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Third Edition
Introduction to
VETERINARY
SCIENCE
James B. Lawhead • meecee Baker
Copyright 2017 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203
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Introduction to
VeterInary ScIence
Introduction to
VETERINARY
SCIENCE
Third Edition
JAMES B. LAWHEAD, V.M.D.
MEECEE BAKER, PH.D.
Australia • Brazil • Mexico • Singapore • United Kingdom • United States
Introduction to Veterinary Science, © 2017, 2009 Cengage Learning
Third Edition
WCN: 01-100-101
James B. Lawhead
MeeCee Baker ALL RIGHTS RESERVED. No part of this work covered by the copyright
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Publisher does not warrant or guarantee any of the products described
herein or perform any independent analysis in connection with any of the
product information contained herein. Publisher does not assume, and ex-
pressly disclaims, any obligation to obtain and include information other
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Printed in the United States of America

Print Number: 01 Print Year: 2015
Dr. Baker dedicates her efforts in producing this text
to her daughter, Elizabeth “Libby” Baker-Mikesell.
CONTENTS
Preface ix
About the Authors xv
Acknowledgments xvii
Unit I ■ Comparative Anatomy and Physiology

CHAPTER 1 Basic Cell Biology 2 CHAPTER 5 The Respiratory System 78
Cell Makeup ..................................................... 3 The Respiratory Tract
T ......................................79
Cell Structure ................................................... 6 Mechanisms of Breathing ...............................83
Cell Function .................................................... 8 Clinical Practice..............................................84
Protein Synthesis............................................10
CHAPTER 6 The Renal System 89
Mitosis and Cancer.........................................11
Renal System Structures ................................91
Mammalian Reproduction ..............................12
Renal System Functions .................................94
Clinical Practice..............................................13
Kidney Structures and Urine Formation and
CHAPTER 2 Tissue Types and Functions 18 Regulation......................................................95
Epithelial Tissues............................................20 Urine and Blood Evaluation ............................98
Connective Tissues .........................................24 Clinical Practice............................................102
Muscle Tissues ...............................................25 CHAPTER 7 The Digestive System 107
Nerve Tissues .................................................28 Digestive System Structures .........................109
Clinical Practice..............................................29 Monogastric Digestion ..................................116
CHAPTER 3 The Musculoskeletal System 33 Species Variation..........................................120
Musculoskeletal System Functions .................34 Clinical Practice............................................123
Bone Structure ...............................................35 CHAPTER 8 The Reproductive System 130
Joint TTypes and Movements ............................36 Male Anatomy and Hormonal Function .........132
Axial and Appendicular Skeletons...................37 Female Anatomy and Hormonal Function .....134
Bone Growth and Remodeling ........................45 Pregnancy and Parturition.............................138
Relation of Bones, Muscles, Clinical Practice............................................143
and Movement ...............................................46
Clinical Practice..............................................48 CHAPTER 9 The Nervous System 151
Neuron Function...........................................153
CHAPTER 4 The Circulatory System 54 Brain Structure and Function........................156
Blood Components and Functions..................56
Anatomy and Function of the
Mammalian Heart Structures..........................60 Spinal Cord ..................................................158
Blood Vessels and Blood Flow ........................61 Sensory Somatic and Autonomic Nervous
Electrocardiograms, Heart Sounds, and Systems .......................................................159
Blood Pressure ...............................................66 Clinical Practice............................................164
Clinical Practice..............................................72
vii
viii Contents
CHAPTER 10 The Endocrine System 169 CHAPTER 11 The Immune System 184
Endocrine System.........................................171 Antigens and Immunity.................................186
Endocrine Glands .........................................172 Immunity and Immune Response .................190
Clinical Practice............................................179 Clinical Practice............................................194
Unit II ■ Nutrition
CHAPTER 12 Basic Nutrients 202 Pet Food Labels............................................221
Nutrients ......................................................204 Equine Nutrition and Fiber Digestion ............225
Clinical Practice............................................214 Ruminant Nutrition and Fiber Digestion ........228
Clinical Practice............................................231
CHAPTER 13 Species Comparison 218
Animal Nutrition ...........................................220
Unit III ■ Diseases

CHAPTER 14 Principles of Infectious Disease 236 CHAPTER 16 Classification of Diseases 273
Koch’s Postulates .........................................238 Disease in Clinical Practice ..........................275
Disease Agents.............................................239 CHAPTER 17 Zoonoses 289
Clinical Practice............................................255 Zoonotic Diseases ........................................291
CHAPTER 15 Disease Prevention 263 Clinical Practice............................................298
Disease Prevention .......................................265 CHAPTER 18 Diagnosis of Disease 301
Vaccines.......................................................267 Disease Diagnosis........................................302
Clinical Practice............................................269 Clinical Practice............................................313
Unit IV ■ Surgery
CHAPTER 19 Principles of Surgery 320 Laceration Healing........................................326
Principles of Surgery.....................................323 Surgical Considerations ................................333
Unit V ■ Professionalism and Careers

CHAPTER 20 Safety 342 Educational Requirements for
Safety Regulations and OSHA.......................343 Veterinary Careers ......................................355
Safety in Veterinary Practice .........................344 Decision Making in Veterinary Practice..........356
CHAPTER 21 Careers and Decision Making

in Veterinary Science 351
Career Profiles .............................................353
Glossary 363
Bibliography 370
Index 375
PREFACE
Agriscience programs vary nationwide and most have These new activities will help instructors reinforce
undergone extensive curricular changes within the student learning using a variety of applications.
past decade. Many include advanced placement-type • The new safety chapter provides guidelines to
coursework, such as veterinary science. While teaching help teachers ensure student safety in the class-
agricultural education at Greenwood High School in room and field laboratories, while another new
Millerstown, Pennsylvania, Dr. Baker searched for ma- chapter further explores veterinary careers.
terials to be used in a new veterinary science course.
• Technical material has been further explained by
After a futile hunt, and hearing similar concerns from
the author, Dr. James Lawhead. These expanded
other instructors, Dr. Baker teamed with Dr. Lawhead,
and updated explanations will help students
a practicing veterinarian who served the local area
grasp more advanced material.
where she taught, in an effort to author a veterinary
science text that was both student and teacher friendly. • Additional photos and figures bring the veteri-
The authors believe that two of the most useful nary practice into the classroom, helping to keep
features in this book are the “A Day in the Life” of students engaged.
a veterinarian, coupled with the “Clinical Practice” • The new edition discusses the most current tech-
chapter features. These two elements tie the real-life nology used in veterinary practice, providing a
work of a veterinarian, which can have less than de- look into recent advances in the field of veterinary
sired outcomes, with the technical and, sometimes, dry medicine.
and difficult text material. Therefore, the next time a
student says, “I want to be a veterinarian,” a venture
into Introduction to Veterinary Science will provide the
learner with a realistic preview of both veterinary work EXTENSION TEACHING/LEARNING
and the academic rigor needed to achieve success in the MATERIALS
profession.
Simply put, the goals of this text are to afford learn- Instructor’s Companion Website
ers a base knowledge of veterinary science by moving
The Instructor Resources are now available on the
through topics ranging from the cell to surgery, and
companion website. Updated for the third edition, this
to provide a view of the practice of veterinary medi-
robust suite of teaching resources includes the follow-
cine through the eyes of an experienced practitioner.
ing components to help minimize instructor prep time
Chapters 1 and 2 begin the text with a comprehensive
and engage students:
investigation of cells and tissues. Following chapters
examine the musculoskeletal, circulatory, respiratory, • Instructor’s Guide to Text—The Instructor’s Guide
renal, digestive, reproductive, nervous, endocrine, and provides answers to the end-of-chapter questions
immune systems. The basic physiology learned in the and additional material to assist the instructor in
beginning of the text is then applied in concluding the preparation of lesson plans.
chapters covering nutrition, species differentiation in • PowerPoint—Chapter outlines with images for
nutrition, principles and prevention of infectious dis- each textbook chapter.
ease, disease classification, zoonotic diseases, disease
• Computerized Test Bank in Cognero—Hundreds
diagnosis, and surgery.
of modifiable questions for exams, quizzes, in-
class work, or homework assignments, in an on-
NEW TO THIS EDITION line platform.
• Image Gallery—Hundreds of images from the
• Additional hands-on activities that use easy- textbook that can be used to easily customize the
to-find materials have been added to the chapters. PowerPoint outlines.
ix
x Preface
CHAPTER 1
Each chapter in the textbook begins with
clear educational objectives to be learned Basic Cell Biology
by the student in the reading, a list of
important key terms, and an introduction
overview of the chapter content.
Objectives
Upon completion of this chapter, you should be able to:
■ Explain the molecular makeup of cells. ■ Discuss mitosis and its clinical significance in diseases
■ Identify the basic structures of the cell and their such as cancer.
corresponding functions. ■ Detail meiosis in mammalian reproduction.
■ Review the basic function of the cell. ■ Connect cellular parts and function to clinical veterinary
■ Describe the process of protein synthesis. practice.
Key Terms
anesthetize glucose metabolism active transport
antibiotics diabetes anabolism endocytosis
cancer glycogen catabolism benign
lipid enzymes homeostasis malignant
hydrophilic antibodies diffusion pathologists
hydrophobic exocytosis osmosis
Introduction
The cell is the basic structure of animal life. However, the body but also many different cell types. The com-
the cell contains other structures and molecules. Cells bination of these cell types makes an animal function.
conduct many functions and are also able to reproduce. This chapter will discuss the structure of cells, and how
Animals not only have millions of cells that comprise they work.
Chapter 4 The Circulatory System 55
A Day in the Life

