Preface |
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xxiii | |
Acknowledgments |
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xxviii | |
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Prologue A Brief History of Eyes |
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1 | (752) |
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The Antiquity of Eyes and Vision |
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1 | (8) |
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Thinking about the eye gave Charles Darwin ``a cold shudder'' |
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1 | (1) |
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The history of the eye is embedded in the history of animals and molecules |
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2 | (1) |
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Several of the eye's critical molecules are ancient |
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3 | (2) |
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Eyes were invented by multicellular animals almost 600 million years ago |
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5 | (1) |
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Eyes arose not once, but numerous times, in different animal groups |
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6 | (2) |
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Eyes are most common in groups of motile animals living in lighted environments |
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8 | (1) |
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The Diversity and Distribution of Eyes |
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9 | (7) |
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The first step in vision is an eye that can sense the direction of incident light |
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9 | (2) |
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At least ten types of eyes can be distinguished by differences in their optical systems |
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11 | (2) |
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Vertebrates always have simple eyes, but invertebrates can have compound eyes, simple eyes, or both |
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13 | (3) |
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Paths and Obstacles to Perfection |
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16 | (5) |
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Simple eyes improve as they become larger |
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16 | (2) |
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Elaborate simple eyes may have evolved rapidly |
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18 | (1) |
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Compound eyes have inherent optical limitations in their performance |
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19 | (2) |
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An Ocular Bestiary: Fourteen Eyes and Their Animals |
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21 | (36) |
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Compound eye---focal apposition, terrestrial variety: Honeybee (Apis mellifica) |
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21 | (2) |
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Compound eye---focal apposition, aquatic variety: Horseshoe crab (Limulus polyphemus) |
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23 | (3) |
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Compound eye---afocal apposition: Monarch butterfly (Danaus plexippus) |
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26 | (3) |
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Compound eye---apposition with neural superposition: Housefly (Musca domestica) |
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29 | (2) |
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Compound eye---refracting superposition: Firefly (Photuris spp.) |
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31 | (2) |
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Compound eye---reflecting superposition: Crayfish |
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33 | (3) |
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Compound eye---parabolic superposition: Crabs |
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36 | (1) |
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Simple eye---pinhole: Nautilus |
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37 | (1) |
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Simple eye---refracting, aquatic variety: Octopus |
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38 | (2) |
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Simple eye---refracting, aquatic variety: Goldfish |
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40 | (3) |
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Simple eye---refracting, terrestrial variety: Pigeon (Columba livia) |
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43 | (2) |
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Simple eye---refracting, terrestrial variety: Jumping spiders (Metaphidippus spp.) |
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45 | (3) |
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Simple eye---reflecting: Scallops |
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48 | (2) |
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Simple eye---refracting, terrestrial variety: Humans (Homo sapiens) |
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50 | (7) |
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Formation of the Human Eye |
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57 | (20) |
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Some Developmental Strategies and Operations |
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57 | (2) |
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Embryogenesis begins with cell proliferation, cell movement, and changes in cell shape |
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57 | (1) |
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Specialized tissues are formed by collections of cells that have become specialized themselves |
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58 | (1) |
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Proliferation, movement, and differentiation in a cell group may require communication with other cells |
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58 | (1) |
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Embryonic Events before the Eyes Appear |
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59 | (5) |
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The blastocyst forms during the first week of embryogenesis |
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59 | (1) |
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The inner cell mass becomes the gastrula, which is divided into different germinal tissues |
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60 | (2) |
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Neurulation begins the development of the nervous system |
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62 | (2) |
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Formation of the Primitive Eye |
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64 | (1) |
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Ocular development begins in the primitive forebrain |
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64 | (1) |
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The optic vesicle induces formation of the lens |
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64 | (1) |
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Elaboration of the Primitive Eye |
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65 | (6) |
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The optic cup and the lens form from different germinal tissues by changes in cell shape |
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65 | (2) |
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The optic cup is initially asymmetric, with a deep groove on its inferior surface |
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67 | (1) |
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Closure of the choroidal fissure completes the optic cup |
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67 | (1) |
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The lens vesicle forms in synchrony with the optic cup |
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67 | (1) |
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The primitive lens is the first ocular structure to exhibit cell differentiation |
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68 | (1) |
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All future growth of the lens comes from the early lens cells, some of which are ``immortal'' stem cells |
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69 | (1) |
