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PREFACE
TO THE THIRD EDITION
I
t is our hope that students and research- by Aunt Lila and Uncle Jack McKean and
ers will continue to ind this atlas a Lola and Chip Wood. Thanks to Jessie
useful resource. In this edition, we have Bacha for proofreading our work, and to
expanded many of the beginning chapters Tristan Bacha for keeping busy while we
that introduce the basic types of tissues to worked! We greatly appreciate the helpful
provide the user with a stronger founda- suggestions from Professor Nancy
tion in histology. The glossary has been Gartland and the students at the University
expanded, and other extras have been Of Pennsylvania School of Veterinary
included that we hope will be valuable. Medicine, and from reviewers and users
Once again, many thanks to all of those of the previous edition. Finally, thanks to
who have made the irst and second Nancy Turner, Erica Judisch, Tracy
edition of our atlas possible! We were able Petersen, Erin Magnani, and all of the
to prepare our page layouts for the third other people at Wiley-Blackwell for their
edition thanks to the scanner given to us role in the production of this edition.
vii
PREFACE
TO THE SECOND EDITION
W
e wish to thank those who have hundred others. Four of the original black
used the irst edition for their and white line drawings have also been
suggestions. We believe the incor- redrawn. Also, a glossary of nearly 750
poration of many of these recommenda- words has been added.
tions will make this edition even more The style, format, and purpose of this
helpful to the user. edition remain essentially unchanged
To this end, we have updated the mate- from the irst edition. We continue to view
rial for the second edition by scanning all the atlas as a useful, benchside reference
of the original kodachromes and relabel- for those interested in understanding and
ing the art. We have added thirteen new interpreting histologic and cytologic
photographs and have enlarged over one preparations.
A
lthough we have written this atlas All photomicrographs and drawings are
primarily to fulill a need of the original. Some drawings were done free-
student of veterinary medicine, we hand, while others were made with the aid
believe that clinicians, private practitio- of a camera lucida. Light microscopy and
ners, and researchers will ind it a useful colored photomicrographs have been used
reference for normal tissues and organs. exclusively. We have chosen color rather
Currently, students rely heavily, if not than black and white because of its corre-
exclusively, on atlases of human histology spondence to stained preparations. With
for guidance in the laboratory. There are, the exception of the few histologic prepa-
of course, similarities between organs and rations loaned to us by generous donors or
tissues of domestic animals and those purchased from a dealer, slides were pre-
of humans. There are also differences, pared by the authors. Fresh organ samples
however, and these are rarely encountered were obtained from a slaughterhouse or
in atlases dealing speciically with human from animals that were euthanized for
histology. various reasons. With the exception of
Our aim has been to compare the his- smear preparations (blood, bone marrow,
tologic structure of organs in a variety of and vaginal), mesenteric spreads, ground
domestic animals. We have used represen- bone, and a single plastic section, slides
tative examples in instances where tissues were prepared using the parafin method.
and organs from different animals share a All slides were stained with hematoxylin
common structure. Wherever differences and eosin unless otherwise noted.
exist, we have tried to provide examples Magniications of photomicrographs are
that are characteristic of a particular total magniications (enlargement of pho-
group of animals. Our selection of animals tograph × objective × projector lens).
includes the dog, cat, horse, cow, sheep, Throughout the atlas, hollow structures,
goat, pig, and chicken because they are for example, blood vessels, kidney tubules,
most frequently referenced in veterinary and alveoli, are usually identiied by label-
school curricula. ing the lumen of the structure.
ix
ACKNOWLEDGMENTS
FROM THE FIRST EDITION
H
elp is often just around the corner. ment. We would also like to express
Dr. Henry Stempen, whose ofice appreciation to the following individuals
was down the hall from ours at from the University of Pennsylvania
Rutgers University in Camden, New Jersey, School of Veterinary Medicine: Mr.
stopped by one day and volunteered his Richard Aucamp and Mrs. Kathy Aucamp,
artistic talents. We’d like to thank him for who provided us with specimens, slides,
his excellent pen and ink drawings of advice, and assistance in a variety of other
various animal parts, which are somewhat ways; Dr. Mark Haskins for kindly making
removed from the fungi he usually draws. available fresh canine and feline material;
Our gratitude also to Ms. Kathleen Carr Dr. John Fyfe and Dr. Vicki Meyers-
for her secretarial services. Special thanks Wallen for supplying us with canine
are extended to Dr. Edward Zambraski, vaginal smears; Dr. and Mrs. Loren Evans
Ms. Kathleen O’Hagan, and Ms. Gail and Dr. David McDevitt for lending us
Thomas of Cook College, Rutgers reference material; Dr. Peter Hand and
University, for making fresh porcine mate- Ms. Graziella Mann for providing mate-
rial available to us, and to Dr. Barry Jesse rial on the nervous system; and Dr. Helen
and Dr. James Harner for supplying us Acland, Dr. Linda Bachin, Mr. James
with sheep parts. Bruce, Dr. Sherrill Davison, Ms. Dawn
Without the unqualiied use of the facili- Dowling, Dr. Robert Dyer, Dr. Robert
ties and equipment of the Biology Eckroade, Dr. George Farnbach, Dr.
