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 PREFACE
TO THE THIRD EDITION
I
t is our hope that students and research-
ers will continue to ind this atlas a
useful resource. In this edition, we have
expanded many of the beginning chapters
that introduce the basic types of tissues to
provide the user with a stronger founda-
tion in histology. The glossary has been
expanded, and other extras have been
included that we hope will be valuable.
Once again, many thanks to all of those
who have made the irst and second
edition of our atlas possible! We were able
to prepare our page layouts for the third
edition thanks to the scanner given to us
vii
by Aunt Lila and Uncle Jack McKean and
Lola and Chip Wood. Thanks to Jessie
Bacha for proofreading our work, and to
Tristan Bacha for keeping busy while we
worked! We greatly appreciate the helpful
suggestions from Professor Nancy
Gartland and the students at the University
Of Pennsylvania School of Veterinary
Medicine, and from reviewers and users
of the previous edition. Finally, thanks to
Nancy Turner, Erica Judisch, Tracy
Petersen, Erin Magnani, and all of the
other people at Wiley-Blackwell for their
role in the production of this edition.
 PREFACE
TO THE SECOND EDITION
W
e wish to thank those who have
used the irst edition for their
suggestions. We believe the incor-
poration of many of these recommenda-
tions will make this edition even more
helpful to the user.
To this end, we have updated the mate-
rial for the second edition by scanning all
of the original kodachromes and relabel-
ing the art. We have added thirteen new
photographs and have enlarged over one
viii COLOR ATLAS OF VETERINARY HISTOLOGY
hundred others. Four of the original black
and white line drawings have also been
redrawn. Also, a glossary of nearly 750
words has been added.
The style, format, and purpose of this
edition remain essentially unchanged
from the irst edition. We continue to view
the atlas as a useful, benchside reference
for those interested in understanding and
interpreting histologic and cytologic
preparations.
viii
 PREFACE
TO THE FIRST EDITION
A
lthough we have written this atlas
primarily to fulill a need of the
student of veterinary medicine, we
believe that clinicians, private practitio-
ners, and researchers will ind it a useful
reference for normal tissues and organs.
Currently, students rely heavily, if not
exclusively, on atlases of human histology
for guidance in the laboratory. There are,
of course, similarities between organs and
tissues of domestic animals and those
of humans. There are also differences,
however, and these are rarely encountered
in atlases dealing speciically with human
histology.
Our aim has been to compare the his-
tologic structure of organs in a variety of
domestic animals. We have used represen-
tative examples in instances where tissues
and organs from different animals share a
common structure. Wherever differences
exist, we have tried to provide examples
that are characteristic of a particular
group of animals. Our selection of animals
includes the dog, cat, horse, cow, sheep,
goat, pig, and chicken because they are
most frequently referenced in veterinary
school curricula.
ix
All photomicrographs and drawings are
original. Some drawings were done free-
hand, while others were made with the aid
of a camera lucida. Light microscopy and
colored photomicrographs have been used
exclusively. We have chosen color rather
than black and white because of its corre-
spondence to stained preparations. With
the exception of the few histologic prepa-
rations loaned to us by generous donors or
purchased from a dealer, slides were pre-
pared by the authors. Fresh organ samples
were obtained from a slaughterhouse or
from animals that were euthanized for
various reasons. With the exception of
smear preparations (blood, bone marrow,
and vaginal), mesenteric spreads, ground
bone, and a single plastic section, slides
were prepared using the parafin method.
All slides were stained with hematoxylin
and eosin unless otherwise noted.
Magniications of photomicrographs are
total magniications (enlargement of pho-
tograph × objective × projector lens).
Throughout the atlas, hollow structures,
for example, blood vessels, kidney tubules,
and alveoli, are usually identiied by label-
ing the lumen of the structure.
 ACKNOWLEDGMENTS
FROM THE FIRST EDITION
H
elp is often just around the corner.
Dr. Henry Stempen, whose ofice
was down the hall from ours at
Rutgers University in Camden, New Jersey,
stopped by one day and volunteered his
artistic talents. We’d like to thank him for
his excellent pen and ink drawings of
various animal parts, which are somewhat
removed from the fungi he usually draws.
Our gratitude also to Ms. Kathleen Carr
for her secretarial services. Special thanks
are extended to Dr. Edward Zambraski,
Ms. Kathleen O’Hagan, and Ms. Gail
Thomas of Cook College, Rutgers
University, for making fresh porcine mate-
rial available to us, and to Dr. Barry Jesse
and Dr. James Harner for supplying us
with sheep parts.
Without the unqualiied use of the facili-
ties and equipment of the Biology
Department of Rutgers, our tissue process-
ing and photomicrography could not have
been accomplished. Our special thanks to
the department for this courtesy.
This book would never have had a
beginning were it not for the generosity of
Dr. Leon Weiss, Department of Animal
Biology, University of Pennsylvania School
of Veterinary Medicine, who invited us to
teach in the veterinary histology labora-
tory and kindly allowed us access to the
slide collection and facilities of the depart-
x
ment. We would also like to express
appreciation to the following individuals
from the University of Pennsylvania
School of Veterinary Medicine: Mr.
Richard Aucamp and Mrs. Kathy Aucamp,
who provided us with specimens, slides,
advice, and assistance in a variety of other
ways; Dr. Mark Haskins for kindly making
available fresh canine and feline material;
Dr. John Fyfe and Dr. Vicki Meyers-
Wallen for supplying us with canine
vaginal smears; Dr. and Mrs. Loren Evans
and Dr. David McDevitt for lending us
reference material; Dr. Peter Hand and
Ms. Graziella Mann for providing mate-
rial on the nervous system; and Dr. Helen
Acland, Dr. Linda Bachin, Mr. James
Bruce, Dr. Sherrill Davison, Ms. Dawn
Dowling, Dr. Robert Dyer, Dr. Robert
Eckroade, Dr. George Farnbach, Dr.
David Freeman, Dr. Wendy Freeman, Dr.
Alan Kelly, Mr. Joseph McGrane, and
Dr. Mary Sommer for their time and con-
sideration in helping us to obtain tissue
specimens.
We are grateful to Dr. Carol Jacobson
and the Department of Anatomy of the
Iowa State University College of Veterinary
Medicine for providing valuable slide
preparations and text material.
Our gratitude is also extended to Hill’s
Pet Products, Topeka, Kansas, and
Pitman-Moore, Inc., Washington Cross-
ing, New Jersey, for their generous inan-
cial assistance.
Many thanks also to: Dr. Caroline
Czarnecki of the University of Minnesota,
College of Veterinary Medicine, for pro-
viding copies of her informative labora-
tory guide; Dr. Deborah Ganster, Dr.
James Lawhead, Dr. Virginia Pierce, Dr.
Maria Salvaggio, Dr. Barbara Strock, and
Dr. Cindi Ward for assisting us in obtain-
ing tissue samples; Mr. Jeff Bringhurst,
Bringhurst Brothers, Tansboro, New
Jersey, for allowing us access to fresh large
animal material; the Longenecker
Hatchery, Elizabethtown, Pennsylvania,
for providing chicken specimens; Ms.
Susan Ulrich, Cornell University Press, for
lending us a dificult-to-obtain reference;
the helpful people at Optical Apparatus
Company Inc., Ardmore, Pennsylvania,
for supplies and for assistance with equip-
ment for the microscope; and Mr. Charles
Behl and Mr. James Durso of Webb and
xi
Company Inc., Cherry Hill, New Jersey,
for their courteous service and helpful
advice.
We are indebted to Mr. William J.
Bacha, Sr., for building a super light box
for us, and to Mr. Thomas H. Wood, Jr.,
for providing black and white prints of
our photomicrographs, which saved us
countless hours of drudgery in the dark-
room. Thanks to Barbara Frasco, Esq.,
for her helpful advice. Our hats are off
also to Snuff, Chew, Chapter Seat, Angel,
Clyde, and all the other animals for their
participation.
We also wish to extend our gratitude to
all at Lippincott Williams & Wilkins
whose efforts helped bring this second
edition into being. We are especially grate-
ful to Carroll C. Cann and Jennifer D.
Weir for their professional advice, cour-
tesy, and assistance.
William J. Bacha, Jr.
Linda M. Bacha
GENERAL PRINCIPLES OF HISTOLOGY xi
 CONTENTS

