A Photographic Atlas of Histology

A Photographic Atlas
of Histology
Michael J. Leboffe, D.A.

San Diego City College
Morton Publishing Company

925 W. Kenyon Ave., Unit 12
Englewood, CO 80110
http://www.morton-pub.com
Book Team
Publisher: Doug Morton
Managing Editor: David Ferguson
Cover and Design: Bob Schram, Bookends Publication Design
Typography: Ash Street Typecrafters, Inc.
Illustrator: Peggy Firth
Graphic Artist: Michael Arnold
Copyright © 2003 by Morton Publishing Company
ISBN 0-89582-605-4
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All rights reserved. No part of this publication may be reproduced,

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Printed in the United States of America

Preface
The primary goal and overriding theme of A Photographic Histology is a challenging, but exciting discipline—
Atlas of Histology is practicality. And while I expect that it once you develop certain skills. Beginners often dread
will get used during home study and test review, I wrote spending hours over a microscope and then have difficulty
with the student sitting at the microscope with a box of “seeing anything.” With practice and experience come a
slides to examine in mind. My hope is that the images in sharpened eye and strategies for successful examination of
this book will assist that student in identifying what needs microscopic preparations, which make the process more re-
to be seen. Toward this end, I used commercially available warding. I have made some suggestions in Chapter 1 to
microscope slides to photograph, so the images represent quicken your arrival to this point. I encourage you to give
the quality and diversity of what a student is actually likely them a try.
to encounter in the laboratory; pathological specimens have By the way, if you begin to imagine
not been used. I also have minimized the inclusion of elec- the cells are looking back at you, you’ve
tron micrographs, because beginning histology students are been studying too hard. It’s time for a
not typically required (or even allowed) to use an electron break.
microscope. Finally, I wrote captions for the images as if I I wish you success on your upcom-
were showing projected images to a class, so they tend to be ing adventure.
lengthy and descriptive.
The textual material is intended to be a concise synop- Mike
sis of histological basics, not an exhaustive treatise on the San Diego, CA
discipline. Further, it is assumed the student has a basic January, 2003
background in anatomy, so gross anatomy is not covered.
Neither is physiology discussed beyond a general descrip-
tion of function. You may want to examine other references
that cover these topics in detail.
iii
A P H O T O G R A P H I C A T L A S O F H I S T O L O G Y
iv
Acknowledgments
Even producing a book of modest size involves a tremendous throughout the book; Peggy Firth, medical illustrator, and
amount of support, cooperation, and encouragement from Michael Arnold, graphic artist, produced original art work;
many people. I would like to take this opportunity to pub- Pamela Jandura, CBC of Leica Microsystems Inc.
licly acknowledge them. (www.leica-microsystems.com) provided the microtome
First, I want to express my gratitude and love to Karen, photograph (Figure 1-1) and led me to John Yorston, Prod-
my wife of 25 years. Being the wife of an author was not uct Support Manager and Jack Vermeulen, International
part of the original deal, was it? Nevertheless, you continue Marketing Manager, both of LEO Electron Microscopy,
to tolerate and love me through it all. You are the best. Ltd. (www.leo-usa.com) who provided Figures 1-4 and 1-6;
Next, thanks to our children who are both beneficiaries Doreen Cantelmo and Stephanie Garguilo of Olympus
and victims of the writing process. Nathan (sorry, no need America, Inc. supplied the photograph for Figure 1-2; Gary
for hand models in this one!), Alicia, and Eric I appreciate D. Wisehart photographed the bone cross section (Figure
how you have shared your dad with his career. I love you 5-13) and many years ago encouraged me to write in the
all and am proud of the people you are developing into. first place—without your courage and ambition as a model,
And yes, Alicia, I know I ended that last sentence with a I wouldn’t have undertaken this and other projects; and
preposition. Laura Donahue and Bridget McCullough, students who
My “day job” is Professor of Biology at San Diego City supplied the blood for the hematocrit in Figure 6-1—you’ll
College. Thanks to the other members of the Biology De- never know which one I used because I don’t remember!
partment for their understanding and help as I served two Thanks to all of you.
masters during this project. In alphabetical order, thanks to Finally, my gratitude goes to Morton Publishing and
Dianne Anderson, James Bartley, David Brady, Dr. Joyce the entire book production team you have assembled. This
Costello, Julie Haugsness-White, Dr. Khasha Khomejany, is my sixth project with Morton Publishing, and my experi-
Burton Pierce, Debra Reed, Brett Ruston, Dr. David Singer, ences have been consistently positive. I have worked with
Dr. Minou Djawdan Spradley, and Gary Wisehart. Your di- Doug Morton, President and Chrissy Morton DeMier,
rect and indirect contributions made this project go more Business Manager since 1995 and appreciate the confidence
smoothly and they, as you, are greatly appreciated. they showed in me when I proposed this project and the trust
The administration of San Diego City College contin- they demonstrated with basically an open-ended budget!
ues to be supportive by making campus facilities available (Can we sign a contract now?) New to me this time around
to me through the Civic Center Program. Thanks to Ter- was Biology Editor David Ferguson, a young, energetic and
rence Burgess, President, Ron Manzoni, Vice President of professional businessman. I appreciate your handling of the
Instruction, Dr. Marianne Tortorici, Dean of Science, Nurs- business matters and understanding that the creative
ing, Health and Athletics, and Carol Dexheimer, Business process is difficult to rush. Joanne Saliger and her crew—
Manager for their support and cooperation. Patricia Govro and Elaine McFarlane—at Ash Street Type-
It is safe to say that this project would not even have crafters, Inc. have done a marvelous job (again) of converting
begun without the assistance of Bob Nazar of Olympus my partial thoughts, chicken scratches and last minute
America, Inc. (www.olympus.com) and David McGhee of changes into an attractive product. Thanks also to Bob
ImagingPlanet (www.imagingplanet.com). Bob made a Schram of Bookends, Inc. for the cover and overall design.
state-of-the-art Olympus BX41TF photomicroscope with I’m proud to be associated with all of you and look for-
top of the line accessories available to me, while David pro- ward to future projects.
vided the software and MagnaFire™ camera needed to cap- With all this help, one might expect a perfect product,
ture the images digitally. Both also generously provided but experience has shown me that mistakes still slip
technical assistance when needed. You have my deepest through and I am ultimately responsible for them. If you
gratitude. have comments, suggestions or corrections, I encourage you
Others who contributed are: J. Jesus Macias and Sylvia to email me at mjpleb@netscape.net or contact me through
Macias (father and daughter, and both former students!) Morton Publishing, http://www.morton-pub.com.
of the University of California, San Diego Medical Center
generously supplied transmission electron micrographs Thank you.
Contents
Chapter 1 Introduction 1 Epithelial Terminology 23

What is Histology? 1 Simple Epithelia 24
Why Study Histology? 1 Stratified Epithelia 24
Histological Techniques— Other Epithelia 24
Specimen Preparation 1 Glandular Epithelia 24
Histological Techniques—Microscopy 2 Chapter 4 Fibrous Connective Tissues 35
Strategies for Studying Histology 2
Basic Characteristics of Connective Tissues 35
Chapter 2 The Cell 7 Functions of Fibrous Connective Tissues 35
Embryonic Connective Tissues—
Cell (Cytoplasmic, Plasma) Membrane 7 Mesenchyme and Mucous Tissue 35
Nucleus 7 Extracellular Matrix
Cytoskeleton: Thin Filaments, Intermediate of Adult Connective Tissues 35
Filaments, and Microtubules 8 Cells of Adult Connective Tissues 36
Ribosomes 8 Classification of Adult
Endoplasmic Reticulum 8 Fibrous Connective Tissues 36
Golgi Apparatus 8
Mitochondria 8 Chapter 5 Cartilage and Bone 45
Surface Modifications: Cilia, Introduction to Skeletal Tissues 45
Flagella, and Microvilli 9 Basic Characteristics of Cartilage 45
Cellular Junctions 9 Types of Cartilage 45
Cellular Inclusions 9 Basic Characteristics of Bone—
The Cell Cycle and Mitosis 9 Matrix and Cells 46
Membranes of Bone 46
Chapter 3 Epithelial Tissue 23 Types of Bone Tissue 46
Basic Characteristics of Epithelial Tissues 23 Bone Growth 46
Epithelial Membranes 23 Synovial Articulations 47
Functions of Epithelia 23 Muscular Attachments to Bone 47
v
vi
Chapter 6 Blood and Bone Marrow 61 Chapter 12 Cardiovascular System 133

