehension in autism spectrum disorders: When language conflicts with world knowledge. Neuropsychologia, 49, 1095-1104. doi:10.1016/j.neuropsychologia.2011.01.018.

    Abstract

    In individuals with ASD, difficulties with language comprehension are most evident when higher-level semantic-pragmatic language processing is required, for instance when context has to be used to interpret the meaning of an utterance. Until now, it is unclear at what level of processing and for what type of context these difficulties in language comprehension occur. Therefore, in the current fMRI study, we investigated the neural correlates of the integration of contextual information during auditory language comprehension in 24 adults with ASD and 24 matched control participants. Different levels of context processing were manipulated by using spoken sentences that were correct or contained either a semantic or world knowledge anomaly. Our findings demonstrated significant differences between the groups in inferior frontal cortex that were only present for sentences with a world knowledge anomaly. Relative to the ASD group, the control group showed significantly increased activation in left inferior frontal gyrus (LIFG) for sentences with a world knowledge anomaly compared to correct sentences. This effect possibly indicates reduced integrative capacities of the ASD group. Furthermore, world knowledge anomalies elicited significantly stronger activation in right inferior frontal gyrus (RIFG) in the control group compared to the ASD group. This additional RIFG activation probably reflects revision of the situation model after new, conflicting information. The lack of recruitment of RIFG is possibly related to difficulties with exception handling in the ASD group.

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  • De Vries, M., Christiansen, M. H., & Petersson, K. M. (2011). Learning recursion: Multiple nested and crossed dependencies. Biolinguistics, 5(1/2), 010-035.

    Abstract

    Language acquisition in both natural and artificial language learning settings crucially depends on extracting information from sequence input. A shared sequence learning mechanism is thus assumed to underlie both natural and artificial language learning. A growing body of empirical evidence is consistent with this hypothesis. By means of artificial language learning experiments, we may therefore gain more insight in this shared mechanism. In this paper, we review empirical evidence from artificial language learning and computational modelling studies, as well as natural language data, and suggest that there are two key factors that help determine processing complexity in sequence learning, and thus in natural language processing. We propose that the specific ordering of non-adjacent dependencies (i.e., nested or crossed), as well as the number of non-adjacent dependencies to be resolved simultaneously (i.e., two or three) are important factors in gaining more insight into the boundaries of human sequence learning; and thus, also in natural language processing. The implications for theories of linguistic competence are discussed.
  • Andics, A., McQueen, J. M., Petersson, K. M., Gál, V., Rudas, G., & Vidnyánszky, Z. (2010). Neural mechanisms for voice recognition. NeuroImage, 52, 1528-1540. doi:10.1016/j.neuroimage.2010.05.048.

    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.
  • Araújo, S., Pacheco, A., Faísca, L., Petersson, K. M., & Reis, A. (2010). Visual rapid naming and phonological abilities: Different subtypes in dyslexic children. International Journal of Psychology, 45, 443-452. doi:10.1080/00207594.2010.499949.

    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.
  • Bramão, I., Faísca, L., Forkstam, C., Reis, A., & Petersson, K. M. (2010). Cortical brain regions associated with color processing: An FMRI study. The Open Neuroimaging Journal, 4, 164-173. doi:10.2174/1874440001004010164.

    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
  • Bramão, I., Faísca, L., Petersson, K. M., & Reis, A. (2010). The influence of surface color information and color knowledge information in object recognition. American Journal of Psychology, 123, 437-466. Retrieved from http://www.jstor.org/stable/10.5406/amerjpsyc.123.4.0437.

    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.
  • Folia, V., Uddén, J., De Vries, M., Forkstam, C., & Petersson, K. M. (2010). Artificial language learning in adults and children. In M. Gullberg, & P. Indefrey (Eds.), The earliest stages of language learning (pp. 188-220). Malden, MA: Wiley-Blackwell.
  • Folia, V., Uddén, J., De Vries, M., Forkstam, C., & Petersson, K. M. (2010). Artificial language learning in adults and children. Language learning, 60(s2), 188-220. doi:10.1111/j.1467-9922.2010.00606.x.

    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.
  • Groen, W. B., Tesink, C. M. J. Y., Petersson, K. M., Van Berkum, J. J. A., Van der Gaag, R. J., Hagoort, P., & Buitelaar, J. K. (2010). Semantic, factual, and social language comprehension in adolescents with autism: An fMRI study. Cerebral Cortex, 20(8), 1937-1945. doi:10.1093/cercor/bhp264.

    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.
  • Meulenbroek, O., Kessels, R. P. C., De Rover, M., Petersson, K. M., Olde Rikkert, M. G. M., Rijpkema, M., & Fernández, G. (2010). Age-effects on associative object-location memory. Brain Research, 1315, 100-110. doi:10.1016/j.brainres.2009.12.011.

    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.
  • Petrovic, P., Kalso, E., Petersson, K. M., Andersson, J., Fransson, P., & Ingvar, M. (2010). A prefrontal non-opioid mechanism in placebo analgesia. Pain, 150, 59-65. doi:10.1016/j.pain.2010.03.011.

