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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
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.
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
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.
Color codes for the bases

A G
C T U
Nontranscribed Strand
5'
5'
3' Transcription
DNA
3'
3'
mRNA Transcribed Strand

(Complementary
Copy of
Transcribed mRNA
DNA Strand)
5'
Codon 1 Codon 2 C
Codon 3 C
Codon 4 C
Codon 5 C
Codon 6

Translation
Amino Amino Amino Amino Amino Amino O

H2N Acid Acid Acid Acid Acid Acid
C
1 2 3 4 5 6 OH
FIGURE 1–15 Translation: The mRNA created in transcription is used to code the amino acid sequence in protein formation.
Translation, which occurs in the ribosomes, is the as the epithelia lining the intestinal tract, divide fre-
process in which the code of bases in the mRNA is con- quently to maintain the integrity of the layer. Other
verted to a series of amino acids. Each series of three cells, such as skeletal muscle, do not divide in an adult.
bases in the mRNA is a codon (Figure 1–15). The codon When these normal controls break down, the cells
provides the signal for a specific amino acid. The mol- can begin to undergo frequent mitosis. Uncontrolled
ecule of mRNA is bound by ribosomes. A molecule of mitosis results in cancer. New cells are produced more
transfer RNA (tRNA) that contains the three complemen- quickly than needed, resulting in an accumulation or
tary bases (anticodon) attaches to the mRNA. Each mole- mass of cells in a region. This mass of rapidly dividing
cule of tRNA carries the amino acid specific to the codon. cells is called a tumor.
Enzymes on the ribosome allow release of the amino In a nondividing cell, the genetic material is called
acid from the tRNA. A peptide bond is created between chromatin. In this form, the chromatin is loosely ar-
adjacent amino acids. This process is repeated along the ranged in the nucleus. The individual chromosomes
length of the mRNA molecule, creating a polypeptide. cannot be seen with a light microscope. These cells are
The proteins created may be used within the cytoplasm described as being in the interphase. In this stage, the
or processed further within the endoplasmic reticulum. cell is in the process of doubling its DNA. The steps of
division are broken down into four phases (Table 1–4).
The phases are identified to help understand the
MITOSIS AND CANCER
Objective
Table 1–4 Stages of Mitosis
■ Discuss Mitosis and its Clinical Significance in Diseases
Such as Cancer 1. Interphase
Cells must reproduce. In mitosis, the cells divide, pro- 2. Prophase

ducing two identical cells. Mitosis is necessary for the 3. Metaphase

growth and maintenance of the animal. Cell division
is controlled by a number of factors present within the 4. Anaphase
cell and the extracellular fluid. The rate of cell division
5. Telophase
is adapted to the needs of the animal. Some cells, such

Interphase Early Prophase Late Prophase Metaphase Anaphase Telophase Interphase
FIGURE 1–16 Mitosis. Interphase: Cell in its normal state, as the chromosomes begin to replicate. Prophase: The chromatin thickens and becomes visible,
taking on an X shape. The nucleoli and nuclear membrane begin to disappear. Metaphase: The spindle forms between two centrioles. The chromosomes
align on the spindle. Anaphase: Chromosomes split at the centromeres, with each half moving to opposite ends. Telophase: The nucleus reforms and a
groove divides the two new cells.
process. However, actual cell division is a continuous the cells. These factors can either stimulate cell divi-
process, as seen in Figure 1–16. sion or inhibit it. Other factors found in the ECF help
Prophase begins as the chromatin thickens into vis- to control cell death in a process called apoptosis. A
ible chromosomes. This is the first time that the individ- classic example of this balance occurs in the cells that
ual chromosomes can be seen with a light microscope. line the gastrointestinal tract. Cells at the base of the
Along with this process, the nucleoli and nuclear mem- lining divide frequently at a rate that balances with the
brane begin to disappear. At this point, the chromosomes cells undergoing apoptosis and death at the surface of
show the doubling that occurred during interphase. The the lining. The programmed cell death is designed to
chromosomes have an X shape. The two identical halves, occur whenever there is significant cell damage such
or chromatids, are joined at a small point called the cen- as mutations.
tromere. Two small organelles, the centrioles, separate Different cells divide at varying rates. Cells found
and move to opposite ends of the cell. in certain areas such as the bone marrow and linings of
In metaphase, a spindle is formed between the the gastrointestinal tract have stem cells that actively
two centrioles. This is a collection of microtubules that divide on a regular basis. Other cells found in organs
stretch between the two centrioles. The chromosomes such as the liver, kidney, and pancreas do not routinely
move to the center of the cell and align themselves on divide. However, following injury or disease, these
the spindle. cells can become activated into frequent mitosis to al-
As anaphase begins, the chromosomes split at the low repair of the organ. A few specialized cells such as
centromere. At this point, the chromosomes are still nerve and muscle cells have very limited or no ability
on the spindle. Each chromatid begins to move out- to divide.
ward. The centromere portion moves first, giving the
chromosome a V shape. The chromosomes move to
opposite ends of the cell. MAMMALIAN REPRODUCTION
Telophase is basically the reverse of prophase.
The chromosomes become loosely organized into chro- Objective
matin. The nuclear membrane and nucleoli return. A ■ Detail Meiosis in Mammalian Reproduction
groove then forms down the center of the cell. This
groove deepens until two identical cells are produced Mammals rely on sexual reproduction for species sur-
in a process called cytokinesis. vival. In sexual reproduction, a sperm cell and egg cell
Mitosis is essential in maintaining the population join to form the new embryo. In this process, half of the
size of cells in the body. The number of cells is estab- genetic material is provided by each of the cells. Mei-
lished on the basis of the frequency of mitosis, the dif- osis is the division in which the resulting cells contain
ferentiation of cells, and cell death. An increase in cell only half of the genetic material.
number can occur if the rate of cell division increases or There are two cell divisions during meiosis, with
the rate of death decreases. In certain instances, a com- only one doubling of the chromatin. The final result is
bination of these two changes has a cumulative effect. the formation of four cells, each with half the number
The rate of cell division is controlled by soluble of chromosomes. Just as in mitosis, meiosis divides into
factors found in the extracellular fluid surrounding phases (Figure 1–17).
The second division of meiosis is similar to mitosis.