Introduction to Blood 61 Introduction to the Cardiovascular System 133
Formed Elements of Blood 61 Basic Blood Vessel Structure 133
Composition of Bone Marrow 62 Types of Blood Vessels 133
Postnatal Development of Blood Cells 62 The Heart 134
Chapter 7 Muscle Tissue 71 Chapter 13 Lymphatic System 143
Introduction to Muscle Tissue 71 Introduction to the Lymphatic System 143
Characteristics of Skeletal Muscle 71 Cells of the Immune System 143
Characteristics of Cardiac Muscle 72 Organs of the Lymphatic System 144
Characteristics of Smooth Muscle 72
Chapter 14 Digestive System 155
Chapter 8 Nervous Tissue and Introduction to the Digestive System 155
Organs of the Nervous System 79 Organs of the Oral Cavity 155
Introduction to Nervous Tissue 79 Organs of the Digestive Tube 156
Cells of Nervous Tissue 80 Glands of the Digestive Tract 158
Other Neural Structures 81
Organs of the Central Nervous System 82 Chapter 15 Respiratory System 177
Introduction to the Respiratory System 177
Chapter 9 Special Sensory Organs 101 Nasal Cavity 177
Introduction to the Senses 101 Larynx 177
Olfactory Receptors 101 Trachea 178
Taste Buds 101 Bronchial Tree 178
The Eye 101 Alveoli 178
The Ear 103
Chapter 16 Urinary System 187
Chapter 10 Endocrine System 111 Introduction 187
Introduction to the Endocrine System 111 Kidneys 187
Pituitary Gland (Hypophysis) 111 Calyces and Renal Pelvis 189
Thyroid Gland 112 Ureters 189
Parathyroid Glands 112 Urinary Bladder 189
Adrenal Glands 112 Urethra 189
Pineal Gland 112
Pancreas 112 Chapter 17 Reproductive Systems 197
Testes 112 Introduction to the Reproductive Systems 197
Ovaries 112 Male Reproductive System 197
Female Reproductive System 199
Chapter 11 Integumentary System 121
Introduction to the Integument 121 Further Reading 215
Layers of the Integument 121
Appendages of the Skin 122 Index 216
Introduction
C H A P T E R
1
What is Histology? chemical treatments, embedding with a supporting medium,
Literally, histology is the study of tissues (histos4tissue; and slicing into thin sections. An overview of the specific
logos4treatise). Tissues are collections of cells that have a steps follows.
similar structure and function. In the adult, there are four ◗ Fixation The specimen is treated with a chemical fixa-
major tissues: tive that alters its chemical composition and slows its
deterioration. Chemical modification may include pro-
◗ Epithelial tissue (epithelium)
tein denaturation and cross-linking, and preservation of
◗ Connective tissue lipids and carbohydrates.
◗ Muscle tissue ◗ Dehydration Beginning with a 50–70% alcohol solu-
◗ Nervous tissue tion, the specimen is bathed in successively more concen-
In modern usage, the term histology has come to include all trated alcohol solutions (up to 100%) to remove water.
levels of microscopic anatomy—from tissues down to cells ◗ Clearing Treatment with xylene makes the specimen
and cell ultrastructure, as well as the ways in which the tis- transparent.
sues are combined to form organs and organ systems. ◗ Embedding The specimen is treated with a supporting
medium, such as paraffin. (Paraffin is insoluble in
Why Study Histology? water and alcohol but is soluble in xylene, thus making
dehydration and clearing steps necessary.) First, the
Histology is the bridge between structure and function. It is
specimen is infiltrated with liquid paraffin, and then it
at the microscopic level that we begin to see how an organ
is embedded in it as the paraffin is allowed to solidify.
actually performs its functions. It is virtually impossible to
Plastics are sometimes used for embedding because they
discuss organ function without first considering its micro-
allow thinner sections (,1 µm) to be made.
scopic structure.
◗ Sectioning The paraffin block containing the specimen
is mounted on an instrument called a microtome (Fig-
Histological Techniques—Specimen Preparation ure 1-1). The microtome has a blade and a mechanism
In some cases, specimens may be examined immediately that moves the block a preset distance (usually 5–10
after removal from the body with staining as the only form µm) after each cut. Thus, each time a cut is made, the
of preparation. A blood smear would be an example. How- paraffin block is moved a distance equal to the thick-
ever, in addition to staining, most specimens must be pre- ness of the next slice. In this fashion, many slices of uni-
pared prior to observation. Preparation involves various form thickness are made.
1
2
◗ Mounting Each slice is applied to a glass slide. Magnification is not the only important feature of a mi-
◗ Staining To provide contrast, specimens are colorized croscope. The ability to produce magnification with good
with various stains. The colored portion of some stain- resolution is what really matters. Basically, resolution is the
ing solutions is acidic. Acidic stains colorize basic (alka- ability of a lens to produce a clear image. Technically, it
line; positively-charged) cellular structures, which are is the ability of a lens to see two separate points as being
said to be acidophilic. Basic stains are used to colorize separate. The closer together the two points are when they
acidic cellular structures, which are said to be baso- can be seen as separate, the better the resolution of the mi-
philic. Hematoxylin and eosin stain (H&E) is used for croscope. The limit of resolution for light microscopes is
most routine preparations and consists of a basic and about 0.1 µm, which allows production of quality images
an acidic component. The basic stain hematoxylin and magnified up to about 1500x.
the acid stain eosin produce the purple and pink speci- The electron microscope (Figure 1-4) uses an electron
mens frequently encountered in a histology laboratory. beam to create an image. The limit of resolution is improved
Table 1-1 lists some common stains and the structures by a factor of 1000 (theoretically down to 0.1 nm, but
they highlight. Since most stains are aqueous solutions, more realistically down to 2 nm) over the light microscope
the paraffin must be removed and the section rehy- and the maximum magnification is about 150,000x. The
drated prior to stain application. Following staining, transmission electron microscope (TEM) produces a two-
the specimen is again dehydrated and a cover slip is dimensional image of an ultrathin section by capturing elec-
permanently mounted over it. trons that have passed through the specimen. The degree of
interaction between the electrons and the heavy metal stain
The whole process may take 48 hours to complete. For
affects the kinetic energy of the electrons, which are col-
specimens that need to be examined quickly (as with surgi-
lected by a fluorescent plate. The light of varying intensity
cal samples) the tissue may be sectioned using a freezing
produced is directly proportional to the electron’s kinetic
microtome. Freezing replaces the embedding process and
energy and is used to produce the image. The TEM is useful
makes dehydration and clearing unnecessary. Freezing
for studying a cell’s interior, its ultrastructure. A sample
microtomes are also used when lipids are to be examined in
transmission electron micrograph is shown in Figure 1-5.
tissue because they are removed when treated with solvents
A scanning electron microscope (SEM) is used to make a
such as xylene.
three-dimensional image of the specimen’s surface. In this
Preparation of specimens for electron microscopy is
technique, a beam of electrons is passed over the stained sur-
similar to preparation described above. The fixing solu-
face of the specimen. Some electrons are reflected (backscat-
tions, such as osmium tetroxide, glutaraldehyde and potas-
ter electrons), whereas other electrons (secondary electrons)
sium permanganate, act as electron-dense stains as well as
are emitted from the metallic stain. These electrons are cap-
serving the primary function of preserving the tissue. Tis-
tured and used to produce the three-dimensional image. A
sues are typically embedded in plastic prior to being cut in
sample scanning electron micrograph is shown in Figure 1-6.
sections 25 to 100 nm thick.
Histological Techniques—Microscopy Strategies for Studying Histology

The light microscope (Figure 1-2) is used to examine speci- I took my first histology class in 1975 as a graduate student.
mens by transillumination, that is, by shining light through My professor told us of a supplemental atlas (Di Fiore’s
them. The mechanical components include focus adjust- Atlas of Human Histology, then in its fourth edition) that
ments, a mechanical stage, and a light source. The optical was available as a “crutch” if we needed it. After one lab
components include a condenser, objective lenses, and ocu- period, I set my grad student pride aside and gladly pur-
lar lenses. The condenser concentrates the light and illumi- chased my crutch. At the time, I just thought it was me.
nates the specimen uniformly. Light from the specimen is However, teaching anatomy and histology for over 25 years
refracted (bent) by the objective lens to produce a magni- has convinced me that this discipline is just plain difficult for
fied image, which is then magnified again as it passes most students to master. Nevertheless, “difficult” does not
through the ocular lens and is transmitted to the retina of equate to “impossible” or “not worthwhile.” Here are some
the eye. A sample light micrograph is shown in Figure 1-3. tips that I hope will help you learn what you need to know.
The amount of magnification produced by each lens is ◗ Read the slide’s label. It will tell you what the specimen
marked on the lens. Total specimen magnification is the prod- is, how it was sectioned (or if it is a “whole mount”),
uct of the power of the objective lens and the ocular lens. and often what stain was used.
Since typical microscopes have 4x (low power), 10x (medium ◗ Realize that most slides have more on them than what
power), 40x (high dry power) and 100x (oil immersion) ob- the label says. One of the most difficult skills to acquire
jectives and 10x oculars, the total magnifications are 40x, in histology is to find the “correct” part of the slide to
100x, 400x, and 1000x, respectively. examine.
C H A P T E R O N E : I N T R O D U C T I O N
3
◗ Examine the slide without the microscope to get an related. When you see something of particular interest,
idea of what the section looks like. Often, you can see a center it and move to the higher powers.
lot without magnification. It also should help you to
“zero-in” on the area of the slide you want to examine ◗ Do not over-illuminate the specimen. Using too much
because you can position the slide on the stage with the light results in loss of detail in the image. Reduce the
relevant part centered. (Remember that the specimen’s light with the iris diaphragm to a point where you can
image will be upside down and backward from its ori- still see the specimen, but it is not too bright. You will
entation on the stage.) need to increase the light at each higher magnification to
◗ Begin microscopic examination at low power. Get a get the same degree of illumination since the field gets
feeling for what is on the slide and how the parts are successively darker as higher power objectives are used.
◗ TABLE 1-1 Commonly Used Histological Stains

