Vasiliki Folia - Academia.edu (original) (raw)

Papers by Vasiliki Folia

Research paper thumbnail of Language comprehension: The interplay between form and content

In a 2x2 event-related FMRI study we find support for the idea that the inferior frontal cortex, ... more 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.

Research paper thumbnail of Assessment of memory skills in illiterates: strategy differences or test artifact?

The Clinical neuropsychologist, 2003

Previous studies have reported that illiterates perform more poorly than literates on a variety o... more Previous studies have reported that illiterates perform more poorly than literates on a variety of neuropsychological measures. We investigated the hypothesis that putative memory deficits in illiterates are an artifact of the assessment tools used rather than a reflection of an 'underdeveloped' ability. In order to accomplish this, we designed two tests, a word list learning test and an object learning test. The illiterate group performed more poorly than semiliterate and literate groups on most variables of the word list learning test, but only on delayed recall and semantic clustering on the object learning test. Our findings suggest that poor memory performance among illiterates can be attributed both to the nature of the task, as well as to the use of different cognitive mechanisms to recall learned information. Presumably, formal education may enhance the innate ability of learning through training individuals in efficient learning and retrieval strategies. We emphasiz...

Research paper thumbnail of Perceived satisfaction from everyday life: are there any predictors in chronic schizophrenics?

Research paper thumbnail of Place of residence-lexical ambiguity: is there any relation between them in schizophrenia?

Research paper thumbnail of Ability of picture naming in chronic schizophrenics

Annals of General Hospital Psychiatry, 2003

Research paper thumbnail of Perceived satisfaction from everyday life among chronic schizophrenics: influences of demographic and clinical factors

Annals of General Hospital Psychiatry, 2003

Research paper thumbnail of Artificial Language Learning in Adults and Children

Research paper thumbnail of Semantic and phonological processing in illiteracy

Journal of the International Neuropsychological Society, 2004

Researchers of cognitive processing in illiteracy have proposed that the acquisition of literacy ... more Researchers of cognitive processing in illiteracy have proposed that the acquisition of literacy modifies the functional organization of the brain. They have suggested that, while illiterate individuals have access only to innate semantic processing skills, those who have learned the correspondence between graphemes and phonemes have several mechanisms available to them through which to process oral language. We conducted 2 experiments to verify that suggestion with respect to language processing, and to elucidate further the differences between literate and illiterate individuals in the cognitive strategies used to process oral language, as well as hemispheric specialization for these processes. Our findings suggest that semantic processing strategies are qualitatively the same in literates and illiterates, despite the fact that overall performance is augmented by increased education. In contrast, explicit processing of oral information based on phonological characteristics appears to be qualitatively different between literates and illiterates: effective strategies in the processing of phonological information depend upon having had a formal education, regardless of the level of education. We also confirmed the differential abilities needed for the processing of semantic and phonological information and related them to hemisphere-specific processing. (JINS, 2004, 10, 818-827.)

Research paper thumbnail of Implicit structured sequence learning: an fMRI study of the structural mere-exposure effect

Frontiers in Psychology, 2014

In this event-related fMRI study we investigated the effect of 5 days of implicit acquisition on ... more In this event-related fMRI study we investigated the effect of 5 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. . Citation: Folia V and Petersson KM (2014) Implicit structured sequence learning: an fMRI study of the structural mere-exposure effect. Front. Psychol. 5:41.

Research paper thumbnail of Lexical Processing in Illiteracy: Effect of Literacy or Education?