Department of Rutgers, our tissue process- David Freeman, Dr. Wendy Freeman, Dr.
ing and photomicrography could not have Alan Kelly, Mr. Joseph McGrane, and
been accomplished. Our special thanks to Dr. Mary Sommer for their time and con-
the department for this courtesy. sideration in helping us to obtain tissue
This book would never have had a specimens.
beginning were it not for the generosity of We are grateful to Dr. Carol Jacobson
Dr. Leon Weiss, Department of Animal and the Department of Anatomy of the
Biology, University of Pennsylvania School Iowa State University College of Veterinary
of Veterinary Medicine, who invited us to Medicine for providing valuable slide
teach in the veterinary histology labora- preparations and text material.
tory and kindly allowed us access to the Our gratitude is also extended to Hill’s
slide collection and facilities of the depart- Pet Products, Topeka, Kansas, and
x
Pitman-Moore, Inc., Washington Cross- Company Inc., Cherry Hill, New Jersey,
ing, New Jersey, for their generous inan- for their courteous service and helpful
cial assistance. advice.
Many thanks also to: Dr. Caroline We are indebted to Mr. William J.
Czarnecki of the University of Minnesota, Bacha, Sr., for building a super light box
College of Veterinary Medicine, for pro- for us, and to Mr. Thomas H. Wood, Jr.,
viding copies of her informative labora- for providing black and white prints of
tory guide; Dr. Deborah Ganster, Dr. our photomicrographs, which saved us
James Lawhead, Dr. Virginia Pierce, Dr. countless hours of drudgery in the dark-
Maria Salvaggio, Dr. Barbara Strock, and room. Thanks to Barbara Frasco, Esq.,
Dr. Cindi Ward for assisting us in obtain- for her helpful advice. Our hats are off
ing tissue samples; Mr. Jeff Bringhurst, also to Snuff, Chew, Chapter Seat, Angel,
Bringhurst Brothers, Tansboro, New Clyde, and all the other animals for their
Jersey, for allowing us access to fresh large participation.
animal material; the Longenecker We also wish to extend our gratitude to
Hatchery, Elizabethtown, Pennsylvania, all at Lippincott Williams & Wilkins
for providing chicken specimens; Ms. whose efforts helped bring this second
Susan Ulrich, Cornell University Press, for edition into being. We are especially grate-
lending us a dificult-to-obtain reference; ful to Carroll C. Cann and Jennifer D.
the helpful people at Optical Apparatus Weir for their professional advice, cour-
Company Inc., Ardmore, Pennsylvania, tesy, and assistance.
for supplies and for assistance with equip-
ment for the microscope; and Mr. Charles William J. Bacha, Jr.
Behl and Mr. James Durso of Webb and Linda M. Bacha
xiii
COL OR ATL AS OF
VETERINARY
HISTOLOGY
Third Edition
1
GENERAL PRINCIPLES OF
HISTOLOGY
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 ric and its appearance is affected by the direction of the
parafin and lattens the section, eliminating wrinkles. The cut.
lattened section is loated onto a slide, which is then The three-dimensional structure of organs and their
placed on a warming table. As the preparation dries, the components also must be considered when examining a
section adheres to the surface of the slide. histologic preparation. Cells are three-dimensional objects
Next, the parafin is removed with xylene or another differing in size and shape. For example, some are long
appropriate solvent and the specimen is rehydrated. It is and thin, some cuboidal, and others ovoid. They may have
then stained, dehydrated, cleared (made transparent) with a random or speciic arrangement within an organ. How
xylene, covered with a resinous mounting medium, and they appear depends on their shape as well as how they
topped with a cover-slip. were cut. Imagine how the spindle-shaped and tall colum-
Various stains are available to the histologist. nar cells shown in Figure 1.2A would look if sectioned in
Hematoxylin and eosin (H&E) is a frequently used com- various planes. Note that the nucleus may or may not be
bination of stains. Hematoxylin imparts a purple color to included in a particular cut through a cell.
substances, but must be linked to a metallic salt called a The histologist examines multicellular structures
mordant before it can function effectively. This combina- having a wide variety of shapes. Some are hollow, some
tion, called a lake, carries a positive charge and behaves as branch repeatedly, some open onto surfaces, etc. Figure
a basic (cationic) stain. The lake combines electrostatically 1.2, B and C, and Figure 1.3 show a variety of three-
with negatively charged radicals such as phosphate groups dimensional structures and how they would appear if cut
of nucleoproteins. Substances that become colored by a at different levels. Examine these carefully. They will help
basic stain are said to be basophilic. Methylene blue, tolu- you to understand situations you will encounter on actual
idine blue, and basic fuchsin are basic stains. Unlike hema- slides.
