Table Of ContentADLER’S
PHYSIOLOGY OF THE EYE
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ADLER’S
PHYSIOLOGY OF THE EYE
E L E V E N T H E D I T I O N
Editors: Managing Editors:
Leonard A. Levin MD PhD Paul L. Kaufman MD
Canada Research Chair of Ophthalmology and Visual Sciences Peter A. Duehr Professor and Chairman
University of Montreal Department of Ophthalmology and Visual Sciences
Montreal, Quebec, Canada University of Wisconsin School of Medicine and Public Health
Professor of Ophthalmology and Visual Sciences Madison, WI, USA
University of Wisconsin
Madison, WI, USA Albert Alm MD
Professor
Siv F. E. Nilsson PhD Department of Neuroscience & Ophthalmology
Lecturer, Department of Medical and Health Sciences, University Hospital
Division of Drug Research Uppsala, Sweden
Linköping University
Linköping, Sweden
James Ver Hoeve MD
Senior Scientist, Department of Ophthalmology and
Visual Sciences
University of Wisconsin School of Medicine and Public Health
Madison, WI, USA
Samuel M. Wu PhD
Camille and Raymond Hankamer Chair in Ophthalmology
Professor of Ophthalmology, Neuroscience and Physiology
Baylor College of Medicine
Houston, TX, USA
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This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted herein).
Chapter 8. Three-Dimensional Rotations of the Eye is in the public domain.
Notices
Knowledge and best practice in this feld are constantly changing. As new research and experience broaden our understanding, changes in
research methods, professional practices, or medical treatment may become necessary. Practitioners and researchers must always rely on their
own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein. In using such
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With respect to any drug or pharmaceutical products identifed, readers are advised to check the most current information provided (i) on
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instructions, or ideas contained in the material herein.
ISBN: 978-0-323-05714-1
British Library Cataloguing in Publication Data
Adler’s physiology of the eye.—11th ed.
1. Eye–Physiology.
I. Physiology of the eye II. Kaufman, Paul L. (Paul Leon), 1943- III. Alm, A. IV. Adler, Francis Heed, 1895-
Physiology of the eye.
612.8’4—dc22
Library of Congress Cataloging in Publication Data
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Preface
The 11th edition of Adler’s sees a signifcant reorganization izing and conveying vast amounts of this new information
based on function and function/structure relationships within in the context of a new organizational strategy has been both
ocular cells, tissues and organs. Prior editions had been challenging and stimulating. We hope that the readers, espe-
organized largely anatomically, but in the eight years since cially our younger scientists and clinicians who represent
the 10th edition, basic, translational and clinical knowledge the future of our feld, will fnd this approach, material and
has increased exponentially. Capturing, synthesizing, organ- presentation conducive to learning, retention and referral.
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List of Contributors
Albert Alm MD Yuzo M Chino PhD B’Ann True Gabelt MS
Professor Benedict-McFadden Professor Distinguished Scientist
Department of Neuroscience & Professor of Vision Sciences Department of Ophthalmology and
Ophthalmology College of Optometry Vision Sciences
University Hospital University of Houston University of Wisconsin
Uppsala Sweden Houston TX USA Madison WI USA
David C Beebe PhD FARVO Darlene A Dartt PhD Juana Gallar MD PhD
Janet and Bernard Becker Professor of Senior Scientist Professor of Human Physiology
Ophthalmology and Visual Science Harold F Johnson Research Scholar Instituto de Neurociencias and
Professor of Cell Biology and Schepens Eye Research Institute Facultad de Medicina
Physiology Associate Professor Universidad Miguel Hernández-CSIC
Department of Ophthalmology and Harvard Medical School San Juan de Alicante Spain
Vision Sciences Boston MA USA
Washington University Adrian Glasser PhD
Saint Louis MO USA Chanukya R Dasari MS Professor of Optometry and Vision
MD Candidate Sciences and Biomedical Engineering
Carlos Belmonte MD PhD School of Medicine Benedict/Pitts Professor
Professor of Human Physiology University of Missouri-Kansas City College of Optometry
Medical School Kansas City MO USA University of Houston
Instituto de Neurociencias de Alicante Houston TX USA
Universidad Miguel Hernandez Daniel G Dawson MD
San Juan Alicante Spain Ophthalmic Researcher Jeffrey L Goldberg MD PhD
Emory University Eye Center Assistant Professor of Ophthalmology,
David M Berson PhD Atlanta GA USA and of Neurosciences
Professor of Neuroscience Bascom Palmer Eye Institute
Department of Neuroscience Henry F Edelhauser PhD University of Miami
Brown University Professor of Ophthalmology Miami FL USA
Providence RI USA Department of Ophthalmology
Emory University Eye Center Gregory J Griepentrog MD