ADR—Ain’t Doin’ Right…
I remember the day in veterinary school when our
stethoscopes arrived. The air filled with excitement as
we listened to our own heartbeats. This instrument be-
came a necessary tool in everyday life as I began to ex-
amine animals. I must admit I felt cool walking around
the hospital in a white lab coat with a stethoscope
Photograph courtesy Richard Musselman.
draped around my neck! It seems like yesterday, even

though more than a few years have passed.
Several months ago I examined a cow that was
ADR—ain’t doin’ right. As I walked into the pen, I
could see she obviously wasn’t feeling well at all. She
appeared quite droopy, had lost a lot of weight, and had
developed a swelling under her jaw. During the phys-
ical, I listened to her heart. It sounded like the noise
FIGURE 4–1 A hedgehog.
from a washing machine in midcycle. The heart made a
sloshing sound with every beat. Using the stethoscope,
I diagnosed hardware disease. The cow had eaten journal article. I had never even met one in real life.
a piece of metal that migrated from the stomach and Therefore, I advised the teacher of my lack of experi- Each
ach chapter features ““A Day in the Life”
lodged close to the heart. The location and structure of ence but agreed to examine Sonic.
the heart provided me with the information necessary to
interpret the symptoms of this disease. Hardware dis-
Sonic arrived at the office in a cage (Figure 4–1).
He looked just like a miniature porcupine. Because
of a veterinarian vignette that relays
ease is often found during my appointed rounds. The
next diagnosis is not.
hedgehogs are nocturnal animals, Sonic was apparently
taking his afternoon nap when he arrived at the office.
James Herriot–type stories with relevance
This week, Dr. Deppen and I were both doing eve-
ning small animal appointments at the office. It was
I disturbed him as I tried to examine his leg. Sonic
jumped and snorted in an attempt to scare me. To be to clinical practice and the real-life work
snowing heavily and we were hoping to finish at a rea- honest, it worked! His prickly quills were quite sharp.
sonable hour. Dr. Deppen was seeing Lucky, a 12-year-
old Schnauzer mix that had a history of having what the
My assistant and I then put on thick leather gloves and
proceeded with the examination. Sonic countered with
of a veterinarian.
owners thought was a seizure. She detected that the another protective measure. He rolled himself into a
dog’s heart rate was too slow and the rhythm was very tight ball, so tight his legs were completely hidden. I
irregular. I had a chance to listen to the dog’s heart as referred to the journal article for help.
well and agreed that we should do more tests to detect Following the recommendations, I anesthetized
the underlying problem. Sonic with an inhalant anesthetic. We placed him in
The author James Herriot portrayed veterinary the large clear mask. The anesthetic was slowly deliv-
work in his best-selling collection of stories, All Crea- ered with every breath. Finally Sonic relaxed enough so
tures Great and Small. Times have changed consider- I was able to have a more thorough look. Once Sonic’s
ably since Herriot practiced. Much more information leg was exposed, the problem was quite obvious. The
and sophisticated medicines and techniques are now rags that Sonic used as a nest had tattered edges with
readily available. Still, I cannot possibly be an expert loose strings. One of these strings had wrapped tightly
on all animals. Last year our office received a call from around his foot and stopped the circulation. The foot
a local school. The sixth grade class mascot, Sonic the had turned dark and was oozing. All mammals rely on
hedgehog, had a sore foot. In this case, my experience circulation to maintain their bodies. What happened to
with hedgehogs was limited to reading just one obscure Sonic’s foot when the blood supply was stopped?
Preface xi
Chapter 3 The Musculoskeletal System 41
Thoracic Vertebrae Lumbar Vertebrae a species. The typical dog has 20 caudal vertebrae, but
this can range from 6 to 23.
The appendicular skeleton includes the bones of
the forelimbs and hind limbs. A study of this part of
the skeleton provides a clear examination of compar-
ative anatomy. Although the same anatomic terms are
used for all mammals, great differences exist in the
numbers and sizes of bones in the mammalian appen-
dicular skeleton. For instance, a dog has four or five
© 2017 Cengage Learning®

toes, whereas a horse has only one.
The forelimb, or thoracic limb, does not have a bony
connection to the axial skeleton. The scapula, or shoul-
der blade, lies flat against the rib cage (Figure 3–15).
The scapula connects to the axial skeleton with a group
of muscles. This attachment allows the scapula to move
Sternum Rib
FIGURE 3–13 Radiograph of a cat, showing the thoracic and lumbar
spine. Ribs and sternum are also visible.
over the rib cage. This rotation ranges as high as
25 degrees in animals such as cats while running. This Each chapter contains combinations
flexibility is also useful in cats as they land after a jump.
attach to these vertebrae, forming a sling that supports
As the cat falls, it extends its front legs fully at both the
scapula and the elbow. As the front feet hit the ground,
of charts, illustrations, photographs,
internal organs.
The sacrum, a group of three sacral vertebrae,
fuses to support the pelvis (Figure 3–14). In addition,
the elbow flexes and the scapula rotates. The cat makes
this very coordinated act look quite graceful. Clinically, radiographs, and the like that help to
this is of significance when cats fall from extreme
the sacrum articulates with the last lumbar vertebra
and the first caudal vertebra. The sacrum then joins
heights. In large cities, this happens often as cats tumble
from balconies or windows of tall buildings. In high-rise
illustrate and enhance the concepts
with the pelvis, allowing the hind limbs to support the
presented.
syndrome, the falling cat rarely breaks a leg; however,
weight of the body. This connection can be damaged. it will often break its lower jaw. The high speed of the
The pelvis may split away from the sacrum when falling cat forces the jaw to contact the ground.
dogs and cats are hit by cars (HBC). During this type
of accident, fracture of the pelvis itself is also com-
mon. Very painful lameness often results from a split
pelvis or pelvic fracture. Many of these fractures heal
if the animal’s activities are restricted. In severe cases,
surgeries may be required.
The final group of vertebrae is called caudal. These
small vertebrae comprise the tail. As mentioned, the
numbers of vertebrae vary among species and within
Lumbar Vertebrae
Pelvis
Spine
FIGURE 3–14 Radiograph of the lumbar spine of a dog. A portion of the

pelvis is also visible. This dog is showing an age-related change called Chapter 1 Basic Cell Biology 5
spondylosis. In spondylosis, bone spurs are formed that can eventually
bridge between vertebrae. FIGURE 3–15 The scapula.
Purines Pyrimidines
NH2 O O
C N C N C
N C HN C HN C C 3
CH
CH CH
HC C C C O C CH
N N N N N
H2N
Thymine
Adenine Guanine
NH2
C
N CH
Base
(Purine or O C CH
Pyrimidine) N
OH Cytosine
HO P O CH2 O
O
O C C
H H C
H H HN CH
C C
O C CH
OH H N
OH Uracil
Deoxyribonucleotide
(In Ribonucleotides)
FIGURE 1–5 Chemical structure of a nucleotide.
allowing veterinarians to diagnose what specific organ- Notice that the bases are the same except for thymine
ism is causing the sickness. and uracil. The order of base combination determines
Nucleic acids provide plans for the differing con- what amino acids are used to make proteins. This infor-
struction of proteins. Nucleic acids are fabricated with mation is stored in the cell’s genetic material.
a series of nucleotides. The nucleotides are made up Both DNA and RNA have a backbone of sugar al-
of a five-carbon sugar, a phosphate group, and a ni- ternating with phosphate. The nitrogenous bases are
trogen-containing base (Figure 1–5). Ribonucleic acid attached to this backbone. In DNA, a double-stranded
(RNA) claims ribose as its sugar, whereas deoxyribo- molecule is formed as the bases are loosely bonded
nucleic acid (DNA) has deoxyribose as its sugar. There together. The molecule has a twisted structure, which
are four different bases for RNA and DNA (Table 1–1). is described as a double helix (Figure 1–6). The bases
join, specifically, thymine to adenine and cytosine
to guanine. Later in the chapter, a process of tran-
scription will be described, in which the sequence of
Table 1–1 RNA and DNA Bases DNA nitrogenous bases is converted to a molecule
DNA Bases RNA Bases of RNA. In this situation, adenine in the DNA mole-
cule bonds to a uracil base of RNA. The sequence of
1. Adenine 1. Adenine nitrogenous bases is used to define the amino acids
2. Cytosine 2. Cytosine used in protein synthesis. A group of three nitroge-

nous bases is the code for a specific amino acid. The
3. Guanine 3. Guanine order of the nitrogenous bases makes up the genetic
code of the animal. Each gene provides the code for
4. Thymine 4. Uracil one peptide chain.
xii Preface
10 Unit 1 Comparative Anatomy and Physiology
Cytoplasm Extracellular enzyme begins at a specific series of bases (thymine,