STAIN COMMENT
AZAN Stains nuclei, nucleoli and RBCs red, muscle orange, and connective
tissue blue.
AZURE B DNA is stained blue-green, RNA is stained purple.
CRESYL VIOLET Typically a bacterial stain, it can be used to stain neuronal Nissl bodies
purple.
DAFANO SILVER METHOD Golgi material stains black.
FEULGEN STAIN DNA is stained reddish violet.
GIEMSA-WRIGHT STAIN Used for staining blood and bone marrow smears. Cytoplasm is stained
light blue; cytoplasmic granules are either pink, purple or unstained;
nuclei are dark purple.
HEMATOXYLIN AND EOSIN (H&E) H&E is a very commonly used stain. Hematoxylin is a basic stain that colors
acidic structures a purplish blue. These include the nucleus and regions with
abundant ribosomes or rough endoplasmic reticulum, in addition to cartilage
matrix. Eosin is an acidic stain and makes basic regions of the cell—that is,
most of it—pink. It also stains collagen fibers pink.
IRON HEMATOXYLIN This stains nuclei, chromosomes, mitochondria, centrioles, and muscle
striations black.
MASSON’S TRICHROME Results in three colors: basophilic structures are stained dark blue,
collagen is stained green or light blue, and cytoplasm, keratin, muscle
and erythrocytes are stained red.
OSMIUM Stains lipid a dark brown color.
PERIODIC ACID-SCHIFF (PAS) Colors complex carbohydrates red (magenta). Cells or cellular structures
that stain red are said to be PAS-positive. Glycogen, brush borders, and
the mucin in goblet cells are PAS-positive.
SILVER STAIN Used for staining delicate structures, such as reticular fibers and neuronal
processes.
TRYPAN BLUE Phagocytic cells are stained blue when they engulf this dye.
4
◗ This one may be the hardest. Don’t be satisfied with one, what you don't see are the dozens of slides, fields,
looking at a single example of each assigned specimen. and photographs that were discarded because of poor
Look at several, then look at some more—especially if quality.
they are from a different organism or manufacturer. It
takes examination of many examples before you will
start to see the common construction.
◗ Most specimens have been cut into thin slices and you
will be expected (eventually) to convert the essentially
two-dimensional image you see on the microscope to a
fully functional, three-dimensional object. This is equiv-
alent to asking a person who has never seen a loaf of
bread to imagine what the loaf looks like after examin-
ing a single slice. This skill will come with practice and
by looking at three-dimensional drawings of the speci-
men as you examine it on the slide. So many structures
are tubular, that it will also be useful to examine Figure
1-7 and see the many ways a tube appears depending
on how it is cut.
◗ Don’t overestimate slide quality. Photos in texts show 1-1
good examples, but not every slide of an organ will
MICROTOME A microtome is used to make thin sections of specimens
show everything you need to see, and neither will slides to be mounted on a microscope slide.
necessarily be of uniform quality. In books such as this (Photograph courtesy of Leica Microsystems Inc.)
Ocular lenses
Arm
Revolving nosepiece
Objective lenses
Stage clip On/off switch
Stage
Light intensity knob
Iris diaphragm
Coarse focus knob

Condenser
Fine focus knob

Lamp
Base
LIGHT MICROSCOPE The light microscope uses Mechanical stage

visible light and lenses to produce a magnified adjustment knobs
image of a specimen.
(Photograph courtesy of Olympus America Inc.) 1-2
C H A P T E R O N E : I N T R O D U C T I O N
5
LIGHT MICROGRAPH A photograph of a microscopic image is a

micrograph. This specimen from the skin was stained with
1-3 H&E and magnified X600 by a light microscope.
ELECTRON MICROSCOPE The

electron microscope produces an
image using either transmitted
electrons (TEM) or reflected
electrons (SEM). The image is
viewed on the monitor. Shown
here is a research grade scanning
electron microscope.
(Photograph courtesy of LEO Electron
1-4 Microscopy Ltd.)
6
1-6
SCANNING ELECTRON MICROGRAPH Like the TEM, the image
produced by the SEM has no color, but it is three dimensional.
This specimen is from mouse liver and was magnified X1800.
(Photograph courtesy of LEO Electron Microscopy Ltd.)
1-5
TRANSMISSION ELECTRON MICROGRAPH The TEM produces images of
sectioned specimens. Since light is not used, the image is not in color. This
cell was magnified X12,500.
(Photograph courtesy of UCSD Medical Center.)
Cross
section
Oblique
section
Longitudinal
section
SECTIONS OF A TUBE Study this

diagram to gain an understanding of
Cut sections in how the cut sections relate to the
various planes
three dimensional whole. This is one
of the toughest skills to learn as a
1-7 histologist.
The Cell
C H A P T E R
2
issues are made of cells, so it is appropriate to study Cell (Cytoplasmic, Plasma) Membrane
T cell structure and ultrastructure before embarking

