Menu Expand
Adler's Physiology of the Eye E-Book

Adler's Physiology of the Eye E-Book

Leonard A Levin | Siv F. E. Nilsson | James Ver Hoeve | Samuel Wu | Paul L. Kaufman | Albert Alm

(2011)

Additional Information

Book Details

Abstract

Drs. Paul L. Kaufman, Albert Alm, Leonard A Levin, Siv F. E. Nilsson, James Ver Hoeve, and Samuel Wu present the 11th Edition of the classic text Adler’s Physiology of the Eye, updated to enhance your understanding of ocular function. This full-color, user-friendly edition captures the latest molecular, genetic, and biochemical discoveries and offers you unparalleled knowledge and insight into the physiology of the eye and its structures. A new organization by function, rather than anatomy, helps you make a stronger connection between physiological principles and clinical practice; and more than 1,000 great new full-color illustrations help clarify complex concepts.

  • Deepen your grasp of the physiological principles that underlie visual acuity, color vision, ocular circulation, the extraocular muscle, and much more.
  • Glean the latest knowledge in the field, including the most recent molecular, genetic, and biochemical discoveries.
  • Make a stronger connection between physiology and clinical practice with the aid of an enhanced clinical emphasis throughout, as well as a new organization by function rather than by anatomy.
  • Better visualize all concepts by viewing 1,000 clear, full-color illustrations.

Table of Contents

Section Title Page Action Price
Front cover cover
Adler's Physiology of the Eye, 11/e i
Copyright page iv
Table of Contents v
Preface vii
List of Contributors viii
Acknowledgements xi
Dedication xii
1 Focusing of an image on the retina 1
1 Optics 1
The young eye 1
Relevant anatomy 1
Axial length 1
Emmetropization 2
Retinal receptors 3
Neural processing 3
Relevant early physiology 4
Recognizing faces 4
Line orientation receptors 5
Monitoring other’s eye movements 5
Recognizing movement 5
Summary – social seeing 6
The image of the human adult eye 6
Tuned to visible light waves 6
Role of the cornea 6
Role of the crystalline lens 7
Accommodation 8
Role of the retina 8
Rhodopsin 8
Receptor size and spacing 9
Visual acuity testing 9
Chart luminance 9
Visual acuity as Log MAR 10
Visual acuity chart contrast 10
Contrast sensitivity testing 10
Definition and units 12
Contrast 12
Contrast sensitivity 12
Targets 12
Sine waves 12
Recording contrast sensitivity 12
Glare, tissue light scattering, and contrast sensitivity 13
Clinical conditions affecting glare and contrast sensitivity 15
Optical conditions 15
Corneal conditions 15
Corneal edema 15
Contact lens wear 15
Keratoconus 15
Nephrotic cystinosis 15
Penetrating keratoplasty 15
Refractive surgery 15
Cataracts and opacified posterior capsules 15
Modulation transfer function 16
Depth of focus 16
Optical aberrations 17
Light scattering 17
Natural defenses against light scattering 19
Chromatic aberrations 20
Spherical aberration 20
Light absorption 21
Summary – a compromise of eye function 21
The aging eye 21
Evolution of ocular components 22
Non-optical brain mechanisms that enhance the retinal image 22
Contrast enhancement 22
Edge sharpening 22
Vernier acuity 23
Removing distractions 24
Refractive errors 24
Prevalence 24
Myopia 25
Pathologic myopia 25
Physiologic, or school, myopia 25
Astigmatism 25
Presbyopia 25
Components of ametropia 25
References 26
2 Optical Aberrations and Wavefront Sensing 28
Introduction 28
Optical aberrations 28
Wavefront optics 28
Optical limitations to vision 30
Aberrations 31
Chromatic aberrations 31
Monochromatic aberrations 31
Measuring optical aberrations 32
Aberrometry and wavefront sensing devices 32
Wavefront sensing devices 34
Correcting higher-order aberrations 34
Visual disturbances associated with HOA 34
Visual performance after correcting HOA 35
Factors which limit the benefit of HOA correction 35
Clinical applications of wavefront aberration correction 37
Corneal ablations 37
Correcting HOA with spectacles, contact lenses and intraocular lenses 37
References 38
3 Accommodation 40
Introduction 40
Accommodation 41
Optics of the eye 42
The optical requirements for accommodation 42
Depth of field 43
Visual acuity 44
The anatomy of the accommodative apparatus 44
The ciliary body 44
The ciliary muscle 44
The zonular fibers 45
The lens capsule 47
The crystalline lens 47
The mechanism of accommodation 48
Accommodative optical changes in the lens and eye 49
The stimulus to accommodate 55
The pharmacology of accommodation 56
Measurement of accommodation 57
Presbyopia 59
Factors contributing to presbyopia 59
Age-related changes in rhesus ciliary muscle 59
Age-related changes in human ciliary muscle 61
Age-related changes in the zonule 61
Age-related changes in the capsule 62
Growth of the crystalline lens 63
Loss of ability of the human lens to accommodate 65
Age-related increase in stiffness of the human lens 66
References 68
2 Physiology of optical media 71
4 Cornea and Sclera 71
Introduction 71
Cornea 71
Embryology, growth, development, and aging 71
Major corneal reference points and measurements 73
Optical properties 77
Light refraction 77
Light transmission 78
Collagen 80
Keratocytes 83
Proteoglycans 85
Corneal nerves 88
Corneal stromal wound healing 89
