Get Slatter s Fundamentals of Veterinary Ophthalmology Fourth Edition David Maggs Bvsc(Hons) Davco PDF ebook with Full Chapters Now

Download the full version of the ebook at
https://ebookultra.com
Slatter s Fundamentals of Veterinary

Ophthalmology Fourth Edition David Maggs
Bvsc(Hons) Davco
https://ebookultra.com/download/slatter-s-
fundamentals-of-veterinary-ophthalmology-fourth-
edition-david-maggs-bvschons-davco/
Explore and download more ebook at https://ebookultra.com

Recommended digital products (PDF, EPUB, MOBI) that
you can download immediately if you are interested.
Notes on Veterinary Ophthalmology 1st Edition Sheila M.

Crispin
https://ebookultra.com/download/notes-on-veterinary-ophthalmology-1st-
edition-sheila-m-crispin/
ebookultra.com
Fundamentals of Matrix Computations 2nd Edition David S.

Watkins
https://ebookultra.com/download/fundamentals-of-matrix-
computations-2nd-edition-david-s-watkins-2/
ebookultra.com
Fundamentals of Matrix Computations 2nd Edition David S.

Watkins
https://ebookultra.com/download/fundamentals-of-matrix-
computations-2nd-edition-david-s-watkins/
ebookultra.com
Biological Performance of Materials Fundamentals of

Biocompatibility Fourth Edition Black
https://ebookultra.com/download/biological-performance-of-materials-
fundamentals-of-biocompatibility-fourth-edition-black/
ebookultra.com
Veterinary Clinical Epidemiology From Patient to
Population Fourth Edition Smith
https://ebookultra.com/download/veterinary-clinical-epidemiology-from-
patient-to-population-fourth-edition-smith/
ebookultra.com
Lishman s Organic Psychiatry A Textbook of Neuropsychiatry

Fourth Edition Anthony David
https://ebookultra.com/download/lishman-s-organic-psychiatry-a-
textbook-of-neuropsychiatry-fourth-edition-anthony-david/
ebookultra.com
Primary Care Ophthalmology Textbook 2nd Edition David

Palay
https://ebookultra.com/download/primary-care-ophthalmology-
textbook-2nd-edition-david-palay/
ebookultra.com
Pediatric Ophthalmology and Strabismus 3rd Edition David

Taylor
https://ebookultra.com/download/pediatric-ophthalmology-and-
strabismus-3rd-edition-david-taylor/
ebookultra.com
BSAVA Manual of Canine and Feline Ophthalmology 3rd

Edition David Gould
https://ebookultra.com/download/bsava-manual-of-canine-and-feline-
ophthalmology-3rd-edition-david-gould/
ebookultra.com
Slatter s Fundamentals of Veterinary Ophthalmology
Fourth Edition David Maggs Bvsc(Hons) Davco Digital
Instant Download
Author(s): David Maggs BVSc(Hons) DAVCO, Paul Miller DVM DACVO,
Ron Ofri DVM PhD DECVO
ISBN(s): 0721605613
Edition: 4
File Details: PDF, 45.04 MB
Year: 2007
Language: english
11830 Westline Industrial Drive
St. Louis, Missouri 63146
SLATTER’S FUNDAMENTALS OF VETERINARY OPHTHALMOLOGY,

EDITION 4 ISBN: 978-0-7216-0561-6
Copyright © 2008, 2001, 1990, 1981 by Saunders, an imprint of Elsevier Inc.
All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any
means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval
system, without permission in writing from the publisher.
Permissions may be sought directly from Elsevier’s Health Sciences Rights Department in Philadelphia, PA,
USA: phone: (+1) 215 239 3804, fax: (+1) 215 239 3805, e-mail: healthpermissions@elsevier.com. You may
also complete your request on-line via the Elsevier homepage (http://www.elsevier.com), by selecting
‘Customer Support’ and then ‘Obtaining Permissions’.
Notice
Knowledge and best practice in this field are constantly changing. As new research and experience broaden
our knowledge, changes in practice, treatment and drug therapy may become necessary or appropriate.
Readers are advised to check the most current information provided (i) on procedures featured or (ii) by the
manufacturer of each product to be administered, to verify the recommended dose or formula, the method
and duration of administration, and contraindications. It is the responsibility of the practitioner, relying on
his or her own experience and knowledge of the patient, to make diagnoses, to determine dosages and the
best treatment for each individual patient, and to take all appropriate safety precautions. To the fullest
extent of the law, neither the Publisher nor the Authors assume any liability for any injury and/or damage
to persons or property arising out or related to any use of the material contained in this book.
The Publisher
Library of Congress Control Number 2007924244
Publishing Director: Linda Duncan

Acquisitions Editor: Anthony Winkel
Developmental Editor: Maureen Slaten
Publishing Services Manager: Patricia Tannian
Senior Project Manager: Kristine Feeherty
Design Direction: Amy Buxton
Printed in China
Last digit is the print number: 9 8 7 6 5 4 3 2 1

CONTRIBUTORS
Itamar Aroch, DVM, DECVIM-CA Gila A. Sutton, DVM, MSc, DACVIM-LA

Senior Lecturer, Small Animal Medicine Clinical Instructor, Equine Internal Medicine
Koret School of Veterinary Medicine Koret School of Veterinary Medicine
Hebrew University of Jerusalem Hebrew University of Jerusalem
Rehovot, Israel Rehovot, Israel
Bradford J. Holmberg, DVM, MS, PhD, DACVO Brian P. Wilcock, DVM, MS, PhD, DACVP
Veterinary Referral Centre Department of Pathobiology
Little Falls, New Jersey Ontario Veterinary College
University of Guelph
Ontario, Canada
v
PREFACE
In September 2005, the veterinary community lost one of the updating them where necessary. Doug was particularly excited
pioneers of veterinary ophthalmology, a gifted surgeon, an about the introduction of color images in the fourth edition of
intuitive scientist, and a prodigious and highly respected this text, and we are delighted that Elsevier has permitted us to
author, when Dr. Douglas H. Slatter died in Australia—his honor, and even expand upon, that wish; other than a few black-
country of birth. Before his death, Doug had asked each of us and-white figures that we have preserved for their historical
to work alongside him as he updated Fundamentals of value or because we considered them irreplaceable, every
Veterinary Ophthalmology for its fourth edition. He explained photograph is now in color.
that it was his goal to diversify the opinions and knowledge As a preface to the first edition of Fundamentals of
contained in the book while avoiding the creation of a large Veterinary Ophthalmology in 1981, Doug wrote, “The aim of
multi-author text. Without hesitation, the three of us joined this text is to provide a basic outline and introduction to clinical
together to continue this effort as a legacy to Doug following ophthalmology for both undergraduate veterinary students and
his death. practitioners.” In that regard, nothing has changed. We hope,
We believe that earlier editions of Dr. Slatter’s book have with this fourth edition of Slatter’s Fundamentals of Veterinary
stood apart from other ophthalmology texts because of the Ophthalmology, that we have built upon the astounding foundation
breadth of species covered; the logical, tissue-based format; Doug created, and we dedicate this book to our friend and
and the precise and detailed original figures. It has been our colleague—Dr. Douglas H. Slatter.
goal to preserve and expand these features while trying to also
bring to the fourth edition something from the diversity of each ACKNOWLEDGMENTS
of our backgrounds and areas of ophthalmic interest, along
with the geographic regions in which we trained and now prac- A project of this magnitude depends on the teamwork and
tice. We also recruited four of the world’s experts in their field assistance of a large number of people—more than we can do
to provide chapters on ophthalmology of exotic pets, ocular justice to or name here. However, some deserve special
manifestations of systemic disease, ocular embryology, and mention. We are indebted to the production crew at Elsevier
ocular pathology, and we are indebted to these authors for their and in particular to Dr. Tony Winkel, our Editor, who oversaw
superb contributions. the delicate transfer of Doug’s text to our care with great
Throughout these updates, we have aimed to retain the crisp diplomacy and kindness; Maureen Slaten, who managed the
and logical organization that Doug began more than 20 years major artwork revisions; and Kristine Feeherty, who oversaw
ago. We begin with chapters on ocular structure and function, the timely production of the text with great maturity and good
congenital anomalies, and the recent pharmacologic advances humor. We also owe heartfelt thanks to Dr. Dan Lavach for his
and finish with a section on ocular emergencies, an appendix sensitive reminiscences about Doug’s life and his contributions
containing breed predispositions to ocular disorders, and a in the dedication of this book. We would also like to thank Dr.
glossary to help readers negotiate the sometimes daunting Robert Barishak for his insightful review of the chapter on
world of ophthalmic terminology. The chapters in between development and congenital abnormalities. Our invited
remain largely tissue-based—“from eyelids to orbit”—and authors—Drs. Itamar Aroch, Bradford Holmberg, Gila Sutton,
within each chapter we have retained a brief description of the and Brian Wilcock—provided their expertise without hesitation,
clinically relevant anatomy, physiology, and pathophysiology, and the text is greatly enhanced by their contributions. In
followed by extensively updated sections on newly recognized addition, much of the work we updated or edited was originally
diseases, diagnostic tests, the latest drugs, and innovative provided by contributors selected by Dr. Slatter, and their
surgical techniques, along with current reading lists. Care has efforts remain an integral part of the current work. Many of the
been taken to ensure that therapies are recommended only figures in the fourth edition were redrawn or extensively
when there is sufficient scientific evidence to support them. All modified by John Doval. His extraordinary skill and patience
the while, we have kept the busy general practitioner and are acknowledged with much gratitude, as is the colorization of
enquiring veterinary student in mind and made extensive use of the figures by Jeanne Robertson. Finally, we thank our families
tables, boxes, and figures. and colleagues who have, in so many ways, written this text
As befitting someone dual-boarded in surgery and ophthal- with us.
mology, Doug created a text known for its exquisite surgical
descriptions and drawings. We have greatly enhanced these David J. Maggs Paul E. Miller Ron Ofri
throughout the book with the introduction of color and by Davis, California Madison, Wisconsin Rehovot, Israel
vii
TRIBUTE
Dr. Douglas H. Slatter was an In 1984, Drs. Slatter and Chambers returned to the United
important and unique individual. As a States. Dr. Slatter had been a visiting Professor of Ophthal-
student he excelled, graduating from mology at the Scheie Eye Institute, University of Pennsylvania.
the veterinary school at the University Drs. Slatter and Chambers eventually acquired the Animal Eye
of Queensland in 1970. He was Clinic in La Habra, California, from retiring Dr. Ralph
immediately accepted into a graduate Vierheller, DAVCO, DACVS. Dr. Slatter and Dr. Vierheller had
program at Washington State the distinction of being board certified in both ophthalmology
University and completed a Master of and surgery. Dr. Slatter continued with academic pursuits while
Science degree. During this time Dr. in private practice. He was an adjunct professor of veterinary
Slatter learned considerable clinical ophthalmology at the Southern California College of Optometry
ophthalmology under the tutelage of in Fullerton, California. Drs. Slatter and Chambers practiced in
Dr. Gary Bryan, DACVO. Next, Dr. La Habra, California, for the remainder of their lives. In
Slatter attended Colorado State University (CSU) in a addition, they operated ophthalmology clinics throughout the
Doctorate of Philosophy Degree program at the Surgical Los Angeles basin; in Bakersfield, California; and in Incline
Metabolic Laboratory directed by Dr. William Lumb. Dr. Village, Nevada. Despite his busy practice schedule, Dr. Slatter
Slatter excelled in this diverse academic environment. He had traveled throughout the world lecturing about ophthalmology
special interests in cardiac surgery and ophthalmology. While and surgery. He made it a priority to vacation in his homeland
at CSU, he spent time with Dr. Glenn Severin, DACVO, of Australia at least twice each year. During these working
DACVIM, furthering Dr. Slatter’s clinical knowledge of vacations, he managed to operate a land development company
veterinary ophthalmology. and a small airplane import business. His passion and skill as a
An important friend at CSU was Mrs. Mary Fischer, BS, private pilot were well respected in both Australia and the
MS. Mrs. Fischer had been the chief laboratory technician in United States. He was also a licensed Justice of the Peace in
the ophthalmology section of the Armed Forces Institute of Australia.
Pathology. She had been hired in the CSU Department of Throughout his years in private practice, Dr. Slatter
Pathology to bring her expertise to the Specialty Pathology continued to pursue academic excellence. He published
Laboratory directed by Dr. Stuart Young, DACVP, Hon revisions to his original ophthalmology text, and he organized
DACVO. Dr. Slatter worked diligently with Dr. Young and and edited the two-volume Textbook of Veterinary Surgery and
Mrs. Fischer in developing retinal digesting techniques to the pocket companion to that book. Each of his textbooks and
advance the study of retinal vascular disease in dogs. His their revisions were translated into several foreign languages.
doctoral thesis, “Effects of Hyperproteinemia and Aging on Dr. Slatter also contributed more than 60 peer-reviewed scientific
Canine Eyes” (1975), was the culmination of this collaboration. manuscripts during his career.
Dr. Slatter was a perfect student. He accomplished an In addition to being a Diplomate in both the American
amazing feat by sitting for his board certification in veterinary Colleges of Veterinary Ophthalmology and Surgery, Dr. Slatter
surgery and veterinary ophthalmology in the same year, was a Diplomate of European College of Veterinary Surgeons,
passing both examinations with high marks. Furthermore, a Fellow of the Royal College of Veterinary Surgeons, a
Dr. Slatter completed his doctoral research project and earned Member of the Australian College of Veterinary Scientists, and
his PhD degree in the same year. During this busy year he one of four founding members of the International Society of
completed rough-draft preparations for the soon-to-be-published Veterinary Ophthalmology.
first edition of this textbook: Fundamentals of Veterinary Dr. Douglas H. Slatter and Dr. Elizabeth D. Chambers were
Ophthalmology. popular speakers, educators, and clinicians who devoted their
After graduate school, Dr. Slatter returned to Australia and lives to improving veterinary ophthalmology and veterinary
taught ophthalmology and small animal surgery at Murdoch surgery. They will be missed.
University, Western Australia. He married Dr. Elizabeth
Chambers, and Dr. Chambers proved to be his soul mate, best J. Daniel Lavach, DVM, DACVO
friend, and fellow collaborator throughout the remainder of Reno, Nevada
their lives.
ix
Chapter
STRUCTURE AND FUNCTION 1

OF THE EYE
Paul E. Miller
FUNDAMENTALS OF VISION VASCULAR ANATOMY AND PERIPHERAL OCULAR REFLEXES

CENTRAL VISUAL PATHWAYS NEUROANATOMY PHYSIOLOGY OF THE AQUEOUS
V ision is a complex phenomenon in which light emanating

from objects in the environment is captured by the eye and
mammals, cats are probably the most efficiently adapted for
nocturnal vision, with a minimum light detection threshold up
to seven times lower than that in humans. Other adaptations
focused onto the retinal photoreceptors (Figures 1-1 and 1-2). that permit cats to function well in nocturnal conditions are a
Electrical signals originating from these cells pass through a tapetum lucidum, which reflects 130 times more light than the
number of cell types in the retina and throughout the central human fundus; a vertical slit pupil, which produces a smaller
nervous system (CNS) before arriving at the visual cortex, aperture in bright light than what is possible with a circular
where the sensation of vision occurs. Numerous species vari- pupil but also allows the pupil to dilate 6 mm more than
ations exist on this basic theme, each allowing the animal to the human pupil; a large cornea, which permits more light to
exploit a particular ecologic niche. The basic similarities enter the eye; a relatively posteriorly located lens, which
among all vertebrate eyes and how they respond to insult allow produces a smaller but brighter image on the fundus; and a
the comparative ophthalmologist to confidently treat a wide retina rich in light-sensitive rod photoreceptors (Figure 1-3).
range of ocular conditions in a diverse array of species. Many of the other domestic mammals have similar but fewer
extreme adaptations for vision in dim light, allowing them to
FUNDAMENTALS OF VISION exploit a photic environment that is not strictly diurnal or
nocturnal.
The act of “seeing” is a complex process that depends on (1) The tapetum is cellular in dogs and cats and collagenous in
light from the outside world falling onto the eye, (2) the eye horses and ruminants, suggesting that the visual advantages this
efficiently transmitting and properly focusing the images of structure offers are of sufficient magnitude that it has evolved
these objects on the retina, (3) the retina detecting these light separately at least twice in mammals (Figure 1-4). In both
rays, (4) transmission of this information via the visual path- cases, the variety of tapetal colors seen during ophthalmoscopy
ways to the brain, and (5) the brain processing this information results from the differential interaction of light with the
so as to make it useful. Differentiating between objects (e.g., a tapetum’s physical structure rather than from the inherent
predator versus its surroundings) is one of the most critical spectral composition, or color, of its pigments. The dorsal
aspects of vision, and because this distinction is so important location of the tapetum may enhance the view of the usually
for survival, normal animals can “see” an object if it differs darker ground, and the ventrally located, usually darkly pig-
sufficiently from its surroundings in any one of five different mented nontapetal region may reduce light scattering originating
aspects: luminance (“brightness”), motion, texture, binocular from the brighter sky. In cats, the tapetum may also absorb light
disparity (depth), and color. In general, objects are differ- in the shorter wavelengths and, via fluorescence, shift it to a
entiated on the basis of their motion, texture, depth, and longer wavelength that more closely approximates the maximal
luminance roughly equally well, but separations based on color sensitivity of the photopigment, rhodopsin. This shift may
are less easily made. Although the individual components of brighten the appearance of a blue-black evening or night sky
vision can be divided into the ability to detect light and motion, and enhance the contrast between other objects in the environ-
visual perspective, visual field of view, depth perception, visual ment and the background sky.
acuity, and the perception of color and form, the complete The rhodopsin photopigment of dogs and cats is tuned to a
visual experience is a synthesis of these parts into a unified slightly different wavelength of light from that in humans and,
perception of the world. as is typical of species adapted to function well in dim light,
takes longer to completely regenerate after extensive exposure
Sensitivity to Light to bright light. The ranges of wavelengths to which rhodopsin
in dogs, cats, and humans is sensitive are similar, however,
The visual system of most domestic mammals has evolved to indicating that vision in dim light is not enhanced by expanding
improve performance under a wide range of lighting conditions the range of detectable wavelengths. The slight wavelength
so that they may exploit specific ecologic niches. Of domestic shifts in the maximal sensitivity of rhodopsin across species
1
2 兩 SLATTER’S FUNDAMENTALS OF VETERINARY OPHTHALMOLOGY
Iris Pupil
Superior
Cilia
punctum
Limbus
Conjunctiva
Medial
(nasal) Lateral
canthus (temporal)
canthus
Inferior
punctum
Third eyelid
Figure 1-1. Frontal view of the external structures of the canine eye.
Figure 1-3. External view of the eye of a normal cat. Nocturnal
adaptations that allow more light to enter the eye include a large cornea,
a deep anterior chamber, and a relatively posteriorly located lens. (From
Czederpiltz JMC, et al. [2005]: Putative aqueous humor misdirection
syndrome as a cause of glaucoma in cats: 32 cases. J Am Vet Med Assoc
Lacrimal gland Retinal veins 227:1476.)
Tarsal gland Retina

Pigment
epithelium suggests that domestic mammals and humans do not perceive
Cilia Tapetum
the world in exactly the same way.
Ciliary Choroid
body Sclera
Zonule Sensitivity to Motion
Dura mater
Cornea Vitreous Arachnoid Although little work has been done on the motion-detecting
Conjunctiva Lens
Lacrimal
Pia mater abilities of most domestic animals, it is probable that the
puncta Optic nerve perception of movement is a critical aspect of their vision and
that they, like people, are much more sensitive to moving objects
Optic disc
Pupil than stationary ones. Rod photoreceptors, which dominate the
Anterior chamber Lamina cribrosa retinas of domestic mammals, are particularly well suited for
Retinal arterioles detecting motion and shapes, and it follows that the motion-
Iris
Vortex vein
detecting abilities of domestic mammals—especially in dim
Cartilage of Ventral (inferior)
nictitating membrane rectus m.
light—would be well developed. In a study of the visual per-
Gland of nictitating
Non–tapetal fundus formance of police dogs, the most sensitive dogs could
membrane recognize a moving object up to 900 m away but could recog-
nize the same object, when stationary, at only 585 m or less.
Because of the superior visual acuity of the human fovea, the
A minimum threshold for motion detection in bright light for cats
is approximately 10 to 12 times greater than that for humans.
Coronal Equatorial Although humans may be better equipped to detect motion
when directly viewing an object in bright light, it is possible
Sagittal that the vision of domestic mammals may be superior in dim
light, when an object is viewed peripherally, or if it is moving
p ole at a certain speed to which the retina is particularly attuned.
Posterior
The ability to detect motion may help explain certain
ole behavior—much of the very large peripheral visual field of the
Anterior p
horse probably supports only the detection of brightness and
motion. When combined with a “prey mentality,” this may cause
the horse to treat every moving object in its peripheral field of
view as dangerous and to be avoided. Similarly, many dogs and
cats ignore static objects, but when these objects move, chase
behavior is elicited, suggesting that the visual system has pre-
Horizontal
ferences for objects moving at certain speeds.
B Sensitivity to Flickering Lights

Figure 1-2. Internal structures of the canine eye (A). Also shown are the
standard reference planes (B). Although not related to motion detection, the point at which
rapidly flickering light fuses into a constantly illuminated light
(flicker fusion) provides insight into the functional charac-
STRUCTURE AND FUNCTION OF THE EYE 兩 3
Figure 1-4. Cellular tapetum of a dog

(A) and fibrous tapetum of a horse (B).
(B from Gilger B [2005]: Equine
Ophthalmology. Saunders, St. Louis.
A and B courtesy Dr. Christopher J.
Murphy.)
A B
A B
Figure 1-5. The effect of visual perspective on vision. The same scene as
viewed by a small dog with eyes located 8 inches above the ground (A),
a tall dog with eyes 34 inches above the ground (B), and a person with
eyes 66 inches above the ground (C). (From Miller PE, Murphy CJ [1995]:
Vision in dogs. J Am Vet Med Assoc 207:1623.)
teristics of rod and cone photoreceptors. The flicker frequency and has a major impact on the perception an animal has of its
at which fusion occurs varies with the intensity and wavelength environment (Figure 1-5). For example, when the visual fields
of the stimulating light. Because dogs can detect flicker at 70 of its two eyes are combined, the horse has a total horizontal
to more than 80 Hz, a television program in which the screen is visual field of up to 350 degrees, with 55 to 65 degrees of
updated 60 times/sec and appears to people as a fluidly moving binocular overlap and a virtually complete sphere of vision
story line may appear to dogs as rapidly flickering. around its body (Figure 1-6). The length of the horse’s nose
interferes with binocular vision, and so a horse views an object
Visual Field of View binocularly until it is about 1 m away, at which point the horse
must turn its head and observe with only one eye. In compar-
The extent of the visual field (i.e., the area that can be seen by ison, humans have a visual field of approximately 180 degrees
an eye when it is fixed on one point) and the height of the eyes (140 degrees of overlap), cats have a 200-degree field of view
above the ground may vary greatly among breeds and species (140-degree overlap), and depending on breed, dogs have 250
with both eyes simultaneously does not guarantee improved

