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PRINCIPLES
OF NEUR AL
SCIENCE

Columns II (left) and IV (right) of the Edwin Smith Surgical Papyrus
This papyrus, transcribed in the Seventeenth Century B.C., is a medical treatise that
contains the earliest reference to the brain anywhere in human records. According to
James Breasted, who translated and published the document in 1930, the word brain
occurs only 8 times in ancient Egyptian, 6 of them on these pages. The papyrus
describes here the symptoms, diagnosis, and prognosis of two patients with compound
fractures of the skull, and compares the surface of the brain to “those ripples that happen in copper through smelting, with a thing in it that throbs and flutters under your
fingers like the weak spot of the crown of a boy before it becomes whole for him.” The
red ink highlights the patients’ ailments and their prognoses. (Reproduced, with permission, from the New York Academy of Medicine Library.)

Men ought to know that from the brain, and
from the brain only, arise our pleasures, joys,
laughter and jests, as well as our sorrows,
pains, griefs and tears. Through it, in
particular, we think, see, hear, and distinguish
the ugly from the beautiful, the bad from the
good, the pleasant from the unpleasant. . . . It
is the same thing which makes us mad or
delirious, inspires us with dread and fear,
whether by night or by day, brings
sleeplessness, inopportune mistakes, aimless
anxieties, absent-mindedness, and acts that are
contrary to habit. These things that we suffer
all come from the brain, when it is not healthy,
but becomes abnormally hot, cold, moist, or
dry, or suffers any other unnatural affection to
which it was not accustomed. Madness comes
from its moistness. When the brain is
abnormally moist, of necessity it moves, and
when it moves neither sight nor hearing are
still, but we see or hear now one thing and
now another, and the tongue speaks in
accordance with the things seen and heard on
any occasion. But when the brain is still, a man
can think properly.
attributed to Hippocrates
Fifth Century, B.C.

Reproduced, with permission, from The Sacred Disease,
in Hippocrates, Vol. 2, page 175, translated by W.H.S.
Jones, London and New York: William Heinemann and
Harvard University Press. 1923.

PRINCIPLES
OF NEUR AL
SCIENCE
Fifth Edition
Edited by

ERIC R. KANDEL
JAMES H. SCHWARTZ
THOMAS M. JESSELL
STEVEN A. SIEGELBAUM
A. J. HUDSPETH
Art Editor

Sarah Mack

New York Chicago San Francisco Lisbon London Madrid Mexico City
Milan New Delhi San Juan Seoul Singapore Sydney Toronto

Copyright © 2013 by The McGraw-Hill Companies, Inc. All rights reserved. Except as permitted under the United States Copyright Act of 1976, no part
of this publication may be reproduced or distributed in any form or by any means, or stored in a database or retrieval system, without the prior written
permission of the publisher.
ISBN: 978-0-07-181001-2
MHID: 0-07-181001-3
The material in this eBook also appears in the print version of this title: ISBN: 978-0-07-139011-8,
MHID: 0-07-139011-1.
All trademarks are trademarks of their respective owners. Rather than put a trademark symbol after every occurrence of a trademarked name, we use names
in an editorial fashion only, and to the benefit of the trademark owner, with no intention of infringement of the trademark. Where such designations appear in
this book, they have been printed with initial caps.
McGraw-Hill eBooks are available at special quantity discounts to use as premiums and sales promotions, or for use in corporate training programs. To
contact a representative please e-mail us at bulksales@mcgraw-hill.com.
Previous editions copyright © 2000 by McGraw-Hill Companies; © 1991 by Appleton & Lange.
Cover image: This image is a lithograph by F. Schima from a drawing by Sigmund Freud of the spinal ganglion of the lamprey Petromyzon. Before he
discovered the unconscious, Freud had a promising career as a neural scientist. The cover thus recognizes that, a century after Freud’s discovery, progress
in the study of cognition has reemphasized the importance of unconscious mental processes for perception and action. (Reproduced, with permission, from
Sigmund Freud, “Über Spinalganglien und Rückenmark der Petromyzon,” Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Classe der
Kaiserlichen Akademie der Wissenschaften, LXXVIII. Band I. Abtheilung, 1878, copyright New York Academy of Medicine.)
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WE WISH TO DEDICATE THIS
FIFTH EDITION of Principles of
Neural Science to our friend and
colleague, James H. Schwartz, one
of the founding editors who died
on March 13, 2006. Jimmy was an
outstanding neuroscientist and
scholar. His talent for science and
his extraordinary erudition were
evident from his days as a medical
student at New York University.
While at NYU he worked with
Werner Maas in the microbiology
department and carried out
an important set of studies on
feedback inhibition in bacterial
James H. Schwartz
metabolism. This work was so
1933–2006
impressive that upon completing
medical school, Jimmy was nominated for the highly selective graduate
program in biology that had just been established at The Rockefeller
University by Detlev Bronk. By the time Jimmy obtained his Ph.D. in
Fritz Lippman’s laboratory and graduated from Rockefeller in 1964, he
had established himself as an outstanding biochemist. He was therefore
eagerly recruited back to NYU in 1965 as an Assistant Professor in the
Department of Microbiology.
There Jimmy turned to studying the nerve cells of the snail Aplysia,
which were so large and uniquely identifiable that they seemed likely
candidates for a study of neuronal biochemical identity. The immediate success of his initial studies encouraged him to devote himself
completely to the nervous system. He rapidly became one of the leading biochemists on the nervous system and one of the leading thinkers
regarding the relationship of brain to behavior.
The idea of going from molecules to behavior was the organizing
theme of the first edition of Principles of Neural Science, which Jimmy
co-edited. He simply loved working on Principles. A superb writer, he
demanded precision in language both in himself and in others. This
made him an exceptional editor. He read and avidly edited every chapter. In addition, Jimmy contributed his sense of historical scholarship.
It was his idea to open Principles with the images of hieroglyphics from
the Egyptian papyrus, the earliest reference to the brain in human
record, which we include as the opening images in this edition as well.
But perhaps most importantly, Jimmy championed the idea that this
book should delineate fundamental principles rather than serve as an
encyclopedia of facts. Thus, Jimmy’s vision and editorial skill greatly
enriched each of the five editions. In his absence we have striven to
make the final product an edition that will continue to meet the high
standards of readability and scholarship he set for all of us.

Notice
Medicine is an ever-changing science. As new research and clinical experience broaden our
knowledge, changes in treatment and drug therapy are required. The authors and the publisher of
this work have checked with sources believed to be reliable in their efforts to provide information
that is complete and generally in accord with the standards accepted at the time of publication.
However, in view of the possibility of human error or changes in medical sciences, neither
the authors nor the publisher nor any other party who has been involved in the preparation
or publication of this work warrants that the information contained herein is in every respect
accurate or complete, and they disclaim all responsibility for any errors or omissions or for the
results obtained from use of the information contained in this work. Readers are encouraged
to confirm the information contained herein with other sources. For example and in particular,
readers are advised to check the product information sheet included in the package of each drug
they plan to administer to be certain that the information contained in this work is accurate
and that changes have not been made in the recommended dose or in the contraindications for
administration. This recommendation is of particular importance in connection with new or
infrequently used drugs.


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