Christian Forkstam | Lund University (original) (raw)

Papers by Christian Forkstam

Cognitive Brain Research, 2002

There is much evidence for the existence of multiple memory systems. However, it has been argued ... more There is much evidence for the existence of multiple memory systems. However, it has been argued that tasks assumed to reflect different memory systems share basic processing components and are mediated by overlapping neural systems. Here we used multivariate analysis of PET-data to analyze similarities and differences in brain activity for multiple tests of working memory, semantic memory, and episodic memory. The results from two experiments revealed between-systems differences, but also between-systems similarities and within-system differences. Specifically, support was obtained for a task-general working-memory network that may underlie active maintenance. Premotor and parietal regions were salient components of this network. A common network was also identified for two episodic tasks, cued recall and recognition, but not for a test of autobiographical memory. This network involved regions in right inferior and polar frontal cortex, and lateral and medial parietal cortex. Several of these regions were also engaged during the working-memory tasks, indicating shared processing for episodic and working memory. Fact retrieval and synonym generation were associated with increased activity in left inferior frontal and middle temporal regions and right cerebellum. This network was also associated with the autobiographical task, but not with living / non-living classification, and may reflect elaborate retrieval of semantic information. Implications of the present results for the classification of memory tasks with respect to systems and / or processes are discussed.

In this study, we used event-related potentials (ERPs) to evaluate the contribution of surface co... more 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.

To clarify whether the neural pathways concerning color processing are the same for natural objec... more To clarify whether the neural pathways concerning color processing are the same for natural objects, for artifacts objects and for non-objects we examined brain responses measured with functional magnetic resonance imaging (FMRI) during a covert naming task including the factors color (color vs. black&white (B&W)) and stimulus type (natural vs. artifacts vs. non-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 objects and artifacts, but not for non-objects. When color objects (both natural and artifacts) were contrasted with color non-objects 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 can improve object recognition (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.

The first objective of this study was to compare the brain network engaged by preference classifi... more 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 repor-ted event-related fMRI data. The results show that preference and gramma-ticality 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 con-trolled by the FOXP2 transcription factor. CNTNAP2 is expressed in lang-uage related brain n...

Cognitive Science, 2004

Brain and Cognition, 2011

The Open Neuroimaging Journal, 2010

Cognitive Brain Research, 2002

Psychopharmacology, 2000

Prepulse inhibition of acoustic startle is the reduction in startle response to an intense audito... more Prepulse inhibition of acoustic startle is the reduction in startle response to an intense auditory stimulus when this stimulus is immediately preceded by a weaker prestimulus. Prepulse inhibition occurs normally in humans and experimental animals, but schizophrenic persons often exhibit a marked impairment in this measure. Previous studies have shown that dopamine (DA)-dependent neuronal mechanisms are involved in the modulation of prepulse inhibition. Experiments were conducted in rats to elucidate further the involvement of DA-ergic mechanisms in prepulse inhibition. In line with previous studies, the indirect DA agonist, amphetamine, was shown to decrease prepulse inhibition. A close reverse relationship over time between DA overflow in the nucleus accumbens and prepulse inhibition was obtained using a technique allowing concomitant measurement of these parameters in awake, freely moving rats. This effect was more pronounced in amphetamine-treated rats compared to rats treated with equimolar doses of cocaine, which increased DA overflow without affecting prepulse inhibition. In other experiments, the combined treatment with subthreshold doses of the selective DA D1 agonist, SKF 38393, and the selective DA D2 agonist, quinpirole, was also shown to decrease prepulse inhibition. Finally, the selective DA D2 antagonist, raclopride, was shown to enhance prepulse inhibition. In line with previous studies, it is concluded that DA neurotransmission is involved in the modulation of prepulse inhibition and that the ventral part of the mesostriatal DA system may serve an important role in this modulation. Furthermore, the possibility is discussed that the discrepant results on prepulse inhibition obtained with amphetamine and cocaine may disclose functionally relevant differences in their mechanisms of action, and that the enhancement of prepulse inhibition induced by some antipsychotics in rats may reflect their propensity to induce adverse mental effects in humans.

Neuropsychologia, 2003

Regions of the prefrontal cortex (PFC) are typically activated in many different cognitive functi... more Regions of the prefrontal cortex (PFC) are typically activated in many different cognitive functions. In most studies, the focus has been on the role of specific PFC regions in specific cognitive domains, but more recently similarities in PFC activations across cognitive domains have been stressed. Such similarities may suggest that a region mediates a common function across a variety of cognitive tasks. In this study, we compared the activation patterns associated with tests of working memory, semantic memory and episodic memory. The results converged on a general involvement of four regions across memory tests. These were located in left frontopolar cortex, left mid-ventrolateral PFC, left mid-dorsolateral PFC and dorsal anterior cingulate cortex. These findings provide evidence that some PFC regions are engaged during many different memory tests. The findings are discussed in relation to theories about the functional contribition of the PFC regions and the architecture of memory.

