The P600 as a continuous index of integration effort.

The integration of word meaning into an unfolding utterance representation is a core operation of incremental language comprehension. There is considerable debate, however, as to which component of the ERP signal-the N400 or the P600-directly reflects integrative processes, with far reaching consequences for the temporal organization and architecture of the comprehension system. Multi-stream models maintaining the N400 as integration crucially rely on the presence of a semantically attractive plausible alternative interpretation to account for the absence of an N400 effect in response to certain semantic anomalies, as reported in previous studies. The single-stream Retrieval-Integration account posits the P600 as an index of integration, further predicting that its amplitude varies continuously with integrative effort. Here, we directly test these competing hypotheses using a context manipulation design in which a semantically attractive alternative is either available or not, and target word plausibility is varied across three levels. An initial self-paced reading study revealed graded reading times for plausibility, suggesting differential integration effort. A subsequent ERP study showed no N400 differences across conditions, and that P600 amplitude is graded for plausibility. These findings are inconsistent with the interpretation of the N400 as an index of integration, as no N400 effect emerged even in the absence of a semantically attractive alternative. By contrast, the link between plausibility, reading times, and P600 amplitude supports the view that the P600 is a continuous index of integration effort. More generally, our results support a single-stream architecture and eschew the need for multi-stream accounts.

Rational Redundancy in Referring Expressions: Evidence from Event-related Potentials.

In referential communication, Grice's Maxim of Quantity is thought to imply that utterances conveying unnecessary information should incur comprehension difficulties. There is, however, considerable evidence that speakers frequently encode redundant information in their referring expressions, raising the question as to whether such overspecifications hinder listeners' processing. Evidence from previous work is inconclusive, and mostly comes from offline studies. In this article, we present two event-related potential (ERP) experiments, investigating the real-time comprehension of referring expressions that contain redundant adjectives in complex visual contexts. Our findings provide support for both Gricean and bounded-rational accounts. We argue that these seemingly incompatible results can be reconciled if common ground is taken into account. We propose a bounded-rational account of overspecification, according to which even redundant words can be beneficial to comprehension to the extent that they facilitate the reduction of listeners' uncertainty regarding the target referent.

Retrieval (N400) and integration (P600) in expectation-based comprehension.

Expectation-based theories of language processing, such as Surprisal theory, are supported by evidence of anticipation effects in both behavioural and neurophysiological measures. Online measures of language processing, however, are known to be influenced by factors such as lexical association that are distinct from-but often confounded with-expectancy. An open question therefore is whether a specific locus of expectancy related effects can be established in neural and behavioral processing correlates. We address this question in an event-related potential experiment and a self-paced reading experiment that independently cross expectancy and lexical association in a context manipulation design. We find that event-related potentials reveal that the N400 is sensitive to both expectancy and lexical association, while the P600 is modulated only by expectancy. Reading times, in turn, reveal effects of both association and expectancy in the first spillover region, followed by effects of expectancy alone in the second spillover region. These findings are consistent with the Retrieval-Integration account of language comprehension, according to which lexical retrieval (N400) is facilitated for words that are both expected and associated, whereas integration difficulty (P600) will be greater for unexpected words alone. Further, an exploratory analysis suggests that the P600 is not merely sensitive to expectancy violations, but rather, that there is a continuous relation. Taken together, these results suggest that the P600, like reading times, may reflect a meaning-centric notion of Surprisal in language comprehension.

When components collide: Spatiotemporal overlap of the N400 and P600 in language comprehension.

