THE SENSES:A COMPREHENSIVE

REFERENCE

THE SENSES:A COMPREHENSIVE

REFERENCE

Volume 6SOMATOSENSATION

Volume Editors

Dr Jon H. KaasVanderbilt University, Nashville, TN, USA

Dr Esther GardnerNew York University, New York, NY, USA

Advisory Board

Dr Allan I. BasbaumUniversity of California, San Francisco, CA, USA

Dr Akimichi KanekoKeio University, Tokyo, Japan

Dr Gordon M. ShepherdYale University, New Haven, CT, USA

Dr Gerald WestheimerUniversity of California, Berkeley, CA, USA

AMSTERDAM BOSTON HEIDELBERG LONDON NEW YORK OXFORDPARIS SAN DIEGO SAN FRANCISCO SINGAPORE SYDNEY TOKYO

Academic Press is an imprint of ElsevierThe Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK

525 B Street, Suite 1900, San Diego, CA 92101-4495, USA

First edition 2008

Copyright ª 2008 Elsevier Inc. All rights reserved

The following article is a US Government work in the public domain and is not subject to copyright:

PSYCHOPHYSICS OF PAINTREATMENT OF HEARING LOSS: VIRAL TRANSFECTION

Copyright ª 2008 A Lalwani

No part of this publication may be reproduced, stored in a retrieval systemor transmitted in any form or by any means electronic, mechanical, photocopying,

recording or otherwise without the prior written permission of the publisher

Permissions may be sought directly from Elsevier’s Science & Technology RightsDepartment in Oxford, UK: phone (þ44) (0) 1865 843830; fax (þ44) (0) 1865 853333;

email: permissions@elsevier.com. Alternatively you can submit your request online byvisiting the Elsevier web site at http://elsevier.com/locate/permissions, and selecting

Obtaining permission to use Elsevier material

NoticeNo responsibility is assumed by the publisher for any injury and/or damage to persons

or property as a matter of products liability, negligence or otherwise, or from any useor operation of any methods, products, instructions or ideas contained in the materialherein, Because of rapid advances in the medical sciences, in particular, independent

verfication of diagnoses and drug dosages should be made

British Library Cataloguing in Publication DataA catalogue record for this book is available from the British Library

Library of Congress Catalog Number: 2007939855

ISBN: 978-012-639482-5

For information on all Elsevier publicationsvisit our website at books.elsevier.com

Printed and bound in Canada

07 08 09 10 11 10 9 8 7 6 5 4 3 2 1

Working together to grow libraries in developing countries

www.elsevier.com | www.bookaid.org | www.sabre.org

Contents

Contents of All Volumes vii

Contributors to All Volumes xiii

Introduction to Volume 6 xxxiii

Dedication xxxv

Somatosensation

6.01 Cutaneous Mechanisms of Tactile Perception: Morphological and ChemicalOrganization of the Innervation to the Skin 1F L Rice and P J Albrecht, Albany Medical College, Albany, NY, USA

6.02 Merkel Cells 33Z Halata and K I Baumann, University of Hamburg, Hamburg, GermanyM Grim, Charles University, Praha, Czech Republic

6.03 Physiological Responses of Sensory Afferents in Glabrous andHairy Skin of Humans and Monkeys 39A W Goodwin and H E Wheat, University of Melbourne, Parkville, Vic, Australia

6.04 Coding of Object Shape and Texture 55S S Hsiao and S Bensmaia, The Johns Hopkins University, Baltimore, MD, USA

6.05 Tactile Sensory Control of Object Manipulation in Humans 67R S Johansson, Umea University, Umea, SwedenJ R Flanagan, Queen’s University, Kingston, ON, Canada

6.06 Physiological Characteristics of Second-Order Somatosensory Circuits in Spinal Cord andBrainstem 87W D Willis Jr, University of Texas Medical Branch, Galveston, TX, USA

6.07 The Somatosensory Thalamus and Associated Pathways 117J H Kaas, Vanderbilt University, Nashville, TN, USA

6.08 Somatosensory Areas of the Cerebral Cortex: Architectonic Characteristics and ModularOrganization 143H-X Qi and J H Kaas, Vanderbilt University, Nashville, TN, USAT M Preuss, Emory University, Atlanta, GA, USA

6.09 Development of the Somatosensory Cortex and Patterning of Afferent Projections 171R S Erzurumlu, University of Maryland School of Medicine, Baltimore, MD, USA

6.10 The Evolution of Parietal Areas Involved in Hand Use in Primates 183L Krubitzer, University of California, Davis, CA, USAE Disbrow, University of California, San Francisco, CA, USA

v

6.11 Role of Primary Somatosensory Cortex in Perceptual Touch Detection and Discrimination 215R Romo, A Hernandez, V de Lafuente, A Zainos, L Lemus, R Luna, V Nacher, and M Alvarez,Universidad Nacional Autonoma de Mexico, Mexico

6.12 Dorsal and Ventral Streams in the Sense of Touch 233Esther P Gardner, Department of Physiology and Neuroscience,New York University School of Medicine, New York, NY, USA

6.13 Plasticity of Somatosensory Function during Learning, Disease and Injury 259C Darian-Smith, Stanford University School of Medicine, Stanford, CA, USA

6.14 Intrinsic Signal Imaging of Somatosensory Function in Nonhuman Primates 299A W Roe, L M Chen, and R Friedman, Vanderbilt University, Nashville, TN, USA

6.15 Twenty-Five Years of Multielectrode Recordings in the Somatosensory System: It is All aboutDynamics 315M C Wiest, E Thomson, and M A L Nicolelis, Duke University, Durham, NC, USA

6.16 Specialized Somatosensory Systems 331J I Johnson, Michigan State University, East Lansing, MI, USAR L Reed, University of Florida, Gainesville, FL, USA

6.17 Somatosensation in Invertebrates 355A Buschges, University of Cologne, Cologne, GermanyR A DiCaprio, Ohio University, Athens, OH, USA

6.18 Visual Deprivation Effects on Somatosensory and Visual Systems: Behavioral andCortical Changes 363H Burton, Washington University School of Medicine, St. Louis, MO, USAD G McLaren, University of Wisconsin, Madison, WI, USA

6.19 Cross-Modal and Multisensory Interactions between Vision and Touch 393S Lacey and K Sathian, Emory University School of Medicine, Atlanta, GA, USA