    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.
  • Reis, A., Petersson, K. M., & Faísca, L. (2010). Neuroplasticidade: Os efeitos de aprendizagens específicas no cérebro humano. In C. Nunes, & S. N. Jesus (Eds.), Temas actuais em Psicologia (pp. 11-26). Faro: Universidade do Algarve.
  • Reis, A., Faísca, L., Castro, S.-L., & Petersson, K. M. (2010). Preditores da leitura ao longo da escolaridade: Um estudo com alunos do 1 ciclo do ensino básico. In Actas do VII simpósio nacional de investigação em psicologia (pp. 3117-3132).

    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.
  • Snijders, T. M., Petersson, K. M., & Hagoort, P. (2010). Effective connectivity of cortical and subcortical regions during unification of sentence structure. NeuroImage, 52, 1633-1644. doi:10.1016/j.neuroimage.2010.05.035.

    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.
  • Uddén, J., Folia, V., & Petersson, K. M. (2010). The neuropharmacology of implicit learning. Current Neuropharmacology, 8, 367-381. doi:10.2174/157015910793358178.

    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.
  • Van Leeuwen, T. M., Petersson, K. M., & Hagoort, P. (2010). Synaesthetic colour in the brain: Beyond colour areas. A functional magnetic resonance imaging study of synaesthetes and matched controls. PLoS One, 5(8), E12074. doi:10.1371/journal.pone.0012074.

    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.
  • Araújo, S., Faísca, L., Petersson, K. M., & Reis, A. (2009). Cognitive profiles in Portuguese children with dyslexia. In Abstracts presented at the International Neuropsychological Society, Finnish Neuropsychological Society, Joint Mid-Year Meeting July 29-August 1, 2009. Helsinki, Finland & Tallinn, Estonia (pp. 23). Retrieved from http://www.neuropsykologia.fi/ins2009/INS_MY09_Abstract.pdf.
  • Araújo, S., Faísca, L., Petersson, K. M., & Reis, A. (2009). Visual processing factors contribute to object naming difficulties in dyslexic readers. In Abstracts presented at the International Neuropsychological Society, Finnish Neuropsychological Society, Joint Mid-Year Meeting July 29-August 1, 2009. Helsinki, Finland & Tallinn, Estonia (pp. 39). Retrieved from http://www.neuropsykologia.fi/ins2009/INS_MY09_Abstract.pdf.
  • Bramão, I., Faísca, L., Forkstam, C., Inácio, K., Petersson, K. M., & Reis, A. (2009). Interaction between perceptual color and color knowledge information in object recognition: Behavioral and electrophysiological evidence. In Abstracts presented at the International Neuropsychological Society, Finnish Neuropsychological Society, Joint Mid-Year Meeting July 29-August 1, 2009. Helsinki, Finland & Tallinn, Estonia (pp. 39). Retrieved from http://www.neuropsykologia.fi/ins2009/INS_MY09_Abstract.pdf.
  • Cavaco, P., Curuklu, B., & Petersson, K. M. (2009). Artificial grammar recognition using two spiking neural networks. Frontiers in Neuroinformatics. Conference abstracts: 2nd INCF Congress of Neuroinformatics. doi:10.3389/conf.neuro.11.2009.08.096.

    Abstract

    In this paper we explore the feasibility of artificial (formal) grammar recognition (AGR) using spiking neural networks. A biologically inspired minicolumn architecture is designed as the basic computational unit. A network topography is defined based on the minicolumn architecture, here referred to as nodes, connected with excitatory and inhibitory connections. Nodes in the network represent unique internal states of the grammar’s finite state machine (FSM). Future work to improve the performance of the networks is discussed. The modeling framework developed can be used by neurophysiological research to implement network layouts and compare simulated performance characteristics to actual subject performance.
  • Folia, V., Forkstam, C., Hagoort, P., & Petersson, K. M. (2009). Language comprehension: The interplay between form and content. In N. Taatgen, & H. van Rijn (Eds.), Proceedings of the 31th Annual Conference of the Cognitive Science Society (pp. 1686-1691). Austin, TX: Cognitive Science Society.

    Abstract

    In a 2x2 event-related FMRI study we find support for the idea that the inferior frontal cortex, centered on Broca’s region and its homologue, is involved in constructive unification operations during the structure-building process in parsing for comprehension. Tentatively, we provide evidence for a role of the dorsolateral prefrontal cortex centered on BA 9/46 in the control component of the language system. Finally, the left temporo-parietal cortex, in the vicinity of Wernicke’s region, supports the interaction between the syntax of gender agreement and sentence-level semantics.
  • Forkstam, C., Jansson, A., Ingvar, M., & Petersson, K. M. (2009). Modality transfer of acquired structural regularities: A preference for an acoustic route. In N. Taatgen, & H. Van Rijn (Eds.), Proceedings of the 31th Annual Conference of the Cognitive Science Society. Austin, TX: Cognitive Science Society.

    Abstract

    Human implicit learning can be investigated with implicit artificial grammar learning, a simple model for aspects of natural language acquisition. In this paper we investigate the remaining effect of modality transfer in syntactic classification of an acquired grammatical sequence structure after implicit grammar acquisition. Participants practiced either on acoustically presented syllable sequences or visually presented consonant letter sequences. During classification we independently manipulated the statistical frequency-based and rule-based characteristics of the classification stimuli. Participants performed reliably above chance on the within modality classification task although more so for those working on syllable sequence acquisition. These subjects were also the only group that kept a significant performance level in transfer classification. We speculate that this finding is of particular relevance in consideration of an ecological validity in the input signal in the use of artificial grammar learning and in language learning paradigms at large.
  • Menenti, L., Petersson, K. M., Scheeringa, R., & Hagoort, P. (2009). When elephants fly: Differential sensitivity of right and left inferior frontal gyri to discourse and world knowledge. Journal of Cognitive Neuroscience, 21, 2358-2368. doi:10.1162/jocn.2008.21163.

