THE SENSES:A COMPREHENSIVE

REFERENCE

THE SENSES:A COMPREHENSIVE

REFERENCE

Volume 4OLFACTION AND TASTE

Volume Editors

Dr Stuart FiresteinColumbia University, New York, NY, USA

Dr Gary K. BeauchampMonell Chemical Senses Center, Philadelphia, PA, 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

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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

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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

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Contents

Contents of All Volumes ix

Contributors to All Volumes xv

Introduction to Volume 4 xxxv

Dedication xxxvii

Olfaction & Taste

4.01 Phylogeny of Chemical Sensitivity 1B W Ache, University of Florida, Gainesville, FL, USAJ M Young, Fred Hutchinson Cancer Research Center, Seattle, WA, USA

4.02 Chemistry of Gustatory Stimuli 27G E DuBois, The Coca-Cola Company, Atlanta, GA, USAJ DeSimone and V Lyall, Virginia Commonwealth University, Richmond, VA, USA

4.03 Insect Gustatory Systems 75John I Glendinning, Barnard College, Columbia University, New York, NY, USA

4.04 Aquatic Animal Models in the Study of Chemoreception 97J Caprio, Louisiana State University, Baton Rouge, LA, USAC D Derby, Georgia State University, Atlanta, GA, USA

4.05 Ultrastructure of Taste Buds 135J C Kinnamon and R Yang, University of Denver, Denver, CO, USA

4.06 Development of the Taste System 157Robin F Krimm, University of Louisville School of Medicine, Louisville, KY, USALinda A Barlow, University of Colorado School of Medicine, Aurora, CO, USA

4.07 The Sweet Taste of Childhood 183J A Mennella, Monell Chemical Senses Center, Philadelphia, PA, USA

4.08 Taste Analgesia in Newborns 189V Anseloni, University of Maryland Dental School, Baltimore, MD, USAM Ennis, University of Tennessee Health Science Center, Memphis, TN, USA

4.09 Taste Receptors 197M Max, Mount Sinai School of Medicine, New York, NY, USAW Meyerhof, German Institute of Human Nutrition Potsdam-Rehbruecke, Nuthetal, Germany

4.10 Taste Transduction 219S C Kinnamon, Colorado State University, Fort Collins, CO, USAR F Margolskee, Mount Sinai School of Medicine, New York, NY, USA

v

4.11 Gustatory Pathways in Fish and Mammals 237M C Whitehead, University of California, San Diego, La Jolla, CA, USAT E Finger, University of Colorado School of Medicine, Aurora, CO, USA

4.12 Neurotransmitters in the Taste Pathway 261R M Bradley, University of Michigan, Ann Arbor, MI, USA

4.13 Functional Magnetic Resonance Imaging (fMRI) Study of Taste 271A Faurion, Neurobiologie Sensorielle, NOPA-NBS, INRA, Jouy en Josas, FranceT Kobayakawa, National Institute of Advanced Industrial Science and Technology (AIST),Tsukuba, JapanB Cerf-Ducastel, San Diego State University, San Diego, CA, USA

4.14 Amiloride-Sensitive Ion Channels 281J A DeSimone and V Lyall, Virginia Commonwealth University, Richmond, VA, USA

4.15 Central Neural Processing of Taste Information 289D V Smith, The University of Tennessee College of Medicine, Memphis, TN, USAS P Travers, The Ohio State University, Columbus, OH, USA

4.16 Neural Ensembles in Taste Coding 329A Fontanini, S E Grossman, B A Revill, and D B Katz, Brandeis University, Waltham, MA, USA

4.17 A Perspective on Chemosensory Quality Coding 339M E Frank, University of Connecticut Health Center, Farmington, CT, USA

4.18 Oral Chemesthesis and Taste 345C T Simons, Global Research and Development Center, Cincinnati, OH, USAE Carstens, University of California, Davis, CA, USA

4.19 Genetics and Evolution of Taste 371J D Boughter, Jr., University of Tennessee Health Science Center, Memphis, TN, USAA A Bachmanov, Monell Chemical Senses Center, Philadelphia, PA, USA

4.20 Propylthiouracil (PROP) Taste 391D J Snyder, Yale University, New Haven, CT, USAV B Duffy, University of Connecticut, Storrs, CT, USAJ E Hayes, Brown University, Providence, RI, USAL M Bartoshuk, University of Florida, Gainesville, FL, USA

4.21 Salt Taste 401G K Beauchamp and L J Stein, Monell Chemical Senses Center, Philadelphia, PA, USA

4.22 Behavioral Analysis of Taste Function in Rodent Models 409S J St. John, Rollins College, Winter Park, FL, USAA C Spector, The Florida State University, Tallahassee, FL, USA