Fluid adenine, cytosine) called a promoter. The RNA poly-
merase moves along the length of the DNA molecule,
creating a complementary strand of RNA. The RNA
bases are added in the specific order that bonds to the
bases of the DNA. The corresponding bases were dis-

cussed earlier in the chapter. This process continues un-
til the polymerase reaches a terminator series of bases
(adenine, thymine, thymine). The mRNA is released
and the DNA helix reconnects.
FIGURE 1–12 Endocytosis: A large particle is engulfed by the cell
membrane and brought into the cytoplasm within a vacuole.
DNA mRNA
Double Helix Strand
Cytoplasm Extracellular
Fluid C G C
G C G
A T A

C G C
A T A
A T A
FIGURE 1–13 Exocytosis: A membrane-bound sac joins with the cell C
C
membrane to release the particle.
T U A
C C G
(Figure 1–12). During endocytosis, the cell membrane
wraps around the particle, pinches off, and moves into T U A
the cytoplasm as a vacuole. Lysosomes then join with
the vacuole, providing the enzymes necessary to break G G C
down the particle. The smaller fragments produced are C C G
then released into the cell. RNA
In cells producing protein, the opposite process oc- T Polymerase U A
curs. In exocytosis, a membrane-bound sac containing A A T
the protein joins with the cell membrane and releases
A
it into the ECF (Figure 1–13). These sacs are produced T U
within the Golgi apparatus. In intestinal cells, fat drop- G G C
lets can be taken into the cell through endocytosis. The
C G RNA
vacuole is transported across the cell and released into C
Nucleotide
the bloodstream by exocytosis. T
A
G C
PROTEIN SYNTHESIS
T A
Objective
C G
■ Describe the Process of Protein Synthesis
G C
As mentioned previously, every cell contains all the
genetic material of the animal. The expression of cer-
tain genes produces specific proteins that allow cell

Cytosine Uracil
specialization. Protein synthesis begins within the nucleus
on the basis of the DNA structure. During transcription, Adenine Thymine
information within the DNA is transferred to a strand of
messenger RNA (mRNA) that moves into the cytoplasm. Guanine
Chapter 3 The Musculoskeletal System An enzyme called RNA polymerase binds to FIGURE 1–14 Transcription of mRNA: RNA polymerase separates the
DNA, causing a separation of the double-helix strands strands of DNA and creates a strand of mRNA coded by the nucleotides
(Figure 1–14). This pulling apart exposes a gene. The of the DNA molecule.
cat from that of a horse. Having muscles closely associ- Articular
ated with the skeleton provides movement of the bones Cartilage
Proximal Epiphysis
at a joint. The movement of bones allows locomotion
Bone Physis
and function of the animal. Marrow Metaphysis
The strength of bones also protects more fragile
tissues. The rib cage gives protection to the heart and Cancellous or
lungs, whereas the skull protects the delicate brain. Spongy Bone
Bone acts as a reservoir for calcium and phosphorus.
In times of need, the minerals are moved from the bone
Medullary Cavity
and sent into the bloodstream. Excess minerals can be
stored in the bone. Calcium plays an essential role in
muscle contraction and enzyme activity. Phosphorus is Artery
necessary for energy metabolism within the cell. Bone,
in response to several hormones, maintains a tight A Compact Diaphysis
regulation on the blood level of these minerals. These Bone Tissue
hormones, calcitonin and parathyroid hormone, will be
Endosteum
discussed in much greater detail in Chapter 10.
The long bones are present in the legs (and arms in
humans). The femur and humerus are classified as long
bones. They have a dense outer shell and a hollow shaft.
Bone marrow is made in this hollow center, the medul-
Periosteum
lary cavity. Bone marrow in turn produces blood cells.
Metaphysis
Physis
BONE STRUCTURE Distal Epiphysis Each chapter is further enhanced by the

Objective
■ Detail the Structure of Bone
addition of repeat objectives to aid in
Splitting a long bone along its length shows the typ-
ical structure of bone (Figure 3–3). The outer shell is
student comprehension.
composed of dense or compact bone. The term cortical
bone is also used for this region. The greater the forces
placed on a bone, the thicker this layer will be. In the
femur, this compact bone is thickest in the middle of
the shaft, where greatest strain occurs.
Within compact bone lies a more loosely arranged
bone, called spongy or cancellous bone. Spongy bone B
is found within the long bones but not inside the flat
bones of the skull or pelvis. It only fills the ends of
these long bones. Spongy bone is made up of tiny
spicules and plates of bone. The spicules look random
but are actually arranged to maximize strength. The
spongy arrangement keeps the weight of the bones

much lighter than that of a solid bone of the same di-
mension. The medullary cavity is located in the hollow
center of the shaft. The bone marrow lies within the
medullary cavity and the spaces of the spongy bone. As
FIGURE 3–3 A. Illustration of bone structure. B. Photograph of the internal
mentioned earlier, bone marrow produces blood cells. structure of bone.
Bones are covered with a thin connective tissue
called the periosteum. The periosteum blends into
tendons and ligaments, binding them to the bone. The provides protection as the bones move against one
periosteum has an extensive blood and nerve supply. another within a joint. The open spaces within bone
Hence trauma to the periosteum is quite painful. The are covered with a similar connective tissue, the en-
portion of bone within the joint is covered with car- dosteum. Both the periosteum and endosteum provide
tilage and not by periosteum. This articular cartilage cells necessary for the repair of damage.
Preface xiii
SUMMARY
Being able to identify respiratory structures and their rate provides a key piece of information to practition-
associated functions, from the nose to the lungs, allows ers when assessing the overall health of animals. The
veterinarians to diagnose and treat such disease condi- status of the respiratory system affects the breathing
tions as pneumonia and roaring. Moreover, respiratory and therefore the total health of animals.
REVIEW QUESTIONS
1. Define any 10 of the following terms: 5. The human larynx is sometimes called the .
respiration 6. The trachea branches into two .
palpated
endotracheal tube
7. Gas exchanges occur in the smallest openings of A chapter summary highlights the topics
the respiratory system. These openings are called
inspiration
expiration
the . that have been presented, and the end of
cyanosis 8. The muscles between the ribs are called the .
pneumonia
pleural friction rub
9. Name the reflex action that occurs when there is each chapter is also followed by a series
an irritation in the nose.
contagious
roaring 10. What substance lines the lungs, making them eas- of review questions and student activities.
heaves ier to inflate?
bronchodilators 11. What controls the rate of respiration?
2. True or False: Mucus lines the epithelial tissue in 12. What is the normal respiration rate for a dog?
the nostrils.
13. What plays a more significant role in the control of
3. True or False: The cartilage rings of the trachea are respiration, oxygen, or carbon dioxide?
shaped like an O.
14. What medical tool is used to evaluate breathing?
4. The is the common area shared by the
15. What species can develop a condition referred to
nose and throat.
as roaring?
ACTIVITIES
Materials needed for completion of the activities: release the balloons yet. Plug the third opening of
stethoscope the Y piece. Hypothesize what will happen when
the balloons are released. Will the large balloon
balloons
Y-shaped polypropylene connecting tubes deflate and fill the smaller balloon to equalize the
size? Or will the smaller balloon deflate into the
1. Use the stethoscope to listen to normal lung other balloon? Surfactant prevents this problem
sounds. Have the “patient” take deep, slow from occurring between alveoli. Even though the
breaths. The patient should breathe quietly, not alveoli may be of different sizes, the pressure in
making noise through the nose and mouth. The each is similar. Without it, the small alveoli would
stethoscope can detect these noises. Listen to dif- deflate.
ferent areas on the chest, from both the front and 3. Observe the respiratory rates of classmates and
the back. pets or livestock. Compare to the normal rates
2. Take two identical balloons and inflate them listed in Table 5–1.
to different sizes. Slip a balloon onto an end of Chapter 21 Careers and Decision Making in Veterinary Science 355
Y-shaped polypropylene connecting tubes. Do not
Courtesy of Dr. David Sweet.