    Abstract

    Both local discourse and world knowledge are known to influence sentence processing. We investigated how these two sources of information conspire in language comprehension. Two types of critical sentences, correct and world knowledge anomalies, were preceded by either a neutral or a local context. The latter made the world knowledge anomalies more acceptable or plausible. We predicted that the effect of world knowledge anomalies would be weaker for the local context. World knowledge effects have previously been observed in the left inferior frontal region (Brodmann's area 45/47). In the current study, an effect of world knowledge was present in this region in the neutral context. We also observed an effect in the right inferior frontal gyrus, which was more sensitive to the discourse manipulation than the left inferior frontal gyrus. In addition, the left angular gyrus reacted strongly to the degree of discourse coherence between the context and critical sentence. Overall, both world knowledge and the discourse context affect the process of meaning unification, but do so by recruiting partly different sets of brain areas.
  • Pacheco, A., Araújo, S., Faísca, L., Petersson, K. M., & Reis, A. (2009). Profiling dislexic children: Phonology and visual naming skills. In Abstracts presented at the International Neuropsychological Society, Finnish Neuropsychological Society, Joint Mid-Year Meeting July 29-August 1, 2009. Helsinki, Finland & Tallinn, Estonia (pp. 40). Retrieved from http://www.neuropsykologia.fi/ins2009/INS_MY09_Abstract.pdf.
  • Petersson, K. M., Ingvar, M., & Reis, A. (2009). Language and literacy from a cognitive neuroscience perspective. In D. Olsen, & N. Torrance (Eds.), Cambridge handbook of literacy (pp. 152-181). Cambridge: Cambridge University Press.
  • Qin, S., Rijpkema, M., Tendolkar, I., Piekema, C., Hermans, E. J., Binder, M., Petersson, K. M., Luo, J., & Fernández, G. (2009). Dissecting medial temporal lobe contributions to item and associative memory formation. NeuroImage, 46, 874-881. doi:10.1016/j.neuroimage.2009.02.039.

    Abstract

    A fundamental and intensively discussed question is whether medial temporal lobe (MTL) processes that lead to non-associative item memories differ in their anatomical substrate from processes underlying associative memory formation. Using event-related functional magnetic resonance imaging, we implemented a novel design to dissociate brain activity related to item and associative memory formation not only by subsequent memory performance and anatomy but also in time, because the two constituents of each pair to be memorized were presented sequentially with an intra-pair delay of several seconds. Furthermore, the design enabled us to reduce potential differences in memory strength between item and associative memory by increasing task difficulty in the item recognition memory test. Confidence ratings for correct item recognition for both constituents did not differ between trials in which only item memory was correct and trials in which item and associative memory were correct. Specific subsequent memory analyses for item and associative memory formation revealed brain activity that appears selectively related to item memory formation in the posterior inferior temporal, posterior parahippocampal, and perirhinal cortices. In contrast, hippocampal and inferior prefrontal activity predicted successful retrieval of newly formed inter-item associations. Our findings therefore suggest that different MTL subregions indeed play distinct roles in the formation of item memory and inter-item associative memory as expected by several dual process models of the MTL memory system.
  • Scheeringa, R., Petersson, K. M., Oostenveld, R., Norris, D. G., Hagoort, P., & Bastiaansen, M. C. M. (2009). Trial-by-trial coupling between EEG and BOLD identifies networks related to alpha and theta EEG power increases during working memory maintenance. Neuroimage, 44, 1224-1238. doi:10.1016/j.neuroimage.2008.08.041.

    Abstract

    PET and fMRI experiments have previously shown that several brain regions in the frontal and parietal lobe are involved in working memory maintenance. MEG and EEG experiments have shown parametric increases with load for oscillatory activity in posterior alpha and frontal theta power. In the current study we investigated whether the areas found with fMRI can be associated with these alpha and theta effects by measuring simultaneous EEG and fMRI during a modified Sternberg task This allowed us to correlate EEG at the single trial level with the fMRI BOLD signal by forming a regressor based on single trial alpha and theta power estimates. We observed a right posterior, parametric alpha power increase, which was functionally related to decreases in BOLD in the primary visual cortex and in the posterior part of the right middle temporal gyrus. We relate this finding to the inhibition of neuronal activity that may interfere with WM maintenance. An observed parametric increase in frontal theta power was correlated to a decrease in BOLD in regions that together form the default mode network. We did not observe correlations between oscillatory EEG phenomena and BOLD in the traditional WM areas. In conclusion, the study shows that simultaneous EEG fMRI recordings can be successfully used to identify the emergence of functional networks in the brain during the execution of a cognitive task.
  • Snijders, T. M., Vosse, T., Kempen, G., Van Berkum, J. J. A., Petersson, K. M., & Hagoort, P. (2009). Retrieval and unification of syntactic structure in sentence comprehension: An fMRI study using word-category ambiguity. Cerebral Cortex, 19, 1493-1503. doi:10.1093/cercor/bhn187.