Sai H S Boddu BPharm MS Atlanta GA USA Clinical Instructor
PhD Candidate and Doctoral Fellow Department of Ophthalmology and
School of Pharmacy Erika D Eggers PhD Visual Sciences
University of Missouri-Kansas City Assistant Professor University of Wisconsin-Madison
Kansas City MO USA Department of Physiology Madison WI USA
University of Arizona
Jamie D Boyd PhD Tucson AZ USA Alecia K Gross PhD
Research Assistant Assistant Professor
Psychiatry, Faculty of Medicine Ione Fine PhD Department of Vision Sciences,
University of British Columbia Assistant Professor of Psychology Biochemistry and Molecular
Vancouver BC Canada Department of Psychology Genetics, Cell Biology and
University of Washington Neurobiology
Vivien Casagrande PhD Seattle WA USA The University of Alabama at
Professor, Cell & Developmental Birmingham
Biology, Psychology, and Laura J Frishman PhD Birmingham AL USA
Ophthalmology & Visual Sciences Professor of Vision Science, Optometry
Departments of Cell & Developmental and Biology Ronald S Harwerth OD PhD
Biology College of Optometry John and Rebecca Moores Professor of
Vanderbilt Medical School University of Houston Optometry
Nashville TN USA Houston TX USA College of Optometry
University of Houston
Houston TX USA
viii
List of Contributors
Horst Helbig MD Ron Krueger MD MSE Peter R MacLeish PhD
Professor of Ophthalmology Professor of Ophthalmology Professor of Neurobiology
Director of the University Eye Hospital Cleveland Clinic Lerner College of Director
Klinik und Poliklinik fur Medicine Neuroscience Institute
Augenheilkunde Medical Director Morehouse School of Medicine
Klinikum der Universitats Regensburg Department of Refractive Surgery Atlanta GA USA
Regensburg Germany Cole Eye Institute
Cleveland Clinic Foundation Clint L Makino PhD
Robert F Hess PhD DSc Cleveland OH USA Associate Professor of Ophthalmology
Professor of Ophthalmology (Neuroscience)
McGill Vision Research James A Kuchenbecker PhD Department of Ophthalmology
McGill University Senior Fellow Massachusetts Eye and Ear Infrmary
Montréal QC Canada Department of Ophthalmology & Harvard Medical School
University of Washington Boston MA USA
Jennifer Ichida PhD WA USA
Postdoctoral Fellow Katherine Mancuso PhD
Department of Ophthalmology & Trevor D Lamb BE ScD FRS FAA Postdoctoral Fellow
Visual Science Distinguished Professor, John Curtin Department of Ophthalmology
Moran Eye Center School of Medical Research University of Washington
University of Utah and Research Director, ARC Centre of Seattle WA USA
Salt Lake City UT USA Excellence in Vision Science
The Australian National University Robert E Marc PhD
Chris A Johnson PhD DSc FARVO Canberra City ACT Australia Professor of Ophthalmology
Professor John A Moran Eye Center
Department of Ophthalmology Dennis M Levi OD PhD University of Utah
University of Iowa Professor of Optometry and Vision Salt Lake City UT USA
Iowa City IA USA Science
Professor, Helen Wills Neuroscience Roan Marion BS
Randy Kardon MD PhD Institute Graduate Student
Professor and Director of Dean, School of Optometry Neuroscience PhD Program
Neuro-ophthalmology University of California, Berkeley Casagrande Vision Research Lab
Pomerantz Family Chair of Berkeley CA USA Vanderbilt University Medical School
Ophthalmology Nashville TN USA
Department of Ophthalmology and Lindsay B Lewis PhD
Visual Sciences Postdoctoral Fellow Joanne A Matsubara BA PhD
University of Iowa and Veterans Department of Ophthalmology Professor and Director of Research
Administration Hospitals McGill Vision Research (Basic Sciences)
Iowa City IA USA Montreal QC Canada Eye Care Centre Department of
Ophthalmology and Visual Sciences
Pradeep K Karla PhD Mark J Lucarelli MD FACS University of British Columbia
Assistant Professor of Pharmaceutical Professor Vancouver BC Canada
Sciences Director of Oculoplastics Service
Department of Pharmaceutical Sciences Department of Ophthalmology and Allison M McKendrick PhD
School of Pharmacy Visual Sciences Senior Lecturer
Howard University University of Wisconsin at Madison Department of Optometry and Vision
Washington DC USA Madison WI USA Sciences
The University of Melbourne
Paul L Kaufman MD Peter D Lukasiewicz PhD Melbourne VIC Australia
Peter A Duehr Professor and Professor of Ophthalmology &
Chairman Neurobiology Linda McLoon PhD
Division of Ophthalmology & Visual Department of Ophthalmology & Professor of Ophthalmology and
Science Visual Sciences Neuroscience
University Wisconsin Madison Washington University School of Departments of Ophthalmology and
Madison WI USA Medicine Neuroscience
St Louis MO USA University of Minnesota
SM Koch MS Minneapolis MN USA
Graduate Student Henrik Lund-Anderson MD DMSc
Neuroscience Graduate Program Professor of Ophthalmology David Miller MD
University of California, San Francisco Department of Ophthalmology Associate Clinical Professor of
San Francisco CA USA University of Copenhagen Ophthalmology
Glostrup Hospital Department of Ophthalmology
Copenhagen Denmark Harvard Medical School
Boston MA USA
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