The net effect of meiosis is the formation of four cells,
each with half of the original number of chromosomes.
Prophase I: Mammals should have an even number
of chromosomes. The chromosomes come in pairs, and
each member of a pair is called a homologue. Before
prophase I, the homologue of each pair replicates and
is formed by two strands (or chromatids). The chro-
Early Prophase I Middle Prophase I Late Prophase I matids are joined by a centromere. Prophase I is very
(Leptotene) (Zygotene) (Diplotene)
complex. The basic process allows the homologues to
pair up near the center of the cell. In this arrangement,
the homologues are joined at several points. At these
points, an exchange of DNA fragments occurs.
Metaphase I: This step is very similar to mitosis.
The nuclear membrane and nucleoli begin to disap-
pear. The paired chromosomes move into alignment
on the spindle. The important distinction in meiosis is
that the homologues align themselves where they will
Metaphase I Anaphase I be divided into opposite cells.
Anaphase I and Telophase I: In anaphase, the chro-
mosomes begin to move to opposite ends of the cell.
In this step, the centromere does not split. Rather, the
pairs of chromosomes are divided.
The length of interphase between the two divisions
Telophase I is variable, and may even be zero (i.e., this phase may
not occur at all). The two cells produced enter into the
second division. The second division of meiosis is basi-
cally the same as mitosis. In this division, the chromo-
somes align on the spindle, separate at the centromere,
Prophase II and send one strand to each new cell. The stages are
named just as they are in mitosis. The final result of
meiosis is the formation of four cells, each with half the
number of chromosomes of the original cell.
Meiosis allows genetic material to be provided
from each parent. The exchange of genetic material be-
Metaphase II
tween homologues in prophase I produces variability
in each cell. Offspring acquire traits from each parent.
With the variation, no two sperm or egg cells will pro-
vide the same genetic material.
Twinning can result in two animals having the
same genetic makeup. Identical twins occur when an
embryo splits. Each half then develops into a new em-
bryo. The resulting offspring begin life with identical
Anaphase II
chromosomes. Even identical twins, however, do not
appear completely identical. There is variation in the
way the genes are expressed.
CLINICAL PRACTICE
Telophase II
Objective
FIGURE 1–17 Meiosis: A two-division process. Prophase I: Homologous ■ Connect Cellular Parts and Function to Clinical Veterinary
chromosomes align in the center of the cell. The homologues exchange Practice
segments of genetic material. Metaphase I: Similar to the step in mitosis,
except that the homologues will separate into opposite cells. Anaphase I: In clinical practice, the appearance of cells is often
The chromosomes move to opposite ends of the cell. Telophase I: The cell is evaluated. A biopsy takes tissues or cells from an
divided into two daughter cells, each with half of the chromosome numbers
animal for microscopic review. This procedure allows
of the original cell.
diagnosis. In tumors, the cells divide without normal
Photograph courtesy Dr. Arthur Hattel, Pennsylvania State University.

control. This leads to a mass in the tissue or in an
organ. Tumors are divided into two major groups,
benign and malignant. Benign tumors are localized
to one area, have a well-defined margin, and do not
spread to other parts of the body. Malignant tumors
are more likely to invade surrounding tissues and
spread to other parts of the body. For example, cells
from a tumor may break away and move into the
bloodstream or lymph vessels and then settle into a
new location. The spread of a tumor from its primary
location is called metastasis. The word tumor, or
neoplasm, can be used to describe either a benign or
malignant mass. The term cancer is typically used in FIGURE 1–18 Photomicrograph (high power) of cancer (squamous cell
reference to a malignant tumor. With biopsy, cells are carcinoma) in the skin of a horse. There is variation in the shape and size
evaluated to determine the type of tumor that is pres- of the nucleus and cells. Many mitotic figures can also be seen.
ent. Penny’s tumor was submitted for biopsy. Fortu-
nately, the pathologists (who interpret and diagnose
changes in cells and tissues) found the tumor to be bloodstream. The walls of the smallest blood vessels
benign and did not find any tumor cells at the margins (capillaries) act as a semipermeable membrane. They
of the sample. This was great news for Penny, and we do not allow the protein within the blood to freely
were optimistic that her tumor would not cause any move to the extracellular fluid. Water, on the other
more problems. hand, is able to move freely in either direction.
Tumors develop because the cells are growing Osmosis occurs across the walls of the capillar-
rapidly and dividing without normal control. Changes ies. With normal blood protein levels, a balance is
that occur in DNA allow a tumor cell to divide, inde- reached between the extracellular fluid and the liq-
pendent of the inhibitory and stimulating control of uid portion of blood. As protein levels decline, there
normal growth factors. The tumor begins from a single is less osmotic pressure to keep water within the
defective cell. Further division can allow even further blood vessels. As protein continues to decline, water
mutations that influence some other aspect of the cell accumulates within the tissues. In sheep with severe
cycle. For example, tumor cells lose susceptibility to parasitism, the most common site for this fluid accu-
factors that control apoptosis. Normal cells require at- mulation to occur is below the jaw.
tachment to other cells and surrounding matrix. Cancer Sadie, a nine-year-old Papillon, presented with a
cells develop the ability to replicate without this at- distended abdomen. Previous workup by a specialist
tachment. This feature is critical to allow for metastasis had discovered a kidney problem that allowed protein
with cells spread from the original tumor being able to to leak into the urine. Table 1–5 shows that Sadie’s
replicate at a new location. total protein had dropped below normal. This was
Many features of cancer can be predicted with primarily due to the decrease in albumin. Sadie’s liver
this information. In cancerous cells, there is often a was unable to make albumin quickly enough to replace
large nucleus with many nucleoli. The chromatin is what was lost. As a result, Sadie developed ascites,
often clumped and visible. There are many more cells an accumulation of free fluid within the abdomen
involved in the process of cell division than in normal (Figure 1–19). On presentation, Sadie weighed four
tissue. Furthermore, many of the dividing cells have pounds, and more than one half pound of fluid was
an abnormal spindle. Only with an understanding of
the normal cell cycle can we interpret these abnormal
findings (Figure 1–18).
Bottle jaw is a term used to describe an accumula- Table 1–5 Blood Results for Sadie, a 9-Year-Old Spayed
tion of fluid within the tissues under the jaw. Edema Female Papillon
describes excess fluid within a tissue. In bottle jaw,
it is described as pitting edema, because finger pres- Sadie Reference Range
sure into the tissue creates a “pit” that only slowly Total protein 3.4 g/dl 5.2–8.2
resolves. Bottle jaw occurs in severely parasitized
sheep. Parasites within the intestinal tract consume Albumin 1.0 g/dl 2.2–3.9
such a large amount of protein that the animal is
Globulin 2.4 g/dl 2.5–4.5
unable to maintain normal protein levels within the