    Abstract

    Sentence comprehension requires the retrieval of single word information from long-term memory, and the integration of this information into multiword representations. The current functional magnetic resonance imaging study explored the hypothesis that the left posterior temporal gyrus supports the retrieval of lexical-syntactic information, whereas left inferior frontal gyrus (LIFG) contributes to syntactic unification. Twenty-eight subjects read sentences and word sequences containing word-category (noun–verb) ambiguous words at critical positions. Regions contributing to the syntactic unification process should show enhanced activation for sentences compared to words, and only within sentences display a larger signal for ambiguous than unambiguous conditions. The posterior LIFG showed exactly this predicted pattern, confirming our hypothesis that LIFG contributes to syntactic unification. The left posterior middle temporal gyrus was activated more for ambiguous than unambiguous conditions (main effect over both sentences and word sequences), as predicted for regions subserving the retrieval of lexical-syntactic information from memory. We conclude that understanding language involves the dynamic interplay between left inferior frontal and left posterior temporal regions.

    Supplementary material

    suppl1.pdf suppl2_dutch_stimulus.pdf
  • Tesink, C. M. J. Y., Buitelaar, J. K., Petersson, K. M., Van der Gaag, R. J., Kan, C. C., Tendolkar, I., & Hagoort, P. (2009). Neural correlates of pragmatic language comprehension in autism disorders. Brain, 132, 1941-1952. doi:10.1093/brain/awp103.

    Abstract

    Difficulties with pragmatic aspects of communication are universal across individuals with autism spectrum disorders (ASDs). Here we focused on an aspect of pragmatic language comprehension that is relevant to social interaction in daily life: the integration of speaker characteristics inferred from the voice with the content of a message. Using functional magnetic resonance imaging (fMRI), we examined the neural correlates of the integration of voice-based inferences about the speaker’s age, gender or social background, and sentence content in adults with ASD and matched control participants. Relative to the control group, the ASD group showed increased activation in right inferior frontal gyrus (RIFG; Brodmann area 47) for speakerincongruent sentences compared to speaker-congruent sentences. Given that both groups performed behaviourally at a similar level on a debriefing interview outside the scanner, the increased activation in RIFG for the ASD group was interpreted as being compensatory in nature. It presumably reflects spill-over processing from the language dominant left hemisphere due to higher task demands faced by the participants with ASD when integrating speaker characteristics and the content of a spoken sentence. Furthermore, only the control group showed decreased activation for speaker-incongruent relative to speaker-congruent sentences in right ventral medial prefrontal cortex (vMPFC; Brodmann area 10), including right anterior cingulate cortex (ACC; Brodmann area 24/32). Since vMPFC is involved in self-referential processing related to judgments and inferences about self and others, the absence of such a modulation in vMPFC activation in the ASD group possibly points to atypical default self-referential mental activity in ASD. Our results show that in ASD compensatory mechanisms are necessary in implicit, low-level inferential processes in spoken language understanding. This indicates that pragmatic language problems in ASD are not restricted to high-level inferential processes, but encompass the most basic aspects of pragmatic language processing.
  • Tesink, C. M. J. Y., Petersson, K. M., Van Berkum, J. J. A., Van den Brink, D., Buitelaar, J. K., & Hagoort, P. (2009). Unification of speaker and meaning in language comprehension: An fMRI study. Journal of Cognitive Neuroscience, 21, 2085-2099. doi:10.1162/jocn.2008.21161.

    Abstract

    When interpreting a message, a listener takes into account several sources of linguistic and extralinguistic information. Here we focused on one particular form of extralinguistic information, certain speaker characteristics as conveyed by the voice. Using functional magnetic resonance imaging, we examined the neural structures involved in the unification of sentence meaning and voice-based inferences about the speaker's age, sex, or social background. We found enhanced activation in the inferior frontal gyrus bilaterally (BA 45/47) during listening to sentences whose meaning was incongruent with inferred speaker characteristics. Furthermore, our results showed an overlap in brain regions involved in unification of speaker-related information and those used for the unification of semantic and world knowledge information [inferior frontal gyrus bilaterally (BA 45/47) and left middle temporal gyrus (BA 21)]. These findings provide evidence for a shared neural unification system for linguistic and extralinguistic sources of information and extend the existing knowledge about the role of inferior frontal cortex as a crucial component for unification during language comprehension.
  • Uddén, J., Araújo, S., Forkstam, C., Ingvar, M., Hagoort, P., & Petersson, K. M. (2009). A matter of time: Implicit acquisition of recursive sequence structures. In N. Taatgen, & H. Van Rijn (Eds.), Proceedings of the Thirty-First Annual Conference of the Cognitive Science Society (pp. 2444-2449).

    Abstract

    A dominant hypothesis in empirical research on the evolution of language is the following: the fundamental difference between animal and human communication systems is captured by the distinction between regular and more complex non-regular grammars. Studies reporting successful artificial grammar learning of nested recursive structures and imaging studies of the same have methodological shortcomings since they typically allow explicit problem solving strategies and this has been shown to account for the learning effect in subsequent behavioral studies. The present study overcomes these shortcomings by using subtle violations of agreement structure in a preference classification task. In contrast to the studies conducted so far, we use an implicit learning paradigm, allowing the time needed for both abstraction processes and consolidation to take place. Our results demonstrate robust implicit learning of recursively embedded structures (context-free grammar) and recursive structures with cross-dependencies (context-sensitive grammar) in an artificial grammar learning task spanning 9 days. Keywords: Implicit artificial grammar learning; centre embedded; cross-dependency; implicit learning; context-sensitive grammar; context-free grammar; regular grammar; non-regular grammar
  • Folia, V., Uddén, J., Forkstam, C., Ingvar, M., Hagoort, P., & Petersson, K. M. (2008). Implicit learning and dyslexia. Annals of the New York Academy of Sciences, 1145, 132-150. doi:10.1196/annals.1416.012.