1 General Principles of Histology 3

2 Epithelium 11
3 Connective Tissue Proper and Embryonic Connective Tissue 19
4 Cartilage 27
5 Bone Tissue 31
6 Blood 41
7 Bone Marrow 53
8 Muscle Tissue 57
9 Nervous System 65
10 Cardiovascular System 77
11 Lymphatic System 89
12 Integument 105
13 Digestive System 139
14 Urinary System 183
15 Respiratory System 195
16 Endocrine System 211
17 Male Reproductive System 225
18 Female Reproductive System 243
19 The Eye 267
20 The Ear 283
Glossary 291
Bibliography 309
Index 311

xiii
COL OR ATL AS OF
VETERINARY
HISTOLOGY
Third Edition
 1

GENERAL PRINCIPLES OF
HISTOLOGY

PREPARATION OF HISTOLOGIC SECTIONS

A
histologic section is a thin slice of tissue, varying, usually, from 0.5 to 10 or more
micrometers (µ) thick. In preparing such a section, a piece of tissue is either inil-
trated with a supporting medium or frozen and is then cut with an instrument
called a microtome. Sections obtained from tissue iniltrated with plastic can be as thin
as 0.5 µ and show superior detail. Excellent preparations as thin as 2 or 3 µ also can be
made from tissue iniltrated with parafin-based embedding media. Sections are afixed
to microscope slides and colored with one or more stains to increase the visibility of
various cellular and intercellular components.
Schematically, Figure 1.1 outlines various steps involved in producing a stained
histologic slide using the parafin procedure. After being removed from an animal, a
tissue or organ is cut into pieces. These pieces are placed into a ixative such as buffered
formalin or Bouin’s, which, ideally, preserves normal morphology and facilitates further
processing. After ixation, the specimen is dehydrated by transferring it through a series
of alcohols of increasing concentrations to 100% alcohol. Next, it is placed into a sub-
stance such as xylene or xylene substitute, which is miscible with both 100% alcohol
and parafin. This intermediate step (called clearing) is essential before iniltrating the
dehydrated tissue with parafin because alcohol and parafin do not mix. During iniltra-
tion, melted parafin completely replaces the xylene. This procedure is done in an oven
at a temperature just above the melting point of the parafin. When iniltration is com-
plete, the specimen is transferred to an embedding mold of fresh parafin, which is
allowed to harden. Then the mold is removed and excess parafin is trimmed away.
The block of parafin is then secured to the microtome and oriented appropriately
with respect to the knife. With each revolution of the microtome handle, the specimen
moves through the blade and a section of the desired thickness is produced. Each suc-
cessive section adheres to the preceding one, forming a continuous ribbon. Subsequently,
one or more sections are carefully separated from the ribbon and transferred to the

3
 Figure 1.1. The various steps involved in producing a histologic slide using the parafin method.

4 CHAPTER 1
surface of warm water in a waterbath. This softens the
parafin and lattens the section, eliminating wrinkles. The
lattened section is loated onto a slide, which is then
placed on a warming table. As the preparation dries, the
section adheres to the surface of the slide.
Next, the parafin is removed with xylene or another
appropriate solvent and the specimen is rehydrated. It is
then stained, dehydrated, cleared (made transparent) with
xylene, covered with a resinous mounting medium, and
topped with a cover-slip.
Various stains are available to the histologist.
Hematoxylin and eosin (H&E) is a frequently used com-
bination of stains. Hematoxylin imparts a purple color to
substances, but must be linked to a metallic salt called a
mordant before it can function effectively. This combina-
tion, called a lake, carries a positive charge and behaves as
a basic (cationic) stain. The lake combines electrostatically
with negatively charged radicals such as phosphate groups
of nucleoproteins. Substances that become colored by a
basic stain are said to be basophilic. Methylene blue, tolu-
idine blue, and basic fuchsin are basic stains. Unlike hema-
toxylin, these stains have molecules that carry a positive
charge of their own and do not require a mordant. Acidic
(anionic) stains carry a negative charge and color cell or
tissue components that bear positive charges. Eosin is an
acid stain. It imparts an orange or red color to acidophilic
substances. Other commonly used acid stains are orange
G, phloxine, and aniline blue.
In addition to the widely used H&E staining proce-
dure, numerous other stain combinations and techniques
are available. Some are especially useful for identifying
certain tissue elements. For example, trichrome procedures
such as Mallory’s and Masson’s speciically stain collage-
nous ibers within connective tissue. Orcein and Weigert’s
resorcin fuchsin are stains used to color elastic ibers, pro-
viding a means of distinguishing them from other ibrous
elements. Reticular ibers and nervous tissue components
such as neurons, myelin, and cells of the neuroglia can be
stained by procedures employing the use of silver. There
are also special histochemical and immunohistochemical
procedures that make possible the localization of various
carbohydrates, lipids, and proteins found in tissue. Lastly,
stains such as Wright’s and Giemsa’s (Romanovsky stains)
are available for differentiating the various cells found in
blood and bone marrow.
INTERPRETING SECTIONS
One must know the gross structure of an organ before a
histologic section from it can be comprehended. It is also
helpful to know how the section was cut, that is, whether
it was a cross section (x.s.), a longitudinal section (l.s.), or
an oblique slice through the organ. Was the cut made
through the entire organ or only through a portion of it?
Frequently, prepared slides are labeled indicating the par-
ticular orientation of the section. This is not important in
an asymmetric organ such as the spleen or liver because
their appearance would be unaffected by the direction of
the cut. Conversely, the small intestine is radially symmet-
ric and its appearance is affected by the direction of the
cut.
The three-dimensional structure of organs and their
components also must be considered when examining a
histologic preparation. Cells are three-dimensional objects
differing in size and shape. For example, some are long
and thin, some cuboidal, and others ovoid. They may have
a random or speciic arrangement within an organ. How
they appear depends on their shape as well as how they
were cut. Imagine how the spindle-shaped and tall colum-
nar cells shown in Figure 1.2A would look if sectioned in
various planes. Note that the nucleus may or may not be
included in a particular cut through a cell.
The histologist examines multicellular structures
having a wide variety of shapes. Some are hollow, some
branch repeatedly, some open onto surfaces, etc. Figure
1.2, B and C, and Figure 1.3 show a variety of three-
dimensional structures and how they would appear if cut
at different levels. Examine these carefully. They will help
you to understand situations you will encounter on actual
slides.
HELPFUL HINTS
Be sure that the lenses of your microscope are clean before
you begin examining slides. Use a piece of lens paper or a
soft, clean cloth such as an old (but clean) linen handker-
chief. If the lenses have been coated with oil or another
substance, remove it using lens tissue moistened sparingly
with a glass cleaner such as Windex. Slides also should be
cleaned using a soft, lint-free cloth or tissue moistened with
glass cleaner.
Every microscope should have a pointer in the ocular.
This is usually supplied by the manufacturer, but can be
made from a short piece of hair. The latter is cemented into
place inside the ocular with a dab of quick-drying glue or
nail polish. Without a pointer, it is not possible to accu-
rately indicate an object in the microscope ield for another
observer.
Before beginning a session at the microscope, make
sure that the ine-adjustment knob is near the middle of its
range of rotation. If you do not, you may ind that the
knob is at the limit of its excursion when you are busily
making observations. At that point, you must stop every-
thing and correct it.
It is also a good habit to examine your slide with the
unaided eye before placing it on the stage of your micro-
scope. By doing so you will gain information about the
gross aspects of the specimen and be more likely to center
it properly over the light source. Centering is especially
important for small specimens that might otherwise be
dificult to locate. Also, make sure that you put the slide
on the stage with the cover glass uppermost. If the slide is
upside down, you will not be able to focus on it with the
high-power lenses. Do not snicker. We have seen this
happen often in the teaching laboratory!
It is always a good idea to start your observations
using the lowest power objective available on your micro-
scope. This is usually the 4x lens. The ield of view will be
GENERAL PRINCIPLES OF HISTOLOGY 5
 Figure 1.2. A. Slices, indicated by numbered planes, taken through two different types of cells would
appear as identiied by the prime numbers. Only if the plane of the cut passes through the nucleus will
the latter be seen. B and C. Planes of section taken from different levels in four separate multicellular
objects are illustrated. Note how the appearance of sections varies with the level of the cut.

6 CHAPTER 1
Figure 1.3. The prime numbers illustrate sections resulting from transverse (4), oblique (1), and longitu-
dinal (2, 3, 5, 6) cuts made through a plate of cells bearing hollow projections (above plate) and invagi-
nations (below plate). Plane 3 differs from the others because it passes only through the cellular wall of
a projection, and not the lumen; therefore, section 3′ appears as a plate of cells rather than a hollow
structure. You should also be aware that structures may often appear unrelated to a surface or another
object, when in fact they are. Compare planes 5 and 6 with sections 5′ and 6′, where continuity of the
invagination with the surface is evident only in 6 and 6′. While not apparent from a single section, such
continuity would be evident if an uninterrupted series of sections through the entire invagination were
made and examined.