Barrier properties 92
Low-permeability barrier: the corneal epithelium 92
High-permeability barrier: the corneal endothelium 96
Leaky barrier function 98
Metabolic pump function 99
Corneal edema 101
Basement membrane and glycocalyx 105
Mechanical properties 106
Corneal stress 106
Corneal stiffness, strength extensibility, and toughness 107
Chronic biomechanical failure of the cornea – ectasia 109
Other functions 112
Drug delivery 112
Ultraviolet light filtration 114
Sclera 115
Embryology, growth, development, and aging 115
Major scleral reference points and measurements 119
Mechanical properties 120
Scleral dehydration and edema 121
Episcleral vasculature 121
Wound healing 122
Drug delivery 122
Acknowledgments 125
References 125
5 The Lens 131
The anatomy of the adult lens 131
The basics of lens refraction and transparency 131
The early development of the lens 133
Lens fiber cell differentiation 135
Lens crystallins 136
The lens fiber cell cytoskeleton 137
Other cellular and biochemical specializations found in lens fiber cells 138
The control of lens growth 139
Communication between lens epithelial and fiber cells 140
Vascular support during lens development 140
The lens as the organizer of the anterior segment 140
Special problems of lens cell metabolism 140
Overview 140
Oxidants within and around the lens 141
Protection against oxidative damage 141
Energy production in the lens 142
Water and electrolyte balance 142
Lens transparency and refraction 143
Changes in the lens with aging 144
The structure and development of the lens sutures 144
The lens capsule 145
The zonules 145
Cataracts 146
Cataract epidemiology 146
General risk factors 146
Age-related nuclear cataracts 148
Age-related cortical cataracts 150
Posterior subcapsular cataracts 151
Mixed cataracts 151
Secondary cataracts 152
Less common types of cataract 152
Overview of age-related cataract formation 155
Perspectives for preventing cataract blindness 155
Acknowledgments 157
References 157
6 The Vitreous 164
Introduction 164
Anatomy 164
Embryology 164
Structural considerations of embryology 164
Molecular and cellular considerations of embryology 164
Anatomy of the mature vitreous body 164
The vitreoretinal interface 165
Ultrastructural, biochemical, and biophysical aspects 166
Ultrastructural and biochemical aspects 166
Biophysical aspects 168
Aging of the vitreous 171
Molecular mechanisms in aging 171
Structural changes 171
Vitreo–retinal interface imaging 172
Diffusion kinetics as an indicator of the biophysical status of the vitreous 172
Physiology of the vitreous body 174
A. Support function for the retina and filling up function of the vitreous body cavity 174
Normal conditions 174
Pathological/pathophysiological correlations 174
Posterior vitreous detachment 174
Development of macular edema 175
B. Diffusion barrier between the anterior and the posterior segments of the eye 176
3 Direction of gaze 182
7 The Extraocular Muscles 182
The bony orbit 182
Normal extraocular muscles 182
Gross anatomy 182
Cranial motor nerve innervation 185
Orbital connective tissue 186
Histological anatomy and physiologic implications 187
Metabolism 191
Proprioception and proprioceptors 195
Development 196
Disease propensity 197
Disorders of eye movements 197
Strabismus 197
Nystagmus 198
Congenital cranial dysinnervation disorders (CCDD) 199
Diseases where EOM are preferentially spared 199
Diseases where EOM are preferentially involved 202
Conclusion 204
References 204
8 Three-Dimensional Rotations of the Eye 208
Eye motility 208
Quantifying eye rotations 208
Nested-axes coordinates 210
Head-fixed coordinates 210
Listing’s law 211
False torsion 213
Neural control of ocular orientation 214
Orbital mechanics can simplify neural control: extraocular pulleys 216
Summary 219
References 219
9 Neural Control of Eye Movements 220
Introduction 220
Three fundamental visual sensory-motor tasks 220
Three components of eye rotation 220
Binocular constraints on eye position control 220
Feedback and feedforward control systems 221
Hierarchy of oculomotor control 222
Final common pathway 222
Cranial nerves: III, IV, & VI and motor nuclei 222
Motor neuron response 223
Functional classification into three general categories 224
I. Stabilization of gaze relative to the external world 224
A. Extra-retinal signals 224
B. Retinal signals 225
C. Neuro-control of stabilization reflexes 226
1. Vestibulo-ocular reflex 226
2. Optokinetic nystagmus 227
II. Foveal gaze lock (maintenance of foveal alignment with stationary and slowly moving targets) 227
A. Static control of eye alignment (fixation) 227
B. Dynamic control of eye alignment (smooth tracking responses to open- and closed-loop stimuli) 229
1. Conjugate smooth pursuit tracking 229
2. Disconjugate smooth vergence tracking 230
3. Adaptable interactions between smooth pursuit and smooth vergence 230
C. Neuro-control of smooth foveal tracking 230
1. Smooth pursuit tracking system 230
2. Smooth vergence tracking system 230
III. Foveal gaze shifts: target selection and foveal acquisition 231
A. Rapid conjugate shifts of gaze direction (saccadic eye movements) 231
B. Disconjugate shifts of gaze distance (the near response in symmetrical convergence) 232
