Cognitive Neuroscience of Language

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“This is an excellent, research-based and evidence-based textbook for cognitive neuroscience of language.”
—Li Hsieh, Associate Professor, Communication Sciences & Disorders, Wayne State University

“A big pro is that this book does not assume much background in either linguistics or cognitive science. I like it
that most chapters start with a discussion of different theoretical viewpoints, and then discuss evidence pros and
cons using data from a wide range of cognitive neuroscience studies.”
—Edith Kaan, Associate Professor, Department of Linguistics, University of Florida

“As a linguist with background but not expertise in psychology, I find the material immensely valuable. Mostly
because it presents an overview of lots of research in cognitive neuroscience and filters that research in an under-
standable way so that the big picture can be perceived. And it relates the cognitive neuroscience research directly
to linguistics in a way that does justice to both fields. It is admirable that the author has such command over two
very different fields.”
—Nancy Hedberg, Professor, Linguistics, Simon Fraser University

“Definitely comprehensive, and careful about framing current debates in a literature that is in many places not
very mature. I think that Kemmerer’s expertise as a linguist and as a cognitive neuroscientist really shines through
making this a valuable text that is informed by broad and deep understanding.”
—Laura Kertz, Assistant Professor, Cognitive, Linguistic & Psychological Sciences, Brown University
Cognitive Neuroscience
of Language

Language is one of our most precious and uniquely human capacities, so it is not surprising that research on its
neural substrates has been advancing quite rapidly in recent years. Until now, however, there has not been a single
introductory textbook that focuses specifically on this topic.
Cognitive Neuroscience of Language fills that gap by providing an up-to-date, wide-ranging, and pedagogi-
cally practical survey of the most important developments in the field. It guides students through all of the major
areas of investigation, beginning with fundamental aspects of brain structure and function, and then proceed-
ing to cover aphasia syndromes, the perception and production of speech, the processing of language in written
and signed modalities, the meanings of words, and the formulation and comprehension of complex expressions,
including grammatically inflected words, complete sentences, and entire stories.
Drawing heavily on prominent theoretical models, the core chapters illustrate how such frameworks are sup-
ported, and sometimes challenged, by experiments employing diverse brain mapping techniques. Although much
of the content is inherently challenging and intended primarily for graduate or upper-level undergraduate students,
it requires no previous knowledge of either neuroscience or linguistics, defining technical terms and explaining
important principles from both disciplines along the way.

David Kemmerer is a professor with a joint appointment in the Department of Speech, Language, and Hearing
Sciences and the Department of Psychological Sciences at Purdue University. He is also an affiliate of the
Interdepartmental Linguistics Program at Purdue. In addition, he has an adjunct appointment in the Division of
Behavioral Neurology and Cognitive Neuroscience at the University of Iowa, where he worked as a postdoctoral
fellow before becoming a faculty member at Purdue. He has taught courses on the neural bases of speech and
language, the broader field of cognitive neuroscience, and diverse topics in linguistics. Professor Kemmerer’s
research focuses on how different kinds of linguistic meaning are mediated by different neural systems, drawing on
behavioral and lesion data from brain-damaged patients as well as behavioral, electrophysiological, and functional
neuroimaging data from healthy subjects. He has published over 40 journal articles and book chapters describing
various aspects of his research.
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Cognitive Neuroscience
of Language
David Kemmerer
First published 2015
by Psychology Press
711 Third Avenue, New York, NY 10017
and by Psychology Press
27 Church Road, Hove, East Sussex BN3 2FA
Psychology Press is an imprint of the Taylor & Francis Group, an informa business
© 2015 Taylor & Francis
The right of David Kemmerer to be identified as author of this work has been
asserted by him in accordance with sections 77 and 78 of the Copyright, Designs
and Patents Act 1988.
All rights reserved. No part of this book may be reprinted or reproduced or utilised
in any form or by any electronic, mechanical, or other means, now known or
hereafter invented, including photocopying and recording, or in any information
storage or retrieval system, without permission in writing from the publishers.
Trademark notice: Product or corporate names may be trademarks or registered
trademarks, and are used only for identification and explanation without intent to
infringe.
Every effort has been made to contact copyright holders. Where this has not been
possible, we would be pleased to hear from the parties concerned.
Library of Congress Cataloging in Publication Data
Kemmerer, David L.
Cognitive neuroscience of language / by David Kemmerer.
pages cm
Includes bibliographical references and index.
1. Cognitive neuroscience. 2. Language acquisition. I. Title.
QP360.5.K455 2014
612.8′233–—dc23
2014017766