GENERAL PRINCIPLES OF HISTOLOGY 7

large, enabling you to locate regions of special interest
more easily. When you locate something you wish to
examine at a higher magniication, center the object in
the middle of the ield of view. Then, when you change
to a stronger lens, the object should be somewhere in the
ield.
Binocular microscopes often have at least one ocular
that can be adjusted to accommodate your vision. It is
important that you adjust this properly if you want to have
a comfortable, headache-free session at the microscope.
Assuming that your microscope is of the binocular type
and that it has at least one adjustable ocular, you should
irst bring the specimen into focus with the ocular that is
not adjustable by using the ine-adjustment knob. When
you have done this, focus the other eye using the adjustable
ocular. Use of this procedure will ensure a proper focus for
both eyes and prevent eye strain.
Yikes! After you turn on the microscope, you do
not see any light as you look through the
oculars! Before you replace the bulb, try the
following:
• Check the light control lever or knob to make sure the light
intensity is not set too low or turned down to zero.
• Check that the objective lens is clicked fully into position.
• Make sure that the plug is itted properly into the electrical
outlet and that the outlet is working.
Bright, even lighting is absolutely essential to effective
microscopy. The best way to achieve this is to use Köhler
illumination. This can be obtained with any microscope
that is equipped with both a condenser aperture diaphragm
(the one in the condenser) and a ield diaphragm (the one
in the light source). If you have such an instrument, proceed
as follows:
1. Center the light source, using the directions you
received with the microscope.
2. Open both the ield and aperture diaphragms fully.
3. Raise the condenser to its uppermost position.
4. Place a specimen on the stage and focus on it using
the 10 × objective.
5. Close the ield diaphragm so that its leaves are clearly
imaged in the ield of view.
6. Center the image of the diaphragm by manipulating
the condenser centering screws, then open the ield
diaphragm until its leaves disappear just beyond the
edge of the ield of view.
7. Remove an ocular and, while looking into the back
aperture of the objective, close the aperture dia-
phragm completely and then open it until it is about
75% of being fully open.
You now have Köhler illumination. If you want to
increase or decrease the light intensity, use the rheostat or
neutral-density ilters, but do not adjust the condenser
aperture diaphragm or ield diaphragm. If the aperture
diaphragm is open to excess, the image will lack some
contrast and be looded with light. If it is closed too far,
8 CHAPTER 1
there will be a loss of resolution and increase in contrast.
This increase in contrast is often confused with sharpness
or high resolution; this is a common error in microscopy.
All of the above adjustments (except for centering the light
source) must be made each time a different objective is
used.
If your microscope lacks a ield diaphragm, you will
not be able to obtain Köhler illumination. You can still
acquire good and useful lighting, however. Place a speci-
men on the stage, open the aperture diaphragm fully, and
adjust the light intensity with the rheostat so that it is
comfortable for your eyes. Be sure that the condenser is
raised to its highest position, or close to it, when you do
this. Now, remove an ocular and look at the back aperture
of the objective. Close the aperture diaphragm fully and
then open it until it is about 75% of being fully open. This
will provide proper lighting for most purposes. If you
should need more or less illumination, make adjustments
only with the rheostat or neutral density ilter; do not use
the aperture diaphragm.
To get the most from a specimen, you must avoid
being a passive microscopist, that is, one who inds an
object and then stares at it admiringly without making
further adjustments of the focus. Get into the habit of
focusing continuously with the ine adjustment as you
peruse a slide, because even though a tissue section may
be only a few micrometers thick, the depth of ield of the
higher power objectives may be less than the thickness of
the specimen. Therefore, if you do not focus repeatedly as
you examine a preparation, you will certainly miss seeing
structural detail that might be important to your work.
You might like to return to a particular location on
your slide preparation at a future time. Remembering land-
marks in the vicinity of the object of interest will aid you
in locating the object later. A more expedient way of relo-
cating structures is by using verniers, which are mounted
on both the X and Y axes of the mechanical stage. A
vernier consists of two parallel, graduated, sliding scales,
one long and one short. The smaller scale is 9 millimeters
(mm) long and is divided into 10 subdivisions (0 to 10).
The larger scale is several centimeters (cm) long and is
graduated in millimeters, for example, 0 to 80 or 100 to
160. To relocate an object on a slide, you must irst center
it in the microscope ield. Once this has been done, you
establish its location by reading each of the verniers (X and
Y). For example, the 0 point on the small scale of the
vernier on the X axis might be located somewhere between
lines 42 and 43 on the larger scale (Figure 1.4). To deter-
mine its speciic location, ind the line on the small scale
that coincides exactly with a line on the longer scale. Then
count, on the smaller scale, the number of spaces between
0 and the point of coincidence. This number is your
decimal point. In the example given (Figure 1.4), the
Figure 1.4. Small and large vernier scales.
decimal is 0.6 and you should read 42.6 as the vernier
value. Do the same for the other vernier (Y) and record
the numbers for both. In the future, if you want to return
to the same location, simply secure the slide to the mechan-
ical stage and move the stage controls until the verniers are
adjusted to the numbers you previously recorded. These
manipulations will have returned the slide to its former
position, and the object you are looking for should be
somewhere within the microscope ield.
By knowing the approximate diameter of a red blood
cell in a section, you can estimate the size of other tissue
components. Therefore, it is useful to know that in tissue
sections prepared by the parafin method the average size
of erythrocytes for each of the following animals is as
follows:
Goat: 2.4 µ diameter (smallest erythrocytes of the
domestic mammals)
Dog: 4.9 µ diameter (largest erythrocytes of the domes-
tic mammals)
Chicken: 9.4 µ long
Each average value is based on a total of 20 to 30 cells
that were measured from ive different slide preparations
of tissues embedded in Paraplast X-TRA (Monoject
Scientiic, Division of Sherwood Medical, St. Louis, MO
63103).
ARTIFACTS
Folds, knife marks, stain precipitate, spaces (where none
belong), shrinkage, and air bubbles are examples of
common imperfections seen in slide preparations. They
were introduced during processing and are called artifacts.
Figures 1.5 through 1.9 are examples of such artifacts.
Troubleshooting a blurred or cloudy image:
• If an image is cloudy or blurry, the oculars and/or objective
lenses may need to be cleaned. To determine if an ocular
needs to be cleaned, turn it as you look through the micro-
scope. A dirty mark or smear will rotate if the ocular is not
clean.
• If the image is still blurred, clean the objective lenses.
• Make sure the slide is resting on the stage properly and is
right side up! Sometimes an image cannot be focused
clearly because the slide is on the stage upside down!
• Are you using the oil immersion lens without a drop of oil
on the slide?
• Be sure that the light source is not partially blocked by
something, such as the electric cord or a ilter holder below
the stage.
GENERAL PRINCIPLES OF HISTOLOGY 9
 3
3
6

Figure 1.5 62.5 Figure 1.9 62.5

2 KEY
1. Dermis 4. Knife mark
2. Epidermis 5. Separation artifact
3. Fold 6. Stain precipitate
5

Figure 1.5. Stain Precipitate, Cartilage, Dog. Occasionally, solu-

1 tions accumulate precipitate that may stick to the surface of tissue
sections during the staining procedure.

Figure 1.6. Separation (Space) Artifact, Skin, Dog. Tissues may

5 be subjected to excessive pressures, tensions, or shrinkage during
processing, resulting in separations within otherwise intact tissue.

Figure 1.6 62.5 Figure 1.7. Crackling Artifact, Thymus, Horse. Highly cellular
tissues, for example, thymus, liver, pancreas, and spleen, often
show numerous tiny cracks throughout. Also note that this speci-
men is not in sharp focus.

Figure 1.8. Knife Marks and Folds, Esophagus, Horse (Masson’s).

Knife marks (scratches) in the tissue section may be caused by
defects in the microtome knife or by accumulations of debris on
the knife edge. Folds occur when the tissue sections fail to spread
properly on the surface of the slide.

Figure 1.9. Fold, Aorta, Pig. In a tissue section, folds are raised
areas that frequently overlap. Note that portions of this picture
are not in sharp focus.

Figure 1.7 62.5

3 4

Figure 1.8 25

10 CHAPTER 1
 2

EPITHELIUM

E
pithelium is one of the four main types of tissue in the body along with connective
tissue, muscle tissue, and nervous tissue.

GENERAL FEATURES
Epithelium is a ubiquitous tissue that covers surfaces, lines cavities, and forms glands of
the body. It is comprised entirely of cells separated only by a thin layer of intercellular
substance that helps hold them together. The cells are supported by a thin extracellular
basement membrane that separates them from the underlying connective tissue.
The free (apical) surfaces of epithelial cells may possess cilia (motile processes),
microvilli (short inger-like projections of the cell membrane), or stereocilia (long
microvilli).
When the epithelium consists of a single layer of cells, it is called a simple epithelium.
If it is formed from two or more layers of cells it is said to be a stratified epithelium.
The individual cells of either type of epithelium may be squamous, cuboidal, or columnar.
In proile view, squamous cells are lat, cuboidal cells are short cells that are approxi-
mately as tall as they are wide, and columnar cells are rectangular cells that are taller
than they are wide.

SIMPLE EPITHELIAL TISSUE

Simple epithelia, formed by a single layer of cells, are found in many different locations
throughout an animal’s body. They line the heart; blood and lymphatic vessels; and

11
pleural, pericardial, and abdominal cavities; and they form
Summary of Types of Epithelial Tissue
the secretory units of many glands. They also line the
stomach, intestines, and portions of the respiratory system. Simple Epithelium
Types of simple epithelia, simple squamous, simple cuboi- Simple squamous
dal, and simple columnar, are named according to the Simple cuboidal
shape of the cells. Pseudostratified columnar is a special Simple columnar
form of simple epithelium. Although it appears in proile Pseudostratiied columnar
Stratiied Epithelium
to be multilayered, it is not. Its apparent stratiication is Stratiied squamous
an illusion that results because the nuclei of cells of differ- Nonkeratinized
ent heights occur at different levels. All of its cells, though, Keratinized
are in contact with the basement membrane and not Stratiied cuboidal
layered on top of each other. Stratiied columnar
Transitional

STRATIFIED EPITHELIAL TISSUE

Stratiied epithelia consist of two or more layers of cells,
with only the bottommost layer in contact with the base-
ment membrane. Each additional layer is added on top of
the previous one and does not contact the basement mem- Word Roots
brane. The total number of layers can vary from two to a
ROOT MEANING EXAMPLE SENTENCE
dozen or more. Classiication of stratiied epithelia depends
upon the shape of the cells in the outermost (surface) layer Column A pillar A columnar cell is tall and slender like a
pillar.
of the epithelium. If the outermost layer is squamous, the
epithelium is stratified squamous; if it is cuboidal, the Epi Upon or Epithelial tissue covers surfaces and lines
over cavities and is above deeper tissues. The
epithelium is stratified cuboidal; if it is columnar, the epi- term epithelium is a noun, and
thelium is stratified columnar. Stratiied epithelia can be epithelial is an adjective. For example,
found in such places as the mouth, esophagus, larynx, one can speak of a simple squamous
epidermis, vagina, and anal canal. Transitional epithelium epithelium, or simple squamous epithelial
tissue. Plural = epithelia.
is the term applied to a special category of stratiied epi-
thelium found only in the urinary system. The shape of its Pseudo False Pseudostratified epithelium appears to be
stratiied, but it is not.
cells changes with the amount of luid pressure applied
against them. For example, its surface cells have a domed Squam A scale Squamous cells, being thin and lat,
resemble scales that cover the body of a ish.
shape when the urinary bladder is empty but become lat-
tened as the bladder ills with urine. Strat A layer A stratified epithelium is layered.