C. Interactions between conjugate and disconjugate eye movements (asymmetric vergence) 233
D. Neuro-control of foveal gaze shifts 235
1. Saccadic gaze shifting system 235
2. Vergence gaze shifting system: the near triad and interactions with saccades 235
Neurological disorders of the oculomotor system 236
I. Strabismus 237
II. Gaze restrictions 237
III. Saccade disorders 239
IV. Nystagmus 240
Acknowledgments 240
References 240
4 Nutrition of the eye 243
10 Ocular Circulation 243
Introduction 243
Anatomy of the ocular circulation 243
Vascular supply of the retina 243
Vascular supply of the choroid 244
Paraoptic pattern 244
Perimacular pattern 244
Fine structure and innervation of retinal and choroidal vessels 246
Vascular supply of the anterior segment 246
Transport through blood–retinal barriers 247
Transcellular pathway (transcytosis) 247
Paracellular pathway 247
Extracellular structures 248
Glycocalyx 248
Extracellular matrix 248
Inner blood–retinal barrier 249
Pericytes 249
Glial cells 249
Outer blood–retinal barrier 249
Fenestrated endothelium 249
Retinal pigment epithelium 249
Bruch’s membrane 249
Blood–aqueous barriers 249
Techniques for measuring ocular blood flow 250
Techniques used in experimental animals 250
Non-invasive techniques used in physiological and clinical research 251
Ocular circulatory physiology 254
General hemodynamic considerations 254
Ocular hemodynamic data under basal physiological conditions 255
Retina 255
Choroid 255
Ciliary circulation 255
Vasomotion 256
Effects of age on ocular blood flow 256
Regulation of ocular BF 256
Autoregulation of ocular blood flow 256
Retina and ONH 256
Choroid 257
Anterior uvea 257
Mechanisms underlying retinal and ONH autoregulation 257
Regulation of blood flow in response to increase in arterial blood pressure (ABP) 257
Static exercises 257
Dynamic exercises 258
Change in posture 258
Regulation of blood flow in response to changes in blood gases 258
Hyperoxia 258
Hypoxia 258
Hypercapnia 259
Metabolic control of retinal blood flow 260
Retinal metabolism and vasoreactivity 260
Blood flow response to visual stimulation 260
Light/dark transition 260
Flicker 261
Control of arterial tone by endothelium or neuro-glial activity 261
Nitric oxide 262
Endothelins 262
Prostaglandins (PGs) 263
Neural, endocrine, and paracrine control 263
Effects of vasoactive nerves 263
Adenosine 264
Endogenous and pharmacological substances 264
Role of administration route 264
Vasoconstrictors 264
Vasodilators 265
Ciliary blood flow regulation 265
Ocular blood flow and its regulation in diseases 265
Diabetes 265
Glaucoma 266
Age-related macular degeneration 266
References 266
11 Production and Flow of Aqueous Humor 274
Aqueous humor formation 274
Physiology of aqueous humor formation 274
Biochemistry of aqueous humor formation 275
Aqueous humor composition 278
Blood–aqueous barrier 278
Active transport 279
Pharmacology and regulation of aqueous humor formation and composition (Box 11.1) 280
Cholinergic mechanisms 280
Adrenergic mechanisms 281
Other agents 281
Aqueous humor drainage 283
Fluid mechanics 283
Structural components 284
Pumping model for trabecular outflow 285
Active involvement of the TM in regulating outflow 286
Outflow obstruction 287
Extracellular matrix accumulation and POAG 287
Cell and other particulates 289
Protein and other macromolecules 289
Pharmacology and regulation of outflow 289
Cholinergic mechanisms 289
Conventional (trabecular) outflow 289
Alterations in cholinergic sensitivity of the outflow apparatus 290
Unconventional (uveoscleral) outflow 291
Adrenergic mechanisms 291
Conventional (trabecular) outflow 291
Unconventional (uveoscleral) outflow 292
Cytoskeletal and cell junctional mechanisms (Box 11.2) 292
Corticosteroid mechanisms 294
Prostaglandin mechanisms (Box 11.3) 296
Cell volume related mechanisms 297
Hyaluronidase and protease-induced facility increases 298
Other agents 298
Physical enhancement of outflow 299
References 299
12 Metabolic Interactions between Neurons and Glial Cells 308
Introduction 308
1. Retinal oxygen distribution and consumption 309
Inner retina 309
Dark and light O2 consumption 309
Outer retina 309
Photoreceptor QO2 in darkness 309
Photoreceptor QO2 in light 309
2. Role of glycolysis underlying retinal function: from whole retina to its parts 310
3. Biochemical specialization of glial cells 311
4. Role of glycogen 313
5. Functional neuronal activity and division of metabolic labor 313
6. Cellular compartmentation of energy substrates other than glucose 314
7. Experimental models used to study the interaction between photoreceptors and glial Müller cells 315
In vitro studies of the retina of the honeybee drone 315
If there are no conventional synapses in drone retina and only the photoreceptors are directly excitable by light, what is the evidence that photoreceptors depend on surrounding glia for their metabolic needs? 315
When bee retinal slices are exposed to the glycolytic poison IAA, the light-induced change in QO2 is gradually abolished. Is this modulation of QO2 a direct effect of IAA in photoreceptors? 316
What is the evidence that photostimulation of drone retinal slices increases the carbohydrate metabolism in the glia to sustain photoreceptor respiration? 316
