The Human Eye Structure and Function

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Format: Paperback
Pub. Date: 1999-06-15
Publisher(s): Sinauer Associates is an imprint of Oxford University Press
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Summary

We are a highly visual species. Most of our information about the world comes to us through our eyes and most of our cultural and intellectual heritage is stored and transmitted as words and images to which our vision gives access and meaning. Knowing more about our eyes and vision is, therefore, one path to better understanding ourselves. And, as it happens, the human eye is a fairly representative vertebrate eye; knowing more about it tells us much about the eyes of other animals and about how they view the world and us.

In more practical terms, a better understanding of the human eye allows us to intervene more intelligently and purposefully as we attempt to correct, modify, or ameliorate disorders of the eye brought on by trauma, disease, or senescence.

Understanding the eye requires an exploration of the relationship between its structure and its function--that is, a consideration not only of how the eye and its parts are constructed, but also of what they do and how they work. Thus, this book considers both the structure and the function of the human eye and how they are related, often using functional issues as a guide to the most meaningful and important features of the anatomy. Limited use of technical terms from the various disciplines that relate to the eye, definitions of terms as they are used, a glossary, and suggestions for additional reading are all included to make the text accessible to readers for whom the subject is new. Boxes interspersed throughout the text discuss methods used to study the structure of the eye and surgical procedures used to alter its structure in beneficial ways.

In addition to the main theme of structure and function, several subthemes make the general point, in different ways, that the eye and our understanding of it are dynamic and changing. Change on a geological timescale is represented by the evolutionary history of eyes generally and the human eye's place among the diversity of eyes in the animal kingdom; these issues are discussed in the Prologue. Change within a human lifetime begins with a chapter about the early stages of development in utero, continues throughout the book with the developmental histories of different parts of the eye, and concludes, in the Epilogue, with accounts of postnatal growth, maturation, and senescence. Change throughout human history in the way we have understood our eyes is another story, fragments of which are contained in a series of "vignettes" about some of the people and ideas that have influenced human thought about the eye over the past several thousand years.

The Human Eye: Structure and Function appeals to a wide audience, including all scientists who are interested in the eye and in vision; optometrists and ophthalmologists; and optometry students and ophthalmology residents.

Author Biography


Clyde W. Oyster is Professor Emeritus in the Department of Physiological Optics of the School of Optometry at The University of Alabama at Birmingham.

Table of Contents

Preface xxiii
Acknowledgments xxviii
Prologue A Brief History of Eyes
1(752)
The Antiquity of Eyes and Vision
1(8)
Thinking about the eye gave Charles Darwin ``a cold shudder''
1(1)
The history of the eye is embedded in the history of animals and molecules
2(1)
Several of the eye's critical molecules are ancient
3(2)
Eyes were invented by multicellular animals almost 600 million years ago
5(1)
Eyes arose not once, but numerous times, in different animal groups
6(2)
Eyes are most common in groups of motile animals living in lighted environments
8(1)
The Diversity and Distribution of Eyes
9(7)
The first step in vision is an eye that can sense the direction of incident light
9(2)
At least ten types of eyes can be distinguished by differences in their optical systems
11(2)
Vertebrates always have simple eyes, but invertebrates can have compound eyes, simple eyes, or both
13(3)
Paths and Obstacles to Perfection
16(5)
Simple eyes improve as they become larger
16(2)
Elaborate simple eyes may have evolved rapidly
18(1)
Compound eyes have inherent optical limitations in their performance
19(2)
An Ocular Bestiary: Fourteen Eyes and Their Animals
21(36)
Compound eye---focal apposition, terrestrial variety: Honeybee (Apis mellifica)
21(2)
Compound eye---focal apposition, aquatic variety: Horseshoe crab (Limulus polyphemus)
23(3)
Compound eye---afocal apposition: Monarch butterfly (Danaus plexippus)
26(3)
Compound eye---apposition with neural superposition: Housefly (Musca domestica)
29(2)
Compound eye---refracting superposition: Firefly (Photuris spp.)
31(2)
Compound eye---reflecting superposition: Crayfish
33(3)
Compound eye---parabolic superposition: Crabs
36(1)
Simple eye---pinhole: Nautilus
37(1)
Simple eye---refracting, aquatic variety: Octopus
38(2)
Simple eye---refracting, aquatic variety: Goldfish
40(3)
Simple eye---refracting, terrestrial variety: Pigeon (Columba livia)
43(2)
Simple eye---refracting, terrestrial variety: Jumping spiders (Metaphidippus spp.)
45(3)
Simple eye---reflecting: Scallops
48(2)
Simple eye---refracting, terrestrial variety: Humans (Homo sapiens)
50(7)
PART ONE Ocular Systems
Formation of the Human Eye
57(20)
Some Developmental Strategies and Operations
57(2)
Embryogenesis begins with cell proliferation, cell movement, and changes in cell shape
57(1)
Specialized tissues are formed by collections of cells that have become specialized themselves
58(1)
Proliferation, movement, and differentiation in a cell group may require communication with other cells
58(1)
Embryonic Events before the Eyes Appear
59(5)
The blastocyst forms during the first week of embryogenesis
59(1)
The inner cell mass becomes the gastrula, which is divided into different germinal tissues
60(2)
Neurulation begins the development of the nervous system
62(2)
Formation of the Primitive Eye
64(1)
Ocular development begins in the primitive forebrain
64(1)
The optic vesicle induces formation of the lens
64(1)
Elaboration of the Primitive Eye
65(6)
The optic cup and the lens form from different germinal tissues by changes in cell shape
65(2)
The optic cup is initially asymmetric, with a deep groove on its inferior surface
67(1)
Closure of the choroidal fissure completes the optic cup
67(1)
The lens vesicle forms in synchrony with the optic cup
67(1)
The primitive lens is the first ocular structure to exhibit cell differentiation
68(1)
All future growth of the lens comes from the early lens cells, some of which are ``immortal'' stem cells
69(1)
The precursors of the future retina, optic nerve, lens, and cornea are present by the sixth week of gestation
69(1)
In general, the eye develops from inside to outside
70(1)
Failures of Early Development
71(1)
``If anything can go wrong, it will''
71(1)
One or both eyes may fail to develop completely
71(1)
Congenital absence of the lens may be an early developmental failure
71(3)
Incomplete closure of the choroidal fissure can produce segmental defects in the adult eye
74
Vignette 1.1 The Eye of Mann
72(5)
Ocular Geometry and Topography
77(34)
Elements of Ocular Structure
77(5)
The human eye is a simple eye
77(1)
The outermost of the three coats of the eye consists of cornea, limbus, and sclera
78(1)
The middle coat---the uveal tract---includes the iris, ciliary body, and choroid
78(1)
The eye's innermost coat---the retina---communicates with the brain via the optic nerve
79(2)
Most of the volume of the eye is fluid or gel
81(1)
Image Quality and Visual Performance
82(8)
Images of point sources are always small discs of light whose size is a measure of optical quality
82(1)
The amount of smear or spread in the image of a point source is related to the range of spatial frequencies transmitted by the optical system
83(3)
The contrast sensitivity function specifies how well different spatial frequencies are seen by the visual system
86(1)
We can see flies when their images subtend about one minute of visual angle
87(3)
The Anatomy of Image Formation
90(7)
The quality of a focused image is affected by pupil size, curvatures of optical surfaces, and homogeneity of the optical media
90(2)
Defocusing produces large changes in the modulation transfer function
92(1)
