Publications

Abstract

An important, but often ignored distinction in the analysis of EEG signals is that between evoked activity and induced activity. Whereas evoked activity reflects the summation of transient post-synaptic potentials triggered by an event, induced activity, which is mainly oscillatory in nature, is thought to reflect changes in parameters controlling dynamic interactions within and between brain structures. We hypothesize that induced activity may yield information about the dynamics of cell assembly formation, activation and subsequent uncoupling, which may play a prominent role in different types of memory operations. We then describe a number of analysis tools that can be used to study the reactivity of induced rhythmic activity, both in terms of amplitude changes and of phase variability.

We briefly discuss how alpha, gamma and theta rhythms are thought to be generated, paying special attention to the hypothesis that the theta rhythm reflects dynamic interactions between the hippocampal system and the neocortex. This hypothesis would imply that studying the reactivity of scalp-recorded theta may provide a window on the contribution of the hippocampus to memory functions.

We review studies investigating the reactivity of scalp-recorded theta in paradigms engaging episodic memory, spatial memory and working memory. In addition, we review studies that relate theta reactivity to processes at the interface of memory and language. Despite many unknowns, the experimental evidence largely supports the hypothesis that theta activity plays a functional role in cell assembly formation, a process which may constitute the neural basis of memory formation and retrieval. The available data provide only highly indirect support for the hypothesis that scalp-recorded theta yields information about hippocampal functioning. It is concluded that studying induced rhythmic activity holds promise as an additional important way to study brain function.

Abstract

In two studies subjects were required to read Dutch sentences that in some cases contained a syntactic violation, in other cases a semantic violation. All syntactic violations were word category violations. The design excluded differential contributions of expectancy to influence the syntactic violation effects. The syntactic violations elicited an Anterior Negativity between 300 and 500 ms. This negativity was bilateral and had a frontal distribution. Over posterior sites the same violations elicited a P600/SPS starting at about 600 ms. The semantic violations elicited an N400 effect. The topographic distribution of the AN was more frontal than the distribution of the classical N400 effect, indicating that the underlying generators of the AN and the N400 are, at least to a certain extent, non-overlapping. Experiment 2 partly replicated the design of Experiment 1, but with differences in rate of presentation and in the distribution of items over subjects, and without semantic violations. The word category violations resulted in the same effects as were observed in Experiment 1, showing that they were independent of some of the specific parameters of Experiment 1. The discussion presents a tentative account of the functional differences in the triggering conditions of the AN and the P600/SPS.

Abstract

To understand spoken language requires that the brain provides rapid access to different kinds of knowledge, including the sounds and meanings of words, and syntax. Syntax specifies constraints on combining words in a grammatically well formed manner. Agrammatic patients are deficient in their ability to use these constraints, due to a lesion in the perisylvian area of the languagedominant hemisphere. We report a study on real-time auditory sentence processing in agrammatic comprehenders, examining
their ability to accommodate damage to the language system. We recorded event-related brain potentials (ERPs) in agrammatic comprehenders, nonagrammatic aphasics, and age-matched controls. When listening to sentences with grammatical violations, the agrammatic aphasics did not show the same syntax-related ERP effect as the two other subject groups. Instead, the waveforms of the agrammatic aphasics were dominated by a meaning-related ERP effect, presumably reflecting their attempts to achieve understanding by the use of semantic constraints. These data demonstrate that although agrammatic aphasics are impaired in their ability to exploit syntactic information in real time, they can reduce the consequences of a syntactic deficit by exploiting a semantic route. They thus provide evidence for the compensation of a syntactic deficit by a stronger reliance on another route in mapping
sound onto meaning. This is a form of plasticity that we refer to as multiple-route plasticity.

Abstract

Syntax is one of the components in the architecture of language processing that allows the listener/reader to bind single-word information into a unified interpretation of multiword utterances. This paper discusses ERP effects that have been observed in relation to syntactic processing. The fact that these effects differ from the semantic N400 indicates that the brain honors the distinction between semantic and syntactic binding operations. Two models of syntactic processing attempt to account for syntax-related ERP effects. One type of model is serial, with a first phase that is purely syntactic in nature (syntax-first model). The other type of model is parallel and assumes that information immediately guides the interpretation process once it becomes available. This is referred to as the immediacy model. ERP evidence is presented in support of the latter model. Next, an explicit computational model is proposed to explain the ERP data. This Unification Model assumes that syntactic frames are stored in memory and retrieved on the basis of the spoken or written word form input. The syntactic frames associated with the individual lexical items are unified by a dynamic binding process into a structural representation that spans the whole utterance. On the basis of a meta-analysis of imaging studies on syntax, it is argued that the left posterior inferior frontal cortex is involved in binding syntactic frames together, whereas the left superior temporal cortex is involved in retrieval of the syntactic frames stored in memory. Lesion data that support the involvement of this left frontotemporal network in syntactic processing are discussed.