FIGURE 21–4 Dr. David Sweet, with assistance of registered veterinary
technician Michele Antoch, examines a surgical incision on a dog.
Courtesy of Dr. Cathy Hanlon.
EDUCATIONAL REQUIREMENTS FOR

VETERINARY CAREERS
Objective
FIGURE 21–3 Dr. Hanlon working with a sedated raccoon that had been ■ Explain the Educational Requirements for a Variety of
captured in a live trap. Veterinary Careers
Veterinary technicians must complete either a two-
methods for prevention of rabies, and responds to year associate degree or four-year bachelor of science
questions about rabies from other public health profes- degree program. Further, they must pass a state
sionals and the public. licensing exam. The number of institutions offering
such coursework has grown significantly over the past
Veterinary Surgeon several years. Veterinary assistants are not required
Dr. David Sweet graduated in 1989 from the University to complete any formal classes. However, increasing
of Pennsylvania School of Veterinary Medicine. Fol- numbers of technical schools and community colleges
lowing his graduation, Dr. Sweet pursued further offer veterinary assistant programs.
training as an intern at the University of Pennsylvania Both technicians and assistants help the veter-
and a surgical residency at the North Carolina State inary practice by performing a wide range of tasks
University. Following that training, Dr. Sweet accepted (Figure 21–5). These individuals may greet patients,
an instructorship at Washington State University and keep records, bill clients, and restrain animals, as well
returned to the University of Pennsylvania as an assis- as feed, exercise, and provide basic health care for
tant professor. During his training, Dr. Sweet met the patients. The responsibilities vary from employer to
The new chapter on careers investigates rigorous qualifications necessary to become a diplomate
in the American College of Veterinary Surgeons. This
employer with technicians performing more technical
duties. Numbers of available jobs for veterinary assis-
occupations in veterinary science, honor earned by Dr. Sweet distinguishes him as a sur-
gical specialist.
tants and technicians will continue to grow with the
demand for veterinarians.
Dr. Sweet works at a referral practice. The cen- Level of degree separates veterinary assistants
including veterinary technicians, veterinary ter employs veterinary specialists in many fields,
including surgery. The veterinary practice provides
from veterinary specialists. Almost 30 programs grant
degrees in veterinary specialties. Most of these pro-
grams deliver master’s and doctorate degrees, al-
assistants, private practitioners, and
a service that allows private practitioners to refer
difficult cases for more specialized treatment. Dr. though a few award associate and bachelor’s degrees.
Sweet performs both soft tissue and orthopedic sur- Specialists may provide such supportive services as
veterinary specialists. gery (Figure 21–4). He performs many complicated
and difficult surgeries. As with all veterinarians, he
nutrition counseling, ration balancing, or radiology ex-
pertise to veterinary clinics. Conversely, other special-
attends continuing education conferences to learn ists may be employed in academia, where they perform
new procedures and information. research or extension duties in veterinary-related
ABOUT THE AUTHORS
Dr. James Lawhead is a veterinarian in a private Harrisburg, Pennsylvania, and Washington, D.C. In
mixed animal practice located in Millerstown, Penn- addition, Dr. Baker serves as an adjunct professor at
sylvania. As lead partner, he works primarily with the North Carolina State University. She earned both
dairy cattle, dogs, and cats. Dr. Lawhead joined this her bachelor’s and doctorate degrees from Pennsyl-
practice in 1987 following graduation from the Univer- vania State University in agricultural education and
sity of Pennsylvania School of Veterinary Medicine. a master’s of science degree from the University of
He gained acceptance to veterinary school following Delaware in agricultural economics. Dr. Baker was
completion of his bachelor’s degree at Juniata College. the first woman to be elected president of the Na-
Dr. Lawhead has a special interest in dairy cattle nu- tional Vocational Agriculture Teachers’ Association
trition, providing nutritional services to a number of (now known as the National Association of Agricul-
his clients. Dr. Lawhead enjoys teaching as well and tural Educators). Formerly, she taught high school
actively supports local school districts with lectures agriculture and worked in the executive office of
and demonstrations. the Pennsylvania Department of Agriculture as co-
ordinator of agricultural education. Dr. Baker lives
Dr. MeeCee Baker owns Versant Strategies, an ag- on her family beef farm with her husband, Jim, and
ricultural and rural affairs firm that serves clients in daughter, Libby.
xv
ACKNOWLEDGMENTS
Although only two authors are listed for this text, the freshen objectives and questions for the third edition.
number of people responsible for the final product is We appreciate the use of reference material supplied
quite large. The authors would like to thank all of those by Mechelle Regester. The veterinary science students
people who supported and contributed to the text, es- at Greenwood High School completed activities, les-
pecially the Cengage Learning Team. Cengage Learn- sons, and accompanying assignments to help fine-
ing deserves special recognition for faith in the authors. tune the text and ancillary material. We appreciate
We would like to thank all the veterinarians and their thoughtful consideration.
staff at Millerstown Veterinary Associates for their In addition, we would like to thank Dr. David
assistance and contributions. Their help in obtaining Sweet, Dr. Cathy Hanlon, Dr. Abby Maxson Sage, and
case material and photographs for the text was in- Dr. Lawrence Hutchinson for their contributions of
valuable. Likewise, we appreciate the support of the photographs and support to the project.
clients who encouraged the use of their case material Having input from experts in various fields helped
for the text. Special thanks are in order for Leesa to strengthen the core material of the text. Our utmost
Landis, Dr. Robert Mikesell, and Krista Pontius for thanks to Dr. William Bacha Jr., Dr. Linda Bacha, and
their long hours of technical help in putting together Dr. Arthur Hattel for the photographic material pro-
the text. Caleb Wright, a Versant intern and newly vided. The histology and pathology photographs are a
minted agricultural education teacher, helped to tremendous benefit to the text.
xvii
Unit I
Comparative Anatomy
and Physiology
CHAPTER 1
Basic Cell Biology
Objectives
Upon completion of this chapter, you should be able to:
■ Explain the molecular makeup of cells. ■ Discuss mitosis and its clinical significance in diseases
■ Identify the basic structures of the cell and their such as cancer.
corresponding functions. ■ Detail meiosis in mammalian reproduction.
■ Review the basic function of the cell. ■ Connect cellular parts and function to clinical veterinary
■ Describe the process of protein synthesis. practice.
Key Terms
anesthetize glucose metabolism active transport
antibiotics diabetes anabolism endocytosis
cancer glycogen catabolism benign
lipid enzymes homeostasis malignant
hydrophilic antibodies diffusion pathologists
hydrophobic exocytosis osmosis
Introduction
The cell is the basic structure of animal life. However, the body but also many different cell types. The com-
the cell contains other structures and molecules. Cells bination of these cell types makes an animal function.
conduct many functions and are also able to reproduce. This chapter will discuss the structure of cells, and how
Animals not only have millions of cells that comprise they work.
2
Chapter 1 Basic Cell Biology 3
A Day in the Life

There Just Never Seems to be a Typical Day . . .
I headed to the office with the thought of doing only cow
work on this particular day. However, those plans were
short lived. Shortly after I arrived at work, two nervous
owners walked through the door with their Labrador re-
triever. Poor Jake had just been run over by the own-
er’s car! Amazingly, Jake was doing very well, although
he was a bit excited. Apart from a couple of cuts on his
jaw, he was ready to go home and play.
Then, at my first farm call of the day, the farmer
wanted me to look at his dog, Millie. Millie had a grape-

fruit-size lump under her jaw. The lump felt like it was
full of fluid. I asked him to bring Millie to the office so
I could work on her there. I finished my farm calls and
headed back to the small animal clinic.
Once there, I anesthetized Millie and made an in-
cision into the skin. Pus flowed from the lump (Figure FIGURE 1–1 Draining an abscess on the side of the face of an
1–1). I flushed the large pocket left behind and started anesthetized cat.
Millie on a course of antibiotics, drugs that fight bacte-
rial infections. Although I do not know why it started, I surgery. The surgery went well, and I was able to re-
do know Millie was fighting an infection with her body’s move the entire lump.
cells. In private practice, cells affect me every day. Today
Next I had the opportunity to remove a tumor from I saw Millie’s cells attacking the bacteria in her neck.
Penny, a 12-year-old cocker spaniel. Last week I gave Penny, on the other hand, had cancer
cancer-causing cells divid-
Penny a physical examination and administered blood ing uncontrollably. To understand how mammals work and
tests. Penny appeared healthy, and we elected to do how to treat them, I first had to learn how cells function.
CELL MAKEUP of these molecules. They possess the basic structure of