12 CHAPTER 2
Simple Squamous Squamous cell Stratified Squamous,
nonkeratinized

Simple Cuboidal Cuboidal cell Bistratified Cuboidal

Simple Columnar Columnar cell Stratified Columnar

Pseudostratified Columnar

Transitional, Transitional,
unstretched stretched

Figure 2.1. Types of Epithelia.

EPITHELIUM 13
 9
9
5
3
2

9 6
8
1
8
Figure 2.5 250
5 KEY
8 1. Blood vessel, lumen 7. Lymphatic vessel, lumen
2 2. Collagenous iber 8. Smooth muscle cell, nucleus
3. Connective tissue 9. Squamous cell, nucleus
9 4. Cuboidal cell, nucleus 10. Tubule, lumen
5. Fibroblast, nucleus 11. Urinary space
1 6. Hepatocytes
9 9
9 Figure 2.2. Simple Squamous Epithelium of Blood Vessels and
Lymphatic Vessels, Submucosa of the Stomach, Pig. Blood
vessels and lymphatic vessels (and the heart) are lined by a simple
squamous epithelium that is speciically called an endothelium in
7 those locations. The cytoplasm of the squamous cells is sparse,
and generally only the nucleus is visible. Characteristically, the
nuclei appear lat in histologic section. If the squamous cells are
9 bunched up, such as those lining a vessel that is contracted, their
nuclei appear round.
9 Figure 2.3. Simple Squamous Epithelium, Kidney, Sheep. The
outer layer of Bowman’s capsule of a renal corpuscle is called
Figure 2.2 180 the parietal layer. It is formed by a single layer of squamous cells.
A magniied example of the parietal layer of Bowman’s capsule
is shown in Figure 2.4. Tubules lined by a simple cuboidal epi-
thelium are evident.
10
4 Figure 2.4. Simple Squamous Epithelium, Kidney, Sheep. The
4 lat nuclei of the squamous cells of the parietal layer of Bowman’s
capsule are visible, forming the outer lining of the urinary space
of the renal corpuscle.
9 11 Figure 2.5. Simple Squamous Epithelium, Mesothelium, Liver,
11 Cat. The surface of the liver is covered by a single layer of squa-
9
mous cells. Mesothelium is a speciic term for the epithelium of
4
2 10 the serous membranes of the body, namely the peritoneum,
pleura, and pericardium.

Helpful Hint
Figure 2.3 125
The nuclei of a squamous epithelium will not always appear
regularly spaced. When a layer of squamous cells of an organ
9 is sectioned, the slice may or may not pass through the nucleus
9 of each cell, resulting in uneven spacing.
4
11
9
Profile View, as seen in a
histologic section

10 Surface View, simple squamous epithelium

The line indicates the plane of the cut.
9
4
10
Figure 2.4 250
 KEY
1. Cuboidal cell, nucleus 3. Tubule, lumen
2. Squamous cell, nucleus

3 1
Figure 2.6. Simple Cuboidal Epithelium, Kidney, Cow
(Trichrome). The lining of the tubules shown here consists of a
single layer of cuboidal cells.

Figure 2.7. Simple Cuboidal and Simple Squamous Epithelium,

1 Medulla of Kidney, Sheep. The boundaries between the cuboidal
3 cells are not distinct in the tubules lined by simple cuboidal epi-
thelium in this image. To determine the shape of the cells, imagine
drawing lines between the nuclei of adjacent cells to approximate
their borders. Note the simple squamous epithelium that lines the
Figure 2.6 250 capillaries and the thin segments of the loop of Henle.

Helpful Hints
1 Organs are formed by various types of tissues, so you may
feel overwhelmed when you begin to look at a histologic
section of an organ to study epithelial tissue.
• Remember that epithelial tissue is found lining a cavity or
covering a surface, so begin to examine an organ at low
magniication and look for epithelium to be bordering the
white space of a cavity or surface.
• Epithelial tissue is formed by cells that are close together,
with little or no extracellular matrix. Look for closely
arranged basophilic nuclei of the epithelial cells to help
1 3 locate an epithelium.
• The shape of nuclei of the cells will help you name the type
2 of epithelium. The nuclei of squamous cells are typically lat
(sometimes round if the cells are bunched up, as in those
lining a contracted vessel) and the cytoplasm is usually not
1 visible. The nuclei are round in cuboidal cells and typically
elongated in columnar cells.
1
3
2

2
Figure 2.7 360

EPITHELIUM 15
 15 17 2
18 7
13
6 13
4 4
12
9
1
5
9
14 3
Figure 2.8 62.5 Figure 2.12 250
15 KEY
15
1. Basement membrane 10. Lymphocyte
2. Cilia 11. Mucus precursor region
3. Collagenous iber 12. Pseudostratiied columnar epithelium
4 4. Columnar cell, nucleus 13. Simple columnar epithelium
5. Gastric gland 14. Smooth muscle tissue
11 4 6. Gastric pit, lumen 15. Stomach, lumen
7. Goblet cell 16. Striated border
13 8. Jejunum, lumen 17. Trachea, lumen
9 9. Lamina propria 18. Similar area magniied in Figure 2.9

Figure 2.8. Simple Columnar Epithelium, Pyloric Stomach, Cat.

13 This low power view shows the location of the simple columnar
6 epithelium that lines the lumen of the stomach and invaginates to
form gastric pits. Figure 2.9 is a magniied view of the epithelium
of the stomach similar to the area outlined by the rectangle in this
Figure 2.9 250 image.
Figure 2.9. Simple Columnar Epithelium, Pyloric Stomach, Dog.
The columnar cells that line the lumen of the stomach and the
gastric pits have apical cup-shaped regions that contain mucus
precursor in living tissue. Note the elongated nuclei that are
typical of columnar cells.
Figure 2.10. Simple Columnar Epithelium, Jejunum, Dog. The
16 jejunum is lined by a simple columnar epithelium. A striated
10 4 7 8 border that consists of numerous microvilli is evident. Goblet cells
and migrating lymphocytes are present among the columnar cells.
13 In the upper part of this image, the simple columnar epithelium
was cut obliquely, making it appear atypical.
Figure 2.11. Ciliated Pseudostratiied Columnar Epithelium,
Trachea, Sheep. A pseudostratiied columnar epithelium is formed
9 by a single layer of cells of different sizes whose nuclei are at
different levels, so that the epitheliums appear stratiied. However,
it is a simple epithelium because all of the epithelial cells contact
Figure 2.10 250 the basement membrane, although this relationship is not visible.
Figure 2.12. Ciliated Pseudostratiied Columnar Epithelium,
17 Trachea, Cow. Goblet cells are evident in this example of a
2 pseudostratiied columnar epithelium.

4 For additional examples of simple epithelia, see

these igures:
12 Simple squamous: 8.2, 10.1 to 10.10 (showing endothe-
lium of blood vessels), 14.9, 14.10
Simple cuboidal: 9.9, 16.15, 17.10, 18.21, 18.30
Simple columnar: 13.83, 13.88, 13.115, 13.139, 18.23
Pseudostratiied columnar: 15.3, 15.4, 15.14, 15.15,
9 15.17, 15.40, 17.10, 18.85

Figure 2.11 250

 8
2
1

12

2 10
6
11 11
7
Figure 2.17 250
Figure 2.13 125
KEY
6 1. Bistratiied cuboidal epithelium 7. Lymphatic vessel, lumen
2. Collagenous iber 8. Pharynx, lumen
3. Dermis 9. Separation artifact
12 4. Esophagus, lumen 10. Smooth muscle tissue
5. Keratinized cells 11. Squamous cell, nucleus
6. Lamina propria 12. Stratiied squamous epithelium
4 11
Figure 2.13. Stratiied Squamous Epithelium, Nonkeratinized,
12 Epiglottis, Goat. Only cells of the basal layer of a stratiied epi-
11 thelium contact the basement membrane. In a nonkeratinized
epithelium, the nuclei of the cells are visible throughout the epi-
thelium, even those of the most supericial cells. Note the simple
squamous epithelium that lines the lymphatic vessel in the underly-
6 ing connective tissue.