Abstract

This study investigated the effects of combined semantic and syntactic violations in relation to the effects of single semantic and single syntactic violations on language-related event-related brain potential (ERP) effects (N400 and P600/ SPS). Syntactic violations consisted of a mismatch in grammatical gender or number features of the definite article and the noun in sentence-internal or sentence-final noun phrases (NPs). Semantic violations consisted of semantically implausible adjective–noun combinations in the same NPs. Combined syntactic and semantic violations were a summation of these two respective violation types. ERPs were recorded while subjects read the sentences with the different types of violations and the correct control sentences. ERP effects were computed relative to ERPs elicited by the sentence-internal or sentence-final nouns. The size of the N400 effect to the semantic violation was increased by an additional syntactic violation (the syntactic boost). In contrast, the size of the P600/ SPS to the syntactic violation was not affected by an additional semantic violation. This suggests that in the absence of syntactic ambiguity, the assignment of syntactic structure is independent of semantic context. However, semantic integration is influenced by syntactic processing. In the sentence-final position, additional global processing consequences were obtained as a result of earlier violations in the sentence. The resulting increase in the N400 amplitude to sentence-final words was independent of the nature of the violation. A speeded anomaly detection task revealed that it takes substantially longer to detect semantic than syntactic anomalies. These results are discussed in relation to the latency and processing characteristics of the N400 and P600/SPS effects. Overall, the results reveal an asymmetry in the interplay between syntax and semantics during on-line sentence comprehension.

Abstract

Mental imagery is a cognitive process crucial to human reasoning. Numerous studies have characterized specific
instances of this cognitive ability, as evoked by visual imagery (VI) or motor imagery (MI) tasks. However, it
remains unclear which neural resources are shared between VI and MI, and which are exclusively related to MI.
To address this issue, we have used fMRI to measure human brain activity during performance of VI and MI
tasks. Crucially, we have modulated the imagery process by manipulating the degree of mental rotation necessary
to solve the tasks. We focused our analysis on changes in neural signal as a function of the degree of mental
rotation in each task.

Abstract

Spoken language comprehension requires rapid integration of information from multiple linguistic sources. In the present study we addressed the temporal aspects of this integration process by focusing on the time course of the selection of the appropriate meaning of lexical ambiguities (“bank”) in sentence contexts. Successful selection of the contextually appropriate meaning of the ambiguous word is dependent upon the rapid binding of the contextual information in the sentence to the appropriate meaning of the ambiguity. We used the N400 to identify the time course of this binding process. The N400 was measured to target words that followed three types of context sentences. In the concordant context, the sentence biased the meaning of the sentence-final ambiguous word so that it was related to the target. In the discordant context, the sentence context biased the meaning so that it was not related to the target. In the unrelated control condition, the sentences ended in an unambiguous noun that was unrelated to the target. Half of the concordant sentences biased the dominant meaning, and the other half biased the subordinate meaning of the sentence-final ambiguous words. The ISI between onset of the target word and offset of the sentence-final word of the context sentence was 100 ms in one version of the experiment, and 1250 ms in the second version. We found that (i) the lexically dominant meaning is always partly activated, independent of context, (ii) initially both dominant and subordinate meaning are (partly) activated, which suggests that contextual and lexical factors both contribute to sentence interpretation without context completely overriding lexical information, and (iii) strong lexical influences remain present for a relatively long period of time.

Abstract

In two ERP experiments, we assessed the impact of discourse-level information on the processing of an unfolding spoken sentence. Subjects listened to sentences like Jane told her brother that he was exceptionally quick/slow, designed such that the alternative critical words were equally acceptable within the local sentence context. In Experiment 1, these sentences were embedded in a discourse that rendered one of the critical words anomalous (e.g. because Jane’s brother had in fact done something very quickly). Relative to the coherent alternative, these discourse-anomalous words elicited a standard N400 effect that started at 150–200 ms after acoustic word onset. Furthermore, when the same sentences were heard in isolation in Experiment 2, the N400 effect disappeared. The results demonstrate that our listeners related the unfolding spoken words to the wider discourse extremely rapidly, after having heard the first two or three phonemes only, and in many cases well before the end of the word. In addition, the identical nature of discourse- and sentence-dependent N400 effects suggests that from the perspective of the word-elicited comprehension process indexed by the N400, the interpretive context delineated by a single unfolding sentence and a larger discourse is functionally identical.