4.23 Flavor Aversion Learning 429I L Bernstein, University of Washington, Seattle, WA, USA

4.24 Roles of Taste in Feeding and Reward 437T Yamamoto and T Shimura, Osaka University, Osaka, Japan

4.25 Dopamine Release by Sucrose 459A Hajnal and R Norgren, Milton S. Hershey Medical Center, Hershey, PA, USA

4.26 The Representation of Flavor in the Brain 469E T Rolls, University of Oxford, Oxford, UK

4.27 The Aging Gustatory System 479S S Schiffman, Duke University Medical Center, Durham, NC, USA

4.28 Signal Transduction in the Olfactory Receptor Cell 499H Takeuchi and T Kurahashi, Osaka University, Osaka, Japan

vi Contents

4.29 Olfactory Cyclic Nucleotide-Gated Ion Channels 511M-C Broillet, University of Lausanne, Lausanne, Switzerland

4.30 Structure, Expression, and Function of Olfactory Receptors 527K Touhara, The University of Tokyo, Chiba, Japan

4.31 Regulation of Expression of Odorant Receptor Genes 545A Tsuboi and H Sakano, University of Tokyo, Tokyo, Japan

4.32 Genomics of Odor Receptors in Zebrafish 553J Ngai and T S Alioto, University of California, Berkeley, CA, USA

4.33 Genomics of Invertebrate Olfaction 561J D Bohbot, R J Pitts, and L J Zwiebel, Vanderbilt University, Nashville, TN, USA

4.34 Regeneration of the Olfactory Epithelium 591J E Schwob, Tufts University School of Medicine, Boston, MA, USAR M Costanzo, Virginia Commonwealth University, Richmond, VA, USA

4.35 Regeneration in the Olfactory Bulb 613P-M Lledo, Pasteur Institute, Paris, France

4.36 Architecture of the Olfactory Bulb 623C A Greer, M C Whitman, L Rela, F Imamura, and D Rodriguez Gil,Yale University School of Medicine, New Haven, CT, USA

4.37 Physiology of the Main Olfactory Bulb 641M Ennis, University of Tennessee Health Science Center, Memphis, TN, USAA Hayar, University of Arkansas for Medical Sciences, Little Rock, AR, USA

4.38 Olfactory Cortex 687D A Wilson, University of Oklahoma, Norman, OK, USA

4.39 Modeling of Olfactory Processing 707C Linster and T A Cleland, Cornell University, Ithaca, NY, USA

4.40 Understanding Olfactory Coding via an Analysis of Odorant-Evoked GlomerularResponse Maps 719B A Johnson and M Leon, University of California, Irvine, CA, USA

4.41 Insect Olfaction 725G Galizia, Universitat Konstanz, Konstanz, Germany

4.42 Odor Plumes and Animal Orientation 771M A Willis, Case Western Reserve University, Cleveland, OH, USA

4.43 Accessory Olfactory System 783Frank Zufall, Trese Leinders-Zufall, and Adam C Puche,University of Maryland School of Medicine, Baltimore, MD, USA

4.44 Genomics of Vomeronasal Receptors 815I Rodriguez, University of Geneva, Geneva, Switzerland

4.45 Human Olfactory Psychophysics 823Brad Johnson, Rehan M Khan, and Noam Sobel, UC Berkeley, Berkeley, CA, USA

4.46 Disorders of Taste and Smell 859R L Doty and K Saito, University of Pennsylvania, Philadelphia, PA, USAS M Bromley, University of Pennsylvania, Philadelphia, PA, USA, UMDNJ-Robert Wood JohnsonMedical School, Camden, NJ, USA

Index 889

Contents vii

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

ix

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

x 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 xi

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

xii 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 xiii

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

xv

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

xvi 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 xvii

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

xviii 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 xix

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

xx 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 xxi

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

xxii 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 xxiii

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

xxiv 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 xxv

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

xxvi 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 xxvii

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

xxviii 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 xxix

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

xxx 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 xxxi

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

xxxii 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 xxxiii

Introduction to Volume 4

This volume of the The Senses: A Comprehensive Reference provides a current review of the chemical senses of tasteand smell. Historically, these were considered the minor senses. Descriptions of them were often combined intextbooks both because the natural stimuli are chemicals and because not much was known about themcompared with vision and audition. Presumably, they were less studied since it did not seem so bad if humanslost their ability to smell or taste; blindness and deafness were much more serious concerns. However, onemight justifiably argue that if all animal life were considered, these are the most important senses. Taste isdevoted to a single overwhelmingly important function: it insures that an organism takes in appropriatenutrients and avoids poison. The sense of smell has more varied functions. It too is involved with recognizingfood and motivating its intake but it also plays a critical role in monitoring the environment for danger and,perhaps most importantly, in regulating social and sexual activities. Thus, without these two senses mostanimals would neither eat nor mate! For humans, we have been learning more about the crucial roles taste andsmell play in regulating food choice and intake, modulating social interactions, surveying the chemicalenvironment, and providing pleasure. Loss or alteration of smell and taste are not trivial afflictions.

Fortunately, research in the chemical senses is no longer neglected. Indeed, the remarkable rate of progressmay, at first blush, even make the idea of a handbook seem absurd. By the time you read these chapters therewill be tens of new publications not covered in this book. But in fact it is all those papers that make acompendium like this useful, even if necessarily incomplete. Not long ago one could start out in the fields ofolfaction or taste and come up to speed in the literature quite easily. Indeed, for some of us that was one of theattractions of the field. The advent of new techniques, their successful application to questions in chemicalsensing, the attraction of investigators from other fields, suddenly transformed the chemical senses from themost mysterious to the most investigated of the sensory systems. Now it is critical to be able to read papers inmolecular biology, anatomy, physiology, imaging, psychophysics, genetics, bioinformatics, genomics. . .

