Liina Pylkkänen

Tracking the time course of word recognition with MEG

Twenty years ago it was discovered that recognition of semantically unexpected words is associated with a special ERP signature - the N400. Pinpointing the precise functional significance of the N400 has, however, been difficult. Recent MEG studies of word processing reveal that, in fact, the N400 decomposes into several functionally distinct subcomponents, allowing for more fine-grained investigation of its significance.

Neuromagnetic Evidence for the Timing of Lexical Activation: An MEG Component Sensitive to Phonotactic Probability but Not to Neighborhood Density ☆

Evidence from electrophysiological measures such as ERPs (event-related potentials) and MEG (magnetoencephalography) suggest that the first evoked brain response component sensitive to stimulus properties affecting reaction times in word recognition tasks occurs at 300-400 ms. The present study used the stimulus manipulation of Vitevich and Luce (1999) to investigate whether the M350, an MEG response component peaking at 300-400 ms, reflects lexical or postlexical processing. Stimuli were simultaneously varied in phonotactic probability, which facilitates lexical activation, and in phonological neighborhood density, which inhibits the lexical decision process. The present results indicate that the M350 shows facilitation by phonotactic probability rather than inhibition by neighborhood density. Thus the M350 cannot be a postlexical component.

Towards a new neurobiology of language.

Theoretical advances in language research and the availability of increasingly high-resolution experimental techniques in the cognitive neurosciences are profoundly changing how we investigate and conceive of the neural basis of speech and language processing. Recent work closely aligns language research with issues at the core of systems neuroscience, ranging from neurophysiological and neuroanatomic characterizations to questions about neural coding. Here we highlight, across different aspects of language processing (perception, production, sign language, meaning construction), new insights and approaches to the neurobiology of language, aiming to describe promising new areas of investigation in which the neurosciences intersect with linguistic research more closely than before. This paper summarizes in brief some of the issues that constitute the background for talks presented in a symposium at the Annual Meeting of the Society for Neuroscience. It is not a comprehensive review of any of the issues that are discussed in the symposium.

The Representation of Polysemy: MEG Evidence

Most words in natural language are polysemous, that is, they can be used in more than one way. For example, paper can be used to refer to a substance made out of wood pulp or to a daily publication printed on that substance. Although virtually every sentence contains polysemy, there is little agreement as to how polysemy is represented in the mental lexicon. Do different uses of polysemous words involve access to a single representation or do our minds store distinct representations for each different sense? Here we investigated priming between senses with a combination of behavioral and magnetoencephalographic measures in order to test whether different senses of the same word involve identity or mere formal and semantic similarity. Our results show that polysemy effects are clearly distinct from similarity effects bilaterally. In the left hemisphere, sense-relatedness elicited shorter latencies of the M350 source, which has been hypothesized to index lexical activation. Concurrent activity in the right hemisphere, on the other hand, peaked later for sense-related than for unrelated target stimuli, suggesting competition between related senses. The obtained pattern of results supports models in which the representation of polysemy involves both representational identity and difference: Related senses connect to same abstract lexical representation, but are distinctly listed within that representation.

Simple Composition: A Magnetoencephalography Investigation into the Comprehension of Minimal Linguistic Phrases

The expressive power of language lies in its ability to construct an infinite array of ideas out of a finite set of pieces. Surprisingly, few neurolinguistic investigations probe the basic processes that constitute the foundation of this ability, choosing instead to focus on relatively complex combinatorial operations. Contrastingly, in the present work, we investigate the neural circuits underlying simple linguistic composition, such as required by the minimal phrase “red boat.” Using magnetoencephalography, we examined activity in humans generated at the visual presentation of target nouns, such as “boat,” and varied the combinatorial operations induced by its surrounding context. Nouns in minimal compositional contexts (“red boat”) were compared with those appearing in matched non-compositional contexts, such as after an unpronounceable consonant string (“xkq boat”) or within a list (“cup, boat”). Source analysis did not implicate traditional language areas (inferior frontal gyrus, posterior temporal regions) in such basic composition. Instead, we found increased combinatorial-related activity in the left anterior temporal lobe (LATL) and ventromedial prefrontal cortex (vmPFC). These regions have been linked previously to syntactic (LATL) and semantic (vmPFC) combinatorial processing in more complex linguistic contexts. Thus, we suggest that these regions play a role in basic syntactic and semantic composition, respectively. Importantly, the temporal ordering of the effects, in which LATL activity (∼225 ms) precedes vmPFC activity (∼400 ms), is consistent with many processing models that posit syntactic composition before semantic composition during the construction of linguistic representations.