(CH2O)n (Figure 1–3). In this formula, n describes the
Objective number of carbon atoms in the molecule. The genetic
■ Explain the Molecular Makeup of Cells
material in the cell has the five-carbon sugars ribose
and deoxyribose. Glucose (blood sugar), a six-carbon
Cells and their structures are composed of molecules. sugar, is used for energy in the cells. The amount of
Biochemistry is the study of these molecules in living glucose in blood is routinely monitored. If there is too
creatures. One goal of this chapter is to identify the dif- much or too little glucose in the blood, the animal will
fering types of molecules and their properties. not function normally. In diabetes, the blood sugar
Lipids or fats combine hydrogen, carbon, and oxy- increases to very high levels, but the animal does not
gen in a form that is poorly dissolvable in water (this utilize it properly. Diabetes requires treatment to lower
is why fat floats to the top of water). Fat consists of the blood sugar.
a molecule of glycerol and three fatty acid molecules Polysaccharides are composed of many monosac-
(Figure 1–2). Fats are stored in the cells of the body as charides. One example of a polysaccharide is starch,
a source of high energy. such as glycogen, which is used to store energy within
Phospholipids are similar but have only two fatty the cell. Glycogen is made when monosaccharides
acid groups and a phosphate group (PO4). This is sig- are taken into the cell and then assembled into a long
nificant because one end of the molecule is attracted to chain. Polysaccharides can be joined with protein mol-
or soluble in water (hydrophilic) and the other end is ecules to form glycoproteins, which assist in building
repelled by water (hydrophobic). These characteristics the cell structure.
of phospholipids are important in the structure of the Proteins play a key role in the structure and func-
cell membrane. tion of cells. Proteins make up 50% of the dry weight of
Carbohydrates supply energy and provide struc- animals. Proteins are large molecules of many amino
ture within the cell. Monosaccharides are the simplest acids. (Twenty-two different amino acids are used to
H NH
3
H C OH CH2
O O
C CH2
H C OH
CH2 CH2
H C OH
CH2 CH2
H
H2N C C OH H2N C C OH
Glycerol
H O H O
O Glutamic Acid (Glu) Lysine (Lys)
HO C R
RLong Hydrocarbon Chain Fatty Acid

OH
C
H O H H H H H H H HC CH
H C O C C C C C C C C H HC CH H3C CH3
C CH2
H H H H H H H
CH2 CH2
O H H H H H
H2N C C OH H2N C C OH

H C O C C C C C C H
H O H O
H H H H H
Tyrosine (Tyr) Leucine (Leu)
O H H H H H
H C O C C C C C C H
FIGURE 1–4 Chemical structure of selected amino acids.

H H H H H H
Lipid
make proteins; Figure 1–4.) A single protein can include
200 to 300 of these amino acids. It was mentioned earlier
that proteins could be joined to sugars. They may also
FIGURE 1–2 Chemical structure of glycerol, a fatty acid, and a typical
be joined with lipids and phosphate groups. Protein
lipid.
molecules are not only very large but also quite com-
plex molecules. Chemical bonding between amino acids
will fold the amino acid chains into a three-dimensional
structure. This complex structure is essential for the
function of certain protein molecules.
Proteins have many functions in cells. Muscle is
CH2OH largely composed of protein that is specially arranged
to allow cells to contract and move. Further, enzymes
C O
H H CH2OH O OH are protein molecules that speed the chemical reactions
H in the body (i.e., enzymes act as catalysts). Proteins also
C C C C
OH H H OH add strength to many of the structures in the body.
HO OH H CH2OH Proteins are found within the cell membrane and are
C C C C
commonly found in the intercellular matrix of tissues.
H OH OH H Protein can bind with other molecules to aid in their

transport in the bloodstream. In addition, proteins
Glucose Fructose found in blood help to carry oxygen, stop bleeding,
and fight off infection. These infection-fighting proteins
are called antibodies. In practice, antibodies specific
FIGURE 1–3 Chemical structure of selected sugars. to different diseases are measured in the blood, thus
Purines Pyrimidines
NH2 O O
C N C N C
N C HN C HN C C 3
CH
CH CH
HC C C C O C CH
N N N N N
H2N
Thymine
Adenine Guanine
NH2
C
N CH
Base
(Purine or O C CH
Pyrimidine) N
OH Cytosine
HO P O CH2 O
O
O C C
H H C
H H HN CH
C C
O C CH
OH H N

OH Uracil
Deoxyribonucleotide
(In Ribonucleotides)
FIGURE 1–5 Chemical structure of a nucleotide.
allowing veterinarians to diagnose what specific organ- Notice that the bases are the same except for thymine
ism is causing the sickness. and uracil. The order of base combination determines
Nucleic acids provide plans for the differing con- what amino acids are used to make proteins. This infor-
struction of proteins. Nucleic acids are fabricated with mation is stored in the cell’s genetic material.
a series of nucleotides. The nucleotides are made up Both DNA and RNA have a backbone of sugar al-
of a five-carbon sugar, a phosphate group, and a ni- ternating with phosphate. The nitrogenous bases are
trogen-containing base (Figure 1–5). Ribonucleic acid attached to this backbone. In DNA, a double-stranded
(RNA) claims ribose as its sugar, whereas deoxyribo- molecule is formed as the bases are loosely bonded
nucleic acid (DNA) has deoxyribose as its sugar. There together. The molecule has a twisted structure, which
are four different bases for RNA and DNA (Table 1–1). is described as a double helix (Figure 1–6). The bases
join, specifically, thymine to adenine and cytosine
to guanine. Later in the chapter, a process of tran-
scription will be described, in which the sequence of
Table 1–1 RNA and DNA Bases DNA nitrogenous bases is converted to a molecule
DNA Bases RNA Bases of RNA. In this situation, adenine in the DNA mole-
cule bonds to a uracil base of RNA. The sequence of
1. Adenine 1. Adenine nitrogenous bases is used to define the amino acids
2. Cytosine 2. Cytosine used in protein synthesis. A group of three nitroge-

nous bases is the code for a specific amino acid. The
3. Guanine 3. Guanine order of the nitrogenous bases makes up the genetic
code of the animal. Each gene provides the code for
4. Thymine 4. Uracil one peptide chain.
P
Protein
Prote
rotein
in
Phospholipid
Phos
Phosph
phoolipid
lipid

R
Repelled
Repe
epelled
epelllle
ed
d by Water
Water
A
Attracted
Attracte
ttracted
d to Water
Water
Phosphate
Sugar
FIGURE 1–7 Illustration of cell membrane. The cell membrane has a dou-
ble layer of phospholipid. In addition, protein molecules are present on
Hydrogen Bonds
and within the phospholipid layers.
the inside of the cell contained. The cell membrane is

so fine that it cannot be seen with a normal light micro-
scope. The cell membrane is about half protein and half
lipid (phospholipid type). One end of phospholipids
is attracted to water, whereas the other end is repelled
by water. The cell membrane, which is surrounded by
water on both sides, has two layers of lipid in its wall
(Figure 1–7). The ends of the lipid that are attracted to
water face outward. Protein is also included in the mem-
brane, both between the lipid molecules and on the sur-
face. The position of the protein molecules is not firmly
established; rather, the molecules are mobile within the
membrane. Cholesterol, another molecule in the cell
membrane, provides stabilization of the membrane.
Cell membranes are semipermeable, meaning they
allow certain substances but not others to pass. Some
molecules, such as water, are able to pass through easily.
The specialized proteins in the cell membrane influence

Cytosine Guanine which molecules are able to pass readily. In addition, the
intrinsic membrane proteins can act as receptors. These
Thymine Adenine
receptors can process a signal from the extracellular
fluid to influence the cell’s interior (e.g., a hormone can
FIGURE 1–6 DNA structure: The structure is described as a double helix.
Phosphate and sugar groups make up the two strands. The strands are trigger a reaction within the cell). Other molecules, such
joined by hydrogen bonds between two nitrogenous bases. as proteins, starches, and some ions, are unable to pass.
Many of the organelles within the cell are also sur-
rounded by a membrane. The basic structure remains
CELL STRUCTURE the same for all the membranes. The specifics of the
makeup differ, depending on function.
Objective Cell contents are divided into the nucleus and the
■ Identify the Basic Structures of the Cell and Their cytoplasm. Cytoplasm generally describes the organelles
Corresponding Functions and fluid in the cell. A nucleus comes as a standard part
of most cells (with a few exceptions such as the red blood
Many cell types exist. These cells not only look dif- cell; Figure 1–8). The nucleus contains the genetic material
ferent but function differently as well. Nevertheless, (i.e., DNA) of the cell, which controls cellular activities
many features are common among cells. Specialized by coding for protein synthesis. The DNA in the nucleus
structures within the cells are called organelles. These is called chromatin. As the cell divides, the chromatin
organelles are present in most but not all cells. Red clumps into chromosomes. Identical DNA is passed to
blood cells, for example, lack a nucleus. all daughter cells. All the cells in the body have the same
The cell membrane (or plasma membrane) is com- chromatin. However, cells take on different roles by using
mon to all cells. It serves as the boundary that keeps certain areas of the chromatin more than others.
Ribosomes Centriole Lysosome
Vacuole Nucleolus
Nucleus
Cytoplasm
Rough Endoplasmic
Reticulum
Golgi Mitochondrion
Apparatus