The major anatomical factors that determine the refractive power of the eye are the curvatures of the cornea and lens and the depth of the anterior chamber
93(2)
Schematic eyes are approximations of the eye's optically relevant anatomy
95(2)
Eye Shape and Size
97(6)
Vertebrate eyes vary considerably in shape
97(1)
Both the cornea and the sclera are aspheric
97(3)
On average, the adult human eye measures twenty-four millimeters in all dimensions
100(1)
Axial lengths and other anatomical features vary among individuals, but most eyes are emmetropic
101(1)
Most refractive error is related to relatively large or small axial lengths
102(1)
The Eye's Axes and Planes of Reference
103
The eyes rotate around nearly fixed points
103(1)
Like the head, the eyes have three sets of orthogonal reference planes
104(2)
The pupillary axis is a measure of the eye's optical axis
106(1)
The line of sight differs from the pupillary axis by the angle kappa
107(1)
The angles kappa in the two eyes should have the same magnitude
107(1)
Eye position is specified by the direction of the line of sight in a coordinate system whose origin lies at the eye's center of rotation
108
Vignette 2.1 The Medieval Eye
80(8)
Box 2.1 Evaluating Visual Resolution
88(10)
Vignette 2.2 Fundamentum Opticum
98(13)
The Orbit
111(22)
The Bony Orbit
111(10)
The orbits are roughly pyramidal
111(1)
The large bones of the face form the orbital margin and much of the orbit's roof, floor, and lateral wall
112(2)
The sphenoid bone fills the apex of the orbital pyramid and contributes to the lateral and medial walls
114(4)
The lacrimal and ethmoid bones complete the medial wall between the maxillary bone in front and the sphenoid in back
118(1)
Three major and several minor foramina permit blood vessels and nerves to enter or exit the orbital cavity
118(1)
Blowout fractures of the orbit are a consequence of the relative weakness of the orbital plates
119(1)
Abnormal positioning of the eye relative to the orbital margin may indicate local or systemic pathology
120(1)
Infection and tumors can enter the orbital cavity through the large sinuses that surround the orbit
120(1)
Connections between the Eye and the Orbit
121(6)
Connective tissue lines the interior surface of the orbit
121(1)
Connective tissue surrounds the eye and extraocular muscles
121(3)
Check ligaments connect Tenon's capsule to the periorbita
124(1)
All structures in the orbital cavity are lined and interconnected with connective tissue
125(1)
Abnormal development of the connective tissue may affect movement of the eyes
125(1)
Fat fills the spaces in the orbital cavity that are not occupied by other structures
126(1)
The septum orbitale prevents herniation of orbital fat into the eyelids
127(1)
Development of the Orbital Bones
127
Many bones form first as cartilage templates
127(1)
Most orbital bones do not have cartilaginous templates
127(3)
The orbital plates begin to form during the sixth week of gestation
130(1)
The capacity of bone for growth, repair, and remodeling lasts many years
130(1)
The eyes and orbits rotate from lateral to frontal positions during development
130(1)
Most developmental anomalies of the orbital bones are associated with anomalies of the facial bones
131
Vignette 3.1 Sovereign of the Visible World
116(6)
Box 3.1 Visualizing the Orbit and Its Contents In Vivo
122(6)
Vignette 3.2 The Anatomy of Vesalius
128(5)
The Extraocular Muscles
133(58)
Patterns of Eye Movement
133(6)
Our eyes are always moving, and some motion is necessary for vision
133(2)
Large, rapid eye movements are used for looking around, for placing retinal images of interest on the fovea
135(1)
Slow eye movements are used to track or follow movement and to compensate for changes in head and body position
136(1)
Eye movement velocities may vary by a factor of 105
137(1)
Since the eyes have overlapping fields of view, their movements must be coordinated
137(1)
Slow movements of the eyes in opposite directions help keep corresponding images on the foveas in both retinas simultaneously
138(1)
Strabismus is a misalignment of the two visual axes under binocular viewing conditions
139(1)
Control of Eye Position
139(14)
The extraocular muscles are arranged as three reciprocally innervated agonist-antagonist pairs
139(2)
The eyes are stationary when the opposing forces exerted by the extraocular muscles are in balance
141(1)
Imbalanced forces produce eye rotations
142(1)
Muscle force is related to muscle length
143(1)
Most of the force that maintains eye position is passive force
144(1)
Equilibrium muscle lengths and forces for different gaze positions are functions of the innervational command
145(1)
Different patterns of innervation are required for fast and slow eye movements
146(1)
Extraocular motor neurons are located in three interconnected nuclei in the brainstem
146(1)
Motor commands are the result of interactions between visual and nonvisual inputs to the motor control centers
147(1)
A copy of the innervational command is used to verify the system's operation
148(1)
Extraocular motor neurons receive inputs from premotor areas in the brainstem to generate appropriate signals for saccadic eye movements
148(1)
The pathways for smooth pursuit movements and for vergences go through the cerebellum, but vergences have a separate control center near the oculomotor nucleus
149(4)
Extraocular Muscle Structure and Contractile Properties
153(12)
Muscle fibers are the units from which muscles are constructed
153(1)
Striated muscle fibers have a parallel arrangement of contractile proteins that interleave to cause contraction
154(1)
Striated muscle fibers differ in structural, histochemical, and contractile properties
155(1)
The extraocular muscles contain muscle fiber types not found in skeletal muscles
156(1)
Thick and thin extraocular muscle fibers differ in their contractile properties
156(1)
Different muscle fiber types are not randomly distributed within the muscles
157(1)
Different muscle fiber types may receive different innervational commands
158(1)
Extraocular muscles have very small motor units
159(2)
Acetylcholine at the neuromuscular junctions depolarizes the cell membrane by opening sodium channels
161(1)
The spread of depolarization along the sarcolemma may differ among muscle fiber types, producing different contractile properties
161(1)
Extraocular muscles exhibit high sensitivity to agents that mimic or block the action of acetylcholine
162(1)
Extraocular muscles often exhibit early symptoms of myasthenia gravis
163(1)
Neurotoxins that interfere with acetylcholine action can be used to alleviate strabismus and blepharospasm
163(2)
Sensory Endings in Extraocular Muscles and Tendons
165(3)
Skeletal muscles have two major types of sensory organs
165(1)
Human extraocular muscles have anatomically degenerate sensory organs and exhibit no stretch reflexes
166(1)
Passive extraocular muscle stretch may produce bradycardia
167(1)
Sensory endings in extraocular muscles probably do not convey information about eye position
167(1)
Sensory signals from the extraocular muscles may be involved in motor learning, motor plasticity, and development
168(1)
Actions of the Extraocular Muscles
168(15)
All of the extraocular muscles except the inferior oblique have their anatomical origins at the apex of the orbit
168(1)
The anatomical origin of the inferior oblique and the functional origin of the superior oblique are anterior and medial in the orbit
169(1)
The four rectus muscles are arranged as horizontal and vertical pairs, all inserting onto the anterior portion of the globe
170(1)
The horizontal recti rotate the eye in the horizontal plane around a vertical axis
170(1)
The vertical recti are responsible for upward and downward rotations of the eye
171(1)
The recti define a muscle cone within the orbital cavity that contains most of the ocular blood vessels and nerves
172(1)
The oblique muscles constitute a third functional pair, inserting onto the posterior portion of the eye
172(2)
Extraocular muscle actions cannot be measured directly
174(1)
The classic description of action of the extraocular muscles is based on the geometry of their origins and insertions
174(2)
Boeder diagrams attempt to describe the actions of the extraocular muscles completely
176(2)
The presence of Tenon's capsule and muscle pulleys invalidates the geometric model of extraocular muscle actions
178(2)
``There is no simple way to describe the action of these muscles on the eye!''