    Abstract

    Several studies have reported an association between dyslexia and implicit learning deficits. It has been suggested that the weakness in implicit learning observed in dyslexic individuals may be related to sequential processing and implicit sequence learning. In the present article, we review the current literature on implicit learning and dyslexia. We describe a novel, forced-choice structural "mere exposure" artificial grammar learning paradigm and characterize this paradigm in normal readers in relation to the standard grammaticality classification paradigm. We argue that preference classification is a more optimal measure of the outcome of implicit acquisition since in the preference version participants are kept completely unaware of the underlying generative mechanism, while in the grammaticality version, the subjects have, at least in principle, been informed about the existence of an underlying complex set of rules at the point of classification (but not during acquisition). On the basis of the "mere exposure effect," we tested the prediction that the development of preference will correlate with the grammaticality status of the classification items. In addition, we examined the effects of grammaticality (grammatical/nongrammatical) and associative chunk strength (ACS; high/low) on the classification tasks (preference/grammaticality). Using a balanced ACS design in which the factors of grammaticality (grammatical/nongrammatical) and ACS (high/low) were independently controlled in a 2 × 2 factorial design, we confirmed our predictions. We discuss the suitability of this task for further investigation of the implicit learning characteristics in dyslexia.
  • Forkstam, C., Elwér, A., Ingvar, M., & Petersson, K. M. (2008). Instruction effects in implicit artificial grammar learning: A preference for grammaticality. Brain Research, 1221, 80-92. doi:10.1016/j.brainres.2008.05.005.

    Abstract

    Human implicit learning can be investigated with implicit artificial grammar learning, a paradigm that has been proposed as a simple model for aspects of natural language acquisition. In the present study we compared the typical yes–no grammaticality classification, with yes–no preference classification. In the case of preference instruction no reference to the underlying generative mechanism (i.e., grammar) is needed and the subjects are therefore completely uninformed about an underlying structure in the acquisition material. In experiment 1, subjects engaged in a short-term memory task using only grammatical strings without performance feedback for 5 days. As a result of the 5 acquisition days, classification performance was independent of instruction type and both the preference and the grammaticality group acquired relevant knowledge of the underlying generative mechanism to a similar degree. Changing the grammatical stings to random strings in the acquisition material (experiment 2) resulted in classification being driven by local substring familiarity. Contrasting repeated vs. non-repeated preference classification (experiment 3) showed that the effect of local substring familiarity decreases with repeated classification. This was not the case for repeated grammaticality classifications. We conclude that classification performance is largely independent of instruction type and that forced-choice preference classification is equivalent to the typical grammaticality classification.
  • Petersson, K. M. (2008). On cognition, structured sequence processing, and adaptive dynamical systems. American Institute of Physics Conference Proceedings, 1060(1), 195-200.

    Abstract

    Cognitive neuroscience approaches the brain as a cognitive system: a system that functionally is conceptualized in terms of information processing. We outline some aspects of this concept and consider a physical system to be an information processing device when a subclass of its physical states can be viewed as representational/cognitive and transitions between these can be conceptualized as a process operating on these states by implementing operations on the corresponding representational structures. We identify a generic and fundamental problem in cognition: sequentially organized structured processing. Structured sequence processing provides the brain, in an essential sense, with its processing logic. In an approach addressing this problem, we illustrate how to integrate levels of analysis within a framework of adaptive dynamical systems. We note that the dynamical system framework lends itself to a description of asynchronous event-driven devices, which is likely to be important in cognition because the brain appears to be an asynchronous processing system. We use the human language faculty and natural language processing as a concrete example through out.
  • De Rover, M., Petersson, K. M., Van der Werf, S. P., Cools, A. R., Berger, H. J., & Fernández, G. (2008). Neural correlates of strategic memory retrieval: Differentiating between spatial-associative and temporal-associative strategies. Human Brain Mapping, 29, 1068-1079. doi:10.1002/hbm.20445.

    Abstract

    Remembering complex, multidimensional information typically requires strategic memory retrieval, during which information is structured, for instance by spatial- or temporal associations. Although brain regions involved in strategic memory retrieval in general have been identified, differences in retrieval operations related to distinct retrieval strategies are not well-understood. Thus, our aim was to identify brain regions whose activity is differentially involved in spatial-associative and temporal-associative retrieval. First, we showed that our behavioral paradigm probing memory for a set of object-location associations promoted the use of a spatial-associative structure following an encoding condition that provided multiple associations to neighboring objects (spatial-associative condition) and the use of a temporal- associative structure following another study condition that provided predominantly temporal associations between sequentially presented items (temporal-associative condition). Next, we used an adapted version of this paradigm for functional MRI, where we contrasted brain activity related to the recall of object-location associations that were either encoded in the spatial- or the temporal-associative condition. In addition to brain regions generally involved in recall, we found that activity in higher-order visual regions, including the fusiform gyrus, the lingual gyrus, and the cuneus, was relatively enhanced when subjects used a spatial-associative structure for retrieval. In contrast, activity in the globus pallidus and the thalamus was relatively enhanced when subjects used a temporal-associative structure for retrieval. In conclusion, we provide evidence for differential involvement of these brain regions related to different types of strategic memory retrieval and the neural structures described play a role in either spatial-associative or temporal-associative memory retrieval.
  • Scheeringa, R., Bastiaansen, M. C. M., Petersson, K. M., Oostenveld, R., Norris, D. G., & Hagoort, P. (2008). Frontal theta EEG activity correlates negatively with the default mode network in resting state. International Journal of Psychophysiology, 67, 242-251. doi:10.1016/j.ijpsycho.2007.05.017.