Figure 2.14 125 Figure 2.14. Stratiied Squamous Epithelium, Nonkeratinized,

Esophagus, Cat. A nonkeratinized stratiied squamous epithelium
5 is visible lining the collapsed lumen of the esophagus.

Figure 2.15. Stratiied Squamous Epithelium, Keratinized, Skin

of Scrotum, Sheep. The supericial layer of the keratinized strati-
ied squamous epithelium is formed by dead, keratinized cells that
appear as thin strands, without visible nuclei, separated from one
3 another. Compare the keratinized cells here with the supericial
cells in the nonkeratinized epithelium in Figures 2.13 and 2.14.
12
Figure 2.16. Stratiied Squamous Epithelium, Keratinized,
5 Wattle, Pig. The wattle is covered by a keratinized stratiied
squamous epithelium.

Figure 2.17. Bistratiied Cuboidal Epithelium, Esophagus, Dog.

This duct of an esophageal gland is lined by a bistratiied cuboi-
Figure 2.15 125 dal epithelium.

Helpful Hints:
5
9
• The cells of a stratiied epithelium will be different shapes
at different levels. The speciic type of stratiied epithelium
12 is named according to the shape of the most supericial
(apical or surface) cells. For example, the surface cells of
a stratiied squamous epithelium are lat.
• The basal layers of a stratiied squamous epithelium look
darker (especially noticeable at lower magniication)
because the deeper cells are smaller with less cytoplasm
and, therefore, their dark basophilic nuclei are closely
3 packed.

Figure 2.16 250

EPITHELIUM 17
 11
7
2
5 9
1 4

Figure 2.18 250 Figure 2.22 125

10 KEY
1. Bistratiied columnar epithelium 7. Squamous cell, nucleus
4 2. Capillary, lumen 8. Stratiied columnar epithelium
3. Collagenous ibers 9. Transitional epithelium
4. Columnar cell, nucleus 10. Urethra, lumen
5. Duct, lumen 11. Urinary bladder, lumen
6. Smooth muscle tissue
8
Figure 2.18. Bistratiied Columnar Epithelium, Duct of Carpal
Gland, Pig. This duct is lined by a bistratiied columnar epithe-
lium. Note the columnar-shaped apical cells (the cells bordering
the lumen) with elongated nuclei. A simple squamous epithelium
3 lines the longitudinal section of a capillary that is visible in this
ield.
Figure 2.19 250 Figure 2.19. Stratiied Columnar Epithelium, Urethra, Goat. This
portion of the urethra is lined by a stratiied columnar epithelium.
6 The surface cells of this stratiied epithelium are columnar.
3
Figure 2.20. Transitional Epithelium, Unstretched, Urinary
Bladder, Pig. A transitional epithelium lines the lumen of the
bladder as well as other parts of the urinary system. Surface cells
of the transitional epithelium are either balloon-shaped or broadly
11 cuboidal when not under tension.
9 Figure 2.21. Transitional Epithelium, Unstretched, Urinary
Bladder, Cat. Note the large rounded surface cells in this
unstretched transitional epithelium.

Figure 2.22. Transitional Epithelium, Stretched, Urinary Bladder,

Cat. Surface cells of this epithelium are lattened and elongated
when the bladder is stretched with urine.
Figure 2.20 125

For additional examples of stratiied epithelia,

11 see these igures:
Bistratiied cuboidal: 12.58
Bistratiied columnar: 12.58
Stratiied squamous, nonkeratinized: 15.9, 15.47,
19.15
9 Stratiied squamous, keratinized: 12.3, 12.14,
12.16, 12.24
Transitional: 14.20, 14.24, 14.26

Figure 2.21 125

18 CHAPTER 2
 3

CONNECTIVE TISSUE PROPER

AND EMBRYONIC
CONNECTIVE TISSUE

O
f the four main types of tissues (epithelial, connective, muscle, and nervous),
connective tissue is the most abundant and includes numerous varieties.
Embryonic connective tissue, connective tissue proper, and the special kinds of
connective tissue are distinguished from one another by the types and numbers of cells
present and the nature of the extracellular matrix. Certain cells, such as ibroblasts, are
common in connective tissue proper and may also be present in some of the special types
of connective tissue. Other cells are unique to one kind, such as osteocytes in bone,
chondrocytes in cartilage, and erythrocytes in blood. The matrix is semiluid in connec-
tive tissue proper; it is gel-like in cartilage, hard in bone, and luid in blood.
Embryonic connective tissue and connective tissue proper are presented in this
chapter. The special types of connective tissue, cartilage, bone, blood, and bone marrow,
will be covered in subsequent chapters.

COMPONENTS OF CONNECTIVE TISSUE PROPER AND

EMBRYONIC CONNECTIVE TISSUE
Connective tissue binds together and supports other tissues. It is a composite of various
cells and ibers in an amorphous ground substance. The latter two components comprise
the extracellular matrix, which typically predominates over the cellular elements.

Extracellular Matrix
Ground Substance
The ground substance, composed largely of glycoproteins and glycosaminoglycans,
forms a well-hydrated gel that ills the spaces between cells, ibers, and vessels of

19
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 "Sticking together," was Master Henly's concise and descriptive answer.

"Glad to hear it," said Willard, repressing a strong inclination to laugh.

"Is Mrs. Tom within?"

"She was when I left the house," said Carl, who seemed determined not
to commit himself.

"Any one with her?" again inquired the young gentleman, looking as
indifferent as possible.

"No, nobody," was the unexpected answer.

"What!" exclaimed Willard, surprised. "I thought I saw a young lady

enter a moment ago!"

"Oh, Christie—she's nobody," said the gallant Mr. Henly.

"Christie—Mrs. Tom's niece—I thought she was away!" exclaimed

Willard.

"So she was, but I went for her this morning; couldn't be bothered doing
her work and my own both any longer," said Carl.

"I suppose I may go in?" said Willard, feeling a sudden thrill of pleasure
at the knowledge that this radiant girl was an inhabitant of the island.

"Yes, I suppose you may, if you like," said Carl, in a tone of the utmost
unconcern.

Thus kindly permitted, Willard advanced and rapped at the door. It was
opened by Mrs. Tom, whose surprise was only equaled by her delight at
being honored by this unexpected visit.

Near the window that overlooked the lodge, stood the golden-haired
vision of the beach. She turned round with a quick, shy glance, and blushed
most enchantingly beneath the deep, dark eyes of the stranger.
 "My niece, Christie, Mr. Drummond," said Mrs. Tom, directing his
attention to her with a wave of her hand. "She got back this mornin'. I allers
find it powerful lonesome here without Christie."

"I have no doubt of it," said Mr. Drummond, seating himself. "But I
have had the pleasure of seeing Miss Christie before."

"Where?" asked Christie, opening her blue eyes in wonder.

"Down on the beach, a few moments ago."

"Oh, yes."

And again Christie blushed vividly, as she recollected how she had been
caught singing.

"Where's Miss Sibyl and Master Guy?" inquired Mrs. Tom.

"Miss Sibyl has gone to N—— with the clergyman's family, and will
not return for a week; and Captain Campbell has gone to Westbrook, where
his vessel is undergoing repairs. So I am left all alone, and came to pay my
respects to you."

"Then you'll stay and spend the evenin'?" said Mrs. Tom, smiling
complacently.

Mr. Drummond professed his willingness; and the little widow,

delighted at the condescension, set about preparing tea instantly, assisted by
Christie, whose wild, shy glances were bent on his face whenever she
fancied herself unobserved. Half pleased, half afraid of him at first, she was
reserved and timid; but as this wore off, he drew her into conversation, and,
to his surprise, found her intelligent and well-educated. This Mrs. Tom
accounted for, by saying she had gone to school for the last five years at
Westbrook, residing there with the friend she had now been visiting.

The evening passed away with the rapidity of magic. Christie, after
much solicitation, consented to sing for him; and if anything was needed to
fairly enchant him, that sweet, clear voice would have done it. Then, too,
Carl added to the general hilarity, by drawing out a rusty Jew's-harp, and
playing a favorite tune of his own composition. Not once during the
evening did he think of Sibyl; her dark, resplendant face, and wild fierce,
black eyes, were forgotten for the golden locks and sweet, fair face of blue-
eyed Christie—this dainty island Peri.

The hour for leaving came all too soon. As he reluctantly rose to go, he
pressed the hand Christie extended, to his lips, with such passionate ardor
that the blood flushed to her very temples, but not with pleasure. Ere he left,
Mrs. Tom cordially invited him to visit her house while he remained on the
island—an invitation he was not loth in accepting.

Christie stood at the window, watching his tall, elegant form as he

walked toward the castle in the bright, clear moonlight.

"I like him, Cousin Christie; don't you?" said Carl, when he had gone.

But Cousin Christie turned away without reply, longing to lay her
burning cheek on the pillow, and muse over the new and delicious joy that
was thrilling her whole heart, and in her slumber to lie dreaming "Love's
young dream."

And Willard Drummond, forgetting his vows, forgetting Sibyl,

forgetting honor, forgetting all but this lovely island-maiden, sought his
couch with but one name on his heart and lips—

"Christie, Christie!"

CHAPTER VII.

THE HEART'S STRUGGLE.