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 HELPFUL HINTS
acid stain. It imparts an orange or red color to acidophilic
substances. Other commonly used acid stains are orange Be sure that the lenses of your microscope are clean before
G, phloxine, and aniline blue. you begin examining slides. Use a piece of lens paper or a
In addition to the widely used H&E staining proce- soft, clean cloth such as an old (but clean) linen handker-
dure, numerous other stain combinations and techniques chief. If the lenses have been coated with oil or another
are available. Some are especially useful for identifying substance, remove it using lens tissue moistened sparingly
certain tissue elements. For example, trichrome procedures with a glass cleaner such as Windex. Slides also should be
such as Mallory’s and Masson’s speciically stain collage- cleaned using a soft, lint-free cloth or tissue moistened with
nous ibers within connective tissue. Orcein and Weigert’s glass cleaner.
resorcin fuchsin are stains used to color elastic ibers, pro- Every microscope should have a pointer in the ocular.
viding a means of distinguishing them from other ibrous This is usually supplied by the manufacturer, but can be
elements. Reticular ibers and nervous tissue components made from a short piece of hair. The latter is cemented into
such as neurons, myelin, and cells of the neuroglia can be place inside the ocular with a dab of quick-drying glue or
stained by procedures employing the use of silver. There nail polish. Without a pointer, it is not possible to accu-
are also special histochemical and immunohistochemical rately indicate an object in the microscope ield for another
procedures that make possible the localization of various observer.
carbohydrates, lipids, and proteins found in tissue. Lastly, Before beginning a session at the microscope, make
stains such as Wright’s and Giemsa’s (Romanovsky stains) sure that the ine-adjustment knob is near the middle of its
are available for differentiating the various cells found in range of rotation. If you do not, you may ind that the
blood and bone marrow. 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
INTERPRETING SECTIONS unaided eye before placing it on the stage of your micro-
scope. By doing so you will gain information about the
One must know the gross structure of an organ before a gross aspects of the specimen and be more likely to center
histologic section from it can be comprehended. It is also it properly over the light source. Centering is especially
helpful to know how the section was cut, that is, whether important for small specimens that might otherwise be
it was a cross section (x.s.), a longitudinal section (l.s.), or dificult to locate. Also, make sure that you put the slide
an oblique slice through the organ. Was the cut made on the stage with the cover glass uppermost. If the slide is
through the entire organ or only through a portion of it? upside down, you will not be able to focus on it with the
Frequently, prepared slides are labeled indicating the par- high-power lenses. Do not snicker. We have seen this
ticular orientation of the section. This is not important in happen often in the teaching laboratory!
an asymmetric organ such as the spleen or liver because It is always a good idea to start your observations
their appearance would be unaffected by the direction of using the lowest power objective available on your micro-
the cut. Conversely, the small intestine is radially symmet- scope. This is usually the 4x lens. The ield of view will be
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.
8 CHAPTER 1
decimal is 0.6 and you should read 42.6 as the vernier ARTIFACTS
value. Do the same for the other vernier (Y) and record
the numbers for both. In the future, if you want to return Folds, knife marks, stain precipitate, spaces (where none
to the same location, simply secure the slide to the mechan- belong), shrinkage, and air bubbles are examples of
ical stage and move the stage controls until the verniers are common imperfections seen in slide preparations. They
adjusted to the numbers you previously recorded. These were introduced during processing and are called artifacts.
manipulations will have returned the slide to its former Figures 1.5 through 1.9 are examples of such artifacts.
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 Troubleshooting a blurred or cloudy image:
sections prepared by the parafin method the average size • If an image is cloudy or blurry, the oculars and/or objective
of erythrocytes for each of the following animals is as lenses may need to be cleaned. To determine if an ocular
follows: needs to be cleaned, turn it as you look through the micro-
Goat: 2.4 µ diameter (smallest erythrocytes of the scope. A dirty mark or smear will rotate if the ocular is not
clean.
domestic mammals) • If the image is still blurred, clean the objective lenses.
Dog: 4.9 µ diameter (largest erythrocytes of the domes- • Make sure the slide is resting on the stage properly and is
tic mammals) right side up! Sometimes an image cannot be focused
Chicken: 9.4 µ long clearly because the slide is on the stage upside down!
Each average value is based on a total of 20 to 30 cells • Are you using the oil immersion lens without a drop of oil
that were measured from ive different slide preparations on the slide?
of tissues embedded in Paraplast X-TRA (Monoject • Be sure that the light source is not partially blocked by
something, such as the electric cord or a ilter holder below
Scientiic, Division of Sherwood Medical, St. Louis, MO the stage.
63103).
2 KEY
1. Dermis 4. Knife mark
2. Epidermis 5. Separation artifact
3. Fold 6. Stain precipitate
5
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 1.9. 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.
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.
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
12 CHAPTER 2
Simple Squamous Squamous cell Stratified Squamous,
nonkeratinized
Pseudostratified Columnar
Transitional, Transitional,
unstretched stretched
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
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.
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
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.
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.
EPITHELIUM 17
11
7
2
5 9
1 4
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.
18 CHAPTER 2
3
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.
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
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mucha paz y quietud acompañados sus largos años.