Visual Perception A Clinical Orientation, 4th Edition

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Visual Perception
A C L I N I C A L O R I E N TAT I O N
FOURTH EDITION

Steven H. Schwartz, OD, PhD

State College of Optometry
State University of New York
New York, New York

New York Chicago San Francisco Lisbon

London Madrid Mexico City Milan New Delhi
San Juan Seoul Singapore Sydney Toronto
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Contents
Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xv

1. Experimental Approaches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
2. Introductory Concepts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Very Basic Ocular Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Basic Retinal Anatomy and Postretinal Pathways . . . . . . . . . . . . . . . . . . . . .6
Electromagnetic Spectrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
Transmission Characteristics of the Eye . . . . . . . . . . . . . . . . . . . . . . . . . . .23
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
3. The Duplex Retina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Basic Distinctions Between Scotopic and Photopic Vision . . . . . . . . . . . . .26
Morphological Distinctions Between Rods and Cones . . . . . . . . . . . . . . . .26
Photopigments in Rods and Cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30
Retinal Distribution of Photoreceptors . . . . . . . . . . . . . . . . . . . . . . . . . . . .36
Dark Adaptation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .37
Light Adaptation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .44
Spatial Resolution and Spatial Summation . . . . . . . . . . . . . . . . . . . . . . . . .47
Temporal Resolution and Temporal Summation . . . . . . . . . . . . . . . . . . . . .51
Stiles–Crawford Effect of the First Kind . . . . . . . . . . . . . . . . . . . . . . . . . . .54
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .57
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .63
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .64
4. Photometry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .65
Photometry and Radiometry: Basic Concepts and Units . . . . . . . . . . . . . . .65
Cosine Diffusers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .72
Inverse Square Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .73

v
vi Contents

Retinal Illumination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .74

Scotopic Units . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .75
Derivation of the Photopic Luminosity Function . . . . . . . . . . . . . . . . . . . .77
Specification of Light Sources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .79
Filters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .82
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .91
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .92
5. Color Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .93
Trichromatic Theory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .93
Absorption Spectra of the Cone Photopigments . . . . . . . . . . . . . . . . . . . .102
Grassman’s Laws of Metamers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .104
Color Labels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .107
Wavelength Discrimination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .109
Bezold–Brücke Phenomenon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .111
Color Constancy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .112
Physiological Basis of Color Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .113
Munsell Color Appearance System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .119
CIE Color Specification System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .120
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .128
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .129
6. Anomalies of Color Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .131
Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .131
Basic Data of Inherited Color Vision Anomalies . . . . . . . . . . . . . . . . . . . .133
Color Labeling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .141
Inheritance of Anomalous Color Vision . . . . . . . . . . . . . . . . . . . . . . . . . .144
Molecular Genetics of Red–Green Color Vision Anomalies . . . . . . . . . . .145
Acquired Color Vision Anomalies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .148
Achromatopsias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .150
Chromatopsias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .150
Standard Color Vision Tests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .151
Nagel Anomaloscope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .157
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .163
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .167
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .168
7. Spatial Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .169
Sine Wave Gratings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .169
Fourier Analysis: Basic Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . .173
Spatial Modulation Transfer Function of a Lens . . . . . . . . . . . . . . . . . . . .174
The Human Contrast Sensitivity Function . . . . . . . . . . . . . . . . . . . . . . . .177
CSF High-Frequency Cutoff . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .177
CSF Low-Frequency Drop-Off . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .182
The Visual System as a Fourier Analyzer . . . . . . . . . . . . . . . . . . . . . . . . . .183
Relationship of CSF to Snellen Acuity . . . . . . . . . . . . . . . . . . . . . . . . . . .187
 Contents vii

Clinical Implications of the CSF . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .190

Clinical Determination of the CSF . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .193
Other Forms of Visual Acuity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .195
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .199
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .200
8. Temporal Aspects of Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .201
Stimulus Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .201
Temporal Modulation Transfer Functions . . . . . . . . . . . . . . . . . . . . . . . . .204
More on the CFF . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .208
Other Temporal Visual Effects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .209
Masking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .214
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .215
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .217
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .218
9. Motion Perception . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .219
Stimuli Used to Study Motion Perception . . . . . . . . . . . . . . . . . . . . . . . . .219
Dorsal Processing Stream . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .223
Roles of Luminance and Color . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .223
Dynamic Visual Acuity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .224
Saccadic Suppression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .225
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .226
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .227
Self-Assessment Question . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .228
10. Depth Perception . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .229
Monocular Depth Cues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .229
Binocular Depth Cues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .236
Size Illusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .237
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .239
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .241
Self-Assessment Question . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .242
11. Psychophysical Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .243
Threshold . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .243
Determination of Threshold . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .245
Signal Detection Theory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .252
Weber’s Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .260
Magnitude of Sensation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .265
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .267
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .268
12. Functional Retinal Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .269
The Retina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .270
Receptive Fields of Ganglion Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .271
Recording from Retinal Neurons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .276
viii Contents