FIGURE 1–19 Photograph of Sadie showing her abdomen distended with ascites.
drawn from the abdomen. This lowered the pressure In peritoneal dialysis, an electrolyte solution is
on her internal organs and provided immediate relief. infused into the abdomen through a catheter. Urea
The low protein level meant the ascites would rapidly is able to diffuse across the membrane that lines the
recur. To slow this progression, Sadie was given a solu- abdomen. Because there is no urea within the infused
tion of hetastarch into her bloodstream. Hetastarch is electrolyte solution, the urea diffuses from the higher
a very large molecule that increases osmotic pressure concentration within the bloodstream into the solution.
within the bloodstream, helping to slow the recurrence The solution is then removed from the body, effectively
of ascites. The hetastarch slowly leaves the blood and reducing the level of urea in the bloodstream. Dialysis
thus provides only a temporary benefit. The goal in this must be repeated to remove the urea that is subse-
case is to improve the osmotic pressure until a more quently produced. Dialysis is used to maintain the
permanent cure can be found. animal in the hope that the kidneys will recover from
Diffusion and osmosis are used therapeutically in the toxic effect.
the process of dialysis. Dialysis is a treatment option in Understanding cellular function is essential for
kidney failure. As will be discussed later in the text, the sound veterinary practice. Although there is great
kidneys serve to maintain fluid and electrolyte balance similarity among all mammals, there are species dif-
within the body. In addition, the kidneys function to ferences. This chapter has discussed enzymes and their
remove many forms of soluble waste products. Ethyl- importance in cellular function. Cells contain enzymes
ene glycol found in certain antifreeze products is ex- that break down medication (that is why medicines
tremely toxic to the kidneys. Unfortunately, pets often must be given more than once). Furthermore, differ-
find the antifreeze palatable (i.e., tasty) and will ingest ences between animals can lead to reactions to med-
enough to severely damage the kidneys. ications. Cats possess much less of certain enzymes
When the kidneys do not function adequately, that break down many medications. Acetaminophen
toxins increase in the bloodstream, and many electro- (Tylenol), a common over-the-counter pain reliever,
lyte levels become imbalanced. The complete process has been proved quite safe for use in humans. Because
of dialysis is quite complex and requires extensive of the differences in the enzymes between species, this
monitoring to be done correctly. Urea will be used as product is very dangerous for cats. As little as half a
an example to describe the function of diffusion in di- tablet designed for adult humans can make a cat sick.
alysis. Urea is a breakdown product in the metabolism Cellular detail seems far removed from a complete an-
of protein. The kidneys normally excrete the urea into imal. However, the details of cells guide treatment of
the urine. With kidney failure the urea level increases animals in clinical practice.
in the bloodstream and acts as a toxin.
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showed he was in full scent again. On, then, by the side of
the now roaring beck, for the snows were melting and the
stream was becoming strong and loud.
Suddenly Laddie stops—ears pricked—head in the air. What

can he hear beside the brawling of the stream? He breaks
out into an almost human cry, and rushes on in spite of
Alice's distressed calls and commands that he should not
leave her. She certainly hears another voice besides his, no
—yes, the joyful barking of two dogs—old Chance's full deep
voice and Laddie's sharp treble. Look, Laddie is coming back
to meet her, trembling all over with intense excitement.
A few more painful efforts, and the young girl sinks

perfectly exhausted beside her senseless brother. But she
has no leisure for fainting, no time for feeling: she has stern
work in hand. Is he dead or alive? That is the only thought.
Chance is still laboriously licking the face and the hands of
his master. Alice lays her hand on his heart, and still—yes,
still—she thinks there is a feeble beating and a little sense
of vital warmth. That is enough for her.
"Off, Laddie," said she earnestly to the dog, waving her