OCULAR VISION
BIN perception of depth. Stereopsis (binocular depth perception)
65° results when the two eyes view the object from slightly different
vantage points and the resulting image is blended or fused into
a single image. If the two images are not fused, double vision
may result. (Such an alteration in vision may occur in animals
with orbital diseases.) Although binocular depth perception is
UNIOCULAR VISI
LAR VISION
superior if the images can be blended into one, monocular

depth perception is also possible. Horses make distance judg-
ments on the basis of static monocular clues; these clues
146°
146°
include relative brightness, contour, areas of light and shadows,
object overlay, linear and aerial perspective, and density of
IOCU
optical texture. In addition, movement of the head results in an

apparent change in the relative positions of the objects viewed
O
UN
N (a phenomenon known as parallax) and produces the sensation

that objects are moving at different speeds, allowing depth to be
estimated (Figure 1-8).
3° Visual Acuity
BLIND AREA Visual acuity refers to the ability to see details of an object
Figure 1-6. The visual field of the horse showing a binocular field separately and in focus. It depends on the optical properties of
(65 degrees) comparable to that of a dog but with much larger panoramic the eye (i.e., the ability of the eye to generate a precisely
monocular fields (146 degrees), and a very small blind area (3 degrees). focused image), the retina’s ability to detect and process images,
and the ability of higher visual pathways to interpret images
degrees (30 to 60 degrees of binocular overlap) (Figure 1-7). The sent to them. In general, visual acuity in most domestic mammals
horse has only a few minor “blind spots,” which are located is limited by the retina and not by the optical properties of the
superior and perpendicular to the forehead, directly below the eyes or by postretinal neural processing in the brain. The latter
nose, in a small oval region in the superior visual field where light two factors can limit visual discrimination in a variety of
strikes the optic nerve itself, and the width of the head directly disease states, such as when the lens is removed or when higher
behind. Clearly, this extensive visual field makes it very difficult CNS visual pathways are impaired.
for a person or potential predator to “sneak up” on a horse.
Optical Factors in Visual Acuity
Depth Perception
The optical media of the eye, namely the cornea, aqueous humor,
Depth perception is enhanced in those regions in which the lens, and vitreous humor, are responsible for creating a properly
visual fields of the two eyes overlap. Merely viewing an object focused image on the retina. The cornea and, to a lesser extent,
OCULAR VISION OCULAR VISION

BIN BIN
140° 60°
UN
N
UN
OCULAR VISIO
IOC
IOCU
ULAR VISION
LAR VISION
90°
90°
A
30°
B
30°
U NI
BLIND AREA BLIND AREA

Figure 1-7. A, Visual field of a cat showing a large binocular field (140 degrees) with a relatively small
monocular field (30 degrees) and a relatively large posterior blind area (160 degrees). B, Monocular and
binocular visual fields in a typical mesocephalic dog. The dog has a modest binocular visual field (60 degrees)
with relatively large monocular visual fields (90 degrees) and a posterior blind area of approximately
120 degrees.
Figure 1-8. A number of cues allow depth to be perceived with one eye
or in a two-dimensional photograph. These cues include apparent size
(the left tower appears closer because it is larger than the right), looming
(cars moving toward the viewer appear to become progressively larger),
interposition (near objects such as the bridge overlay the more distant
hills), aerial perspective (water vapor and dust in the air make the more
distant hills less distinct and relatively color-desaturated), shading
(shadows on the tower suggest depth), perspective (the parallel roadways
appear to converge toward the horizon), relative velocity (the nearer cars
appear to move faster than more distant ones), and motion parallax (if the
eye is fixed on the center of the bridge, the images of near objects appear
to move opposite to the direction the observer moves the head, whereas Figure 1-9. Top, The image is properly focused on the retina
distant objects move in the same direction as the head). (From Gilger B (emmetropia). Middle, The image is focused in front of the retina, making
[2005]: Equine Ophthalmology. Saunders, St. Louis.) the eye nearsighted (myopia). Bottom, The image is in focus at a plane that
is behind the retina, making the eye farsighted (hyperopia). (Modified
from Miller PE, Murphy CJ [1995]: Vision in dogs. J Am Vet Med Assoc
207:1623.)
the lens are the principal refracting surfaces of the eye, and
their ability to bend (refract) light is determined by their radii
of curvature and the differences between their refractive index errors of cats and horses are somewhat conflicting, it appears that
and that of the adjacent air or fluid. If the focal length of the the average refraction for these species approximates emmetropia,
focusing structures of the eye does not equal the length of the although deviations of 1 to 2 D do regularly occur.
eye, a refractive error is present. In a normally focused In addition to myopia and hyperopia, other optical aberra-
(emmetropic) eye, parallel rays of light (effectively anything tions (e.g., astigmatism) may result from imperfections in the
20 feet or more away from the eye) are accurately focused on refractive media such as the cornea or lens and lead to degradation
the retina. If parallel rays of light are focused in front the retina, of the image formed on the retina. Astigmatism occurs when
myopia (nearsightedness) results. If they are focused behind the different regions of the optical system (especially cornea or
retina, hyperopia (farsightedness) results (Figure 1-9). Such lens) do not focus light in a uniform fashion, resulting in
errors in refraction are usually expressed in units of optical warping of the image, an extreme example of which can be
power called diopters (D). The extent of the error can be found in the irregular mirrors found at carnivals. Spontaneous
expressed by the formula D = 1/f, where f equals the focal astigmatism is generally uncommon in dogs but has been
length (in meters) of either the lens or the optical system as a observed in a variety of breeds. Astigmatism commonly
whole. Therefore if an eye is 2 D myopic at rest, it is focused at accompanies corneal diseases that result in scarring and
a plane located 0.5 m in front of the eye. Similarly, an eye that distortion of the corneal curvature (Figure 1-10).
is emmetropic at rest but can accommodate (change focus) 3 D Although visual acuity requires that optical portions of the
is capable of clearly imaging objects on the retina that range eye be transparent and that optical blur from refractive errors or
from as far away as the visual horizon (infinity) to as near as astigmatism be limited, an adjustable focusing (accommodative)
0.33 m in front of the eye. mechanism is needed if objects at different distances are to be
The average resting refractive state of the dog is within 0.25 D seen with equal clarity. Accommodation in dogs and cats may
of emmetropia. There are individuals, however, that are signifi- be brought about by altering the curvature of the lens surface
cantly myopic, and breed predispositions to myopia are found in or, more likely, by moving the lens anteriorly (Figure 1-11).
German shepherds and Rottweilers. In one study, 53% of German The accommodative range for most domestic animals is quite
shepherds were myopic by –0.5 D or more in a veteri-nary clinic limited and does not generally exceed 2 to 3 D for dogs, 4 D for
population, but only 15% of German shepherds in a guide dog cats, and less than 2 D for horses. This finding suggests that
program were myopic, suggesting that dogs with visual dogs are capable of accurately imaging objects on the retina
disturbances such as nearsightedness do not perform as well as that are within 50 to 33 cm of their eyes but that objects nearer
normally sighted dogs. It may be reasonable to screen dogs that than this will be blurred. Hence, dogs use other senses, such as
will be expected to perform visually demanding tasks, or those on smell or taste, to augment vision in the investigation of very
which human life relies for refractive errors, before embarking on near objects. For comparison, young children are capable of
extensive training programs. Although studies of the refractive accommodating approximately 14 D, or to about 7 cm.
light as occurs in dogs typically requires that a greater number

of photoreceptors (primarily rods) synaptically converge on a
single ganglion cell. This results in reduced visual acuity, just
as high-speed photographic film produces a “grainy” image in
bright daylight. Additionally, the tapetum also scatters light and
further degrades visual acuity in bright light. Retinas with
excellent resolving power have a high ratio of ganglion cells to
photoreceptors, a large number of ganglion cells and optic
nerve fibers, and a high density of photoreceptors and usually
lack a tapetum. In primates, the fovea has one ganglion cell per
cone, whereas in cats, the peak ratio is one ganglion cell for
every four cones. In all species, there are fewer ganglion cells
in the periphery of the retina than in the center, and the ratio
may decline to 1:16 in primates and 1:20 in cats, explaining the
reduced visual acuity of their peripheral visual fields.
Figure 1-10. Corneal astigmatism after a corneal laceration with Domestic mammals lack the highly developed primate fovea
subsequent scarring. Note the irregular flash artifact on the cornea but, instead, have a generally oval visual streak that contains
indicating that light is being unevenly focused on the retina. (From Gilger the greatest density of photoreceptors, ganglion cells, and
B [2005]: Equine Ophthalmology. Saunders, Philadelphia. Courtesy
Dr. Ellison Bentley.) rhodopsin and thereby affords the greatest visual acuity. The visual
streak, located in the tapetal region slightly superior and tem-
poral to the optic nerve, has approximately linear, short
Loss of the lens, as occurs after cataract surgery, results in temporal and longer nasal extensions (Figure 1-12). The oval
severe hyperopia (farsightedness), with objects being focused temporal part of the visual streak is relatively free of blood
approximately 14 D behind infinity, and a reduction in visual vessels larger than capillaries, and nerve fibers take a curved
acuity to 20/800 or worse. This means that aphakic eyes are course to the optic disc dorsal and ventral to the visual streak,
unable to image any object clearly, whether near or far away, presumably to avoid interfering with light reaching the photo-
and are unable to accommodate. Although the aphakic dog is receptors. The temporal, oval portion of the streak may facili-
extremely “farsighted,” it must be kept in mind that, for objects tate binocular vision, whereas the nasal, linear portion may be
of similar size, objects that are closer to the dog will create a used to scan the horizon and better use the wider field of view
much larger image on the retina than objects that are located far available to the domestic mammals.
away. Therefore the aphakic dog may be able to better visually Wolves, presumably the ancestors of modern-day dogs, have
orient to near objects despite being “farsighted.” Surprisingly, a pronounced visual streak with a dense central area and exten-
although this degree of hyperopia is markedly debilitating to sions far into the temporal and nasal portions of the retina. In
some dogs, most dogs are still able to visually orient adequately contrast, domesticated dogs, even of the same breed, have
in their environment without correction. either a similar pronounced visual streak or a smaller, less
densely packed, moderately pronounced visual streak. Wolves
Retinal Factors in Visual Acuity also generally have a greater maximum density of ganglion
cells (12,000 to 14,000/mm2) than do most dogs (6,400 to
The retina may be the limiting factor in visual acuity for normal 14,400/mm2). This difference implies that the visual acuity of
domestic animals, and its architecture may provide clues to the wolves may be better than that of some dogs, and that the
potential visual abilities of the eye. Enhanced vision in dim constancy of the form of the visual streak in wolves may be a
Meridional fibers
Radial fibers
Figure 1-11. Classic accommodation in primates. Left,
Distant vision. Relaxation of the ciliary muscle increases Circular fibers
tension on the lens zonules, which flattens the lens and Zonular fibers
Iris (suspensory ligaments)
brings distant objects into focus. Right, Near vision.
Contraction of the ciliary muscle reduces tension on the Lens Lens
zonules, which allows the elastic lens capsule to assume a Cornea relaxed accommodated
more spheric shape. The resulting increase in lens power
allows near objects to be brought into focus on the retina.
The importance of this mechanism of accommodation in
most domestic mammals is debated. (Modified From Getty
R [1975]: Sisson and Grossman’s The Anatomy of the
Domestic Animals, 5th ed. Saunders, Philadelphia.) Ciliary process
Fibers of ciliary Fibers of ciliary
muscle relaxed muscle contracted
A B
Figure 1-12. Diagram of retinal ganglion cell densities from the right retinas of a German shepherd with a very
pronounced wolflike visual streak (A) and a beagle with a moderately pronounced visual streak (B). Retinas were
cut radially to flatten them and are displayed at the same magnification. The intensity of the dots reflects varying
ganglion cell densities. The irregular shape in the center of each retina is the region of the optic nerve head.
Ganglion cells could not be seen in this area because of thick, overlying nerve fiber layer. (From Miller PE,
Murphy CJ [1995]: Vision in dogs. J Am Vet Med Assoc 207:1623.)
result of environmental selective pressures that were altered by a moving cotton ball) test the motion sensitivity of virtually the
domestication. It is unclear whether there are differences in the entire retina, and positive responses are still present even though
visual acuity of dog breeds that have been developed to hunt by visual acuity may be very poor (up to 20/20,000). Visually
sight (sight hounds) and breeds that have been developed to distinguishing the fine details in objects is less important for most
hunt by smell (scent hounds), although the finding of a large domestic mammals (even working animals) than it is for most
number of beagles (a scent hound) with a pronounced visual people. The trade-off of improved vision in dim light for less
streak suggests that there are insignificant differences between acute vision in bright light allows such animals to exploit
these two groups of dogs, despite their uses. ecologic niches inaccessible to people and aids in both seeking
prey and avoiding predators.
Estimates of Visual Acuity
Color Vision
The most familiar indicator of visual acuity for the human eye
is the Snellen fraction, which relates the ability of a subject to Color vision in domestic mammals has been the subject of
distinguish between letters or objects at a fixed distance numerous studies with conflicting results. More recent, well-
(usually 20 feet, or 6 m) with a standard response. Snellen controlled studies suggest that most domestic mammals possess,
fractions of 20/20, 20/40, and 20/100 mean that the test subject and use, color vision.
needs to be 20 feet away from a test image to discern the details The presence of cone photoreceptors in domestic mammals
that the average person with normal vision could resolve from suggests the potential for color vision, although the numbers
20, 40, and 100 feet away, respectively. This test actually measures and types of cones are smaller than those in humans. Cones
the ability of the area of greatest visual acuity (the fovea) to dis- constitute less than 10% of the visual streak in the dog, whereas
criminate between objects of high contrast. Peripheral visual they occupy almost 100% of the human fovea. Additionally,
acuity in humans is typically quite poor (i.e., 20/100, 20/200, or instead of three types of cones (“red,” “green,” and “blue”) found
worse), presumably because the photoreceptor density is lower in humans with normal color vision, dogs have only two
and the ratio of photoreceptors to ganglion cells is higher in functional cone types. One type of canine cone is maximally
these regions of the retina than in the fovea. The visual acuity sensitive to light at 429 to 435 nm (“violet” to normal humans
of the normal dog ranges between 20/50 and 20/140, with and corresponding to the “blue” cone), and another type has
20/75 or so being the likely average. Feline visual acuity has maximal sensitivity to light at 555 nm (“yellow-green” to normal
been estimated to be between 20/100 and 20/200, whereas the humans) with extension into the red end of the color spectrum
larger eye of the horse (hence a greater numbers of photo- (corresponding to the “red” cone). Dogs lack, or do not use,
receptors) may result in a visual acuity of 20/30. The visual “green” cones and appear to confuse red and green colors (red-
acuity of cattle, however, is unclear, because the size of their green color blindness, or deuteranopia). This means that dogs
eyes and density of ganglion cells would suggest they possess are unable to differentiate middle to long wavelengths of light,
a visual acuity comparable to that of horses but behavioral which appear to people as green, yellow-green, yellow, orange,
studies that depend on the cooperation of the animal have or red.
documented a visual acuity of only 20/240 to 20/440. Although it is not known whether the dog’s “blue” and “red”
Most commonly used procedures to determine vision in cones perceive colors in the same way as those of humans, the
animals (e.g., determination of menace responses by moving a canine visible spectrum may be divided into two hues: one in
hand across the visual field or having the animal’s eyes follow the human violet and blue-violet range (430 to 475 nm), which
is probably seen as blue by dogs, and a second in the human distinguish red, orange, yellow, and green objects solely on the
greenish-yellow, yellow, and red range (500 to 620 nm), which basis of color. In these cases, other visual clues, such as relative
is probably seen as yellow by dogs. Dogs also appear to have a brightness and contrast, or the other senses—smell, sound,
narrow region (475 to 485 nm, blue-green to humans) that taste, and touch—are required to differentiate objects that appear
appears colorless. Light in this spectral neutral point probably similar in color. Additionally, dogs, and probably most other
appears to be white or a shade of gray to dogs. In people with domestic mammals, are able to differentiate perfectly between
deuteranopia, however, the neutral point is in a greener (505-nm) closely related shades of gray indistinguishable to the human
region of the spectrum, so dogs are not exactly the same as red- eye. This ability is far more valuable in exploiting their ecologic
green color blind humans. Wavelengths at the two ends of the niche than color vision, because it increases visual discri-
spectrum (blue at one end and yellow at the other) probably mination when insufficient light may be present to effectively
provide the most saturated colors. Intermediate wavelengths stimulate cones.
are less intensely colored, appearing as if they were blends with
white or gray. CENTRAL VISUAL PATHWAYS
The cat has a limited but detectable capacity for color vision
and can distinguish between two stimuli if they differ greatly in The eye is only the first step in “seeing” (Figure 1-15). Vision
spectral content, especially if the stimuli are also large. Cats is not simply a recording of each pixel in a scene, as a camera
appear to have the physical capacity (based on the presence of would make, because that would quickly overwhelm the visual
three types of cones) for trichromacy like humans, although system with massive amounts of information that may not be
behavioral studies have not demonstrated this ability and it is, pertinent to the animal’s survival or lifestyle. The brain does
at best, a pale copy of human trichromacy. Horses appear to not, and cannot, consciously pay attention to the flood of infor-
have both a short-wavelength-sensitive (blue) cone with a peak mation it receives from the eyes, but instead categorizes the
sensitivity of approximately 428 nm, and a second cone with a information into specific “topics” that are channeled to specific
peak sensitivity between the human red and green cones (539 nm, areas of the brain for further processing. These “topics” are
and called a middle–long-wavelength-sensitive cone). Therefore “internal” features such as texture and contrast, the direction
although horses have only two cones like dogs, orange and blue and velocity of the object’s movement, its overall orientation as
colors appear similar (shades of gray) to horses, whereas red and represented on the retinal surface, its shape, its color, and many
green appear similar (as shades of gray) to dogs (Figures 1-13 other aspects. Additionally, unlike a camera, the brain compares
and 1-14). Cattle and swine also appear to have two functional the current image with previous images, images from the other
cone pigments. The yellow tint to the equine lens probably eye, and input from other senses such as hearing, smell, and
filters out blue wavelengths, diminishing certain optical aberra- touch. Once this comparison is completed, only the information
tions as well as glare and increasing the contrast of certain that is relevant for the task at hand, or the animal’s survival,
objects on select backgrounds. rises to the level of conscious attention. Therefore the act of
Restrictions in color vision are probably of limited con- seeing depends not only on the function and health of the eye
sequence to domestic mammals, as it is likely that they react but also on the cognitive processes in the brain that decide what
only to colors of biologic importance to them. Problems may information merits conscious attention and what is to remain
arise when one is teaching hunting and working dogs to subconscious or ignored.
The central visual pathway begins with the retina, which is
in effect an extension of the brain. In this tissue, information is
processed in three functional stages. The first stage occurs in
the rod and cone photoreceptors. These cells have varying
sensitivity to different wavelengths of light and require
different numbers of photons to strike them in order to elicit a
A 0.7 ␮m 0.6 ␮m 0.5 ␮m 0.4 ␮m response. The second stage of retinal processing occurs in the
Infrared Ultraviolet outer plexiform layer. At this level the photoreceptors, bipolar
cells, and horizontal cells synaptically interact, and the
responses of some photoreceptors are modulated by what is
happening to other photoreceptors (so-called “on” and “off” or
“center” and “surround” responses). These responses are
associated with the static and spatial aspects of an object and
serve to better define an object’s brightness and borders by
altering its contrast with surrounding objects. The third stage
occurs in the synaptic interactions in the inner plexiform layer,
which is more concerned with the dynamic and temporal
B C aspects of vision. At this stage, transient responses that may
underlie motion and direction sensitivity may occur in the
Figure 1-13. A, The color spectrum and corresponding wavelengths of
light. B and C, Difference between dichromatic color vision of the horse amacrine and retinal ganglion cells. The amount of retinal
and normal human color vision. B, Color wheel representing the processing of an image before it gets to the brain varies greatly
spectrum of colors perceived by the trichromatic human visual system. by species.
C, Reducing the number of types of cone from three to two results in Retinal ganglion cell nerve fibers then form the nerve fiber
dichromatic color vision and an enormous reduction in the number of
different colors seen. (A from Gilger B [2005]: Equine Ophthalmology.
layer of the retina, converge at the optic disc, turn posteriorly,
Saunders, Philadelphia. B and C from Carrol J, et al. [2001]: Photopigment gain a myelin sheath, and pass through the sievelike opening in
basis for dichromatic color vision in the horse. J Vis Sci 1:80.) the sclera, the lamina cribrosa. The fibers pass via the optic
A B
C D
Figure 1-14. Simulation of the visual acuity and color vision of the horse. Original image (A, B). Images adjusted
to reflect the visual acuity and color vision abilities of the horse (C, D). (From Carrol J, et al. [2001]:
Photopigment basis for dichromatic color vision in the horse. J Vis Sci 1:80.)
nerve to the optic chiasm (see Figures 1-2, 1-15, and 1-16). humans—as the eyes become more frontally directed. This
Fibers coming from different parts of the retina maintain definite finding is clinically relevant because species with extensive
positions within the optic nerve and throughout the path to the binocular overlap will not necessarily bump into objects if only
visual cortex. Fibers from both optic nerves enter the optic one eye is blind or there is a lesion on only one side of
chiasm, where partial decussation, or “crossing over” of fibers the brain.
from one side to the other, may occur (see Figure 1-15). The optic chiasm receives the optic nerves as they enter the
The proportion of optic nerve fibers that decussate in the cranial vault via the optic foramen and canals (Figure 1-17).
chiasm is related to the relative laterofrontal positioning of the The chiasm lies at the base of the brain adjacent and anterior to
orbit and eye in the skull and degree of binocular overlap. the hypophysis, which sits in the pituitary fossa of the
Animals with laterally directed eyes and no overlap between postsphenoid bone. This relationship between the pituitary
the visual fields of the two eyes exhibit complete decussation at and the chiasm and optic tracts is important in considering
the chiasm, and the information from the right (or left) visual the potential effect of space-occupying masses of the pituitary
field is processed entirely by the opposite visual cortex. As on vision.
the eyes become more frontally directed in different species, From the optic chiasm, fibers enter the left and right optic
however, it becomes possible for an object to be seen with both tracts, which then pass laterally from the chiasm, anterior to the
eyes. For example, an object in the animal’s right visual field hypophysis, and beneath the ventral surface of the cerebral
(on its right side) falls on the nasal area of the right retina and peduncle. The tracts then curve dorsally and posteriorly, between
the temporal area of the left retina. In order for the same side of the cerebral peduncle to which they are attached laterally and the
the brain (the left side in this example) to continue to process pyriform lobe. The tracts thus pass to the lateral geniculate body.
all the information from the right visual field, some optic nerve Before reaching the lateral geniculate body, some 20% to 30% of
fibers must remain ipsilateral and must not decussate at the the fibers leave the tracts and enter the pretectal area. Some of
chiasm. This percentage varies by species, but in horses and these fibers enter the superior colliculus directly, and others pass
cattle, which have relatively laterally directed eyes, 83% to via the tracts and lateral geniculate body to the colliculus
87% of the optic nerve fibers cross, whereas the percentage indirectly. The majority of fibers entering the lateral geniculate
decreases—75% for dogs, 67% for cats, and 50% for body synapse here with the third ascending neuron in the visual
system, which passes without further synapse to the visual