 "She loves, but knows not whom she loves,
Nor what his race, nor whence he came;
Like one who meets in Indian groves
Some beauteous bird without a name,
Brought by the last ambrosial breeze
To show his plumage for a day
To wondering eyes, then wing away."—LALLA ROOKH.

Pale, feverish, and unrefreshed, after a night of restless dreams, Willard

Drummond arose from a vision of Christie to hail a new day.

Passion and principle were at war already. Bound by every pledge of

love, by every vow, to Sibyl, his whole soul was steeped in this new, all-
absorbing passion that had taken possession of him. He had fancied he
loved her, until he beheld radiant, dazzling, bewildering Christie and from
that moment he could have yielded heaven for her. Every feeling of his
inmost heart and soul was up, in arms. Every feeling of honor bade him fly
from this intoxicating siren, whose power he felt growing stronger each
moment over him; but the voice of passion cried: "Remain! love her, if you
will! What right has Sibyl to stand between you and the heaven of your
dreams?" And, like all who allow the struggle between right and wrong to
wage its warfare in their bosom, Willard Drummond was lost. For, with his
hot, fervid, Southern nature, worldly considerations, former vows, reason,
principle, justice, even honor, were swept away, like a wall of smoke,
before the fierce impetuosity of passion.

With a head throbbing, and pulse quick and feverish with the inward
conflict, Willard descended to breakfast.

Captain Campbell stood in the sitting-room, awaiting his coming. With

a courteous "good morning," he advanced to meet him; but started back in
surprise at beholding his extreme pallor.

"My dear fellow," he exclaimed, in a tone of solicitude, "you are ill—

very ill, I am afraid. What in the world is the matter?"
 "Nothing. I had bad dreams, and did not sleep well," said Drummond,
with a forced smile. "A cup of Aunt Moll's excellent coffee will set me all
right again."

"I don't know about that," said Captain Campbell, with his eyes fixed
anxiously on his face, "You are looking terribly feverish, and you were
complaining of a headache yesterday. I hope you are not going to be ill."

"I assure you it is nothing," said Willard, in a tone of slight impatience.

"You are needlessly alarming yourself. A bad night's rest is the cause of
all."

"Well, if it is not, I will have to call up Mrs. Tom to nurse you till Sibyl
comes. And, by the way, I regret exceedingly that I shall be obliged to leave
you solitary and alone for some days. Important business, that cannot be
postponed, demands my immediate attention."

Willard's heart suddenly bounded—he would scarcely have

acknowledged to himself the reason—at the words.

"It seems hardly courteous or hospitable to leave you thus," continued

the young captain; "but I know you will excuse me, my dear fellow, when I
tell you it cannot possibly be helped."

"Oh, certainly, certainly!" interrupted Drummond, cordially. "Go, by all

means. I will get along well enough in your absence. When do you leave?"

"Immediately after breakfast. It is an affair that cannot be postponed. In

fact, I will not have time even to go and see Sibyl; but, as you will probably
be there during the day, you can tell her. Perhaps you will come over to the
mainland with me?"

"No, I think not," said Willard, with affected carelessness. "I may go
during the course of the day."

"But how? I will take the boat."

"Oh, with Carl Henly. He has one, I believe."

 "Well, suit yourself. And now I'm off. Take care of yourself, my boy;
and au revoir!"

"Good-by!" said Willard, accompanying him to the door. "Aunt Moll

and I will keep bachelor's hall till you come back."

Captain Guy laughed, and hurried down to the beach. And when he was
gone, Willard arranged his slightly disordered dress and disheveled locks,
and sauntering out, almost mechanically took the road to the cottage.

It came in sight at last—this little, quaint, old house, that held all of
heaven to him now.

"Shall I enter—shall I thrust myself into temptation?" was his thought.

"If I look again on this fairy sylph I am lost!"

He thought of Sibyl, and her dark, bright, menacing eyes arose before
him, as if to warn him back.

"For your honor's sake—for your life's sake—for your soul's sake—go
not there!" said the threatening voice of conscience.

"And have I not a right to love whom I please? Why should I offer
violence to myself in leaving this bright enchantress, for that dark, wild
Amazon? Go, go, and be happy," said passion.

And, as if to overthrow his last good resolution, the image of Christie,

radiant, dazzling, and beautiful, as he had beheld her first, in the bright
flush of the fading sun light, arose before him, and once again passion
conquered.

He approached and entered the cottage.

Mrs. Tom sat near the window, spinning and singing to herself.
Willard's eyes wandered around in search of another; but bright Christie
was not to be seen.

The widow arose, smiling, to welcome her guest, and placed a chair for
him near herself. And still Willard's eyes went wandering round the room.
 "She will appear presently," he thought, not yet liking to inquire for her.

"What a venerable-looking affair your wheel is, Mrs. Tom," he said,

surveying it, with its hard polished wood and bright brass rings.

"Yes, it's as old as the hills," said Mrs. Tom, resuming her work; "and
has been in our family since the flood. I think I spun on that there wheel all
the yarn that makes the socks, mittens, and comforters for half the county
round; besides making sheets, blankets, and lots of other things for
ourselves," said Mrs. Tom, with conscious pride.

"You deserve a premium for industry, Mrs. Tom," said Willard.

"Well, you may be jokin' now, and I dare say you are; but it is true, for
all that. Many a true word is spoke in jest, you know," said Mrs. Tom, as her
wheel went merrily round. "There ain't many women in this place o' my age
and means, can do, or does do more work than me, though I say it as hadn't
oughter. I knit, and spin, and sew, wash, brew, bake, sow, and reap, and fifty
other things, too numerous to mention, besides. Carl, if I go out there I'll put
an end to your lazin', you idle, good-for-nothin' vagabone, you!" she added,
breaking off in sudden wrath, as she espied Carl, leaning on the spade with
which he should have been digging in the garden.

"You should make Carl do these things, Mrs. Tom," said Willard, still
impatiently watching the door and wondering why Christie did not come.

"Carl?" said Mrs. Tom, with a short laugh. "Lor'-a-massy! he ain't worth
his salt; that there's the laziest, most worthlessest young scape-goat ever any
living 'oman was plagued with. I hain't a minute's peace with him night nor
day; and if scolding was a mite of good, the Lord knows he might have
been a saint by this time, for he gets enough of it."

Willard laughed. And in such conversation the morning slipped away—

very rapidly to Mrs. Tom, but each moment an age to our impatient lover.
For Christie was absent still; and a strange reluctance, for which he could
not account, still prevented Willard from asking for her. It was an inward
sense of guilt that troubled him; for, feeling toward her as he did, he felt he
had no right even to mention her name.
 At last, in despair, he arose to go. Mrs. Tom relieved his mind by
saying:

"Christie will be disappointed at not seeing you," said the old lady,
following him out. "She went out berrying to the woods this morning, and
hain't got home yet."

Willard started at the information; and, inwardly cursing the folly that
had detained him so many hours talking to a foolish old woman, he darted
off, with a rapidity that quite amazed Mrs. Tom, in the direction of the pine
woods.

"What a confounded fool I have been," he exclaimed, savagely, "to stay

there listening to the way to make butter, and flannel, and 'yarb tea,' as if the
old beldame thought I was going to be somebody's housekeeper, or a female
doctress; and all the time this enchanting little blue-eyed witch was
wandering alone by herself. What an opportunity I have lost! and now I
suppose I may search for an hour and not find her."

He turned an abrupt angle in the winding path, and stifled a sudden

exclamation of surprise and delight; for there before him, reclining on the
grass, with half-veiled eyes, and soft, musing smile, sat the object of all his
thoughts, wishes, and desires.

He paused for a moment to contemplate the picture before him, for, if

Christie had seemed beautiful when he first beheld her, oh, doubly lovely
did she appear now in her attitude of unstudied grace.

Her dress was a loose, light muslin robe, fitting to perfection her
rounded waist and swelling bust. Her straw hat lay on the ground beside
her, and her golden, sunshiny hair floated, with all its wealth of rippling
ringlets, round her ivory throat. How dazzlingly fair looked that smooth,
snowy brow, contrasted with the full crimson lips and delicately flushed
cheeks—how enchanting the long curved lashes, falling over the deep-blue
eyes—how beautiful that faultless form, that soft, gentle, happy smile of
guileless girlhood.
 Willard Drummond's breath came and went, quick and short, as he
gazed, and his dark eyes filled with a subdued fire.

He advanced toward her. His shadow, falling on the grass at her feet,
was the first token she had of his coming. With a quick, startled cry, she
sprang to her feet in terror; but when she saw who it was that stood before
her, she stopped short, while the color flushed gloriously to her rounded
cheeks. Her first impression was: He has read my thoughts in my face, and
knows I was thinking of him.

"Have I disturbed you, bright Christie?" he asked, coming nearer.

"Oh, no!" she answered, blushingly. "I was only waiting to rest a little
while before going home."

"And dreaming, I perceive," said Willard. "May I ask, of what—of

whom?"

"I wasn't dreaming," said Christie, innocently. "I was wide awake all the
time."

"Day-dreaming, I mean," said Drummond, with a smile. "Do you know,

fairest Christie, I have been at your cottage all the morning, waiting to see
you?"

"To see me?" said Christie, with another quick, glad blush.