Cortex, 2006

Difficulties in phonological processing in illiterates have been attributed to their limited phon... more Difficulties in phonological processing in illiterates have been attributed to their limited phonological awareness, a consequence of their lack of literacy. We sought to explore the potential influence of education on auditory lexical processing above and beyond literacy per se. In order to achieve this goal, we compared a lexical decision making paradigm with a repetition paradigm using words and pseudo-words. We based this choice of tasks on previous research, which has shown that pseudo-word repetition is dependent on the phonological loop; such studies have thus demonstrated a literacy effect on repetition. Instead, lexical decision making is known to depend on the size of one's vocabulary, which is influenced by the level of education attained. Our sample comprised three groups: illiterate no education, literate/low education and literate/high education, individuals. The pattern of our findings confirmed that literacy has an effect on the capacity of the phonological loop, as our illiterate group alone had difficulty with repetition, as compared with both literate/educated groups. Also, our findings suggested an education effect on lexical decision making, as we found a gradation in the performance of the three groups. Therefore, we succeeded in dissecting the effect of literacy and education on auditory lexical processing through the application and comparison of two simple paradigms.

Research paper thumbnail of The inferior frontal cortex in artificial syntax processing: An rTMS study

Brain Research, 2008

The human capacity to implicitly acquire knowledge of structured sequences has recently been inve... more 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. a v a i l a b l e a t w w w. s c i e n c e d i r e c t . c o m w w w. e l s e v i e r. c o m / l o c a t e / b r a i n r e s ARTICLE IN PRESS Please cite this article as: Uddén, J. et al., The inferior frontal cortex in artificial syntax processing: An rTMS study, Brain Res. (2008),

Research paper thumbnail of What artificial grammar learning reveals about the neurobiology of syntax

Brain and Language, 2012

In this paper we examine the neurobiological correlates of syntax, the processing of structured s... more 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.

Research paper thumbnail of Implicit Learning and Dyslexia

Annals of the New York Academy of Sciences, 2008

Several studies have reported an association between dyslexia and implicit learning deficits. It ... more 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. resentation and relate this to a visuographic input-output code. This is typically achieved by means of a supervised learning process (i.e., teaching), in contrast to natural language acquisition, which is largely a spontaneous, non-supervised, and self-organized acquisition process ). Aspects of language can also be an object of metalinguistic awareness: the intentional and explicit control over aspects of phonology, syntax, semantics,

Research paper thumbnail of The Neuropharmacology of Implicit Learning

Current Neuropharmacology, 2010

Two decades of pharmacologic research on the human capacity to implicitly acquire knowledge as we... more 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.

Research paper thumbnail of Language comprehension: The interplay between form and content

In a 2x2 event-related FMRI study we find support for the idea that the inferior frontal cortex, ... more 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.

Research paper thumbnail of Assessment of memory skills in illiterates: strategy differences or test artifact?

The Clinical neuropsychologist, 2003

Previous studies have reported that illiterates perform more poorly than literates on a variety o... more Previous studies have reported that illiterates perform more poorly than literates on a variety of neuropsychological measures. We investigated the hypothesis that putative memory deficits in illiterates are an artifact of the assessment tools used rather than a reflection of an 'underdeveloped' ability. In order to accomplish this, we designed two tests, a word list learning test and an object learning test. The illiterate group performed more poorly than semiliterate and literate groups on most variables of the word list learning test, but only on delayed recall and semantic clustering on the object learning test. Our findings suggest that poor memory performance among illiterates can be attributed both to the nature of the task, as well as to the use of different cognitive mechanisms to recall learned information. Presumably, formal education may enhance the innate ability of learning through training individuals in efficient learning and retrieval strategies. We emphasiz...

Research paper thumbnail of Perceived satisfaction from everyday life: are there any predictors in chronic schizophrenics?

Research paper thumbnail of Place of residence-lexical ambiguity: is there any relation between them in schizophrenia?

Research paper thumbnail of Ability of picture naming in chronic schizophrenics

Annals of General Hospital Psychiatry, 2003

Research paper thumbnail of Perceived satisfaction from everyday life among chronic schizophrenics: influences of demographic and clinical factors