Temporal Nasal Nasal Temporal cortex. The first two synapses are the photoreceptor-bipolar and
field field field field bipolar-ganglion cell interfaces.
Those fibers that leave the optic tracts before entering the
lateral geniculate body pass to the pretectal area, carrying
afferent impulses of the pupillary light reflex. In the pretectal
area much decussation occurs, and the fibers pass to the midline
Edinger-Westphal nuclei of the oculomotor nerve (see Figure
1-15). Efferent impulses pass from these nuclei to the pupillary
sphincter muscle in each iris.
LEFT EYE RIGHT EYE A positive pupillary light reflex does not mean that the eye can
see. Fibers that mediate the reflex arc leave the optic tracts
Temporal Nasal before the tracts enter the lateral geniculate body.
retina retina
Ciliary From the lateral geniculate body, fibers pass forward and lateral
ganglion to the lateral ventricle as the fanlike optic radiation, which
Optic
nerve enters the occipital or visual cortex, where interpretation of
some visual stimuli occurs in domestic animals (Figure 1-18).
Optic Optic Increases in intraventricular pressure (hydrocephalus) can
tract chiasm
affect the visual pathway at this point.
In dogs and cats the visual cortex is not the sole center of
Edinger-Westphal interpretation of visual stimuli. If the cortex is removed, light
nucleus
Midbrain perception and discrimination of light intensity are retained, but
familiarity of surroundings is lost. Subcortical integration is
Lateral
geniculate believed to occur in the superior colliculus.
Superior colliculus body
Pupillary
fibers and pretectal area
VASCULAR ANATOMY AND PERIPHERAL
NEUROANATOMY
Further details of orbital anatomy may be found in Chapter 17.
Optic
radiation
Arterial Supply
Left occipital lobe Right occipital lobe The major arterial supply of the eye is from the external
Figure 1-15. The visual pathway. ophthalmic artery, a branch of the internal maxillary artery,
N. olfactorius l Olfactory bulb Olfactory peduncle
A. ethmoidalis interna Olfactory tracts:

Medial
A. communicans rostralis Lateral
N. opticus ll Rostral perforated substance
A. cerebri rostralis Optic chiasm
A. cerebri media Optic tract
A. carotis interna, cut stump Infundibulum of hypophysis
N. oculomotorius lll Piriform lobe
A. cerebri caudalis Mammillary body

Figure 1-16. Ventral view of the brain and cranial
nerves in the dog. Structures important to the A. communicans caudalis Caudal rhinal sulcus
visual system are highlighted. (From Done SH, et Interpeduncular fossa
al. [1996]: Color Atlas of Veterinary Anatomy, A. cerebelli rostralis
Cerebral peduncle
Volume 3. Mosby, St. Louis.) N. trigeminus V
N. trochlearis lV
N. abducens Vl
Pons
N. facialis Vll
Paraflocculus
N. vestibulocochlearis Vlll
Flocculus
N. vagus X and
N. glossopharyngeus lX Cerebellar hemisphere
A. cerebelli caudalis Trapezoid body
N. accessorius Xl A. basilaris
N. hypoglossus Xll Pyramid
N. cervicalis l Ventral median fissure
Palatine fissure
Infraorbital foramen
Fossa for lacrimal sac
Cribriform plate
Maxillary foramen
Lateral part of frontal sinus
Alveolar foramen
Salcus chiasmatis Figure 1-17. Skull of the dog with calvaria
Optic canal removed, dorsal aspect. Structures important
Rostral clinoid process to the visual system are highlighted.
Orbital fissure (Modified from Evans HE [1993]: Miller’s
Hypophyseal fossa Anatomy of the Dog, 3rd ed. Saunders,
Philadelphia. © Cornell University 1964.)
Foramen rotundum
Caudal clinoid process
Foramen ovale
Dorsum sellae
Crista petrosa Canal for trigeminal nerve
Canal for transverse sinus Internal acoustic meatus
Cerebellar fossa Jugular foramen
Hypoglossal canal Condyloid canal
F = face
A = arm-forelimb
T = trunk
L = leg-hindlimb
M = mouth and tongue
N = nose and lips
Auditory area
Prefrontal area
Motor area
Somatosensory
area
Visual area
Figure 1-18. Motor and sensory areas of the cerebral cortex of the dog. (Modified from Hoerlein BF [1978]:
Canine Neurology, 3rd ed. Saunders, Philadelphia.)
which arises from the external carotid artery (Figure 1-19). There is no central retinal artery in domestic species. Single
The contribution from the internal carotid artery is small, medial and lateral long posterior ciliary arteries pass around
unlike the situation in primates, and is via an internal the globe horizontally, within the sclera, to supply the ciliary
ophthalmic artery, which arises from the circle of Willis. The body. Muscular branches of the orbital artery, which supplies
internal ophthalmic artery enters the orbit through the optic the extraocular muscles, also enter the globe near the insertions
canal with the optic nerve. From the external ophthalmic artery, of these muscles. These anterior ciliary arteries anastomose
numerous short posterior ciliary arteries arise (Figure 1-20) with the long posterior ciliary arteries to form the ciliary
and penetrate the sclera around the optic nerve head. These arterial supply. When the globe is prolapsed, these muscular
arteries supply the retina and choroid. branches may be destroyed, decreasing the available supply to
M. rectus dorsalis
M. levator palpebrae
Dorsal muscular br.
M. retractor bulbi
Ext. ethoidal a.
Lacrimal gland Ext. ophthalmic a.
Int. ophthalmic a.
M. rectus lateralis
Rostral cerebral a.
M. obliquus ventralis Middle cerebral a.
Caud. communicating a.
M. rectus ventralis Int. carotid a.
Malar a.
Middle meningeal a.
Maxillary a.
Anastomotic ramus
Br. to m. rectus medialis
Ext. ophthalmic a.
Ventral muscular br.
Rostral deep temporal a.
Figure 1-19. Arteries of the orbit and extrinsic ocular muscles in the dog, lateral aspect. (From Evans HE [1993]:
Miller’s Anatomy of the Dog, 3rd ed. Saunders, Philadelphia.)
Anterior ciliary artery

Anterior ciliary vein
Long posterior
ciliary artery
Vortex vein
Anterior ciliary vein
Retinal vein
Retinal arterioles
Ciliary process
Long posterior
ciliary artery
Figure 1-20. Vascular supply of the canine eye.
the anterior segment of the eye. Branches from the ciliary The eyelids are supplied by the superficial temporal artery, a
arterial network form the major arterial circle of the iris. The deep branch of the external carotid artery, and by the malar artery, a
conjunctival arterioles at the limbus anastomose with the anterior branch of the infraorbital artery.
ciliary arteries before they enter the sclera, and also with arterioles
in the ciliary body. Vascular events of clinical importance (e.g., Venous Drainage
inflammation in one area of this network of anastomosing vessels)
can often be seen in other parts, and their origin must be The retina is drained by the retinal veins and venules, which
distinguished clinically if possible (see Figure 1-20). run from the peripheral retina toward the optic nerve head (see
Orbital plexus the vertebral sinuses, external jugular vein, and internal maxillary
vein. Venous blood thus passes posteriorly from the orbit via
Ophthalmic vein this route. It may also pass anteriorly via anastomoses between
Vortex veins the ophthalmic veins and the malar, angularis oculi, and facial
veins to the external maxillary and external jugular veins.
Angular vein
Considerable species variation exists in the vascular supply
and drainage of the eye and orbit.
Nerve Supply of the Eye and Adnexa

The general plan of nerve supply to the eye is shown in
Internal maxillary vein Figure 1-22. For further details, see Chapter 16.
Deep facial vein Facial vein
Optic Nerve (Cranial Nerve II)
External jugular vein External maxillary vein
Figure 1-21. The venous drainage of the eye and orbit of the dog.
The optic nerve and meninges pass from the globe, through the
(Modified from Startup FG [1969]: Diseases of the Canine Eye. Williams cone formed by the retractor bulbi muscles, via the optic canal
& Wilkins, Baltimore.) to the optic chiasm. The dura covering the nerve is continuous
with the outer layers of the sclera. The optic nerve consists of
ganglion cells, whose cell bodies lie in the ganglion cell layer
Figures 1-20 and 1-21). The venous circle they form at the optic of the retina. It is a tract of the CNS, not a peripheral nerve.
disc may be complete or incomplete in the dog. The venous
circle drains posteriorly through the sclera via the posterior Oculomotor Nerve (Cranial Nerve III)
ciliary veins to a dilation in the orbital vein, the superior
(dorsal) ophthalmic vein. The nucleus of the oculomotor nerve lies in the brainstem and
The choroid is drained by approximately four vortex veins, has several components serving different extraocular muscles,
which leave the globe near the equator and join the superior the ventral rectus, dorsal rectus, medial rectus, inferior
and inferior ophthalmic veins. The ciliary body is drained by oblique, and levator palpebrae superioris muscles. The nerve
the anterior ciliary veins to the same superior and inferior also carries parasympathetic fibers originating from the
ophthalmic veins that drain to the orbital venous plexus at the Edinger-Westphal nucleus, which lies near the other nuclei of
apex of the orbit. This plexus drains to the cavernous venous the oculomotor nerve, and serves the sphincter pupillae and
sinus within the cranial vault. The cavernous sinus drains via ciliary muscles. The oculomotor nerve thus contains efferent
Optic nerve (sensory)

Retina
Optic foramen
(fissure) Oculomotor (motor d
orsal div
i sion
Trochle
r ner
ve (m Ciliar y ga
Levator palpebra and dorsal rectus muscle
Ophthalmic n otor ng
lion
erve
(sen
) Ventral divi Medial ventral rectus muscle
Ab d so r sion
uc e y) Ventral oblique muscle
ns
ne
rve Su
pra Dorsal oblique muscle
(m orb
Inf ital
rao ner
ve
ot
rbita
or
La l ner
ve Skin of upper lid and forehead
)
crim
al n
2. Eth
mo
erve Medial canthus and third eyelid
1. ida
l ne
rve Lacrimal gland
Cili
M ary Nasal mucosa via ethmoid foramen
Round foramen ax
gang
lion
illa
ry Cornea, iris, and ciliary body
ne
rv
e
Extraocular muscles
(se
ns Lateral rectus muscle
or y
) Retractor bulbi muscle
Zyg
Stylomastoid foramen om atis nerve
Upper eyelid
Lower eyelid
Fa ebral ner ve
Facial area
cia lo-palp
l ner Auricu Lacrimal gland
ve (motor) VII
Orbicularis oculi muscle
1. Round and orbital foramina fuse in the pig and

ruminants, forming the foramen orbitorotundum
2. Only orbital branches shown here
Figure 1-22. Nerve supply to the eye.

motor fibers to the striated extraocular muscles, of mesodermal and adjacent skin (Figure 1-25). In horses, cattle, sheep, and
origin, and parasympathetic fibers to the smooth muscles of the pigs, the nerve reaches the upper lid via the supraorbital
iris and ciliary body, of neuroectodermal origin. foramen, but in dogs and cats, it passes beneath the orbital
The oculomotor nerve leaves the brainstem on its ventro- ligament.
medial surface (see Figure 1-16), and passes ventral to the optic The lacrimal nerve supplies the lacrimal gland. The
tracts, through the cavernous sinus, and enters the orbit via the nasociliary nerve, the major continuation of the ophthalmic
orbital fissure (foramen orbitorotundum in cattle, sheep, and nerve in the orbit, gives rise to the ethmoidal and infratrochlear
pigs). In the orbit, the nerve divides into dorsal and ventral nerves. The ethmoidal nerve passes through the ethmoidal
rami. A branch from the ventral ramus passes to the ciliary foramen to supply the mucous membranes of the nasal cavity.
ganglion, where the preganglionic parasympathetic fibers The infratrochlear nerve passes beneath the trochlear, pene-
synapse. For more details on the fibers leaving the ciliary trates the septum orbitale, and innervates the medial canthus,
ganglion, see Figures 1-23 and 1-24. third eyelid, and adjacent lacrimal system (see Figure 1-25).
Within the orbit, the nasociliary nerve gives off the long ciliary
Trochlear Nerve (Cranial Nerve IV) nerve, which enters the globe near the optic nerve to provide
sensory innervation to the globe itself.
The trochlear nerve leaves the brainstem on the dorsal surface The maxillary nerve passes through the round foramen and
and runs lateral to the tentorium cerebelli to the orbital fissure. via the alar canal to the pterygopalatine fossa. It gives rise to
It passes through the fissure with the oculomotor nerve and the the zygomatic nerve, which divides into zygomaticotemporal
ophthalmic branch of the trigeminal nerve. The trochlear nerve and zygomaticofacial branches within the orbit. The zygomatico-
innervates the dorsal oblique muscle only. temporal branch supplies sensory innervation to the lateral upper
lid and rostral temporal area. The zygomaticofacial branch
Trigeminal Nerve (Cranial Nerve V) emerges from the periorbita ventral to the lateral canthus and
supplies the lateral lower lid and surrounding skin. Post-
The sensory branches of the trigeminal nerve receive the ganglionic sympathetic fibers from the cranial cervical
majority of the input from the orbit and periocular area. The ganglion may also be distributed to the orbit via the branches
nerve has both motor and sensory roots (see Figure 1-16), of the maxillary nerve, which has no other branches of ophthalmic
which pass in a common sheath through the petrous temporal significance.
bone to the trigeminal ganglion. The three branches of the
nerve—ophthalmic, maxillary, and mandibular—arise from the Abducent Nerve (Cranial Nerve VI)
trigeminal ganglion. The ophthalmic nerve leaves the cranial
cavity via the orbital fissure, and the maxillary nerve via the The abducent nerve leaves the ventral surface of the medulla
round foramen (see Figure 1-22). oblongata (see Figure 1-16) and passes through the wall of the
Once in the orbit, the ophthalmic nerve divides into the cavernous sinus, forward via the orbital fissure (see Figure 1-22),
supraorbital (frontal), lacrimal, and nasociliary nerves. The to enter the orbit and supply the retractor bulbi and lateral
supraorbital nerve is sensory to the middle of the upper eyelid rectus muscles.
Major petrosal nerve Tensor tympani muscle
Trigeminal nerve Chorda tympani
Abducent nerve Minor petrosal nerve

Head of malleus
Geniculate ganglion
Short crus of incus
Facial nerve
Stapes
Vestibular nerve
Vestibule
Cochlear nerve
Chorda tympani
Glossopharyngeal nerve Nerve to stapedius muscle
Proximal ganglion of X Auricular branches of VII
Vagus nerve Location of stapedius muscle
Hypoglossal nerve Auricular branch of X
Accessory nerve
Condyloid canal
Figure 1-23. The canine petrous temporal bone, sculptured to show the path of the facial nerve, dorsal aspect.
(Modified from Evans HE [1993]: Miller’s Anatomy of the Dog, 3rd ed. Saunders, Philadelphia. © Cornell
University 1964.)
AUTONOMIC INNERVATION
SYMPATHETIC
Cranial nerve ganglion Dilator pupillae muscle
Lacrimal gland
PARASYMPATHETIC
Ciliary ganglion Sphincter pupillae muscle
via oculomotor nerve Ciliary muscle
Pterygopalatine ganglion
Lacrimal gland
A via facial nerve
Pretectal nucleus
Edinger-Westphal nucleus
Hypothalamus
Thoracic segments
1 2 3
Ciliary ganglion
Cranial cervical Sympathetic chain ganglia

B ganglion
Figure 1-24. A, Autonomic innervation of the eye. B, The motor pathways to the iris: parasympathetic constrictor
and sympathetic dilator. (B modified from Hoerlein BF [1978]: Canine Neurology, 3rd ed. Saunders,
Philadelphia.)
Facial Nerve (Cranial Nerve VII) Autonomic Innervation

The mixed facial nerve contains somatic motor and para- Action of the Ocular Muscles with Autonomic
sympathetic fibers, innervating the orbicularis oculi and Innervation (Table 1-1)
retractor anguli muscles and the lacrimal gland. Cell bodies of The pupillary dilator and sphincter muscles are antagonistic to
the motor fibers are found in the facial nucleus in the medulla each other and control the size of the pupil. As one muscle
oblongata. The parasympathetic cell bodies are located in the contracts, the other relaxes. If either muscle fails to function,
rostral salivatory nucleus in the medulla. The nerve leaves the the effects of the remaining muscle predominate; for example,
brainstem lateral to the origin of the abducent nerve (see Figure paralysis of the dilator muscle alone results in a small pupil
1-16) and, with the vestibulocochlear nerve, enters the petrous (miosis) because of the unbalanced action of the sphincter muscle.
temporal bone near the acoustic meatus (see Figure 1-23), a The ciliary muscle varies in size and orientation among
point of clinical significance to be discussed later with species but in general is composed of one set of smooth muscle
hemifacial spasm (see Chapter 16). The facial nerve enters the fibers arranged so as control tension on the lens zonules and the
facial canal of the temporal bone, where the geniculate refractive power of the lens and another set that controls the
ganglion is situated. relative width of the iridocorneal angle, through which aqueous
From the ganglion arises the major petrosal nerve. Joined by humor exits the eye. Müller’s muscle elevates the upper eyelid
the deep petrosal (sympathetic) nerve, the nerve of the together with the levator palpebrae superioris muscle. If either
pterygoid canal is formed and passes via the pterygoid canal to muscle is defective or denervated, the upper eyelid droops
the pterygopalatine fossa, ending as the pterygopalatine (ptosis).
ganglion. The parasympathetic fibers synapse here, and some
pass to the lacrimal gland. Table 1-1 兩 Actions of the Ocular Muscles with
The facial nerve passes from the geniculate ganglion and Autonomic Innervation
emerges from the stylomastoid foramen to give numerous
branches. The facial trunk terminates as the auriculopalpebral SYMPATHETIC PARASYMPATHETIC
nerve, which crosses the temporal region and zygomatic arch
M. dilator pupillae M. sphincter pupillae
(see Figure 1-25). The palpebral branch supplies the orbicularis Müller’s muscle Ciliary muscle
oculi and retractor anguli oculi muscles.
To scutuloauricularis superficial dorsal muscle

To scutuloauricularis prof. major muscle
Rostral internal
auricular nerve Rostral auricular nerve
Palpebral nerve Rostral auricular plexus
Middle internal
auricular nerve Zygomaticotemporal nerve
Supraorbital nerve (frontal)
Caudal internal
auricular nerve Infratrochlear nerve
Zygomaticofacial nerve
Caudal auricular
branch to platysma Infraorbital nerve
(retroauricular nerve)
Great auricular nerve
IIC, ventral branch
Caudal auricular Ramus 1

branches:
Ramus 2
Facial nerve VII
External jugular vein
Mental nerves
To digastricus muscle
Buccalis nerve
Branches of mylohyoid nerve
To parotidoauricularis muscle
Dorsal buccal branch
To stylohyoideus muscle Nerves to tactile and sinus hairs
Lateral interior auricular nerve Parotid duct (cut)
Cervical branch Auriculotemporal nerve
Auriculopalpebral nerve Ventral buccal branch
Figure 1-25. Superficial branches of the facial and trigeminal nerves in the dog, lateral aspect. (Modified from
Evans HE [1993]: Miller’s Anatomy of the Dog, 3rd ed. Saunders, Philadelphia. © Cornell University 1964.)
Parasympathetic Supply OCULAR REFLEXES

Parasympathetic fibers arise from the Edinger-Westphal All ocular reflexes and responses may be thought of as having
nucleus of the oculomotor nerve and pass via the nerve to five parts: (1) a receptor, (2) an afferent neuron, (3) interneuronal
synapse in the ciliary ganglion (see Figures 1-24 and 1-26). connections, (4) an efferent neuron, and (5) an effector.
Other fibers that pass into the ciliary ganglion but do not Knowledge of the pathways involved in each of the reflexes and
synapse in it are as follows: responses is invaluable for the localization of neuroophthalmic
lesions. Table 1-2 summarizes these reflexes and responses.
• Postganglionic sympathetic fibers from the cranial cervical
ganglion Pupillary Light Reflex (See Figures 1-24 and 1-27)
• Sensory fibers from the ophthalmic branch of the trigeminal
nerve (V) Constriction of the pupil on the same (ipsilateral) side as the
stimulus is termed the direct pupillary reflex, whereas pupillary
Sympathetic Supply constriction on the opposite (contralateral) side is called the
consensual light reflex. In order to obtain reliable reflexes, a
Sympathetic fibers from the brain pass down the cervical spinal bright penlight or transilluminator must be used. Both direct
cord and leave via spinal nerves of segments T1 and T2 and consensual responses are present in normal animals, although
(see Figures 1-24 and 1-26). The fibers leave the nerves, pass to considerable differences exist in the speed of the reflexes
the sympathetic trunk, and pass cranially with it, in its com- between small and large animal species.
mon sheath with the vagus nerve. The sympathetic trunk
terminates cranially at the cranial cervical ganglion, where Palpebral/Corneal Reflex
many of the fibers synapse. Postganglionic fibers pass via a
variety of pathways to the pupillary dilator muscle and to The palpebral/corneal reflex is elicited by touching either the
Müller’s muscle. periocular skin (palpebral) or the cornea (corneal). This reflex
Cell body of preganglionic

parasympathetic fiber in the
Edinger-Westphal nucleus
Nasociliary nerve
Optic nerve Trigeminal