    Abstract

    We used simultaneously recorded EEG and fMRI to investigate in which areas the BOLD signal correlates with frontal theta power changes, while subjects were quietly lying resting in the scanner with their eyes open. To obtain a reliable estimate of frontal theta power we applied ICA on band-pass filtered (2–9 Hz) EEG data. For each subject we selected the component that best matched the mid-frontal scalp topography associated with the frontal theta rhythm. We applied a time-frequency analysis on this component and used the time course of the frequency bin with the highest overall power to form a regressor that modeled spontaneous fluctuations in frontal theta power. No significant positive BOLD correlations with this regressor were observed. Extensive negative correlations were observed in the areas that together form the default mode network. We conclude that frontal theta activity can be seen as an EEG index of default mode network activity.
  • Tendolkar, I., Arnold, J., Petersson, K. M., Weis, S., Brockhaus-Dumke, A., Van Eijndhoven, P., Buitelaar, J., & Fernandez, G. (2008). Contributions of the medial temporal lobe to declarative memory retrieval: Manipulating the amount of contextual retrieval. Learning and Memory, 15(9), 611-617. doi:10.1101/lm.916708.

    Abstract

    We investigated how the hippocampus and its adjacent mediotemporal structures contribute to contextual and noncontextual declarative memory retrieval by manipulating the amount of contextual information across two levels of the same contextual dimension in a source memory task. A first analysis identified medial temporal lobe (MTL) substructures mediating either contextual or noncontextual retrieval. A linearly weighted analysis elucidated which MTL substructures show a gradually increasing neural activity, depending on the amount of contextual information retrieved. A hippocampal engagement was found during both levels of source memory but not during item memory retrieval. The anterior MTL including the perirhinal cortex was only engaged during item memory retrieval by an activity decrease. Only the posterior parahippocampal cortex showed an activation increasing with the amount of contextual information retrieved. If one assumes a roughly linear relationship between the blood-oxygenation level-dependent (BOLD) signal and the associated cognitive process, our results suggest that the posterior parahippocampal cortex is involved in contextual retrieval on the basis of memory strength while the hippocampus processes representations of item-context binding. The anterior MTL including perirhinal cortex seems to be particularly engaged in familiarity-based item recognition. If one assumes departure from linearity, however, our results can also be explained by one-dimensional modulation of memory strength.
  • Uddén, J., Folia, V., Forkstam, C., Ingvar, M., Fernández, G., Overeem, S., Van Elswijk, G., Hagoort, P., & Petersson, K. M. (2008). The inferior frontal cortex in artificial syntax processing: An rTMS study. Brain Research, 1224, 69-78. doi:10.1016/j.brainres.2008.05.070.

    Abstract

    The human capacity to implicitly acquire knowledge of structured sequences has recently been investigated in artificial grammar learning using functional magnetic resonance imaging. It was found that the left inferior frontal cortex (IFC; Brodmann's area (BA) 44/45) was related to classification performance. The objective of this study was to investigate whether the IFC (BA 44/45) is causally related to classification of artificial syntactic structures by means of an off-line repetitive transcranial magnetic stimulation (rTMS) paradigm. We manipulated the stimulus material in a 2 × 2 factorial design with grammaticality status and local substring familiarity as factors. The participants showed a reliable effect of grammaticality on classification of novel items after 5days of exposure to grammatical exemplars without performance feedback in an implicit acquisition task. The results show that rTMS of BA 44/45 improves syntactic classification performance by increasing the rejection rate of non-grammatical items and by shortening reaction times of correct rejections specifically after left-sided stimulation. A similar pattern of results is observed in FMRI experiments on artificial syntactic classification. These results suggest that activity in the inferior frontal region is causally related to artificial syntax processing.
  • Van Wingen, G. A., Van Broekhoven, F., Verkes, R. J., Petersson, K. M., Bäckström, T., Buitelaar, J. K., & Fernández, G. (2008). Progesterone selectively increases amygdala reactivity in women. Molecular Psychiatry, 13, 325-333. doi:doi:10.1038/sj.mp.4002030.