Glucose is not the principal energy substrate used by photoreceptors, so what is the identity of the energy metabolite maintaining photoreceptor function and respiration? 316
What is the biochemical evidence for the metabolic effects of NH4+and glutamate in glia? 317
Overall scheme for metabolic compartmentation and metabolic trafficking in honeybee drone retina 317
Experimental models in vertebrates 318
8. Metabolic interactions between vertebrate photoreceptors and Müller glial cells 319
9. Metabolic interaction between photoreceptors and retinal pigment epithelia 319
10. Metabolic factors in the regulation of retinal blood flow 320
11. Metabolic pathway leading to nitric oxide release 321
References 322
13 The Function of the Retinal Pigment Epithelium 325
Introduction 325
Absorption of light 325
Transepithelial transport 325
Transport from the blood side to the photoreceptor side 325
Transport from the retinal side to the blood side 326
Capacitative compensation of fast changes in the ion composition in the subretinal space 327
Visual cycle 328
Phagocytosis of photoreceptor outer segments 329
Secretion 330
Structural integrity of neighboring tissues 330
Immune privilege of the eye 331
References 331
5 Protection of the eye 333
14 Functions of the Orbit and Eyelids 333
Introduction 333
Orbital anatomy and function 333
Orbit osteology 333
The orbital apex 335
Orbital soft tissues 337
Periorbital fascia 337
Orbital fat 338
Orbital nerves 338
Vascular anatomy 339
Arterial supply 339
Venous drainage 340
Orbital lymphatic drainage 340
Facial and eyelid anatomy and function 340
The eyebrow and forehead 340
The midface 341
The eyelid 343
The eyelid margin 343
Eyelid musculature 344
Blinking 345
Eyelid fat 345
Eyelid vasculature 345
Eyelid lymphatics 346
Eyelid innervation 346
References 347
15 Formation and Function of the Tear Film 350
1. Tear film overview 350
2. Glycocalyx 350
A. Structure 350
B. Function 352
3. Mucous layer 353
A. Structure 353
B. Conjunctival goblet cells 353
C. Regulation of goblet cell mucin production 353
Overview 353
Regulation of goblet cell secretion 353
Regulation of goblet cell proliferation 354
D. Regulation of conjunctival electrolyte and water secretion 354
E. Mucous layer function 356
4. Aqueous layer 356
A. Overview 356
B. Lacrimal gland structure 356
C. Lacrimal gland innervation 356
D. Protein secretion regulation 356
Types of protein secretion 356
Cholinergic agonists 357
VIP 357
α1-Adrenergic agonists 357
EGF 358
Interaction of pathways 358
E. Regulation of electrolyte and water secretion 358
Mechanism of acinar electrolyte and water secretion 358
Mechanism of ductal electrolyte and water secretion 359
Neural activation of electrolyte and water secretion 360
F. Lacrimal gland fluid composition 360
G. Aqueous layer function 360
5. Lipid layer 360
A. Structure of meibomian glands and mechanism of lipid production 360
B. Regulation of meibum secretion 361
Neural regulation 361
Hormonal regulation 361
C. Function 361
References 361
16 Sensory Innervation of the Eye 363
Introduction 363
1. Anatomy of ocular sensory nerves 363
1.1 Origin of the ocular sensory nerves 363
Trigeminal ganglion neurons 363
The ophthalmic nerve and its branches 363
1.2 Distribution of sensory nerve fibers within the eye 364
1.3 Architecture of corneal sensory nerves 365
Corneal stromal nerves 365
Subepithelial nerve plexus 365
Sub-basal nerve plexus 365
Intraepithelial nerve terminals 365
1.4 Central sensory pathways 367
2. Development and remodeling of corneal innervation 367
2.1 Development of corneal nerves 367
2.2 Dynamic remodeling of adult corneal innervation 369
2.3 Regeneration of injured corneal nerves 369
3. Functional characteristics of ocular sensory innervation 369
3.1 Trigeminal ganglion neurons 369
3.1.1 Sensory fibers of the cornea and conjunctiva 369
Polymodal nociceptors 371
Mechano-nociceptors 371
Cold thermal receptors 371
“Silent” nociceptors 372
3.1.2 Sensory fibers of the sclera, iris, and ciliary body 374
3.1.3 Ocular trigeminal ganglion neurons 374
3.2 Central pathways 374
4. Inflammation and injury effects on ocular sensory neurons 374
4.1 Local inflammation 374
4.2 Nerve injury 375
5. Trophic effects of ocular sensory nerves 375
6. Sensations arising from the eye 377
6.1 Techniques for testing ocular surface sensitivity 377
6.2. Psychophysics of corneal and conjunctival sensations 377
6.3 Sensitivity of the injured cornea 377
7. Ocular pain 379
7.1 Superficial ocular pain 381
7.2 Deep ocular pain 381
7.3 Pain referred to the eye 382
8. Drugs acting on ocular sensory nerves 382
Topical anesthetics 382
Anti-inflammatory drugs 382
Cycloplegic agents 382
Analgesics 382
Prevention of surgical pain 382
References 383
17 Outward-Directed Transport 385
Introduction 385
Efflux transporters – brief history 385
Efflux transporters in ocular tissues 386
P-gp 386
Mode of action and structure of P-gp 387
MRP 387
Mode of action and structure of MRP 389
BCRP 389
Mode of action and structure of BCRP 390
LRP 390
Localization of transporters 390
Discussion 390
Clinical correlates from literature 390
Strategies to evade efflux transporters 391
Acknowledgment 392
References 392
6 Photoreception 394
18 Biochemical Cascade of Phototransduction 394
Overview 394
Location and compartmentalization of rods and cones 394