Eyeball ganglion
Short ciliary Cranial

nerves Ciliary ganglion cervical ganglion
Figure 1-26. Autonomic innervation of the eye. (Modified from Getty R [1975]: Sisson and Grossman’s The
Anatomy of the Domestic Animals, 5th ed. Saunders, Philadelphia.)
Table 1-2 兩 Neuroophthalmic Reflexes and Responses

AFFERENT EFFERENT
REFLEX STIMULUS RECEPTOR NEURON INTERNEURON NEURON EFFECTOR RESPONSE
PLR Light Photoreceptors II Subcortical III Iris Constrict

Sphincter pupil
Menace Hand motion Photoreceptors II Cortical VII OOM Blink
response Cerebellum VI RBM Retract globe
XI BCM Turn
head/neck
Dazzle Bright light Photoreceptors II Subcortical VII OOM Blink
Palpebral Touch lids Touch receptors skin V (Ophth) Subcortical VII OOM Blink
Corneal Touch cornea Touch receptors cornea V (Ophth) Subcortical VII OOM Blink
VI RBM Retract globe
Doll’s eye Head motion Semicircular canals VIII Subcortical III Extraocular Maintain line
(VOR) Cerebellum muscles of sight
BCM, Brachiocephalic muscles; OOM, orbicularis oculi muscle; PLR, pupillary light reflex; RBM, retractor bulbi muscles; VOR, vestibuloocular reflex.
Menace Response
The menace response is a learned response rather than a true
reflex, and as such, it may be absent in normal young, naïve, or
stoic animals. Therefore lack of this response in such animals
should not be taken as definitive proof that they are blind. The
stimulus is hand movement across the animal’s visual field, and
care must be taken to prevent air currents from stimulating a
corneal or palpebral reflex and causing a false-positive
response. Motion parallel to the ocular surface (as opposed to
directly toward the eye) or application of the stimulus behind a
sheet of glass or plastic may prevent air currents from
stimulating the corneal reflex and causing a false-positive
Figure 1-27. The pupillary light reflex. The direct pupillary light reflex is response. Such false-positive and false-negative responses are
the response shown by the eye being illuminated. The consensual often misinterpreted by owners as evidence of sight or
pupillary light reflex is the response shown by the contralateral eye, which blindness in an animal.
is not being illuminated.
PHYSIOLOGY OF THE AQUEOUS

is important to protecting the eye, and interference with it (e.g.,
facial paralysis, trigeminal palsy, local anesthesia) often results Aqueous humor fills the aqueous compartment, which consists
in severe ocular damage. Closure of the lids of the stimulated of the anterior chamber between the iris and cornea, and the
eye is referred to as the direct corneal reflex, and closure of the posterior chamber, between the posterior iris surface and the
contralateral lids is termed the consensual corneal reflex. In anterior lens surface. The posterior chamber should not be
normal animals, the direct reflex is typically more pronounced confused with the vitreous compartment, which is located pos-
than the consensual reflex. terior to the lens (Figure 1-28).
Anterior chamber in the inner layer of the unpigmented ciliary epithelium and
may be associated with the “sodium pump” that probably accounts
Posterior chamber for the majority of actively formed aqueous.
Aqueous humor is also produced via the enzyme carbonic
anhydrase, which catalyzes the formation of carbonic acid
from carbon dioxide and water. Carbonic acid dissociates,
Vitreous
allowing negatively charged bicarbonate ions to pass to the
aqueous. Positively charged sodium ions, and eventually water,
then follow bicarbonate into the posterior chamber. Drugs that
inhibit carbonic anhydrase therefore result in decreased
aqueous production and a reduction in intraocular pressure
Figure 1-28. The chambers of the eye. The aqueous compartment is
(IOP; see later). Aqueous carries nutrients for the tissues it
subdivided into two chambers by the iris diaphragm. The anterior bathes (i.e., iris, cornea) and receives constant contributions of
chamber is anterior to the plane of the iris and pupil (blue), whereas the waste products of metabolism. Thus the composition changes
posterior chamber (green) is posterior to the iris-pupil plane but anterior as it passes from the ciliary body to the drainage angle.
to the vitreous (white). The retina and optic nerve are in yellow. (Modified Aqueous may then leave the eye via several routes. In the
from Fine S, Yanoff M [1972]: Ocular Histology: A Text and Atlas. Harper
& Row, New York.) conventional or traditional outflow route, aqueous humor
passes through the pupil into the anterior chamber, and from
there it enters the trabecular meshwork in the drainage
Formation and Composition (iridocorneal) angle, the blood-free venous angular aqueous
plexus, and eventually the systemic venous circulation through
In order to maintain the optical clarity of the eye, a series of a plexus of small veins in the sclera, the scleral venous plexus.
blood-ocular barriers are present in the normal eye. The blood- Contraction of smooth muscle fibers of the ciliary muscle that
aqueous barrier reduces aqueous humor protein concentrations insert into the trabecular meshwork are probably capable of
to about 0.5% of plasma concentrations and also prevents many increasing drainage of aqueous from the eye by increasing
substances, including drugs, from entering the aqueous. Most the size of the spaces in the trabecular meshwork. The vast
large molecules, including proteins, are unable to pass through majority of aqueous humor leaves the eye via the traditional
or between cells in the two layers of the ciliary epithelium over- outflow route.
lying the ciliary processes because of tight intercellular junctions An alternative route of drainage—the uveoscleral route—
between cells. Similarly, a blood-retinal barrier formed by the normally accounts for about 3% to 15% of aqueous outflow in
retinal capillary endothelial cells and their basement membrane most species, and probably more in the horse. Aqueous passes
also limits passage of substances into the retina so as to prevent through the ciliary body and choroid via the supraciliary
distortion of the photoreceptors. The exact anatomic location of and suprachoroidal spaces, and from there it passes through
the barrier, however, is probably different for different substances the sclera into the orbit. Outflow via this route may be sub-
(e.g., capillary endothelial cells, endothelial basement membrane, stantially increased in certain disease states and in response
or intercellular junctions). Inflammation and certain diseases to some antiglaucoma drugs, such as the prostaglandin
frequently disrupt the blood-ocular barrier and allow higher derivatives.
amounts of proteins, including immunoglobulins and fibrinogen,
to enter the eye. Pressure Dynamics
Aqueous is produced in the ciliary body, by passive
(diffusion and ultrafiltration) and active (selective transport Equilibrium between formation (production) and drainage
against a concentration gradient) processes. Fluid from ciliary (outflow) of aqueous results in a relatively constant intraocular
capillaries passes into the stroma of the ciliary processes, pressure (IOP) (Table 1-3). IOP is affected by factors such as
through the ciliary epithelium, and into the posterior chamber. age, species, mean arterial pressure, central venous pressure,
An energy-dependent transport mechanism similar to that in the blood osmolality, and episcleral venous pressure. IOP typically
renal epithelium is present in the ciliary body and results in exceeds the pressure in the draining venous system (the epi-
higher concentrations of certain substances (e.g., ascorbic acid) scleral veins) because aqueous humor is actively produced and
in the aqueous than in the plasma. Sodium and chloride ions are the trabecular meshwork slows its departure (provides
actively pumped into the aqueous and draw water passively resistance) from the eye. The ease with which aqueous humor
along a concentration gradient. Na-K–activated ATPase is present leaves the eye is called the facility of outflow (C). The various
Table 1-3 兩 Aqueous Humor Statistics

RATE OF FORMATION FACILITY OF OUTFLOW UVEOSCLERAL EPISCLERAL VENOUS
SPECIES (µL/min) (µL/min/mm Hg) OUTFLOW (%) PRESSURE (mm Hg)
Dog 2.5 0.25 (normal) 15 (normal) 10-12

0.13 (glaucoma) 3 (glaucoma)
Cat 15.0 0.19 3 8
Rabbit 4.0 0.23 13-25 9
Human 2.0 0.28 (normal) 4-14 5-15
0.18 (glaucoma)
Modified from Gum G (1991): Physiology of the eye, in Gelatt KN (editor): Veterinary Ophthalmology, 2nd ed. Lea & Febiger, Philadelphia.
features of IOP dynamics may be related to each other with the BIBLIOGRAPHY
following equation:
Budras KD, et al. (2002): Anatomy of the Dog: An Illustrated Text.
Aqueous secretion Schlütersche, Hannover, Germany.
IOP = + Episcleral venous pressure Duke-Elder S (1958, 1968): System of Ophthalmology: Vol. 1: The Eye in
Outflow facility
or Evolution, and Vol. IV: Physiology of the Eye and of Vision. Henry
Kimpton, London, pp. 605-706.
Po = (F/C) + Pe Evans HE (1993): Miller’s Anatomy of the Dog, 3rd ed. Saunders, Philadelphia.
where IOP (Po) is expressed in mm Hg, F is the rate of aqueous Jacobs GH (1993): The distribution and nature of color vision among the
mammals. Biol Rev 68:413.
formation in µL/min, C is the facility of aqueous outflow in Kaufman PL, Alm A (2003): Adler’s Physiology of the Eye, 10th ed. Mosby,
mL/min/mm Hg via both the conventional and uveoscleral out- St. Louis.
flow routes, and Pe is the episcleral venous pressure in mm Hg. Miller PE, Murphy CJ (1995): Vision in dogs. J Am Vet Med Assoc
These relationships make it clear that there are multiple 207:1623.
Miller PE, Murphy CJ (2005): Equine vision: normal and abnormal, in Gilger
avenues for decreasing IOP, including decreasing aqueous BC (editor): Equine Ophthalmology. Saunders, St. Louis, pp. 371-408.
humor secretion, increasing conventional or uveoscleral outflow, Neitz J, et al. (1989): Color vision in the dog. Vis Neurosci 3:119.
and potentially lowering episcleral venous pressure. They also Peichl L (1992): Topography of ganglion cells in the dog and wolf retina.
suggest that (1) IOP can be reduced only to 8 to 9 mm Hg J Comp Neurol 324:603.
(episcleral venous pressure) by methods that improve outflow Prince JH, et al. (1960): Anatomy and Histology of the Eye and Orbit of
Domestic Animals. Charles C. Thomas, Springfield, IL.
via the conventional pathways but that (2) in theory, improving Samuelson D, et al. (1989): Morphologic features of the aqueous humor
outflow via the uveoscleral pathway can achieve very low IOPs drainage pathways in horses. Am J Vet Res 50:720.
because this route drains into the orbit, which has a pressure of Walls GL (1963): The Vertebrate Eye and Its Adaptive Radiation. Hafner,
only 1 to 3 mm Hg. New York.
20 兩 SLATTER’S FUNDAMENTALS OF VETERINARY OPHTHALMOLOGY Chapter
DEVELOPMENT AND CONGENITAL 2

ABNORMALITIES*
Ron Ofri
DEVELOPMENT CONGENITAL ABNORMALITIES
DEVELOPMENT With the closure of the anterior end of the neural tube,
intratubular fluid accumulates and its pressure causes the
Formation of Optic Primordia evagination of the optic grooves and their transformation into
Broadly speaking, the embryonic and fetal development of the the two optic vesicles (Figures 2-3 and 2-4). This marks the
eye occurs in three stages: beginning of organogenesis. In the dog this event occurs on the
fifteenth day of gestation. The lumen of the neural tube remains
• Embryogenesis: Segregation of the primary layers of the
connected to the cavities of the optic vesicles by two optic
developing embryo. The period begins with fertilization and
stalks (see Figure 2-4, D). Under the pressure of the
ends with differentiation of the primary germ layers.
intratubular (intraventricular) fluid, the optic vesicles continue
• Organogenesis: Separation into the general pattern of
to enlarge and bulge, eventually coming in contact with the
various organs.
surface ectoderm. At the site of contact with the optic vesicle
• Differentiation: Detailed development of the characteristic
the surface ectoderm thickens and forms the lens placode
structure of each organ.
(Figures 2-3; 2-4, C; and 2-5). The contact of the optic vesicle
It is assumed that the embryonic development of the eye is with the surface ectoderm serves as an induction for the optic
similar in sequence for all mammalian species and that inter- vesicle to start invaginating, thus forming the double-layered
species differences pertain mostly to the duration of gestation optic cup (see Figures 2-3 and 2-4, E and F).
and the age of the various anatomic end points—for example, The invagination of the vesicle progresses from inferior to
regression of embryonic vasculature or eyelid opening. superior but is not completed on the ventral side of the optic
The optic primordia (rudimentary eye) develops from that cup, where a fissure, called the embryonic optic fissure, remains.
portion of the embryo that later forms the anterior part of the The double layers of the optic cup are aligned on both sides
central nervous system (CNS). The first step in the embryogenesis of the fissure, which extends posteriorly under the optic stalk.
of the future eye takes place at the embryonic plate stage, when This fissure allows the secondary mesenchyme present around
the ectoderm invaginates along the posterior-anterior axis to the cup to penetrate into the cavity of the optic cup to form the
form the neural groove. The two neural lips of the groove hyaloid vascular system (twenty-fifth day of gestation in the
subsequently fuse, thus turning the groove into the neural tube dog) (see Figure 2-4, G and H). This fissure gradually closes
(Figures 2-1 and 2-2). At the site of fusion, between the ecto- leaving a small aperture at the anterior end of the optic stalk,
derm and the neuroepithelium (the epithelium of the neural through which the hyaloid artery passes (Figure 2-6). The
tube), neuroepithelial cells proliferate to form the neural crest hyaloid artery supplies the inner layers of the optic cup and
cells and migrate sideways into the paraxial and lateral meso- developing lens vesicle. The two lips of the optic fissure fuse
derm. These neural crest cells mix with mesodermal cells and anteriorly to the optic stalk. The fusion process progresses
form the secondary mesenchyme; this secondary mesenchyme anteriorly and posteriorly, eventually causing closure of the
forms the main mesodermal structures of the eye. Therefore the optic cup and allowing intraocular pressure to build up (see
eye develops from the neural ectoderm, neural crest, and surface Figure 2-6).
ectoderm, with minor contributions from the mesoderm. The lens placode thickens to become the lens vesicle.
The anterior end of the neural tube enlarges and bends down Following the invagination of the optic vesicle, the lens vesicle
to form the primordia of the CNS. On its outer surface, on both finds itself embedded inside the cavity of the cup (see Figure 2-4,
sides, appear two small pits called the optic grooves or optic F and H). Anteriorly, the hyaloid artery gives branches, the
pits. These pits, which appear on day 13 of gestation in the dog, tunica vasculosa lentis, which cover the posterior and lateral
are the anlage of the eyes (see Figure 2-2, G). faces of the lens. This vascular network supplies the metabolic
requirements of the lens during development. The hyaloid
vascular system disappears at advanced stages of the develop-
*The author wishes to acknowledge the contribution of Dr. Robert Barishak, ment or during the postnatal period. In dogs remnants of the
and to thank him for his input throughout the years and to this chapter. hyaloid system might remain visible until the fourth postnatal
20
DEVELOPMENT AND CONGENITAL ABNORMALITIES 兩 21
Optic vesicle Optic cup Lens vesicle
Neural crest
Ectoderm
Notochord Mesoderm
Endoderm
Figure 2-3. Formation of the optic vesicle, optic cup, and lens vesicle.
Figure 2-1. Developing embryo at the start of neural tube formation. (From Yanoff M, Duker J [2004]: Ophthalmology, 2nd ed. Mosby,
(From Yanoff M, Duker J [2004]: Ophthalmology, 2nd ed. Mosby, St. Louis.)
St. Louis.)
Cut edge of
amnion
Level of
Neural fold section B
Neural groove
Somite
Primitive node
A Primitive streak
E
Ectoderm
Mesoderm
B Endoderm
Surface ectoderm
Neural crest
Mesoderm
Neural tube of
F neural ectoderm
C
Optic pit
Neural folds
D G
Figure 2-2. Formation of neural tube. A, Dorsal surface of embryo as seen from above. B, Vertical section
through three-layered embryonic disc. C, Neural groove forms in neural plate area of ectoderm. D, Neural
groove invaginates and neural folds are formed. E, Neural folds continue to grow toward each other. F, Neural
crest cells separate from ectoderm of neural folds as the folds fuse; neural tube is formed (of neural ectoderm);
and surface ectoderm is again continuous. G, Evaginations in area of forebrain form the optic pits. (From
Remington LA [2005]: Clinical Anatomy of the Visual System, 2nd ed. Butterworth-Heinemann, St. Louis.)
month, whereas in cattle they may persist until 12 months of age. is reviewed in the following sections, beginning with the
In humans, but not in domestic animals, the caudal portions of posterior parts of the eye and progressing anteriorly.
the hyaloid artery and vein transform themselves into the
central retinal artery and vein. Retina
At this stage of development, organogenesis has been
completed and the general structure of the eye has been The optic cup is lined by two layers of epithelium of neuro-
determined. It is followed by a period of differentiation as the ectodermal origin (see Figure 2-4, H). The inner layer, facing
specific structures of the eye begin to form. Their development the vitreous, is nonpigmented, but the outer layer, facing the
Optic groove
Level of section B Optic groove Neural fold
Neural fold
Mesenchyme
Neural groove
A B Surface ectoderm
Neural tube
Notochord Optic stalk

Lens placode
Forebrain
Mesenchyme Lens pit
Lens placode
Surface ectoderm
Optic vesicle
C D Early stage of optic cup
Mesenchyme
Midbrain
Surface ectoderm Outer layer of optic cup
Inner layer
of optic cup
Forebrain Lens vesicle
Optic fissure
Hyaloid artery
Level of section G
E Optic cup F
Hyaloid vein
Optic fissure
Hyaloid artery
Lumen of optic stalk Lens vesicle
Mesenchyme
Wall of brain
Hyaloid artery
Intraretinal space
Hyaloid vein in
G optic fissure H
Figure 2-4. Early eye development. A, Dorsal view of the cranial end of 22-day embryo, showing the first
indication of eye development. B, Transverse section through neural fold, showing an optic groove. C, Forebrain
and its covering layers of mesenchyme and the surface ectoderm from an embryo of about 28 days. D, F, and H,
Sections of the developing eye, illustrating early stage in the development of the optic cup and lens vesicle.
E, Lateral view of the brain of an embryo at about 32 days, showing the external appearance of the optic cup.
G, Transverse section through the optic stalk, showing the optic fissure and its contents. (From Remington LA
[2005]: Clinical Anatomy of the Visual System, 2nd ed. Butterworth-Heinemann, St. Louis.)
(future) sclera, is pigmented. The anterior rim of the cup will The outer epithelial layer of the posterior optic cup forms
form the anterior uvea (ciliary body and iris), and the posterior the pigment epithelium of the retina. The inner epithelial layer
part of the cup will form the retina. The two epithelial layers of forms the sensory retina (Figure 2-7). The two layers of the
the optic cup will form the two epithelial layers of the retina posterior optic cup are separated by the intraretinal space
and the anterior uvea. representing the cavity of the optic vesicle, which has gradually
Neural and an inner neuroblastic layer (see Figure 2-7). These outer
ectoderm Retinal disc and inner neuroblastic layers are separated by the fiber layer of
Surface
Mesenchyme ectoderm Lens placode
Chievitz. The cells of the outer neuroblastic layer differentiate
into cones and rods externally and horizontal cells internally.
The cells of the inner neuroblastic layer differentiate into
ganglion cells, amacrine cells, bipolar cells, and Müller’s cells.
The rods and cones (i.e., the photoreceptors) form the outer
retina and are adjacent to the choroid and sclera (Figure 2-8).
The ganglion cell layer, which originated in the inner neuro-
blastic layer, is called the inner retina because it is adjacent to
the vitreous. The resulting retina is called an inverted retina
because the photoreceptors are close to the outer layers of the
eye and light must pass through all of the retinal layers to reach
the photoreceptors. The reason for this arrangement is to place
the photoreceptors next to the choroid, thus giving these cells,
which have very high metabolic requirements, their own
“private” blood supply.
There is species variation in the degree of retinal development
present at birth. In dogs it is possible to record electrophysiologic
activity of the photoreceptors during the first postnatal week;
signals reach adult amplitude by 5 to 8 weeks of age. Similarly,
Figure 2-5. Light micrograph of 6-mm pig embryo showing thickening of
differentiation of the rod and cone inner and outer segments is
lens placode. (From Remington LA [2005]: Clinical Anatomy of the Visual evident histologically during the first 8 weeks of life.
System, 2nd ed. Butterworth-Heinemann, St. Louis.)
Optic Nerve
been obliterated during the invagination of the optic vesicle.
Diseases of the posterior segment of the eye cause retinal Axons from the ganglion cells grow toward the optic stalk, thus
detachment in this space as the sensory retina separates from forming the nerve fiber layer, the innermost layer of the retina
the pigment epithelium. (see Figures 2-6, B and D; and 2-8). Axons from throughout the
The common neuroblastic layer, which is the nuclear portion entire retina converge on the optic disc, where they form into
of the sensory retina, divides into an outer neuroblastic layer bundles collectively known as the optic nerve (cranial nerve II)
Lumen of optic stalk

Lens
Inner layer of optic stalk

Hyaloid vessels (containing axons of
in optic fissure ganglion cells)
Mesenchyme
B
Level of section B
A
Optic stalk
Walls of optic stalk continuous
with the wall of the brain and
the layers of the optic cup
Lens
Axons of ganglion cells

