Brain Research in Language LITERACY STUDIES VOLUME 1 Series Editor: R. Malatesha Joshi,Texas A&M University, USA Editorial Board: Linnea Ehri, CUNY Graduate School, USA George Hynd, Purdue University, USA Richard Olson, University of Colorado, USA Pieter Reitsma, Vrije University Amsterdam, the Netherlands Rebecca Treiman, Washington University in St. Louis, USA Usha Goswami, University of Cambridge, UK Jane Oakhill, University of Sussex, Brighton, JK Philip Seymour, University of Dundee, UK Guinevere Eden, Georgetown University Medical Center, USA Catherine McBride Chang, Chinese University of Hong Kong, China While language defines humanity, literacy defines civilization. Understandably, illiteracy or difficulties in acquiring literacy skills have become a major concern of our technological society. A conservative estimate of the prevalence of literacy problems would put the figure at more than a billion people in the world. Because of the seriousness of the problem, research in literacy acquisition and its breakdown is pursued with enormous vigor and persistence by experts from diverse backgrounds such as cognitive psychology, neuroscience, linguistics and education. This, of course, has resulted in a plethora of data, and consequently it has become difficult to integrate this abundance of information into a coherent body because of the artificial barriers that exist among different professional specialties. The purpose of the proposed series is to bring together the available research studies into a coherent body of knowledge. Publications in this series are intended for use by educators, clinicians and research scientists in the above-mentioned specialties. Some of the titles suitable for the Series are: fMRI, brain imaging techniques and reading skills, orthography and literacy; and research based techniques for improving decoding, vocabulary, spelling, and comprehension skills. The titles published in this series are listed at the end of this volume. Zvia Breznitz (Editor) Brain Research in Language Zvia Breznitz University of Haifa Haifa 31905 Israel [email protected] Series Editor: R. Malatesha Joshi Texas A&M University College Station, TX 77843 USA Library of Congress Control Number: 2007935911 ISBN-13: 978-0-387-74979-2 e-ISBN-13: 978-0-387-74980-8 Printed on acid-free paper. (cid:164) 2008 Springer Science+Business Media, LLC. All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer Science+Business Media, LLC., 233 Spring Street, New York, NY 10013, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use in this publication of trade names, trademarks, service marks and similar terms, even if they are not identified as such, is not to be taken as an expression of opinion as to whether or not they are subject to proprietary rights. 9 8 7 6 5 4 3 2 1 springer.com CONTENTS Introduction: Brain Research in Language .............................................................1 Zvia Breznitz 1. Neurobiological Correlates of Dyslexia..............................................................7 Zvia Breznitz and Lilach Lebovitz 2. Event-Related Potentials (ERPs) in the Study of Dyslexia.............................51 Shelley Shaul 3. Auditory P2 is Reduced in 5 Month Old Infants from Dyslexic Families................................................................................................................93 Pieter H. Been, Theo H. van Leeuwen, Marieke van Herten, Ben Maassen, Aryan van der Leij and Frans Zwarts 4. Electrophysiological Functional Imaging of Auditory Processing in Humans.........................................................................................................111 Hillel Pratt 5. Overlapping Tasks Methodology as a Tool for Investigating Language Perception........................................................................................125 Werner Sommer and Annette Hohlfeld 6. A Sampler of Event-Related Brain Potential (ERP) Analyses of Language Processing...................................................................................153 Marta Kutas and Katherine A. Delong 7. Processing the Grammatical Function of Words in Sentence Reading..............................................................................................................187 Mark Leikin vi Contents 8. Timing the Metaphoric Brain.........................................................................205 Abraham Goldstein, Yossi Arzouan and Miriam Faust 9. EEG Correlates of Hemispheric Word Recognition.....................................225 Eran Zaidel, Andrew Hill and Scott Weems 10. Brain Mapping of Language Using High Frequency Gamma Oscillations in the Electrocorticography........................................................247 Alon Sinai Index........................................................................................................................275 BRAIN RESEARCH IN LANGUAGE Zvia Breznitz Neurocognitive Research Laboratory, Faculty of Education, University of Haifa, Haifa 31905 Israel The human brain has existed for approximately 60,000 years while the alphabetic code has been in use for only 5000 years. It can therefore be argued that the ability to read is not part of our evolutionary heritage, as no brain system was specifically developed for the purposes of reading. The reading process is a highly composite cognitive task, which relies on brain systems that were originally devoted to other functions. In most cases, the reading process is successful, but in some cases it is not. Reading involves the decoding and comprehension of printed materials. Word decoding implies the activation of different brain entities such as the visual and auditory modalities and the orthographic, phonological, and semantic systems. Each entity has a different function in the reading