180(2)
A realistic model of the extraocular muscle system is important for the diagnosis and treatment of muscle paresis
182(1)
Development of the Extraocular Muscles
183
Each muscle develops from several foci in the mesoderm surrounding the optic cup
183(2)
The extraocular muscles appear after the optic cup, but before the orbital bones
185(1)
Different muscle fiber types form late in gestation and continue to develop postnatally
185(1)
Most developmental anomalies are associated with the connective tissue of the muscles or with their innervation
186(1)
The oculomotor system is not fully operational at birth
186
Box 4.1 Detecting Ocular Misalignment
140(12)
Box 4.2 Changing the Effects of Extraocular Muscle Contraction
152(12)
Vignette 4.1 Locating the Extraocular Muscles
164(20)
Vignette 4.2 In the Service of the Eye
184(7)
The Nerves of the Eye and Orbit
191(56)
Elements of Neural Organization
191(4)
The brain deals with information about the external world and the body
191(1)
Neurons are the anatomical elements of neural systems
191(2)
Neural circuits consist of neurons linked mostly by unidirectional chemical synapses
193(1)
The direction of neural information flow distinguishes between sensory and motor nerves
194(1)
Motor outputs are divided anatomically and functionally into somatic and autonomic systems
194(1)
The autonomic system is subdivided into the sympathetic and parasympathetic systems
194(1)
The Optic Nerve and the Flow of Visual Information
195(17)
In the optic nerve, the location of axons from retinal ganglion cells corresponds to their location on the retina
195(3)
Axons from the two optic nerves are redistributed in the optic chiasm
198(1)
The decussation of axons in the chiasm is imperfect
199(1)
Spatial ordering of axons changes in the optic tracts
200(1)
In the lateral geniculate nuclei, which are primary targets of axons in the optic tracts, inputs from the two eyes are separated into different layers
200(1)
Axons terminating in the lateral geniculate nuclei are spatially ordered
201(2)
Some axons leave the optic tracts for other destinations
203(1)
Axons terminating in the superior colliculi form discontinuous retinotopic maps
204(1)
Axons forming the afferent part of the pupillary light reflex pathway terminate in the pretectal nuclear complex
204(1)
Retinal inputs to the accessory optic system may help coordinate eye and head movement
205(1)
Retinal axons may provide inputs to a biological clock
205(2)
Lesions of the optic nerves and tracts produce defects in the visual fields
207(5)
Lesions in the secondary visual pathways can be observed only as motor deficits
212(1)
The Trigeminal Nerve: Signals for Touch and Pain
212(7)
Two of the three trigeminal divisions carry signals from the eye and surrounding tissues
212(1)
All somatosensory information from the eye is conveyed by the nasociliary nerve to the ophthalmic division of the trigeminal
213(1)
Sensory nerve fibers from the cornea, conjunctiva, limbus, and anterior sclera join to form the long ciliary nerves
213(2)
Stimulation of corneal or conjunctival nerve endings elicits sensations of touch or pain, a blink reflex, and reflex lacrimation
215(1)
Other sensory fibers from the eye are conveyed by the short ciliary nerves and the sensory root of the ciliary ganglion
215(1)
Most other branches of the ophthalmic nerve carry somatosensory fibers from the skin of the eyelids and face
215(2)
A few branches of the maxillary nerve pass through the orbit from the facial skin and the maxillary sinus
217(1)
Lesions in the branching hierarchy of the ophthalmic nerve produce anesthesia that helps identify the lesion site
218(1)
Viral infection of the trigeminal system can produce severe corneal damage
219(1)
The Extraocular Motor Nerves
219(7)
The three cranial nerves that innervate the extraocular muscles contain axons from clusters of cells in the brainstem
219(1)
Cells in different parts of the oculomotor nerve nucleus innervate the levator, the superior and inferior recti, the medial rectus, and the inferior oblique
219(2)
Axons destined for different muscles run together in the oculomotor nerve until it exits the cavernous sinus just behind the orbit
221(3)
The oculomotor nerve contains parasympathetic fibers bound for the ciliary ganglion
224(1)
Cells in the trochlear nerve nucleus innervate the contralateral superior oblique
224(1)
Abducens nerve cells innervate the ipsilateral lateral rectus
225(1)
All of the oculomotor nerves pass through the cavernous sinus on their way to the orbit
225(1)
The extraocular motor nerves probably contain sensory axons from muscle spindles and tendon organs
226(1)
Innervation of the Muscles of the Eyelids
226(2)
Three sets of muscles are associated with the eyelids
226(1)
The orbicularis is innervated by the facial nerve
227(1)
The superior and inferior tarsal muscles are innervated by the sympathetic system
227(1)
Ptosis may result from either oculomotor or sympathetic lesions
228(1)
Autonomic Innervation of Smooth Muscle within the Eye
228(7)
The superior cervical ganglion is the source of most sympathetic innervation to the eye
228(2)
Sympathetic fibers enter the eye in the short ciliary nerves
230(1)
Sympathetic innervation of the dilator muscle acts at alpha-adrenergic receptors to dilate the pupils
230(1)
The arterioles in the uveal tract receive sympathetic innervation that produces vasoconstriction
231(1)
Horner's syndrome is the result of a central lesion in the sympathetic pathway
231(1)
Parasympathetic fibers entering the eye originate in the ciliary or the pterygopalatine ganglion
231(1)
Axons from cells in the ciliary ganglion innervate the sphincter and the ciliary muscle
232(1)
Axons from the pterygopalatine ganglion cells innervate vascular smooth muscle in the choroid
233(1)
Accommodation and pupillary light reflexes share efferent pathways from the Edinger-Westphal nuclei to the eyes; pupillary reflexes are mediated by retinal signals reaching the Edinger-Westphal nuclei through the pretectal complex
233(1)
Deficient pupillary reflexes may be associated with midbrain lesions
234(1)
Innervation of the Lacrimal Gland
235(1)
Axons from cells in the pterygopalatine ganglion reach the lacrimal gland via the zygomatic and lacrimal nerves
235(1)
The efferent pathway for lacrimal innervation begins in the facial nerve nucleus
235(1)
Basal tear production may require tonic innervation of the lacrimal gland
236(1)
Some Issues in Neural Development
236(5)
Specialized growth cones guide the extension of axons and dendrites
236(1)
Pathfinding by growth cones depends on recognition of local direction signs
237(1)
Target recognition and acquisition may require specific markers produced by the target cells
238(1)
Many early neurons are eliminated as mature patterns of connectivity are established
238(1)
Adult connectivity patterns are not always complete at birth, and postnatal development is subject to modification
239(1)
Ocular albinism is associated with a pathfinding error in the development of optic nerve axons
239(1)
Anomalous innervation of the extraocular muscles may be the result of pathfinding or target recognition errors
240(1)
Some forms of amblyopia may be related to problems with postnatal establishment and maintenance of synaptic connections
240(1)
Innervation of the extraocular muscles begins early in gestation, sensory innervation much later
241(1)
Postnatal Neuron Growth and Regeneration
241