Optic fissure closed
Hyaloid vessels
Level of section D D Optic fissure closing

C
Hyaloid vessels
in optic fissure
Figure 2-6. Closure of optic fissure. A and C, Views of inferior surface of optic cup and stalk, showing
progressive stages in closure of optic fissure. B and D, Transverse sections through optic stalk, showing successive
stages in closure of optic fissure. Note that the lumen of optic stalk is obliterated gradually. (From Remington LA
Vitreous
The embryonic vitreous consists of the primary, secondary, and
tertiary vitreous (Figure 2-11). The primary vitreous develops
with the hyaloid vasculature (Figure 2-12). It has
mesenchymal, neuroectodermal, and ectodermal components.
The mesenchymal elements enter posteriorly with the hyaloid
vessels, and anteriorly through the space between the anterior
Deeper, Inner, rim of the optic cup and the lens vesicle. The ectodermal
nucleated nonnucleated elements are the fibrils produced by the posterior face of the
layer layer lens. The primary vitreous also contains neuroectodermal
elements, which consist of the fibrils produced by the inner
limiting membrane of the retina. The secondary vitreous is the
Figure 2-7. The two retinal walls. The inner retinal wall shows two
zones—an inner, nonnucleated layer and a deeper, nucleated layer. (From
“definitive” vitreous that will persist into adulthood. It is
Yanoff M, Duker J [2004]: Ophthalmology, 2nd ed. Mosby, St. Louis.) denser, is more homogeneous and avascular, and is laid down
around the primary vitreous (see Figures 2-11, B, and 2-13). It
is also secreted by the inner limiting membrane of the retina.
(Figure 2-9). The axons of optic nerve extend posteriorly to The tertiary vitreous is secreted by the ciliary epithelium.
form the optic chiasm and optic tracts before making their first Bundles of fibers extend from the ciliary epithelium toward
synapse at the lateral geniculate nucleus. the lens equator, covering the secondary vitreous anteriorly
As the ganglion cell axons collect at the optic disc they dis- (see Figure 2-11, C). In the adult they persist as lens zonules
place primitive neuroectodermal cells forward, into the vitreous (suspensory ligament of the lens).
cavity. These displaced cells form a glial sheath around the
hyaloid artery. At the disc, the same cells may form an Lens
agglomeration called Bergmeister’s papilla, which protrudes
into the vitreous. The papilla may persist into adult life As noted earlier, thickening of the lens placode (on the
(especially in ruminants), or it may atrophy, thus forming a seventeenth day of gestation in the dog) occurs as a result of
depression, known as the physiologic optic cup, in the optic induction by the optic vesicle. The placode then invaginates,
disc (Figure 2-10). In newborns this physiologic optic cup may and by day 25 it forms the lens vesicle (see Figures 2-3 and
be confused with a coloboma of the optic disc. In patients with 2-4, D, F, and H). This vesicle is lined by surface ectodermal
glaucoma the physiologic optic cup may enlarge owing to the cells, the apex of which is directed toward the center of the lens
forces of the increased intraocular pressure on this region of vesicle cavity (Figures 2-14 and 2-15). The base of the cells
the eye. forms the primitive lens capsule. The anterior cells of the lens
Retinal pigment
epithelium
Photoreceptor
A layer
External limiting
membrane
Sclera
Outer nuclear layer
Choroid Outer plexiform layer
Inner nuclear layer

Retina
Inner plexiform layer

Ganglion cell
layer
Nerve fiber layer
Internal limiting
membrane
Figure 2-8. Photomicrographs of the posterior section of the eye. A, The three posterior layers of the eye. The
sclera is the outermost layer, whereas the retina faces the vitreous. B, An enlargement of the 10 layers of
the retina. The retinal pigment epithelium and the photoreceptors are the outermost layer of the retina, facing the
choroid. The ganglion cell layer and nerve fiber layer are the innermost layers of the retina, facing the vitreous.
(From Remington LA [2005]: Clinical Anatomy of the Visual System, 2nd ed. Butterworth-Heinemann, St Louis.)
Ganglion cell
layer of the retina
Axons of ganglion cells Sheath of the optic nerve (continuous

with the meninges of the brain and the
choroid and sclera)
A
Optic stalk
Central
Lens artery and
vein of the retina
Optic nerve Axons of ganglion cells
Level of section C C Optic fissure closed
B Central vein and artery of the retina

Figure 2-9. Formation of optic nerve. A, Axons of ganglion cells of the retina grow through optic stalk to brain.
B, Transverse section through the optic stalk, showing the formation of the optic nerve. The optic nerve is formed
after closure of the optic fissure (this stage follows the stage shown in Figure 2-6, C). C, The lumen of the optic
stalk is obliterated as axons of ganglion cells accumulate in the inner layer of the stalk. (From Remington LA
Figure 2-10. Optic nerve

heads of a dog (A) and a
sheep (B). The dark spot in
the center of the canine optic
nerve head is the physiologic
optic cup (arrow). The pink
tuft in the center of the sheep
optic nerve head is the ovine
Bergmeister’s papilla (arrow).
(Courtesy University of
California, Davis, Veterinary
Ophthalmology Service Col-
lection.)
A B
vesicle remain cuboidal, but the posterior cells elongate, become life from those equatorial epithelial cells that maintain their
columnar, and form the primary lens fibers (Figure 2-16). lifelong mitotic activity. Successive layers of fibers are
These fibers extend anteriorly, thus filling the cavity of the deposited on top of preexisting fibers, like the layers of an
vesicle with lens fibers (see Figures 2-14, D and E; and 2-16, C onion. As a result the embryonal nucleus is surrounded by the
and D). Their nuclei disappear, and these fibers constitute the fetal nucleus, which in turn is surrounded by the adult nucleus
embryonal nucleus of the lens. As a result, the posterior aspect and cortex (Figure 2-17).
of the adult lens is devoid of cells and is composed only of a Because none of the lens fibers is quite long enough to reach
lens capsule. The anterior cuboidal cells, on the other hand, fully from pole to pole, and because the cells are too thick at the
remain as the adult lens epithelium. ends for all to meet in a single point, they meet in a Y-shaped
The junction between the anterior lens epithelium and structure known as the lens suture. The anterior lens suture is
the primary lens fibers extends along the equator of the lens an upright Y, and the posterior suture is inverted (Figures 2-16,
and forms the equatorial zone. Epithelial cells in this area F and G; and 2-18).
form the secondary lens fibers; these fibers extend anteriorly The lens capsule is secreted anteriorly by the anterior lens
along the lens epithelium and posteriorly along the lens epithelium. Its formation continues throughout life, and there-
capsule. Secondary lens fibers continue to form throughout fore its thickness increases with age. The posterior lens capsule,
Primary vitreous Secondary vitreous
Primary vitreous
(hyloid artery)
A
Primary Primary Secondary
vitreous vitreous vitreous
Remains of primary vitreous Figure 2-13. Vitreous development. The primary vitreous and hyaloid
(Cloquet’s canal) artery fill the optic cup. The primary vitreous retracts and the hyaloid
artery regresses, while the secondary avascular vitreous develops. (From
Yanoff M, Duker J [2004]: Ophthalmology, 2nd ed. Mosby, St. Louis.)
Secondary (adult) vitreous
B
Presumptive
A
fibers
Tertiary vitreous
(lens zonules)
B
Remains of primary
vitreous (Cloquet’s canal)
C
Secondary (adult)
vitreous C
Elongating
Figure 2-11. Scheme of main features in vitreous development and posterior
regression of hyaloid system, shown in drawings of sagittal sections. epithelium
A, Hyaloid vessels and branches occupy much of the space between lens D
and neural ectoderm, forming the primary vitreous. B, An avascular
secondary vitreous of fine fibrillar composition fills the posterior part of
the eye. The primary vitreous shown in A is condensed into Cloquet’s Lens
canal as the hyaloid vessels atrophy. C, Vessels of hyaloid system atrophy fibers
progressively. Zonular fibers (tertiary vitreous) begin to stretch from
growing ciliary region toward lens capsule. (Modified from Duke-Elder S E
[editor] [1963]: System of Ophthalmology. Vol III: Normal and Abnormal
Development, Part 1. Embryology. Henry Kimpton, London.) Figure 2-14. A, Formation of lens placode. B, Invagination forming lens
vesicle. C to E, Development of embryonic nucleus. C, Hollow lens vesicle
is lined with epithelium. D, Posterior cells elongate, becoming primary
lens fibers. E, Primary lens fibers fill lumen, forming embryonic nucleus.
Tunica Curved line formed by cell nuclei is called the lens bow. Anterior epi-
vasculosa thelium remains in place. (From Remington LA [2005]: Clinical Anatomy
lentis of the Visual System, 2nd ed. Butterworth-Heinemann, St. Louis.)
which is much thinner, is formed by the basal membrane of the

Hyaloid
elongating primary lens fibers.
artery
Primitive Vascular System
The hyaloid artery, a branch of the internal ophthalmic artery,
enters the optic cup through the embryonal optic fissure (see
Figure 2-6, A). Its branches continue forward, reach the anterior
margin of the optic cup, and anastomose with the annular vessel
Figure 2-12. Light micrograph of 25-mm pig embryo showing the hyaloid (see Figure 2-6, C) formed by the choriocapillaris, which is
arterial system filling future vitreal cavity. Vessels are evident extending
through the optic stalk, and the vascular network attached to posterior the capillary plexus that surrounds the optic cup. The hyaloid
lens is evident. (Modified from Remington LA [2005]: Clinical Anatomy of artery is called vasa hyaloidea propria; its anterior dividing
the Visual System, 2nd ed. Butterworth-Heinemann, St. Louis.) branches form a net around the lens called the lateral and
Lens
vesicle
cavity
Posterior A B
epithelial
cells
elongating
into primary
lens fibers
Figure 2-15. Light micrograph of 15-mm pig embryo showing lens vesicle
filling with primary lens fibers; lens bow configuration is evident. (Modified
from Remington LA [2005]: Clinical Anatomy of the Visual System, 2nd
ed. Butterworth-Heinemann, St. Louis.) C D
posterior tunica vasculosa lentis (Figure 2-19). The anterior

tunica vascuolosa lentis is formed by branches from the
annular vessel. These tunicas are responsible for vascular
supply to the lens during embryonic development. During
the last stages of embryologic development or soon after
birth, as the aqueous humor takes over this metabolic function,
the hyaloid vasculature atrophies, regresses, and is replaced by E F
the pupillary membrane (see also next section, Ciliary Body
and Iris).
The mesenchyme around the optic cup forms the choroid,
which surrounds the choriocapillaris. The nasal and temporal
long posterior ciliary arteries branch off the ophthalmic artery
and advance forward in the horizontal plane through the choroid
to supply the future ciliary body. At the level of the ciliary body
they anastomose to form the major vascular circle of the iris.
Posteriorly, the short ciliary arteries arrange themselves around
the entrance of the optic nerve into the globe (Figure 2-20). The
vessels form an anastomosing plexus, called the Haller-Zinn
vascular circle, which plays a role in the vascular supply of the G H
optic nerve head. Figure 2-16. Stages of development of the lens. A, Elongation and
anterior growth of posterior cuboidal epithelial cells to form primary lens
Ciliary Body and Iris fibers. B, Elongation of primary lens fibers to fill the cavity in the lens
vesicle, and formation of the lens bow of cuboidal cell nuclei. C and D,
The adult ciliary body is lined with two layers of epithelium of Secondary lens fibers proliferate from the equatorial region of the lens,
covering the primary lens fibers and scattered cuboidal cell nuclei. E, The
neuroectodermal origin. The inner layer, close to the vitreous, adult lens. F and G, Appearance of the Y sutures. H, New layers of
is unpigmented. This layer is the anterior extension of the secondary lens fibers are laid down around the central primary lens fibers.
sensory neuroretina, though it contains no neural elements. Growth continues throughout life. (A modified from Severin GA [2000]:
The outer layer is pigmented and is the anterior continuation of Severin’s Veterinary Ophthalmology Notes, 3rd ed. Severin, Ft. Collins,
CO.)
the retinal pigment epithelium. The ciliary epithelium forms
folds called ciliary processes (Figure 2-21). These processes
are the production site of the aqueous humor; they also serve as derived from the neuroectoderm of the anterior rim of the optic
the anchoring site of the lens zonules, which suspend the lens cup. The iris stroma originates from the neural crest–derived
in the eye. The underlying ciliary muscle and stroma of the secondary mesenchyme (see Figure 2-21). This mesenchyme
ciliary body originate from the neural crest–derived secondary continues to grow over the anterior part of the lens, covering
mesenchyme; the power of the muscle’s contraction and relax- the hole that will be the future pupil with a membrana pupil-
ation is transferred through the ciliary processes and zonules to laris, which replaces the anterior tunica vasculosa lentis at the
the lens, changing its refraction and the focusing of the eye. same location (see Figures 2-19 and 2-22). This membrane
The anterior rim of the optic cup forms the iris, which has contains branches from the major arterial circle of the iris that
two pigmented epithelial layers on its posterior face. These form a vascular net over the iris and the pupil. However, strands
epithelial layers are continuous with the two layers of the of the membrana pupillaris may remain attached to the anterior
epithelium of the ciliary body. The sphincter and dilator smooth surface of the iris. These strands, known as persistent pupillary
muscles of the mammalian iris, which control the constriction membranes, are inherited as a homozygous recessive trait in the
and dilation of the pupil through their antagonistic actions, are basenji.
Anterior capsule Major circle of the iris

Anterior
Cortex epithelium
Adult nucleus Anterior ciliary artery

Long posterior
Equator ciliary artery
Fetal nucleus
Muscular artery
Embryonal n.
to lateral rectus
muscle
Equatorial
epithelium
transforming
into secondary Choroidal blood Muscular vein
lens fibers vessels
Retinal blood vessels
Posterior capsule
Figure 2-17. Adult lens showing the successive layers of the lens that are Vortex vein
laid around the embryonal nucleus throughout life. (From Remington LA
[2005]: Clinical Anatomy of the Visual System, 2nd ed. Butterworth- Long posterior ciliary artery
Heinemann, St. Louis.) Short posterior ciliary arteries
Central retinal artery
Central retinal vein
Elongation of posterior epithelium Secondary lens fiber migration Figure 2-20. Horizontal section of the eye showing ciliary circulation.
(From Remington LA [2005]: Clinical Anatomy of the Visual System, 2nd
ed. Butterworth-Heinemann, St. Louis.)
Cornea and Anterior Chamber

The outer corneal epithelium derives from the surface ecto-
derm, but the inner layers, which include the corneal stroma
and corneal endothelium, derive from the secondary mesen-
chyme. Descemet’s membrane is secreted by the endothelial cells
(see Figure 2-22). Further ingrowth of secondary mesenchyme
occurs between the epithelium and endothelium, forming the
corneal stroma.
Figure 2-18. Lens embryogenesis. Left, Elongation of the posterior
epithelium results in obliteration of the lens lumen. Right, Secondary lens Between the cornea and the lens two spaces develop: the
fiber migration leads to the formation of the Y sutures. (From Yanoff M, posterior chamber between the iris and the lens, and the anterior
Duker J [2004]: Ophthalmology, 2nd ed. Mosby, St. Louis.) chamber between the iris and the cornea (Figure 2-23). After
the regression of the pupillary membrane, aqueous may flow
from the posterior chamber to the anterior chamber through the
pupil.
Anterior vascular capsule

(pupillary membrane)
Sclera and Extraocular Muscles
Neural crest–derived mesenchyme surrounds the optic cup and
forms two layers. The inner layer, which is adjacent to the
retina, is the choroid, and the outer layer is the sclera. Con-
densation of the sclera begins anteriorly, near the ciliary body,
and proceeds posteriorly to the optic nerve, where it is continu-
Posterior ous with the dura mater of the optic nerve. Extraocular muscles
vascular
capsule form in the neural crest–derived secondary mesenchyme of
the orbit.
Capsulopupillary
portion
Eyelids and Third Eyelid
The lower eyelid and the third eyelid are formed by the
maxillary process. The upper eyelid is formed by the paraxial
Hyaloid artery
mesoderm. During development the upper and lower eyelids
are fused (Figure 2-24). With time, these fused eyelids separate,
Figure 2-19. Hyaloid vasculature and primary vitreous during embryo- although the age at which separation occurs varies among
logic ocular development. (From Yanoff M, Duker J [2004]: Ophthalmology, species. In horses, cattle, sheep, and pigs, lids open at 7 to 10 days
2nd ed. Mosby, St. Louis.) postpartum. During the formation of the eyelids their inner surface
Cornea
Conjunctiva
Outer, pigmented Iris sphincter
epithelium Iris stroma Iris
Ciliary body
Ciliary processes
Sclera Lens
Posterior,
pigmented Figure 2-21. A diagram (A) and a photomicrograph (B) of the
iris epithelium periphery of the anterior segment of the eye. The nonpigmented
epithelium is the innermost layer of the ciliary body, facing the
Inner, nonpigmented Zonules vitreous, and is continuous with the sensory retina (not shown).
epithelium
A The pigmented epithelium is the outer layer, facing the sclera, and
is continuous with the retinal pigment epithelium (not shown).
These two epithelial layers continue anteriorly as the pigmented
epithelium on the posterior aspect of the iris. The zonules (which
are the tertiary vitreous) suspend the lens from the ciliary
processes, and their remnants can be seen in B. (Modified from
Remington LA [2005]: Clinical Anatomy of the Visual System, 2nd
ed. Butterworth-Heinemann, St. Louis.)
Future iris Optic cup Corneal epithelium Fused Anterior Posterior

Cornea lids chamber chamber
4 mm
Hyaloid
artery
Future cornea
Anterior
Lens chamber
Descemet
endothelium Retina
(mesothelium)
Pupillary
Posterior membrane
chamber
Mesoderm
Figure 2-22. Formation of the anterior chamber and cornea. Note the
pupillary membrane, which replaced the anterior tunica vasculosa lentis,
Figure 2-23. Section through eye and orbit of 48-mm human embryo
covering the future pupil.
(approximately 9.5 weeks). (Modified from Remington LA [2005]: Clinical
Anatomy of the Visual System, 2nd ed. Butterworth-Heinemann,
St. Louis.)
(and the anterior sclera) is lined with palpebral conjunctiva tarsus and dermis of the lids, but mesoderm contributes to the
derived from the surface ectoderm. This ectoderm also contri- formation of eyelid muscles.
butes to the formation of lid epidermis, cilia, and a number of
glands: the lacrimal and nictitating glands, which produce the Nasolacrimal System
aqueous portion of the tear film; the tarsal meibomian glands,
which produce the lipid component of the tear film; and Zeiss The nasolacrimal groove separates the lateral nasal fold from
(sebaceous) and Moll (sweat) glands. Neural crest–derived the maxillary processes. At the bottom of the groove a solid
secondary mesenchyme contributes to the development of the cord of ectodermal cells forms and gets buried as the maxillary
including microphthalmia, cataract, retinal dysplasia, anterior

segment dysgenesis, and optic nerve hypoplasia. It is only by
Cornea Fused detailed study of anomalies that etiology and pathogenesis
eyelids
of the lesions are understood, and subsequent diagnosis and
evaluation simplified.
Anterior
chamber
The stage of development (e.g., embryogenesis, organogenesis,
or fetal period) at which a teratogenic factor acts is most
Iris
Extraocular important in determining the final effects on the eye.
muscles
In this section common congenital abnormalities of the
whole eye of domestic animals are considered. Abnormalities
of the individual parts of the eye are discussed in the relevant
chapters.
Anophthalmos and Microphthalmos