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The precursors of the future retina, optic nerve, lens, and cornea are present by the sixth week of gestation |
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69 | (1) |
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In general, the eye develops from inside to outside |
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70 | (1) |
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Failures of Early Development |
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71 | (1) |
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``If anything can go wrong, it will'' |
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71 | (1) |
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One or both eyes may fail to develop completely |
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71 | (1) |
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Congenital absence of the lens may be an early developmental failure |
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71 | (3) |
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Incomplete closure of the choroidal fissure can produce segmental defects in the adult eye |
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74 | |
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Vignette 1.1 The Eye of Mann |
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72 | (5) |
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Ocular Geometry and Topography |
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77 | (34) |
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Elements of Ocular Structure |
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77 | (5) |
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The human eye is a simple eye |
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77 | (1) |
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The outermost of the three coats of the eye consists of cornea, limbus, and sclera |
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78 | (1) |
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The middle coat---the uveal tract---includes the iris, ciliary body, and choroid |
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78 | (1) |
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The eye's innermost coat---the retina---communicates with the brain via the optic nerve |
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79 | (2) |
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Most of the volume of the eye is fluid or gel |
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81 | (1) |
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Image Quality and Visual Performance |
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82 | (8) |
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Images of point sources are always small discs of light whose size is a measure of optical quality |
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82 | (1) |
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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 |
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83 | (3) |
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The contrast sensitivity function specifies how well different spatial frequencies are seen by the visual system |
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86 | (1) |
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We can see flies when their images subtend about one minute of visual angle |
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87 | (3) |
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The Anatomy of Image Formation |
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90 | (7) |
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The quality of a focused image is affected by pupil size, curvatures of optical surfaces, and homogeneity of the optical media |
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90 | (2) |
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Defocusing produces large changes in the modulation transfer function |
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92 | (1) |
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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 |
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93 | (2) |
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Schematic eyes are approximations of the eye's optically relevant anatomy |
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95 | (2) |
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97 | (6) |
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Vertebrate eyes vary considerably in shape |
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97 | (1) |
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Both the cornea and the sclera are aspheric |
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97 | (3) |
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On average, the adult human eye measures twenty-four millimeters in all dimensions |
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100 | (1) |
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Axial lengths and other anatomical features vary among individuals, but most eyes are emmetropic |
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101 | (1) |
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Most refractive error is related to relatively large or small axial lengths |
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102 | (1) |
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The Eye's Axes and Planes of Reference |
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103 | |
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The eyes rotate around nearly fixed points |
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103 | (1) |
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Like the head, the eyes have three sets of orthogonal reference planes |
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104 | (2) |
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The pupillary axis is a measure of the eye's optical axis |
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106 | (1) |
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The line of sight differs from the pupillary axis by the angle kappa |
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107 | (1) |
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The angles kappa in the two eyes should have the same magnitude |
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107 | (1) |
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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 |
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108 | |
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Vignette 2.1 The Medieval Eye |
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80 | (8) |
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Box 2.1 Evaluating Visual Resolution |
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88 | (10) |
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Vignette 2.2 Fundamentum Opticum |
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98 | (13) |
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111 | (22) |
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111 | (10) |
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The orbits are roughly pyramidal |
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111 | (1) |
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The large bones of the face form the orbital margin and much of the orbit's roof, floor, and lateral wall |
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112 | (2) |
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The sphenoid bone fills the apex of the orbital pyramid and contributes to the lateral and medial walls |
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114 | (4) |
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The lacrimal and ethmoid bones complete the medial wall between the maxillary bone in front and the sphenoid in back |
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118 | (1) |
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Three major and several minor foramina permit blood vessels and nerves to enter or exit the orbital cavity |
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118 | (1) |
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Blowout fractures of the orbit are a consequence of the relative weakness of the orbital plates |
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119 | (1) |
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Abnormal positioning of the eye relative to the orbital margin may indicate local or systemic pathology |
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120 | (1) |
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Infection and tumors can enter the orbital cavity through the large sinuses that surround the orbit |
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120 | (1) |
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Connections between the Eye and the Orbit |
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121 | (6) |
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Connective tissue lines the interior surface of the orbit |
