Contributors Jocelyne Bachevalier Emory University, Atlanta, GA Anirvan Ghosh University of California, San Diego, James F. Baker Northwestern University Medical School, Chicago, IL Andrea C. Gore University of Texas at Austin Floyd E. Bloom The Scripps Research Institute, La Austin, TX Jolla, CA Jacqueline P. Gottlieb Columbia University, New Scott T. Brady University of Illinois at Chicago, York. ny Chicago, IL James L. Gould Princeton University, Princeton, NJ Marianne Bronner-Fraser Caltech, Pasadena, CA Sten Grillner Karolinska Institute, Stockholm Peter J. Brophy University of Edinburgh, Edinburgh, Sweden William A. Harris University of Cambridge, M. Christian Brown Harvard Medical School Cambridge, United Kingdom Boston ma Volker Hartenstein University of California, Los Steven J Burden NYU Medical Center, New York, NY Angeles, CA Ania Busza University of Massachusetts Medical Mary E. Hatten The Rockefeller University, New School, Worcester, MA John H. Byrne University of Texas Medical School at Stewart H. Hendry Johns Hopkins University, Houston, Houston, TX David N. Caplan Massachusetts General Hospital, J. Allan Hobson Harvard Medical School, Boston, Boston, ma MA J. Patrick Card University of Pittsburgh, Pittsburgh, Patrick R Hof Mount Sinai School of Medicine, New York,, NY Luz Claudio Mount Sinai School of Medicine, New Steven S. Hsiao Johns Hopkins University, York, NY Baltimore mD Hollis Cline Cold Spring Harbor Laboratory, Cold Yuh-Nung Jan University ofCalifornia,San Spring Harbor, NY Francisco, San Francisco, CA Carol L. Colby University of Pittsburgh, Pittsburgh, Jon H. Kaas Vanderbilt University, Nashville, TN Sabine Kastner Princeton University, Princeton, David R Colman Montreal Neurological Institute, Montreal, Quebec, Canada Grahame Kidd Lerner Research Institute, Cleveland Ariel Y. Deutch Vanderbilt University Medical Clinic, Cleveland, OH Chris Kintner The Salk Institute for Biological Howard B Eichenbaum Boston University, Boston Studies, San Diego, CA Christof Koch California Institute of Technology, Patrick Emery University of Massachusetts Medical Pasadena, CA School, Worcester Alex Kolodkin Johns Hopkins University School of arry J. Everitt University of Cambridge, Cambridge, Medicine, Baltimore, MD United Kingdom Eric I Knudsen Stanford University School of Medi- Jack L Feldman David Geffen School of Medicine at cine, Stanford, CA UCLA, Los Angeles, CA George F Koob The Scripps Research Institute, La Mary Kay Floeter National Institute of Neurological Jolla, CA Disorders and Stroke, Bethesda, MD Richard J. Krauzlis The Salk Institute, La Jolla, CA
Contributors xix Jocelyne Bachevalier Emory University, Atlanta, GA James F. Baker Northwestern University Medical School, Chicago, IL Floyd E. Bloom The Scripps Research Institute, La Jolla, CA Scott T. Brady University of Illinois at Chicago, Chicago, IL Marianne Bronner-Fraser Caltech, Pasadena, CA Peter J. Brophy University of Edinburgh, Edinburgh, Scotland M. Christian Brown Harvard Medical School, Boston, MA Steven J. Burden NYU Medical Center, New York, NY Ania Busza University of Massachusetts Medical School, Worcester, MA John H. Byrne University of Texas Medical School at Houston, Houston, TX David N. Caplan Massachusetts General Hospital, Boston, MA J. Patrick Card University of Pittsburgh, Pittsburgh, PA Luz Claudio Mount Sinai School of Medicine, New York, NY Hollis Cline Cold Spring Harbor Laboratory, Cold Spring Harbor, NY Carol L. Colby University of Pittsburgh, Pittsburgh, PA David R. Colman Montreal Neurological Institute, Montreal, Quebec, Canada Ariel Y. Deutch Vanderbilt University Medical Center, Nashville, TN Howard B. Eichenbaum Boston University, Boston, MA Patrick Emery University of Massachusetts Medical School, Worcester, MA Barry J. Everitt University of Cambridge, Cambridge, United Kingdom Jack L. Feldman David Geffen School of Medicine at UCLA, Los Angeles, CA Mary Kay Floeter National Institute of Neurological Disorders and Stroke, Bethesda, MD Anirvan Ghosh University of California, San Diego, La Jolla, CA Andrea C. Gore University of Texas at Austin, Austin, TX Jacqueline P. Gottlieb Columbia University, New York, NY James L. Gould Princeton University, Princeton, NJ Sten Grillner Karolinska Institute, Stockholm, Sweden William A. Harris University of Cambridge, Cambridge, United Kingdom Volker Hartenstein University of California, Los Angeles, CA Mary E. Hatten The Rockefeller University, New York, NY Stewart H. Hendry Johns Hopkins University, Baltimore, MD J. Allan Hobson Harvard Medical School, Boston, MA Patrick R. Hof Mount Sinai School of Medicine, New York, NY Steven S. Hsiao Johns Hopkins University, Baltimore, MD Yuh-Nung Jan University of California, San Francisco, San Francisco, CA Jon H. Kaas Vanderbilt University, Nashville, TN Sabine Kastner Princeton University, Princeton, NJ Grahame Kidd Lerner Research Institute, Cleveland Clinic, Cleveland, OH Chris Kintner The Salk Institute for Biological Studies, San Diego, CA Christof Koch California Institute of Technology, Pasadena, CA Alex Kolodkin Johns Hopkins University School of Medicine, Baltimore, MD Eric I. Knudsen Stanford University School of Medicine, Stanford, CA George F. Koob The Scripps Research Institute, La Jolla, CA Richard J. Krauzlis The Salk Institute, La Jolla, CA