hand in the direction of Scarf Beck Farm; "off, and bring
them here;" pointing to his master.
The dog understood and was off. What cannot a shepherd's

dog understand, when, at his master's bidding, he will hunt
up, collect and bring home, a whole flock of lost and
scattered sheep, without missing a single one?
The party returning from the old Man were coming along
the pathway which led to Scarf Beck Farm, when Laddie saw
them at a distance—dark figures on the white snow—and
rushed wildly up to them, entreating them by every
argument, short of speech to follow him.
"What's to do with the young dog?" said old Geordie in an
excited voice. "He has found out something for sure. Look
at his ways."
They hurried along after him as though he were a human

guide, and before long they came, sure enough, upon that
sad group in the cave-like hollow which was scooped out of
the side of the great snow-drift.
"She has found him," exclaimed Mark; "thank God for that."
Alice was sitting on the snow holding her brother in her

arms and clasping him tight to her warm heart. She had
wound her plaid around him besides; and the old dog was
still leaning against him, licking his blue hands and wrists.
Alice's face was buried in her brother's hair; but at the
sound of Mark's exclamation, she looked up hastily, and
saying:
"Oh, Mark," burst into an agony of tears, the first she had
shed. "Oh, do something to save him. I know he is not
dead. I am sure I can feel his breath."
They knelt around the seemingly lifeless figure, and Old

Geordie took out a little flask of brandy, which he had put
into his pocket on starting from home, saying to Old Ann,
"I'll take this, however; no one knows but there may be
sore need of it."
He now managed to get a little of the restorative within the

livid lips; and after what seemed a long, long time, there
was an evident attempt to swallow. It was more like a
spasm of pain than a natural effort; but it showed that the
living spark had not quite gone out. They all now set to
work, rubbing the hands, chafing the feet, bathing the
temples with brandy, and again contriving to get a little of
the liquid swallowed.
"We must carry him home," said Geordie; "we shall never
get him round here, in the cold wind. Sister will wrap him
up right well, and we men will carry him."
And so the melancholy-looking procession moved away

from the shining snow-cavern, the dogs trailing along
behind in a state of deep depression, because they did not
like the look of the long muffled figure helplessly borne
along by the four silent men. Alice tried to get on in
advance, to give warning to her mother, and to prepare the
hot bed, hot blankets, and hot drinks, on which she rested
her hopes. But with all her efforts, her spent strength could
make small progress.
"Don't distress yourself so, Alice," said Mark Wilson, who

was anxiously watching her spasmodic efforts; "we shall be
there as soon as you."
"I must do something to help," she replied, with quivering

lip.
"Thou go and help the lile maiden," said Geordie

compassionately; "she'll drop soon, and we shall e'en have
them both to carry home to mother. One is bad enough."
But for this encouragement from the old servant, Mark's shy
reserve would have withheld him from aiding the poor girl.
She had dropped behind by this time, in utter prostration of
her over-strained powers. It was well that this strong and
willing arm came to the rescue. How confidingly she leaned
on him! How she trusted her weakness to his strength! She
felt as if she could have done anything with that arm to aid,
that voice to encourage, that look of understanding
sympathy, tender and true, to comfort her. Will they ever
forget that mournful walk? Never—as long as they live.
Young Mat had by this time returned from his bold but
fruitless expedition to the distant hamlet, and had caught
sight of the dark group of figures winding along over the
white fields. He thought that all was indeed over. It looked
like nothing but one of those mournful processions which he
was accustomed to see creeping along the side of the hills,
up out of one valley, and down into another, on the old
paved pathways leading to the common centre in the
church-yard which are expressively called in the language of
the country, "corpse-roads."
"They are coming along, mother," said he, entering the

kitchen, and gently going up to his mother's chair; "they
have found Miles, I think, but I don't rightly know how."
The widow looked searchingly in the boy's face, and

trembled all over. "Nevertheless," said she, "I'll have
everything ready for life. In His hands are the issues. Warm
the lad's bed, Ann, and heat the blankets—ay, roast them
brown, if thou likest. Put the peppermint tea on the hob. Do
thou try warmth, and I'll try prayer;" and the aged women
betook themselves to their several offices.
The party soon entered in silence; for they dared not raise
the mother's hopes over so very doubtful a case of revival.
"Take him straight to his own warm bed," she said, "and
place me and my chair beside him. Who knows but the Lord
may hear the cry of the destitute and not despise their
prayer? It seems borne in upon my mind that it shall be
well with the lad, and that his spirit shall come to him
again."
Two long hours afterwards, Alice glided downstairs, and

going to Mark, who was leaning his head on both his hands,
with his elbows on the kitchen table, said with beaming
eyes and glowing face, "Oh, Master Wilson, the color is
come again into Miles' face, and he is quite warm, and
sleeping like a child."
He started up, took both her hands in his, and pronounced

a solemn thanksgiving.
"Alice," he added, "let us pray that our brother may in very

truth be alive from the dead. Surely that was repentance
last night. I thought I heard the sob of a broken and a
contrite spirit. But his going out in that strange way is what
puzzles me."
"Wait—wait, and see," said the loving sister; "I think it will
all be made clear. He was so tender to mother last night; so
very gentle and kind to me."
"That is no particular sign of good that I know of," said