on a study of the tissues that comprise organs.
Cells were first discovered and studied using the light
While a line in the region of the cell membrane is often vis-
ible with the light microscope (Figure 2-3), it wasn’t until
the electron microscope was used that membranes could be
microscope. For centuries, our understanding of cell struc- visualized in detail. The electron microscope also revealed
ture was based on the view provided by the light micro- the presence of numerous membrane bound organelles in
scope. However, due to the physical limitations posed by the cytoplasm.
using light to create the specimen’s image, a point was Membranes are a double layer of phospholipids ori-
reached where cell biologists had basically “seen it all.” Fig- ented with their hydrophobic fatty acid tails towards the in-
ure 2-1 illustrates two light micrographs of cells stained terior of the membrane and the hydrophilic phosphate
with different dyes. Notice that the cells appear to be little heads on each surface (Figure 2-4). Proteins are embedded
more than granular cytoplasm surrounded by a membrane in the phospholipid bilayer or are positioned on its surface.
with a nucleus inside. These proteins may be enzymes, transport proteins, recep-
With the development of the electron microscope in the tors, electron carriers or form channels that penetrate the
middle 20th Century and its ability to produce much higher membrane. Other lipids and carbohydrates are also found
magnification with greater resolution, cells could be visual- in membranes. Functionally, the cytoplasmic membrane is
ized with a new degree of clarity. Structural detail (cell ultra- the primary permeability barrier between the cell’s interior
structure) that was inaccessible before was now revealed and the external environment.
and opened the door to further study of cell structure and In electron micrographs, membranes appear as two
function. An artist’s rendition of a generalized animal cell dark lines separated by a lighter band in the center, with a
based on electron microscopy is shown in Figure 2-2. total thickness of approximately 7 nm (Figure 2-5). While
While the electron microscope is used for most research membranes throughout the cell have the same appearance,
applications, it is still the light microscope that is employed they differ in the particular phospholipids, proteins, and
in introductory histology courses and routine examination other components, depending on their functions.
of tissues in clinical settings. In this chapter, we briefly re-
view cell ultrastructure as understood from electron micro- Nucleus
graphs, but the main emphasis is on what can be seen with The nucleus (Figure 2-1) is the most prominent cellular
the light microscope. For complete structural and func- structure when viewed with the light microscope. It
tional details, the reader is referred to a standard reference contains the cell’s hereditary material—deoxyribonucleic
text on general biology, histology, or cytology. acid (DNA). Each DNA molecule (of which there are 46 in
7
8
a normal human cell) is composed of two long, thin Actin filaments are also involved in pinching the cytoplasm
polynucleotides wound in a double helix. Figure 2-6, which into two parts during cell division.
is stained to show DNA, gives no indication of the poly- Electron micrographs show microtubules to be hollow
meric nature of the molecule, but does illustrate that DNA protein cylinders. They form the interior of structures such
is localized in the nucleus. The genes of DNA contain infor- as cilia and flagella (see page 9), comprise the centrioles of
mation for production of cellular structures as well as the the centrosome (Figure 2-17), and are involved in move-
enzymes that catalyze metabolic reactions. ment of chromosomes during mitosis as the spindle appara-
In a dividing cell, each DNA molecule and its asso- tus (Figure 2-18).
ciated proteins is tightly coiled and becomes visible as a Intermediate filaments are involved in maintaining cell
chromosome (Figures 2-7 and 2-8). Chromosomes are an shape and are intermediate in size between thin filaments
extremely compact and efficient form in which to distribute and microtubules. In association with actin filaments and
genetic material to the daughter cells produced by division other proteins, intermediate filaments form the terminal
(see the section on mitosis at the end of this chapter). web in the cytoplasm near the surface of epithelial cells.
In a nondividing cell, the DNA is in a dispersed form The cytoskeleton visible in Figure 2-14 is made primarily of
called chromatin. Chromatin granules are often visible in intermediate filaments (neurofilaments).
light micrographs of the nucleus, especially at the periphery
(Figure 2-9). Electron micrographs are able to differentiate Ribosomes
between denser heterochromatin and less dense euchroma- Ribosomes are small, electron-dense structures composed
tin (Figure 2-10). The former is inactive, whereas the latter of ribosomal RNA (rRNA) and protein. In eukaryotes, the
is actively being transcribed into ribonucleic acid (RNA). In ribosome is made of a large subunit and a small subunit,
females, one entire X-chromosome is inactive and is some- distinguishable by their sedimentation coefficients (60S and
times visible in light micrographs as a “drumstick” at the 40S, respectively). Some ribosomes are attached to endo-
periphery of the nucleus of certain white blood cells (Figure plasmic reticulum (Figure 2-19), whereas others are free in
2-11). the cytoplasm (Figure 2-20). The former produce secreted
Also often visible in the nucleus are the nucleolus and proteins, whereas the latter are involved in synthesis of pro-
nucleoplasm (Figure 2-12). The nucleolus appears as a dark teins for internal use. Ribosomes are not visible individually
region and is involved in rRNA synthesis. The nucleoplasm with the light microscope, but because of their RNA com-
comprises the remaining contents of the nucleus. position, they are basophilic. Many protein-secreting cells
Surrounding the nucleus is the nuclear envelope, often have a dark purple or blue cytoplasm due to the abundance
visible as a line in light micrographs (Figure 2-12), but of ribosomes (Figure 2-21).
shown to be a double layer of membrane in electron micro-
Endoplasmic Reticulum
graphs (Figure 2-13). The outer nuclear membrane is con-
tinuous with the rough endoplasmic reticulum. Regions in The cytoplasm of eukaryotic cells is traversed by a network
the nuclear envelope, called nuclear pores, are formed of membranous tubules called endoplasmic reticulum.
where the two membranes of the envelope join. Simple dif- Smooth endoplasmic reticulum (SER) appears to be in-
fusion and ATP-dependent transport mechanisms move volved in steroid, cholesterol and triglyceride synthesis. The
materials between the nuclear and cytoplasmic compart- membranes of rough endoplasmic reticulum (RER) have ri-
ments of the cell through the pores. bosomes (Figure 2-19) involved in synthesis of proteins for
external use. The endoplasmic reticulum is continuous with
the outer nuclear membrane. The dark staining regions in
Cytoskeleton: Thin Filaments, Intermediate
the micrographs in Figure 2-21 actually show the regions of
Filaments, and Microtubules
RER.
The cytoskeleton is composed of protein filaments that are
responsible for movement and maintaining cell shape. It is Golgi Apparatus
difficult to view with the light microscope, but in some cells The Golgi apparatus (Figure 2-22) is made of a stack of
where the filaments are especially abundant, its presence curved, membranous sacs called cisternae. The Golgi is in-
can be demonstrated (Figure 2-14). volved in carbohydrate synthesis, and modifying, sorting,
Thin filaments (microfilaments) are composed of the and packaging proteins. The Golgi apparatus is visible with
protein actin. Actin filaments, along with the thicker an appropriate stain or occasionally as a pale-staining
myosin filaments, are present in skeletal and cardiac muscle region in the cytoplasm of some cells (Figure 2-23).
where they form the contractile apparatus. While the indi-
vidual filaments are not visible in light micrographs, their Mitochondria
highly organized arrangement is reflected in the banding Mitochondria (Figure 2-24) are double-membrane or-
pattern seen in skeletal muscle fibers (Figures 2-15 and 2-16). ganelles that are the primary site of ATP production in the
C H A P T E R T W O : T H E C E L L
9
cell. The space bounded by the inner membrane is the mito- been shown to be made of highly interdigitated cell mem-
chondrial matrix and possesses the enzymes of the Krebs branes and junctions resembling zonula adherans, desmo-
cycle. Embedded in the inner membrane are the compo- somes, and gap junctions.
nents of the mitochondrial electron transport system. Mito-
chondria are barely visible with the light microscope and Cellular Inclusions
appropriate stain. Some cells are specialized to store carbon and energy as
either lipid or glycogen. These materials can be demonstrated
Surface Modifications: Cilia, with special stains. Figure 2-33 shows cells storing lipid and
Flagella, and Microvilli glycogen.
Cilia and flagella are easily viewed with the light micro-
scope (Figures 2-25 and 2-26). Both are long, thin cellular The Cell Cycle and Mitosis
projections involved in motility by producing sweeping mo- The cell cycle marks the events of a cell between the time it
tions. Typically, cilia are more numerous and shorter than comes into existence until it divides into two new cells.
flagella (which are found singly in humans). When viewed These events are divided into two major parts: interphase
with the electron microscope, it was seen that both have the and mitosis/cytokinesis. Interphase (Figure 2-34a) com-
same basic construction. Both are surrounded by cell mem- prises the majority of the cell cycle and is recognized histo-
brane and contain nine pairs of microtubules (doublets) logically by a typically appearing nucleus. It is subdivided
around two single microtubules (singlets) (Figure 2-27). into three stages: G1 (first gap) phase, S phase, and G2 (sec-
This 912 arrangement of microtubules constitutes the axo- ond gap) phase. During G1, the cell grows and begins syn-
neme. At the base of each flagellum and cilium is a basal thesizing macromolecules necessary for DNA replication,
body composed of nine microtubule triplets with no central the hallmark event of the S phase. The G2 phase is devoted
microtubules (identical to centrioles) into which the axo- to synthesis of materials necessary for division.
neme inserts. Mitosis is the part of the cell cycle where replicated
Microvilli (Figure 2-28) are tiny projections that increase nuclear material is distributed to opposite ends of the cell
the surface area of cells involved in absorption. While not prior to division of the cytoplasm, cytokinesis. In this way,
individually visible with the light microscope, they do appear the two daughter cells receive complete and identical
as a striated or brush border on cells of the small intestine, amounts of nuclear DNA. Mitosis is divided into five
colon, and kidney tubules (Figure 2-29). Actin filaments in- stages.
side each microvillus anchor it to the terminal web. ◗ The first is prophase (Figure 2-34b), in which the chro-
Stereocilia are long microvilli found in the epididymis matin condenses into visible chromosomes, nucleoli
(Figure 2-30) and on the hair cells of the inner ear, where disappear, and the spindle apparatus (made of micro-
they are involved in producing a signal in response to tubules) begins to form. At this point, the chromo-
sound waves. somes each possess two sister chromatids attached at a
centromere.
Cellular Junctions ◗ During prometaphase, the nuclear envelope disinte-
The first evidence that adjacent cells are connected by spe- grates and the spindle fibers attach to each chromo-
cialized attachments was provided by light micrographs of some’s centromere.
intestinal epithelium, which often exhibit terminal bars ◗ During metaphase (Figure 2-34c), the chromosomes are
(Figure 2-31). The electron microscope has since revealed a positioned at the cell’s equator to form the metaphase
number of structurally different types of attachments found plate.
not only in epithelium, but also in cardiac and smooth
◗ In anaphase (Figures 2-34d and 2-34e), centromeres
muscle. These include the following:
split and sister chromatids move to opposite poles of
◗ occluding or tight junctions that bind adjacent epithe- the cell.
lial cells tightly and seal the intercellular spaces; also
◗ Mitosis ends with telophase (Figure 2-34f), in which the
known as zonula occludens
events of prophase are undone: chromosomes disperse
◗ adhering junctions which include zonula adherans, a as chromatin, the nucleoli reappear, the spindle appara-
continuous ring of attachment, and desmosomes which tus disintegrates, and the nuclear envelope reforms.
are spot attachments,
During late anaphase and telophase, cytokinesis begins
◗ communicating or gap junctions are made of tiny pores with the formation of a cleavage furrow (Figure 2-34g).
through which adjacent cells can transfer materials.
The cleavage furrow eventually pinches the cytoplasm into
In epithelia, these collectively form the junctional com- two separate compartments—the daughter cells—and the
plex. Intercalated discs (Figure 2-32) of cardiac muscle have division is complete.
10
The specimens used to illustrate the stages of mitosis in Some cells in mitotically active tissue might not present
Figures 2-34b through 2-34f were prepared exclusively for chromosome arrangements that fit the defined categories.
that purpose, but mitotically-dividing cells may be observed In some instances, it may be difficult to differentiate be-
in various tissues (Figure 2-35). However, the cell cycle is tween late anaphase and early telophase, for instance. This
not uniform for all body cells. Some cells continually divide, is to be expected, since mitosis is a continuous process and
entering G1 of interphase immediately following comple- we see the cells stopped in the middle of their activity. Fur-
tion of mitosis/cytokinesis. Skin cells and the intestinal epi- ther, the orientation of the division plane may complicate
thelium are examples. Others, such as liver cells, enter a G0 interpretation. Figure 2-34h shows a cell whose division
phase and divide only when disease or damage make it nec- plane is parallel with the page, so we are observing it from
essary. Terminally differentiated cells, such as neurons, have one end, not above. This makes placing it in a particular
become so specialized that they have lost their ability to stage very difficult.
divide.
CELLS AS VIEWED WITH THE LIGHT

MICROSCOPE (a) This light micro-
graph, stained with H&E, allows
* identification of cell membranes
(CM), nuclei (N), and nucleoli
(Nu), but the cytoplasm (C) just
appears as a grainy, amorphous
* material. Notice the cells that
CM *
apparently lack a nucleus (*). Their
nuclei were not in the plane of the
N section so they appear to be absent.
C
(X600) (b) A silver stain was used
to prepare this specimen from the
Nu spinal cord. In it, the neuron’s
nucleolus is the most prominent
feature with the nucleus staining
lighter than the cytoplasm. (X250)
2-1a
N Nu
2-1b
11
Smooth
Nuclear Nucleolus endoplasmic
envelope Golgi apparatus
reticulum
Pore Cilia
Chromatin
Microfilaments
Lysosome
Microvilli
Plasma
membrane
Pinocytotic
vesicle
Free
ribosomes
Ribosomes
Rough
endoplasmic Centriole Mitochondrion
reticulum
2-2
AN ARTIST’S RENDITION OF ANIMAL CELL ULTRASTRUCTURE This illustrates the general features found in animal cells, but not all would be seen in
a single cell.
CM
CM
CELL MEMBRANE AS SEEN

WITH THE LIGHT MICROSCOPE
Membranes (CM) are pretty
obvious between the cells of the
tubule running across the field,
but they are less distinct on many
of the other cells in the field.
2-3 (X530)
12
Integral protein
with channel
Glycolipid
Integral Cholesterol
glycoproteins
Hydrophobic
tails of
phospholipids Peripheral Hydrophilic
protein heads
Grated channel
2-4
AN ARTIST’S RENDITION OF A TYPICAL MEMBRANE Membranes are composed of a phospholipid bilayer associated with various proteins and
carbohydrates. The specific molecules and their locations are related to the membrane’s function. Notice the orientation of the phospholipids,
with their hydrophilic heads on each surface and their hydrophobic tails projecting to the membrane’s interior.
2-5 2-6
DNA STAIN DNA appears reddish when treated with a Feulgen stain. Notice that
ELECTRON MICROGRAPH OF A MEMBRANE In this TEM, the
the DNA is found in the nuclei. (X300)
membrane (arrow) looks like a black line. At higher
magnification, membranes typically appear as two dark
bands surrounding a lighter band on electron micrographs.
(X140,000) (Courtesy of UCSD Medical Center)
13
2-7
CHROMOSOME SPREAD Chromosomes are compact structures that are
an efficient way of distributing the long DNA molecules to the two
daughter cells during division. They are not visible during most of the 2-8
cell cycle. These chromosomes are from a male cell grown in tissue
culture. (X650) CHROMOSOMES Chromosomes are visible in tissues where the cells are
actively dividing, as in these cells from the testes. (X650)
Nu
2-9
CHROMATIN GRANULES Dense regions of chromatin in the nucleus are
often visible in light micrographs and may be of use in cell identification.
(X530)
EUCHROMATIN AND HETEROCHROMATIN In this