CONTRIBUTORS Jeff w. Lichtman Molecular and Cellular Biology, Robert H. Roth Yale University School Harvard University, Cambridge, MA New Haven, CT John C Longhurst University of California, Irvine, Joseph Santos-Sacchi Yale University School of Medicine, New Haven, CT Andrew Lumsden MRC Centre for Developmental Peter Scheiffele Columbia University, New York, NY Neurobiology, Kings College London, U.K Marc H. Schieber University of rochester School of Pierre J. Magistretti University of Lausanne, Medicine and Dentistry, Rochester, NY Lausanne, Switzerland Howard Schulman Stanford University Medical Joseph R Manns Emory University, Altanta, GA Center, Stanford, CA Michael D Mauk University of Texas Health Science Thomas L Schwarz Childrens Hospital, Boston, MA Center at Houston Houston Tx Kristin Scott University of California, Berkeley, David A. McCormick Yale University School of Medicine, New Haven, CT Gordon M. Shepherd Yale University School of Donald R McCrimmon Feinberg School of Medi- Medicine. New Haven, CT cine, Northwestern University, Chicago, IL Robert Stickgold Harvard Medical School, Boston, George Z Mentis The Porter Neuroscience Center, MA INDS, NIH Bethesda, MD Edward M. Stricker University of Pittsburg, Earl K. Miller Massachusetts Institute of Technol- Pittsburg, PA ogy, Cambridge, MA arry w. Swanson University of Southern Jonathan W Mink University of Rochester School of California, Los Angeles, CA Medicine and Dentistry, Rochester, NY Juan C. Tapia Molecular and Cellular Biology, Robert Y Moore University of Pittsburgh School of Harvard University, Cambridge, MA Medicine, Pittsburgh, PA Marc Tessier-Lavigne Genentech, Inc, South San Esther A Nimchinsky Rutgers University, Newark, Francisco, CA W. Thomas Thach Washington University School of Dennis D. M. OLeary The Salk Institute, La Jolla Medicine, St Louis, MO R oger B H. Tootell Martinos Center for Biomedical Carl R. Olson Carnegie Mellon University, Imaging, Hosp Pittsburgh, PA Charlestown, MA Ronald w. Oppenheim Wake Forest University Bruce D. Trapp Cleveland Clinic Foundation, School of Medicine, Winston-Salem, NC Cleveland Clinic, Cleveland, OH Edward F. Pace-Schott Harvard Medical School, Leslie G. Ungerlieder National Institute of Mental Boston, Ma Health, Bethesda, MD Luiz Pessoa Department of Psychological and Brain w. Martin Usrey University of California, Davis, CA Sciences Indiana University, Bloomington, Bloom- Jean de vellis University of California, Los Angeles, ington, IN CA Terry L Powley Purdue University, West Lafayette, Joseph G. Verbalis Georgetown University Medical enter Todd M. Preuss University of Louisiana at Lafayette, Christopher S von Bartheld University of Nevada New Iberia, LA School of Medicine, Reno, NV Peter R Rapp Mount Sinai School of Medicine, New Jonathan D. Wallis University of California at Berkeley, Berkeley, CA R Clay Reid Harvard Medical School, Boston, MA M. Neal Waxham University of Texas Health Science Steven M. Reppert University of Massachusetts Center, Houston, TX Medical School, Worcester MA David R. Weaver University of Massachusetts John H. Reynolds The Salk Institute, La Jolla, CA Medical School, Worcester, MA Trevor w. Robbins University of Cambridge, Stephen C Woods University of Cincinnati Medical Cambridge, United Kingdom Center, Cincinnati, OH