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121 | (1) |
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Connective tissue surrounds the eye and extraocular muscles |
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121 | (3) |
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Check ligaments connect Tenon's capsule to the periorbita |
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124 | (1) |
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All structures in the orbital cavity are lined and interconnected with connective tissue |
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125 | (1) |
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Abnormal development of the connective tissue may affect movement of the eyes |
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125 | (1) |
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Fat fills the spaces in the orbital cavity that are not occupied by other structures |
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126 | (1) |
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The septum orbitale prevents herniation of orbital fat into the eyelids |
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127 | (1) |
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Development of the Orbital Bones |
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127 | |
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Many bones form first as cartilage templates |
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127 | (1) |
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Most orbital bones do not have cartilaginous templates |
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127 | (3) |
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The orbital plates begin to form during the sixth week of gestation |
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130 | (1) |
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The capacity of bone for growth, repair, and remodeling lasts many years |
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130 | (1) |
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The eyes and orbits rotate from lateral to frontal positions during development |
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130 | (1) |
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Most developmental anomalies of the orbital bones are associated with anomalies of the facial bones |
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131 | |
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Vignette 3.1 Sovereign of the Visible World |
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116 | (6) |
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Box 3.1 Visualizing the Orbit and Its Contents In Vivo |
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122 | (6) |
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Vignette 3.2 The Anatomy of Vesalius |
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128 | (5) |
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133 | (58) |
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133 | (6) |
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Our eyes are always moving, and some motion is necessary for vision |
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133 | (2) |
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Large, rapid eye movements are used for looking around, for placing retinal images of interest on the fovea |
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135 | (1) |
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Slow eye movements are used to track or follow movement and to compensate for changes in head and body position |
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136 | (1) |
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Eye movement velocities may vary by a factor of 105 |
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137 | (1) |
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Since the eyes have overlapping fields of view, their movements must be coordinated |
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137 | (1) |
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Slow movements of the eyes in opposite directions help keep corresponding images on the foveas in both retinas simultaneously |
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138 | (1) |
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Strabismus is a misalignment of the two visual axes under binocular viewing conditions |
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139 | (1) |
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139 | (14) |
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The extraocular muscles are arranged as three reciprocally innervated agonist-antagonist pairs |
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139 | (2) |
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The eyes are stationary when the opposing forces exerted by the extraocular muscles are in balance |
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141 | (1) |
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Imbalanced forces produce eye rotations |
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142 | (1) |
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Muscle force is related to muscle length |
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143 | (1) |
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Most of the force that maintains eye position is passive force |
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144 | (1) |
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Equilibrium muscle lengths and forces for different gaze positions are functions of the innervational command |
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145 | (1) |
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Different patterns of innervation are required for fast and slow eye movements |
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146 | (1) |
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Extraocular motor neurons are located in three interconnected nuclei in the brainstem |
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146 | (1) |
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Motor commands are the result of interactions between visual and nonvisual inputs to the motor control centers |
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147 | (1) |
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A copy of the innervational command is used to verify the system's operation |
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148 | (1) |
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Extraocular motor neurons receive inputs from premotor areas in the brainstem to generate appropriate signals for saccadic eye movements |
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148 | (1) |
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The pathways for smooth pursuit movements and for vergences go through the cerebellum, but vergences have a separate control center near the oculomotor nucleus |
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149 | (4) |
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Extraocular Muscle Structure and Contractile Properties |
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153 | (12) |
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Muscle fibers are the units from which muscles are constructed |
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153 | (1) |
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Striated muscle fibers have a parallel arrangement of contractile proteins that interleave to cause contraction |
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154 | (1) |
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Striated muscle fibers differ in structural, histochemical, and contractile properties |
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155 | (1) |
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The extraocular muscles contain muscle fiber types not found in skeletal muscles |
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156 | (1) |
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Thick and thin extraocular muscle fibers differ in their contractile properties |
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156 | (1) |
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Different muscle fiber types are not randomly distributed within the muscles |
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157 | (1) |
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Different muscle fiber types may receive different innervational commands |
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158 | (1) |
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Extraocular muscles have very small motor units |
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159 | (2) |
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Acetylcholine at the neuromuscular junctions depolarizes the cell membrane by opening sodium channels |
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161 | (1) |