Photoreceptors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .276
Horizontal Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .279
Bipolar Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .279
Amacrine Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .284
Ganglion Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .285
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .290
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .290
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .291
13. Parallel Processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .293
What the Frog’s Eye Tells Its Brain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .294
X- and Y-Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .295
Parvo and Magno Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .296
Functions of the Parvo and Magno Pathways . . . . . . . . . . . . . . . . . . . . . .300
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .302
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .306
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .307
14. Striate Cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .309
Basic Organization of the Cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .309
Simple and Complex Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .311
Edges or Gratings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .318
Striate Cortical Architecture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .318
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .324
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .324
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .325
15. Information Streams and Extrastriate Processing . . . . . . . . . . . . . . . . . . .327
Information Diverges Beyond Striate Cortex . . . . . . . . . . . . . . . . . . . . . .327
Cortical Modularity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .328
Role of Area MT/V5 in Motion Perception . . . . . . . . . . . . . . . . . . . . . . .330
IT and Form Perception . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .332
Role of V4 in Binocular Rivalry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .333
A Word About Visual Attention . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .333
Visual Agnosias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .338
Phantom Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .340
Are There Grandmother Cells? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .341
Can Vision Be Trained? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .341
A Word of Caution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .342
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .343
Self-Assessment Question . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .344
16. Gross Electrical Potentials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .345
Electrooculogram . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .345
Electroretinogram . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .348
Visually Evoked Potentials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .357
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .362
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .363
 Contents ix

17. Development and Maturation of Vision . . . . . . . . . . . . . . . . . . . . . . . . . .365

Deprivation Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .365
Amblyopia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .370
Meridional Amblyopia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .376
Perceptual Consequences of Bilateral Visual Deprivation . . . . . . . . . . . . .379
Development of Refractive Errors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .380
Development of Grating Acuity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .382
Development of Other Visual Attributes . . . . . . . . . . . . . . . . . . . . . . . . . .387
Additional Clinical Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .391
Vision in the Later Years . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .392
Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .400
Answers to Self-Assessment Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .401
Chapter 3: The Duplex Retina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .401
Chapter 4: Photometry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .402
Chapter 5: Color Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .404
Chapter 6: Anomalies of Color Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . .405
Chapter 7: Spatial Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .406
Chapter 8: Temporal Aspects of Vision . . . . . . . . . . . . . . . . . . . . . . . . . . .409
Chapter 9: Motion Perception . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .409
Chapter 10: Depth Perception . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .409
Chapter 11: Psychophysical Methodology . . . . . . . . . . . . . . . . . . . . . . . . .410
Chapter 12: Functional Retinal Physiology . . . . . . . . . . . . . . . . . . . . . . . .410
Chapter 13: Parallel Processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .410
Chapter 14: Striate Cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .411
Chapter 15: Information Streams and Extrastriate Processing . . . . . . . . .411
Chapter 16: Gross Electrical Potentials . . . . . . . . . . . . . . . . . . . . . . . . . . .412
Chapter 17: Development and Maturation of Vision . . . . . . . . . . . . . . . .412
Practice Examinations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .415
Practice Examination 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .415
Practice Examination 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .425
Practice Examination 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .435
Answers to Practice Examinations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .447
Answers to Practice Examination 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .447
Answers to Practice Examination 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .448
Answers to Practice Examination 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .449
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475
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Foreword
If we believed that we must try to find out what is not known, we should be
better and braver and less idle than if we believed that what we do not know
is impossible to find out and that we need not even try.
—Socrates, The Meno