Mark, smiling at her flushed and eager defence of her
brother; "how could he be anything else than kind, and
loving to thee?"
"Oh, Master Wilson, you don't understand me at all."
"Master? We can never be master and pupil in that old

distant way after having shared so much together yesterday
and to-day. I shall never be able to separate the thought of
thee from anything now, Alice."
But she was gone, fluttering away like a startled bird,

before he had finished the sentence which it had cost him
so much emotion to pronounce.
"She is gone," said he, despairingly. "I believe I don't

understand women's minds as well as school-boys. Now,
one would have thought that she would at least have waited
to hear what I should say next."
CHAPTER VI.
THE MOUNTAIN ECHOES.
"An idle word—a lowly prayer—

A gentle 'Bless thee!' fitly
spoken,
May live, and echo through the
air,
As if its life could ne'er be
broken."
It was days before Miles Lawson rallied. His powers were so

utterly prostrated, that dull heavy sleep seemed to be the
only resource of nature. They could scarcely rouse him,
even to take the needful nourishment; and at such times he
took no notice of any one, but drank his mother's
concoctions, or Alice's less distasteful preparations, in a
mere mechanical manner, without raising his dim eyes to
see what hand was ministering to him. All this while he
never spoke, or showed the least consciousness of what had
passed; and his watchful attendants wisely refrained from
forcing any exciting subject upon his over-strained nerves.
"He will mind all about it as soon as he has got strength to

look back over his shoulder," said the old lady; and so she
folded her hands patiently, and waited another day, and yet
another, for the signs of reviving consciousness.
In the meantime, all were not idle at The Yews. Mr. Knibb
had been extremely excited by the news of the discovery of
the secret still on the mountainside, and had made more
than one effort to set off for the lofty spot on the same
evening. But Madam was wholly of another mind, and
showed so much partiality for the warm stable, and such an
objection to be mounted on four hard and slippery balls of
snow, that the impatient old gentleman had to yield the
point. Madam was quite in the right; nothing could then
have been attempted with success, or even with safety.
Early on the following morning a strong party was

mustered, headed by Mr. Knibb and the schoolmaster, and
composed of Geordie Garthwaite, young Mat Lawson, the
farm lads, and Bella Hartley's brothers. There were two
carts in attendance, and these were picketed at a chosen
spot, as far up the mountainside as wheels could reach. The
old exciseman, leaving Madam with the rear guard, carefully
covered with his own drab top-coat, labored up the steep
path with marvellous spirit and speed.
Once or twice they caught sight of a small figure,

apparently that of a young boy, which was hovering about
at a considerable distance from them, and hiding in nooks
and crevices, as if stealthily watching their proceedings.
"What is that? A lad or a dog?" asked Mr. Knibb, too much

excited to observe narrowly. "Catch it, my lads, whatever it
is. Depend upon it, it is contraband."
They gave chase for a few minutes; but the little object
darted away like a mountain sheep, and disappeared.
"Never heed," said the schoolmaster; "I don't much like

catching a poor child; even if we could, and torturing him
with fright into telling all he knows, and perhaps more than
he knows."
Besides his compassionate feeling, Mark was very anxious,

for his friend's sake, to get through the affair as quickly and
easily as possible, consistently with his strong sense of the
just and the right. And although he would have done
everything which he believed to be required of him as a
good subject, yet he was not sorry to see the little scout re-
appear at an amazing distance down the steep, and joining
a larger group of moving objects, wholly vanish together
with them behind a broad spur of the mountain. Pursuit was
hopeless, as the unknown figures had the advantage of
ground and of great distance.
"There go your potters, I should say, Mr. Knibb," said Mark.
"Likely enough," was the reply, in a tone of mortification.

"Much if they have not been at the still in the night and
carried off all the stock."
"We had best hurry on, and see about it," said Old Geordie
eagerly.
They did hasten on, gained the mouth of the adit, and
looked anxiously round to see if anything had been
disturbed. No; everything wore the same look as on the
previous day.
"Here we are, just in time," remarked Mr. Knibb, rubbing his

hands; "my potter friends were just then coming to strip
the place, and old George Knibb has for once got the start."
He was perfectly charmed with the completeness of the

little establishment, and went about sniffing and tasting
with marvellous zest, making entries in his note-book of
everything which the cave contained.
Whilst this regular inspection was going on, and whilst Mr.
Knibb was making his official inventory, Mark looked
anxiously round to see if he could discover any traces of his
friend's complicity in the lawless affair. Suddenly, he espied
a sheet of writing paper, which was stuffed into a corner,
and which looked as if it had been used to wrap up some
little refreshment, such as bread and cheese, or the like. A
thought struck him; that scrap of paper may afford
evidence of guilt.
He stepped across to the side where it lay, and, on looking

at it more closely, thought he recognized his own hand-
writing upon it. His own writing? How could this possibly
be? He looked at it again: "My dear Miles." Here, then, was
proof fearfully clear. This must be a letter which he had
written to Miles Lawson a week or two before, in reference
to the period of his regularly revolving visit to The Yews,
and which he had sent by a careful hand, not likely to have
failed in its due delivery.
He hastily seized the paper, and, crumpling it up, thrust it

into his breast; but this was done with such an undisguised
look of pale terror, that Old Geordie, who, like himself, was
on the look-out for traces of his young master's former
presence in this unlawful spot, saw at a glance that
something was wrong.
With his wonted almost feudal attachment to his house, he

hurried to the schoolmaster's help by calling out in a loud
voice, in order to divert attention. "To think that we should
be such a bad disloyal lot, as never to have minded that 'tis
our first duty to drink the king's health in his own 'mountain
dew,' with 'three times three,' my lads!"
"Ay, to be sure, so we ought," exclaimed the farm servants.