Index 405

vi Contents

Contents of All Volumes

Volume 1 Vision I

1.01 The Visual System and Its Stimuli1.02 Evolution of Vertebrate Eyes1.03 Vision in Birds1.04 Vision in Fish1.05 Phototransduction in Microvillar Photoreceptors of Drosophila and Other Invertebrates1.06 Central Processing of Visual Information in Insects1.07 Color in Invertebrate Vision1.08 Visual Ecology1.09 Mammalian Photopigments1.10 Phototransduction in Rods and Cones1.11 Mammalian Rod Pathways1.12 Decomposing a Cone’s Output (Parallel Processing)1.13 Contributions of Horizontal Cells1.14 Contributions of Bipolar Cells to Ganglion Cell Receptive Fields1.15 Amacrine Cells1.16 The P, M and K Streams of the Primate Visual System: What Do They Do for Vision?1.17 Neural Mechanisms of Natural Scene Perception1.18 Seeing in the Dark: Retinal Processing and Absolute Visual Threshold1.19 Direction-Selective Cells1.20 Melanopsin Cells1.21 Blue-ON Cells1.22 Mosaics, Tiling and Coverage by Retinal Neurons1.23 Circuit Functions of Gap Junctions in the Mammalian Retina1.24 Plasticity of Retinal Circuitry1.25 Retinal Ganglion Cell Types and Their Central Projections1.26 Pupillary Control Pathways1.27 The Suprachiasmatic Nucleus1.28 The Visual Thalamus1.29 Functional Maps in Visual Cortex: Topographic, Modular, and Columnar Organizations1.30 Organization of Human Visual Cortex

Index to Volumes 1 and 2

Volume 2 Vision II

2.01 Temporal Coherence: A Versatile Code for the Definition of Relations2.02 High-Level Visual Processing2.03 Luminance Sensitivity and Contrast Detection2.04 Lightness Perception and Filling-In2.05 Nocturnal Vision2.06 Spectral Sensitivity2.07 Chromatic Detection and Discrimination

vii

2.08 Color Appearance2.09 Motion Detection Mechanisms2.10 Cortical Processing of Visual Motion2.11 Cortical Mechanisms for the Integration of Visual Motion2.12 Optic Flow2.13 Biological Motion Perception2.14 Transparency and Occlusion2.15 Three-Dimensional Shape: Cortical Mechanisms of Shape Extraction2.16 Visual Search2.17 Object-Based Attention2.18 Visual Attention and Saccadic Eye Movements2.19 Saliency2.20 Perceptual Learning2.21 Face Recognition2.22 The VOR: A Model for Visual-Motor Plasticity

Index to Volumes 1 and 2

Volume 3 Audition

3.01 Phylogeny and Evolution of Ciliated Mechanoreceptor Cells3.02 Insect Ears3.03 High-Frequency Hearing3.04 Sensory Ecology of Hearing3.05 Genetics of Mechanoreceptor Evolution and Development3.06 Molecular Anatomy of Receptor Cells and Organ of Corti3.07 Genetic Hearing Loss3.08 Homeostasis of the Inner Ear3.09 Meniere’s Disease3.10 Mechano-Acoustical Transformations3.11 Evolution of the Middle Ear and Inner Ear in Vertebrates3.12 Biophysics of Chordotonal Organs3.13 Interconnections between the Ears in Nonmammalian Vertebrates3.14 Underwater Hearing3.15 Otoacoustic Emissions3.16 Hair Cell Transduction and Adaptation: Physiology and Molecular Mechanisms3.17 Amplification and Feedback in Invertebrates3.18 Tinnitus3.19 Prestin3.20 Cochlear Receptor Potentials3.21 Manifestations of Cochlear Events in the Auditory Brain-stem Response and

Its Clinical Applications3.22 Afferent Synaptic Mechanisms3.23 Perspectives on Auditory Neuropathy: Disorders of Inner Hair Cell, Auditory Nerve, and

Their Synapse3.24 Efferent System3.25 Overview of Treatment of Hearing Loss3.26 Cochlear Implants3.27 Hearing Loss and Hearing Aids: A Perspective3.28 Sensory Regeneration in the Vertebrate Ear3.29 Treatment of Hearing Loss: Viral Transfection3.30 Vertebrate Auditory Pathways3.31 Invertebrate Auditory Pathways3.32 Biophysical Specializations of Neurons that Encode Timing3.33 Central Synapses that Preserve Auditory Timing3.34 Acoustic Startle in Mice and Rats3.35 Encoding of Interaural Timing for Binaural Hearing

viii Contents of All Volumes

3.36 Encoding of Interaural Level Differences for Sound Localization3.37 Monaural Sound Localization Using Spectral Cues3.38 The Bat Cochlea3.39 Auditory Processing in the Bat Medial Superior Olive3.40 Brain Mechanisms of Sound Localization in Barn Owls3.41 Sound Localization in Insects3.42 Inputs to the Inferior Colliculus3.43 The Nuclei of the Lateral Lemniscus: Two Functional Systems3.44 Auditory Map Plasticity in Juvenile and Adult Owls3.45 The Functional Neuroanatomy of the Auditory Cortex3.46 Sound Localization and the Auditory Cortex3.47 Pitch Perception3.48 Perception of Speech Sounds3.49 Auditory Scene Analysis3.50 Human Auditory Development3.51 Sleep and Memory Consolidation in Audition

Index

Volume 4 Olfaction & Taste

4.01 Phylogeny of Chemical Sensitivity4.02 Chemistry of Gustatory Stimuli4.03 Insect Gustatory Systems4.04 Aquatic Animal Models in the Study of Chemoreception4.05 Ultrastructure of Taste Buds4.06 Development of the Taste System4.07 The Sweet Taste of Childhood4.08 Taste Analgesia in Newborns4.09 Taste Receptors4.10 Taste Transduction4.11 Gustatory Pathways in Fish and Mammals4.12 Neurotransmitters in the Taste Pathway4.13 Functional Magnetic Resonance Imaging (fMRI) Study of Taste4.14 Amiloride-Sensitive Ion Channels4.15 Central Neural Processing of Taste Information4.16 Neural Ensembles in Taste Coding4.17 A Perspective on Chemosensory Quality Coding4.18 Oral Chemesthesis and Taste4.19 Genetics and Evolution of Taste4.20 Propylthiouracil (PROP) Taste4.21 Salt Taste4.22 Behavioral Analysis of Taste Function in Rodent Models4.23 Flavor Aversion Learning4.24 Roles of Taste in Feeding and Reward4.25 Dopamine Release by Sucrose4.26 The Representation of Flavor in the Brain4.27 The Aging Gustatory System4.28 Signal Transduction in the Olfactory Receptor Cell4.29 Olfactory Cyclic Nucleotide-Gated Ion Channels4.30 Structure, Expression, and Function of Olfactory Receptors4.31 Regulation of Expression of Odorant Receptor Genes4.32 Genomics of Odor Receptors in Zebrafish4.33 Genomics of Invertebrate Olfaction4.34 Regeneration of the Olfactory Epithelium4.35 Regeneration in the Olfactory Bulb4.36 Architecture of the Olfactory Bulb