The core of any profession is its basis in scientific knowledge, and for those in the
eye and vision care professions, we are indeed fortunate that our core is so rich and
deep and broad. Vision science draws its facts and its theories from a wide range of
sources—physics, optics, anatomy, physiology, biochemistry, psychology, and
cognitive neurosciences. Although much of our core knowledge traces back to
Leonardo da Vinci, Thomas Young, Helmholtz, Hering, and Wheatstone, vision
science is a dynamic field, and our current knowledge also owes a great debt to more
recent contributions by such modern workers as Granit, Wald, Hubel and Wiesel,
Enroth-Cugell and Robson, and Nathans. Over the more than 20 years since
Cornsweet’s excellent book on the topic, there has been an explosion in our
knowledge and understanding of visual neurophysiology, genetics, development and
aging, and many other key aspects of visual function.
Although there are many books dealing with aspects of visual science in great
depth, there are few books written on the topic with a view to the clinical impor-
tance of the topic for those in the eye and vision care professions. In this book,
Dr. Steven Schwartz has, I believe, succeeded in covering a broad range of subjects
in visual science and in bringing home the clinical relevance of each section to the
reader.
No single volume can address all of visual science; however, Visual Perception:
A Clinical Orientation covers an extraordinarily broad range of clinically important
topics, including color vision and its defects, spatial vision, temporal aspects of
vision, psychophysics, physiology, and development and aging. The fourth edition
is a real advance. All the figures are now in stunning color, and there is a host of
new images that are clinical in nature (including, for example, color vision tests,

xi
xii Foreword

fundus photographs, etc.). Each chapter has been revised to keep up with advances
in the basic sciences, while increasing the clinical orientation. In certain respects,
the revised Chapter 15, with its discussion of higher level processing and neurol-
ogy, represents the largest thematic change from the original text. And throughout,
the linkage between basic psychophysics and clinical practice has been strengthened.
Dr. Schwartz has a keen understanding of both the fundamental aspects of vision
and their clinical implications, and his unique approach, which brings the science
and its applications together, will make this book a very popular one in teaching
visual science to optometry students.
Dr. Schwartz has a real skill for explaining difficult concepts in very simple (but
technically accurate) terms. Moreover, this book goes well beyond the basics, includ-
ing information from anatomy to perception, and always highlights the clinical
implications. Throughout the book, Dr. Schwartz manages to convey much of the
excitement of important new developments, while providing the reader with suit-
able caution as to which aspects of the unfolding stories are still controversial.
Dr. Schwartz displays a real empathy for his readers. For example, in Chapter 5,
“Color Vision,” after accurately describing the very difficult concepts related to the
CIE system, Dr. Schwartz assures readers by telling them, “If you are having diffi-
culty understanding this system, do not feel alone.” This is a nice touch, and
Dr. Schwartz goes on to provide helpful suggestions for gaining deeper under-
standing. The discussion questions (and the “hints”) are also likely to be helpful to
both teachers of vision science and to the students using the book. And students
(and teachers) will be pleased to note that the number of multiple-choice questions
in the three practice examinations, which are intended to simulate the National
Board of Examiners in Optometry examination, is now at 201.
In summary, this book represents a comprehensive text on visual science, pro-
viding fundamental concepts in an engaging and interesting style. This information
does not exist in any other single volume, and the close links forged between the
basic knowledge and the clinical applications make the book particularly appealing
for optometric and ophthalmologic students, faculty, and researchers.

Dennis M. Levi, OD, PhD

University of California, Berkeley
Preface
A patient with incipient cataracts complains of a marked reduction in vision that
interferes with her daily activities. When we measure visual acuity, however, little
or no reduction is found. How do we explain the apparent lack of consistency
between the patient’s symptoms and her near-normal acuity? On the basis of our
knowledge of visual perception, how do we best treat this patient?
Eye care professionals routinely diagnose and treat conditions that profoundly
affect visual perception. But how well do ophthalmic clinicians understand the phys-
iology of visual perception and its application to clinical practice? My experience
tells me that patient care would benefit from greater utilization of the rich body of
knowledge developed in recent decades.
Although there are several fine textbooks that address vision and its physiologi-
cal basis, none are written for the eye care professional. This text covers those essen-
tials of monocular visual perception on which successful clinical practice is
predicated, with a strong emphasis on physiologically based models. A patient may
present with a sensory/perceptual complaint, but the precipitating condition affects
the structures of the visual system. Clinically useful models of visual perception link
perception with anatomy and physiology.
In addition to providing a basic science background, each chapter discusses the
clinical relevance and application of the material. Clinical information is presented
in such a manner that beginning clinicians and other readers (e.g., graduate students
and experimental psychologists) will benefit from the discussions. As in prior edi-
tions, an effort has been made to present complex and sophisticated concepts in a
manner that is concise, comprehensible, and clinically relevant, while maintaining
an appropriate degree of scientific rigor.
The fourth edition has been expanded to include certain of the myriad advances
in visual perception that have occurred in recent years. Material of clinical interest
is highlighted throughout the book. Figures have been updated and colorized, and
color photographs, many showing clinical conditions, have been added. The book
now includes more than 250 colored diagrams and photographs.