    Abstract

    The acute neural effects of progesterone are mediated by its neuroactive metabolites allopregnanolone and pregnanolone. These neurosteroids potentiate the inhibitory actions of c-aminobutyric acid (GABA). Progesterone is known to produce anxiolytic effects in animals, but recent animal studies suggest that pregnanolone increases anxiety after a period of low allopregnanolone concentration. This effect is potentially mediated by the amygdala and related to the negative mood symptoms in humans that are observed during increased allopregnanolone levels. Therefore, we investigated with functional magnetic resonance imaging (MRI) whether a single progesterone administration to healthy young women in their follicular phase modulates the amygdala response to salient, biologically relevant stimuli. The progesterone administration increased the plasma concentrations of progesterone and allopregnanolone to levels that are reached during the luteal phase and early pregnancy. The imaging results show that progesterone selectively increased amygdala reactivity. Furthermore, functional connectivity analyses indicate that progesterone modulated functional coupling of the amygdala with distant brain regions. These results reveal a neural mechanism by which progesterone may mediate adverse effects on anxiety and mood.
  • Bramão, I., Mendonça, A., Faísca, L., Ingvar, M., Petersson, K. M., & Reis, A. (2007). The impact of reading and writing skills on a visuo-motor integration task: A comparison between illiterate and literate subjects. Journal of the International Neuropsychological Society, 13(2), 359-364. doi:10.1017/S1355617707070440.

    Abstract

    Previous studies have shown a significant association between reading skills and the performance on visuo-motor tasks. In order to clarify whether reading and writing skills modulate non-linguistic domains, we investigated the performance of two literacy groups on a visuo-motor integration task with non-linguistic stimuli. Twenty-one illiterate participants and twenty matched literate controls were included in the experiment. Subjects were instructed to use the right or the left index finger to point to and touch a randomly presented target on the right or left side of a touch screen. The results showed that the literate subjects were significantly faster in detecting and touching targets on the left compared to the right side of the screen. In contrast, the presentation side did not affect the performance of the illiterate group. These results lend support to the idea that having acquired reading and writing skills, and thus a preferred left-to-right reading direction, influences visual scanning. (JINS, 2007, 13, 359–364
  • Gisselgard, J., Uddén, J., Ingvar, M., & Petersson, K. M. (2007). Disruption of order information by irrelevant items: A serial recognition paradigm. Acta Psychologica, 124(3), 356-369. doi:10.1016/j.actpsy.2006.04.002.

    Abstract

    Irrelevant speech effect (ISE) is defined as a decrement in visually presented digit-list short-term memory performance due to exposure to irrelevant auditory material. Perhaps the most successful theoretical explanation of the effect is the changing state hypothesis. This hypothesis explains the effect in terms of confusion between amodal serial order cues, and represents a view based on the interference caused by the processing of similar order information of the visual and auditory materials. An alternative view suggests that the interference occurs as a consequence of the similarity between the visual and auditory contents of the stimuli. An important argument for the former view is the observation that ISE is almost exclusively observed in tasks that require memory for serial order. However, most short-term memory tasks require that both item and order information be retained in memory. An ideal task to investigate the sensitivity of maintenance of serial order to irrelevant speech would be one that calls upon order information but not item information. One task that is particularly suited to address this issue is serial recognition. In a typical serial recognition task, a list of items is presented and then probed by the same list in which the order of two adjacent items has been transposed. Due to the re-presentation of the encoding string, serial recognition requires primarily the serial order to be maintained while the content of the presented items is deemphasized. In demonstrating a highly significant ISE of changing versus steady-state auditory items in a serial recognition task, the present finding lends support for and extends previous empirical findings suggesting that irrelevant speech has the potential to interfere with the coding of the order of the items to be memorized.
  • Marklund, P., Fransson, P., Cabeza, R., Petersson, K. M., Ingvar, M., & Nyberg, L. (2007). Sustained and transient neural modulations in prefrontal cortex related to declarative long-term memory, working memory, and attention. Cortex, 43(1), 22-37. doi:10.1016/S0010-9452(08)70443-X.

    Abstract

    Common activations in prefrontal cortex (PFC) during episodic and semantic long-term memory (LTM) tasks have been hypothesized to reflect functional overlap in terms of working memory (WM) and cognitive control. To evaluate a WM account of LTM-general activations, the present study took into consideration that cognitive task performance depends on the dynamic operation of multiple component processes, some of which are stimulus-synchronous and transient in nature; and some that are engaged throughout a task in a sustained fashion. PFC and WM may be implicated in both of these temporally independent components. To elucidate these possibilities we employed mixed blocked/event-related functional magnetic resonance imaging (fMRI) procedures to assess the extent to which sustained or transient activation patterns overlapped across tasks indexing episodic and semantic LTM, attention (ATT), and WM. Within PFC, ventrolateral and medial areas exhibited sustained activity across all tasks, whereas more anterior regions including right frontopolar cortex were commonly engaged in sustained processing during the three memory tasks. These findings do not support a WM account of sustained frontal responses during LTM tasks, but instead suggest that the pattern that was common to all tasks reflects general attentional set/vigilance, and that the shared WM-LTM pattern mediates control processes related to upholding task set. Transient responses during the three memory tasks were assessed relative to ATT to isolate item-specific mnemonic processes and were found to be largely distinct from sustained effects. Task-specific effects were observed for each memory task. In addition, a common item response for all memory tasks involved left dorsolateral PFC (DLPFC). The latter response might be seen as reflecting WM processes during LTM retrieval. Thus, our findings suggest that a WM account of shared PFC recruitment in LTM tasks holds for common transient item-related responses rather than sustained state-related responses that are better seen as reflecting more general attentional/control processes.
  • Nieuwland, M. S., Petersson, K. M., & Van Berkum, J. J. A. (2007). On sense and reference: Examining the functional neuroanatomy of referential processing. NeuroImage, 37(3), 993-1004. doi:10.1016/j.neuroimage.2007.05.048.