transmission electron micrograph, dense regions
of inactive chromatin, called heterochromatin
(H), are visible at the periphery of the nucleus
(N). DNA that is being transcribed forms the
lighter euchromatin (E). The medium gray region
near the center is the nucleolus (Nu). (X6,000)
(Courtesy of UCSD Medical Center) 2-10
14
2-12a
2-11
A “DRUMSTICK” The inactive X-chromosome in females is
often visible as an appendage (arrow) of the nucleus in white
blood cells called neutrophils. (X650)
2-12b
NUCLEOLUS AND NUCLEOPLASM The nucleolus
is a prominent dark region in light micrographs
of the nucleus. It should not be confused with
chromatin granules, which are usually toward
the periphery. The nucleoplasm is the lighter,
background material in the nucleus. The nuclear
envelope is also visible as a dark line on the
outside of the nucleus. (a) Testicular interstitial
cells. (X650) (b) Liver cells. (X650)
NUCLEAR ENVELOPE AND PORES In this transmission electron

micrograph, the nucleus is toward the right and the cytoplasm
is toward the left. The double membrane of the nuclear enve-
lope runs vertically in the center of the field. A nuclear pore
is also visible (arrow). (X31,500)
2-13 (Courtesy of UCSD Medical Center)
15
2-14
MICROFILAMENTS The cytoskeleton is composed of many different kinds of
protein filaments. The thin lines (arrows) in the cytoplasm of this neuron are
actin filaments. (X530)
2-15
SKELETAL MUSCLE The banding seen in these skeletal muscle
cells is due to the highly organized arrangement of the protein
filaments actin and myosin. (X530)
M A
2-16
TEM OF ACTIN AND MYOSIN FILAMENTS IN SKELETAL MUSCLE Thick myosin 2-17
(M) and thin actin (A) filaments are visible in this skeletal muscle cell where
CENTRIOLES AND THE CENTROSOME Centrioles are paired,
they form the contractile apparatus. (X40,000) (Courtesy of UCSD Medical Center)
cylindrical structures composed of 9 triplets of microtubules.
(One triplet is indicated by an arrow.) The centrioles are
oriented at 90° to one another and together form the
centrosome. The single centriole shown in this transmission
electron micrograph is cut in cross section. (X48,000)
(Courtesy of UCSD Medical Center)
16
M
R
RER
2-18
MICROTUBULES OF THE SPINDLE APPARATUS Microtubules 2-19
(M) are barely visible with the light microscope. Here they
are shown forming the spindle apparatus of mitosis. The TEM OF RIBOSOMES ON ROUGH ENDOPLASMIC RETICULUM Ribosomes (R) are
spindle fibers radiate outward from the centrosome (C) small, electron dense particles that are the site of protein synthesis. Ribosomes
region, which can be localized because of its position at responsible for producing proteins for secretion are bound to the endoplasmic
the apex of spindle fibers. (X530) reticulum to form rough endoplasmic reticulum (RER). (X20,000)
ZC
MC
2-21a
2-20
TEM OF FREE RIBOSOMES These free ribosomes (R) are not bound to
membrane and produce proteins for internal use. (X31,500)
NB
RER AFFECTS THE APPEARANCE OF THE CYTOPLASM

The light microscope cannot resolve the detail of
RER. However, if it is abundant, the cytoplasm
appears granular and dark staining. (a) Zymogenic
cells (ZC) produce proteins (enzymes) for secretion.
Contrast these with the pale staining mucus cells
(MC) visible in the field. Because mucus is rich in
polysaccharides, RER is not required in its pro-
duction. (X530) (b) Regions of RER in neurons
appear as Nissl bodies (NB) with the light
microscope. (X530) 2-21b
17
2-23a
2-22
GOLGI APPARATUS AS VIEWED WITH THE TEM This membrane
bound organelle is responsible for processing and packaging
materials for secretion. (X24,000) (Courtesy of UCSD Medical Center)
2-23b
GOLGI APPARATUS AS VIEWED WITH THE LIGHT MICROSCOPE
(a) The Golgi apparatus is occasionally visible as a light
staining region near the nucleus of some cells. The cell
indicated is an antibody secreting plasma cell. (X650) (b)
This specimen has been prepared by the DaFano Silver
method, a stain that makes Golgi bodies dark. (X650)
MITOCHONDRIA Mitochondria (M) are the

organelles that house the enzymes and electron
transport chain of aerobic respiration. The inner
and outer membranes are clearly visible in this
2-24 TEM. (X31,500) (Courtesy of UCSD Medical Center)
18
2-25
CILIA Cilia (C) are thin, motile projections responsible for moving materials
across the cell’s surface. They are more numerous and shorter than flagella.
(Compare to Figure 2-26). These cilia are from the respiratory tract where they
sweep inhaled materials away from the lungs. Do not confuse cilia with a brush
border of microvilli. Microvilli are usually shorter and individual ones cannot be
seen, as is often the case with cilia. (X530) 2-26
FLAGELLA Sperm are the only human cells that have flagella.
The tails of these sperm cells are the flagella. (X650)
2-27
CROSS SECTION OF CILIA The 912 microtubular
arrangement is apparent in this TEM of cilia in cross
section. Flagella have exactly the same arrangement.
(X40,000) (Courtesy of UCSD Medical Center)
2-28
MICROVILLI Microvilli (arrows) are foldings of the surface membrane that increase
surface area for absorption. (X15,000) (Courtesy of UCSD Medical Center)
19
BB
2-29
BRUSH BORDER Individual microvilli are not visible with the light mi-
croscope. Rather, a fuzzy band—the brush border (BB)—is seen. These
cells are from the small intestine. Do not confuse a brush border with
cilia. Typically, individual cilia can be discerned, and often they are
clumped, like bent teeth on a comb. (X530)
2-30
STEREOCILIA The misnamed stereocilia are actually long microvilli.
These are from the epididymis. (X240)
BB
ID
ID
2-31
TERMINAL BARS The junctions between adjacent intestinal epithelial
cells are sometimes seen as terminal bars. Use of the electron micro- 2-32
scope has uncovered several different types of intercellular junctions.
Also note the brush border (BB). (X530) INTERCALATED DISCS The junctions between adjacent cardiac muscle
cells are complex and include highly interdigitated cell surfaces joined
by several types of intercellular attachments. In the light microscope,
these appear as the intercalated discs (ID). Do not confuse intercalated
discs with the striated pattern produced by the organized arrangement
of actin and myosin filaments. (X530)
20
2-33a
CELLULAR INCLUSIONS Some storage products,
such as glycogen and lipid, appear in the
cytoplasm of cells. (a) This liver specimen has
been stained for lipids with osmium tetroxide,
which makes them appear brownish. (X650)
(b) PAS positive glycogen in this liver specimen
appears reddish. (X650)
2-33b
21
2-34a 2-34b 2-34c
CF
2-34d 2-34e 2-34f

MITOSIS IN WHITEFISH EMBRYO CELLS While
not human, this tissue provides many ex-
amples of mitotic figures for identification
and is frequently studied in introductory
classes. All specimens have been magnified
X530. (a) Interphase These cells have
obvious nuclear membranes and the nuclear
material is still dispersed as chromatin. (b)
Prophase Several cells are seen in various
stages of prophase. Note the presence/
absence of a nuclear membrane and the
spindle apparatus, and the chromatin
beginning to condense as visible chromo-
somes. (c) Metaphase The two cells in the
center are in metaphase. In each, notice
the chromosomes are aligned at the cell’s
2-34g 2-34h equator and the well-formed spindle appa-
ratus. (d) Early Anaphase The spindle
apparatus has begun to separate the chromosomes in the upper cell. The cell near the bottom of the field is in metaphase. (e) Late Anaphase The
chromosomes are close to the centrosomes, which means they have moved apart just about as far as they will go. (f) Telophase The chromosomes
have reached opposite poles of the cell and the spindle fibers have begun to break down. The cleavage furrow (CF) has also begun to form. (g)
Late Telophase and Cytokinesis The cleavage furrow has nearly divided the two cells, so cytokinesis is almost complete. Telophase will continue
with the chromosomes dispersing as chromatin and the nuclear envelope reforming. (h) Polar View The cell in the center doesn’t fit any of the
descriptions because it is being viewed from one end (pole) rather than from above.
22
MITOSIS IN LYMPHATIC TISSUE