Annals of General Hospital Psychiatry, 2003

Research paper thumbnail of Artificial Language Learning in Adults and Children

Research paper thumbnail of Semantic and phonological processing in illiteracy

Journal of the International Neuropsychological Society, 2004

Researchers of cognitive processing in illiteracy have proposed that the acquisition of literacy ... more Researchers of cognitive processing in illiteracy have proposed that the acquisition of literacy modifies the functional organization of the brain. They have suggested that, while illiterate individuals have access only to innate semantic processing skills, those who have learned the correspondence between graphemes and phonemes have several mechanisms available to them through which to process oral language. We conducted 2 experiments to verify that suggestion with respect to language processing, and to elucidate further the differences between literate and illiterate individuals in the cognitive strategies used to process oral language, as well as hemispheric specialization for these processes. Our findings suggest that semantic processing strategies are qualitatively the same in literates and illiterates, despite the fact that overall performance is augmented by increased education. In contrast, explicit processing of oral information based on phonological characteristics appears to be qualitatively different between literates and illiterates: effective strategies in the processing of phonological information depend upon having had a formal education, regardless of the level of education. We also confirmed the differential abilities needed for the processing of semantic and phonological information and related them to hemisphere-specific processing. (JINS, 2004, 10, 818-827.)

Research paper thumbnail of Implicit structured sequence learning: an fMRI study of the structural mere-exposure effect

Frontiers in Psychology, 2014

In this event-related fMRI study we investigated the effect of 5 days of implicit acquisition on ... more In this event-related fMRI study we investigated the effect of 5 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. . Citation: Folia V and Petersson KM (2014) Implicit structured sequence learning: an fMRI study of the structural mere-exposure effect. Front. Psychol. 5:41.

Research paper thumbnail of Lexical Processing in Illiteracy: Effect of Literacy or Education?

Cortex, 2006

Difficulties in phonological processing in illiterates have been attributed to their limited phon... more Difficulties in phonological processing in illiterates have been attributed to their limited phonological awareness, a consequence of their lack of literacy. We sought to explore the potential influence of education on auditory lexical processing above and beyond literacy per se. In order to achieve this goal, we compared a lexical decision making paradigm with a repetition paradigm using words and pseudo-words. We based this choice of tasks on previous research, which has shown that pseudo-word repetition is dependent on the phonological loop; such studies have thus demonstrated a literacy effect on repetition. Instead, lexical decision making is known to depend on the size of one's vocabulary, which is influenced by the level of education attained. Our sample comprised three groups: illiterate no education, literate/low education and literate/high education, individuals. The pattern of our findings confirmed that literacy has an effect on the capacity of the phonological loop, as our illiterate group alone had difficulty with repetition, as compared with both literate/educated groups. Also, our findings suggested an education effect on lexical decision making, as we found a gradation in the performance of the three groups. Therefore, we succeeded in dissecting the effect of literacy and education on auditory lexical processing through the application and comparison of two simple paradigms.

Research paper thumbnail of The inferior frontal cortex in artificial syntax processing: An rTMS study

Brain Research, 2008

The human capacity to implicitly acquire knowledge of structured sequences has recently been inve... more 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. a v a i l a b l e a t w w w. s c i e n c e d i r e c t . c o m w w w. e l s e v i e r. c o m / l o c a t e / b r a i n r e s ARTICLE IN PRESS Please cite this article as: Uddén, J. et al., The inferior frontal cortex in artificial syntax processing: An rTMS study, Brain Res. (2008),

Research paper thumbnail of What artificial grammar learning reveals about the neurobiology of syntax

Brain and Language, 2012

In this paper we examine the neurobiological correlates of syntax, the processing of structured s... more 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.

Research paper thumbnail of Implicit Learning and Dyslexia

Annals of the New York Academy of Sciences, 2008

Several studies have reported an association between dyslexia and implicit learning deficits. It ... more 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. resentation and relate this to a visuographic input-output code. This is typically achieved by means of a supervised learning process (i.e., teaching), in contrast to natural language acquisition, which is largely a spontaneous, non-supervised, and self-organized acquisition process ). Aspects of language can also be an object of metalinguistic awareness: the intentional and explicit control over aspects of phonology, syntax, semantics,

Research paper thumbnail of The Neuropharmacology of Implicit Learning

Current Neuropharmacology, 2010

Two decades of pharmacologic research on the human capacity to implicitly acquire knowledge as we... more 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.