Most postnatal neuron growth is interstitial growth
241(1)
Neurons do not undergo mitosis postnatally
242(1)
Spinal neurons in peripheral nerves can regenerate after being damaged
242(1)
Central nervous system neurons do not regenerate following major damage
242(1)
Corneal nerve endings will regenerate following local damage
243(1)
Neuronal degeneration can affect other, undamaged neurons
243
Vignette 5.1 The Integrative Action of the Nervous System
196(10)
Box 5.1 Tracing Neural Pathways: Degeneration and Myelin Staining
206(4)
Vignette 5.2 Seeing One World with Two Eyes: The Problem of Decussation
210(12)
Box 5.2 Tracing Neuronal Connections: Axonal Transport Methods
222(25)
Blood Supply and Drainage
247(44)
Distributing Blood to Tissues
247(8)
Arteries control blood flow through capillary beds, and veins regulate blood volume
247(1)
Blood flow through capillary beds can be controlled locally or systemically
248(3)
Capillary beds in a tissue may be independent or interconnected
251(1)
The interchange between blood and cells depends partly on the structure of the vascular endothelium
252(1)
Capillary endothelium is renewable, and capillary beds can change
253(1)
Neovascularization is a response to altered functional demands
253(1)
Structurally weakened capillaries may be prone to excessive neovascularization
254(1)
The Ophthalmic Artery and Ophthalmic Veins
255(5)
The ophthalmic artery distributes blood to the eye and its surroundings
255(2)
Blood supplied to tissues by the ophthalmic artery is drained to the cavernous sinus by the ophthalmic veins
257(3)
Supply and Drainage of the Eye
260(19)
Muscular arteries supply both the extraocular muscles and the anterior segment of the eye
260(1)
The anterior ciliary arteries contribute to the episcleral and intramuscular arterial circles
261(1)
The conjunctiva and corneal arcades are supplied by branches from the episcleral arterial circle and drained by the episcleral and anterior ciliary veins
262(1)
The system of episcleral veins drains the conjunctiva, corneal arcades, and limbus
263(1)
The posterior ciliary arteries divide into long and short posterior ciliary arteries that supply different regions
264(1)
The intramuscular and major arterial circles are formed in the ciliary body by branches from the anterior and long posterior ciliary arteries
265(3)
Blood supply to the anterior segment is redundant, but there is some segmentation in supply to the iris
268(1)
The short posterior ciliary arteries terminate in the choriocapillaris, which supplies the retinal photoreceptors
269(3)
The short posterior ciliary arteries contribute to the supply of the optic nerve through the circle of Zinn
272(2)
The short posterior ciliary arteries sometimes contribute to the supply of the inner retina
274(1)
The central retinal artery enters the eye through the optic nerve and ramifies to supply the inner retina
275(2)
The central retinal vein exits the eye through the optic nerve
277(1)
The vortex veins drain most of the uveal tract
277(1)
The vortex veins have segmented drainage fields, but they are heavily anastomotic
278(1)
Supply and Drainage of the Eyelids and Surrounding Tissues
279(5)
The lacrimal gland is supplied by the lacrimal artery and drained by lacrimal veins
279(1)
The eyelids are supplied by branches of the lacrimal, ophthalmic, facial, and infraorbital arteries
279(3)
The terminal branches of the ophthalmic artery leave the orbit to supply the skin and muscles of the face
282(1)
The infraorbital artery runs under the orbital floor
283(1)
The orbital veins are connected to the veins of the face, the pterygoid plexus, and the nose
283(1)
Development of the Ocular Blood Vessels
284
Primitive embryonic blood vessels appear very early in the eye's development
284(1)
Several parts of the early ocular vasculature are transient and do not appear in the mature eye
284(3)
The anterior ciliary system forms later than the posterior ciliary system
287(1)
Remnants of normally transient, embryological vasculature may persist in the mature eye
287
Vignette 6.1 Circulation of the Blood
256(14)
Box 6.1 Tracing Hidden Blood Vessels: Vascular Casting
270(10)
Vignette 6.2 Blood Vessels inside the Eye
280(11)
The Eyelids and the Lacrimal System
291(34)
Structure and Function of the Eyelids
291(16)
Structural rigidity of the lids is provided by the tarsal plates
291(1)
The tarsal plates are made of dense connective tissue in which glands are embedded
292(2)
The palpebral fissure is opened by muscles inserting onto or near the edges of the tarsal plates
294(2)
The palpebral fissure is closed by contraction of the orbicularis
296(2)
Blinking may be initiated as a reflex response or as a regular, spontaneous action
298(1)
Lid movements during spontaneous blinks move tears across the cornea
299(1)
Overaction of the orbicularis may appear as blepharospasm or as entropion
299(1)
Paresis of the orbicularis produces ectropion and epiphora
300(1)
Other glands in the lids are associated with the eyelashes
301(1)
The skin on the lids is continuous with the conjunctiva lining the posterior surface of the lids and covering the anterior surface of the sclera
302(1)
The orbital septum is a connective tissue sheet extending from the orbital rim to the tarsal plates
303(1)
The shape and size of the palpebral fissure vary
304(2)
The overall structure of the lids consists of well-defined planes or layers of tissue
306(1)
Tear Supply and Drainage
307(8)
Most of the tear fluid is supplied by the main lacrimal gland
307(1)
Secretion by the lacrimal gland is regulated by autonomic inputs operating through a second-messenger system
308(1)
The composition of the lacrimal gland secretion varies with the secretion rate
309(1)
Dry eye may result from a decreased amount of tears, abnormal tear composition, or both
310(1)
Tears are drained off at the medial canthus and deposited in the nasal cavity
311(1)
Pressure gradients created by contraction of the orbicularis during blinks move tears through the canaliculi into the lacrimal sac
312(3)
Formation of the Eyelids and the Lacrimal System
315(10)
The eyelids first appear as folds in the surface ectoderm, which gives rise to the lid glands
315(1)
The lacrimal gland and the lacrimal drainage system derive from surface ectoderm
316(2)
Most developmental anomalies in the eyelids and lacrimal system are problems in lid position or blockage of the drainage channels
318(1)
Anomalous innervation can produce eyelid movements linked to contraction of muscles in the jaw
318(7)
PART TWO Components of the Eye
The Cornea and the Sclera
325(54)
Components and Organization of the Cornea and Sclera
325(24)
The cornea, sclera, and limbus are made primarily of collagen fibrils
325(1)
Collagen is embedded in a polysaccharide gel that forms the extracellular matrix
326(1)
The fibroblasts in the corneal and scleral stroma constitute a small fraction of the stroma's volume
327(1)
Collagen fibrils in the cornea are highly organized; those in the sclera are not
328(3)
The structure of the corneal stroma is altered in Bowman's layer
331(1)
Corneal transparency is a function of its regular structure
332(2)
Collagen organization in the stroma and corneal transparency depend on intact epithelium and endothelium
334(1)
The corneal epithelium is a multilayered, renewable barrier to water movement into the cornea
335(4)
The corneal endothelium is a single layer of metabolically active cells
339(1)