Mitotic figures can also be seen
by careful observers in many
different tissues. This specimen is
from a lymph node. Several cells
are undergoing mitosis (arrows),
but most are in interphase. Notice
the chromatin granules and the
nucleoli in these nondividing cells.
2-35 (X650)
Epithelial Tissue
C H A P T E R
3
Basic Characteristics of Epithelial Tissues Functions of Epithelia
Epithelial tissues or epithelia cover most surfaces in the The structures of epithelia vary depending on their func-
body. Typically, the cells are joined by junctional complexes tions. Some functions of epithelia are:
(see Chapter 2) into sheets with little intercellular material ◗ Mechanical protection from abrasive forces (e.g., the
(Figure 3-1). (For a graphic example, recall the last time epidermis).
you peeled after a sunburn!) Epithelia rest on a basement ◗ Absorption of substances from the lumen (inner por-
membrane that separates them from the underlying con- tion) of a tubular organ (e.g., intestinal epithelium).
nective tissue (Figure 3-2). Electron micrographs show that The luminal surface is usually modified with microvilli
the basement membrane is composed of a basal lamina to increase the surface area and improve efficiency of
(derived from the epithelial cells) and a lamina reticularis absorption.
(produced by fibroblast cells of the connective tissue).
◗ Secretion of materials (e.g., intestinal epithelium). This
Blood vessels do not penetrate the basement membrane,
epithelium often has single-celled, mucigen-secreting
making epithelia avascular.
goblet cells (see page 24) in it.
◗ Lubrication of surfaces (e.g., mesothelium of pericard-
Epithelial Membranes ium).
Since epithelia are avascular, they rely on nourishment from ◗ Formation of a diffusion membrane (e.g., alveoli of
the capillaries of the underlying connective tissues. This lung).
results in a structural and functional association between
an epithelium and its underlying connective tissue: an Epithelial Terminology
epithelial membrane. Epithelial membranes are of three Some important terminology associated with epithelia is
types. Mucous membranes line surfaces open to the illustrated in Figure 3-4. If the cell is actually on the surface
external environment and often secrete mucus (Figure 3-3). (and is not buried in the epithelium) the surface edge of the
Examples are the membranes lining the digestive, respira- cell is called the free, apical, or luminal edge. The sides
tory, urinary and reproductive tracts. Serous membranes contacting other epithelial cells are called lateral edges, and
line the ventral body cavities—pericardial, pleural, and the edge in contact with the basement membrane is called
peritoneal cavities—as pericardium, pleura, and peritoneum, the basal edge. If the epithelium consists of more than one
respectively. The epithelial component of serous mem- layer, the basal cells are closest to the basement membrane.
branes is called mesothelium. Cutaneous membrane is the Epithelia are named based on two main criteria: the
skin. number of cell layers and the shape of the cells at the
23
24
surface. Simple epithelia have a single layer of cells, so all Other Epithelia
cells contact the basement membrane. Stratified epithelia Some epithelia do not conform very well to the conven-
are made of two or more cell layers and only the basal cells tional naming criteria. These are considered in this section.
contact the basement membrane. Squamous cells are flat Transitional epithelium, found lining the urinary blad-
with flattened nuclei. (Note: The shape of the nucleus is der and ureters, is stratified and specialized for stretching.
important in identifying an epithelium because nuclei tend When in the stretched state (Figure 3-20), all cell layers are
to be the most obvious part of a cell.) Cuboidal cells are flattened. When relaxed (Figures 3-21 and 3-22), the cells
cube-shaped with spherical nuclei positioned in the cell’s take on an irregular appearance, especially in the most
center. Columnar cells are taller than wide and have an superficial layer where they often bulge into the lumen,
elongated, basal nucleus. To completely name an epithe- giving the surface a scalloped appearance.
lium, both the number of cell layers and the shape of cells Pseudostratified ciliated columnar epithelium—which
at the surface must be included. is shortened to PSCC—is not a truly stratified epithelium
since all cells are in contact with the basement membrane
Simple Epithelia (Figures 3-2, 3-23, and 3-24). However, not all cells reach
Simple squamous epithelium is composed of a single layer the surface; some cells are short, some are tall, and others
of flat cells. It forms the alveoli of the lungs (Figure 3-5) are intermediate in height. This variation in cell height
where it participates in forming the respiratory membrane results in nuclei being seen at different levels in the
through which gases diffuse, and glomerular capsules of the epithelium, giving the false impression of stratification—
kidney where blood filtration occurs (Figure 3-6). The hence “pseudostratified.” And, since the cells that reach the
simple squamous lining of blood vessels and the heart is surface are taller than wide, the epithelium is considered to
called endothelium (Figure 3-7). Mesothelium is the epithe- be columnar. It is most associated with the respiratory tract
lial component of serous membranes and secretes a lubri- where it lines the nasal cavity, trachea, and bronchi. Goblet
cating fluid (Figures 3-1 and 3-8). cells produce mucus that traps dust and other particles in
Simple cuboidal epithelium is composed of a single inspired air, then the cilia sweep the mucus toward the
layer of cube-shaped cells. It is found in kidney tubules pharynx where it is swallowed.
(Figure 3-9) where it is involved in secretion and absorp-
tion, and ducts of glands (Figure 3-10), among other places.
Simple columnar epithelium (Figure 3-11) is made of a
Glandular Epithelia
single layer of tall cells with basally positioned, elongated Glandular epithelia are specialized for secreting materials.
nuclei. It is typically involved in absorption and secretion, They develop from typical lining epithelium that grows
and often has microvilli and goblet cells (Figure 3-12). down (invaginates) into deeper tissues and as such is not
found on the surface. Exocrine glands remain connected to
the surface and deliver their secretion to it by way of a
duct. They are the subjects of this section. Endocrine glands
Stratified Epithelia (Chapter 10) lose the connection to the surface during
Stratified squamous epithelium is composed of several to development, so the secretion is transported by the blood.
many layers of cells with the superficial layers being flattened. Unicellular glands are represented by the abundant
Since it is so thick, it is generally found in places subjected goblet cells of the respiratory and digestive tracts (Figures
to abrasion. Keratinized stratified squamous (Figures 3-13 3-25 and 3-26). Goblet cells secrete mucigen, which con-
through 3-16) is found on dry surfaces (e.g., the skin). As verts to mucin when hydrated. Mucin is a major com-
the epithelial cells produced at the base get pushed to the ponent of mucus.
surface, they die and undergo changes, including accumu- Multicellular exocrine glands are more complex, with
lation of the protein keratin. The dead cells so-formed specialized duct cells and secretory cells (Figure 3-27). Photo-
provide a waterproof, microbe-proof, abrasion barrier. micrographic examples are provided in Figures 3-28
Nonkeratinized stratified squamous (Figure 3-17) is through 3-34. Multicellular glands are classified according
found in moist areas of the body not subjected to as much to their duct(s) and their secretory portions. If the duct is un-
abrasion (e.g., oral cavity, esophagus, and vagina). Unlike branched, the gland is simple; if it branches, the gland is
the keratinized variety, the surface cells are living and have compound. If the secretory portion is about the same size as
normal nuclei. the duct, the gland is tubular. If the secretory cells are larger
Stratified cuboidal and stratified columnar epithelia than the duct, it is an acinar (or alveolar) gland. Some
generally consist of only a couple of layers. They are found glands have both acinar and tubular secretory portions.
in the ducts of some glands (Figures 3-18 and 3-19), with The secreting cells and the ducts comprise the paren-
the cell height corresponding to the duct’s size. chyma of the gland. In addition, there may be a connective
C H A P T E R T H R E E : E P I T H E L I A L T I S S U E
25
tissue stroma in the gland that supports the parenchyma. In Myoepithelial cells (Figure 3-35) are associated with
larger glands, the connective tissue capsule on the surface some acini. These are contractile epithelial cells that push
sends branches into the gland and divides it into lobes and the gland’s secretion into the duct. They are found in
lobules. salivary and sweat glands.
3-1 3-2
EPITHELIUM IS A CELLULAR TISSUE This is a whole mount of meso- LIGHT MICROGRAPH OF A BASEMENT MEMBRANE Shown is an epi-
thelium, an epithelium composed of a single layer of flat cells. Viewed thelium from the respiratory tract that has a prominent basement
from the surface, the very cellular and sheet-like nature of epithelial membrane (arrow). In sections of most epithelia, the basement
tissue is apparent. Most specimens you will encounter are viewed in membrane is not as obvious. In many cases, all that is visible is the
section, so are seen from the side. (X300) junction between the epithelium and the underlying connective tissue.
(X240)
AN EPITHELIAL MEMBRANE All epithelial membranes

(mucous, serous, and cutaneous membranes) are com-
posed of an epithelium and its underlying connective
tissue. In each, the avascular epithelium relies on the
capillaries of the connective tissue for its oxygen and
nutrients, and thus forms a functional as well as a
structural unit. Illustrated here is the colon’s mucous
membrane. Notice the numerous, pale staining mucus
3-3 cells in the epithelium. (X70)
26
Cilia on luminal surface
Terminal
bar
Surface
layer
Epithelium
Basal layer Basement