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The spread of depolarization along the sarcolemma may differ among muscle fiber types, producing different contractile properties |
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161 | (1) |
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Extraocular muscles exhibit high sensitivity to agents that mimic or block the action of acetylcholine |
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162 | (1) |
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Extraocular muscles often exhibit early symptoms of myasthenia gravis |
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163 | (1) |
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Neurotoxins that interfere with acetylcholine action can be used to alleviate strabismus and blepharospasm |
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163 | (2) |
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Sensory Endings in Extraocular Muscles and Tendons |
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165 | (3) |
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Skeletal muscles have two major types of sensory organs |
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165 | (1) |
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Human extraocular muscles have anatomically degenerate sensory organs and exhibit no stretch reflexes |
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166 | (1) |
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Passive extraocular muscle stretch may produce bradycardia |
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167 | (1) |
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Sensory endings in extraocular muscles probably do not convey information about eye position |
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167 | (1) |
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Sensory signals from the extraocular muscles may be involved in motor learning, motor plasticity, and development |
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168 | (1) |
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Actions of the Extraocular Muscles |
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168 | (15) |
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All of the extraocular muscles except the inferior oblique have their anatomical origins at the apex of the orbit |
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168 | (1) |
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The anatomical origin of the inferior oblique and the functional origin of the superior oblique are anterior and medial in the orbit |
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169 | (1) |
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The four rectus muscles are arranged as horizontal and vertical pairs, all inserting onto the anterior portion of the globe |
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170 | (1) |
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The horizontal recti rotate the eye in the horizontal plane around a vertical axis |
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170 | (1) |
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The vertical recti are responsible for upward and downward rotations of the eye |
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171 | (1) |
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The recti define a muscle cone within the orbital cavity that contains most of the ocular blood vessels and nerves |
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172 | (1) |
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The oblique muscles constitute a third functional pair, inserting onto the posterior portion of the eye |
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172 | (2) |
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Extraocular muscle actions cannot be measured directly |
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174 | (1) |
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The classic description of action of the extraocular muscles is based on the geometry of their origins and insertions |
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174 | (2) |
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Boeder diagrams attempt to describe the actions of the extraocular muscles completely |
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176 | (2) |
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The presence of Tenon's capsule and muscle pulleys invalidates the geometric model of extraocular muscle actions |
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178 | (2) |
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``There is no simple way to describe the action of these muscles on the eye!'' |
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180 | (2) |
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A realistic model of the extraocular muscle system is important for the diagnosis and treatment of muscle paresis |
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182 | (1) |
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Development of the Extraocular Muscles |
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183 | |
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Each muscle develops from several foci in the mesoderm surrounding the optic cup |
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183 | (2) |
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The extraocular muscles appear after the optic cup, but before the orbital bones |
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185 | (1) |
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Different muscle fiber types form late in gestation and continue to develop postnatally |
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185 | (1) |
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Most developmental anomalies are associated with the connective tissue of the muscles or with their innervation |
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186 | (1) |
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The oculomotor system is not fully operational at birth |
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186 | |
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Box 4.1 Detecting Ocular Misalignment |
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140 | (12) |
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Box 4.2 Changing the Effects of Extraocular Muscle Contraction |
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152 | (12) |
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Vignette 4.1 Locating the Extraocular Muscles |
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164 | (20) |
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Vignette 4.2 In the Service of the Eye |
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184 | (7) |
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The Nerves of the Eye and Orbit |
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191 | (56) |
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Elements of Neural Organization |
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191 | (4) |
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The brain deals with information about the external world and the body |
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191 | (1) |
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Neurons are the anatomical elements of neural systems |
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191 | (2) |
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Neural circuits consist of neurons linked mostly by unidirectional chemical synapses |
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193 | (1) |
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The direction of neural information flow distinguishes between sensory and motor nerves |
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194 | (1) |
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Motor outputs are divided anatomically and functionally into somatic and autonomic systems |
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194 | (1) |
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The autonomic system is subdivided into the sympathetic and parasympathetic systems |
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194 | (1) |
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The Optic Nerve and the Flow of Visual Information |
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195 | (17) |
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In the optic nerve, the location of axons from retinal ganglion cells corresponds to their location on the retina |
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195 | (3) |
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Axons from the two optic nerves are redistributed in the optic chiasm |
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198 | (1) |
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The decussation of axons in the chiasm is imperfect |
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199 | (1) |
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Spatial ordering of axons changes in the optic tracts |