ISBN: 978-1-84872-620-8 (hbk)

ISBN: 978-1-84872-621-5 (pbk)
ISBN: 978-1-315-76406-1 (ebk)

Typeset in Galliard
by Book Now Ltd, London
Contents

Preface xix
Acknowledgments xxi

PART I FUNDAMENTALS 1
1 The Human Brain 3
2 Brain Mapping Methods 29

PART II APHASIA 69
3 Classic Aphasia Syndromes 71
4 Primary Progressive Aphasia Syndromes 93

PART III THE PERCEPTION AND PRODUCTION OF SPEECH 109

5 Speech Perception 111
6 Speech Production 145
7 Prosody 189

PART IV OTHER MODALITIES OF LANGUAGE USE 213

8 Reading and Writing 215
9 Sign Language 247

PART V THE MEANINGS OF WORDS 271

10 Object Nouns 273
11 Action Verbs 309
12 Abstract Words 335

PART VI MORPHOLOGY, SYNTAX, AND DISCOURSE 361

13 Morphology 363
14 Sentence Production 391
15 Sentence Comprehension 419
16 Discourse 471

References 489
Author Index 555
Subject Index 583
Page Intentionally Left Blank
Detailed Contents

PART I FUNDAMENTALS 1
1 The Human Brain 3
Introduction 3
Neurons: The Basic Signaling Units of the Brain 4
Anatomy 5
Physiology 6
Representation 8
Navigating the Neural Landscape 9
Building the Brain from the Bottom Up 10
Brainstem and Thalamus 10
Hippocampus and Amygdala 11
Basal Ganglia and Cerebellum 12
The Cerebral Cortex 13
The Major Lobes–—Visible and Hidden 14
Gyral–Sulcal Organization 14
Cytoarchitectonic Organization 15
Connectional Organization 20
Sensory, Motor, and Higher-Order Systems 21
Box 1.1: Born for Phonetics? 23
Language-Related Regions: Broca’s Area, Wernicke’s Area,
and Beyond 25
Summary and Key Points 26
Recommended Reading 28

2 Brain Mapping Methods 29

Introduction 29
Neuropsychology 30
Single and Double Dissociations 30
Groups and Individuals 33
Visualizing the Brain: The Basic Mechanics of Magnetic
Resonance Imaging 34
Types of Brain Damage 35
Stroke 36
Traumatic Brain Injury 36
x Detailed Contents

Neurodegenerative and Infectious Diseases 37

Tumors 38
Relationships Between Behavioral Data and Lesion Data 39
Lesion Overlap and Subtraction Analysis 39
Voxel-Based Lesion–Symptom Mapping (VLSM) 40
A Few Caveats About Neuropsychological Research on Structure–Function
Relationships 41
Functional Neuroimaging 43
Two Techniques 44
Positron Emission Tomography (PET) 44
Functional Magnetic Resonance Imaging (f MRI) 44
Standardized Three-Dimensional Coordinates for Defining Stereotaxic Brain Space 46
Box 2.1: What’s in a Voxel? 47
Blocked Versus Event-Related Designs 48
Some Basic Experimental Paradigms 49
Subtraction 49
Correlation 51
Multivariate Pattern Analysis 54
Electrophysiology 56
Stimulation 56
Recording 58
Intracranial 58
Extracranial 60
Box 2.2: Magnetoencephalography 62
Transcranial Magnetic Stimulation 63
How It Works 63
Applications to Language 65
Major Strengths and Weaknesses of the Different Methods 66
Summary and Key Points 66
Recommended Reading 68