(Complex)
Plasma Membrane
Smooth Endoplasmic Reticulum

FIGURE 1–8 Illustration of cell structure.
A membrane made of two lipid bilayers surrounds Smooth endoplasmic reticulum (SER) has no
the nucleus. This membrane is often joined to other ribosomes attached. This form is not as common. Some
organelles, such as the endoplasmic reticulum and liver cells contain a large amount of SER. The SER in
ribosomes. Such a close association helps the nucleus these cells produces glycogen and lipids, and removes
control cell function. toxins.
In cells not dividing, a nucleolus is often seen in the The Golgi apparatus is formed with large amounts
nucleus. The nucleolus produces RNA that forms the of folded membrane that looks similar to SER. The
ribosomes, which in turn produce protein. Cells with Golgi apparatus produces polysaccharides and special
large nucleoli actively produce protein. protein sacs called lysosomes. Protein produced in
Ribosomes are small granular-like structures that the RER is moved to the Golgi apparatus. The Golgi
can be found in the cytoplasm. They contain roughly apparatus then changes the protein and collects it in
60% RNA and 40% other protein. Ribosomes manufac- the lysosomes. These sacs are pinched from the Golgi
ture the protein used in the cell. Growing cells require apparatus and then moved to the surface of the cell and
large amounts of protein and, therefore, have a greater released.
number of ribosomes. The specific proteins produced The proteins contained in the lysosomes are
by a cell are governed by the nucleus. enzymes (remember, enzymes are molecules that help
The endoplasmic reticulum (ER) is a collection speed chemical reactions in the body). Lysosomes con-
of folded membrane. This membrane attaches to the tain enzymes that help to break down other mole-
membrane of the nucleus. The ribosomes often line cules. Varying enzymes match differing molecules.
this membrane, giving it a bumpy appearance and The membrane surrounding lysosome prevents the
therefore its name, rough endoplasmic reticulum enzymes from attacking other parts of the cell.
(RER). Protein produced by the ribosomes is then Lysosomes are used to digest food taken in by the
deposited into the RER. These proteins can be further cell and to destroy cell structures no longer needed. In
changed in the RER. This protein may be used by the Millie, the dog with the abscess, her white blood cells
cell or moved to the surface of the cell for secretion. were using lysosomes to destroy bacteria. Cells that
The protein is moved through the membrane in a pro- die in the body are eliminated when enzymes within
cess called exocytosis, which will be discussed later lysosomes are released into the cytoplasm. This process
in the chapter. of autolysis makes room for replacement cells.
Mitochondria are small rod-shaped organelles Table 1–2 Components of the Extracellular Fluid
found in varying numbers in cells. The more active the
cell, the more mitochondria are present. Mitochondria
1. Water
have a double membrane, similar to the cell membrane. 2. Dissolved gases: oxygen, carbon dioxide
The outer membrane is smooth and forms the shape of
the mitochondria. The inner membrane is highly folded. 3. Inorganic ions
These shelflike infolded ridges are called cristae. Macrominerals: sodium, potassium, chloride, phosphate,
The role of mitochondria is well defined. The mito- calcium, bicarbonate
chondria convert food substances into energy that can Trace minerals: copper, zinc, manganese, cobalt,
be used by the cell. Mitochondria contain the enzymes selenium, fluoride, iron
necessary for this process. Because of this role, mito- 4. Organic compounds (carbon-containing compounds):
chondria are called the powerhouses of the cell. The proteins, amino acids, lipids, carbohydrates, vitamins

mitochondria are found within cells at their areas of
highest activity. 5. Hormones: compounds produced by glands to influence
metabolism of cells
CELL FUNCTION 6. Waste products
Objective eliminated by the ECF. Without elimination, the waste

■ Review the Basic Function of the Cell products actually become toxic to the cell.
Many of the products in ECF must be maintained
The cell constantly reacts to its environment. Metabolism at constant normal concentrations. Cells will be unable
describes all the reactions going on in cells. Metabo- to function properly if there is too much or too little of
lism can be categorized into two main types. Anabolism certain products. Glucose provides an excellent exam-
describes reactions in which smaller molecules are ple. Small puppies can become low in blood sugar if
combined into larger ones. The joining of amino acids they have too many parasites robbing them of nutri-
to form proteins serves as an example. Catabolism, the ents. When the sugar in ECF becomes too low, the cells
opposite, occurs when large molecules are broken down do not have adequate energy. The puppy can become
into smaller ones. The breaking down of glycogen to weak or, in severe cases, develop a seizure. Homeo-
release energy is an example of catabolism. stasis is the maintenance of ECF. Homeostasis allows
A liquid called extracellular fluid (ECF) surrounds maintenance of normal concentrations of molecules in
living cells. The ECF supplies cells with all the prod- spite of a wide variety of external conditions.
ucts necessary for their functions. ECF is derived from Cells must be able to obtain products from the
blood. The outermost skin cells are not covered in liq- ECF. It is not enough that the chemicals just exist in
uid; however, they are no longer living. the ECF; there must be means for their exchange with
Other cells exposed to the surface, such as those of the cell. Table 1–3 summarizes the mechanisms by
the eye, need moisture. In the eye, tears produced by which materials are exchanged across the cell mem-
glands act as the source of moisture and nutrients. The brane. The first mechanism is a process called diffusion
eyelids help to sweep the tears across the surface of the (Figure 1–9), in which molecules move from higher to
eye. Certain breeds of dogs, such as the pug, have eyes lower concentrations. Because molecules are always
that bulge from the eye socket. The bulging can be so moving, there is a greater chance that they will move
severe that the eyelids cannot keep the surface of the toward areas of lower concentration. This movement
eye moist with tears. This results in a disease condition continues until the concentrations are equalized.
on the surface of the eye. Artificial tears are often used The cell membrane does not allow totally free
to keep the surface moist. diffusion. Diffusion is influenced by the size of the
Table 1–2 summarizes the makeup of ECF. Water molecule, its charge, and its ability to dissolve in lipid.
is the major component of ECF. Oxygen passes to the In general, the smaller the molecule, the more easily
cells through the ECF. Conversely, carbon dioxide
passes from the cells through it. There are many inor-
Table 1–3 Mechanisms of Cellular Exchange
ganic ions in the ECF. Some ions, macrominerals, are
present in large amounts. Trace minerals are present in 1. Diffusion
much smaller amounts. Both macrominerals and trace
minerals are essential for cellular function. Many of 2. Osmosis
the trace minerals are needed for enzymes to function. 3. Active transport
Organic compounds, including the lipids, proteins,
and carbohydrates, are also delivered by the ECF. 4. Endocytosis
Metabolism produces waste products, which must be
5. Exocytosis
removed from the cells. These waste products are
Molecule Hypotonic Solution

Hypertonic Solution
Water

Selectively
Permeable Membrane
FIGURE 1–10 Osmosis: The semipermeable membrane prevents the
passage of large molecules. In this situation, water moves across the
membrane to equalize the concentration.
In certain situations, a cell may require a higher

concentration of a molecule than is found in the ECF.
For example, red blood cells have higher levels of
potassium than the surrounding fluid. Diffusion con-
stantly attempts to equalize the concentrations (e.g.,
potassium continually diffuses from the cell). In this
case, the potassium is pumped back into the cell, and
the higher concentration is maintained. This process
is referred to as active transport (Figure 1–11). Active
transport requires the cell to burn energy and use en-
zymes to aid the process. Many different cell types per-
form the function. Another example occurs in intestinal
cells, which transport glucose into the bloodstream,
where it is present at higher levels.
Large molecules, such as proteins, must be moved

through the membrane in a process called endocytosis
Molecule at
Equilibrium Receptor Site
FIGURE 1–9 Diffusion: Random movement of molecules allows Outside
equalization of concentrations across a membrane.
the diffusion occurs. Some large molecules such as

proteins are unable to diffuse through the membrane
and must be transported in other ways.
As previously learned, the property of allowing Cell
Membrane
only certain molecules to diffuse through the mem-
brane is called semipermeability. This characteristic sets
the stage for a special type of diffusion, called osmosis.
A solvent (in the following case, water) moves across
the membrane to equalize the concentration; however,

the molecules dissolved in the water (called solutes)
cannot pass through the membrane (Figure 1–10). This Energy
process can be observed in red blood cells when they Inside
are placed in a concentrated solution. The water from Transport Channel
the cell moves outward into the solution. Microscopi- FIGURE 1–11 Active transport: Energy is used to actively pump molecules
cally, the red blood cells can be seen to shrink. into a region of higher concentration.
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*** START OF THE PROJECT GUTENBERG EBOOK CHARLES

DICKENS AND OTHER VICTORIANS ***
By Sir Arthur Quiller-Couch
On the Art of Writing

On the Art of Reading
Studies in Literature
(first series)
Studies in Literature
(second series)
Adventures in Criticism
Charles Dickens and
Other Victorians
Charles Dickens
And Other Victorians
Charles Dickens
And Other Victorians
By
Sir Arthur Quiller-Couch, M.A.
Fellow of Jesus College
King Edward VII Professor of English Literature
in the University of Cambridge
G.P. Putnam’s Sons

New York & London
The Knickerbocker Press
1925
Copyright, 1925
by
Sir Arthur Quiller-Couch
The
Knickerbocke
r
Press
New York
Made in the United States of America