So the value of a handbook is as a quick but inclusive reference that will bring even senior investigatorsrapidly up to speed in an unfamiliar area. In this respect, the contributors to this volume have done anadmirable job. Chapters cover all of the above topics in the context of specific systems in olfaction and taste,they cover historical literature (now anything published before 1995 it seems), and provide the kind ofbackground that will facilitate appreciation of the up-to-date advances that appear monthly, if not weekly, inour dynamic field.

This handbook will also, we hope, serve new entrants in the field, especially students and postdoctoralfellows. Each chapter has extensive citations that are an excellent guide to the current literature, and willremain so for many years. The authors have endeavored not only to review the current state of the field, butalso to identify important questions and remaining mysteries. Although there have been amazing advances inthe past decade and a half, there are even more questions, and more interesting questions, than there were whenthe last edition of this Handbook appeared.

xxxv

So, how about the next edition? What will it contain? Will it appear in print or only electromagnetically?What are the advances that will be chronicled in that edition? Who will the chapter authors be? This is aremarkable era in the chemical senses. Our bet is that it is only beginning. We thank the authors for their workto chronicle its current progress and to set the stage for future discoveries.

Gary K. Beauchamp

xxxvi Introduction to Volume 4

Dedication

David V. Smith (1943–2006)

David Smith was prominent among a cohort of taste physiologists born in the 1940s on three continents, who

have collectively defined – or trained those who defined – gustatory activity in the central nervous system.

They learned from the founders of our discipline: Yngve Zotterman, Carl Pfaffmann, Lloyd Beidler, and

Masayasu Sato. Equipped with self-styled microelectrodes, they extended recordings from peripheral axons to

the small, medial neurons of this ancient sense and taught us how taste selects from a perilous chemical

environment to compose a healthy body.Even as David made his way from his boyhood home in Memphis to study psychology in Knoxville

forces were aligning in a competition that was to become the central motif of his career: labeled-line versus

patterning. It was a binary that always concerned David, but never consumed him, as it did others of his era. It

was our primary topic of professional conversation when David and I explored the South Pacific for a month in

the early 1970s, and still the focus of a chapter we wrote three decades later. However voluminous the data,

however sophisticated the analyses, they have never been sufficient to seal a victory, and now the issue, still

unresolved if indeed a resolution exists, lies exhausted at the periphery of the field (see Chapter 4.17 A

Perspective on Chemosensory Quality Coding by M. Frank)David was always respectful of the coding arguments from both sides, as he was of those who made them.

However, he could never divorce his thinking from the central discovery of gustatory electrophysiology – that

taste cells are broadly tuned – and permit himself to favor a labeled-line strategy that would seem poorly suited

to that finding. Thus, David remained an advocate of patterning even as he vowed unsuccessfully, three times in

my presence, never to entertain the topic again.David trained with Don McBurney in psychophysics, then with Pfaffmann in the electrophysiological

techniques that became central to his life’s work. He experimented on blowflies, frogs, mice, rats, rabbits,

cats, and humans, but David’s primary focus was on the hamster hindbrain. Over 30 years, David generated a

body of data from the hamster that complemented each component that others had revealed in rats: anatomical

connections, membrane qualities, coding principles, neurotransmitters, and centrifugal influences (see Chapter 4.12

xxxvii

Neurotransmitters in the Taste Pathway by R. Bradley). David’s thinking was creative and original; histechniques precise; his analyses sophisticated and unbiased. He did not work in large groups – across all hispublications David has a mean of fewer than 1.5 co-authors – but over time he worked with scores ofcolleagues, learning their techniques, sharing his, and always pressing for deeper understanding (see Chapter4.15 Central Neural Processing of Taste Information by D. Smith and S. Travers).

His objective pursuit of information absent personal agendas made David a trusted colleague and leader.The Association for Chemoreception Sciences (AChemS; Minneapolis) elected him Executive Chairperson(now ‘President’) in 1985. David directed the neuroscience program at the University of Wyoming in the 1970sand 1980s, the Taste and Smell Center at Cincinnati in the 1980s and 1990s, served as Vice-Chair of Anatomyand Neurobiology at the University of Maryland School of Medicine in the 1990s and 2000s, then completedhis life cycle, returning to Memphis as endowed Department Chair of Anatomy and Neurobiology at theTennessee Health Science Center. In each role, David was fair, collegial, yet demanding, as he was as ExecutiveEditor of Chemical Senses.

David was in the fullness of his personal and professional life, living in the city that had called him home,surrounded by appreciative colleagues and by his wife Michiko, whose devotion David requited. In my last chatwith the healthy David in 2005, he expressed as much satisfaction with his life as his modesty would permit. Itwas all too brief.

Thomas R. Scott

xxxviii Dedication