Jeff W. Lichtman Molecular and Cellular Biology, Harvard University, Cambridge, MA John C. Longhurst University of California, Irvine, CA Andrew Lumsden MRC Centre for Developmental Neurobiology, King’s College London, U.K. Pierre J. Magistretti University of Lausanne, Lausanne, Switzerland Joseph R. Manns Emory University, Altanta, GA Michael D. Mauk University of Texas Health Science Center at Houston, Houston, TX David A. McCormick Yale University School of Medicine, New Haven, CT Donald R. McCrimmon Feinberg School of Medicine, Northwestern University, Chicago, IL George Z. Mentis The Porter Neuroscience Center, NINDS, NIH, Bethesda, MD Earl K. Miller Massachusetts Institute of Technology, Cambridge, MA Jonathan W. Mink University of Rochester School of Medicine and Dentistry, Rochester, NY Robert Y. Moore University of Pittsburgh School of Medicine, Pittsburgh, PA Esther A. Nimchinsky Rutgers University, Newark, NJ Dennis D. M. O’Leary The Salk Institute, La Jolla, CA Carl R. Olson Carnegie Mellon University, Pittsburgh, PA Ronald W. Oppenheim Wake Forest University School of Medicine, Winston-Salem, NC Edward F. Pace-Schott Harvard Medical School, Boston, MA Luiz Pessoa Department of Psychological and Brain Sciences Indiana University, Bloomington, Bloomington, IN Terry L. Powley Purdue University, West Lafayette, IN Todd M. Preuss University of Louisiana at Lafayette, New Iberia, LA Peter R. Rapp Mount Sinai School of Medicine, New York, NY R. Clay Reid Harvard Medical School, Boston, MA Steven M. Reppert University of Massachusetts Medical School, Worcester, MA John H. Reynolds The Salk Institute, La Jolla, CA Trevor W. Robbins University of Cambridge, Cambridge, United Kingdom Robert H. Roth Yale University School of Medicine, New Haven, CT Joseph Santos-Sacchi Yale University School of Medicine, New Haven, CT Peter Scheiffele Columbia University, New York, NY Marc H. Schieber University of Rochester School of Medicine and Dentistry, Rochester, NY Howard Schulman Stanford University Medical Center, Stanford, CA Thomas L. Schwarz Children’s Hospital, Boston, MA Kristin Scott University of California, Berkeley, Berkeley, CA Gordon M. Shepherd Yale University School of Medicine, New Haven, CT Robert Stickgold Harvard Medical School, Boston, MA Edward M. Stricker University of Pittsburg, Pittsburg, PA Larry W. Swanson University of Southern California, Los Angeles, CA Juan C. Tapia Molecular and Cellular Biology, Harvard University, Cambridge, MA Marc Tessier-Lavigne Genentech, Inc., South San Francisco, CA W. Thomas Thach Washington University School of Medicine, St. Louis, MO Roger B.H. Tootell Martinos Center for Biomedical Imaging, Massachusetts General Hospital, Charlestown, MA Bruce D. Trapp Cleveland Clinic Foundation, Cleveland Clinic, Cleveland, OH Leslie G. Ungerlieder National Institute of Mental Health, Bethesda, MD W. Martin Usrey University of California, Davis, CA Jean de Vellis University of California, Los Angeles, CA Joseph G. Verbalis Georgetown University Medical Center, Washington, DC Christopher S. von Bartheld University of Nevada School of Medicine, Reno, NV Jonathan D. Wallis University of California at Berkeley, Berkeley, CA M. Neal Waxham University of Texas Health Science Center, Houston, TX David R. Weaver University of Massachusetts Medical School, Worcester, MA Stephen C. Woods University of Cincinnati Medical Center, Cincinnati, OH xx CONTRIBUTORS
SECTION I NEUROSCIENCE
SECTION I NEUROSCIENCE
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CHAPTER Fundamentals of neuroscience A BRIEF HISTORY OF NEUROSCIENCE participated at the societys 36th annual meeting at which 14, 268 research presentations were made The field of knowledge described in this book is euroscience, the multidisciplinary sciences that analyze the nervous system to understand the biological basis for behavior. Modern studies of the nervous system THE TERMINOLOGY OF NERVOUS have been ongoing since the middle of the nineteenth SYSTEMS IS HIERARCHICAL century. Neuroanatomists studied the brains shape, DISTRIBUTED, DESCRIPTIVE AND s cellular structure, and its circuitry; neurochemists HISTORICALLY BASED studied the brains chemical composition, its lipid and proteins; neurophysiologists studied the brains Beginning students of neuroscience justifiably could bioelectric properties; and psychologists and neu- find themselves confused. Nervous systems of many ropsychologists investigated the organization and organisms have their cell assemblies and macroscopi neural substrates of behavior and cognition cally visible components named by multiple overlap- The term neuroscience was introduced in the mid ping and often synonymous terms With a necessarily 1960s, to signal the beginning of an era in which each gracious view to the past, this confusing terminolo of these disciplines would work togethe er Co0 could be viewed as the intellectual cost of focused tively, sharing a common language, common concepts, discourse with predecessors in the enterprise. The and a common goal-to understand the structure and nervous systems of invertebrate organisms often are function of the normal and abnormal brain. Neuro- designated for their spatially directed collections of science today spans a wide range of research