Dark-adapted rods 394
The resting dark-adapted state 396
The membrane potential 396
The dark current and the cGMP-gated channel 396
Ca2+ and the exchanger 397
Control of [cGMP] by guanylate cyclase and PDE6 398
Rhodopsin 398
G-protein, Gt 399
Importance of lipid milieu 400
The activation phase of a light response 401
Photoisomerization of rhodopsin 401
G-protein activation 402
PDE6 activation 402
Channel closing 402
Slowing of neurotransmitter release 402
The recovery phase 402
Rhodopsin phosphorylation, retinoid recycling and regeneration 402
Arrestin binding 404
cGMP restoration by guanylate cyclase activation 404
G-protein and PDE6 inactivation by RGS9-1 404
Amplification 404
Responses to saturating light levels 405
Adaptation to changing levels of ambient lighting (see Chapter 20) 405
Turnover of guanine nucleotides 405
Comparison of cones and rods 406
Similarities and differences of phototransduction molecules 406
Physiological differences 406
Phototransduction and disease 406
Retinal degeneration and night blindness 407
What we don’t know 407
Where the field is headed 408
References 408
19 Photoresponses of Rods and Cones 411
Photovoltage response to flashes 411
Photocurrent response to flashes 411
Detecting single photons 413
Photocurrent response to steady light 415
Action spectra of rods and cones 417
CNG channel and Na+/K+,Ca2+ exchanger 420
Role of inner segment conductances 422
Delayed rectifier potassium current, IKV 423
Hyperpolarization-activated current, IH 423
Voltage-activated calcium current, ICa 423
Calcium-activated potassium current, IK(Ca) 424
Calcium-activated anion current, ICl(Ca) 424
Electrotonic coupling 425
Summary 426
Acknowledgment 426
References 426
20 Light Adaptation in Photoreceptors 429
1. Vision from starlight to sunlight 429
Light adaptation versus dark adaptation 429
Purposes of light adaptation 429
2. Performance of the photopic and scotopic divisions of the visual system 430
Photopic vision: the cone system is the workhorse of vision 430
The responses of cones are rapid and moderately sensitive 430
Comparison of photopic and scotopic light adaptation 430
Scotopic vision: the rod system provides specialization for night vision 430
3. Light adaptation of the electrical responses of cones and rods 432
Saturation of the electrical response in rods and its avoidance in cones 432
Desensitization and acceleration of the photoreceptor’s electrical response 433
Unaltered rising phase but accelerated recovery 433
Dependence of sensitivity on background intensity: Weber’s Law 434
Extremely rapid recovery of human cone photocurrent 435
4. Molecular basis of photoreceptor light adaptation 436
The phototransduction cascade 436
Photoreceptor light adaptation independent of calcium 436
Accelerated turnover of cGMP 437
Calcium-dependent mechanisms of rapid light adaptation: re-sensitization through prevention of saturation 437
Powerful negative feedback loop mediated by calcium 437
Guanylyl cyclase activation 438
Shortened R* lifetime 438
Channel reactivation 439
Molecular basis of the cone’s incredibly rapid recovery from light exposure 439
Cone avoidance of saturation 439
Modeling of human cone light adaptation 440
5. Slow changes in rods: light adaptation or dark adaptation? 440
Light-induced change in the dominant time constant 440
Light-induced translocation of proteins 440
6. Dark adaptation of the rods: very slow recovery from bleaching 440
References 442
7 Visual processing in the retina 443
21 The Synaptic Organization of the Retina 443
Kinds of neurons 443
The multipolar neuron phenotype 444
The gliaform cell phenotype 444
True glia and vasculature 445
Basic synaptic communication 446
1. Photoreceptor ribbon synapses: small-volume multi-target signaling 446
2. BC ribbon synapses: semi-precise target signaling 447
3. AC and AxC conventional fast synapses: precise presynaptic → postsynaptic signaling 448
4. AC, AxC, and efferent slow transmitter synapses: large volume signaling 448
5. HC non-canonical signaling 449
6. Coupling types and coupling patterns 449
Fast, focal neurochemistry, synaptic currents, and amplification 449
Global neurochemistry and modulation 450
Modulation by transporters 450
Signal processing 451
Sign-conserving (>) and sign-inverting (>i, >m) transfers 451
Synaptic chains and polarity 451
Feedback, feedforward, and nested feedback/feedforward 452
Caveats 452
Networks 452
The synaptology of center-surround organization 452
The synaptology of mammalian rod pathways – evolution of a new amplification scheme 452
The synaptology of motion – AC surrounds from afar 454
The synaptology of color – HC surrounds again? 454
R/G opponency 454
B/Y opponency 455
Revising the retinal synaptic networks with disease 455
References 457
22 Signal Processing in the Outer Retina 459
Electrical synapses (coupling) between photoreceptors 459
Glutamatergic synapses between photoreceptors and second-order retinal neurons 460
Horizontal cell responses 460
Horizontal cell output synapses 462
Rod and cone pathways and bipolar cell output synapses 464
Bipolar cell responses and center-surround antagonistic receptive field (CSARF) organization 465
Acknowledgments 469
References 469
23 Signal Processing in the Inner Retina 471