Anophthalmos means the total absence of an eye. It may be
Figure 2-24. Light micrograph of 45-mm pig embryo. Eyelids are fused, caused by the suppression of the optic primordia during the
extraocular muscle is evident, and axons are evident in optic nerve. development of the forebrain, may be caused by the abnormal
(Modified from Remington LA [2005]: Clinical Anatomy of the Visual
System, 2nd ed. Butterworth-Heinemann, St. Louis.)
development of the forebrain, or may be due to the degeneration
of the optic vesicles after they have already formed as a result
of a teratogenic insult.
process grows over it to fuse to the lateral nasal fold. Two True anophthalmos is very rare, and its diagnosis is made
ectodermal buds grow from the proximal end of the buried cord after histologic examination of the orbital contents has not shown
toward the upper and lower lid folds near the nasal canthus. These the presence of any ocular structure. Most instances of presumed
buds form the superior and inferior lacrimal puncta. The distal end clinical anophthalmos are cases of extreme microphthalmos,
of the cord enters the ventral nasal meatus. The entire cord because some histologic evidence of a rudimentary eye can
becomes the nasolacrimal duct by a process of canalization. usually be found.
Incomplete canalization is common in domestic animals, resulting Microphthalmos is an eye that is smaller than normal
in obstruction of the tear drainage. In dogs the puncta and the (Figure 2-25). Microphthalmos is most frequent in pigs and
upper half of the nasolacrimal duct are most commonly affected.
In horses the nasal meatus of the duct may be imperforate.
CONGENITAL ABNORMALITIES
Teratology is the branch of embryology that deals with abnor-
mal development and congenital malformations. It is important
to remember that not all congenital abnormalities are necessarily
inherited, as some may result from toxicity or disease during
development. Conversely, not every inherited abnormality is
necessarily congenital. Many inherited disorders (e.g., cataract, A
progressive rod cone degeneration) may be manifested later
in life.
The most important determining aspect of the character of a
deformity is the stage of development at which the etiologic
agent acts. Factors acting during the early period of embryo-
genesis are generally lethal. Those occurring during organogenesis
result in gross deformities affecting the whole eye (e.g.,
anophthalmia, microphthalmia, and cyclopia). If the factor acts
during the fetal period, when the most fundamental and active
stage has been completed, minor defects of individual parts of
the eye caused by arrests in development and associated
deformities due to aberrant growth may occur. Because much
of the development and differentiation of the eye occurs very
early in gestation (during the first 2 weeks in the dog), events
initiated during this time may result in malformations in struc- B
tures that do not fully mature until much later. Studies of the Figure 2-25. A, Bilateral microphthalmia as part of a multiple ocular
effect of exposure to teratogens on ocular development have defects syndrome, which also includes developmental defects in the iris,
identified narrow, critical periods for induction of malformations; lens, retina and embryonic hyaloid apparatus. B, Close-up of the right eye
of the same dog, highlighting the microphthalmia and iris abnormalities.
for example, exposure during gastrulation (formation of the (Courtesy University of California, Davis, Veterinary Ophthalmology
mesodermal germ layer) results in a spectrum of malformations Service Collection.)
Table 2-1 兩 Anomalies Associated with Microphthalmos In cyclopia the prosencephalon does not show cleavage;
in Dogs there is one midbrain, one dorsal cyst, and a single optic nerve
and optic canal. The frontonasal process presents a proboscis
ANOMALY DOG
(displaced nose) above the single orbit. The lids of the two eyes
Anterior segment dysgenesis Saint Bernard are fused around the single orbit. Cyclopia has been reported,
Doberman in Idaho and Utah, in lambs born to ewes that grazed on Veratrum
Cataract Old English sheepdog californicum on the fourteenth day of gestation and, in Western
Miniature schnauzer
Cavalier King Charles
Australia, in lambs born to ewes that grazed on unknown toxic
spaniel plants.
Persistent hyperplastic primary vitreous Irish wolfhound
Retinal dysplasia Saint Bernard
Doberman
Coloboma
Modified from Cook C (1995): Embryogenesis of congenital eye Coloboma is a condition in which a portion of the eye, usually
malformations. Vet Comp Ophthal 5:110. a portion of the uvea, is lacking. Most colobomas (typical
colobomas) are due to an incomplete closure of the embryonic
optic fissure (Figure 2-27). These colobomas are usually
dogs. In pigs, vitamin A deficiency in the dam is the most situated in the inferonasal portion of the eye. The extent of the
common cause. In dogs, microphthalmos occurs frequently as coloboma may vary. Severe colobomas are associated with the
part of the collie eye anomaly. Administration of griseofulvin formation of an orbital cyst (microphthalmos with orbital cyst),
to pregnant cats for treatment of dermatomycosis has resulted because the optic fissure failed to close and form a vesicle.
in anophthalmos or microphthalmos in their kittens. In white Moderate colobomas may involve numerous ocular structures,
shorthorn cattle hereditary microphthalmos is associated with whereas mild cases may manifest as only a simple notch in the
large lids and third eyelid, resulting in entropion because the lower nasal quadrant of the pupil.
small globe does not support the elongated lids. Atypical colobomas are not associated with the incomplete
Microphthalmos may occur in eyes that are otherwise closure of the embryonic fissure and are not located in the
(functionally) normal, if all the internal eye structures remain lower nasal quadrant. They are usually due to lack of induction
proportional in size. It may also occur in eyes with multiple of one tissue by another. For example, lack of induction by
ocular anomalies, including cataract, retinal dysplasia, and retinal pigment epithelium may cause colobomas in the choroid
anterior segment dysgenesis (Table 2-1). In Jersey calves, an and sclera, whereas lack of induction by the anterior rim of the
autosomal recessive condition causes congenital blindness with optic cup may result in aniridia, or lack of iris.
microphthalmos, aniridia (lack of iris), microphakia (small Colobomas of the optic nerve head may be seen in dogs
lens), ectopia lentis (malpositioned lens), and cataract. Lambs affected with collie eye anomaly (Figure 2-28) and in basenjis
grazing on seleniferous pasture in Wyoming were afflicted with that present with persistent pupillary membranes. In cats,
microphthalmos, ectopia lentis or aphakia, optic nerve hypo-
plasia, persistent pupillary membrane, uveal coloboma, and
nonattachment of the retina. In Hereford cattle an encephalopathy- Mesodermal vascular tissue
microphthalmos syndrome is inherited as a simple autosomal
recessive hereditary trait. Animals present with a domed skull, Retina
degeneration of skeletal muscles, small palpebral fissures, small
orbits, retinal dysplasia, vitreous syneresis, microphakia, and
bilateral microphthalmos.
Cyclopia and Synophthalmus

In cyclopia there is a single eye. In synophthalmus the eyes are
fused in the midline (Figure 2-26). These conditions are
incompatible with life. Pigment epithelium
Retina
Pigment epithelium
Figure 2-27. The closure of the fetal cleft. The margins come together
accurately (top), but subsequently an excessive growth of the inner
(retinal) layer leads to its eversion (bottom), causing a coloboma of the
Figure 2-26. Cyclopia in lambs whose dam grazed on Veratrum retina and posterior uvea. (Modified from Duke-Elder S [editor] [1963]:
californicum. (Courtesy University of Wisconsin–Madison Veterinary System of Ophthalmology, Vol III: Normal and Abnormal Development.
Ophthalmology Service Collection.) Part 2: Congenital Deformities. Henry Kimpton, London.)
Bistner SI, et al. (1973): Development of the bovine eye. Am J Vet Red 34:7.
Boroffka SA (2005): Ultrasonographic evaluation of pre- and postnatal
development of the eyes in beagles. Vet Radiol Ultrasound 46:72.
Collinson JM, et al. (2004): Analysis of mouse eye development with
chimeras and mosaics. Int J Dev Biol 48:793.
Cook C (1995): Embryogenesis of congenital eye malformations. Vet Comp
Ophthal 5:110.
Cook C, et al. (1993): Prenatal development of the eye and its adnexa, in
Tasman Jaeger W (editor): Duane’s Foundations of Clinical
Ophthalmology. JB Lippincott, Phildelphia, p. 1.
Cook CS (1989): Experimental models of anterior segment dysgenesis.
Ophthalmic Paediatr Genet 10:33.
Cvekl A, Tamm ER (2004): Anterior eye development and ocular
mesenchyme: new insights from mouse models and human diseases.
Bioessays 26:374.
Duddy JA, et al. (1983): Hyaloid artery patency in neonatal beagles.
Am J Vet Res 44:2344.
Gould DB, et al. (2004): Anterior segment development relevant to glaucoma.
Int J Dev Biol 48:1015.
Gum GG, et al. (1984): Maturation of the retina of the canine neonate as
determined by electroretinography and histology. Am J Vet Res 45:1166.
Figure 2-28. A coloboma of the optic nerve head and sclera as part of the Jubb KF, Kennedy PC (1993): Pathology of the Domestic Animals, 4th ed,
collie eye anomaly syndrome. Vol II. Academic Press, New York.
McAvoy JW, et al. (1999): Lens development. Eye 13:425.
Mey J, Thanos S (2000): Development of the visual system of the chick:
colobomas in the lateral segments of the upper eyelids are I. Cell differentiation and histogenesis. Brain Res Brain Res Rev 32:343.
common. These must be surgically corrected, because they Pichaud F, Desplan C (2002): Pax genes and eye organogenesis. Curr Opin
allow facial hair to irritate the cornea and conjunctiva. Genet Dev 12:430.
Provis JM (2001): Development of the primate retinal vasculature. Prog Retin
Eye Res 20:799.
BIBLIOGRAPHY Reza HM, Yasuda K (2004): Lens differentiation and crystallin regulation:
a chick model. Int J Dev Biol 48:805.
Achiron R, et al. (2000): Axial growth of the fetal eye and evaluation of the Rutledge JC (1997): Developmental toxicity induced during early stages of
hyaloid artery: in utero ultrasonographic study. Prenat Diagn 20:894. mammalian embryogenesis. Mutat Res 12:113.
Aguirre G, et al. (1972): The development of the canine eye. Am J Vet Res Sengpiel F, Kind PC (2002): The role of activity in development of the visual
33:2399. system. Curr Biol 12:R818.
Bailey TJ, et al. (2004): Regulation of vertebrate eye development by Spencer WH (1996): Ophthalmic Pathology, 4th ed. Saunders, Philadelphia.
Rx genes. Int J Dev Biol 48:761. Stromland K, et al. (1991): Ocular teratology. Surv Ophthalmol 35:429.
Baker CV, Bronner-Fraser M (2001): Vertebrate cranial placodes I: embryonic Zieske JD (2004): Corneal development associated with eyelid opening.
induction. Dev Biol 232:1. Int J Dev Biol 48:903.
Barishak RY (2001): Embryology of the Eye and Its Adnexa. S. Karger,
Basel, Switzerland.
Chapter
OCULAR PHARMACOLOGY AND 3

THERAPEUTICS
David J. Maggs
THERAPEUTIC FORMULATIONS IMMUNOMODULATING THERAPY PROSTAGLANDIN ANALOGUES

ROUTES OF ADMINISTRATION (IMMUNOSUPPRESSANTS AND LOCAL ANESTHETICS
ANTIBACTERIAL DRUGS IMMUNOSTIMULANTS) ENZYMES AND ENZYME INHIBITORS
ANTIFUNGAL DRUGS MAST CELL STABILIZERS AND TEAR REPLACEMENT PREPARATIONS
ANTIVIRAL DRUGS ANTIHISTAMINES (“ARTIFICIAL TEARS”)
CORTICOSTEROIDS HYPEROSMOTIC AGENTS MISCELLANEOUS THERAPEUTIC AGENTS
NONSTEROIDAL ANTIINFLAMMATORY AUTONOMIC DRUGS PHYSICAL THERAPY
DRUGS CARBONIC ANHYDRASE INHIBITORS
THERAPEUTIC FORMULATIONS ROUTES OF ADMINISTRATION

Although topical application of solutions, suspensions, and The main factors governing choice of the route of adminis-
ointments is most common in ocular medicine, parenteral tration are as follows:
methods of administration via a systemic (i.e., intravenous,
• Inherent properties of the drug
intramuscular, subcutaneous) or local (i.e., subconjunctival,
• Site of desired action (surface or intraocular structures)
intraorbital, intracameral, intravitreous) route are also used.
• Frequency of administration possible
Drugs for ocular administration are prepared in various ways.
• Drug concentration required at target tissue
The topical use of powders for ocular treatment is detrimental
• Vascularity of the target tissue
to the eye and outmoded. Solutions, suspensions, and oint-
ments for topical application must have physical charac- Some drugs, because of their properties, are restricted as to
teristics within a relative narrow range to be well tolerated. the routes by which they can be given. For example, polymyxin
Of these the most important characteristics are tonicity B cannot be given systemically because of nephrotoxicity
and pH. These parameters must also be considered by or by subconjunctival injection because of local irritation.
compounding pharmacists when formulating drugs for topical Drugs required in high concentration in the cornea or conjunctiva
ophthalmic use. are usually administered by frequent topical application or sub-
Ophthalmic preparations must be sterile, especially if they conjunctival injection. If high concentrations are required in the
enter the interior of the eye. Bacterial filtering and the addition anterior uveal tract (i.e., iris or ciliary body), subconjunctival
of preservatives such as benzalkonium chloride are used to limit injection, systemic administration, or frequent topical application
contamination of multidose containers. However, these preser- of drugs that will pass through the intact cornea are used. Drugs
vatives are also toxic to mammalian cells. This fact has a that do not pass through the blood-ocular barrier still reach high
number of important clinical implications: concentrations in the highly vascular anterior uvea (iris and
ciliary body), posterior uvea (choroid), and sclera. With inflam-
• Topical drug use is not benign and should always be limited
mation the blood-ocular barrier may be reduced, and drugs that
to the lowest effective concentration, frequency, and
cannot normally enter the aqueous or vitreous humor may do
duration.
so. If high concentrations are required within orbital tissues,
• Drugs designed for topical use should not be used
systemic administration is usually used. Choice of route is
intraocularly or injected, especially subconjunctivally.
summarized in Figure 3-1.
• Preservatives in ophthalmic drugs and diagnostic agents
The cornea may be considered a trilaminar (lipid-water-
may interfere with diagnostic attempts to isolate and grow
lipid) “sandwich,” in which the epithelium and endothelium are
microbes from the ocular surface.
relatively lipophilic and hydrophobic, whereas the stroma is
Although topical application of drugs provides excellent relatively hydrophilic and lipophobic. Lipid-soluble drugs (e.g.,
drug concentrations at the ocular surface, there are two critical chloramphenicol) penetrate more readily, whereas electrolytes
barriers to penetration of drugs into the eye. These are the and water-soluble drugs (e.g., neomycin, bacitracin, and peni-
blood-ocular barrier (which, like the blood-brain barrier, is cillin) penetrate poorly if at all after topical application. The
impermeable to most drugs unless there is significant intra- lipophilic properties of the epithelium may be partially bypassed
ocular inflammation) and the cornea (which becomes more by subconjunctival injection, provided that other properties of
permeable when ulcerated). the drug are suitable for administration by this route.
33
Figure 3-1. Sites of drug administration. A, Topically applied

C
ointments, suspensions, and solutions achieve high drug
concentrations on the corneal and conjunctival surface. Not all
penetrate through the cornea and so are not uniformly useful for
treating intraocular disease. Penetration to the posterior segment
is extremely limited for all topical medications. B, Sub-
conjunctival injections facilitate high drug concentrations at the
ocular surface and provide a variable-duration depot effect. Not
all drugs are tolerated at this site. Intraocular penetration varies B
with drug type. C, Intraocular (intracameral or intravitreal)
injection of drugs is used rarely because general anesthesia is
needed, serious ocular risks are associated with this route of
therapy, and alternative routes often provide equivalent
intraocular drug concentrations. D, Systemically administered
drugs (oral or parenteral) reach all vascular ocular structures but
are not delivered in useful concentrations to the avascular A
structures, such as the cornea and lens. Intraocular penetration of
systemically administered drugs depends on the patency of the
blood-ocular barrier and the lipophilicity of the drug.
D
Higher or more prolonged drug concentrations and therapeutic Solutions and Suspensions (“Drops”)
effects may be achieved with the following approaches:
Ophthalmic solutions and suspensions (or “drops”) are com-
• Increasing drug concentration in the topical preparation; monly used for topical treatment of ocular disease. They are
for example, 1% prednisolone suspension results in higher usually easily instilled in dogs and cats but not in large animals.
intraocular concentrations than does 0.5% suspension The correct method for instilling eyedrops is shown in Figure 3-3.
(Figure 3-2). Drops permit the delivered dose to be controlled and varied
• Increasing the frequency of application; for example, easily, and they are alleged to interfere less with repair of
topical administration every 10 minutes for an hour results corneal epithelium than ointments, although this last feature is
in higher concentrations than does a single application. unlikely to be clinically significant. Drops are quickly diluted
• Slowing absorption. Drugs released over a long period can and eliminated from the eye by tears, so greater frequency of
be used to maintain drug concentrations. This is one application or drug concentration may be required, especially
mechanism whereby subconjunctival administration leads to with increased lacrimation. It is important to note that systemic
higher topical concentrations of drug because of delayed absorption of drugs from the conjunctival sac after topical
release (“leakage”) of drug back along the injection tract. application is rapid and may result in notable blood
Penetration of topically administered drugs can also be concentrations. This may be of clinical significance with use of
enhanced through the use of preparations that maintain longer phenylephrine (producing systemic hypertension) and long-term
contact with the eye before being washed away by the tears corticosteroid use (inducing iatrogenic hyperadrenocorticism).
(e.g., ointments, suspensions, or more viscous solutions).
• Facilitation of passage of drug between epithelial cells by
limited and controlled damage of the intercellular adhesions
of the epithelium with a surface-active preservative such as
benzalkonium chloride (an additive in some drugs).
Fludrocortisone 0.1%
Cortisone 2.5%
Hydrocortisone 2.5%
Prednisolone 0.5%
Dexamethasone 0.1%
Figure 3-3. Correct method of instilling an eyedrop. The lower eyelid is
Prednisolone 1.0% held open with the hand being used to restrain the patient’s head. The
upper eyelid is retracted with the hand holding the medication. The
Figure 3-2. Relative antiinflammatory action of various corticosteroid medication container is held 1 to 2 cm from the eye, and a single drop is
preparations. Note that prednisolone (1.0%) and dexamethasone are instilled. Care must be taken to avoid the bottle touching the eye because
relatively more potent than hydrocortisone. (Modified from Havener WH this may injure the ocular surface or cause contamination of the drug
[1994]: Ocular Pharmacology, 6th ed. Mosby, St. Louis.) remaining in the bottle.
OCULAR PHARMACOLOGY AND THERAPEUTICS 兩 35
Continuous or Intermittent Ocular Surface Subpalpebral lavage systems placed in the medial aspect
Lavage Systems of the lower lid are associated with less common and less severe
ocular complications than those placed centrally and dorsally, even
With frequent treatment or in horses with painful eyes, a lavage when left in place and used by owners for up to 55 days after
system allows medications to be conveniently, safely, and discharge from hospital. The lavage tube leads back to the
frequently delivered into the conjunctival sac. Originally, such shoulder, where it is secured at the mane and where drugs can be
systems were placed within the nasolacrimal duct and medi- administered with less risk of injury to the eye or the operator.
cations were instilled in a retrograde fashion. More recently Drugs are injected into the tube and either slowly propelled to the
subpalpebral lavage systems have been described that are eye with a gently administered bolus of air from a syringe or
simply placed and avoid nasal irritation and risk of dislodge- continuously propelled by a gravity-fed bottle or small mechanical
ment. A two-hole technique through the skin of the upper infusion pump connected to the tube. This method of therapy is
eyelid has now been replaced by single-hole systems, owing to usually reserved for horses with severe corneal or uveal disease. A
the commercial availability of lavage systems with footplates to protective eyecup can be applied over the lavage tube for
prevent inadvertent removal. The original one-hole system was protection of the eye and apparatus. Ointments (and some more
placed in the central upper lid but is associated with a relatively viscous suspensions) cannot be applied through lavage systems.
high risk of complications, most notably corneal ulceration due
to rubbing of the footplate on the cornea. Placement of the sub- Ointments
palpebral lavage system in the ventromedial conjunctival fornix
may be preferred because of the natural corneal protection In horses the orbicularis oculi muscle is very powerful, and
provided by the third eyelid at that point (Figure 3-4). it is impossible to tilt the head to allow a drop of solution to
A B
C D
Figure 3-4. Placement of a subpalpebral lavage system in the medial