"And not finding you there, I have come in search of you," he

continued.

"And found me," she said, laughing. "If I had known you were coming I
should have staid at home."

"Perhaps it is better as it is, bright one; for I have found you alone. It is
very pleasant to have found so fair companion on this lonely isle."

"Yes, it is a lonely place," said Christie, musingly; "and yet I like it

better than Westbrook, or any other place I have ever been in. Only I would
like always to have a friend with me to talk to; and that, you know, I cannot
have here. Aunt Tom is always too busy to go out; and Carl don't care about
the trouble of talking, much less walking, so I always have to go alone."

"And if he would go, I fancy Master Carl is hardly the kind of

companion Miss Christie would select," said Willard.

"Not if I could find any better," said Christie, with a laugh; "but I have
grown so accustomed to being alone now that I do not mind it at all, as I
used to."

"And so you are perfectly happy here, fairest Christie, reigning queen of
this fairy isle?"

"Ah, no! beautiful Miss Sibyl is queen of the isle. I am only her most
loyal subject," said Christie, gayly; "you ought to know that, having paid
her your allegiance."

"What if I should say that the subject is more lovely than the queen?"
said Willard, in a low voice, and in a tone that brought the hot blood
flushing to Christie's face.

"I should say you were laughing at me, as of course you would be.
Certainly no one would ever think of me while Miss Sibyl was near. Oh!
how I wish she would always stay here, and then I would have a
companion."

"Ah, bright one! if I were in her place, what would I not surrender for
such a privilege!"

"Would you?" said Christie, looking at him in unfeigned surprise; "then

why not stay? I am sure I should be glad to have you here always."

Her innocent words, her enticing beauty, her child-like candor, were a
strong temptation. For one moment he was about to fall before her, to clasp
her in his arms, to hold her there forever, while he breathed forth his mad,
passionate love, and told her nothing on earth should ever part them now.
But again rose before him the dark, warning face of Sibyl to allay the fever
in his blood. It seemed to him he could see her black, fierce eyes gleaming
on them through the trees—he could almost hear her voice shouting
"Traitor!"

All unconscious of the struggle raging in his breast, Christie stood

leaning against a tree, her curved crimson lips half parted—her blue eyes
fixed on a cloud drifting slowly over the sky, little dreaming of the far
darker clouds gathering rapidly, now, over the horizon of her life.

And still in Willard's heart went on the struggle. He dared not look at
her as she stood before him—-bright, radiant, bewildering—lest the last
lingering remains of fidelity and honor should be swept away by the fierce
impetuosity of passion in his unstable heart.

But his good angel was in the ascendant still, for at that moment the
voice of Carl was heard calling loudly;

"Christie! Christie!"

"Here, Carl! Here I am," she answered; and in another instant honest
Carl stood before them.

"Aunt Tom sent me looking for you," said the young gentleman, rather
sulkily; "and I've been tramping through the woods this half-hour, while
you were taking it easy here," said Carl, wiping the perspiration from his
brow.

"It was all my fault, my good Carl," said Willard, as Christie hastily
snatched up her hat and basket and fled, having a just terror of Mrs. Tom's
sharp tongue. "Make my excuses to your good aunt, and here is something
for yourself."

Carl's dull face brightened wonderfully as Willard drew a gold piece

from his pocket and pressed it into his hand, and then turned his steps
slowly in the direction of Campbell Castle, thinking all earthly happiness
lay centered in the opposite direction.

Mrs. Tom's reproaches fell unheeded, for the first time, on Christie's ear
that day. She heard not a word of the long lecture delivered with more than
the good widow's usual eloquence, for she was thinking of another voice,
whose lowest tone had power already to thrill to the innermost recesses of
her heart. She loved without knowing it, without wishing to define the new,
delicious feeling filling her breast, only conscious she had never been so
happy before in her life, and longing for the time when she should see him
again. Ah, well had it been for her had they never met more.

CHAPTER VIII.

THE TRIUMPH OF PASSION.

"All other passions have their hour of thinking,
And hear the voice of reason. This alone
Sweeps the soul in tempests."

"Well," said Aunt Moll, to her son and heir, Lem, as he entered the long,
high kitchen of Campbell's Lodge, "I would like ter know what dat ar
Master Drummin's up ter? I doesn't understan' dese yer new-fangled young
men 'tall. Fust he comes a courtin' of our Miss Sibyl, and jes' as soon as her
back's turned he goes rite off an' takes up wid dat ar Miss Chrissy."

"'Tain't no business your'n, ole woman," said Lem, gruffly. "I 'spec's as
how Marse Drummin' knows what he's about."

"Yes, honey; but 'pears to me I ought to tell Miss Sibyl 'bout it. Ef he is
her beau, he oughtn't to be takin' up wid dat ar Miss Chrissy.'

"Better let Miss Sibyl look arter her own beau," replied her dutiful son.
"How does ye know he's a courtin' Miss Chrissy?"
 "'Cause I seed dem, chile—yes I did—las' night, down on de shore. De
moon was shinin' jes' as bright as a new pin, an' I took dat ar litter o' kittens
down to de shore to drown 'em, when I seed Marse Drummin' a walkin'
along wid Miss Chrissy, and he had his head stooped down jes' so,"—and
Aunt Moll ducked her woolly head to illustrate it—"an' was whisperin' soft
stuff, jes' as folks do when dey're in love."

"Well, what den?" asked Lem, growing interested.

"Well dey come up ah' seed me, in course, an', Lor', Lor'! I jes' wish you
seed de look Marse Drummin' give me. 'Peared as ef he'd a liked to knocked
my ole head off. But I warn't afeared, 'deed I warn't, chile; so I jes' stood
still an' drapped a courtesy, an' Miss Chrissy, she got red rite up to de roots
ob her hair. 'Good-evenin', marse and young miss,' sez I; 'don't be skeered; I
only wants to drown dese little kittens,' sez I, for I thought as I might be
perlite jest as well as not.

"'Oh, how does yer do, Aunt Moll?' sez Miss Chrissy, a laughin' an'
blushin'; 'how is Lem and yer rheumatiz, dese times?'

"'T'ank you, honey,' sez I, 'dey's pretty well, bof ob dem.'

"An' den Marse Drummin' he pulled her arm right troo his'n, and
marched her off wid him; an' den I pitched de kittens right in de water an'
come home."

"Well, dat ar warn't much," said the skeptical Lem. "Dey might be
walkin' on de beach, but that ain't by no means courtin'. Marse Drummin'
walk wid her, 'cause Miss Sibyl's gone, an' he ain't got nobody else to talk
to."

"'Cisely so, chile; but dat ain't all," says Aunt Moll. "Dis berry mornin',
as I was passin' troo de hall, de sittin'-room door was open, and I heered
voices a talkin' dere; so I listened and peeked in, an' dar was Marse
Drummin', rampin' up and down, a talking to hisself."

"Well, dat ain't nothin', eider," said the still contradictory Lem. "I've
hearn dat ar Carl talk to hisself when Miss Tom sent him out to work; an' he
ain't in love wid no one."

"But listen, honey, and don't you be puttin' me out so, 'cause 'tain't
'spectful—'deed it ain't," said Aunt Moll, getting slightly indignant. "As I
was sayin', I clapt my ear to de door, an' I heered him sayin' jes' as plain as
nothin' 'tall;

"'Oh, dischanting, onwildering Chrissy! ef I had nebber met you, I

might yet be happy!' Dar, what he say dat for ef he warn't in lub?"

This last was a settler. Lem felt that his mother had the best of the
argument, and unwilling to seem defeated, he went out, leaving the old lady
to enjoy her triumph.

Three days had passed since the departure of Sibyl, and certainly
Willard's conduct seemed to justify Aunt Moll's suspicions. Unable to break
the thrall which bound him, wishing, yet unable to fly from the spell of the
enchantress, he lingered still by her side. There were shame, dishonor, sin,
in remaining, but oh! there were death, misery, and desolation in going. All
worldly considerations, her unknown birth, her obscure connections, her
lowly rank, were swept away like walls of cobweb before the fierce torrent
of passion that overwhelmed, conquered every other feeling in its
impetuous tide.

And she loved him, this angel of beauty, this fairy princess of the isle;
he could see it in the quick flush of joy at his approach, the quick, burning
glances shot from her beautiful eyes, more quickly averted when they met
his—her low, impassioned tones, her bright, beautiful blushes. There was
joy, there was rapture in the thought; and yet, unless he forgot honor, vows,
all that should have been sacred, what did this love avail?

And so, like a tempest-tossed bark on a tempest-tossed sea, he strove

with passion and honor, love and remorse, right and wrong.

Once only, fearing lest her suspicions might be aroused by his absence,
he had visited Sibyl, whose rapturous greeting and confiding love made him
feel far more of a villain than ever. He looked forward with dread to the
period of her return, fearing for the discovery of his falsity; but, more than
all, fearing for the effects of her fierce wrath on Christie, knowing well
what must be the strength of Sibyl's passion when unchained.

And so, when Mrs. Brantwell proposed that Sibyl should remain with
her another week, instead of returning to the dreary isle, instead of feeling
irritated now, he backed the proposal, saying that perhaps it would be better
for her to do so, more especially during her brother's absence.