Bipolar cells form parallel pathways and provide excitatory input to the IPL 471
Synaptic mechanisms shape excitatory signals in the IPL 471
Amacrine cells mediate inhibition in the IPL 473
GABAergic feedback inhibition changes the timecourse of bipolar cell signaling 475
GABAergic inputs to the bipolar cell axon terminals contribute to surround signaling in the retina 475
The contributions of the inner and outer retina to ganglion cell receptive field surround organization 475
Glycinergic inhibition plays several different roles in the IPL 476
Neuromodulators in the IPL 476
Parallel ganglion cell output pathways 476
Ganglion cells encode color information 476
Directional selective ganglion cells 477
Intrinsically photosensitive ganglion cells 477
Ganglion cell types form a retinal mosaic 478
Conclusions 478
References 478
24 Electroretinogram of Human, Monkey and Mouse 480
Introduction 480
Generation of the ERG 480
Radial current flow 480
Glial currents 482
Stimulus conditions 483
Non-invasive recording of the ERG 483
Classical definition of components of the ERG 484
Slow PIII, the c-wave and other slow components of the direct current (dc)-ERG 484
Full-field dark-adapted (Ganzfeld) flash ERG 484
Dark-adapted a-wave 485
Negative ERGs 485
Modeling 486
Mixed rod-cone a-wave 486
Timecourse of the rod photoreceptor response 487
Dark-adapted b-wave (PII) 487
Scotopic threshold response (STR) 488
Light-adapted, cone-driven ERGs 489
Isolating cone-driven responses 489
Light-adapted a-wave 491
Light-adapted b-wave 492
Light-adapted d-wave 493
Flicker ERG 494
Oscillatory potentials 495
Photopic negative response 495
Pattern ERG 496
Multifocal ERG 497
Closing comments 498
References 499
8 Non-perceptive vision 502
25 Regulation of Light through the Pupil 502
The neuronal pathway of the pupil light reflex and near pupil response 503
Afferent arm of the pupil light reflex 504
The interneuron arm of the pupil light reflex 506
The efferent arm of the pupil light reflex 507
The pupil near reflex and accommodation 508
Pupil reflex dilation: central and peripheral nervous system integration 508
Other neuronal input to the iris 509
Structure of the iris 509
Iris sphincter, iris dilator, and iris color 509
Properties of light and their effect on pupil movement 510
Relative afferent pupillary defects 511
Clinical observation of the pupil light reflex 511
Computerized pupillometry 514
Pupil perimetry 514
Efferent pupillary defects 514
Anisocoria 514
Pupil inequality that increases in the dark 515
Pharmacologic diagnosis of Horner syndrome with cocaine or apraclonidine 518
Pharmacologic localization of the denervation in Horner syndrome 519
Congenital and childhood Horner syndrome 519
Pupil inequality that is increased in bright light 520
Examination of the iris with high magnification using the slit-lamp biomicroscope 520
Pharmacologic response of the iris sphincter to cholinergic drugs 520
Cholinergic supersensitivity 520
Subsensitivity of the iris sphincter to cholinergic testing 521
Adie’s tonic pupil: postganglionic parasympathetic denervation 521
Pupil involvement in third nerve palsy 522
Aberrant regeneration in the third nerve 523
Light-near dissociation: evaluation of the near response 523
When the pupil fails to dilate 523
References 524
26 Ganglion-Cell Photoreceptors and Non-Image-Forming Vision 526
Overview 526
Historical roots 526
Discovery of melanopsin and ganglion-cell photoreceptors 527
Distinctive functional properties of ipRGCs 528
Melanopsin chromophore and pigment bistability 528
Spectral tuning 528
Invertebrate-like phototransduction cascade 529
Depolarizing photoresponse with action potentials 529
Sensitivity 531
Kinetics 531
Morphology, retinal distribution and receptive field 531
Resistance to pathological states 532
Synaptic input 533
Bipolar cell input 533
Amacrine cell input 535
Color coding of synaptic inputs 536
Synaptic output and physiological functions 536
Intraretinal output 536
Central projections 537
The pupillary light reflex 538
Circadian photoentrainment and photic modulation of the pineal 538
Acute regulation of activity and sleep 540
Lateral geniculate nucleus and conscious light perception 541
Development 541
References 541
9 Visual processing in the brain 545
27 Overview of the Central Visual Pathways 545
Targets of the retinal projections 545
Visual field lesions 547
References 549
28 Optic Nerve 550
Introduction 550
Optic nerve anatomy 550
Retinal ganglion cell axons within the nerve fiber layer 550
Intrascleral optic nerve 550
Intraorbital optic nerve 550
The optic canal 551
Intracranial optic nerve and the optic chiasm 552
The optic tract and lateral geniculate nucleus 552
Optic nerve axon counts and dimensions 553
Microscopic anatomy and cytology 553
Axons 553
Oligodendrocytes and myelin 553
Astrocytes 553
Microglia 554
Meninges and meningothelial cells 555
Blood supply 555
Optic nerve head 555
Intraorbital optic nerve and optic canal 556
Intracranial optic nerve, chiasm, and optic tract 556
Vascular biology 556
Optic nerve development 556
Generation of optic nerve oligodendrocytes and myelination 556
Generation of optic nerve astrocytes 557
Development of optic nerve meninges 557
Axon number 557
Axon growth 557
Axon guidance 557
Optic nerve physiology 558