Abstract

In two experiments, we explored the use of event-related brain potentials to selectively track the processes that establish reference during spoken language comprehension. Subjects listened to stories in which a particular noun phrase like "the girl" either uniquely referred to a single referent mentioned in the earlier discourse, or ambiguously referred to two equally suitable referents. Referentially ambiguous nouns ("the girl" with two girls introduced in the discourse context) elicited a frontally dominant and sustained negative shift in brain potentials, emerging within 300–400 ms after acoustic noun onset. The early onset of this effect reveals that reference to a discourse entity can be established very rapidly. Its morphology and distribution suggest that at least some of the processing consequences of referential ambiguity may involve an increased demand on memory resources. Furthermore, because this referentially induced ERP effect is very different from that of well-known ERP effects associated with the semantic (N400) and syntactic (e.g., P600/SPS) aspects of language comprehension, it suggests that ERPs can be used to selectively keep track of three major processes involved in the comprehension of an unfolding piece of discourse.

Abstract

This study tests the recent claim that Broca’s aphasics are impaired in automatic lexical access, including the retrieval of word meaning. Subjects are required to perform a lexical decision on visually presented prime target pairs. Half of the word targets are preceded by a related word, half by an unrelated word. Primes and targets are presented with a long stimulus-onset-asynchrony (SOA) of 1400 msec and with a short SOA of 300 msec. Normal priming effects are observed in Broca’s aphasics for both SOAs. This result is discussed in the context of the claim that Broca’s aphasics suffer from an impairment in the automatic access of lexical–semantic information. It is argued that none of the current priming studies provides evidence supporting this claim, since with short SOAs priming effects have been reliably obtained in Broca’s aphasics. The results are more compatible with the claim that in many Broca’s aphasics the functional locus of their comprehension deficit is at the level of postlexical integration processes.

Abstract

Stimulus-related changes in cerebral blood oxygenation were measured using high-resolution functional magnetic resonance imaging sequentially covering visual occipital areas in contiguous sections. During dynamic imaging, healthy subjects silently viewed pseudowords, single false fonts, or length-matched strings of the same false fonts. The paradigm consisted of a sixfold alternation of an activation and a control task. With pseudowords as activation vs single false fonts as control, responses were seen mainly in medial occipital cortex. These responses disappeared when pseudowords were alternated with false font strings as the control and reappeared when false font strings instead of pseudowords served as activation and were alternated with single false fonts. The string-length contrast alone, therefore, is sufficient to account for the activation pattern observed in medial visual cortex when word-like stimuli are contrasted with single characters.

Abstract

In this study the N400 component of the event-related potential was used to investigate spoken sentence understanding in Broca's and Wernicke's aphasics. The aim of the study was to determine whether spoken sentence comprehension problems in these patients might result from a deficit in the on-line integration of lexical information. Subjects listened to sentences spoken at a normal rate. In half of these sentences, the meaning of the final word of the sentence matched the semantic specifications of the preceding sentence context. In the other half of the sentences, the sentence-final word was anomalous with respect to the preceding sentence context. The N400 was measured to the sentence-final words in both conditions. The results for the aphasic patients (n = 14) were analyzed according to the severity of their comprehension deficit and compared to a group of 12 neurologically unimpaired age-matched controls, as well as a group of 6 nonaphasic patients with a lesion in the right hemisphere. The nonaphasic brain damaged patients and the aphasic patients with a light comprehension deficit (high comprehenders, n = 7) showed an N400 effect that was comparable to that of the neurologically unimpaired subjects. In the aphasic patients with a moderate to severe comprehension deficit (low comprehenders, n = 7), a reduction and delay of the N400 effect was obtained. In addition, the P300 component was measured in a classical oddball paradigm, in which subjects were asked to count infrequent low tones in a random series of high and low tones. No correlation was found between the occurrence of N400 and P300 effects, indicating that changes in the N400 results were related to the patients' language deficit. Overall, the pattern of results was compatible with the idea that aphasic patients with moderate to severe comprehension problems are impaired in the integration of lexical information into a higher order representation of the preceding sentence context.

Abstract

The temporal properties of semantic and phonological processes in speech production were investigated in a new experimental paradigm using movement-related brain potentials. The main experimental task was picture naming. In addition, a 2-choice reaction go/no-go procedure was included, involving a semantic and a phonological categorization of the picture name. Lateralized readiness potentials (LRPs) were derived to test whether semantic and phonological information activated motor processes at separate moments in time. An LRP was only observed on no-go trials when the semantic (not the phonological) decision determined the response hand. Varying the position of the critical phoneme in the picture name did not affect the onset of the LRP but rather influenced when the LRP began to differ on go and no-go trials and allowed the duration of phonological encoding of a word to be estimated. These results provide electrophysiological evidence for early semantic activation and later phonological encoding.