And Sibyl, in her deep love and woman's trust, suspecting nothing,
fearing nothing, consented, to the inward joy and sincere relief of her false
lover.

Resolving to visit her frequently, and so allay any suspicions that his
absence might give rise to, Willard Drummond returned to the island and to
—Christie, yielding himself without further effort to the witching spell of
her love.

Mrs. Tom suspected nothing of the contraband courting carried on under

her very eyes. It was the most natural thing in the world, she thought, that,
in the absence of Sibyl and her brother, the young man should spend whole
days with them, for it was not pleasant having no one to talk to but a couple
of negroes, as she very well knew. Then, it was not to be wondered at, that
he preferred talking and walking with Christie to any of the rest, for she was
"book-l'arned" like himself, which neither she nor Carl was. She did wonder
a little sometimes, and said as much to Christie, why he should stay on the
island at all, in the absence of the other.

"But, I suppose," was always her conclusion, "It's because it's Miss
Sibyl's home, and, for her sake, he stays there until she comes."

But Christie, though she only blushed and was silent, was of a different
opinion—one that she would scarcely own to her own heart. As to his being
in love with Christie, Mrs. Tom would have scouted the idea with scorn and
unbelief, had she heard it. Every circumstance was against such a
conclusion. He was rich, highly connected, and proud as a prince of the
blood; she was poor, unknown, and, compared with him, uneducated.
Besides, in the good widow's opinion, she was a child in feeling, as she
certainly was in years, scarcely knowing the meaning of the word love.
 Ah! she had been till he came; and his fervid, impassioned words, his
burning glances, his thrilling touch, had swept away the glamour of
childhood and simplicity, and revealed to her the passionate woman's heart
within her. His words, his looks, his tones, were all new revelations to the
artless, island maiden, changing her, as if by magic, from a child to a
woman. She revered him as the embodiment of all that was brave, generous,
and noble; worshipped him as a god, and loved him with all the affection of
her fresh, young heart, with all the ardor of a first, deep love.

As yet, she knew not whether that love was returned; for, unfaithful as
he was in thought to Sibyl, passion had not yet so totally conquered his
reason as to make him sin in words. He had never said, "Christie, I love
you;" but, ah, how often had his eyes said this, and much more; and how
long would this slight barrier stand before the fiery impetuosity of unstable
youth?

And so that day passed, and the next, and the next, and the next, and
with every passing hour the temptation grew stronger and harder to be
resisted. Matters must come to a crisis now, or never. Sibyl, in a day or two,
would be home, and this wild frenzy of his could be hidden no longer. If she
should come, as matters stood now, all would be lost.

And thus, torn between conflicting emotions, Willard sought Christie,

on the day before Sibyl was expected home, with the determination of
bringing this struggle to an end, then and there.

It was a glorious August afternoon. The island wore its bright dress of
green, and nestled in the blue shining river like an emerald set in sapphire.
The birds in the deep pine forest were filling the air with their melody, and
the odor of the wild roses came floating softly on the summer breeze.

But Willard Drummond was in no mood to admire the beauties of

Nature. The morning had been spent in pacing up and down his room,
hesitating, resolving, doubting, wishing, yet undecided still. For, when duty
and principle would appear for a moment victorious, the waving golden
hair, the beautiful blue eyes, the gentle, loving face of Christie would arise
before him, scattering all his good resolutions to the winds. And, mingled
with this, there was a sort of superstitious foreboding of evil to come. He
thought of his dream, and of the yellow-haired siren luring him on to
destruction; and of Sibyl, fiery daughter of a fiery race, fierce, vindictive,
and implacable in her wrongs.

"Oh, that I had never met this dark, passionate girl!" he murmured,
distractedly, "who now stands between me and the heaven of my dreams; or
would that I had seen this beautiful, enchanting Christie first! Oh, for that
angel as my wife! And but for those fatal vows once made to Sibyl, she
might be mine. I was mad, crazed, to mistake my fancy for that dark, wild-
eyed girl for love! And now, for that one mistake, am I to be wretched for
life? Shall I give up this beautiful, radiant creature, who loves me, for one I
care for no longer? No; the struggle is past. Christie shall be my bride, and I
will brave the worst that may follow!"

He set his teeth hard; and, as if fearing second reflection might make
him change his mind, he left the house and hurried out to meet Christie.

Down on the shore, under the shade of an overhanging willow, he knew

Christie had a favorite seat, where, on pleasant days, she used to take her
work. Here he was sure of finding her, and in this direction he bent his
steps.

She sat, sewing, under the shade of the drooping willow, singing softly
to herself, and looking like some sylvan goddess of a sylvan scene; or some
beautiful sea-nymph, just risen from her grotto of coral and chrystal.

Radiant and bewildering was the smile and blush with which she
welcomed him—a smile and blush that might have been found too strong
even for more potent principles than his.

He seated himself beside her, with a look of moody abstraction, all

unusual with him, watching her covertly from under his eyelashes, as she
bent smiling and happy over her work.

For a time, Christie chatted gayly on various commonplace matters; but,

at last, catching her tone from his, she, too, grew silent and thoughtful. She
bent lower over her work, wondering if she had offended him, and
involuntarily sighed.
 He heard it, and said:

"And wherefore that sigh, Christie! Are you unhappy?"

"No not unhappy; but troubled."

"And why should you be troubled, bright one? What can there be to
grieve one so fair?"

"I—I—feared I had offended you," she answered, timidly. "You appear

out of spirits."

"You offend me, gentle one—you who never offended any one in your
life? No, no; it is not that."

"Then you are unhappy," she said, shyly.

"Yes, I am miserable—wretched!" he cried, vehemently. "I wish to

Heaven I had never been born!"

"Oh, Mr. Drummond! what has happened!" she cried, laying her hand
on his, and looking up wistfully in his face.

Her touch, her tone, her look swept away every remaining trace of
fidelity. He forgot everything he should have remembered—his vows, his
honor, his truth—and saw nothing but the bright, radiant, bewildering
vision before him. In an instant he was on his knees at her feet, exclaiming,
with impassioned vehemence:

"Christie! Christie! my life, my dream, my hope, I love you. See, I am

at your feet, where my heart, my name, my fortune, long have been. With
my whole heart, and soul, and life, I love you with a love stronger than
death or the grave. All the devotion and hopes of my life I offer you, if you
can only say you love me."

He was pale and panting; his eyes were fierce and burning; his tones
low, thrilling, and passionate.
 Trembling, shrinking, blushing, yet, with a deep, intense, fervent joy
thrilling through her heart and being, Christie listened. The blood swept in
torrents to her face, neck, and bosom, which rose and fell with her rapid
breathing. She dare not look up to meet his ardent, burning, gaze.

"Christie, Christie! my love, my life! look up; speak—answer me—tell

me that you love me!"

Still no reply, only those downcast eyes, deepest blushes, and quick,
hurried breathing.

"Speak! speak! my beautiful love! only one word from those sweet lips;
but one touch of your dear hand to tell me I may live," he cried, growing
more wild and impassioned.

With a low, glad cry of intense joy, she buried her blushing face on his
shoulder.

"Thanks! my heart's thanks for this sweetest, loveliest Christie!" he

cried, with exultant joy, pressing her yielding form to his bounding heart.
"My life, with all its hopes, energies, and ambitions, shall be devoted to but
one purpose now—that of rewarding you for your priceless love."

"Oh, Mr. Drummond, your love is all the reward I ask!" she said, in the
deep, earnest voice of perfect trust.

"Not Mr. Drummond now, sweetest Christie. I am Willard to you, now

and forever. Let me always hear that name in music from your lips, and
earth has no higher boon in store for me."

"But oh! can you love me thus—me, a poor, little, nameless, uncultured
girl, while you are rich, distinguished, and highly connected. Oh, Willard,
will you not, some day, repent this choice—you, who might win the highest
and fairest in the land?"

"Repent! never—never! Perish my heart, if it ever admit of any love but

thine; palsied be my arm, if it ever encircle any form but this; accursed be
my lips, if they ever perjure the words I have spoken now; lost forever be
my soul, when it is false to thee!" he cried, with passionate vehemence.

"Oh, Willard! dearest, hush! I do not doubt you—Heaven forbid. I

should die, if I thought you could be false to me."

"Speak not of death; it is not for such as you, bright, beautiful Christie.
And now, only one thing is wanting, to make me the happiest of men."

She lifted her radiant face with a look of earnest inquiry.

"Christie, one little word from you, and ere the sun rises on a new day,
my joy will be complete—my cup of earthly happiness will be filled to the
brim."

Still the same earnest, anxious gaze.

"Dearest love, you will not refuse? It will be but a small matter to you,
and will make me supremely blessed."

"And that?" she inquired, wonderingly.

"Brightest Christie, be my bride—my wife!" he cried, folding her closer

in his arms, and speaking in a thrilling whisper.

Again the eloquent blood swept over her stainless neck and bosom, but
she did not reply.

"You will not refuse me, my own Christie, this last, greatest favor?
Comply now—to-day; for if the present opportunity passes, it may never
occur again."

"But how—how can we be wedded here?" she said, shyly, lifting her
eyes to his impassioned ones, and dropping them in brightest blushes.

"Christie, yonder lies a boat; it is three hours to sunset; long before that
time we can reach Westport; there we can find a clergyman, and there you
can become my own for life!"