An meg study of silent meaning

Although research on the neural bases of language has made significant progress on how the brain accesses the meanings of words, our understanding of sentence-level semantic composition remains limited. We studied the magnetoencephalography (MEG) responses elicited by expressions whose meanings involved an element not expressed in the syntax, which enabled us to investigate the brain correlates of semantic composition without confounds from syntactic composition. Sentences such as the author began the book, which asserts that an activity was begun although no activity is mentioned in the syntax, were contrasted with control sentences such as the author wrote the book, which involved no implicit meaning. These conditions were further compared with a semantically anomalous condition (the author disgusted the book). MEG responses to the object noun showed that silent meaning and anomaly are associated with distinct effects, silent meaning, but not anomaly, eliciting increased amplitudes in the anterior midline field (AMF) at 350450 msec. The AMF was generated in ventromedial prefrontal areas, usually implicated for social cognition and theory of mind. Our results raise the possibility that silent meaning interpretation may share mechanisms with these neighboring domains of cognition.

Sensitivity to syntax in visual cortex

One of the most intriguing findings on language comprehension is that violations of syntactic predictions can affect event-related potentials as early as 120 ms, in the same time-window as early sensory processing. This effect, the so-called early left-anterior negativity (ELAN), has been argued to reflect word category access and initial syntactic structure building (Friederici, 2002). In two experiments, we used magnetoencephalography to investigate whether (a) rapid word category identification relies on overt category-marking closed-class morphemes and (b) whether violations of word category predictions affect modality-specific sensory responses. Participants read sentences containing violations of word category predictions. Unexpected items varied in whether or not their word category was marked by an overt function morpheme. In Experiment 1, the amplitude of the visual evoked M100 component was increased for unexpected items, but only when word category was overtly marked by a function morpheme. Dipole modeling localized the generator of this effect to the occipital cortex. Experiment 2 replicated the main results of Experiment 1 and eliminated two non-morphology-related explanations of the M100 contrast we observed between targets containing overt category-marking and targets that lacked such morphology. Our results show that during reading, syntactically relevant cues in the input can affect activity in occipital regions at around 125 ms, a finding that may shed new light on the remarkable rapidity of language processing.

Syntactic Structure Building in the Anterior Temporal Lobe during Natural Story Listening.

The neural basis of syntax is a matter of substantial debate. In particular, the inferior frontal gyrus (IFG), or Brocas area, has been prominently linked to syntactic processing, but the anterior temporal lobe has been reported to be activated instead of IFG when manipulating the presence of syntactic structure. These findings are difficult to reconcile because they rely on different laboratory tasks which tap into distinct computations, and may only indirectly relate to natural sentence processing. Here we assessed neural correlates of syntactic structure building in natural language comprehension, free from artificial task demands. Subjects passively listened to Alice in Wonderland during functional magnetic resonance imaging and we correlated brain activity with a word-by-word measure of the amount syntactic structure analyzed. Syntactic structure building correlated with activity in the left anterior temporal lobe, but there was no evidence for a correlation between syntactic structure building and activity in inferior frontal areas. Our results suggest that the anterior temporal lobe computes syntactic structure under natural conditions.

Early Occipital Sensitivity to Syntactic Category Is Based on Form Typicality

Syntactic factors can rapidly affect behavioral and neural responses during language processing; however, the mechanisms that allow this rapid extraction of syntactically relevant information remain poorly understood. We addressed this issue using magnetoencephalography and found that an unexpected word category (e.g., “The recently princess . . . ”) elicits enhanced activity in visual cortex as early as 120 ms after exposure, and that this activity occurs as a function of the compatibility of a word’s form with the form properties associated with a predicted word category. Because no sensitivity to linguistic factors has been previously reported for words in isolation at this stage of visual analysis, we propose that predictions about upcoming syntactic categories are translated into form-based estimates, which are made available to sensory cortices. This finding may be a key component to elucidating the mechanisms that allow the extreme rapidity and efficiency of language comprehension.

Basic Linguistic Composition Recruits the Left Anterior Temporal Lobe and Left Angular Gyrus During Both Listening and Reading

Language is experienced primarily through one of two mediums--spoken words and written text. Although substantially different in form, these two linguistic vehicles possess similar powers of expression. Consequently, one goal for the cognitive neuroscience of language is to determine where, if anywhere, along the neural path from sensory stimulation to ultimate comprehension these two processing streams converge. In the present study, we investigate the relationship between basic combinatorial operations in both reading and listening. Using magnetoencephalography, we measured neural activity elicited by the comprehension of simple adjective-noun phrases (red boat) using the same linguistic materials and tasks in both modalities. The present paradigm deviates from previous cross-modality studies by investigating only basic combinatorial mechanisms--specifically, those evoked by the construction of simple adjective-noun phrases. Our results indicate that both modalities rely upon shared neural mechanisms localized to the left anterior temporal lobe (lATL) and left angular gyrus (lAG) during such processing. Furthermore, we found that combinatorial mechanisms subserved by these regions are deployed in the same temporal order within each modality, with lATL activity preceding lAG activity. Modality-specific combinatorial effects were identified during initial perceptual processing, suggesting top-down modulation of low-level mechanisms even during basic composition.