    Abstract

    In an event-related fMRI study, we examined the cortical networks involved in establishing reference during language comprehension. We compared BOLD responses to sentences containing referentially ambiguous pronouns (e.g., “Ronald told Frank that he…”), referentially failing pronouns (e.g., “Rose told Emily that he…”) or coherent pronouns. Referential ambiguity selectively recruited medial prefrontal regions, suggesting that readers engaged in problem-solving to select a unique referent from the discourse model. Referential failure elicited activation increases in brain regions associated with morpho-syntactic processing, and, for those readers who took failing pronouns to refer to unmentioned entities, additional regions associated with elaborative inferencing were observed. The networks activated by these two referential problems did not overlap with the network activated by a standard semantic anomaly. Instead, we observed a double dissociation, in that the systems activated by semantic anomaly are deactivated by referential ambiguity, and vice versa. This inverse coupling may reflect the dynamic recruitment of semantic and episodic processing to resolve semantically or referentially problematic situations. More generally, our findings suggest that neurocognitive accounts of language comprehension need to address not just how we parse a sentence and combine individual word meanings, but also how we determine who's who and what's what during language comprehension.
  • Petersson, K. M., Silva, C., Castro-Caldas, A., Ingvar, M., & Reis, A. (2007). Literacy: A cultural influence on functional left-right differences in the inferior parietal cortex. European Journal of Neuroscience, 26(3), 791-799. doi:10.1111/j.1460-9568.2007.05701.x.

    Abstract

    The current understanding of hemispheric interaction is limited. Functional hemispheric specialization is likely to depend on both genetic and environmental factors. In the present study we investigated the importance of one factor, literacy, for the functional lateralization in the inferior parietal cortex in two independent samples of literate and illiterate subjects. The results show that the illiterate group are consistently more right-lateralized than their literate controls. In contrast, the two groups showed a similar degree of left-right differences in early speech-related regions of the superior temporal cortex. These results provide evidence suggesting that a cultural factor, literacy, influences the functional hemispheric balance in reading and verbal working memory-related regions. In a third sample, we investigated grey and white matter with voxel-based morphometry. The results showed differences between literacy groups in white matter intensities related to the mid-body region of the corpus callosum and the inferior parietal and parietotemporal regions (literate > illiterate). There were no corresponding differences in the grey matter. This suggests that the influence of literacy on brain structure related to reading and verbal working memory is affecting large-scale brain connectivity more than grey matter per se.
  • Qin, S., Piekema, C., Petersson, K. M., Han, B., Luo, J., & Fernández, G. (2007). Probing the transformation of discontinuous associations into episodic memory: An event-related fMRI study. NeuroImage, 38(1), 212-222. doi:10.1016/j.neuroimage.2007.07.020.

    Abstract

    Using event-related functional magnetic resonance imaging, we identified brain regions involved in storing associations of events discontinuous in time into long-term memory. Participants were scanned while memorizing item-triplets including simultaneous and discontinuous associations. Subsequent memory tests showed that participants remembered both types of associations equally well. First, by constructing the contrast between the subsequent memory effects for discontinuous associations and simultaneous associations, we identified the left posterior parahippocampal region, dorsolateral prefrontal cortex, the basal ganglia, posterior midline structures, and the middle temporal gyrus as being specifically involved in transforming discontinuous associations into episodic memory. Second, we replicated that the prefrontal cortex and the medial temporal lobe (MTL) especially the hippocampus are involved in associative memory formation in general. Our findings provide evidence for distinct neural operation(s) that supports the binding and storing discontinuous associations in memory. We suggest that top-down signals from the prefrontal cortex and MTL may trigger reactivation of internal representation in posterior midline structures of the first event, thus allowing it to be associated with the second event. The dorsolateral prefrontal cortex together with basal ganglia may support this encoding operation by executive and binding processes within working memory, and the posterior parahippocampal region may play a role in binding and memory formation.
  • Reis, A., Faísca, L., Mendonça, S., Ingvar, M., & Petersson, K. M. (2007). Semantic interference on a phonological task in illiterate subjects. Scandinavian Journal of Psychology, 48(1), 69-74. doi:10.1111/j.1467-9450.2006.00544.x.

    Abstract

    Previous research suggests that learning an alphabetic written language influences aspects of the auditory-verbal language system. In this study, we examined whether literacy influences the notion of words as phonological units independent of lexical semantics in literate and illiterate subjects. Subjects had to decide which item in a word- or pseudoword pair was phonologically longest. By manipulating the relationship between referent size and phonological length in three word conditions (congruent, neutral, and incongruent) we could examine to what extent subjects focused on form rather than meaning of the stimulus material. Moreover, the pseudoword condition allowed us to examine global phonological awareness independent of lexical semantics. The results showed that literate performed significantly better than illiterate subjects in the neutral and incongruent word conditions as well as in the pseudoword condition. The illiterate group performed least well in the incongruent condition and significantly better in the pseudoword condition compared to the neutral and incongruent word conditions and suggest that performance on phonological word length comparisons is dependent on literacy. In addition, the results show that the illiterate participants are able to perceive and process phonological length, albeit less well than the literate subjects, when no semantic interference is present. In conclusion, the present results confirm and extend the finding that illiterate subjects are biased towards semantic-conceptual-pragmatic types of cognitive processing.

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