"Come, then, all of you, out to the mouth of the hole," said
the old shepherd leading the way—an invitation which was
but too willingly followed by the whole party, with the single
exception of Mark Wilson.
"Come, schoolmaster," said Mr. Knibb, "I hope you teach

loyalty to all your lads."
"Yes; but not by making them worthless subjects," replied

Mark, recovering from his ill-concealed consternation.
"You don't mean to say you are not going to drink his
majesty's health and long reign on this extraordinary
occasion?" expostulated the excited old exciseman.
"My voice shall swell the cheer," said Mark, smiling; "but my
lips shall never touch the white-fire, which might turn me
into a fool or a madman."
"The master is right, as he is a'most times," said Geordie;

"an old man had so well keep a clear head upon his
shoulders, when he has such a scramble before him a'down
over t' crags, with a heavy jar or two upon his back. I'll join
Master Wilson in a dry toast."
"But, Mr. Knibb," said Mark, "the spirits are not ours."
"Suppose then we honor the king, and give the cheer

without the whiskey. It will be a new sort of toast, but not
the worse for that." The exciseman agreed to the proposal
and gave the words, "Long live the king;" whereupon arose
a cheer from the mouth of that strange hollow, which the
mountain echoes took up and repeated, one crag tossing it
on: like a bounding ball to the next crag, the ledge taking it
up and flinging it on into the hollow, the ridge sharpening it
afresh before it could die away in the depths of the gorge,
and the ravine sending it softly down in a silvery whisper to
the sleepy lake below. The group of mountaineers was
almost startled by the marvellous effects of this ebullition of
their loyalty; for the excited echoes seemed to go on
repeating that hearty cheer as if they would never let it die
out—as if determined that it should become a living, abiding
voice of the hills.
However, time pressed, and they returned to business. It

was no trifling undertaking to remove such cumbrous goods
and chattels down the steep sides of a craggy mountain;
and more than once a disastrous slip occurred in the
experience of the heavy-footed farm servants, which sent a
jar bounding, leaping, curveting down the face of the rocks,
the liquid contents flying off in sparkling jots as it went,
insulting the innocent little juniper bushes which were
taking their long winter sleep beneath the snow, and
demoralizing the pure and spotless snows themselves.
Mr. Knibb was uncommonly exasperated by these accidents,

and was led to waste large stores of the "King's English" on
the occasion. "Why, it is direct treason. It is his majesty's
property that the fellow is destroying in that reckless way. I
declare I will report him—I'll prosecute him. There it goes
again. Next time I'll send you after it, and we shall see how
you will relish trundling in that fashion from rock to rock, all
down the precipice."
At length, the bivouac was reached, the carts were loaded

and despatched; and the weary procession again ascended
the mountain, to return heavily laden as before. The cavern
was at last completely dismantled; and the schoolmaster's
pen was speedily employed by Mr. Knibb in drawing up a
brilliant narrative of "The remarkable discovery of an illicit
still on Coniston Old Man."
A reward was afterwards offered for the apprehension of all
the offenders; but Miner Jack and Broom Tim had
succeeded in effecting their escape "over sands," though
chased by a far more relentless foe than the old exciseman,
even by that treacherous tide which walks up so calmly and
yet with such rapid steps, taking noiseless possession of the
sandy plain.
But to return to The Yews: Miles' lethargy lay so long and so

heavily upon him, that his patient mother at last thought it
would be desirable to rouse him. One evening, when she
had been administering her last resource (a burning hot tea
made of Cayenne pepper), which had appeared to
stimulate, not to say excite him, more than any previous
means that had been tried, she thought she would venture
to mention the name of Bella Hartley. But she was not
prepared for the suddenness of the effect produced.
Miles started up in bed, exclaiming, "I must go—I must go

and save her; no wrong shall ever come to her door
through me. The snow is deep, very deep; but nevertheless,
I must go, traitor or no traitor, informer or not."
With this he made a feeble effort to spring out of bed; but

the widow laid her hand upon his arm, settled his head
again on his pillow, and said, "Bella is safe; no harm has
come to her; the snow is all gone, and thou art lying quietly
in thy own bed with thy old mother sitting beside thee,
Miles, my son."
He looked first puzzled, then troubled; but the dull light in

his eyes was clearing, and gleam after gleam of intelligence
was passing slowly over his face. It was evident that the
heavy clouds were gathering up and rolling away, and that
the distempered mists and unhealthy fogs which lay over
the past were being dispelled by the reviving rays of
memory. But it was manifest to Mark's eye (for Mark, too,
was sitting beside his friend) that the backward views which
Miles was obtaining through these rents in the clouds were
often very painful ones. He changed color again and again,
and passed his hand across his forehead, as if to wipe out
their remembrance.
At last he took his mother's hand, and smiled such a smile

as she had not seen on her son's face for many a long day—
a smile of confiding love and of almost child-like simplicity.
"I shall have to tell thee all things that ever I did, I think,
mother; and thou must make thy heart ready to listen to a
sad story."
"If we confess our sins, God is faithful and just to forgive us

our sins, and to cleanse us from all unrighteousness," was
his mother's reply; "confess thy sins to God, and ask Jesus
to reconcile thee to thy Father and his Father, to thy God
and his God. Never mind the telling me all about them."
"I have confessed them to him, mother, and I do believe he

has put away their iniquity. But it will be right for me to
humble myself to the dust, after sinning with such a high
hand: and I wish, I really do wish that someone else, you
know who, was here to know all about it, too."
"No," said Mark, hastily, "if you mean the young woman,
she never need know all the sad particulars about the past,
now. Miles will lead a new life; he will uphold the good old
family motto; he will 'Feare God, and worke ryteousnesse,'
and the young woman's respect and love never need be
shaken, I should think."
"She will only love him the more for all he has suffered,"
said a gentle voice; but when they looked round, Alice
flushed and went to the window.
"Alice is right," said Mark Wilson, looking at her with fond
pride; "there is nothing like honesty, and openness and
truth. It was only to spare her and to spare him that I took
the weaker part. We always walk safest and surest in broad
daylight."
Miles held out his hand to his old friend and said, "That was
the doctrine you always taught me; but I have been a bad
scholar indeed. However, I shall tell Bella everything when I
am strong enough; and then we shall see whether she will
ever trust me again."
CHAPTER VII.
CHANGING SEASONS.
"Life is astir beneath dead Nature's snows