PREFACE
ALL save one of the papers here collected were written as lectures
and read from a desk at Cambridge; the exception being that upon
Trollope, contributed to The Nation and the Athenaeum and
pleasantly provoked by a recent edition of the “Barsetshire” novels.
To these it almost wholly confines itself. But a full estimate of
Trollope as one of our greatest English novelists—and perhaps the
raciest of them all—is long overdue, awaiting a complete edition of
him. His bulk is a part of his quality: it can no more be separated
from the man than can Falstaff’s belly from Falstaff. He will certainly
come to his own some day, but this implies his coming with all his
merits and all his defects: and this again cannot happen until some
publisher shows enterprise. The expensive and artificial vogue of the
three-volume-novel did wonders for Trollope in one generation, to kill
him for another: since no critic can talk usefully about books to many
of which his hearers have no access. But we shall see Trollope
reanimated.
The papers on Dickens and Thackeray attempt judgment on
them as full novelists. Those on Disraeli and Mrs. Gaskell merely
take a theme, and try to show how one theme, taking possession,
will work upon two very different minds. Much more could have been
said generally upon both authors, and generically upon the “idea” of
a novel.
As usual, with a few corrections, I leave these lectures as they
were written and given, at intervals and for their purpose. They
abound therefore with repetitions and reminders which the reader
must try to forgive.
ARTHUR QUILLER-COUCH.
January 5, 1925.
CONTENTS
PAGE
Preface v
Dickens
I 3
II 24
III 42
IV 62
V 81
Thackeray
I 100
II 119
III 137
The Victorian Background 158
Disraeli 180
Mrs. Gaskell 199
Anthony Trollope
The Barsetshire Novels 219
Index 235
Charles Dickens
and Other Victorians
DICKENS (I)
I
IF anything on this planet be great, great things have happened in
Westminster Hall: which is open for anyone, turning aside from
London’s traffic, to wander in and admire. Some property in the oak
of its roof forbids the spider to spin there, and now that architects
have defeated the worm in beam and rafter it stands gaunt and clean
as when William Rufus built it: and I dare to say that no four walls
and a roof have ever enclosed such a succession of historical
memories as do these, as no pavement—not even that lost one of
the Roman Forum—has been comparably trodden by the feet of
grave men moving towards grave decisions, grand events.
The somewhat cold interior lays its chill on the imagination. A
romantic mind can, like the spider, spin its cobwebs far more easily
in the neighbouring Abbey, over the actual dust to which great men
come—
Here the bones of birth have cried—

“Though gods they were, as men they died.”
Here are sands, ignoble things
Dropt from the ruin’d sides of kings.
But in the Abbey is finis rerum, and our contemplation there the
common contemplation of mortality which, smoothing out place
along with titles, degrees and even deeds, levels the pyramids with
the low mounds of a country churchyard and writes the same moral
over Socrates as over our Unknown Soldier—Vale, vale, nos te in
ordine quo natura permittet sequamur. In Westminster Hall (I am
stressing this with a purpose) we walk heirs of events in actual play,
shaping our destiny as citizens of no mean country: in this covered
rood of ground have been compacted from time to time in set conflict
the high passions by which men are exalted to make history. Here a
king has been brought to trial, heard and condemned to die; under
these rafters have pleaded in turn Bacon, Algernon Sidney, Burke,
Sheridan. Here the destinies of India were, after conflict, decided for
two centuries. Through that great door broke the shout, taken up,
reverberated by gun after gun down the river, announcing the
acquittal of the Seven Bishops.
II
So, if this tragic comedy we call life be worth anything more than
a bitter smile: if patriotism mean anything to you, and strong opposite
wills out of whose conflict come great issues in victory or defeat, the
arrest, the temporary emptiness of Westminster Hall—a sense of
what it has seen and yet in process of time may see—will lay a
deeper solemnity on you than all the honoured dust in the Abbey.
But, as men’s minds are freakish, let me tell you of a solitary
figure I see in Westminster Hall more vividly even than the ghosts of
Charles I and Warren Hastings bayed around by their accusers: the
face and figure of a youth, not yet twenty-two, who has just bought a
copy of the Magazine containing his first appearance in print as an
author. “I walked down to Westminster Hall,” he has recorded, “and
turned into it for half an hour, because my eyes were so dimmed with
joy and pride that they could not bear the street and were not fit to be
seen there.”
Now the paper which opened the fount of these boyish tears
(here, if you will, is bathos) was entitled A Dinner at Poplar Walk.
You may find it to-day under another title, “Mr. Minns and his Cousin”
among Sketches by Boz: reading it, you may pronounce it no great
shakes; and anyhow you may ask why anyone’s imagination should
select this slight figure, to single it out among the crowd of ghosts.
Well, to this I might make simple and sufficient answer, saying that
the figure of unbefriended youth, with its promise, a new-comer
alone in the market-place, has ever been one of the most poignant in
life, and, because in life, therefore in literature. Dickens himself, who
had been this figure and remembered all too well the emotion that
choked its heart, has left us a wonderful portrait-gallery of these lads.
But indeed our literature—every literature, all legend, for that matter
—teems with them: with these youngest brothers of the fairy-tales,
these Oedipus’s, Jasons, these Dick Whittingtons, Sindbads,
Aladdins, Japhets in search of their Fathers; this Shakespeare
holding horses for a groat, that David comely from the sheepfold with
the basket of loaves and cheeses. You remember De Quincey and
the stony waste of Oxford Street? or the forlorn and invalid boy in
Charles Lamb’s paper on The Old Margate Hoy who “when we
asked him whether he had any friends where he was going,” replied,
“he had no friends.” Solitariness is ever the appeal of such a figure;
an unbefriendedness that “makes friends,” searching straight to our
common charity: this and the attraction of youth, knocking—so to
speak—on the house-door of our own lost or locked-away ambitions.
“Is there anybody there?” says this Traveller, and he, unlike the older
one (who is oneself), gets an answer. The mid-Victorian Dr. Smiles
saw him as an embryonic Lord Mayor dazed amid the traffic on
London Bridge but clutching at his one half-crown for fear of pick-
pockets. I myself met him once in a crowded third-class railway
carriage. He was fifteen and bound for the sea: and when we came
in sight of it he pushed past our knees to the carriage window and
broke into a high tuneless chant, all oblivious of us. Challenge was in
it and a sob of desire at sight of his predestined mistress and
adversary. For the sea is great, but the heart in any given boy may
be greater: and
these things are life

And life, some think, is worthy of the Muse.
III
But I am a Professor, and ought to have begun by assuring you
that this figure in Westminster Hall has a real historical interest in
connexion with your studies “on the subject of English Literature.”
Well, then, it has. The date of the apparition is New Year’s Day,
1834, and by New Year’s Day, 1838, Charles Dickens was not only
the most popular of living authors, but in a fair way to become that
which he remained until the end in 1870—a great National
Institution.
I use no exaggerated term. Our fathers of the nineteenth century
had a way (and perhaps not altogether a bad way) of considering
their great writers as national institutions; Carlyle was one, Ruskin
another. It was a part of their stout individualism, nowadays derided.
And it was, if you will consider, in the depths of its soul [say, if you
will, its Manchester Soul] a high-polite retort upon such a sworn
enemy as Ruskin. “Curse us, Sir: but we and no Government make
you a demigod.” You will never understand your fathers, Gentlemen,
until you understand their proud distrust of Government save by
consent. Take a favourite term of theirs—say “The Liberty of the
Press.” By that they meant liberty from interference by Government.
We, using that term to-day, should mean nothing of the sort. We
should mean “liberty from control by capitalists.”
I interrogate my youthful memories and am confident that, in a
modest country household these men—Carlyle, Ruskin—were, with
decent reverence, though critically, read for prophets. Tennyson, too,
and Browning had their sacred niches; and Darwin and Huxley, and
Buckle, who perished young attempting a History of Civilisation in
Europe: John Stuart Mill, also, and Kingsley, Maurice, George Eliot,
and Thackeray. These names leap to memory as names of
household gods. A few weeks ago, rummaging over some family
papers I came upon the following entry:
1848, June 20. I received a visit from Mr. Alfred Tennyson,