Contents of All Volumes ix

4.37 Physiology of the Main Olfactory Bulb4.38 Olfactory Cortex4.39 Modeling of Olfactory Processing4.40 Understanding Olfactory Coding via an Analysis of Odorant-Evoked Glomerular

Response Maps4.41 Insect Olfaction4.42 Odor Plumes and Animal Orientation4.43 Accessory Olfactory System4.44 Genomics of Vomeronasal Receptors4.45 Human Olfactory Psychophysics4.46 Disorders of Taste and Smell

Index

Volume 5 Pain

5.01 The Adequate Stimulus5.02 Pain Theories5.03 Anatomy of Nociceptors5.04 Molecular Biology of the Nociceptor/Transduction5.05 Zoster-Associated Pain and Nociceptors5.06 Ectopic Generators5.07 Sodium Channels5.08 Physiology of Nociceptors5.09 Itch5.10 Thermal Sensation (Cold and Heat) through Thermosensitive TRP Channel Activation5.11 The Development of Nociceptive Systems5.12 Appropriate/Inappropriate Developed ‘‘Pain’’ Paths5.13 Pain Control: A Child-Centered Approach5.14 Assaying Pain-Related Genes: Preclinical and Clinical Correlates5.15 Evolutionary Aspects of Pain5.16 Redheads and Pain5.17 Autonomic Nervous System and Pain5.18 Sympathetic Blocks for Pain5.19 Sprouting in Dorsal Root Ganglia5.20 Vagal Afferent Neurons and Pain5.21 Sex, Gender, and Pain5.22 Neurotrophins and Pain5.23 Morphological and Neurochemical Organization of the Spinal Dorsal Horn5.24 Spinal Cord Physiology of Nociception5.25 What is a Wide-Dynamic-Range Cell?5.26 Spinal Cord Mechanisms of Hyperalgesia and Allodynia5.27 Glycine Receptors5.28 Pain Following Spinal Cord Injury5.29 Long-Term Potentiation in Pain Pathways5.30 Immune System, Pain and Analgesia5.31 Mechanisms of Glial Activation after Nerve Injury5.32 Trigeminal Mechanisms of Nociception: Peripheral and Brainstem Organization5.33 Migraine – A Disorder Involving Trigeminal Brainstem Mechanisms5.34 Tooth Pain5.35 Ascending Pathways: Anatomy and Physiology5.36 Dorsal Columns and Visceral Pain5.37 Visceral Pain5.38 Irritable Bowel Syndrome5.39 Pain in Childbirth5.40 Urothelium as a Pain Organ5.41 The Brainstem and Nociceptive Modulation

x Contents of All Volumes

5.42 Emotional and Behavioral Significance of the Pain Signal and the Role of the MidbrainPeriaqueductal Gray (PAG)

5.43 The Thalamus and Nociceptive Processing5.44 Psychophysics of Sensations Evoked by Stimulation of the Human Central Nervous System5.45 Nociceptive Processing in the Cerebral Cortex5.46 Phantom Limb Pain5.47 Human Insular Recording and Stimulation5.48 The Rostral Agranular Insular Cortex5.49 Descending Control Mechanisms5.50 Diffuse Noxious Inhibitory Controls (DNIC)5.51 Fibromyalgia5.52 Pain Perception – Nociception during Sleep5.53 Pharmacological Modulation of Pain5.54 Forebrain Opiates5.55 Neuropathic Pain: Basic Mechanisms (Animal)5.56 Animal Models and Neuropathic Pain5.57 Neuropathic Pain: Clinical5.58 Neurogenic Inflammation in Complex Regional Pain Syndrome (CRPS)5.59 Complex Regional Pain Syndromes5.60 Poststroke Pain5.61 Psychophysics of Pain5.62 Consciousness and Pain5.63 Assessing Pain in Animals5.64 Psychological Modulation of Pain5.65 The Placebo Effect5.66 Hypnotic Analgesia

Index

Volume 6 Somatosensation

6.01 Cutaneous Mechanisms of Tactile Perception: Morphological and ChemicalOrganization of the Innervation to the Skin

6.02 Merkel Cells6.03 Physiological Responses of Sensory Afferents in Glabrous and

Hairy Skin of Humans and Monkeys6.04 Coding of Object Shape and Texture6.05 Tactile Sensory Control of Object Manipulation in Humans6.06 Physiological Characteristics of Second-Order Somatosensory Circuits in Spinal Cord and

Brainstem6.07 The Somatosensory Thalamus and Associated Pathways6.08 Somatosensory Areas of the Cerebral Cortex: Architectonic Characteristics and Modular Organization6.09 Development of the Somatosensory Cortex and Patterning of Afferent Projections6.10 The Evolution of Parietal Areas Involved in Hand Use in Primates6.11 Role of Primary Somatosensory Cortex in Perceptual Touch Detection and Discrimination6.12 Dorsal and Ventral Streams in the Sense of Touch6.13 Plasticity of Somatosensory Function during Learning, Disease and Injury6.14 Intrinsic Signal Imaging of Somatosensory Function in Nonhuman Primates6.15 Twenty-Five Years of Multielectrode Recordings in the Somatosensory System: It is All about Dynamics6.16 Specialized Somatosensory Systems6.17 Somatosensation in Invertebrates6.18 Visual Deprivation Effects on Somatosensory and Visual Systems: Behavioral and

Cortical Changes6.19 Cross-Modal and Multisensory Interactions between Vision and Touch