Retinal ganglion cell electrophysiology and synaptic transmission 558
Axonal conduction 559
Action potentials 559
Role of oligodendrocytes and myelin 559
Role of astrocytes 559
Axonal transport 560
Optic nerve injury 560
Clinical implications 560
Types of optic nerve injury 560
Traumatic optic neuropathy 560
Ischemic optic neuropathy 560
Optic neuritis and inflammation 561
Compressive optic neuropathy 561
Glaucoma 561
Papilledema 562
Retinal ganglion cell death after optic nerve injury 562
Time course 563
Apoptosis 563
Signaling of axonal injury 563
Phagocytosis and immune activation 564
Gliosis 564
Failure of axon regeneration 564
Glial inhibition of neurite extension 564
Neuron-intrinsic limitations to axon regeneration 565
Optic nerve repair 565
Optic nerve remyelination 565
Neuroprotection and retinal ganglion cell survival 565
Regeneration of RGC axons 566
“Neuroenhancement” of retinal ganglion cell function 566
Conclusions 567
Acknowledgments 567
References 567
29 Processing in the Lateral Geniculate Nucleus (LGN) 574
The lateral geniculate nucleus: the gateway to conscious visual perception 574
Overview of lateral geniculate anatomy 574
Layers and maps 574
Cell classes 575
Inputs: the retina 576
Inputs: extraretinal sources and cortical feedback 576
Outputs: projections to V1 and beyond 577
LGN circuits: How are visual signals regulated? 578
Feedback and feedforward pathways 578
Circuit neurochemistry 579
Signal processing in the LGN 580
Receptive field properties and parallel processing 580
Physiology of M, P, and K cells 581
The influence of the “extra-classical” surround 581
The impact of feedback 581
The LGN and arousal, attention and conscious vision 583
The LGN and motor planning 583
The LGN and binocular rivalry and visual awareness 584
Conclusions 584
References 585
30 Processing in the Primary Visual Cortex 586
Overview: The primary visual cortex constructs local image features 586
Overview of cortical organization: a general road map 586
Layers, connections, and cells of V1: The inputs, outputs, and general wiring 588
LGN inputs 588
Other inputs to V1 589
Cell classes and connections within V1 589
Output pathways from V1 590
Receptive field properties: How is V1 different from the LGN? 591
Columns and modules: Outlining the functional architecture of V1 593
How do parallel inputs relate to parallel outputs? 594
Does V1 Do More? 595
The importance of time 595
The importance of context 596
Conclusion 597
References 597
31 Extrastriate Visual Cortex 599
What is extrastriate visual cortex? 599
Methods used to identify extrastriate areas in monkeys and humans 599
Histology 599
Retinotopic mapping 599
Connection patterns 601
Functional specificity 601
Comparing visual areas in monkeys and humans 602
Processing streams in extrastriate cortex 603
V2 603
Areas of the dorsal stream 605
MT/V5 and related areas 605
V3 606
V3A 607
PO/V6 607
Parietal lobe areas 607
Areas of the ventral stream 607
V4 607
Inferotemporal cortex 608
PIT/TEO 608
TE 609
References 609
10 Visual perception 613
32 Early Processing of Spatial Form 613
Introduction 613
Foveal window of visibility 613
What limits our contrast sensitivity? 614
What is the relationship between the contrast sensitivity function and the response of single cortical cells? 614
The contribution of M & P pathways to contrast sensitivity 615
Do these two parallel systems carry the same or different contrast sensitivity information? 615
The contribution of different cortical areas to contrast sensitivity 616
The effect of disease on contrast sensitivity 616
Peripheral window of visibility 618
Is the periphery specialized for detecting anything? 619
Why does contrast sensitivity for high spatial frequencies decline with eccentricity? 619
Luminance 619
Chromatic sensitivity 621
Suprathreshold sensitivity 623
Conclusion 624
Acknowledgments 625
References 625
33 Visual Acuity 627
Defining and specifying visual acuity 627
Minimum visible acuity 627
Minimum resolvable acuity 627
Minimum recognizable acuity 629
Minimum discriminable acuity 629
Limiting factors in visual acuity 630
Optical quality of the eye 630
What limits optical image formation of the eye? 632
Refractive error and defocus results in a marked loss of image quality 632
Photoreceptor size and spacing; aperture size; the “Nyquist” limit; aliasing 633
Cone to ganglion cell convergence 633
Eccentricity 634
Crowding in peripheral vision 636
Luminance 637
Contrast 637
Time 637
Motion 637
Anisotropies 638
Visual acuity and reading 638
Spatial vision with low contrast 639
The contrast sensitivity function represents our window of visibility 639
The CSF in peripheral vision 640
Clinical testing of visual acuity 642
Visual acuity chart design considerations 642
Clinical tests for CSF 643
Glare 643
Development of spatial vision 643
Development of visual acuity and CSF 643
Development of hyperacuity 644
Visual acuity through the lifespan 644
Amblyopia 645
Crowding and amblyopia 645
References 646
34 Color Vision 648
Molecular genetics of color vision and color deficiencies 648
Tests of color vision 650
Color appearance 650
Blue-yellow circuitry 650
Red-green circuitry 652
Black-white circuitry 653
Future directions 653
References 654
35 The Visual Field 655
Introduction 655
The psychophysical basis for perimetry 655