process, and each has a different biological structure. Each is activated in different brain areas, and each processes information in a different manner and at a different speed. In addition, as a cognitive process, reading is based on information processing mechanisms and requires highly complex information processing skills. As such, this cognitive activity varies along the different stages of the information processing system. During the input stage, attention, perception and discrimination of alphabetic symbols in the visual and acoustic modalities are required. In the processing stage, activation of these symbols in short term memory (STM) and working memory (WM) is necessary, as are interpretations and retrieval of alphabetic representations from long 2 Zvia Breznitz term memory (LTM) and the mental lexicon (ML). At this stage, the orthographic, phonological and semantic systems are activated. During the output stage, reading requires activation of various motor systems. This process ends with the reader’s responses. The complex nature of this process poses a major challenge for the human brain, which proves too much for some readers. The majority of studies in this area have implemented behavioral methodologies, which provide information concerning the entire cognitive sequence at the conclusion of processing only, in the reader’s output. The reader arrives at this stage following the completion of sensory, cognitive and motor processes (Bentin, 1989). As such, behavioral measures cannot specify all of the covert component opera- tions that contribute to reading, nor can they determine the relative processing times required by the individual stages. Furthermore, they cannot determine which processes occur serially, which occur in parallel and which overlap in time (Brandeis & Lehmann, 1994; Johnson, 1995). This makes it difficult to determine, on the basis of behavioral measures alone, the extent to which dysfunction or slowness at any particular stage of processing contributes to reading deficits. However, recent advancements in the field of neuroscience and cognitive development have added a new dimension with regard to the research into the universal and domain specific aspects of reading with the advent of innovative neurophysiological measurement tech- niques. The most common are electroencephalography (EEG) and functional magnetic resonance imaging (fMRI). These two methods provide researchers with the opportunity to examine, in-depth, the neural correlates of the reading processing with precise temporal and spatial resolutions, respectively. EEG techniques assess on-line processing of the cognitive activity utilizing Event Related Potential (ERP) methodology. This method permits direct observation of infor- mation processing at different levels of analysis, and can provide crucial information by means of real-time imaging of the neural system’s responses to sensory stimulation (Bentin, 1989). Thus, it enables us to trace on-line the speed at which information is processed during the various cognitive stages of the reading activity. The fMRI method focuses on the source of brain activity with time resolution that is not sufficient to trace temporal brain activation during the reading process. Even though these two methodologies are complimentary, it is important to note that precise time measurement Brain Research in Language 3 of temporal resolution is a critical factor in reading research, and as such, the advantage of EEG measurement is clear. Tracing the source of brain activity during reading is crucial for understanding both the accurate and impaired process and led various researchers to focus on developing new approaches for estimating the source localization of brain activity from ongoing EEG measures during performance of cognitive tasks such as reading, i.e., Low Resolution Electromagnetic Tomography (LORETA). The extent to which this estimation is similar to fMRI results remains unclear. The current book contains chapters based on a series of lectures given at the annual conference held by the Department of Learning Disabilities at the University of Haifa in June 2006 on “The Use of Brain Activity Measures in Reading Research.” The lectures presented data obtained from various studies employing behavioral, electro- physiological and imaging methodologies in different languages focusing on the regular reading process and the dyslexic population. The first two chapters review the use of imaging and electro- physiological techniques employing the Evoked Related Potential (ERP) methodology in reading research. Data from studies focusing on measuring the brain activity of regular and dyslexic readers is presented. The first contribution by Breznitz and Lebovitz offers evidence from post-mortem, structural and functional imaging studies in which the brains of dyslexic individuals are compared to those of regular readers. As phonological processing is a key concept in our under- standing of the regular and impaired reading process, the majority of studies in the field focus primarily on the nature of neural activity within the phonological system. Data concerning the effect of phonological intervention training on the brain activity of dyslexics are discussed. Moreover, based on the fact that reading is an act of perceiving printed materials from the printed page via the visual system, data from studies focusing on activation in the visual system are also presented. The second chapter contributed by Shaul and Breznitz surveys the different ERP components characteristic of brain activity during the reading process as well as the compelling results stemming from studies implementing this technique. The aim of this review is to gain a better understanding of the underlying causes of reading as a