Index

Contents of All Volumes xi

Contributors to All Volumes

B W AcheUniversity of Florida, Gainesville, FL, USA

P J AlbrechtAlbany Medical College, Albany, NY, USA

J M AlexanderUniversity of Wisconsin–Madison, Madison, WI, USA

T S AliotoUniversity of California, Berkeley, CA, USA

M AlvarezUniversidad Nacional Autonoma de Mexico, Mexico

B L AndersonUniversity of New South Wales, Sydney, NSW, Australia

D E AngelakiWashington University School of Medicine, St. Louis, MO, USA

V AnseloniUniversity of Maryland Dental School, Baltimore, MD, USA

A V ApkarianNorthwestern University, Chicago, IL, USA

K M ArmstrongStanford University School of Medicine, Stanford, CA, USA

K BowmakerUniversity College London, London, UK

A A BachmanovMonell Chemical Senses Center, Philadelphia, PA, USA

C A BagleyJohns Hopkins Hospital, Baltimore, MD, USA

R BandlerUniversity of Sydney, Sydney, NSW, Australia

L A BarlowUniversity of Colorado School of Medicine, Aurora, CO, USA

R BaronChristian-Albrechts-Universitat Kiel, Kiel, Germany

xiii

L M BartoshukUniversity of Florida, Gainesville, FL, USA

K I BaumannUniversity of Hamburg, Hamburg, Germany

G K BeauchampMonell Chemical Senses Center, Philadelphia, PA, USA

O BehrendHumboldt-University, Berlin, Germany

K W BeiselCreighton University, Omaha, NE, USA

F BenedettiUniversity of Turin Medical School, Turin, Italy

S BensmaiaThe Johns Hopkins University, Baltimore, MD, USA

D A BereiterUniversity of Minnesota, Minneapolis, MN, USA

J BerganStanford University School of Medicine, Stanford, CA, USA

I L BernsteinUniversity of Washington, Seattle, WA, USA

D M BersonBrown University, Providence, RI, USA

T BertaUniversity of Lausanne, Lausanne, Switzerland

K BielefeldtUniversity of Pittsburgh, Pittsburgh, PA, USA

L A BirderUniversity of Pittsburgh School of Medicine, Pittsburgh, PA, USA

F BirkleinUniversity of Mainz, Mainz, Germany

J D BohbotVanderbilt University, Nashville, TN, USA

R T BornHarvard Medical School, Boston, MA, USA

J D Boughter Jr.University of Tennessee Health Science Center, Memphis, TN, USA

S BradesiUniversity of California, Los Angeles, CA, USA

R M BradleyUniversity of Michigan, Ann Arbor, MI, USA

A S BregmanMcGill University, Montreal, QC, Canada

K H BrittenUniversity of California, Davis, CA, USA

xiv Contributors to All Volumes

M-C BroilletUniversity of Lausanne, Lausanne, Switzerland

S M BromleyUniversity of Pennsylvania, Philadelphia, PA, USA, UMDNJ-Robert Wood Johnson Medical School,Camden, NJ, USA

R M BurgerLehigh University, Bethlehem, PA, USA

H BurtonWashington University School of Medicine, St. Louis, MO, USA

M R ByersUniversity of Washington, Seattle WA, USA

A BuschgesUniversity of Cologne, Cologne, Germany

S W CaddenUniversity of Dundee, Dundee, UK

J N CampbellJohns Hopkins University, Baltimore, MD, USA

J CaprioLouisiana State University, Baton Rouge, LA, USA

C E CarrUniversity of Maryland, College Park, MD, USA

J CarrollMedical College of Wisconsin, Milwaukee, WI, USA

E CarstensUniversity of California, Davis, CA, USA

M J CaterinaJohns Hopkins School of Medicine, Baltimore, MD, USA

B Cerf-DucastelSan Diego State University, San Diego, CA, USA

F CerveroMcGill University, Montreal, QC, Canada

L M ChenVanderbilt University, Nashville, TN, USA

J Christensen-DalsgaardUniversity of Southern Denmark, Odense, Denmark

T A ClelandCornell University, Ithaca, NY, USA

T J CoderreMcGill University, Montreal, QC, Canada

D CopenhagenUniversity of California San Francisco, CA, USA

R M CostanzoVirginia Commonwealth University, Richmond, VA, USA

Contributors to All Volumes xv

E CoveyUniversity of Washington, Seattle, WA, USA

A D CraigBarrow Neurological Institute, Phoenix, AZ, USA

W CroninUniversity of Maryland, Baltimore, MD, USA

C Darian-SmithStanford University School of Medicine, Stanford, CA, USA

R Davis-TaberGlobal Pharmaceutical Research and Development, Abbott Park, IL, USA

J W DawsonCarleton University, Ottawa, ON, Canada

Y De KoninckCentre de recherche Universite Laval Robert-Giffard, Quebec, QC, Canada

V de LafuenteUniversidad Nacional Autonoma de Mexico, Mexico

I DecosterdUniversity of Lausanne, Lausanne, Switzerland

P H DelanoUniversidad de Chile, Santiago, Chile

C D DerbyGeorgia State University, Atlanta, GA, USA

S W G DerbyshireUniversity of Birmingham, Birmingham, UK

J A DeSimoneVirginia Commonwealth University, Richmond, VA, USA

J DeSimoneVirginia Commonwealth University, Richmond, VA, USA

M DevorHebrew University of Jerusalem, Jerusalem, Israel

R A DiCaprioOhio University, Athens, OH, USA

E DisbrowUniversity of California, San Francisco, CA, USA

J O DostrovskyUniversity of Toronto, Toronto, ON, Canada

R L DotyUniversity of Pennsylvania, Philadelphia, PA, USA

A DrayAstraZeneca Research and Development, Montreal, PQ, Canada

R DubnerUniversity of Maryland, Baltimore, MD, USA

G E DuBoisThe Coca-Cola Company, Atlanta, GA, USA

xvi Contributors to All Volumes

B DuchaineUniversity College London, London, UK

V B DuffyUniversity of Connecticut, Storrs, CT, USA

J D DurrantUniversity of Pittsburgh, Pittsburgh, PA, USA

P L Edds-WaltonParmly Hearing Institute, Chicago, IL, USA

E EliavUMDNJ-New Jersey Dental School, Newark, NJ, USA

M EnnisUniversity of Tennessee Health Science Center, Memphis, TN, USA

R S ErzurumluUniversity of Maryland School of Medicine, Baltimore, MD, USA

R T Eskew Jr.Northeastern University, Boston, MA, USA

T EulerMax-Planck-Institute for Medical Research, Heidelberg, Germany

A FaurionNeurobiologie Sensorielle, NOPA-NBS, INRA, Jouy en Josas, France

R R FayLoyola University Chicago, Chicago, IL, USA

D J FellemanUniversity of Texas Medical School, Houston, TX, USA

A S FengUniversity of Illinois at Urbana-Champaign, Urbana, IL, USA

K M FennUniversity of Chicago, Chicago, IL, USA

R D FernaldStanford University, Stanford, CA, USA

J FerraroUniversity of Kansas Medical Center, Kansas City, KS, USA

R B FillingimUniversity of Florida College of Dentistry, Community Dentistry and Behavioral Science Gainesville, FL, USA