The endothelial cell tiling changes with time because cells that die cannot be replaced
340(5)
Descemet's membrane separates the endothelium from the stroma
345(2)
Nerve endings in the cornea give rise to sensations of touch or pain, a blink reflex, and reflex lacrimation
347(1)
The epithelium contains a dense array of free terminals of nerve fibers from the long ciliary nerves
347(2)
Corneal sensitivity can be measured quantitatively
349(1)
The dense innervation of the cornea makes it subject to viral infection
349(1)
The Cornea as a Refractive Surface
349(16)
The optical surface of the cornea is the precorneal film covering the surface of the epithelium
349(2)
The cornea's outline is not circular, its thickness is not uniform, and its radius of curvature is not constant
351(2)
The shape of the cornea is determined by comparison to a sphere
353(1)
The cornea does not have a single, specifiable shape
354(2)
Contact lenses can affect corneal shape and structure directly or indirectly
356(2)
The shape and optical properties of the cornea can be permanently altered
358(1)
Surgically reshaped corneas may change with time
359(1)
Corneal shape can be changed by removing tissue
360(1)
Stromal reshaping leaves the epithelium intact
361(1)
Corneal grafts are used to repair optically damaged corneas
361(4)
Corneal Healing and Repair
365(6)
The epithelium heals quickly and completely
365(3)
Corneal healing may require limbal transplants
368(1)
Repair of damage to the stroma produces translucent scar tissue
368(1)
The endothelium repairs itself by cell expansion and migration
369(1)
Corneal graft incisions are repaired by the normal healing processes
369(1)
Radial keratotomy incisions are repaired by epithelial hyperplasia and collagen formation
370(1)
Photorefractive keratectomy ablations are healed mostly by the epithelium
371(1)
Growth and Development of the Cornea
371
The epithelium and endothelium are the first parts of the cornea to appear
371(1)
The stroma is derived from neural crest cells associated with the mesoderm
372(1)
The regular arrangement of the stromal collagen appears soon after collagen production begins
372(2)
Corneal growth continues for a few years postnatally
374(1)
Anomalous corneal development can produce misshapen or opaque corneas
375
Box 8.1 Biomicroscopy of the Cornea
342(8)
Vignette 8.1 The Invisible Made Visible
350(12)
Box 8.2 Some Reservations about Corneal Refractive Surgery
362(4)
Vignette 8.2 The Art of William Bowman
366(13)
The Limbus and the Anterior Chamber
379(32)
The Anterior Chamber and Aqueous Flow
379(11)
The anterior chamber is the fluid-filled space between the cornea and the iris
379(1)
The angle of the anterior chamber varies in magnitude
380(1)
Aqueous is formed by the ciliary processes and enters the anterior chamber through the pupil
381(2)
Aqueous drains from the eye at the angle of the anterior chamber
383(2)
Intraocular pressure depends on the rate of aqueous production and the resistance to aqueous outflow
385(5)
The Anatomy of Aqueous Drainage
390(15)
The scleral spur is an anchoring structure for parts of the limbus and the ciliary body
390(2)
The trabecular meshwork is made of interlaced cords of tissue extending from the apex of the angle to the margin of the cornea
392(1)
Schwalbe's ring separates the trabecular meshwork from the cornea
393(1)
The trabecular cords have a collagen core wrapped with endothelial cells
393(2)
The major source of outflow resistance is the juxta-canalicular tissue separating the canal of Schlemm from the trabecular spaces
395(1)
The canal of Schlemm encircles the anterior chamber angle
395(2)
Aqueous enters the canal of Schlemm by way of large vacuoles in the endothelial lining of the canal
397(2)
Aqueous drains out of the canal into venous plexuses in the limbal stroma
399(1)
Pilocarpine reduces intraocular pressure, probably by an effect of ciliary muscle contraction on the structure of the trabecular meshwork
400(2)
An effective way to reduce intraocular pressure seems to be to increase the uveoscleral outflow
402(1)
Surgery for glaucoma aims to increase aqueous outflow
402(1)
The outer surface of the limbus is covered with episcleral tissue and a heavily vascularized conjunctiva
403(2)
Development of the Limbus
405
The anterior chamber is defined by the iris growing between the developing cornea and lens
405(1)
The angle of the anterior chamber opens during development as the root of the iris shifts posteriorly
406(2)
The trabecular meshwork develops between the fourth and eighth months
408(1)
Most developmental anomalies in the limbus are associated with structural anomalies that affect other parts of the anterior chamber
408
Box 9.1 Through the Looking Glass: Gonioscopy
382(6)
Box 9.2 Estimating the Pressure Within: Tonometry
388(23)
The Iris and the Pupil
411(36)
Functions of the Iris and Pupil
411(9)
The iris is an aperture stop for the optical system of the eye
411(1)
The entrance pupil is a magnified image of the real pupil
412(1)
Variation of pupil size changes the amount of light entering the eye, the depth of focus, and the quality of the retinal image
413(1)
Pupil size varies with illumination level, thereby helping the retina cope with large changes in illumination
414(2)
Pupil size varies with accommodation and accommodative convergence
416(1)
The pupillary near response is smaller in children than in adults
417(1)
The pupil is in constant motion, and it reacts quickly to changes in retinal illumination
418(1)
Decreased iris pigmentation in ocular albinism affects the optical function of the iris
419(1)
Structure of the Iris
420(12)
The pupils in the two eyes are normally the same size and are decentered toward the nose
420(1)
The iris is constructed in layers and regional differences in the iris are related to the different muscles within them
421(1)
The anterior border layer is an irregular layer of melanocytes and fibroblasts interrupted by large holes
422(1)
The iris stroma has the same cellular components as the anterior border layer, but loosely arranged
423(2)
Small blood vessels run radially through the stroma, anastomosing to form the minor arterial circle and supply the iris muscles
425(1)
The sphincter and dilator occupy different parts of the iris and have antagonistic actions
426(1)
The sphincter is activated by the parasympathetic system, the dilator by the sympathetic system
427(1)
The anterior pigmented epithelium is a myoepithelium, forming both the epithelial layer and the dilator muscle
428(2)
The posterior epithelial cells contact the anterior surface of the lens
430(2)
Surgery for closed-angle glaucoma often involves the iris rather than the limbus
432(1)
Some Clinically Significant Anomalies of the Iris and Pupil
432(8)
Changes in iris color after maturity are potentially pathological
432(1)
Differences between the two eyes in pupil size or pupillary responses to light are commonly associated with neurological problems
433(2)
Anisocoria and unresponsive pupils are often associated with defects in the efferent part of the innervational pathways
435(2)
Clinically useful drugs affecting pupil size fall into four functional groups
437(3)
Development of the Iris
440(7)
The iris stroma forms first by migration of undifferentiated neural crest cells
440(1)
The epithelial layers and the iris muscles develop from the rim of the optic cup and are therefore of neuroectodermal origin
440(2)
The pupil is the last feature of the iris to appear