the Poet. He came into Cornwall along the North Coast, and
from about Camelford crossed over to Fowey, where I called on
him on the 19th. He came to Polperre in a boat, with Mr. Peach
and others; and after viewing our scenery in all directions and
taking tea at our house, they all rowed back to Fowey late in the
evening. I find him well-informed and communicative. I believe a
good Greek scholar with some knowledge of Hebrew. His
personal appearance is not prepossessing; having a slouch in
his gait and rather slovenly in his dress tho’ his clothes were new
and good. He confesses to this. He admired the wildness of our
scenery, deprecated the breaking in of improvements, as they
are termed. He enquired after traditions, especially of the great
Arthur: his object in visiting the County being to collect materials
for a poem on that Chief. But he almost doubted his existence.
He show’d me a MS. sketch of a history of the Hero: but it was
prolix and modern.
You see, hinted in this extract from a journal, how our ancestors, in
1848 and the years roundabout, and in remote parts of England,
welcomed these great men as gods: albeit critically, being
themselves stout fellows. But above all these, from the publication of
Pickwick—or, to be precise, of its fifth number, in which (as Beatrice
would say) “there was a star danced” and under it Sam Weller was
born—down to June 14, 1870, and the funeral in Westminster
Abbey, Dickens stood exalted, in a rank apart. Nay, when he had
been laid in the grave upon which, left and right, face the
monuments of Chaucer, Shakespeare and Dryden, and for days
after the grave was closed, the stream of unbidden mourners went
by. “All day long,” wrote Dean Stanley on the 17th, “there was a
constant pressure on the spot, and many flowers were strewn on it
by unknown hands, many tears shed from unknown eyes.”
Without commenting on it for the moment, I want you to realise
this exaltation of Dickens in the popular mind, his countrymen’s and
countrywomen’s intimate, passionate pride in him; in the first place
because it is an historical fact, and a fact (I think) singular in our
literary history; but also because, as a phenomenon itself unique—
unique, at any rate, in its magnitude—it reacted singularly upon the
man and his work, and you must allow for this if you would
thoroughly understand either.
IV
To begin with, you must get it out of your minds that it resembled
any popularity known to us, in our day: the deserved popularity of Mr.
Kipling, for example. You must also (of this generation I may be
asking a hard thing, but it is necessary) get it out of your minds that
Dickens was, in any sense at all, a cheap artist playing to the gallery.
He was a writer of imperfect, or hazardous, literary education: but he
was also a man of iron will and an artist of the fiercest literary
conscience. Let me enforce this by quoting two critics whom you will
respect. “The faults of Dickens,” says William Ernest Henley,
were many and grave. He wrote some nonsense; he sinned

repeatedly against taste; he could be both noisy and vulgar; he
was apt to be a caricaturist where he should have been a
painter; he was often mawkish and often extravagant; and he
was sometimes more inept than a great writer has ever been.
But his work, whether good or bad, has in full measure the
quality of sincerity. He meant what he did; and he meant it with
his whole heart. He looked upon himself as representative and
national—as indeed he was; he regarded his work as a universal
possession; and he determined to do nothing that for lack of
pains should prove unworthy of his function. If he sinned, it was
unadvisedly and unconsciously; if he failed it was because he
knew no better. You feel that as you read....
He had enchanted the public without an effort: he was the
best beloved of modern writers almost from the outset of his
career. But he had in him at least as much of the French artist as
of the middle-class Englishman; and if all his life he never
ceased from self-education, but went unswervingly in pursuit of
culture, it was out of love for his art and because his conscience
as an artist would not let him do otherwise.
Now let me add this testimony from Mr. G. K. Chesterton:
Dickens stands first as a defiant monument of what happens

when a great literary genius has a literary taste akin to that of the
community. For the kinship was deep and spiritual. Dickens was
not like our ordinary demagogues and journalists. Dickens did
not write what the people wanted. Dickens wanted what the
people wanted.... Dickens never talked down to the people. He
talked up to the people. He approached the people like a deity
and poured out his riches and his blood. He had not merely
produced something they could understand, but he took it
seriously, and toiled and agonised to produce it. They were not
only enjoying one of the best writers, they were enjoying the best
he could do. His raging and sleepless nights, his wild walks in
the darkness, his note-books crowded, his nerves in rags, all this
extraordinary output was but a fit sacrifice to the ordinary man.
“The good, the gentle, high-gifted, ever-friendly, noble Dickens,”

wrote Carlyle of him, on hearing the news of his death,—“every inch
of him an honest man.” “What a face it is to meet,” had said Leigh
Hunt, years before; and Mrs. Carlyle, “It was as if made of steel.”
V
I shall endeavour to appraise with you, by and by, the true worth
of this amazing popularity. For the moment I merely ask you to
consider the fact and the further fact that Dickens took it with the
seriousness it deserved and endeavoured more and more to make
himself adequate to it. He had—as how could he help having?—an
enormous consciousness of the power he wielded: a consciousness
which in action too often displayed itself as an irritable
conscientiousness. For instance, Pickwick is a landmark in our
literature: its originality can no more be disputed than the originality
(say) of the Divina Commedia. “I thought of Pickwick”—is his
classical phrase. He thought of Pickwick—and Pickwick was. But just
because the ill-fated illustrator, Seymour—who shot himself before
the great novel had found its stride—was acclaimed by some as its
inventor, Dickens must needs charge into the lists with the hottest,
angriest, most superfluous, denials. Even so, later on, when he finds
it intolerable to go on living with his wife, the world is, somehow or
other, made acquainted with this distressing domestic affair as
though by a papal encyclical. Or, even so, when he chooses (in
Bleak House) to destroy an alcoholised old man by “spontaneous
combustion”—quite unnecessarily—a solemn preface has to be
written to explain that such an end is scientifically possible. This
same conscientiousness made him (and here our young novelist of
to-day will start to blaspheme) extremely scrupulous about
scandalising his public—I use the term in its literal sense of laying a
stumbling-block, a cause of offence. For example, while engaged
upon Dombey and Son, he has an idea (and a very good idea too,
though he abandoned it) that instead of keeping young Walter the
unspoilt boyish lover that he is, he will portray the lad as gradually
yielding to moral declension, through hope deferred—a theme which,
as you will remember, he afterwards handled in Bleak House: and he
seriously writes thus about it to his friend Forster:
About the boy, who appears in the last chapter of the first
number—I think it would be a good thing to disappoint all the
expectations that chapter seems to raise of his happy
connection with the story and the heroine, and to show him
gradually and naturally trailing away, from that love of adventure
and boyish light-heartedness, into negligence, idleness,
dissipation, dishonesty and ruin. To show, in short, that common,
every day, miserable declension of which we know so much in
our ordinary life: to exhibit something of the philosophy of it, in
great temptations and an easy nature; and to show how the
good turns into the bad, by degrees. If I kept some notion of
Florence always at the bottom of it, I think it might be made very
powerful and very useful. What do you think? Do you think it
may be done without making people angry?
George Gissing—in a critical study of Dickens which cries out for

reprinting—imagines a young writer of the ’nineties (as we may
imagine a young writer of to-day) coming on that and crying out upon
it.
What! a great writer, with a great idea, to stay his hand until
he has made grave enquiry whether Messrs. Mudie’s
subscribers will approve it or not! The mere suggestion is
infuriating.... Look at Flaubert, for example. Can you imagine him
in such a sorry plight? Why, nothing would have pleased him
better than to know he was outraging public sentiment! In fact, it
is only when one does so that one’s work has a chance of being
good.
All which, adds Gissing, may be true enough in relation to the

speaker. As regards Dickens, it is irrelevant. And Gissing speaks the
simple truth; “that he owed it to his hundreds of thousands of readers
to teach them a new habit of judgment Dickens did not see or begin
to see.” But that it lay upon him to deal with his public scrupulously
he felt in the very marrow of his bones. Let me give you two
instances:
When editing Household Words he receives from a raw
contributor a MS. impossible as sent, in which he detects merit. “I
have had a story,” he writes to Forster, “to hack and hew into some
form this morning, which has taken me four hours of close attention.”
“Four hours of Dickens’ time,” comments Gissing, “in the year 1856,
devoted to such a matter as this!—where any ordinary editor, or
rather his assistant, would have contented himself with a few
blottings and insertions, sure that ‘the great big stupid heart of the
public,’ as Thackeray called it, would be no better pleased, toil how
one might.”

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Unit I ■ Comparative Anatomy and Physiology

Unit III ■ Diseases

Unit V ■ Professionalism and Careers

CHAPTER 21 Careers and Decision Making

Chapter 4 The Circulatory System 55

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draped around my neck! It seems like yesterday, even

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FIGURE 3–14 Radiograph of the lumbar spine of a dog. A portion of the

FIGURE 1–5 Chemical structure of a nucleotide.

2. Cytosine 2. Cytosine used in protein synthesis. A group of three nitroge-

10 Unit 1 Comparative Anatomy and Physiology

Cytoplasm Extracellular enzyme begins at a specific series of bases (thymine,

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BONE STRUCTURE Distal Epiphysis Each chapter is further enhanced by the

spongy arrangement keeps the weight of the bones

88 Unit 1 Comparative Anatomy and Physiology

Courtesy of Dr. David Sweet.

EDUCATIONAL REQUIREMENTS FOR

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CELL MAKEUP of these molecules. They possess the basic structure of

RLong Hydrocarbon Chain Fatty Acid

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FIGURE 1–4 Chemical structure of selected amino acids.

H OH OH H Protein can bind with other molecules to aid in their

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FIGURE 1–5 Chemical structure of a nucleotide.

2. Cytosine 2. Cytosine used in protein synthesis. A group of three nitroge-

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the inside of the cell contained. The cell membrane is

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Smooth Endoplasmic Reticulum

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Objective eliminated by the ECF. Without elimination, the waste

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RLong Hydrocarbon Chain Fatty Acid