T E FingerUniversity of Colorado School of Medicine, Aurora, CO, USA

N B FinnerupAarhus University Hospital, Aarhus, Denmark

M F FitzgeraldUniversity College London, London, UK

J R FlanaganQueen’s University, Kingston, ON, Canada

H FlorCentral Institute of Mental Health, Mannheim, Germany

Contributors to All Volumes xvii

A FontaniniBrandeis University, Waltham, MA, USA

D H FosterUniversity of Manchester, Manchester, UK

M E FrankUniversity of Connecticut Health Center, Farmington, CT, USA

M A FreedUniversity of Pennsylvania School of Medicine, Philadelphia, PA, USA

A S FrenchDalhousie University, Halifax, NS, Canada

R FriedmanVanderbilt University, Nashville, TN, USA

B FritzschCreighton University, Omaha, NE, USA

M FrotINSERM U879, Bron France

T FukushimaThe University of Tokyo School of Medicine, Tokyo, Japan

D N FurnessKeele University, Keele, UK

G GaliziaUniversitat Konstanz, Konstanz, Germany

J L GallantHelen Wills Neuroscience Institute, Berkeley, CA, USA

P D R GamlinUniversity of Alabama at Birmingham, Birmingham, AL, USA

E P GardnerDepartment of Physiology and Neuroscience, New York University School of Medicine, New York, NY, USA

G F GebhartUniversity of Pittsburgh, Pittsburgh, PA, USA

C D GilbertThe Rockefeller University, New York, NY, USA

D Rodriguez GilYale University School of Medicine, New Haven, CT, USA

J I GlendinningBarnard College, Columbia University, New York, NY, USA

P J GoadsbyUniversity of California, San Francisco, CA, USA

P GocheeUniversity of Kansas Medical Center, Kansas City, KS, USA

M S GoldUniversity of Pittsburgh, Pittsburgh PA, USA

A W GoodwinUniversity of Melbourne, Parkville, Vic, Australia

xviii Contributors to All Volumes

J GottliebColumbia University, New York, NY, USA

R H GracelyUniversity of Michigan Health System, VAMC, Ann Arbor, MI, USA

C A GreerYale University School of Medicine, New Haven, CT, USA

M Gridi-PappUniversity of California, Los Angeles, CA, USA

M GrimCharles University, Praha, Czech Republic

S E GrossmanBrandeis University, Waltham, MA, USA

B GrotheLudwig-Maximilians-University, Munich, Germany

M C GopfertUniversity of Cologne, Cologne, Germany

T A HackettVanderbilt University, Nashville, TN, USA

C M HackneyUniversity of Cambridge, Cambridge, UK

A HajnalMilton S. Hershey Medical Center, Hershey, PA, USA

Z HalataUniversity of Hamburg, Hamburg, Germany

R HallworthCreighton University, Omaha, NE, USA

R C HardieUniversity of Cambridge, Cambridge, UK

K M HargreavesUniversity of Texas Health Science Center, San Antonio, TX, USA

I A HarringtonAugustana College, Rock Island, IL, USA

J P HarrisUniversity of California, San Diego, CA, USA

G J HathwayUniversity College London, London, UK

S E HausseltMax-Planck-Institute for Medical Research, Heidelberg, Germany

A HayarUniversity of Arkansas for Medical Sciences, Little Rock, AR, USA

J E HayesBrown University, Providence, RI, USA

D HeCreighton University, Omaha, NE, USA

Contributors to All Volumes xix

B HedwigUniversity of Cambridge, Cambridge, UK

H E HeffnerUniversity of Toledo, Toledo, OH, USA

R S HeffnerUniversity of Toledo, Toledo, OH, USA

M M HeinricherOregon Health & Science University, Portland, OR, USA

A HernandezUniversidad Nacional Autonoma de Mexico, Mexico

A HirshUniversity of Florida, Gainesville, FL, USA

J R HoltUniversity of Virginia School of Medicine, Charlottesville, VA, USA

P HonoreGlobal Pharmaceutical Research and Development, Abbott Park, IL, USA

S S HsiaoThe Johns Hopkins University, Baltimore, MD, USA

J W HuUniversity of Toronto, Toronto, ON, Canada

J IglesiasCuban Center for Neuroscience, Habana, Cuba

F ImamuraYale University School of Medicine, New Haven, CT, USA

S L IngramWashington State University, Vancouver, WA, USA

J IsnardLyon I University and INSERM U879, Bron, France

G H JacobsUniversity of California, Santa Barbara, CA, USA

W JanigPhysiologisches Institut, Christian-Albrechts-Universitat zu Kiel, Germany

W JanigChristian-Albrechts-Universitat zu Kiel, Kiel, Germany

L JasminNeurosurgery and Gene Therapeutics Research Institute, Los Angeles, CA, USA

T S JensenAarhus University Hospital, Aarhus, Denmark

R S JohanssonUmea University, Umea, Sweden

S J St. JohnRollins College, Winter Park, FL, USA

B A JohnsonUniversity of California, Irvine, CA, USA

xx Contributors to All Volumes

B JohnsonUC Berkeley, Berkeley, CA, USA

J I JohnsonMichigan State University, East Lansing, MI, USA

J H KaasVanderbilt University, Nashville, TN, USA

T KamigakiThe University of Tokyo School ofMedicine,Tokyo, Japan

E KaplanThe Mount Sinai School of Medicine, NewYork, NY, USA

H KasaharaThe University of Tokyo School of Medicine, Tokyo, Japan

D B KatzBrandeis University, Waltham, MA, USA

B J B KeatsLouisiana State University Health Sciences Center, New Orleans, LA, USA