aspect of a horse’s lower eyelid. A, A local (subcutaneous) injection of
lidocaine is administered. B, The palpebral and fornicial conjunctival
surfaces are anesthetized with proparacaine-soaked cotton-tipped
applicators held in place for 1 or 2 minutes. C, A trocar is used to
penetrate the lower lid from the conjunctival fornix, and the lavage tube
is threaded through it. D, The trocar is removed, and the lavage tube is
pulled down until it lies snugly in the ventral conjunctival fornix between
the third eyelid and lower lid. E, The lavage tube is sutured in place with
adhesive tape tabs, and an injection port is placed at its terminus near the
mane on the same side as the affected eye.
E
enter without contaminating the bottle. Ointments are preferred auriculopalpebral nerve block to produce akinesia of the upper
in such patients. Ointments also allow longer contact between lid (see Chapter 5). A few drops of topical ophthalmic anes-
the drug and surface tissues. Finally, less drug enters the thetic (e.g., proparacaine) are instilled into the conjunctival sac.
nasolacrimal apparatus. Therefore ointments achieve higher Conjunctival anesthesia is facilitated by a cotton-tipped
tissue concentrations than solutions or suspensions do. A applicator soaked in topical anesthesia and placed against the
soothing effect occurs on instillation, they are a more stable conjunctiva at the planned injection site for about 30 seconds.
medium for labile antibiotics and drugs, and they also provide The solution for subconjunctival injection then is administered
physical lubrication and protect against desiccation better than through a 25- to 27-gauge needle with a 1-mL tuberculin or
solutions do. However, because oily ointment bases cause insulin syringe under the bulbar conjunctiva as close as poss-
severe intraocular inflammation, and because application of ible to the lesion being treated (Figure 3-6). Injection under
ointments may result in ocular trauma from the tube itself, the palpebral conjunctiva is not effective. The needle is rotated
ointments are not recommended when globe perforation has on withdrawal to limit leakage through the needle tract. Up
occurred or is likely. Because they make tissue handling diffi- to 1 mL of drug can be given beneath the bulbar conjunctiva,
cult and cause granulomatous inflammation if they penetrate but most injections do not exceed 0.5 mL. Slight hemorrhage
ocular surface structures, ointments should also not be used into the injection site occasionally occurs but is absorbed
before ocular or adnexal surgery. Finally, owners tend to over- within 7 to 10 days. Injections of depot preparations should be
medicate when using ointments, resulting in loss of medication, avoided at this site because they often lead to granuloma
higher cost, and lower compliance with treatment regimens. As formation.
little as 0.5 cm of ointment from a fine nozzle is sufficient to
medicate an eye. Supplies for subconjunctival injection are topical anesthetic, a
cotton-tipped applicator, a 1-mL syringe (tuberculin or insulin),
Subconjunctival, Subtenons, and Retrobulbar Injection a 25- to 27-gauge needle, and solution for injection.
Subconjunctival injection permits a portion of the administered Retrobulbar injection is used rarely and only for treatment of
drug to bypass the barrier of the corneal epithelium and disease processes in the orbit or posterior half of the globe. These
penetrate transsclerally. However, a notable proportion of the areas can usually be treated adequately with safer and simpler
injected drug leaks back out the injection tract and is absorbed systemic routes of treatment. Therefore this route of therapy is
as if it had been administered topically. Subconjunctival now generally limited to the injection of local anesthetic into the
administration is used to facilitate high drug concentrations in muscle cone behind the globe for removal of the bovine eye.
anterior regions of the eye, whereas deeper injections beneath
Tenon’s capsule allow greater diffusion of drugs through the Systemic Drug Administration
sclera and into the eye. Mydriatics (for pupillary dilation),
antibiotics, and corticosteroids are the main groups of drugs Although there are rare exceptions, systemically administered
administered by this route. Some irritating drugs (e.g., drugs should be considered to reach only the vascular tissues of
polymyxin B) or any topical drug containing a preservative the eye and surrounding structures—that is, not the cornea, the
cannot be given subconjunctivally. Drugs with potent systemic lens, or (in the presence of an intact blood-ocular barrier)
sympathomimetic or vasopressor effects also should not be the aqueous or vitreous humor. This knowledge may be used to
given in this manner. the clinician’s advantage. For example, systemic administration
In cooperative patients, subconjunctival injections can be of a corticosteroid for control of uveitis in the presence of
given using topical anesthesia only. Handheld lid retractors corneal ulceration is safe and effective, because the target tissue
may be helpful in all species (Figure 3-5). For horses and (the uvea) is vascular, but the drug will not reach the avascular
cattle, one should also consider tranquilization, appropriate cornea in quantities sufficient to retard healing. Equally, the
restraint (a twitch for horses and nose grips for cattle), and an systemic administration of an antibiotic for treatment of
A B
Figure 3-5. A, Handheld lid retractors. B, Retractor in use.
A B
Figure 3-6. Subconjunctival injection technique. A, After application of a topical anesthestic, a 25- to 27-gauge
needle is inserted beneath the bulbar conjunctiva. Care must be taken not to penetrate the globe. B, The
medication injected forms a noticeable bleb that reduces in size over the next few minutes to hours. (Courtesy
Dr. David Ramsey.)
an ulcer in a nonvascularized cornea is of little value, and often are used interchangeably. Antibacterial agents may be
topical administration is most effective. Therefore systemically classified as bactericidal (destroying bacteria) or bacteriostatic
administered drugs should be reserved for treatment of diseases of (inhibiting bacterial growth and reproduction; Box 3-1). Some
the eyelids, conjunctiva, sclera, uvea (iris, ciliary body, choroid), antibiotics may act in either manner, depending on concen-
retina, optic nerve, extraocular muscles, and orbital contents. tration. Combining bactericidal and bacteriostatic antibiotics
As with other body systems, intravenous, subcutaneous, and may result in antagonism between the agents, although the
intramuscular injections provide relatively high plasma con- clinical importance of this effect is debated. In particular, com-
centrations of a drug to the vascular components of the eye. binations of bactericidal drugs infrequently used elsewhere in
However, because continuous treatment is necessary for many the body are commonly employed in topical treatment of the
ocular disorders, oral drug administration by owners is used eye. This practice allows a wider spectrum of activity than with
most frequently, particularly in dogs and cats. Continuous intra- single drugs and reduces the chance of drug resistance. The
muscular administration is used occasionally in large animals. combination of neomycin, polymyxin B, and bacitracin (or
Intravenous therapy is rarely used for ocular disease, with gramicidin) as so-called triple antibiotic is very useful. For
the important exception of the administration of mannitol for resistant infections combinations of agents with differing
reduction of intraocular pressure (IOP) and vitreous volume in mechanisms are sometimes used—for example, a penicillin or
acute glaucoma. cephalosporin (which inhibits cell wall synthesis) with an amino-
Intraocular drug concentrations attainable by systemic routes glycoside (which inhibits intracellular protein synthesis).
depend on the following three important factors:
• Absorption of the drug from the injection site or Selection and Administration of Antibiotics
gastrointestinal tract, and the plasma concentrations reached
The following factors must be considered in the selection of an
• Vascularity of the target tissue
antibiotic:
• Properties of the drug with respect to the blood-ocular
barrier. Systemically administered drugs will not reach
• The offending organism and its sensitivity
those areas of the eye “protected” by an intact blood-ocular
• Location of the organism
barrier (the vitreous, aqueous humor, and retina). In
• Penetration of available drugs to that site
inflamed eyes, this barrier is usually much less effective.
• Pharmacokinetics of the available drugs
Some examples reinforce these general points. Although • Spectrum of activity of available drugs
penicillin is readily absorbed by intramuscular injection, peni-
cillin G is poorly absorbed orally because it is destroyed by
Box 3-1 兩 Classification of common ophthalmic
gastric acid. Even when adequate plasma concentrations are
antibiotics
achieved, it penetrates the blood-ocular barrier poorly. By con-
trast, chloramphenicol is well absorbed by dogs after oral
administration and, once in the plasma, passes the blood- Bactericidal Bacteriostatic
aqueous barrier well. From these examples, it follows that the Aminoglycosides Chloramphenicol
clinician must understand the properties of the individual drugs Bacitracin Cephalosporins
used in order to predict their applicability for ophthalmic use. Erythromycin* Erythromycin*
Fluoroquinolones Tetracyclines
Potentiated sulfonamides
ANTIBACTERIAL DRUGS Neomycin
Penicillins
Antibacterial drugs act by altering cell wall synthesis, protein
Polymyxins
synthesis, or cell wall permeability in bacteria but may also Vancomycin
have undesirable effects on the cells of the patient. Although
not strictly accurate, the terms antibiotics and antibacterials *Depending on concentration and organism.
• Toxicity of available drugs Table 3-1 兩 Normal Flora of the Canine Conjunctival Sac
• Owner compliance
PERCENTAGE OF CASES
AREA AND FLORA WITH POSITIVE CULTURES
The ideal basis for selection of an ocular antibiotic consists of
identification of the responsible organism and its antibiotic sensi- WESTERN UNITED STATES*
tivity. However, obtaining this information often cannot be Diphtheroids 75.0
justified because of expense or because treatment must be Staphylococcus epidermidis 46.0
Staphylococcus aureus 24.0
instituted before the results of such testing are available. Therefore Bacillus spp. 12.0
knowledge of the most likely organisms, their sensitivity, and the Gram-negative organisms 7.0
most likely effective antibiotics is necessary. Treating infections on (Acinetobacter, Neisseria,
such an empirical basis, although practical and often unavoidable, Moraxella, and Pseudomonas spp.)
does not always lead to a satisfactory result. A more rational Streptococcus spp. (a-hemolytic) 4.0
Streptococcus spp. (b-hemolytic) 2.0
choice of therapeutic agent can be made after examination of the
staining and morphologic characteristics of organisms seen on a MIDWESTERN UNITED STATES†
Gram-stained or Diff-Quik–stained sample of the affected tissue S. epidermidis 55.0
S. aureus 45.0
and is essential in severe or nonresponsive infections. In more Streptococcus spp. (a-hemolytic) 34.0
severe infections (e.g., stromal corneal ulcers or endophthalmitis), Diphtheroids 30.0
the organism should be identified, and a combination of routes of Neisseria spp. 26.0
administration and synergistic drugs should be considered. If Pseudomonas spp. 14.0
ocular infections persist or recur despite treatment with the Streptococcus spp. (b-hemolytic) 7.3
appropriate antibiotic (based on results of culture and sensitivity EASTERN AUSTRALIA‡
testing), the infection may be secondary to an underlying disorder S. aureus 39.0
or pathologic process. Alternatively, a fastidious bacterium or Bacillus spp. 29.0
Corynebacterium spp. 19.0
nonbacterial microbe not originally cultured (Chlamydophila S. epidermidis 16.0
spp., Mycoplasma spp., fungus, virus, etc.) may be present. Yeasts 5.0
Organisms commonly isolated from the conjunctival sacs of Streptococcus spp. (a-hemolytic) 3.0
normal and diseased animals are given in Tables 3-1 through Streptococcus spp. (nonhemolytic) 3.0
3-9. These tables highlight a number of important general Micrococcus spp. 3.0
Neisseria spp. 2.0
points about ocular surface flora that are relevant when inter- Streptococcus spp. (b-hemolytic) 1.0
preting these data in an individual patient: Pseudomonas spp. 1.0
Nocardia spp. 1.0
• There are marked species, individual, geographic, and Escherichia coli 1.0
seasonal variations in the normal ocular surface flora. Clostridium spp. 1.0
• The normal conjunctival sac often contains potential pathogens. Enterobacter spp. 1.0
• Because of the variety of organisms present, empirical Flavobacterium spp. 1.0
Branhamella catarrhalis 1.0
treatment with standard antibiotics may be unsuccessful.
• Care is advisable in interpretation of frequency and *Data from Bistner SI, et al. (1969): Conjunctival bacteria: clinical
sensitivity data from different geographic areas. appearances can be deceiving. Mod Vet Pract 50:45.
• In vitro sensitivity data may not necessarily reflect in vivo †Data from Urban W, et al. (1972): Conjunctival flora of clinically normal
dogs. Am J Vet Med Assoc 161:201.
experience when one is using topically applied antibiotics, ‡Data from McDonald PJ, Watson ADJ (1976): Microbial flora of normal
because high surface concentrations can be achieved. canine conjunctivae. J Small Anim Pract 17:809.
• Previously untreated infections seen in general practice may
have a different spectrum of sensitivity from those reported
Because it is unstable at low pH, oral administration is not
in referral hospital populations.
possible, so the agent is administered by injection. Because of
The following sections provide a summary of some impor- its high water solubility, penicillin G does not penetrate the
tant properties of antibiotics used commonly in veterinary intact cornea when applied topically and does not pass through
ophthalmology, and Table 3-10 lists the typical Gram-staining the intact blood-ocular barrier. It is most effective against gram-
characteristics of, and the antibiotics of choice for, common positive organisms, but it is susceptible to b-lactamase. For all
organisms. of these reasons penicillin G does not have a great number of
uses in veterinary ophthalmology, with perhaps the exception
Penicillins of subconjunctival injection for treatment of susceptible
infections and in situations in which frequent applications
The penicillins form a large family of natural and synthetic of medication are inconvenient (e.g., infectious bovine
derivatives of 6-aminopenicillanic acid that range considerably keratoconjunctivitis).
in stability, solubility, spectrum of activity, ocular penetration,
and resistance to b-lactamase. Sodium Methicillin
Penicillin G Methicillin is resistant to b-lactamase and is used by intra-

venous infusion for resistant staphylococci. Renal excretion is
Penicillin G is soluble in water, attains high concentrations in rapid. Because the drug is unstable in solution, it should be
blood, and is excreted in urine in 4 to 6 hours. Penicillin G is dissolved just before use. Methicillin can be used topically or
available in crystalline, procaine, and benzathine forms. by subconjunctival injection for corneal infections. Methicillin
Table 3-2 兩 Organisms Cultured from Dogs with External Table 3-4 兩 Fungal Flora from Normal Horses (50 Horses)
Ocular Disease PERCENTAGE OF CASES WITH
PERCENTAGE OF ORGANISM POSITIVE CULTURES
CASES WITH
AREA AND FLORA POSITIVE CULTURES Aspergillus spp. 36.0
Cladosporium spp. 34.0
THE NETHERLANDS* Positive but no quantitative data given: Alternaria, Fusarium,
Streptococcus canis 20.3 Monotospira, Paecilomyces, Phoma, Pullularia, Scopulariopsis,
No growth 18.7 Streptomyces, Trichoderma, Verticullium spp.
Staphylococcus epidermidis 14.0
Modified from Smith PJ, et al. (1997): Identification of sclerotomy sites for
posterior segment surgery in the dog. Vet Comp Ophthalmol 7:180.
Other nonpathogenic Streptococcus spp. 7.8
Nocardia spp. 7.8
Absidia ramosa (fungus) 7.8 Table 3-5 兩 Organisms Cultured from Horses with
Pseudomonas aeruginosa 6.1
Corynebacterium spp. 4.6
External Ocular Disease (123 Eyes)
Other pathogenic S. canis spp. 3.1 PERCENTAGE OF CASES WITH
Proteus vulgaris 3.1 ORGANISM POSITIVE CULTURES
Clostridium perfringens 1.5
Candida spp. 1.5 Streptococcus spp. (total) 43.9
COLORADO† b-Hemolytic 26.0
S. aureus 68.0 Other hemolytic 17.9
S. epidermidis 27.0 Staphylococcus spp. 24.4
Streptococcus spp. (b-hemolytic) 19.0 Pseudomonas spp. 13.8
Streptococcus spp. (a-hemolytic) 17.0 Bacillus spp. 10.6
Proteus mirabilis 11.0 Enterobacter spp. 6.5
Escherichia coli 10.0 Escherichia coli 4.0
Bacillus spp. 5.0 Corynebacterium spp. 3.2
Corynebacterium spp. 3.0 Proteus spp. 3.2
P. aeruginosa 2.0 Aspergillus spp. 2.4
Klebsiella spp. 1.0 Klebsiella spp. 2.4
Moraxella spp. 2.4
ILLINOIS‡ Pasteurella spp. 2.4
Staphylococcus spp. 39.4 Mima spp. 1.6
Coagulase positive 29.0 Diplococcus spp. 0.8
Staphylococcus intermedius 17.0 Flavobacterium spp. 0.8
S. epidermis 11.0 Fusarium spp. 0.8
Streptococcus spp. 25.2 Neisseria spp. 0.8
b-Hemolytic streptococci 17.0 Nocardia spp. 0.8
Pseudomonas spp. 9.4 Penicillium spp. 0.8
Rhizopus spp. 0.8
*Data from Verwer MAJ, Gunnick JW (1968): The occurrence of bacteria in Trichosporon spp. 0.8
chronic purulent eye discharge. J Small Anim Pract 9:33.
†Data from Murphy JM, et al. (1978): Survey of conjunctival flora in dogs Data from McLaughlin SA, et al. (1983): Pathogenic bacteria and fungi
with clinical signs of external eye disease. J Am Vet Med Assoc 172:66. associated with extraocular disease in the horse. J Am Vet Med Assoc
‡Data from Gerding PA, et al. (1988): Pathogenic bacteria and fungi 182:241.
associated with external ocular diseases in dogs: 131 cases (1981-1986).
J Am Vet Med Assoc 193:242.
Table 3-6 兩 Normal Flora of the Bovine Conjunctival Sac
PERCENTAGE OF CASES
Table 3-3 兩 Normal Flora of the Feline Conjunctival Sac AREA AND FLORA WITH POSITIVE CULTURES
PERCENTAGE OF CASES
NORTHEASTERN AUSTRALIA
WITH POSITIVE CULTURES Unidentified gram-positive cocci 54.4
AREA AND FLORA CONJUNCTIVA LIDS Corynebacterium spp. 27.4
Moraxella nonliquefaciens 26.9
WESTERN UNITED STATES Streptococcus faecalis 20.0
Staphylococcus epidermidis 16.3 13.3 No growth 13.4
Staphylococcus aureus 10.4 8.8 Neisseria (Branhamella) catarrhalis 10.5
Mycoplasma spp. 5.0 — (nonhemolytic)
Bacillus spp. 2.9 1.7 Unidentified gram-negative rods 8.5
Streptococcus spp. (a-hemolytic) 2.5 1.7 Acinetobacter spp. 8.0
Corynebacterium spp. 1.3 — Moraxella bovis 6.5
Escherichia coli — 0.4 Coliforms 6.5
Data from Campbell L (1973): Ocular bacteria and mycoplasma of the Moraxella liquefaciens 2.2
clinically normal cat. Feline Pract Nov-Dec:10. Bacillus spp. 1.3
Unclassified Moraxella 1.0
Actinobacillus spp. 0.7
Proteus spp. 0.0
Data from Wilcox G (1970): Bacterial flora of the bovine eye with special
reference to Moraxella and Neisseria. Aust Vet J 46:253.
Table 3-7 兩 Normal Flora of the Ovine Conjunctival Sac Amoxicillin and Ampicillin
PERCENTAGE OF CASES
AREA AND FLORA WITH POSITIVE CULTURES
Ampicillin is a broad-spectrum penicillin that is often effective
against Escherichia coli and Proteus spp. and may be given
EASTERN AUSTRALIA orally, intramuscularly, or subconjunctivally. Although it enters
No growth 60.0 the aqueous humor to some extent, ampicillin is not the agent
Neisseria ovis 24.0 of first choice for gram-negative infections, because the high
Micrococcus spp. NQD
Streptococcus spp. NQD
inhibitory concentrations necessary are not always reached in
Corynebacterium spp. NQD the aqueous humor. Amoxicillin has a spectrum of activity
Achromobacter spp. NQD similar to that of ampicillin but is better absorbed from the
Bacillus spp. NQD gastrointestinal tract than ampicillin. Both ampicillin and
Moraxella spp. NQD amoxicillin are susceptible to b-lactamase. Amoxicillin reaches
Data from Spradbrow P (1968): The bacterial flora of the ovine conjunctival blood concentrations two to three times higher than those
sac. Aust Vet J 44:117. of ampicillin after oral administration, but both drugs enter
NQD, No quantitative data (present in small numbers, but NQD given). the uninflamed eye to about the same degree. Because it
inhibits b-lactamases, clavulanic acid is added to preparations
Table 3-8 兩 Normal Bacterial Flora of South American
of amoxicillin. This preparation is useful in initial treatment of
Camelid Eyes (88 Animals) chronic staphylococcal blepharitis—in which the staphylococci
PERCENTAGE OF CASES are often b-lactamase producers—and orbital cellulitis,
ORGANISM WITH POSITIVE CULTURES which may involve anaerobic bacteria implanted from the
oral cavity.
GRAM-POSITIVE
Staphylococcus spp. 58
Bacillus spp. 28 Cephalosporins
Streptomyces spp. 18
a-Hemolytic Streptococcus spp. 13 A range of cephalosporins is available. They are generally
Corynebacterium spp. 8 similar to the penicillins in mechanism of action and pharma-
Micrococcus/Planococcus spp. 7
cology but are less susceptible to staphylococcal b-lactamases.
GRAM-NEGATIVE A type of b-lactamase (cephalosporinase) produced by some
Pseudomonas spp. 41 gram-negative organisms may inactivate them. Cephalosporins
Pasteurella ureae 9 have had excellent results when administered subconjunctivally
Klebsiella spp. 2
Escherichia coli 2 or topically via lavage tube to horses with corneal ulceration,
particularly those infected with Streptococcus spp. They are
Data include llama, alpaca, and guanaco eyes and are from Gionfriddo JR, also very useful for bacterial blepharitis. Cefazolin is the anti-
et al. (1991): Bacterial and mycoplasmal flora of the healthy camelid biotic of choice for perioperative antimicrobial prophylaxis in
conjunctival sac. Am J Vet Res 52:1061.
small animal surgery.
Table 3-9 兩 Normal Fungal Flora of South American
Camelid Eyes (127 Animals) Chloramphenicol
PERCENTAGE OF CASES Chloramphenicol is a broad-spectrum bacteriostatic antibiotic
ORGANISM WITH POSITIVE CULTURES* effective against a wide range of gram-positive and gram-
Aspergillus spp. 20-43
negative organisms, Rickettsia, spirochetes, and Chlamydophila
Fusarium spp. 2-30 spp. However, Pseudomonas aeruginosa is often resistant.
Rhinocladiella spp. 8-35 Because of its lipid solubility, chloramphenicol passes the
Penicillium spp. 3-33 blood-ocular barrier and through corneal epithelium better than
Mucor spp. 5-14 most water-soluble antibiotics. Chloramphenicol may be
Dematiaceous fungi 12-33
administered orally, intramuscularly, subcutaneously, intra-
Data include llama, alpaca, and guanaco eyes and are from Gionfriddo JR, venously, subconjunctivally, or topically. Because absorption
et al. (1992): Fungal flora of the healthy camelid conjunctival sac. Am J Vet after oral administration results in high blood concentrations,
Res 53:643. this is the route of choice for infections in the posterior
*Percentages varied according to season and camelid species.
globe and orbit. Despite controversial toxicity studies in
cats, the drug has been used clinically over many years
with few ill effects except anorexia and occasional pyrexia
may be expected to enter the aqueous humor in therapeutic in some cats after systemic administration, provided that
concentrations when the blood-ocular barrier is disrupted by administration is not prolonged. Because of its wide spectrum
inflammation. of activity and intraocular penetration, chloramphenicol as
an ointment or drops was widely used in veterinary practice
Sodium Oxacillin for ocular surface injuries and infections, but its use is
declining in favor of bactericidal antibiotics. Polymyxin B is
Oxacillin is resistant to b-lactamase; it is also acid-stable and often added to aid in control of gram-negative organisms,
may be used orally. Unfortunately, much of it is bound to especially Pseudomonas spp. It is still useful in the topical
plasma protein in the circulation and cannot enter the aqueous treatment of suspected Chlamydophila infections in cats,
humor, even in an inflamed eye. Oxacillin is useful in orbital although systemically administered drugs are preferable (see
and adnexal infections when given orally. Chapter 7).
Table 3-10 兩 Antibiotics of Choice for Common Organisms

ORGANISM DRUG OF CHOICE ORGANISM DRUG OF CHOICE
GRAM-POSITIVE COCCI FUNGI AND YEASTS

Staphylococcus spp. Neomycin Fusarium spp. Natamycin
Bacitracin Thiabendazole
Amoxicillin Aspergillus spp. Amphotericin B
Cephalosporins Ketoconazole
Erythromycin Itraconazole
Fluoroquinolones Flucytosine
Staphylococcus aureus Gentamicin Nystatin
Oxacillin Candida spp. Nystatin
Methicillin Amphotericin B
Cephalosporins Flucytosine
Fluoroquinolones Ketoconazole
Staphyococcus epidermidis Neomycin Itraconazole
Gentamicin Cryptococcus spp. Ketoconazole
Erythromycin Itraconazole
Fluoroquinolones Flucytosine
Streptococcus spp. Penicillin Penicillium spp. Natamycin
Chloramphenicol Blastomyces and Histoplasma spp. Ketoconazole
Amoxicillin Itraconazole
Cephalosporins Amphotericin B
Microsporon spp. Ketaconazole
GRAM-NEGATIVE COCCI Trichophyton spp. Itraconazole
Neisseria spp. Penicillin
Epidermophyton spp. Griseofulvin
Tetracyclines
Sulfonamides (± trimethoprim) ACTINOMYCETES
Actinomyces spp. Penicillin
GRAM-POSITIVE RODS Tetracyclines
Corynebacterium spp. Penicillin
Nocardia spp. Chloramphenicol
Tetracyclines
(± streptomycin)
Sulfonamides (± trimethoprim)
(± isoniazid)
GRAM-NEGATIVE RODS CHLAMYDIA
Pseudomonas aeruginosa Polymixin B
Chlamydia and Chlamydophila spp. Azithromycin
Gentamicin
Doxycycline
Tobramycin
Tetracyclines
Amikacin
Chloramphenicol
Fluoroquinolones
Escherichia coli Chloramphenicol MYCOPLASMA
Tetracyclines Mycoplasma spp. Tetracyclines
Gentamicin Erythromycin
Fluoroquinolones Chloramphenicol
Enterobacter spp. Amoxicillin (± streptomycin)
Proteus spp. Gentamicin
Fluoroquinolones
Tobramycin
Amikacin
Chloramphenicol
Hemophilus spp. Amoxicillin
Tetracyclines
Moraxella spp. Penicillin
Tetracyclines
Aminoglycosides as a topical agent of first choice for bacterial prophylaxis.