PART II APHASIA 69
3 Classic Aphasia Syndromes 71
Introduction 71
Historical Background 72
Box 3.1: Handedness and Language Lateralization 73
How Should Different Types of Aphasia be Classified? 73
Broca’s Aphasia 76
Production 76
Comprehension 78
Repetition 78
Lesion Correlates 78
Box 3.2: New Analyses of the Brains of Broca’s Original Patients 79
Wernicke’s Aphasia 80
Production 80
Box 3.3: A Rare Case of Acute, Selective, and Temporary Dysfunction of Broca’s Area 81
 Detailed Contents xi

Comprehension 82
Repetition 83
Lesion Correlates 83
Conduction Aphasia 84
Production 84
Comprehension 85
Repetition 85
Lesion Correlates 85
Global Aphasia 86
Anomic Aphasia 86
Production 86
Comprehension 88
Repetition 88
Lesion Correlates 88
Transcortical Aphasia Syndromes 88
Transcortical Motor Aphasia 88
Transcortical Sensory Aphasia 89
Mixed Transcortical Aphasia 90
Flow Chart for Classifying the Aphasia Syndromes 90
Box 3.4: Aphasia in Bilinguals and Polyglots 91
Summary and Key Points 91
Recommended Reading 92

4 Primary Progressive Aphasia Syndromes 93

Introduction 93
Historical Background 94
Progressive Nonfluent/Agrammatic Aphasia (PNFA) 96
Production 96
Comprehension 97
Repetition 97
Lesion Correlates and Other Biomarkers 97
Associated Neurological Deficits 99
Box 4.1: Creativity in Art and Music: The Positive Side of PPA? 99
Semantic Dementia (SD) 101
Production 101
Comprehension 102
Repetition 103
Lesion Correlates and Other Biomarkers 103
Associated Neurological Deficits 104
Logopenic Progressive Aphasia (LPA) 104
Production 104
Comprehension 105
Repetition 105
Lesion Correlates and Other Biomarkers 105
Associated Neurological Deficits 105
PPA as a Window on the Neurobiology of Language 106
Summary and Key Points 107
Recommended Reading 107
xii Detailed Contents

PART III THE PERCEPTION AND PRODUCTION OF SPEECH 109

5 Speech Perception 111
Introduction 111
Box 5.1: Some Basic Properties of Speech Sounds 111
Early Cortical Stages of Speech Perception 113
Hierarchical Organization 114
Box 5.2: From Cochlea to Cortex 115
Bilateral Organization 117
Both Hemispheres Contribute to Speech Perception 117
The Two Hemispheres Have Partially Different Temporal Windows for
Speech Perception 122
Box 5.3: The Neural Substrates of Auditory–Visual Integration During Speech Perception:
A Combined fMRI and TMS Study of the McGurk Effect 125
Summary 127
A Double Dissociation Between Comprehension and Repetition: Initial Evidence for
Separate Streams of Speech Processing 127
The Ventral “What” Stream: From Sound to Meaning 130
The Lexical Interface 130
The Combinatorial Network 133
Summary 134
The Dorsal “How” Stream: From Sound to Action 135
The Sensorimotor Interface 135
The Articulatory Network 138
Box 5.4: Might Articulatory Activation During Speech Perception Facilitate
Turn-Taking? 142
Summary 143
Summary and Key Points 144
Recommended Reading 144

6 Speech Production 145

Introduction 145
The Lemma Model of Lexical Selection and Form Encoding 146
Box 6.1: “Close to Scientific Paradise” 146
Box 6.2: “Where Wings Take Dream” 147
The Architecture of the Model 148
Conceptual Focusing and Perspective-Taking 148
Lemma Selection 151
Interlude: Crossing the Rift 152
Retrieving Morphemic and Phonological Codes 152
Prosodification and Syllabification 154
Phonetic Encoding and Articulation 154
Self-Monitoring 155
Neurobiological Evidence for the Model 155
A Meta-Analysis of 82 Brain Mapping Experiments Involving
Word Production 155
Conceptual Focusing and Perspective-Taking 157
Lemma Selection 159
 Detailed Contents xiii