Spring's quickened pulse is bounding
through the earth.
Lo, in the wakened heart a life-stream
flows:
Old things are dead; behold a second
birth."
TIME passed on. Months had slipped away since the mind of
Miles Lawson had awakened to the consciousness of the
past. But the recovery of his bodily powers had not kept
pace with the renewal of the mental. A low fever, the
natural rebound of the death-like chill, had been lying
heavily upon him, completely prostrating his strength, and
refining his rude manly frame into something too ethereal
and shadow-like for a young mountaineer.
His brown and sinewy hands had become white and almost
transparent; his cheek hollow and pale, save for the small
bright spot which lighted it up, while his eyes looked
prominent and lustrous as lamps. But the expression of his
whole countenance and manner had as decidedly changed
for the better as his health had altered for the worse: the
countenance was now open, winning, and thoughtfully
intelligent, instead of sullen and unhappy: the manner was
gentle and deferential, instead of capricious and intolerant.
No one who knew anything of the changing signs of the

moral seasons—seed-time and harvest, winter and spring—
could doubt these outward evidences of the inward work of
grace. That change had been going on which is described in
Scripture language as the turning from darkness to light,
from the power of Satan unto God. The depths of
repentance had been passed through, not once only, but
again and again; the sweetness of reconciliation had been
known with a justly offended God through the free and full
atonement offered for sinners by the Son sent by the
Father: and thus delivered from the bondage of sin and
Satan, the new creature in Christ Jesus was seeking to live
a new life through the power of the sanctifying Spirit.
But Miles Lawson, after his many slips and wanderings, had
found it good for his soul's health to linger long in the
"valley of humiliation," and he found it to be such as John
Bunyan describes it, "as fruitful a place as any the crows fly
over."
"I have known many laboring men," says Bunyan, "that

have got good estates in the valley; for indeed, it is very
fruitful soil, and doth bring forth by handfuls."
A beautiful change in the outward aspect of the dale, and in

the habits of dale life, had been going on during the same
period. It is scarcely like the same region. True, the noble
outlines remain immovable—the mountains drawing their
fine forms against the sky; the lower fells crossing each
other in those graceful intersecting lines which the eye so
loves to follow; the valley biding with such shy and shady
reserve under the glooms of the projecting crags; and the
stream finding its way with its lovely curves and bends,
forever humming its mountain melodies.
But everything else has changed: the coppice wood at the

mouth of the glen is one sheet of varied and delicate
greens; the rough leaves of the hazels are intermixed with
the silver stems and small bright foliage of the birch;
sycamores are shaking out their broad leaves, creased and
puckered with their tight foldings in the buds, the oaks are
sunning their finely-cut leaves; and the ashes, last to come
and first to go, are waving their sprays in the breeze; while
the larches have long ago hung out their light green tassels,
and are now creeping up the sides of the mountain with
pointed crests, and in close array.
All color has changed, saving the dull dark hues of the time-
worn pines, and the grand and sombre masses of the
ancestral yews: what is the short summer of the dales to
them? It may be a fleeting joy for the ephemeral foliage
around them. It may make a holiday for the golden brooms,
and the hedges of snowy thorn, and the festal plumes of
the bird cherry. It may cause a flutter of excitement in the
sensitive sprays of the aspen, and make the green moss-
beds first all silvery with snowdrops, then all golden with
nodding daffodils and starry primroses, and again all blue
with bell-hung hyacinths, and pearly with the shy wood
anemone. But to them—to these dignified sires of vegetable
life, what is the fleeting influence of season? Hoary winter,
song-resounding spring, festal summer, golden autumn—
these can scarcely impart an added furrow, or wreathe their
stately brows with any passing glory.
Everything else, however, seems young and jubilant. Look

at the lambs upon the springy turf of the fells. They are
playing like kittens. No, better than that they are getting up
regular games of their own. There is system in that fun of
theirs. That fat, saucy fellow, white as snow, save for his
black nose and his legs, which seem to dance all the lighter
for their little black worsted stockings—he is evidently
master of the revels. He marshals his band on the top of
that old gray rock that bares its forehead from amidst the
elastic turf on the mountainside. There are some of the
young rebels who are determined to scale the height from
beneath in a wholly unauthorized way. That will never do;
the leader and his lambs line the ramparts, and butt and
push at every black nose that aspires upwards. The
aspirants are beaten back; and then down comes the whole
garrison, leaping, bounding up in the air "all fours" at once;
and sweeping away the opposing force, the whole lamb
community careers away in one troop down the green slope
of the fell.
This is thought to be rather too wild work by those

respectable old ewes, who, in their staid sobriety, have
altogether forgotten the days of their youth. They lift their
plain and anxious countenances from cropping the scented
turf, and, with mouths full of thyme and heath, utter a few
warning remonstrances, which only seem to stimulate the
wild frolics of the young folks. Surely they don't mean to
send that black, perfectly black lamb "to Coventry"? The
insolent little rogues, it really looks so. They won't let him
join in the fun, forgetting their own legs and noses: for they
are but quadroons themselves, at the best: and so he
retires to the genial society of his mother, in whose eyes he
seems to be white and comely as a lily. Really, it is rather
trying to think that these charming lambs will grow up into
those ungainly and uninteresting old sheep.
Alice's garden is brilliant with flowers. "'Tis but a common

