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      A hierarchy of linguistic predictions during natural language comprehension

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          Significance

          Theorists propose that the brain constantly generates implicit predictions that guide information processing. During language comprehension, such predictions have indeed been observed, but it remains disputed under which conditions and at which processing level these predictions occur. Here, we address both questions by analyzing brain recordings of participants listening to audiobooks, and using a deep neural network to quantify the predictions evoked by the story. We find that brain responses are continuously modulated by linguistic predictions. We observe predictions at the level of meaning, grammar, words, and speech sounds, and find that high-level predictions can inform low-level ones. These results establish the predictive nature of language processing, demonstrating that the brain spontaneously predicts upcoming language at multiple levels of abstraction.

          Abstract

          Understanding spoken language requires transforming ambiguous acoustic streams into a hierarchy of representations, from phonemes to meaning. It has been suggested that the brain uses prediction to guide the interpretation of incoming input. However, the role of prediction in language processing remains disputed, with disagreement about both the ubiquity and representational nature of predictions. Here, we address both issues by analyzing brain recordings of participants listening to audiobooks, and using a deep neural network (GPT-2) to precisely quantify contextual predictions. First, we establish that brain responses to words are modulated by ubiquitous predictions. Next, we disentangle model-based predictions into distinct dimensions, revealing dissociable neural signatures of predictions about syntactic category (parts of speech), phonemes, and semantics. Finally, we show that high-level (word) predictions inform low-level (phoneme) predictions, supporting hierarchical predictive processing. Together, these results underscore the ubiquity of prediction in language processing, showing that the brain spontaneously predicts upcoming language at multiple levels of abstraction.

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          Most cited references100

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          FieldTrip: Open Source Software for Advanced Analysis of MEG, EEG, and Invasive Electrophysiological Data

          This paper describes FieldTrip, an open source software package that we developed for the analysis of MEG, EEG, and other electrophysiological data. The software is implemented as a MATLAB toolbox and includes a complete set of consistent and user-friendly high-level functions that allow experimental neuroscientists to analyze experimental data. It includes algorithms for simple and advanced analysis, such as time-frequency analysis using multitapers, source reconstruction using dipoles, distributed sources and beamformers, connectivity analysis, and nonparametric statistical permutation tests at the channel and source level. The implementation as toolbox allows the user to perform elaborate and structured analyses of large data sets using the MATLAB command line and batch scripting. Furthermore, users and developers can easily extend the functionality and implement new algorithms. The modular design facilitates the reuse in other software packages.
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            Nonparametric statistical testing of EEG- and MEG-data.

            In this paper, we show how ElectroEncephaloGraphic (EEG) and MagnetoEncephaloGraphic (MEG) data can be analyzed statistically using nonparametric techniques. Nonparametric statistical tests offer complete freedom to the user with respect to the test statistic by means of which the experimental conditions are compared. This freedom provides a straightforward way to solve the multiple comparisons problem (MCP) and it allows to incorporate biophysically motivated constraints in the test statistic, which may drastically increase the sensitivity of the statistical test. The paper is written for two audiences: (1) empirical neuroscientists looking for the most appropriate data analysis method, and (2) methodologists interested in the theoretical concepts behind nonparametric statistical tests. For the empirical neuroscientist, a large part of the paper is written in a tutorial-like fashion, enabling neuroscientists to construct their own statistical test, maximizing the sensitivity to the expected effect. And for the methodologist, it is explained why the nonparametric test is formally correct. This means that we formulate a null hypothesis (identical probability distribution in the different experimental conditions) and show that the nonparametric test controls the false alarm rate under this null hypothesis.
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              Predictive coding in the visual cortex: a functional interpretation of some extra-classical receptive-field effects.

              We describe a model of visual processing in which feedback connections from a higher- to a lower-order visual cortical area carry predictions of lower-level neural activities, whereas the feedforward connections carry the residual errors between the predictions and the actual lower-level activities. When exposed to natural images, a hierarchical network of model neurons implementing such a model developed simple-cell-like receptive fields. A subset of neurons responsible for carrying the residual errors showed endstopping and other extra-classical receptive-field effects. These results suggest that rather than being exclusively feedforward phenomena, nonclassical surround effects in the visual cortex may also result from cortico-cortical feedback as a consequence of the visual system using an efficient hierarchical strategy for encoding natural images.
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                Author and article information

                Journal
                Proc Natl Acad Sci U S A
                Proc Natl Acad Sci U S A
                pnas
                PNAS
                Proceedings of the National Academy of Sciences of the United States of America
                National Academy of Sciences
                0027-8424
                1091-6490
                3 August 2022
                9 August 2022
                3 February 2023
                : 119
                : 32
                : e2201968119
                Affiliations
                [1] aDonders Institute, Radboud University , 6525 EN Nijmegen, The Netherlands;
                [2] bMax Planck Institute for Psycholinguistics , 6525 XD Nijmegen, The Netherlands
                Author notes
                1To whom correspondence may be addressed. Email: micha.heilbron@ 123456donders.ru.nl .

                Edited by Stanislas Dehaene, Commissariat a l’ Énergie Atomique et aux Énergies Alternatives, Gif-sur-Yvette, France; received February 11, 2022; accepted June 28, 2022

                Author contributions: M.H., P.H., and F.P.d.L. designed research; M.H., K.A., and J.-M.S. performed research; M.H. contributed new reagents/analytic tools; M.H. analyzed data; and M.H., K.A., J.-M.S., P.H., and F.P.d.L. wrote the paper.

                Author information
                https://orcid.org/0000-0003-3039-4007
                https://orcid.org/0000-0003-0923-6610
                https://orcid.org/0000-0001-7280-7549
                https://orcid.org/0000-0002-6730-1452
                Article
                202201968
                10.1073/pnas.2201968119
                9371745
                35921434
                eedf898d-0a79-40f5-b5c3-5f69608e1071
                Copyright © 2022 the Author(s). Published by PNAS.

                This article is distributed under Creative Commons Attribution-NonCommercial-NoDerivatives License 4.0 (CC BY-NC-ND).

                History
                : 28 June 2022
                Page count
                Pages: 12
                Categories
                424
                431
                Biological Sciences
                Neuroscience
                Social Sciences
                Psychological and Cognitive Sciences

                language,prediction,eeg,meg,computational modeling
                language, prediction, eeg, meg, computational modeling

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