Box 6.3: Happy Faces Are Named Faster than Neutral Faces 160
Retrieving Morphemic and Phonological Codes 162
Prosodification and Syllabification 164
Phonetic Encoding and Articulation 165
Self-Monitoring 165
Some Challenges Facing the Model 166
The Lemma Dilemma 166
Discrete Versus Interactive Processing 167
The DIVA Model of Speech Motor Control 168
How Do You Say “Good Doggie”? 168
Feedforward Control 171
Box 6.4: A Brain–Machine Interface Restores Rudimentary Speech in a
Patient with Locked-In Syndrome 173
Box 6.5: What the Brain Does Before the Tongue Slips 174
Box 6.6: When the Will Is Gone 175
A Quick Look at the Roles of Forward and Inverse Models in Motor Control 176
Auditory Feedback Control 177
Somatosensory Feedback Control 179
Box 6.7: Using the DIVA Model to Simulate Stuttering 180
Some Challenges Facing the Model 181
The Island of Reil 181
Are the Auditory and Somatosensory Feedback Circuits Integrated
in the Planum Temporale? 183
Peripheral Mechanisms of Speech Production 183
Summary and Key Points 186
Recommended Reading 188

7 Prosody 189
Introduction 189
Emotional Prosody 190
Perception 190
The Right Mid to Anterior Superior Temporal Cortex: Auditory Integration 191
The Amygdala: Relevance Detection 192
The Right Ventral Frontoparietal Cortex: Emotion Simulation 192
The Basal Ganglia: Emotion Simulation, Sequence Decoding, and/or
Response Triggering 194
The Bilateral Orbitofrontal and Inferior Frontal Cortices: Cognitive Evaluation 195
Summary 198
Production 198
Box 7.1: Sex Differences in the Perception of Emotional Prosody 199
Linguistic Prosody 201
Perception 201
The Syntactic Domain 202
The Lexical Domain 204
The Tonal Domain 206
Box 7.2: Correlations Between the Geographic Distributions of Tone Languages and
Genes for Brain Development 208
xiv Detailed Contents

Summary 209
Production 209
Summary and Key Points 211
Recommended Reading 212

PART IV OTHER MODALITIES OF LANGUAGE USE 213

8 Reading and Writing 215
Introduction 215
Reading 216
Hierarchical Coding of Letter Strings in the Ventral Occipitotemporal Cortex 217
Box 8.1: Reading Printed Letters Activates the Motor Region
for Writing the Same Letters 219
The Visual Word Form Area (VWFA) 222
Normal Response Properties 222
Effects of Damage 223
Developmental Origins: The Neuronal Recycling Hypothesis 226
From Print to Sound and Meaning 229
A Cognitive Model 229
Evidence from Acquired Dyslexia 230
Neural Substrates 231
Summary 235
Writing 236
From Sound and Meaning to Print 236
A Cognitive Model 236
Evidence from Acquired Dysgraphia 238
Neural Substrates 239
Summary 242
Summary and Key Points 243
Recommended Reading 245

9 Sign Language 247

Introduction 247
Structural Aspects of Sign Language 248
Phonology 248
Morphology 250
Syntax 251
Nonmanual Signs 251
Summary 252
Left-Hemisphere Dominance 252
Wada Testing 252
Dissociations Between Sign Language and Visuospatial Cognition 253
Dissociations Between Sign Language and Symbolic Gesture 254
Summary 256
Functional–Anatomical Organization Within the Left Hemisphere 256
Different Sign Language Aphasias Associated with Anterior and Posterior Lesions 256
The Production of Signs 259
The Perception of Signs 260
Box 9.1: The Plasticity of Left-Hemisphere “Auditory” Areas in Congenitally
Deaf Brains Reflects Sign Language Rather than Sensory Deprivation 262
Summary 263
 Detailed Contents xv

Right-Hemisphere Contributions 263

Activation During Sentence Comprehension: A Mysterious and
Controversial Phenomenon 263
Involvement in Classifier Constructions 265
Summary 268
Summary and Key Points 269
Recommended Reading 269