endeav- neurons responsible for local control of operations ors from the molecular biology of nerve cells (i.e, the such as the thoracic or abdominal ganglia, which genes encoding the proteins needed for nervous system receive sensations and direct motoric responses for function) to the biological basis of normal and disor- specific body segments, all under the general control dered behavior, emotion, and cognition(i. e, the mental of a cephalic ganglion whose role includes sensing the properties by which individuals interact with each external environment other and with their environments). For a more com- In vertebrates, the components of the nervous plete, but concise, history of the neurosciences see system were named for both their appearance and Kandel and Squire(2000 their location. As noted by Swanson, and expanded Neuroscience is currently one of the most rapidly upon in Chapter 2 of this volume, the names of the growing areas of science. Indeed, the brain is some- major parts of the brain were based on creative inter times referred to as the last frontier of biology. In 1971, pretations of early dissectors of the brain, attributing 1100 scientists convened at the first annual meeting of names to brain segments based on their appearance ir the Society for Neuroscience. In 2006, 25, 785 scientists the freshly dissected state: hippocampus(shaped like Neuroscience. Third Edition 92008,2003.1999 Elsevier Inc
CHAPTER 1 Fundamentals of Neuroscience A BRIEF HISTORY OF NEUROSCIENCE The fi eld of knowledge described in this book is neuroscience, the multidisciplinary sciences that analyze the nervous system to understand the biological basis for behavior. Modern studies of the nervous system have been ongoing since the middle of the nineteenth century. Neuroanatomists studied the brain’s shape, its cellular structure, and its circuitry; neurochemists studied the brain’s chemical composition, its lipids and proteins; neurophysiologists studied the brain’s bioelectric properties; and psychologists and neuropsychologists investigated the organization and neural substrates of behavior and cognition. The term neuroscience was introduced in the mid- 1960s, to signal the beginning of an era in which each of these disciplines would work together cooperatively, sharing a common language, common concepts, and a common goal—to understand the structure and function of the normal and abnormal brain. Neuroscience today spans a wide range of research endeavors from the molecular biology of nerve cells (i.e., the genes encoding the proteins needed for nervous system function) to the biological basis of normal and disordered behavior, emotion, and cognition (i.e., the mental properties by which individuals interact with each other and with their environments). For a more complete, but concise, history of the neurosciences see Kandel and Squire (2000). Neuroscience is currently one of the most rapidly growing areas of science. Indeed, the brain is sometimes referred to as the last frontier of biology. In 1971, 1100 scientists convened at the fi rst annual meeting of the Society for Neuroscience. In 2006, 25,785 scientists participated at the society’s 36th annual meeting at which 14,268 research presentations were made. THE TERMINOLOGY OF NERVOUS SYSTEMS IS HIERARCHICAL, DISTRIBUTED, DESCRIPTIVE, AND HISTORICALLY BASED Beginning students of neuroscience justifi ably could fi nd themselves confused. Nervous systems of many organisms have their cell assemblies and macroscopically visible components named by multiple overlapping and often synonymous terms. With a necessarily gracious view to the past, this confusing terminology could be viewed as the intellectual cost of focused discourse with predecessors in the enterprise. The nervous systems of invertebrate organisms often are designated for their spatially directed collections of neurons responsible for local control of operations, such as the thoracic or abdominal ganglia, which receive sensations and direct motoric responses for specifi c body segments, all under the general control of a cephalic ganglion whose role includes sensing the external environment. In vertebrates, the components of the nervous system were named for both their appearance and their location. As noted by Swanson, and expanded upon in Chapter 2 of this volume, the names of the major parts of the brain were based on creative interpretations of early dissectors of the brain, attributing names to brain segments based on their appearance in the freshly dissected state: hippocampus (shaped like Fundamental Neuroscience, Third Edition 3 © 2008, 2003, 1999 Elsevier Inc