Given its relatively narrow effective spectrum, and because this
Neomycin spectrum encompasses gram-negative organisms, which are not
Neomycin is a particularly useful bactericidal agent for ocular use commonly found as part of the ocular surface flora, such use
and is active against gram-positive and gram-negative bacteria, is questionable. As a result of this over-use, the value of
including Staphylococcus aureus. Bacterial resistance develops gentamicin for treatment of more resistant organisms is now
less readily to neomycin than to streptomycin, and neomycin is markedly diminished. Gentamicin is variably effective against
more effective against Proteus vulgaris than is polymyxin B. many strains of S. aureus, Pseudomonas spp., E. coli,
Because of nephrotoxicity and ototoxicity, neomycin is Aerobacter, Klebsiella spp., and Proteus spp. Topical appli-
administered only topically or by subconjunctival injection. cation does not result in high intraocular concentrations, and
Topical hypersensitivity to neomycin occasionally develops. although some drug enters the eye after subconjunctival or
intravenous injection, vitreous penetration is poor regardless of
Gentamicin route of administration. Long-term systemic therapy is limited
by ototoxicity and nephrotoxicity. Gentamicin (like other
Gentamicin (perhaps only because of its relatively low cost) aminoglycosides) causes cataract and severe retinal degener-
has unfortunately been used frequently and indiscriminately ation when injected intraocularly.
Exploring the Variety of Random
Documents with Different Content
was therefore adopted, July 27, 1918. In this the volume of the
absorbent was reduced from 450 cc. to 300 cc. It was packed in two
layers, ⅔ in the bottom and ⅓ in the top. One pad was placed
between the layers and one on top. This change gave a lowering of
the resistance of 27 per cent (to 2.5 inches) at a sacrifice of 50 per
cent of the length of life of the canister, but not of protection during
the shortened life. Type L differed from this only in having 325 cc. of
absorbent, a change made to decrease leakage about the top cotton
pad.
Fig. 57.—U. S. Army Canister, Type J.
The following table shows the relative efficiency of various

canisters:
U. S.,
p. p. British, French,
Type German
m. S. B. R. A. R. S.
H
Chloropicrin 1000 770 17 2 43
Phosgene 2500 85 54 5 16
Hydrocyanic
500 70 90 — 10
acid
Mustard gas 100 1800 — 35 195
The figures represent time in minutes till the first traces of gas
begin to come through.
Fig. 58.—Type J Canister and Contents.
Manufacture
The following description of the manufacture of the gas mask at
the Long Island plant is taken from an article by Col. Bradley
Dewey[26]:
“Incoming Inspection—A thorough 100 per cent inspection was
made of each part before sending it to the Assembly Department.
The inspectors were carefully chosen and were sent to a school for
training before they were assigned to this important work. Every
feature found to be essential to the manufacture of a perfect gas
mask was carefully checked.
“The incoming inspection of the flexible rubber hose leading from
the canister to the facepiece can be taken as an illustration. Each
piece of hose was given a visual inspection for buckles or blisters in
the ends or in the corrugations; for cuts, air pockets, or other defects
on the interior; for loose seams where fabric covering was cemented
to the rubber tube; for weaving defects in the fabric itself; and for
careless application of the cement. Special tests were conducted for
flexibility, as a stiff hose would produce a strain on the soldier’s
mouth; for permanent set to insure that the hose was properly cured;
for the adhesion of the fabric covering to the hose; and for kinking
when the hose was doubled on the fingers. Finally each piece was
subjected to a test for leaks under water with a pressure of 5 lbs. per
sq in.
“Each eyepiece and the three-way metal connection to the
facepiece were subjected to a vacuum test for leakage. The delicate
exhalation valve was carefully examined for defects which would be
liable to cause leakage. Fabric for the facepiece was given a high-
tension electrical test on a special machine developed at the plant to
overcome the difficulty met in the inspection of this most important
material. It was of course necessary that the facepiece fabric be free
from defects but just what constituted a defect was the source of
much discussion. The electrical test eliminated all personal views
and gave an impartial test of the fabric. The machine consisted of
two steel rolls between which a potential difference of 4,000 volts
was maintained; the fabric was led through the rolls and wherever
there was a pinhole or flaw the current arced through and burned a
clearly visible hole.
“Preliminary Facepiece Operations—Blanks were died out
from the facepiece fabric in hydraulic presses. Each face blank was
swabbed to remove bloom and the eye washers were cemented
about the eyeholes. The pockets for holding the noseclips were also
cemented to the blanks. The bands which formed a gas-tight seal of
the mask about the face were died out from rubberized fabric to
which a felt backing was attached. The harness consisting of elastic
and cotton tapes was also sewed together at this point.
“Facepiece Operations—The sewing machine operations were
next performed. First the died out blanks were pleated to form the
facepiece. The operator had to register the various notches in the
blank to an accuracy of ¹/₃₂ in. and to locate the stitches in some
cases as closely as ¹/₆₄ in. The band was next sewed to the
periphery of the facepiece after which the harness was attached.
The stitches on the outside of the facepiece were covered with liquid
dope, which filled the needle holes and made the seams gas-tight.
“In addition to the inspection of each operation, the completed
facepiece was submitted to a control inspection to discover any
defects that might escape the attention of the inspectors on the
various operations.
“Assembly Operations—The facepieces were now ready for
assembly and were sent for insertion of the eyepieces, which was
done in specially designed automatic presses. The eyepieces had to
be carefully inserted so that the facepiece fabric extended evenly
around the entire circumference.
“Before manufacture began on a large scale, the most
satisfactory method of conducting each assembly operation was
worked out and the details standardized, so that operators could be
quickly and efficiently trained. No detail was considered too small if it
improved the quality of the mask. The assembly operations
proceeded as follows:
“The exhalation valve was first joined to the three-way metal tube
which formed the connection between the facepiece, flexible hose,
and mouthpiece. Each valve was then tested for leakage under a
pressure difference of a one inch head of water. No valve was
accepted which showed leakage in excess of 10 cc. per min. under
these conditions.
“The metal guard to protect the exhalation valve was next
assembled, followed by the flexible hose. The three-way tube was
then assembled to the facepiece by means of a threaded connection
and the rubber mouthpiece attached. To illustrate the attention to
details the following operation may be cited:
“The contact surfaces between each rubber and metal part were
coated with rubber cement before the parts were assembled. The
connection was then tightly wired, care being taken that none of the
turns of wire should cross and finally the wire was covered with
adhesive tape so that no sharp edges would be exposed.
“The masks, completely assembled except for the canisters, were
inspected and hung on racks on specially designed trucks which
prevented injury in transit, and were delivered to the Finishing
Department.
“Canister Filling—Meanwhile the canisters were being filled, in
another building.
“The chemicals were first screened in such a way that the fine
and coarse materials were separated from the correctly sized
materials. They were then carried on a belt conveyor to the storage
bins, whence they were fed by gravity through pipes to various
mixing machines. A special mixing machine was developed to mix
the carbon and granules in the proper proportions for use in the
canister. The mixed chemicals were then led to the canister-filling
machines. There was a separate mixing machine for each filling
machine, of which there were eighteen in all.
“The can-filling department was laid out in six units. Each unit
had a capacity of 20,000 cans per day. A system of double belt
conveyors was installed to conduct empty canisters to the machines
and carry away the filled ones.
“Each filling operation was carefully inspected and special stops
were placed on the belt conveyors so that a canister could not go to
the next operation without having been inspected. Operators and
inspectors were stationed on opposite sides of the belt. The
chemicals were placed in the canister in three equal layers which
were separated by pads of cotton wadding. The first layer was
introduced from the filling machine (which delivered automatically the
proper volume of chemicals), the canister was shaken to pack the
chemicals tightly, the cotton baffle inserted, the second layer of
chemicals introduced and so forth. On top of the top layer of
chemicals were placed a wire screen and a specially designed
spring which held the contents of the canister securely in place. The
metal top was then fitted and securely soldered.
“Each canister was tested under water for possible leaks in joints
or soldering, with an air pressure of 5 lbs. per sq. in. A test was also
made for the resistance which it offered to breathing, a rate of flow of
air through the canister of 85 liters per min. being maintained and the
resistance being measured in inches of water.
“The filled canisters were then painted a distinctive color to
indicate the type of filling.
“Finishing Department—In the finishing department, the filled
canisters, were conducted down the middle of the finishing tables
and assembled to masks.
Fig. 59.
“The finished masks were then inspected, placed in unit boxes,

ten to a box, and returned for the final inspection.
“Final Inspection—Final inspection of the completely
assembled masks was as rigid as could be devised, and was closely
supervised by army representatives. Only the most painstaking, and
careful women were selected for this work and the masks were
examined in every detail to discover any defect that might have
escaped previous inspection. Finally, each mask was inspected over
a bright light in a dark booth for small pinholes which the ordinary
visual inspection might not have detected.
“As a check on the quality on the final inspectors’ work a
reinspection of 5 per cent of the passed masks was conducted.
Where it was found that a particular inspector was making numerous
mistakes, her eyes were examined to see whether it was due to
faulty eyesight or careless work. Masks containing known defects
were purposely sent to these inspectors to determine whether they
were capable of continuing the inspection work. In this way the
desired standard was maintained.
“A daily report of the final inspection was sent back to each of the
assembly departments involved so that defects might be eliminated
immediately and the percentage of rejects kept as low as possible.
“After the final inspection the masks were numbered, packed in
knapsacks, and the filled knapsacks placed in packing cases,
twenty-four to a case.”
Tissot Mask
The French, as has already been pointed out, early recognized
that certain classes of fighting men, as the artillerymen, needed the
maximum of protection with the minimum decrease in efficiency. The
result of this was the Tissot Mask. Before the United States entered
the war, the British standard box respirator had reached a greater
degree of perfection, with far greater ruggedness and portability. It
was therefore adopted as the American standard. At the time of the
invention of the British box respirator and practically up to the time
the United States entered the war, masks were worn only during the
sporadic gas attacks then occurring and only for a brief period at a
time. As the war progressed, the men were compelled to wear their
masks for much longer periods (eight hours was not uncommon). It
was then seen that more comfort was needed, even at the expense
of a little safety.
The principle of the Tissot mask was correct so far as comfort
was concerned, since it did away with the irritating mouthpiece and
noseclip, but the chief danger in the French mask arose from the fact
that the facepiece was made of thin, pure gum rubber. The Research
Division, together with the Gas Defense Division, developed two
distinct types of Tissot masks. The first of these was the Akron
Tissot, the second the Kops Tissot. The best features of these have
been combined in the 1919 Model.
Fig. 60.—American Tissot Mask,
Early Type.
Fig. 61.—American Tissot Mask,
Interior View.
1919 Model American Mask

Facepiece. This facepiece is made of rubberized stockinet about
one-tenth inch in thickness. The stockinet is on the outside only and
is for the purpose of strengthening and protecting the rubber which is
of very high grade. The facepiece is died out as a single flat piece
from the stockinet which is furnished in long rolls. The die is of such
shape that when the facepiece is sewed there is but one seam, and
that between the angle tube opening and the edge under the chin.
This seam is sewed with a zigzag stitch with the stockinet sides flat
together. The seam is then stretched over a jig, so as to form a flat
butt joint. This seam is then cemented with rubber cement and
taped, inside and out, to make it thoroughly gas-proof.
The eyepiece openings are of oval shape with the longer axes
horizontal and considerably smaller than the finished eyepieces. The
eyepieces being circular, the cloth is stretched to accommodate
them, giving the necessary bulge to keep the cloth and metal of the
eyepieces away from the face. The harness has three straps on
each side. Instead of the single strap over the top of the head, two
straps lead from directly over the eyes, both being made of elastic
the same as the other straps. All six straps are brought together
around a pad of felt and cloth about 2½ × 3½ inches at the back of
the head. This pad makes the harness much more comfortable.
The rubberized stockinet is reinforced on the inner or rubber side
with thin bits of cloth at all points where the straps are sewed on.
The strap across the temples just above the ears is sewed at two
points, one about one-half inch from the edge and the other about
two inches from the edge. This is for the purpose of helping press
the cloth against the temples, thereby adding to the gas-tightness for
those heads that have a tendency to be hollow at the temples. The
lower strap is just above the chin and is for the purpose of giving
gas-tightness in that vicinity. All of the straps except the two over the
top of the head are attached to the pad with buckles, and are thus
capable of exact adjustment.
The eyepieces are of triplex glass in metal rings with rubber
gaskets. In pressing the rings home, the rubberized stockinet is
turned and held securely so that there is no possibility of pulling
them out. The angle tube containing the outlet valve and the
connection to the corrugated tube connecting with the canister is the
same as with the latest model R. F. K. mask. The only difference as
regards the corrugated tube is that a greater length is needed with
the new carrier under the left shoulder. The total length of the tube
for this model is about 24 inches. On the inside of the facepiece and
connected to the angle tube inlet is a butterfly baffle of rubber, so
arranged that the incoming air is thrown upward and over the
eyepieces, thus keeping them clear no matter how much the exertion
or what the temperature, except in certain rare cases when the
temperature is down at zero F. or below.
Fig. 62.—1919 Model American Mask.
Canister
The canister is radically different from the canisters used in the R.
F. K. and earlier types. In the first place, it is longer, the total length
finished being 8 inches. It has two inlet valves at the top end
protected by a tin cover instead of the single inlet valve at the bottom
of the earlier types. The two inlet valves are each ⅝ inch in diameter
and are made up of square flat valves on the end of a short rubber
tube. The rubber tube is fitted over a short metal tube. Gas-tightness
is obtained both by the pressing of the valve against the round edge
of the metal tube and by the pressure of the edges against each
other. These valves, while delicate, are proving very satisfactory, and
being simply check valves to prevent the air going back through the
canister, they are not vital. In case of failure, the eyepieces would fog
somewhat and the dead air space be increased by that held in the
inlet tube.
The canister consists really of two parts—an outer casing that is
solid and an inner perforated tin casing. Around the perforated tin is
fitted a filter of wool felt ³/₁₆ of an inch in thickness. This wool felt is
very securely fastened by turning operations to solid pieces of tin,
top and bottom, so that no air can get into the chemicals without
passing through the filter. Thus the air coming through the inlet
valves at the top circulates around the loosely fitting outside
corrugated case to all parts of the filter and after passing through the
filter continues through the perforations of the tin into the charcoal
and soda-lime granules.
The chemicals are packed around a central wedge-shaped tube
extending about two-thirds the length of the can. The wedge is
enlarged at the top and made circular where it passes through the
top of the can to connect with the corrugated tube. The wedge-
shaped inner piece is made of perforated tin and is covered with thin
cloth to prevent dust from the chemicals passing into the tube and
thus into the lungs. The cans are filled from the bottom and are
subjected to two mechanical jarring operations in order to settle the
chemicals thoroughly before the spring which holds them in place is
added. The outer tin cap protecting the inlet valves has two openings
on each side but none at the ends of the canister.
Fig. 63.—1919 Model American Mask
after Adjustment.
The carrier is a simple canvas case nearly rectangular, about one

foot wide and 15 inches in length. The width is just sufficient at the
back to hold the canister and the front part to hold the extra length of
corrugated tube and the facepiece. There are two straps, one
passing over the right shoulder and the other around the body. The
one passing over the right shoulder has two “V” shaped seams at the
top so as to change the direction of the strap over the shoulder in
order that it will pull directly downward instead of against the neck.
The flap closing the case opens outward.
It has the usual automobile curtain fasteners. A secondary
fastener at the top of the opening is arranged so that when the tube
is adjusted to the proper length and the mask is adjusted to the face
of the wearer, the flap can be buttoned tightly over the corrugated
tube and held tightly. This prevents water from entering the case.
Figures 62 and 63 show the position of the carrier both with the
facepiece in the carrier and after adjustment. It will be noted that the
carrier does not interfere with the pack nor with anything on the front
of the body. The left arm hangs almost entirely natural over the case.
It has been thoroughly tried out by the Infantry, Cavalry, Artillery and
Special Gas Troops and adopted as eminently satisfactory.
Special Canisters
Navy. The early Navy canister is a drum much like the German
canister. The container is a slightly tapered metal cylinder, 9 cm. in
diameter at the bottom. The most satisfactory filling for this drum
consists of two layers, 98 cc. in each, of a standard mixture of
charcoal and soda-lime, separated by cotton wadding pad. The filling
is 6-20 mesh, instead of 8-14 mesh. A later type is shown in Figure
41.
Carbon Monoxide. This canister is discussed in Chapter XI.
Ammonia. Ammonia respirators were needed by the Navy and
also by the workmen in refrigeration plants. Early protection was
obtained by the use of pumice stone impregnated with sulfuric acid.
This had many disadvantages, such as the amount of heat evolved,
the caustic fumes produced, high resistance and corrosion of the
canister. To overcome these, the “Kupramite” canister was
developed. The filling consists of pumice stone impregnated with
copper sulfate. Pumice stone, 8 to 14 mesh, and technical copper
sulfate are placed in an evaporating pan in the ratio of one part by
weight CuSO₄·5H₂O to 1.5 parts pumice, and the whole is covered
with sufficient water to dissolve the salt at boiling temperature. The
mixture is then boiled down with constant stirring until crystallization
takes place on the pumice and the crystals are nearly dry. The
pumice thus treated is then removed from the dish, spread out and
allowed to dry in the air. The fines are then screened out on a 14-
mesh sieve. Care must be taken in the evaporating process that the
absorbent is still slightly moist when taken from the pan.
Fig. 64.—Early Type Navy Mask.

Contains noseclip and mouthpiece.
In packing the standard Army canister with kupramite a layer of
toweling is placed on top of the absorbent to filter out any fine
particles which might be drawn up from the absorbent, and the whole
is held in place by the usual heavy wire screen and spring. This
method of packing is to be used with the present mouthpiece type of
army mask. If the new Tissot type mask is used, a modification of the
packing is desirable in order to eliminate the trouble due to moisture
given off by the absorbent during service condensing on the
eyepieces of the mask and thus impairing the vision of the wearer.
To remedy this defect a 1-in. layer of kupramite at the top of the
canister is replaced by activated charcoal or silica gel, preferably
silica gel. This decreases the humidity of the effluent air sufficiently
to prevent dimming of the eyepieces. If charcoal is used, a 2-8 cotton
pad (Eastern Star Furrier Co., Pawtucket, R. I.) is substituted for the
toweling in order to remove charcoal dust. The canister complete
weighs about 1.7 lbs.
Fig. 65.—Ammonia Canister—“Kupramite.”

A canister containing 45 cu. in. of this material will protect a man
breathing at rest for at least 5 hours against 2 per cent ammonia and
for 2½ hours against 5 per cent ammonia. Its advantages are large
capacity and activity, negligible heat of absorption, and cheapness.
Physiological Features of the Mask

For some time after the introduction of gas warfare, the gases
used were of the so-called non-persistent type. Under these
conditions it was necessary to wear the mask for only relatively short
periods, after which the cloud dissipated. With the increasing use of
gas and the introduction of the more persistent gases, particularly
mustard gas, it not only became necessary to wear the mask for long
periods of time but also to do relatively heavy physical work, such as
serving artillery, when wearing the mask.
Fig. 66.—Ammonia Mask,
Showing Relative Size of Canister.
Under these conditions, it became evident that the wearing of the

mask caused a very great reduction in the military efficiency of the
soldier. The reasons for this reduction in efficiency have been made
the subject of extensive research by a group of the foremost
physiologists and psychologists of the country. As a result of their

Get Slatter s Fundamentals of Veterinary Ophthalmology Fourth Edition David Maggs Bvsc(Hons) Davco PDF ebook with Full Chapters Now

Uploaded by

Copyright:

Available Formats

Get Slatter s Fundamentals of Veterinary Ophthalmology Fourth Edition David Maggs Bvsc(Hons) Davco PDF ebook with Full Chapters Now

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Get Slatter s Fundamentals of Veterinary Ophthalmology Fourth Edition David Maggs Bvsc(Hons) Davco PDF ebook with Full Chapters Now

Uploaded by

Copyright:

Available Formats

Download the full version of the ebook at

Slatter s Fundamentals of Veterinary

Explore and download more ebook at https://ebookultra.com

Notes on Veterinary Ophthalmology 1st Edition Sheila M.

Fundamentals of Matrix Computations 2nd Edition David S.

Fundamentals of Matrix Computations 2nd Edition David S.

Biological Performance of Materials Fundamentals of

Lishman s Organic Psychiatry A Textbook of Neuropsychiatry

Primary Care Ophthalmology Textbook 2nd Edition David

Pediatric Ophthalmology and Strabismus 3rd Edition David

BSAVA Manual of Canine and Feline Ophthalmology 3rd

SLATTER’S FUNDAMENTALS OF VETERINARY OPHTHALMOLOGY,

Copyright © 2008, 2001, 1990, 1981 by Saunders, an imprint of Elsevier Inc.

Library of Congress Control Number 2007924244

Publishing Director: Linda Duncan

Last digit is the print number: 9 8 7 6 5 4 3 2 1

Itamar Aroch, DVM, DECVIM-CA Gila A. Sutton, DVM, MSc, DACVIM-LA

STRUCTURE AND FUNCTION 1

FUNDAMENTALS OF VISION VASCULAR ANATOMY AND PERIPHERAL OCULAR REFLEXES

V ision is a complex phenomenon in which light emanating

Tarsal gland Retina

B Sensitivity to Flickering Lights

Figure 1-4. Cellular tapetum of a dog

with both eyes simultaneously does not guarantee improved

superior if the images can be blended into one, monocular

optical texture. In addition, movement of the head results in an

N (a phenomenon known as parallax) and produces the sensation

OCULAR VISION OCULAR VISION

BLIND AREA BLIND AREA

light as occurs in dogs typically requires that a greater number

system, which passes without further synapse to the visual

N. olfactorius l Olfactory bulb Olfactory peduncle

A. ethmoidalis interna Olfactory tracts:

A. cerebri caudalis Mammillary body

A. cerebelli caudalis Trapezoid body

Crista petrosa Canal for trigeminal nerve

Canal for transverse sinus Internal acoustic meatus

Cerebellar fossa Jugular foramen

Hypoglossal canal Condyloid canal

Lacrimal gland Ext. ophthalmic a.

M. obliquus ventralis Middle cerebral a.

M. rectus ventralis Int. carotid a.

Anterior ciliary artery

Figure 1-20. Vascular supply of the canine eye.

Nerve Supply of the Eye and Adnexa

Optic nerve (sensory)

1. Round and orbital foramina fuse in the pig and

Figure 1-22. Nerve supply to the eye.

Major petrosal nerve Tensor tympani muscle

Trigeminal nerve Chorda tympani

Abducent nerve Minor petrosal nerve

via oculomotor nerve Ciliary muscle

Cranial cervical Sympathetic chain ganglia

Facial Nerve (Cranial Nerve VII) Autonomic Innervation

To scutuloauricularis superficial dorsal muscle

IIC, ventral branch

Caudal auricular Ramus 1

Facial nerve VII

External jugular vein

Parasympathetic Supply OCULAR REFLEXES