The problem of spatiotemporal overlap between event-related potential (ERP) components is generally acknowledged in language research. However, its implications for the interpretation of experimental results are often overlooked. In a previous experiment on the functional interpretation of the N400 and P600, it was argued that a P600 effect to implausible words was largely obscured - in one of two implausible conditions - by an overlapping N400 effect of semantic association. In the present ERP study, we show that the P600 effect of implausibility is uncovered when the critical condition is tested against a proper baseline condition which elicits a similar N400 amplitude, while it is obscured when tested against a baseline condition producing an N400 effect. Our findings reveal that component overlap can result in the apparent absence or presence of an effect in the surface signal and should therefore be carefully considered when interpreting ERP patterns. Importantly, we show that, by factoring in the effects of spatiotemporal overlap between the N400 and P600 on the surface signal, which we reveal using rERP analysis, apparent inconsistencies in previous findings are easily reconciled, enabling us to draw unambiguous conclusions about the functional interpretation of the N400 and P600 components. Overall, our results provide compelling evidence that the N400 reflects lexical retrieval processes, while the P600 indexes compositional integration of word meaning into the unfolding utterance interpretation.

Neurobehavioral Correlates of Surprisal in Language Comprehension: A Neurocomputational Model.

Expectation-based theories of language comprehension, in particular Surprisal Theory, go a long way in accounting for the behavioral correlates of word-by-word processing difficulty, such as reading times. An open question, however, is in which component(s) of the Event-Related brain Potential (ERP) signal Surprisal is reflected, and how these electrophysiological correlates relate to behavioral processing indices. Here, we address this question by instantiating an explicit neurocomputational model of incremental, word-by-word language comprehension that produces estimates of the N400 and the P600-the two most salient ERP components for language processing-as well as estimates of "comprehension-centric" Surprisal for each word in a sentence. We derive model predictions for a recent experimental design that directly investigates "world-knowledge"-induced Surprisal. By relating these predictions to both empirical electrophysiological and behavioral results, we establish a close link between Surprisal, as indexed by reading times, and the P600 component of the ERP signal. The resultant model thus offers an integrated neurobehavioral account of processing difficulty in language comprehension.

Splitting event-related potentials: Modeling latent components using regression-based waveform estimation.

Event-related potentials (ERPs) provide a multidimensional and real-time window into neurocognitive processing. The typical Waveform-based Component Structure (WCS) approach to ERPs assesses the modulation pattern of components-systematic, reoccurring voltage fluctuations reflecting specific computational operations-by looking at mean amplitude in predetermined time-windows. This WCS approach, however, often leads to inconsistent results within as well as across studies. It has been argued that at least some inconsistencies may be reconciled by considering spatiotemporal overlap between components; that is, components may overlap in both space and time, and given their additive nature, this means that the WCS may fail to accurately represent its underlying latent component structure (LCS). We employ regression-based ERP (rERP) estimation to extend traditional approaches with an additional layer of analysis, which enables the explicit modeling of the LCS underlying WCS. To demonstrate its utility, we incrementally derive an rERP analysis of a recent study on language comprehension with seemingly inconsistent WCS-derived results. Analysis of the resultant regression models allows one to derive an explanation for the WCS in terms of how relevant regression predictors combine in space and time, and crucially, how individual predictors may be mapped onto unique components in LCS, revealing how these spatiotemporally overlap in the WCS. We conclude that rERP estimation allows for investigating how scalp-recorded voltages derive from the spatiotemporal combination of experimentally manipulated factors. Moreover, when factors can be uniquely mapped onto components, rERPs may offer explanations for seemingly inconsistent ERP waveforms at the level of their underlying latent component structure.

Event-related potentials index lexical retrieval (N400) and integration (P600) during language comprehension.

The functional interpretation of two salient language-sensitive ERP components - the N400 and the P600 - remains a matter of debate. Prominent alternative accounts link the N400 to processes related to lexical retrieval, semantic integration, or both, while the P600 has been associated with syntactic reanalysis or, alternatively, to semantic integration. The often overlapping predictions of these competing accounts in extant experimental designs, however, has meant that previous findings have failed to clearly decide among them. Here, we present an experiment that directly tests the competing hypotheses using a design that clearly teases apart the retrieval versus integration view of the N400, while also dissociating a syntactic reanalysis/reprocessing account of the P600 from semantic integration. Our findings provide support for an integrated functional interpretation according to which the N400 reflects context-sensitive lexical retrieval - but not integration - processes. While the observed P600 effects were not predicted by any account, we argue that they can be reconciled with the integration view, if spatio-temporal overlap of ERP components is taken into consideration.