442(1)
Most postnatal development of the iris is an addition of melanin pigment
443(1)
Segmental defects and holes in the iris result from unsynchronized or failed growth of the optic cup rim
443(2)
An ectopic pupil is improperly centered in an otherwise normal iris
445(1)
A persistent pupillary membrane may be the result of either insufficient tissue atrophy or tissue hyperplasia
445(2)
The Ciliary Body and the Choroid
447(44)
Anatomical Divisions of the Ciliary Body
447(2)
The ciliary processes characterize the pars plicata
447(2)
The ciliary muscle extends through both pars plicata and pars plana
449(1)
The Ciliary Processes and Aqueous Formation
449(12)
The ciliary processes are mostly filled with blood vessels
449(2)
The capillaries in the ciliary processes are highly permeable
451(1)
Two layers of epithelium lie between the capillaries and the posterior chamber
452(1)
Aqueous formation involves metabolically driven transport systems
452(2)
The ciliary epithelium is anatomically specialized as a blood-aqueous barrier
454(1)
Ions are transported around the band of tight junctions to produce an osmotic gradient in the basal folds of the unpigmented epithelium
455(1)
Aqueous production varies during the day and declines with age
456(2)
The major classes of drugs used to reduce aqueous production interact either with adrenergic membrane receptors or with the intracellular formation of bicarbonate ions
458(1)
The pars plana is covered by epithelial layers that are continuous with the epithelial layers of the pars plicata
459(2)
The Ciliary Muscle and Accommodation
461(16)
The ciliary muscle has three parts with a complex geometry
461(2)
Contraction of the ciliary muscle produces movement inward toward the lens so that the muscle behaves like a sphincter
463(2)
The zonule provides a mechanical linkage between ciliary muscle and lens
465(3)
Accommodation is a result of ciliary muscle contraction
468(1)
The primary stimulus to accommodation is retinal image blur
469(3)
Accommodative amplitude decreases progressively with age
472(1)
Presbyopia is not a consequence of reduced innervation to the ciliary muscle
473(1)
Aging of the ciliary muscle is unlikely to be a significant factor in presbyopia
474(3)
The Choroid
477(5)
The choroidal stroma consists of loose connective tissue and dense melanin pigment
477(1)
Blood vessels that supply and drain the capillary bed supplying retinal photoreceptors make up the main part of the choroid
478(1)
The choriocapillaris is heavily anastomotic but has local functional units
479(1)
The choriocapillaris varies in capillary density and in the ratio of arterioles to venules
480(1)
Capillaries in the choriocapillaris are specialized for ease of fluid movement across the capillary endothelium
481(1)
Bruch's membrane lies between capillaries and pigmented epithelium in both the choroid and the pars plana of the ciliary body
481(1)
Development of the Ciliary Body and Choroid
482
The ciliary epithelium arises from the optic cup, the ciliary muscle from neural crest cells
482(1)
Formation of the ciliary epithelium may be induced by the lens
483(1)
Formation of the ciliary muscle may be induced by the ciliary epithelium
484(1)
The ciliary muscle begins to form during the fourth month and continues to develop until term
485(1)
The muscles associated with the eye originate from different germinal tissues
486(1)
The ciliary processes form in synchrony with the vascular system in the ciliary body
486(1)
The zonule is produced by the ciliary epithelium
486(2)
The choroidal vasculature has two developmental gradients: center to periphery and inside to outside
488
Vignette 11.1 The Source
470(21)
The Lens and the Vitreous
491(54)
Structure of the Lens
492(21)
Some unusual proteins, the crystallins, are the dominant structural elements in the lens
492(2)
Dense, uniform packing of the crystallins within lens cells is responsible for lens transparency
494(1)
Crystallins are highly stable molecules, making them some of the oldest proteins in the body, but they can be changed by light absorption and altered chemical environments
494(2)
a-Crystallins may play a special role in maintaining native crystallin structure over time
496(1)
The lens is formed of long, thin lens fibers arranged in concentric shells to form a flattened spheroid
497(1)
Lens fibers in each shell meet anteriorly and posteriorly along irregular lines
498(1)
Lens shells are bound together with miniature locks and keys, a kind of biological Velcro
499(2)
The anterior epithelium is the source of new cells for the lens
501(2)
Elongating epithelial cells at the equator become long lens fibers that form new shells in the lens
503(1)
The size of the lens and the number of lens fibers increase throughout life
503(1)
Each new lens shell has one more fiber than the previous shell and about five new shells are added each year after the age of five
504(1)
An aged lens has about 2500 shells and 3.6 million lens fibers
505(2)
The lens capsule encloses the lens shells and epithelium
507(3)
The locations at which the zonule inserts onto the lens change with age
510(3)
The Lens as an Optical Element
513(17)
The refractive index of the ocular lens varies from one part of the lens to another
513(1)
Lens transparency is related not only to protein regularity but also to water content, which is maintained by ion pumping in the epithelium
514(1)
The lens contains several different optical zones
515(1)
The lens surfaces are parabolic and therefore flatten gradually from the poles to the equator
516(3)
Both anterior and posterior lens surfaces become more curved with accommodation, but the anterior surface change is larger
519(1)
The lens thickens with age and its curvatures increase, but unaccommodated lens power does not increase with age
520(2)
The increased lens surface curvatures in accommodation are primarily a consequence of tissue elasticity
522(1)
The presbyopic lens is aging, fat, and unresponsive
523(2)
Presbyopia is largely, if not solely, associated with age-related changes in the lens
525(1)
Cataracts, most of which are age-related, take different forms and can affect any part of the lens
526(4)
The Vitreous
530(9)
The vitreous is the largest component of the eye
530(1)
The primary structural components of the vitreous are collagen and hyaluronic acid
530(2)
The external layer of the vitreous---the vitreous cortex---attaches the vitreous to surrounding structures
532(1)
Inhomogeneity of the vitreous structure produces internal subdivisions in the vitreous
533(1)
The vitreous changes with age
533(3)
Shrinkage of the vitreous gel may break attachments to the retina
536(1)
Altered activity of cells normally present in the vitreous or the introduction of cells from outside the vitreous may produce abnormal collagen production and scar formation
537(1)
Vitrectomy removes abnormal portions of the vitreous
538(1)
Development of the Lens and Vitreous
539
The lens forms from a single cell line
539(1)
Most failures of lens development are manifest as congenital cataracts
540(1)
The primary vitreous forms around the embryonic hyaloid artery
541(1)
The secondary vitreous, initially acellular, forms outside the vasa hyaloidea propria
541(1)
Most developmental anomalies in the vitreous represent incomplete regression of the hyaloid artery system
541
Vignette 12.1 Putting the Lens in Its Proper Place
512(16)
Box 12.1 Cataract Surgery
528(17)