K KeayUniversity of Sydney, Sydney, NSW, Australia

V KefalovWashington University School of Medicine, St. Louis, MO, USA

D R KettenWoods Hole Oceanographic Institution, Woods Hole, MA, USA

R M KhanUC Berkeley, Berkeley, CA, USA

M C KillionEtymotic Research Ltd., Elk Grove Village, IL, USA

J C KinnamonUniversity of Denver, Denver, CO, USA

S C KinnamonColorado State University, Fort Collins, CO, USA

K R KluenderUniversity of Wisconsin–Madison, Madison, WI, USA

E KnudsenStanford University School of Medicine, Stanford, CA, USA

T KobayakawaNational Institute of Advanced Industrial Science and Technology (AIST), Tsukuba, Japan

H KomatsuNational Institute for Physiological Sciences, Okazaki, Japan

M KonishiCalifornia Institute of Technology, Pasadena, CA, USA

H G KrappImperial College London, London, UK

B KrekelbergRutgers University, Newark, NJ, USA

Contributors to All Volumes xxi

R F KrimmUniversity of Louisville School of Medicine, Louisville, KY, USA

L KrubitzerUniversity of California, Davis, CA, USA

T KurahashiOsaka University, Osaka, Japan

M KosslJohann Wolfgang Goethe Universitat, Frankfurt/Main, Germany

S LaceyEmory University School of Medicine, Atlanta, GA, USA

R LadherRIKEN Centre for Developmental Biology, Kobe, Japan

A K LalwaniNew York University School of Medicine, New York, NY, USA

G J LavigneUniversite de Montreal, Montreal, QC, Canada

H C LawsonJohns Hopkins Hospital, Baltimore, MD, USA

D Le BarsINSERM U-713, Paris, France

B B LeeSUNY College of Optometry, New York, NY, USA

S LeeKorea Institute of Science and Technology, Seoul, Korea

T Leinders-ZufallUniversity of Maryland School of Medicine, Baltimore, MD, USA

A LelliUniversity of Virginia School of Medicine, Charlottesville, VA, USA

L LemusUniversidad Nacional Autonoma de Mexico, Mexico

F A LenzJohns Hopkins Hospital, Baltimore, MD, USA

M LeonUniversity of California, Irvine, CA, USA

A R LightUniversity of Utah, Salt Lake City, UT, USA

D LimaUniversidade do Porto, Porto, Portugal

C LinsterCornell University, Ithaca, NY, USA

W LiThe Rockefeller University, New York, NY, USA

P-M LledoPasteur Institute, Paris, France

xxii Contributors to All Volumes

E R LoewCornell University, Ithaca, NY, USA

R LunaUniversidad Nacional Autonoma de Mexico, Mexico

D-G LuoJohns Hopkins University School of Medicine, Baltimore, MD, USA

V LyallVirginia Commonwealth University, Richmond, VA, USA

H MachelskaCharite – Universitatsmedizin Berlin, Campus Benjamin Franklin, Berlin, Germany

E A MacphersonUniversity of Michigan, Ann Arbor, MI, USA

S F MaierUniversity of Colorado at Boulder, Boulder, CO, USA

H MaijaHelsinki University Hospital, Helsinki, Finland

P B ManisThe University of North Carolina at Chapel Hill, Chapel Hill, NC, USA

G A ManleyTechnische Universitat Munchen, Garching, Germany

I MarcUniversite Laval, Quebec City, QC, Canada

D MargoliashUniversity of Chicago, Chicago, IL, USA

R F MargolskeeMount Sinai School of Medicine, New York, NY, USA

G R MartinUniversity of Birmingham, Birmingham, UK

S C MasseyUniversity of Texas Medical School, Houston, TX, USA

F MauguiereLyon I University and INSERM U879, Bron, France

M MaxMount Sinai School of Medicine, New York, NY, USA

B J MayThe Johns Hopkins University School of Medicine, Baltimore, MD, USA

E A MayerUniversity of California, Los Angeles, CA, USA

C H McCoolUniversity of California, Davis, CA, USA

D H McDougalUniversity of Alabama at Birmingham, Birmingham, AL, USA

P A McGrathThe University of Toronto, Toronto, ON, Canada

Contributors to All Volumes xxiii

E M McLachlanPrince of Wales Medical Research Institute, Randwick, NSW, Australia

D G McLarenUniversity of Wisconsin, Madison, WI, USA

L M MendellState University of New York, Stony Brook, NY, USA

J A MennellaMonell Chemical Senses Center, Philadelphia, PA, USA

S MenseInstitut fur Anatomie und Zellbiologie, Universitat Heidelberg, Heidelberg, Germany

W MeyerhofGerman Institute of Human Nutrition Potsdam-Rehbruecke, Nuthetal, Germany