membrane
Basement
membrane Connective
tissue
Connective
tissue
3-4
EPITHELIAL TERMINOLOGY Illustrated are standard descriptive terms associated with epithelia. Notice the importance of the basement membrane
and surface as points of reference.
3-5 3-6
SIMPLE SQUAMOUS EPITHELIUM OF THE LUNG Notice that the squamous SIMPLE SQUAMOUS EPITHELIUM OF THE KIDNEY The renal corpuscles of
cells are so thin, only their nuclei are visible (arrows). (X380) the kidney are lined with a simple squamous epithelium (arrow).
Again, the flattened nuclei are the most prominent feature of the
epithelium. (X530)
27
BV
BV
3-7
SIMPLE SQUAMOUS ENDOTHELIUM The inside of all blood vessels and
3-8
the heart are lined with a simple squamous endothelium (arrow). This MESOTHELIUM The outer surface of organs in the ventral body cavity is
micrograph is of a small vein. (X320) lined with a serous membrane made of a connective tissue and a simple
squamous mesothelium (arrows). Also note the simple squamous
endothelium lining the various blood vessels (BV). (X150)
3-9
SIMPLE CUBOIDAL EPITHELIUM These renal tubules are lined with a 3-10
simple cuboidal epithelium. The lateral membranes are not visible
between all cells, but their cuboidal shape is still obvious. The cells A TALL SIMPLE CUBOIDAL EPITHELIUM The pink cells are considered
apparently missing nuclei are the result of their nuclei not being in the simple cuboidal epithelium rather than simple columnar because of the
plane of the section. The pinkish line at the base of the epithelium is the shape and position of their nuclei. In some cases, classification between
basement membrane, prominently stained by the PAS method. (X320) cuboidal and columnar becomes a judgment call. These are ducts in a
salivary gland. (X320)
28
BB
GC
3-11
SIMPLE COLUMNAR FROM THE STOMACH These simple columnar cells
are characterized by a tall shape and a basal, elongated nucleus. The 3-12
basement membrane is not apparent in this specimen, but it is still SIMPLE COLUMNAR EPITHELIUM WITH GOBLET CELLS AND A BRUSH
clear where the epithelium ends and the connective tissue begins BORDER This intestinal epithelium is simple columnar. The large pale
(arrow). The region indicated by the bracket is a curved surface lined staining cells are goblet cells (GC) that secrete mucus. A brush border
by the same epithelium, but cut tangentially. Notice the similarity of (BB) is clearly visible over most of the surface. (X320)
the light region to the luminal portion of the other cells. (X320)
3-13
THIN KERATINIZED STRATIFIED SQUAMOUS EPITHELIUM The flat surface 3-14
cells are dead and form the shredded keratinized layer. This specimen is
from abdominal skin. The whole epithelium is only a few cells thick. MORE THIN KERATINIZED STRATIFIED SQUAMOUS EPITHELIUM This
(X320) specimen illustrates the convention that epithelia are named according
to the cell shape at the surface. Note how tall the basal cells are.
(X320)
29
K
K
BM
BV
BV BV
BV
3-15
THICK KERATINIZED STRATIFIED SQUAMOUS EPITHELIUM This specimen is 3-16
from palmar skin. Notice the absence of nuclei in the thick keratinized
layer (K). Also, notice the wavy basement membrane (BM) typical of MORE THICK KERATINIZED STRATIFIED SQUAMOUS EPITHELIUM In this
this epithelium—so typical, in fact, that it is often identifiable at low higher magnification, some cell and nuclear outlines are still visible in
power without even seeing the individual cells. (X50) the keratinized layer (K). Notice the wavy basement membrane and the
prickly appearance of the living cells due to intercellular attachments
(desmosomes). Also, notice the abundant blood vessels (BV) in the
deeper connective tissue and their absence in the epithelium—
remember that epithelia are avascular. The simple squamous
endothelium of the blood vessels is also visible. (X210)
3-17 BV
NONKERATINIZED STRATIFIED SQUAMOUS EPITHELIUM In this specimen
from the vagina, the cells are flat at the surface and are still alive, as
evidenced by the presence of nuclei. (The slight shredding of the
surface is not a keratinized layer—the cells are alive.) The wavy 3-18
basement membrane is also visible. (X100)
STRATIFIED CUBOIDAL EPITHELIUM Ducts of glands may be lined with a
stratified cuboidal epithelium. This stratified epithelium is only two
cells thick. Notice the simple squamous endothelium of the blood
vessel (BV) in the lower right corner of the field. (X320)
30
3-20
3-19 TRANSITIONAL EPITHELIUM—STRETCHED Transitional epithelium is
stratified and is found in the urinary bladder and ureters. It is
STRATIFIED COLUMNAR EPITHELIUM Larger ducts of glands may be lined specialized for stretching. When stretched, the cells flatten, as in this
with a stratified columnar epithelium. Notice that the basal cells are micrograph. This might be confused with stratified squamous, but
cuboidal and only the surface layer is columnar. (X210) notice the straight basement membrane (arrow). (X210)
3-21 3-22
TRANSITIONAL EPITHELIUM—RELAXED When the urinary bladder empties, MORE TRANSITIONAL EPITHELIUM—RELAXED The rounded apical surface
the wall relaxes and the epithelium assumes the shape shown in this of the superficial cells is prominently displayed in this specimen of
micrograph. Most of the cells are vertically elongated and the apical transitional epithelium. The arrow indicates the level of the basement
edge of the surface cells is rounded. The arrow indicates the level of the membrane. (X210)
basement membrane. (X320)
31
C
C GC
GC
BM
BM
3-23
PSEUDOSTRATIFIED CILIATED COLUMNAR EPITHELIUM (PSCC) PSCC is 3-24
found in the respiratory tract. This specimen is from the trachea. Short, ANOTHER PSCC Since the actual cell boundaries are not very clear
intermediate, and tall cells, all of which touch the basement membrane, in this specimen of PSCC, one might confuse this with a stratified
characterize it. It is the various heights of nuclei that give the impres- columnar epithelium. However, stratified columnar epithelium usually
sion of a stratified epithelium. Also present are cilia (C) on the tallest has only a couple of cell layers and the nuclear layers are fairly distinct,
cells, mucus secreting goblet cells (GC), and a prominent basement not jumbled as in this micrograph. Globlet cells (GC), cilia (C), and the
membrane (BM). (X320) basement membrane (BM) are clearly shown. (X320)
BB GC
N N
3-25 3-26
GOBLET CELLS OF THE COLON The numerous goblet cells in the colon’s GOBLET CELLS STAINED BY PAS METHOD Due to their high polysaccharide
simple columnar epithelium are visible. One cell, indicated by the arrow, content, the mucin granules of the goblet cells (GC) are PAS positive
has released its secretion, mucin, which when hydrated becomes mucus. and appear red. Also notice the basal nuclei (N) of the goblet cells and
Flat, dense staining goblet cell nuclei (N) are visible below the mucin the brush border (BB) on the simple columnar epithelium. This
droplets in a couple of cells. (X320) specimen is from the small intestine. (X530)
32
Simple tubular Simple branched tubular Simple coiled tubular
Simple acinar
Simple branched acinar
Compound acinar
Compound tubular
3-27
MULTICELLULAR GLANDS Multicellular glands are composed of a duct (lighter purple) and a secretory portion (darker purple). If the duct is
unbranched, the gland is simple; if branched, the gland is compound. In tubular glands, the secretory cells are about the same size as the duct
cells. If the secretory cells are larger than the duct cells, the gland is alveolar or acinar. All combinations of simple/compound and tubular/acinar
glands are possible.
SIMPLE TUBULAR GLANDS OF THE COLON

The crypts of Lieberkühn in the colon
are simple tubular glands. The simple
columnar epithelium also has abundant
3-28 goblet cells. (X100)
33
SwG
3-30
3-29 BRANCHED TUBULAR GLANDS The mucosal glands of the stomach
COILED TUBULAR GLAND OF THE SKIN A relatively straight duct (D) (gastric glands) are branched tubular glands (G). The straight, tubular
connects the coiled secretory part of a sweat gland (SwG) to the skin’s nature of the glands is apparent even at this magnification. Several
surface. Several sweat glands and their ducts are visible in this field. glands branch off each mucosal depression at the level of the arrows
Also note the keratinized stratified squamous epithelium. (X50) and extend to the base of the mucosa. (X50)
SbG
SbG
BG
3-31 3-32
BRANCHED ACINAR GLAND Sebaceous glands (SbG), associated with COMPOUND BRANCHED TUBULAR GLAND Mucus secreting Brünner’s
hair follicles, are branched alveolar glands. Two glands connected to glands (BG) of the duodenum are compound branched tubular
the follicle are shown in this field. On the left are pieces of other glands—that is, both the ducts and secretory parts are branched. The
branches that connect in a different plane. The secretory cells release glands are the lighter staining cells in the middle third of the field.
their secretion by disintegrating, which is apparent in this specimen. Their uniform diameter and branching are apparent in this specimen.
(X100) Notice the goblet cells in the simple columnar epithelium. (X100)
34
3-33
COMPOUND ACINAR GLAND The exocrine portion of the pancreas is made of compound acinar
glands. The smaller of the two ducts (D) visible in the center of the field is probably a branch of
the larger one. Clusters of secretory cells called acini surround the ducts. One acinus is circled.
(X250)
3-34 3-35
COMPOUND TUBULOACINAR GLAND This salivary gland has tubular and MYOEPITHELIAL CELLS In sweat glands (as in this specimen) and sali-
acinar secretory portions, as well as numerous branched ducts. The vary glands, myoepithelial cells (arrow) occupy the region between the
gray secretory cells produce mucus, whereas the darker, more granular secretory cells and the basement membrane. Their contraction assists
cells produce enzymes. The ducts (D) are lined with a simple columnar movement of the secretion into the duct. (X530)
epithelium and have an obvious lumen. (X200)
Fibrous Connective Tissues
C H A P T E R
4
Basic Characteristics of Connective Tissues derived from embryonic mesoderm. The cells of mesenchy-
Connective (supporting) tissues anchor organs and join the mal tissue are unspecialized and have the potential to differ-
other tissues of the body into a structurally integrated whole. entiate into the cells typical of adult connective tissues
In fact, most organs have a connective tissue covering that (Figure 4-2). Further, adult connective tissues appear to re-
often penetrates it and binds the whole organ together. Un- tain some mesenchymal cells that act as a source of cells in
like epithelia, connective tissues generally have abundant case repair is necessary. (Some authors attribute the source
extracellular (intercellular) matrix and relatively few cells. of cells to pericytes found associated with capillaries.) Mes-
The matrix consists of protein fibers, a ground substance, enchymal cells are angular or spindle-shaped and form a
and other supporting biochemicals. Connective tissues may loose mesh that is functionally a rudimentary connective
be vascular or avascular. tissue. Fibers are absent.
Fibrous connective tissues (Connective Tissue Proper) Mucous tissue (Figure 4-3) is found only in the umbili-
are the subject of this chapter. Cartilage and bone are cov- cal cord and a few other places in the embyro. It has fibro-
ered in Chapter 5, and Blood and Bone Marrow are cov- blasts and very few fibers coursing through a jelly-like
ered in Chapter 6. ground substance (Wharton’s jelly).
Functions of Fibrous Connective Tissues Extracellular Matrix of Adult Connective Tissues