xiii
xiv Preface

Self-assessment questions are presented at the conclusion of many chapters, with

answers given at the end of the book. These questions are intended to develop and
reinforce key concepts. This new edition also includes 201 multiple-choice ques-
tions, divided into three practice examinations. The questions are of varying levels
of difficulty and should prove useful to the reader for determining his or her mas-
tery of the material. Answers to these questions are also provided.
Acknowledgments
Data and models presented in this book result from the hard work of the many vision
and clinical scientists referenced herein, and it is to them that I owe my deepest
gratitude. In the process of writing and revising this book over the past 15 years, I
have been fortunate to receive input from a diverse group of scientists and clinicians.
Below are listed those individuals who generously reviewed or provided feedback
on this or earlier editions, along with their affiliations at that time. Any shortcomings
of the book are, of course, my responsibility.

Benjamin T. Backus, PhD Jeff Hovis, OD, PhD

State College of Optometry School of Optometry
State University of New York University of Waterloo
New York, New York Waterloo, Ontario, Canada
Michael C. Barris, PhD Ralph J. Jensen, PhD
Fredonia, New York Center for Innovative Visual
Rehabilitation
James P. Comerford, PhD, OD
Boston VA Medical Center
New England College of Optometry
Boston, Massachusetts
Boston, Massachusetts
James Kundart, OD, MEd
Mike Fendick, OD, PhD
College of Optometry
College of Optometry
Pacific University
Nova Southeastern University
Forest Grove, Oregon
Fort Lauderdale, Florida
David Lee, OD, PhD
Peter F. Hitchcock, PhD
Illinois College of Optometry
W. K. Kellogg Eye Center
Chicago, Illinois
University of Michigan
Ann Arbor, Michigan

xv
xvi Acknowledgments

Dennis M. Levi, OD, PhD Thomas O. Salmon, OD, PhD

School of Optometry College of Optometry
University of California Northeastern State University
Berkeley, California Tahlequah, Oklahoma
Michael S. Loop, PhD Scott B. Steinman, OD, PhD
School of Optometry/The Medical Southern College of Optometry
Center Memphis, Tennessee
University of Alabama at Birmingham
David Troilo, PhD
Birmingham, Alabama
State College of Optometry
Leon Nehmad, OD, MSW State University of New York
State College of Optometry New York, New York
State University of New York
Dean Yager, PhD
New York, New York
State College of Optometry
Joel Pokorny, PhD State University of New York
The University of Chicago New York, New York
Chicago, Illinois
Robert L. Yolton, OD, PhD
Jeff Rabin, OD, PhD College of Optometry
College of Optometry Pacific University
Pacific University Forest Grove, Oregon
Forest Grove, Oregon
Alan Riezman, OD
VA Pacific Islands Healthcare System
Honolulu, Hawaii
 1
Experimental Approaches
The visual system extracts information from the environment, transforming it into
a neural code that results in perception. Chromatic, motion, detail, form, and depth
information is encoded early in the visual system, further analyzed along cortical
processing streams, and ultimately synthesized with other senses and memory in
higher cortical areas to result in the rich world that we perceive. Diseases of the
visual system can disrupt these processes at any stage, leading to muted or erroneous
perceptions. By understanding the mechanisms that underlie visual perception,1 the
clinician is better able to make sense of patient symptoms that may otherwise be
bewildering.
The data presented in this book were obtained through anatomical, neurophysi-
ological, imaging, and psychophysical experiments. To understand the utility of these
approaches, conceptualize the visual system as a novel and complex electronic
machine with operations that we wish to understand. How do we go about under-
standing such a device?
The machine is opened and its structure determined. Diagrams are made that
illustrate the locations of various components and their interconnections. As with
anatomical studies, structure is determined with the expectation that it will provide
insight regarding operations.
We ascertain the function of the individual components that constitute the device.
Electronic probes are used to record activity from various elements and circuits.
Similar to electrophysiological methodology, local circuits are studied to determine
how information is processed.
Looking inside the intact machine with imaging devices allows us to see activity
in its various components while it is engaged in specific functions. Which areas are
busy, for example, when it performs a given task, and which are busy when it per-
forms a very different task? Akin to using noninvasive procedures to view the human

1. Just as there are numerous approaches to the study of visual perception, there are myriad defini-
tions. For this book, we define visual perception as the conscious expression of neural activity within
the neural structures of the visual system.