R A MeyerJohns Hopkins University, Baltimore, MD, USA

H J MichalewskiUniversity of California, Irvine, CA, USA

J C MiddlebrooksUniversity of Michigan, Ann Arbor, MI, USA

E D MilliganUniversity of Colorado at Boulder, Boulder, CO, USA

Y MiyashitaThe University of Tokyo School of Medicine, Tokyo, Japan

J S MogilMcGill University, Montreal, QC, Canada

T MooreStanford University School of Medicine, Stanford, CA, USA

T MoserUniversity of Goettingen, Goettingen, Germany

V NacherUniversidad Nacional Autonoma de Mexico, Mexico

P M NarinsUniversity of California, Los Angeles, CA, USA

J NgaiUniversity of California, Berkeley, CA, USA

M A L NicolelisDuke University, Durham, NC, USA

R NorgrenMilton S. Hershey Medical Center, Hershey, PA, USA

P T OharaUniversity of California, San Francisco, CA, USA

S OharaJohns Hopkins Hospital, Baltimore, MD, USA

K OkuraTokushima Graduate School, Tokushima, Japan

xxiv Contributors to All Volumes

D OliverUniversitat Freiburg, Freiburg, Germany

G A OrbanK.U. Leuven Medical School, Leuven, Belgium

D OsorioUniversity of Sussex, Brighton, UK

M H OssipovUniversity of Arizona, Tucson, AZ, USA

C C PackMcGill University School of Medicine, Montreal, PQ, Canada

G E PickardColorado State University, Fort Collins, CO, USA

R J PittsVanderbilt University, Nashville, TN, USA

G S PollackMcGill University, Montreal, QC, Canada

A N PopperUniversity of Maryland, College Park, MD, USA

F PorrecaUniversity of Arizona, Tucson, AZ, USA

C V PortforsWashington State University, Vancouver, WA, USA

M PostmaUniversity of Cambridge, Cambridge, UK

R J PrengerUniversity of California, Berkeley, CA, USA

T M PreussEmory University, Atlanta, GA, USA

D D PriceUniversity of Florida, Gainesville, FL, USA

I ProvencioUniversity of Virginia, Charlottesville, VA, USA

A C PucheUniversity of Maryland School of Medicine, Baltimore, MD, USA

S PuriaStanford University, Stanford, CA, USA

H-X QiVanderbilt University, Nashville, TN, USA

P RainvilleUniversite de Montreal, Montreal, QC, Canada

S N RajaJohns Hopkins University, Baltimore, MD, USA

R RajimehrMassachusetts General Hospital, Charlestown, MA, USA

Contributors to All Volumes xxv

R L ReedUniversity of Florida, Gainesville, FL, USA

B E ReeseUniversity of California, Santa Barbara, CA, USA

L RelaYale University School of Medicine, New Haven, CT, USA

K RenUniversity of Maryland, Baltimore, MD, USA

B A RevillBrandeis University, Waltham, MA, USA

J ReynoldsThe Salk Institute for Biological Studies, San Diego, CA, USA

A Ribeiro-da-SilvaMcGill University, Montreal, QC, Canada

F L RiceAlbany Medical College, Albany, NY, USA

F RiekeUniversity of Washington, Seattle, WA, USA

M RingkampJohns Hopkins University, Baltimore, MD, USA

H L RittnerCharite – Universitatsmedizin Berlin, Campus Benjamin Franklin, Berlin, Germany

D RobertUniversity of Bristol, Bristol, UK

W M RobertsUniversity of Oregon, Eugene, OR, USA

M E RobinsonUniversity of Florida, Gainesville, FL, USA

L RoblesUniversidad de Chile, Santiago, Chile

V RodrıguezCuban Center for Neuroscience, Habana, Cuba

I RodriguezUniversity of Geneva, Geneva, Switzerland

A W RoeVanderbilt University, Nashville, TN, USA

E T RollsUniversity of Oxford, Oxford, UK

R RomoUniversidad Nacional Autonoma de Mexico, Mexico

E W RubelUniversity of Washington, Seattle, WA, USA

I RussellUniversity of Sussex, Brighton, UK

xxvi Contributors to All Volumes

M A RutherfordUniversity of Oregon, Eugene, OR, USA

K SaitoUniversity of Pennsylvania, Philadelphia, PA, USA

H SakanoUniversity of Tokyo, Tokyo, Japan

A N SaltWashington University School of Medicine, St. Louis, MO, USA

J SandkuhlerMedical University of Vienna, Vienna, Austria

K SathianEmory University School of Medicine, Atlanta, GA, USA

R J SchaferStanford University School of Medicine, Stanford, CA, USA

S S SchiffmanDuke University Medical Center, Durham, NC, USA

M SchmelzUniversity of Heidelberg, Mannheim, Germany

J SchouenborgLund University, Lund, Sweden

B A SchulteMedical University of South Carolina, Charleston, SC, USA

I SchwetzMedical University, Graz, Austria

J E SchwobTufts University School of Medicine, Boston, MA, USA

V E ScottGlobal Pharmaceutical Research and Development, Abbott Park, IL, USA

R V ShannonHouse Ear Institute, Los Angeles, CA, USA

A SharmaColumbia University, New York, NY, USA

L T SharpeUniversity College London, London, UK

S M ShermanThe University of Chicago, Chicago, IL, USA

T ShimuraOsaka University, Osaka, Japan

J SiegelNorthwestern University, Evanston, IL, USA

C T SimonsGlobal Research and Development Center, Cincinnati, OH, USA

W SingerMax Planck Institute for Brain Research, Frankfurt, Germany

Contributors to All Volumes xxvii

D V SmithThe University of Tennessee College of Medicine, Memphis, TN, USA

M T SmithJohn Hopkins Medical School, Baltimore, MD, USA

R G SmithUniversity of Pennsylvania, Philadelphia, PA, USA

J B Snow Jr.University of Pennsylvania, Philadelphia, PA, USA

D J SnyderYale University, New Haven, CT, USA

N SobelUC Berkeley, Berkeley, CA, USA

P J SollarsColorado State University, Fort Collins, CO, USA

A C SpectorThe Florida State University, Tallahassee, FL, USA

H StaeckerUniversity of Kansas Medical Center, Kansas City, KS, USA

A StarrUniversity of California, Irvine, CA, USA

R StaudUniversity of Florida, Gainesville, FL, USA

E A StaufferUniversity of Virginia School of Medicine, Charlottesville, VA, USA

G C SteckerUniversity of Washington, Seattle, WA, USA

C R SteeleStanford University, Stanford, CA, USA

C SteinCharite – Universitatsmedizin Berlin, Campus Benjamin Franklin, Berlin, Germany