Retina I: Photoreceptors and Functional Organization
545(50)
The Retina's Role in Vision
545(4)
The retina detects light and tells the brain about aspects of light that are related to objects in the world
545(1)
Objects are defined visually by light and by variations in light reflected from their surfaces
546(1)
The retina makes sketches of the retinal image from which the brain can paint pictures
547(2)
Functional Organization of the Retina
549(11)
Photoreceptors catch photons and produce chemical signals to report photon capture
549(2)
Photoreceptor signals are conveyed to the brain by bipolar and ganglion cells
551(1)
Lateral pathways connect neighboring parts of the retina
552(2)
Recurrent pathways may assist in adjusting the sensitivity of the retina
554(1)
The retina has anatomical and functional layers
555(5)
Catching Photons: Photoreceptors and Their Environment
560
Each photoreceptor contains one of four photopigments, each of which differs in its spectral absorption
560(2)
Color vision requires more than one photopigment
562(2)
The photopigments are stacked in layers within the outer segments of the photoreceptor
564(1)
Light absorption produces a structural change in the photopigments
565(1)
Structural change in the photopigment activates an intracellular second-messenger system using cGMP as the messenger
566(2)
A decrease in cGMP concentration closes cation channels, decreases the photocurrent, and hyperpolarizes the photoreceptor
568(2)
Absorption of one photon can produce a detectable rod signal
570(2)
Photocurrent in the outer segment decreases in proportion to the number of absorbed photons
572(1)
Photopigments activated by photon absorption are inactivated, broken down, and then regenerated
573(2)
Photoreceptor sensitivity is modulated by intracellular Ca2+
575(3)
Changes in photoreceptor sensitivity account for less than half of the retina's sensitivity increase in the dark and sensitivity decrease in the light
578(1)
The tips of photoreceptor outer segments are surrounded by pigment epithelial cell processes
579(1)
The pigment epithelium and the interphotoreceptor matrix are necessary for photopigment regeneration
580(3)
Both rods and cones undergo a continual cycle of breakdown and renewal
583(2)
The inner segments of photoreceptors assemble the proteins to construct the outer segment membranes
585(1)
The inner segments form tight junctions with Muller's cells; these junctions are the external limiting membrane
586(4)
Photoreceptors signal light absorption by decreasing the rate of glutamate release from their terminals
590(2)
Glutamate release from a photoreceptor is subject to modification by activity in other photoreceptors
592
Vignette 13.1 ``Everything in the Vertebrate Eye Means Something''
558(37)
Retina II: Editing Photoreceptor Signals
595(54)
The Editing Process
595(2)
Interactions among Photoreceptors, Horizontal Cells, and Bipolar Cells
597(13)
Horizontal cells integrate photoreceptor signals
597(4)
Horizontal cells receive inputs from photoreceptors and send signals of opposite sign back to the photoreceptor terminals, using GABA as the neurotransmitter
601(1)
Horizontal cell connections emphasize differences in illumination between different photoreceptors
602(3)
Different glutamate receptors on cone bipolar cells cause increases and decreases in light intensity to be reported by ON and OFF bipolar cells, respectively
605(1)
Signals from both red and green cones go to midget bipolar cells, which are specific for cone type, and to diffuse bipolar cells, which are not cone specific
606(2)
Blue cones have their own bipolar cells
608(1)
Rods have sign-inverting synapses to rod bipolar cells, which do not contact ganglion cells but send signals to the cone pathways through an amacrine cell
608(2)
Interactions among Bipolar Cells, Amacrine Cells, and Ganglion Cells
610(14)
Bipolar cell terminals in the inner plexiform layer release glutamate at synapses to amacrine or ganglion cells and receive inputs from amacrine cells
610(1)
Bipolar cells terminate at different levels within the inner plexiform layer, thereby creating functional sublayers
611(5)
Amacrine cells vary in the extent over which they promote lateral interactions among vertical pathways and in the levels of the inner plexiform layer in which they operate
616(2)
Amacrine cells exert their effects mainly at glycine and GABA synapses, while several other neurotransmitters or neuromodulators play subsidiary roles
618(2)
The effects of neurotransmission depend on postsynaptic receptors
620(1)
Amacrine cell connections centering on the AII amacrine cells illustrate difficulties in understanding amacrine cell operations
621(3)
Ganglion Cell Signals to the Brain: Dots for the Retinal Sketches
624
Most ganglion cells are midget or parasol cells
624(4)
The small region of the world seen by a ganglion cell is its receptive field
628(2)
The concentric organization of excitation and inhibition makes ganglion cells sensitive to contrast rather than to average light intensity
630(1)
Ganglion cell receptive fields can be thought of as filters that modify the retinal image
631(5)
Sensitivity functions of ganglion cell receptive fields differ in size and in the strength of their inhibitory components
636(1)
Ganglion cell signals differ in their reports on stimulus duration and on the rate of intensity change
637(2)
Midget ganglion cells have wavelength information embedded in their signals, but only small bistratified cells are known to convey specific wavelength information
639(3)
Axons from midget and parasol ganglion cells go to different layers in the lateral geniculate nucleus
642(1)
Ganglion cell responses are the elements of retinal sketches
643
Vignette 14.1 The Retina Comes to Light
612(14)
Vignette 14.2 The Shoemaker's Apprentice
626(6)
Box 14.1 Studying Individual Neurons
632(17)
Retina III: Regional Variation and Spatial Organization
649(52)
Making Retinal Sketches out of Dots: Limits and Strategies
649(11)
The detail in a sketch is limited by dot size and spacing, and cones set the dot size in the central retina
649(3)
The entire retinal image cannot be sketched in great detail
652(1)
Most retinas are organized around points or lines
653(1)
Retinal sketches should be continuous, with no unnecessary blank spots
654(3)
Tilings do not need to be regular, and tiles do not have to be the same size
657(2)
Tilings formed by axonal or dendritic arbors at different levels of the retina need not match precisely
659(1)
Spatial Organization of the Retina
660(31)
The fovea is a depression in the retina where the inner retinal layers are absent
660(1)
The spatial distribution of a pigment in and around the fovea is responsible for entoptic images associated with the fovea
661(2)
Photoreceptor densities vary with respect to the center of the fovea, where cones have their maximum density and rods are absent
663(1)
The human retina varies from center to periphery in terms of the spatial detail in the retinal sketch
664(2)
Maximum cone densities vary among different retinas by a factor of three
666(1)
The human retina has about 4.5 million cones and 91 million rods
666(1)
Blue cones have a different distribution than red and green cones have the center of the fovea is dichromatic
667(1)
There are more red cones than green cones, and more green cones than blue cones
668(1)
The distribution of different types of cones is neither regular nor random
668(4)