PART V THE MEANINGS OF WORDS 271

10 Object Nouns 273

Introduction 273
Perceptual and Motor Features of Object Concepts 274
Theoretical Background 274
Color Features 276
Box 10.1: What Is a Violin? 276
Shape Features 278
Motion Features 280
Motor Features 281
Auditory Features 282
Gustatory and Olfactory Features 284
Summary 285
A Semantic Hub for Object Concepts 286
Box 10.2: The Concept of a Nest in the Brain of a Mouse 287
Evidence from Semantic Dementia 289
Evidence from fMRI and TMS 292
Summary 295
Domains of Object Concepts 296
Three Major Domains of Selective Semantic Impairment 296
Animal Concepts 296
Box 10.3: The Influences of Gender and Culture on Concepts for Animals and
Fruits/Vegetables 297
Fruit/Vegetable Concepts 300
Tool Concepts 301
Explanatory Approaches 303
Summary 306
Summary and Key Points 306
Recommended Reading 308

11 Action Verbs 309

Introduction 309
Perceptual and Motor Features of Action Concepts 310
Visual Features 310
Motor Features 315
Box 11.1: Hockey Players Do It Better 318
Summary 322
A Semantic Hub for Action Concepts 322
Domains of Action Concepts 325
xvi Detailed Contents

Transitive and Intransitive Verbs 325

Involvement of Temporal and Parietal Regions 326
Involvement of Broca’s Area 329
Box 11.2: Do Action Concepts Have a Default Directionality? 331
Summary 332
Summary and Key Points 332
Recommended Reading 333

12 Abstract Words 335

Introduction 335
Cognitive and Neural Distinctions Between Concrete and Abstract Concepts 335
Theoretical Background 335
Box 12.1: Do Abstract Concepts Have Metaphorical Foundations? 336
Evidence from PET and fMRI 339
Evidence from Neuropsychology and rTMS 340
Summary 343
A Semantic Hub for Abstract Concepts 343
Evidence from Semantic Dementia 344
Evidence from rTMS 350
Summary 351
Domains of Abstract Concepts 351
Emotions 351
Box 12.2: Good and Bad in Right- and Left-Handers 355
Numbers 356
Summary 359
Summary and Key Points 359
Recommended Reading 360

PART VI MORPHOLOGY, SYNTAX, AND DISCOURSE 361

13 Morphology 363
Introduction 363
Box 13.1: How to Build a Noun in Tabasaran 364
Morphosyntactic and Morphophonological Aspects of Inflection 365
Sahin et al.’s (2006) fMRI Study 366
Sahin et al.’s (2009) Intracranial Electrophysiological Study 368
Summary 371
Noun and Verb Inflection: A Closer Look at Morphosyntax 371
A Common Pathway in Broca’s Area 371
A Neuropsychological Double Dissociation 373
Neural Correlates of Noun-Specific and Verb-Specific Morphosyntactic Processing 375
What About Comprehension? 376
Summary 378
Regular and Irregular Inflection: A Closer Look at Morphophonology 378
Theoretical Background 378
Box 13.2: Connectionist Computer Simulations of the Regular/Irregular
Distinction 380
The Perspective from Neuropsychology 382
 Detailed Contents xvii

Box 13.3: Do the Basal Ganglia Contribute to Regular Inflection? 383

The Perspective from Functional Neuroimaging 385
What About Comprehension? 385
Summary 388
Summary and Key Points 388
Recommended Reading 389

14 Sentence Production 391

Introduction 391
Syntax: A Short Tutorial 392
Devices for Indicating Participant Roles 392
Hierarchical Structure 393
Argument Structure 393
Closed-Class Elements 395
Complex Sentences 395
Syntactic Encoding: A Rudimentary Model 395
Insights from Neuropsychology 398
Agrammatism: A Complex Syndrome Steeped in Controversy 398
Box 14.1: Similar Symptoms, Different Causes 401
Sentence Production Deficits in Vascular Aphasia 403
Deficits Involving the Functional Level 403
Deficits Involving the Positional Level 406
Sentence Production Deficits in Primary Progressive Aphasia 407
Summary 410
Box 14.2: Selection for Position: A Possible Contribution of Broca’s Area to Syntactic Encoding 411
Insights from PET and fMRI 412
Indefrey et al.’s (2001, 2004) PET Studies 412
Haller et al.’s (2005) fMRI Study 415
Summary 416
Summary and Key Points 417
Recommended Reading 417