The physiologic basis for perimetry 656
Types of perimetric testing 656
Kinetic perimetry 657
Static perimetry 657
Suprathreshold static perimetry 658
Detection of perimetric sensitivity loss and interpretation of results 659
Patterns of visual field loss associated with different pathologic conditions 662
Determination of visual field progression 664
A guide for interpretation of visual field information 665
New perimetric test procedures 667
Short wavelength automated perimetry (SWAP) 667
Frequency doubling technology (FDT) perimetry 667
Flicker and temporal modulation perimetry 669
Motion perimetry 669
High pass resolution perimetry 670
Rarebit perimetry 670
Multifocal visual evoked potentials (mfVEP) 671
Conclusions 673
References 674
36 Binocular Vision 677
Introduction 677
Two eyes are better than one 677
Visual direction 677
Normal retinal correspondence 679
Abnormal retinal correspondence 681
Binocular (retinal) disparity 683
Stereopsis 685
Quantitative and qualitative stereopsis 686
Stereoacuity 687
Stereoacuity with refractive defocus 690
Spatial frequency and contrast effects on stereopsis 690
Spatial distortions from aniseikonia 690
Motion-in-depth 693
Suppression in normal binocular vision 694
Summary 695
References 695
37 Temporal Properties of Vision 698
Temporal summation and the critical duration 698
Factors affecting the critical duration 699
Temporal sensitivity to periodic stimuli 699
Critical flicker fusion frequency 699
Effect of stimulus luminance on CFF 700
Effect of stimulus chromaticity on CFF 701
Effect of eccentricity on CFF 701
Effect of stimulus size on CFF: the Granit–Harper law 701
Temporal contrast sensitivity 702
Chromatic temporal sensitivity 703
Spatial effects on temporal sensitivity 704
Mechanisms underlying temporal sensitivity 704
Surround effects on temporal sensitivity 705
Differences between mean-modulated and luminance-pedestal flicker 705
The effects of flicker on perception 706
Temporal phase segmentation 706
Clinical applications of temporal sensitivity measurements 707
Motion processing 708
Psychophysical and perceptual evidence for unique motion processing 708
The neural encoding of motion 708
Clinical applications of motion processing 709
References 710
11 Development and deprivation of vision 713
38 Development of Vision in Infancy 713
Methodologies for assessing infant vision and their interpretation 713
Preferential looking 713
Visual evoked potentials 713
Ocular following movements 713
Hierarchy of visual processing 714
Spatio-temporal vision 715
Temporal resolution 716
Grating acuity 716
Vernier acuity 717
Optotype acuity 718
Motion 718
Motion direction asymmetries 718
Binocular vision 719
Fusion 719
Stereopsis and disparity sensitivity 720
Age of onset of disparity sensitivity 720
Development of disparity sensitivity 720
Summary 722
References 722
39 Development of Retinogeniculate Projections 725
Retinogeniculate projections are refined during development 725
Activity-dependent refinement of retinogeniculate projections 726
What parameters of activity drive refinement? 726
Synaptic inputs change strength with segregation 728
Molecular mechanisms involved in activity-dependent axonal segregation 729
Molecular mechanisms guiding the formation of eye-specific axonal territories 729
Summary 730
Acknowledgments 730
References 730
40 Developmental Visual Deprivation 732
Effects of early monocular form deprivation 732
Constant monocular form deprivation 732
Perceptual deficits 732
Neural changes 733
Intermittent monocular deprivation 734
Alternating monocular deprivation 734
Reverse occlusion 734
Brief unrestricted vision during monocular deprivation 734
Critical period 735
Critical period for monocular form deprivation 735
Molecular mechanisms of ocular dominance plasticity 736
Effects of early monocular defocus 739
Constant monocular defocus 739
Perceptual deficits 739
Neural changes 739
Alternating defocus 740
Effects of early strabismus 742
Perceptual deficits 742
Animal models of strabismus 742
Neural changes 742
Effects of onset age and duration of strabismus 742
Onset age 744
Duration 744
Eye movement anomalies 745
Amblyopia 745
Perceptual deficits 746
Neural changes 746
Improved visual performance in adult subjects with developmental disorders 747
Summary 747
References 747
41 The Effects of Visual Deprivation After Infancy 750
Introduction 750
The neuronal effects of visual deprivation 750
Cross-modal processing in visually normal development 750
Cross-modal processing in visually normal adults 751
Cross-modal processing in early blind individuals 752
Tactile performance 752
Braille tactile processing 755
Non-Braille tactile processing 756
Auditory processing 756
Auditory localization 758
Auditory language 759
Cross-modal connectivity within occipital cortex of early blind individuals 759
Neuronal structure within occipital cortex of early blind individuals 761
Differences in cross-modal processing between the periphery and the fovea 761
Blindfolding studies 761
Restoration of vision 762
Concluding remarks 764
Acknowledgments 765
References 765
Index 767
A 767
B 769
C 770
D 773
E 774
F 776
G 777
H 778
I 778
J 780
K 780
L 780
M 781
N 783
O 784
P 785
Q 788
R 788
S 790
T 792
U 793
V 794
W 795
X 795
Y 795
Z 795