Cone pedicles probably tile the retina in and near the fovea, but rod spherules probably never form a single-layered tiling
672(2)
The pedicles of cones in and near the fovea are displaced radially outward from the cone inner segments, but spatial order is preserved
674(1)
The density of horizontal cells is highest near the fovea and declines in parallel with cone density
675(1)
Neither H1 nor H2 horizontal cells form tilings
676(1)
All types of cone and rod bipolar cells are distributed like their photoreceptor types
676(2)
The different types of bipolar cells provide different amounts of coverage with their dendrites
678(1)
All bipolar cell terminals form tilings at different levels in the inner plexiform layer
679(2)
AII amacrine cells tile the retina, varying in density as ganglion cells do
681(1)
Medium- and large-field amacrine cells are low-density populations whose processes generate high coverage factors
682(3)
Ganglion cell density declines steadily from the parafovea to the periphery of the retina
685(2)
Midget and parasol ganglion cell dendrites tile at different levels in the inner plexiform layer
687(3)
Spatial resolution is limited by cone spacing in the fovea and parafovea and by midget ganglion cells elsewhere in the retina
690(1)
A Final Look at Three Small Pieces of Retina: Dots for the Retinal Sketches
691
A sampling unit is the smallest retinal region containing at least one representative from each type of ganglion cell
691(1)
Sampling units are smallest at the foveal center and are dominated by cone signals
692(2)
Rods and blue cones become significant in the parafoveal sampling units
694(1)
Rods and rod pathways dominate in peripheral sampling units
695(3)
The problem of understanding how the retina works can be reduced to the problem of understanding its sampling units
698(1)
The central representation of a sampling unit depends on the number of ganglion cells it contains
698
Box 15.1 Locating Species of Molecules: Immunohistochemistry
670(31)
The Retina In Vivo and the Optic Nerve
701(52)
Electrical Signals and Assessment of Retinal Function
702(6)
A difference in electrical potential exists between the vitreal and choroidal surfaces of the retina and between the front and back of the eye
702(1)
The electroretinogram measures a complex change in voltage in response to retinal illumination
702(1)
The a-wave and off effect are generated by the photoreceptors, the c-wave by the pigment epithelium
703(1)
The b-wave is either a direct reflection of ON bipolar cell activity or is indirectly related to their activity by a secondary potential arising from Muller's cells
704(2)
The ERG is useful as a gross indicator of photoreceptor function
706(1)
Multifocal ERGs provide assessments of retinal function within small areas of the retina
707(1)
The Retinal Vessels and Assessment of Retinal Health
708(11)
The retina in vivo is invisible
708(3)
Since the choroidal circulation is usually not directly visible, irregularities and nonuniformities on the fundus are commonly indicators of pathology
711(1)
The central retinal artery is an end-arterial system
712(1)
The capillaries supplied by the central retinal artery ramify in the inner two-thirds of the retina
713(1)
Retinal detachment separates photoreceptors from their blood supply
714(1)
The foveal center lacks capillaries
715(1)
Retinal capillaries are specialized to create a blood-retina barrier
715(2)
Retinal blood flow is autoregulated
717(1)
The arterial and venous branches on the retinal surface can be distinguished ophthalmoscopically
717(1)
Drainage of the inner retina is segmental
718(1)
The Optic Nerve
719(13)
All ganglion cell axons and all branches of the central retinal artery and vein converge at the optic nerve head
719(1)
The nerve head and the optic nerve consist primarily of axon bundles separated by sheaths of glial cells and connective tissue
719(2)
The blood supply and drainage differ between the pre- and postlaminar portions of the nerve head
721(1)
Ganglion cell axons form a stereotyped pattern as they cross the retina to the optic nerve head
722(1)
Axons from many widely separated ganglion cells are collected in bundles in the nerve fiber layer
723(1)
Axon bundles have an orderly arrangement in the nerve head
724(3)
Scotomas observed in advanced stages of glaucoma correspond to those produced by lesions along the superior and inferior temporal margin of the nerve head
727(1)
The lamina cribrosa is weaker than the rest of the sclera
727(1)
Field defects in glaucoma may be due to blockage of axonal transport secondary to deformation of the lamina cribrosa
728(2)
Ganglion cell loss in experimental glaucoma does not appear to be selective by cell type or axon diameter
730(2)
Development of the Retina and Optic Nerve
732
The retina develops from the two layers of the optic cup
732(1)
Retinal development proceeds from the site of the future fovea to the periphery
733(1)
Retinal neurons have identifiable birthdays
733(1)
Ganglion cells, horizontal cells, and cones are the first cells in the retina to be born
733(2)
As distance from the fovea increases, the firstborn cells appear at progressively later dates
735(1)
Synapse formation has a center-to-periphery gradient superimposed on a gradient from inner retina to outer retina
736(1)
The location of the future fovea is specified very early; the pit is created by cell migration
737(1)
Foveal cones are incomplete at birth
737(2)
Photoreceptor densities are shaped by cell migration and retinal expansion
739(1)
Ganglion cell density is shaped by migration, retinal expansion, and cell death
740(1)
The spatial organization of the retina may depend on specific cell-cell interactions and modifications of cell morphology during development
741(2)
Retinal blood vessels develop relatively late
743(1)
Developing vessels are inhibited by too much oxygen
744(1)
The optic nerve forms as tissue in the optic stalk is replaced with developing ganglion cell axons and glial cells
745(1)
Fusion of the optic stalks produces the optic chiasm, where pioneering axons must choose the ipsilateral or contralateral path
746(1)
The last stages of development in the optic nerve are axon loss and myelination
746(2)
The inner retina seems relatively immune to congenital anomalies
748(1)
The most common developmental anomalies are failures to complete embryonic structures or eliminate transient structures
748
Box 16.1 Fluorescein Angiography and the Adequacy of Circulation
710(43)
Epilogue Time and Change
753
Postnatal Growth and Development
753(6)
The newborn eye increases in overall size for the next 15 years
753(1)
Refractive error is quite variable among newborn infants, but the variation decreases with growth
754(2)
Visual functions mature at different rates during the first 6 years of life
756(2)
Changes in the lens and vitreous that begin in infancy continue throughout life
758(1)
Maturation and Senescence
759
The average refractive error is stable from ages 20 to 50, but the eye becomes more hyperopic and then more myopic later in life
759(1)
Although the gross structure of the eye is stable after the age of 20, tissues and membranes are constantly changing
760(1)
Retinal illuminance and visual sensitivity decrease with age
761(2)
Visual acuity declines after age 50, largely because of optical factors
763
Historical References and Additional Reading 1(1)
Glossary 1(1)
Index 1

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