15 Sentence Comprehension 419

Introduction 419
A Large-Scale Neural Network for Sentence Comprehension: Anatomical Organization 420
Lesion and Connectivity Studies by Dronkers et al. (2004) and Turken and
Dronkers (2011) 421
Dronkers et al.’s (2004) Lesion Study 421
Turken and Dronkers’ (2011) Connectivity Study 424
Convergent Results from Other Studies 427
Summary 429
A Large-Scale Neural Network for Sentence Comprehension: Functional Considerations 429
Possible Contributions of the pMTG 430
Snijders et al.’s (2009) Investigation 431
Tyler et al.’s (2011) and Papoutsi et al.’s (2011) Investigations 432
Summary 435
Possible Contributions of the aSTG and Some Adjacent Regions 435
Syntactic Analysis 436
A Combinatorial Syntactic–Semantic Network 438
xviii Detailed Contents

Some Challenging Data from Neuropsychology 439

Summary 440
Possible Contributions of the pSTS/BA39 and Some Adjacent Regions 440
Associations Between Auditory–Verbal STM and Sentence Comprehension 441
Dissociations Between Auditory–Verbal STM and Sentence Comprehension 445
Summary 447
Possible Contributions of Broca’s Area and Some Adjacent Regions 447
Is Broca’s Area Necessary for Sentence Comprehension? 447
Hypotheses that Emphasize Sequential and Hierarchical Processing 448
Box 15.1: Shared Syntax for Producing and Comprehending Sentences in Broca’s Area 449
Hypotheses that Emphasize Auditory–Verbal STM and Cognitive Control 453
Summary 457
Summary 458
Box 15.2: Subcortical Contributions to Sentence Comprehension 459
Major ERP Components Associated with Sentence Comprehension 459
The N400 459
The P600 462
Early and Sustained Negativities 466
Summary 469
Summary and Key Points 469
Recommended Reading 470

16 Discourse 471
Introduction 471
Story Production 472
Frog Stories: A Window onto the Neural Substrates of Narrative Coherence 472
Box 16.1: A Normal Frog Story 474
Ash et al.’s (2006) Neuropsychological Study 474
Troiani et al.’s (2008) fMRI Study 476
Story Comprehension 478
Electrophysiological Evidence for the Rapid Incorporation of Words into
the Discourse Context 479
A Meta-Analysis of 12 PET and fMRI Studies of the Comprehension of
Coherent Versus Incoherent Narratives 481
The Anterior Temporal Lobes: Integrating Semantic Information 481
The Medial Parietal Cortex: Establishing and Updating Situation Models 482
The Dorsomedial Prefrontal Cortex: Drawing Inferences 483
The Temporoparietal Junction: Attributing Thoughts to Protagonists 485
Summary 487
Summary and Key Points 488
Recommended Reading 488