cottage garden," you will say: no exotics, no rare and
delicate plants. But here are England's dear old favorites,
her best and choicest flowers. What can surpass those regal
"cabbage roses," so round and so full, or those moss rose-
buds growing under the lee of the white porch? Then there
are "sweet-williams," deep red and variegated, very stiff
and very handsome in their way: there are showy orange-
colored lilies and queenly white ones: there are purple
columbines, and great red peonies, and tall "Jacob's
ladders," and grave "Solomon's seals": there are graceful
sweet peas clinging for support to anything they can reach;
and a little bed of spicy pinks, scenting the air like an island
of the eastern seas. The beds are all edged either with box,
or with double red and white daisies, or with the little fragile
"witch's thimbles," chiming their small blue and white bells
to every passing breeze.
But there is something going on under the broad shadow of

the yew trees. On the smooth grass-plot which spreads
before Alice's rustic seat, stands the long kitchen table,
which must have been moved out of doors for festal
purposes, because it is covered with a clean white table-
cloth, and a number of chairs are placed round it. Plates of
piled-up bread and butter, cut and buttered currant "wigs,"
a massive-looking cake, whose consistency is very much
that of cold "figgy pudding," a beautiful dish of ripe
strawberries, dressed out with leaves worthy of a ducal
coronet *, and another dish of cherries from off the walls of
the house, furnish the entertainment. There is a tea-tray
also; and Mrs. Lawson's dozen of little old-fashioned silver
teaspoons have been taken out of the old oak chest and
rubbed up for the occasion.
* The strawberry-leaf is the ducal symbol, and is placed

on the coronet.
But who are the guests? The widow is there, sitting up in

her chair in considerable state as hostess; Alice, in simple
gala costume, is flitting about with a brilliant flush of
pleased expectation on her artless countenance; Mat is
gone in to rub up his merry face until it shines like a rosy-
cheeked apple, to comb his light hair down over his sun-
burnt forehead, and to put on his best red waistcoat, with
blue glass buttons, and his bright green neck-tie. Chance
and Laddie have made no such distinguished toilette, but
they are particularly on the alert, barking little gala salvos,
and pricking their ears at every sound.
There is a sound now in the distance—a slow rumble of

wheels in the direction of Green Gap. The dogs are violently
excited by this, because a friend, a great buff sheep dog,
with an enormous white plumy tail, always comes in
company with the rumbling "shandry" * from Scarf Beck
Farm. They are right in their apprehensions.
Mr. and Mrs. Hartley, a particularly comfortable, not to say

jolly-looking couple, are sitting up in the high-backed seat,
with Bella between them, and the young Hartley brothers
are walking in front. This is evidently a state visit; tea out
of doors at half-past three o'clock, and supper looming in
the background at seven.
* The "travelling carriage" of the country—half-cart, half-

gig.
We must have a photograph of Bella Hartley; for she is

called by many, the Queen of the Dales. She is taller by
half-a-head than young Alice Lawson, and of a far more
noble figure and carriage. Her features are finely-cut; and
her head, with its bountiful profusion of bright brown hair,
sits with remarkable grace on her long neck and finely
sloping shoulders. Her dress is a simple light print, with a
colored ribbon round her throat.
Ah! there is a little story attached to that rather faded

ribbon, and some amount of sentiment; or else it would not
have been selected for this especial occasion. She has a
grave, thoughtful, perhaps anxious look; but it has not been
always there. Her face was as sparkling as her own Scarf
Beck, until some twelve months ago, when, after a long and
painful talk with Miles Lawson, that care-worn expression
took up its settled abode on her lofty brow, and in her deep
shadowy eyes.
On that summer evening, a year ago, she had said to him,

"No, Miles, you must give me back my promise; for I will
never be the wife of a man that I cannot trust out of my
sight for half-an-hour."
And so they parted—he to grow more thoughtless and

reckless than ever, because he held himself to be an injured
man; she to watch over him from a distance with the folded
hands of prayer, and to weep and mourn in secret; while
she carried before her little mountain world a brow so calm,
and a manner so serene and collected, that folks said she
was unfeeling, fickle, and heartless.
Other young farmers, and one or two land or mine agents,

had paid their court in the meantime; but no one had been

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CHAPTER 21 Careers and Decision Making

Chapter 4 The Circulatory System 55

A Day in the Life

draped around my neck! It seems like yesterday, even

Chapter 3 The Musculoskeletal System 41

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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,

© 2017 Cengage Learning®

© 2017 Cengage Learning®

© 2017 Cengage Learning®

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

A Day in the Life

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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-

© 2017 Cengage Learning®

the inside of the cell contained. The cell membrane is

The specialized proteins in the cell membrane influence

Ribosomes Centriole Lysosome

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Smooth Endoplasmic Reticulum

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CELL FUNCTION 6. Waste products

Objective eliminated by the ECF. Without elimination, the waste

Molecule Hypotonic Solution

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In certain situations, a cell may require a higher

Large molecules, such as proteins, must be moved

the diffusion occurs. Some large molecules such as

the membrane to equalize the concentration; however,

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

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Color codes for the bases

mRNA Transcribed Strand

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Amino Amino Amino Amino Amino Amino O

RLong Hydrocarbon Chain Fatty Acid