NeuroImage, 2006

The human brain supports acquisition mechanisms that extract structural regularities implicitly f... more The human brain supports acquisition mechanisms that extract structural regularities implicitly from experience without the induction of an explicit model. It has been argued that the capacity to generalize to new input is based on the acquisition of abstract representations, which reflect underlying structural regularities in the input ensemble. In this study, we explored the outcome of this acquisition mechanism, and to this end, we investigated the neural correlates of artificial syntactic classification using event-related functional magnetic resonance imaging. The participants engaged once a day during an 8-day period in a short-term memory acquisition task in which consonantstrings generated from an artificial grammar were presented in a sequential fashion without performance feedback. They performed reliably above chance on the grammaticality classification tasks on days 1 and 8 which correlated with a corticostriatal processing network, including frontal, cingulate, inferior parietal, and middle occipital/occipitotemporal regions as well as the caudate nucleus. Part of the left inferior frontal region (BA 45) was specifically related to syntactic violations and showed no sensitivity to local substring familiarity. In addition, the head of the caudate nucleus correlated positively with syntactic correctness on day 8 but not day 1, suggesting that this region contributes to an increase in cognitive processing fluency. D

Proceedings of the …, 2009

Human implicit learning can be investigated with implicit artificial grammar learning, a simple m... more 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 ...

Proceedings of the Cognitive Science …, 2005

In this paper we investigate the neural correlates of syntactic classification of an acquired gra... more In this paper we investigate the neural correlates of syntactic classification of an acquired grammatical sequence structure in an event-related FMRI study. During acquisition, participants were engaged in an implicit short-term memory task without performance feedback. We manipulated the statistical frequency-based and rule-based characteristics of the classification stimuli independently in order to investigate their role in artificial grammar acquisition. The participants performed reliably above chance on the classification task. We observed a partly overlapping corticostriatal processing network activated by both manipulations including inferior prefrontal, cingulate, inferior parietal regions, and the caudate nucleus. More specifically, the left inferior frontal BA 45 and the caudate nucleus were sensitive to syntactic violations and endorsement, respectively. In contrast, these structures were insensitive to the frequency-based manipulation.

Proceedings of the …, 2009

A dominant hypothesis in empirical research on the evolution of language is the following: the fu... more 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.

Current Opinion in Neurology, 2005

The human brain supports acquisition mechanisms that can extract structural regularities implicit... more The human brain supports acquisition mechanisms that can extract structural regularities implicitly from experience without the induction of an explicit model. Reber defined the process by which an individual comes to respond appropriately to the statistical structure of the input ensemble as implicit learning. He argued that the capacity to generalize to new input is based on the acquisition of abstract representations that reflect underlying structural regularities in the acquisition input. We focus this review of the implicit learning literature on studies published during 2004 and 2005. We will not review studies of repetition priming (&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;#39;implicit memory&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;#39;). Instead we focus on two commonly used experimental paradigms: the serial reaction time task and artificial grammar learning. Previous comprehensive reviews can be found in Seger&amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;amp;#39;s 1994 article and the Handbook of Implicit Learning. Emerging themes include the interaction between implicit and explicit processes, the role of the medial temporal lobe, developmental aspects of implicit learning, age-dependence, the role of sleep and consolidation. The attempts to characterize the interaction between implicit and explicit learning are promising although not well understood. The same can be said about the role of sleep and consolidation. Despite the fact that lesion studies have relatively consistently suggested that the medial temporal lobe memory system is not necessary for implicit learning, a number of functional magnetic resonance studies have reported medial temporal lobe activation in implicit learning. This issue merits further research. Finally, the clinical relevance of implicit learning remains to be determined.

Cognitive Science, 2004

In the present study, using event-related functional magnetic resonance imaging, we investigated ... more In the present study, using event-related functional magnetic resonance imaging, we investigated a group of participants on a grammaticality classification task after they had been exposed to well-formed consonant strings generated from an artificial regular grammar. We used an implicit acquisition paradigm in which the participants were exposed to positive examples. The objective of this study was to investigate whether brain regions related to language processing overlap with the brain regions activated by the grammaticality classification task used in the present study. Recent meta-analyses of functional neuroimaging studies indicate that syntactic processing is related to the left inferior frontal gyrus (Brodmann's areas 44 and 45) or Broca's region. In the present study, we observed that artificial grammaticality violations activated Broca's region in all participants. This observation lends some support to the suggestions that artificial grammar learning represents a model for investigating aspects of language learning in infants [TICS 4 (2000) 178] and adults [

Brain Research, 2008

Human implicit learning can be investigated with implicit artificial grammar learning, a paradigm... more 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 forcedchoice preference classification is equivalent to the typical grammaticality classification. ava 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 ev i e r. c o m / l o c a t e / b r a i n r e s

Cognitive Brain Research, 2002

Cognitive Science, 2004

Brain and Cognition, 2011

The Open Neuroimaging Journal, 2010

Cognitive Brain Research, 2002

Psychopharmacology, 2000

Neuropsychologia, 2003

NeuroImage, 2006

Proceedings of the …, 2009

Proceedings of the Cognitive Science …, 2005

Proceedings of the …, 2009

Current Opinion in Neurology, 2005

Cognitive Science, 2004

Brain Research, 2008