On the predictability of event boundaries in discourse: An ERP investigation.

When reading a text describing an everyday activity, comprehenders build a model of the situation described that includes prior knowledge of the entities, locations, and sequences of actions that typically occur within the event. Previous work has demonstrated that such knowledge guides the processing of incoming information by making event boundaries more or less expected. In the present ERP study, we investigated whether comprehenders' expectations about event boundaries are influenced by how elaborately common events are described in the context. Participants read short stories in which a common activity (e.g., washing the dishes) was described either in brief or in an elaborate manner. The final sentence contained a target word referring to a more predictable action marking a fine event boundary (e.g., drying) or a less predictable action, marking a coarse event boundary (e.g., jogging). The results revealed a larger N400 effect for coarse event boundaries compared to fine event boundaries, but no interaction with description length. Between 600 and 1000 ms, however, elaborate contexts elicited a larger frontal positivity compared to brief contexts. This effect was largely driven by less predictable targets, marking coarse event boundaries. We interpret the P600 effect as indexing the updating of the situation model at event boundaries, consistent with Event Segmentation Theory (EST). The updating process is more demanding with coarse event boundaries, which presumably require the construction of a new situation model.

Teasing apart coercion and surprisal: Evidence from eye-movements and ERPs.

Previous behavioral and electrophysiological studies have presented evidence suggesting that coercion expressions (e.g., began the book) are more difficult to process than control expressions like read the book. While this processing cost has been attributed to a specific coercion operation for recovering an event-sense of the complement (e.g., began reading the book), an alternative view based on the Surprisal Theory of language processing would attribute the cost to the relative unpredictability of the complement noun in the coercion compared to the control condition, with no need to postulate coercion-specific mechanisms. In two experiments, monitoring eye-tracking and event-related potentials (ERPs), respectively, we sought to determine whether there is any evidence for coercion-specific processing cost above-and-beyond the difficulty predicted by surprisal, by contrasting coercing and control expressions with a further control condition in which the predictability of the complement noun was similar to that in the coercion condition (e.g., bought the book). While the eye-tracking study showed significant effects of surprisal and a marginal effect of coercion on late reading measures, the ERP study clearly supported the surprisal account. Overall, our findings suggest that the coercion cost largely reflects the surprisal of the complement noun with coercion specific operations possibly influencing later processing stages.

Non coding RNAs in aortic aneurysmal disease.

An aneurysm is a local dilatation of a vessel wall which is >50% its original diameter. Within the spectrum of cardiovascular diseases, aortic aneurysms are among the most challenging to treat. Most patients present acutely after aneurysm rupture or dissection from a previous asymptomatic condition and are managed by open surgical or endovascular repair. In addition, patients may harbor concurrent disease contraindicating surgical intervention. Collectively, these factors have driven the search for alternative methods of identifying, monitoring and treating aortic aneurisms using less invasive approaches. Non-coding RNA (ncRNAs) are emerging as new fundamental regulators of gene expression. The small microRNAs have opened the field of ncRNAs capturing the attention of basic and clinical scientists for their potential to become new therapeutic targets and clinical biomarkers for aortic aneurysm. More recently, long ncRNAs (lncRNAs) have started to be actively investigated, leading to first exciting reports, which further suggest their important and yet largely unexplored contribution to vascular physiology and disease. This review introduces the different ncRNA types and focus at ncRNA roles in aorta aneurysms. We discuss the potential of therapeutic interventions targeting ncRNAs and we describe the research models allowing for mechanistic studies and clinical translation attempts for controlling aneurysm progression. Furthermore, we discuss the potential role of microRNAs and lncRNAs as clinical biomarkers.