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200 | (1) |
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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 |
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200 | (1) |
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Axons terminating in the lateral geniculate nuclei are spatially ordered |
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201 | (2) |
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Some axons leave the optic tracts for other destinations |
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203 | (1) |
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Axons terminating in the superior colliculi form discontinuous retinotopic maps |
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204 | (1) |
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Axons forming the afferent part of the pupillary light reflex pathway terminate in the pretectal nuclear complex |
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204 | (1) |
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Retinal inputs to the accessory optic system may help coordinate eye and head movement |
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205 | (1) |
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Retinal axons may provide inputs to a biological clock |
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205 | (2) |
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Lesions of the optic nerves and tracts produce defects in the visual fields |
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207 | (5) |
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Lesions in the secondary visual pathways can be observed only as motor deficits |
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212 | (1) |
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The Trigeminal Nerve: Signals for Touch and Pain |
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212 | (7) |
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Two of the three trigeminal divisions carry signals from the eye and surrounding tissues |
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212 | (1) |
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All somatosensory information from the eye is conveyed by the nasociliary nerve to the ophthalmic division of the trigeminal |
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213 | (1) |
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Sensory nerve fibers from the cornea, conjunctiva, limbus, and anterior sclera join to form the long ciliary nerves |
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213 | (2) |
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Stimulation of corneal or conjunctival nerve endings elicits sensations of touch or pain, a blink reflex, and reflex lacrimation |
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215 | (1) |
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Other sensory fibers from the eye are conveyed by the short ciliary nerves and the sensory root of the ciliary ganglion |
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215 | (1) |
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Most other branches of the ophthalmic nerve carry somatosensory fibers from the skin of the eyelids and face |
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215 | (2) |
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A few branches of the maxillary nerve pass through the orbit from the facial skin and the maxillary sinus |
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217 | (1) |
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Lesions in the branching hierarchy of the ophthalmic nerve produce anesthesia that helps identify the lesion site |
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218 | (1) |
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Viral infection of the trigeminal system can produce severe corneal damage |
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219 | (1) |
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The Extraocular Motor Nerves |
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219 | (7) |
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The three cranial nerves that innervate the extraocular muscles contain axons from clusters of cells in the brainstem |
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219 | (1) |
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Cells in different parts of the oculomotor nerve nucleus innervate the levator, the superior and inferior recti, the medial rectus, and the inferior oblique |
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219 | (2) |
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Axons destined for different muscles run together in the oculomotor nerve until it exits the cavernous sinus just behind the orbit |
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221 | (3) |
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The oculomotor nerve contains parasympathetic fibers bound for the ciliary ganglion |
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224 | (1) |
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Cells in the trochlear nerve nucleus innervate the contralateral superior oblique |
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224 | (1) |
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Abducens nerve cells innervate the ipsilateral lateral rectus |
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225 | (1) |
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All of the oculomotor nerves pass through the cavernous sinus on their way to the orbit |
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225 | (1) |
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The extraocular motor nerves probably contain sensory axons from muscle spindles and tendon organs |
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226 | (1) |
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Innervation of the Muscles of the Eyelids |
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226 | (2) |
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Three sets of muscles are associated with the eyelids |
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226 | (1) |
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The orbicularis is innervated by the facial nerve |
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227 | (1) |
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The superior and inferior tarsal muscles are innervated by the sympathetic system |
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227 | (1) |
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Ptosis may result from either oculomotor or sympathetic lesions |
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228 | (1) |
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Autonomic Innervation of Smooth Muscle within the Eye |
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228 | (7) |
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The superior cervical ganglion is the source of most sympathetic innervation to the eye |
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228 | (2) |
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Sympathetic fibers enter the eye in the short ciliary nerves |
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230 | (1) |
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Sympathetic innervation of the dilator muscle acts at alpha-adrenergic receptors to dilate the pupils |
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230 | (1) |
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The arterioles in the uveal tract receive sympathetic innervation that produces vasoconstriction |
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231 | (1) |
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Horner's syndrome is the result of a central lesion in the sympathetic pathway |
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231 | (1) |
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Parasympathetic fibers entering the eye originate in the ciliary or the pterygopalatine ganglion |
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231 | (1) |
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Axons from cells in the ciliary ganglion innervate the sphincter and the ciliary muscle |
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232 | (1) |
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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) |
|
|
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) |
|
|
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) |
|
|
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) |
|
|
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) |
|
|
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 | |
|
|
470 | (21) |
|
The Lens and the Vitreous |
|
|
491 | (54) |
|
|
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) |
|
|
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) |
|
|
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) |
|
|
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) |
|
|
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 | |