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ADVANCES I N PHOTORECEPTION
Proceedings of a Symposium on
Frontiers of Visual Science
Committee on Vision
Commission on Behavioral and Social Sciences and Education
National Research Council
National Academy Press
Washington? D.C. 1990
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NOTICE: The project that is the subject of this report was approved by the Governing Board of
the National Research Council, whose members are drawn from the councils of the National Academy
of Sciences, the National Academy of Engineering, and the Institute of Medicine. The menders of
the committee responsible for the report were chosen for their special competences and with regard
for appropriate balance.
This report has been reviewed by a group other than the authors according to procedures
approved by a Report Review Committee consisting of members of the National Academy of
Sciences, the National Academy of Engineering, and the Institute of Medicine.
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COMMITTEE ON VISION
SUZANNE MCKEE (Chair), Smith-Kettlewell Eye Research Foundation,
San Francisco
ROBERT BOYNTON (NAS), Department of Psychology, University of
California, San Diego
LYNN COOPER, Department of Psychology, Columbia University
RUSSELL LEE DEVALOIS (NAS), Department of Psychology,
University of California, Berkeley
MERTON CLYDE FLOM, College of Optometry, University of Houston
DAVID L. GUYTON, Wilmer Ophthalmological Institute, Johns Hopkins
University
DONALD HOOD, Department of Psychology, Columbia University
JAMES LACKNER, Ashton Graybiel Spatial Orientation Laboratory,
Brandeis University
ROBERT SHAPLEY, Department of Psychology, New York University
LOUIS SILVERSTEIN, Honeywell, Inc., Phoenix, Ariz.
KENT ~ STEVENS, Department of Computer and Information Science,
University of Oregon
ANDREW B. WATSON, NASA Ames Research Center, Moffett Field,
Calif.
PAMELA EBERT FLATTAU, Study Director
JOANNE ALBANES, Research Assistant
CAROL METCALF, Administrative Secretary
ROSE WHITE, Secretary
. . .
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SYMPOSIUM PARTICIPANTS
ANTHONY J. ADAMS, School of Optometry? University of California,
Berkeley
JAMES K. BOWMAKER, School of Biological Sciences, Queen Mary
College, University of London
DONALD C. HOOD, Department of Psychology, Columbia University
WALTER MAKOUS, Center for Visual Science, University of Rochester
WILLIAM MILLER, School of Medicine, Yale University
HUGH PERRY, Department of Experimental Psychology, University
of Oxford
EDWARD N. PUGH, Jr., Department of Psychology, University of
Pennsylvania
ROBERT SHAPLEY, Department of Psychology, New York University
JULIE L. SCHNAPF, Department of Ophthalmology, University of
California, San Francisco
DAVID R. WILLIAMS, Center for Visual Science, University of
Rochester
JOHN I. YELLOWS, Cognitive Science Department, University of
California, Irvine
1V
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Preface
The Committee on Vision is a standing committee of the National
Research Council's Commission on Behavioral and Social Sciences and
Education. The committee provides analysis and advice on scientific issues
and applied problems involving vision. It also attempts to stimulate the
development of visual science and to provide a forum in which basic and
applied scientists, engineers, and clinicians can interact. Working groups
of the committee study questions that may involve engineering and equip-
ment, physiological and physical optics, neurophysiology, psychophysics,
perception, environmental effects on vision, and visual disorders.
From time to time, the committee sponsors public meetings that feature
papers on advances in vision research. The meetings are designed to aid
the newcomer in reaching a preliminary understanding of the utility of the
latest approaches to vision research and to challenge more experienced
scientists, engineers, and clinicians alike to consider the appropriate role
for these findings in the advancement of vision research and its application
to practical problems.
In March 1988 the committee sponsored a Symposium on Frontiers of
Visual Science at the National Academy of Sciences in Washington, D.C.
The committee brought together 10 leading investigators in vision research
to discuss advances in photoreception. Speakers included visual physiol-
ogists and psychophysicists, who discussed wavelength specificity of visual
pigments, receptor transduction in health and disease, adaptation of pho-
toreceptors to light, variations of the photoreceptor lattice across the retina,
and photoreceptor sampling of visual images and aliasing. Comparisons of
human and animal retinas also were discussed.
Funds for the symposium were provided from the general budget of the
committee, which receives support from the departments of the Army, the
v
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Navy, and the Air Force; the National Eye Institute; the National Institute
on Aging; the National Aeronautics and Space Administration; the National
Science Foundation, the Veterans Administration; the American Academy
of Ophthalmology; and the American Academy of Optometry.
The committee gratefully acknowledges the efforts of the vision scien-
tists who took time from their demanding schedules to participate in the
symposium. The committee also thanks its staff officer, Pamela Ebert Flat-
tau, for organizing the meeting and preparing the final report. Production
of the report was effectively assisted by Carol Metcalf of the committee
staff. Freelance editor Barbara ~ Bodling helped improve the style and
clarity of the report. To all these, we express our gratitude.
Suzanne McKee, Chair
Committee on Vision
V1
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Contents
Introduction, 1
Robert Shapley
Optical and Molecular Design of Rods, 2
William H. Miller
COLOR AND WAVELENGTH SPECIFICITY
OF RECEPTORS .............................
Cone Visual Pigments in Monkeys and Humans, 19
James K. Bowmaker
Photosensitivity of Primate Photoreceptors, 31
Julie ~ Schnapf
TRANSDUCTION, ADAPTATION, AND DISEASE
PROCESSES ........................................
The ERG and Sites and Mechanisms of Retinal Disease, Adaptation,
and Development, 41
Donald C. Hood
Phototransduction in Vertebrate Rods: The Electrophysiological
Approach to the cGMP Cascade TheoIy, 59
Edward At Pugh, Jr.' OH. Cobbs, and J.D. Tanner
Partitioning Visual Processes, 78
Walter Makous
. .
V11
. . .39
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SPATIAL EFFECTS OF PHOTORECEPTOR MOSAIC
The Distribution of Cones in the Primate Retina, 105
~ Hugh Peny
. . .. . .
The Photoreceptor Mosaic as an Image Sampling Device, 117
John I. Yellott
The Invisible Cone Mosaic, 135
David R Williams
. . .
vail
. . .
....103