L J SteinMonell Chemical Senses Center, Philadelphia, PA, USA

A StockmanUniversity College London, London, UK

R StormsVeterans Administration Medical Center, Kansas City, MO, USA

E StrettoiNeuroscience Institute, Pisa, Italy

H TakeuchiOsaka University, Osaka, Japan

E ThomsonDuke University, Durham, NC, USA

N TianYale University, New Haven, CT, USA

xxviii Contributors to All Volumes

D J TollinUniversity of Colorado Health Sciences Center, Aurora, CO, USA

M TominagaNational Institutes of Natural Sciences, Okazaki, Japan

R TootellMassachusetts General Hospital, Charlestown, MA, USA

K TouharaThe University of Tokyo, Chiba, Japan

S P TraversThe Ohio State University, Columbus, OH, USA

R D TreedeJohannes Gutenberg-University, Mainz, Germany

R D TreedeRuprecht-Karls-University Heidelberg, Heidelberg, Germany

N F TrojeQueen’s University, Kingston, ON, Canada

L O TrussellOregon Health and Science University, Portland, OR, USA

A TsuboiUniversity of Tokyo, Tokyo, Japan

M J Valdes-SosaCuban Center for Neuroscience, Habana, Cuba

D I VaneyThe University of Queensland, Brisbane, QLD, Australia

M VaterUniversitat Potsdam, Golm, Germany

M VorobyevUniversity of Queensland, Brisbane, QLD, Australia

E T WaltersUniversity of Texas at Houston, Medical School, Houston, TX, USA

M E WarcholWashington University School of Medicine, St. Louis, MO, USA

E WarrantUniversity of Lund, Lund, Sweden

W H WarrenBrown University, Providence, RI, USA

L R WatkinsUniversity of Colorado at Boulder, Boulder, CO,USA

L A WernerUniversity of Washington, Seattle, WA, USA

U WesselmannThe Johns Hopkins University School of Medicine, Baltimore, MD, USA

G WestheimerUniversity of California, Berkeley, CA, USA

Contributors to All Volumes xxix

K N WestlundUniversity of Texas Medical Branch, Galveston, TX, USA

H E WheatUniversity of Melbourne, Parkville, Vic, Australia

M C WhiteheadUniversity of California, San Diego, La Jolla, CA, USA

M C WhitmanYale University School of Medicine, New Haven, CT, USA

M WickleinUniversity College London, London, UK

M C WiestDuke University, Durham, NC, USA

J C WillerINSERM U-731, Paris, France

M A WillisCase Western Reserve University, Cleveland, OH, USA

W D Willis JrUniversity of Texas Medical Branch, Galveston, TX, USA

J F WillottUniversity of South Florida, Tampa, FL

D A WilsonUniversity of Oklahoma, Norman, OK, USA

M WilsonUniversity of California, Davis, CA, USA

J M WolfeBrigham and Women’s Hospital & Harvard Medical School, Cambridge, MA, USA

J N WoodUniversity College London, London, UK

H WassleMax-Planck-Institute for Brain Research, Frankfurt/Main, Germany

J E YackCarleton University, Ottawa, ON, Canada

T YamamotoOsaka University, Osaka, Japan

R YangUniversity of Denver, Denver, CO, USA

K-W YauJohns Hopkins University School of Medicine, Baltimore, MD, USA

R P YezierskiComprehensive Center for Pain Research and The McKnight Brain Institute, University of Florida, Gainesville, FL, USA

W A YostLoyola University Chicago, Chicago, IL, USA

J M YoungFred Hutchinson Cancer Research Center, Seattle, WA, USA

xxx Contributors to All Volumes

G YovelTel Aviv University, Tel Aviv, Israel

A ZainosUniversidad Nacional Autonoma de Mexico, Mexico

H U ZeilhoferUniversity of Zurich, Zurich, Switzerland

D M ZeitlerNew York University School of Medicine, New York, NY, USA

F G ZengUniversity of California, Irvine, CA, USA

J-K ZubietaUniversity of Michigan, Ann Arbor, MI, USA

F ZufallUniversity of Maryland School of Medicine, Baltimore, MD, USA

L J ZwiebelVanderbilt University, Nashville, TN, USA

Contributors to All Volumes xxxi

Introduction to Volume 6

When we read the word ‘touch’ we think of hands: an empty hand extended in friendship, a lover’s caress, a

mother cuddling her infant child, the ‘high fives’ of teammates slapping each others’ hand celebrating victory in

sporting events. Touch means contact between our body and another person, object, plant, or animal. Contact

deforms the skin in ways that convey information to the brain about the identity of external entities; their size,

shape, compliance, texture, and temperature. Because the hand is the most densely innervated part of the

human body, it is not surprising that we use the hand more than any other body part to glean information

beyond ourselves that allows us to interact with our surroundings in meaningful ways.The sense of touch is unique in that it is not merely receptive, but it is crucial for guiding motor behavior.

We interact with the environment to acquire tactile information, and use that knowledge to modify the world.

Touch is essential for guiding the behavior of our hands. This dual sensory and motor role of touch has been

beautifully depicted by the seventeenth century Spanish artist Jusepe de Ribera (1591–1652) in a canvas

entitled ‘The Sense of Touch’ (Figure 1). Here we see a blind man holding a sculpted head in his left hand as he

palpates the face with his right hand. The surface contours and features of the head that are sensed by receptors

in the hand are transformed by the brain into a mental image of the object. However, this object is not found in

this form in nature, but was actually created by the hand of the sculptor, guided by touch. The painting itself is,

Figure 1 The Sense of Touch, c. 1615–16 by Jusepe de Ribera. Reproduced with permission from The Norton Simon

Foundation ª 2002.

xxxiii

of course, another work of the hand in which direct vision of the world, or the world of the artist’s imagination,is transformed by the actions of the hand into an image on canvas delineated by skilled hand movements.

In this volume we have attempted to summarize current knowledge about the sense of touch. The variouschapters have been selected to provide insight into the neurobiological mechanisms that underlie tactilesensations. We begin with an examination of the psychophysics of touch, the ability of humans to perceivethe world through deformation of the skin by external stimuli. We then describe the various sensory receptorsin the skin that transduce mechanical and thermal events into a pattern of nerve impulses that convey discretebits of tactile information to the central nervous system. We explore how the information is integrated andtransformed in the brain to give rise to conscious sensations of touch. In higher centers of the cerebral cortex,the sensory signals are further transformed by cognitive mechanisms such as attention, experience, andprediction, as well as by motor behaviors as we interact with the world in meaningful and useful manners.These subjective actions allow us to feel what we are particularly interested in, and to shape the information inways that are useful for accomplishing the desired goals.

Most of the chapters in this volume deal with the sensory capabilities of the hand of humans and monkeys.However, we have also included chapters on touch in other parts of the body, and in other species, to highlightthose properties of touch unique to primate hands, and those generalizable to touch as a sensory modality. Wehope that this volume will provide a valuable resource to students encountering this subject for the first time, aswell as to the expert audience who have contributed so much to our understanding of the sense of touch.

Jon H. Kaas

xxxiv Introduction to Volume 6

Dedication

During the course of one year, the somatosensory community lost three major colleagues: Kenneth Johnson,Stanley Bolanowski, and Tim Pons. All three had been invited to contribute to this Volume, but sadly theypassed away before this was possible. References to their work abound in various chapters, but do not substitutefor their unique achievements and voices. We all miss them dearly, and would like to dedicate this Volume totheir memory.

Esther P. Gardner and Jon H. Kaas

xxxv