References 489
Author Index 555
Subject Index 583
Preface

Welcome to Cognitive Neuroscience of Language. actions, and abstract notions (Part V); and the
During the past few decades, research on how our formulation and comprehension of complex expres-
brains enable us to perceive and produce language has sions, including grammatically inflected words,
been advancing at a dramatic pace. As a result, the field complete sentences, and entire stories (Part VI).
now contains a wealth of fascinating findings about this Substantial space is devoted to elaborating recent dis-
uniquely human capacity. Broadly speaking, the pur- coveries about the neural substrates of these central
pose of this book is to give students, teachers, research- aspects of language. But in order to keep the length
ers, and clinicians a solid, accessible introduction to this of the book within manageable limits, several other
material. More narrowly, the text has been deliberately topics are not covered, such as development, reha-
designed for use in courses offered to graduate students bilitation, bilingualism, figurative language, conversa-
and upper-level undergraduate students. Although tion, and co-speech gesture.
much of the content is inherently challenging, no pre- A distinctive feature of the text is that almost all of
vious knowledge of either neuroscience or linguistics is the chapters in Parts III–VI draw upon prominent the-
required, since technical terms and important princi- oretical models that characterize the various linguistic
ples from both disciplines are explained along the way. domains at both cognitive and neurobiological levels
In fact, during the roughly three-year period when I of analysis. In addition, these chapters illustrate how
was writing the book, I used the accumulating chapters the different components of the models are supported,
in my own courses at Purdue University, and many stu- and in some cases challenged, by experiments employ-
dents with little to no background in the neurobiology ing diverse brain mapping techniques. A special effort
of language managed to perform quite well, emerging has been made to describe these experiments in consid-
with a deep appreciation of the current state of the art erable detail, providing information about their goals,
in this exciting field of study. methods, results, and implications. The rationale for
The book begins by discussing human functional such an in-depth approach is that it may help students
neuroanatomy and brain mapping methods (Part I) understand not only how empirical studies are con-
as well as classical and progressive aphasia syndromes ducted, but also how they contribute to the dynamic
(Part II). The subsequent chapters then build on that interplay between theory and data.
background by focusing on the following topics: the A website with supplementary material is available at
perception and production of speech (Part III); the www.psypress.com/cw/kemmerer. Here teachers and
processing of language in written and signed modali- students will find PowerPoint slides for each chapter as
ties (Part IV); the meanings of words for objects, well as internet links and test questions.
Page Intentionally Left Blank
Acknowledgments

I am extremely grateful to my editors, Paul Dukes and Piñango, David Poeppel, Liina Pylkkänen, Meredith
Fred Coppersmith, for recognizing the potential of Saletta, and Stephen Wilson.
my original proposal and gradually bringing this book Many other individuals helped me keep the project
to fruition. They shepherded the manuscript through moving forward by answering questions, sending papers,
countless drafts while skillfully orchestrating a compli- and providing encouragement. I’m grateful to all of them:
cated review process. Daniel Abrams, Sharon Ash, Marina Bedny, Jeff Binder,
I would also like to acknowledge the assistance of Richard Binney, Anjan Chatterjee, Oren Civier, Evelyn
the following scholars, each of whom provided valu- Ferstl, Ed Fox, Guido Gainotti, Murray Grossman, Paul
able feedback on batches of several chapters: Joana Hoffman, Matt Lambon Ralph, Lotte Meteyard, Silke
Acha, Sheila Blumstein, Jonathan Brennan, Eva Paulmann, Steve Pinker, Friedemann Pulvermüller,
Fernández, Christopher Grindrod, Nancy Hedberg, Brenda Rapp, Jamie Reilly, Myrna Schwartz, Peter Sörös,
Greg Hickok, Li Hsieh, Edith Kaan, Laura Kertz, Cindy Thompson, Michael Ullman, Cosimo Urgesi, and
Ellen Lau, Filip Loncke, Michele Miozzo, Raksha Dirk Wildgruber.
Anand Mudar, Jodi Tommerdahl, and Matthew Special thanks goes to my mentor, collaborator, and
Wagers. friend, Dan Tranel, who taught me how to not only
In addition, I am indebted to the following people survive, but thrive, in the competitive world of cogni-
for serving as expert reviewers of single chapters: Juliana tive neuroscience.
Baldo, Larry Barsalou, Mike Bonner, Rutvik Desai, And last but certainly not least, I thank my wife and
Karen Emmorey, Jack Gandour, Lisa Goffman, Argye soul mate, Natalya Kaganovich, for her inspiration,
Hillis, Brad Mahon, Alex Martin, Loraine Obler, Maria reassurance, love, and companionship.
Page Intentionally Left Blank
 Fundamentals
PART I
Chapter 1: The Human Brain

Chapter 2: Brain Mapping Methods