The functions of fibrous connective tissues are many and The properties of a connective tissue are largely due to the
varied. Some examples are: properties of its matrix. Comprising matrix are the ground
substance, fibers, and structural glycoproteins.
◗ Binding and support (e.g., ligaments)
Ground substance is an amorphous, gel-like material
◗ Defense (e.g., macrophages and mast cells of various composed of charged glycosaminoglycans—GAGs—(mu-
tissues) copolysaccharides) and proteoglycans (mucoproteins) of
◗ Storage (e.g., adipose tissue) various types. Both are polymers of disaccharide subunits,
◗ Transport of materials between blood and other tissues but the latter are also bonded to proteins. Their charges
(e.g., tissue fluid in loose connective tissue and others) make them hydrophilic, resulting in tissue fluid mixing read-
ily with the ground substance and helping create the gel-like
Embryonic Connective Tissues— state. Because of this association, the ground substance is an
Mesenchyme and Mucous Tissue essential participant in the transport of nutrients and wastes
Connective tissues of the adult are derived from an em- between the blood and other tissues. Ground substance is
bryonic tissue called mesenchyme (Figure 4-1), which is not easily visualized in histological preparations.
35
36
Fibers are made of protein and come in two basic types: cells (Figure 4-11b). Multilocular fat cells (Figure 4-12) are
collagen and elastic. Collagen fibers (Figure 4-4) are made smaller, store fat in many droplets, and have a spherical
of the protein collagen and are the primary fibers of con- nucleus.
nective tissues. At least 19 types of collagen have been iden- Other leukocytes (white blood cells) of various types
tified. All exhibit a high tensile strength and are inelastic. may be seen in connective tissues, especially at the site of
Type III collagen fibers (Figure 4-5) are thin and branched, infection or inflammation. Most commonly seen in healthy
and were formerly known as reticular fibers. These fibers tissue preparations are lymphocytes and plasma cells (Fig-
form a framework of the liver, lymphatic tissue and bone ures 4-13 and 4-14). Lymphocytes have a dark staining nu-
marrow. Elastic fibers (Figures 4-4, 4-6, and 4-7) are made cleus surrounded by a thin layer of cytoplasm. Plasma cells
of the protein elastin. The elasticity they confer is an impor- are derived from lymphocytes and secrete antibodies. They
tant property of organs that can be deformed and then re- have a purplish cytoplasm (due to abundant RER necessary
turn to their original shape, such as larger arteries and the for antibody production) and an eccentric nucleus. A pale
skin. region next to the nucleus may also be visible. It is the site
Structural glycoproteins are involved in anchoring and of the Golgi apparatus. Neutrophils have a granular cyto-
fastening cells to extracellular material, including basement plasm and a segmented nucleus. They are phagocytic.
membranes.
Classification of Adult Fibrous Connective Tissues
Cells of Adult Connective Tissues Fibrous connective tissues have traditionally been classified
according to the arrangement and density of fibers. Regular
There are many cell types in fibrous connective tissues. Fi-
connective tissues have the fibers oriented in the same direc-
broblasts, mast cells, macrophages, adipocytes, lympho-
tion; the fibers of irregular connective tissues are oriented in
cytes and plasma cells are the most commonly encountered
all directions. The fibers of loose connective tissues occur
ones.
singly, whereas the fibers of dense connective tissue occur in
Fibroblasts (Figures 4-4, 4-6, and 4-8) are derived from
bundles and are tightly packed together. While these cate-
mesenchymal cells and are responsible for synthesis and
gories will be used here, it should be realized that they rep-
maintenance of the matrix. They are elongated in the direc-
resent extremes and that intermediate connective tissue
tion of fiber orientation and have a granular nucleus with a
types exist. Further, some authors consider adipose tissue
prominent nucleolus.
and reticular tissue as loose connective tissues, but they will
Macrophages (Figure 4-9) are derived from blood
be considered separately in this chapter.
monocytes and are found in a variety of tissues. Their func-
tion is to phagocytose (engulf) foreign, dead and dying Loose Areolar Tissue
cells, and cellular debris. In their role as antigen presenting Loose areolar tissue (Figures 4-4, 4-6, 4-8, 4-15 and 4-16)
cells (APCs) they present antigens to lymphocytes as part of is the connective tissue component of serous and mucous
the immune response. Resident (fixed) macrophages (Figure membranes, acts as filler between muscles and between
4-9a) are regular inhabitants of a particular tissue whereas muscles and skin, and is found in various other locations
elicited (wandering or free) macrophages (Figure 4-9b) cir- in the body. Collagen and elastic fibers are present, as are
culate in the blood and migrate to where they are needed. fibroblasts, mast cells, macrophages, and most other con-
In the light microscope, macrophages have an irregular nective tissue cells. It is vascular.
shape, a basophilic, finely granular cytoplasm, and an oval-
to kidney-shaped nucleus. Dense Irregular Connective Tissue
Mast cells (Figure 4-10) originate in the bone marrow, Dense irregular connective tissue (Figures 4-17 and 4-18) is
but occupy various connective tissues. In many ways, they characterized by densely packed collagen bundles oriented
resemble blood basophils, but are not developmentally re- in all directions. It is vascular and fibroblasts are the pre-
lated to them. They are large with prominent cytoplasmic dominant cell type. It comprises the dermis and capsules of
membrane-bound granules. The granules contain chemicals many organs.
(such as heparin, histamine, chemotactic factors, and many
others) which are involved in the inflammatory response. Dense Regular Connective Tissue
Release of the granules’ contents is called degranulation. Dense regular connective tissue is an avascular tissue that
Adipocytes are derived from mesenchymal cells and, has fiber bundles arranged in parallel fashion. This fiber
perhaps, fibroblasts. They are specialized to store fat. arrangement confers great tensile strength on the tissue.
Unilocular fat cells (Figure 4-11) store the fat as a single, Fibroblasts are compressed and appear elongated between
large droplet that pushes the nucleus and cytoplasm to the the compact fibers.
cell’s periphery. Often, the fat is dissolved from the adipo- Tendons provide an example of dense regular collage-
cytes during slide preparation, so all that is seen is empty nous connective tissue (Figure 4-19). Dense regular elastic
C H A P T E R F O U R : F I B R O U S C O N N E C T I V E T I S S U E S
37
connective tissue has an abundance of elastic fiber bundles

and is found in larger blood vessels and some vertebral liga-
ments (Figure 4-20).
Adipose Tissue
Adipose tissue is a vascular tissue and differs from other
connective tissues in that it is very cellular with little inter-
cellular material. White adipose tissue (Figures 4-11 and
4-21) is composed of unilocular adipocytes and is com-
monly found in the subcutaneous regions as well as in
serous membranes. Fixing slides by standard methods dis-
solves the fat, so “empty” adipocytes are what are typically
seen. In some procedures, the fat is retained and can be
stained, as with the oil-soluble dye Sudan red (Figure
4-11a).
Brown adipose tissue (Figure 4-22) is present in the em-
bryo and possibly persists in modified form in the adult. It
is composed of multilocular adipocytes and is much more
vascular than white adipose tissue. It is involved in genera-
tion of body heat.
Reticular Connective Tissue

Reticular connective tissue (Figure 4-23) is a loose connec-
tive tissue with an abundance of reticular fibers (Type III
collagen). The fibers are not stained with H&E, but can be
visualized with silver stains. Reticular connective tissue 4-1
forms the framework of bone marrow, lymphoid organs,
MESENCHYME Mesenchymal tissue is a primitive connective tissue
and the liver sinusoids. lacking in fibers. Its cells are angular and have the potential to develop
into more specialized connective tissue cells. (X610)
4-2
SCHEME OF CONNECTIVE TISSUE CELL DEVELOPMENT Connective tissues develop from embryonic mesoderm which gives rise to mesenchymal
cells and pluripotent hemopoietic stem cells (HSC). Mesenchymal cells give rise to fibroblasts and adipocytes of connective tissue proper
(covered in this chapter), and cartilage and bone cells (covered in Chapter 5). The HSC gives rise to all formed elements in blood
(covered in Chapter 6).
38
EF
CF
4-4
COLLAGEN FIBERS Thick, strong collagen fibers (CF) are seen in this
specimen of loose areolar tissue. The thin fibers are elastic fibers (EF),
and most of the cells are fibroblasts. (X100)
4-3
MUCOUS TISSUE Mucous tissue is found only in the umbilical cord
and a few other sites in the embryo. It contains fibroblasts (F) and a
few fibers, so it is more specialized than mesenchyme. The ground
substance is like jelly. (X380) EF
MF
RF
CF F
4-6
4-5
ELASTIC FIBERS IN LOOSE AREOLAR TISSUE The thin, dark lines are
RETICULAR FIBERS (TYPE III COLLAGEN FIBERS) Reticular fibers (RF) are elastic fibers (EF) in this loose areolar tissue. Also visible are
thin and branched. This specimen is a lymph node, but they also form the collagen fibers (CF), fibroblasts (F), and macrophages (MF).
structural framework of the spleen, bone marrow, and some other organs. (X210)
These fibers do not show up when stained with H&E; a silver stain must be
used to visualize them. (X320)
39
ELASTIC FIBERS IN ELASTIC TISSUE This specimen

is from the aorta. Its wall has abundant elastic
fibers (dark lines) arranged circularly around
the vessel. The elasticity allows the aorta to
stretch when blood is pumped into it, then
recoil and push the blood further when the
heart relaxes. (X100)
4-7
4-8a
FIBROBLASTS The most commonly seen cell in fibrous
connective tissues is the fibroblast. They have an
irregular shape and (usually) a prominent nucleolus.
(a) Most cells in this spread of loose areolar tissue are
fibroblasts. Collagen and elastic fibers are also visible
in this specimen. (X530) (b) A single fibroblast
illustrating the irregular shape is shown here. (X760)
4-8b
40
MF
K
4-9b
MACROPHAGES Macrophages may either be fixed or wandering. (a) The
4-9a
Kupffer cells (K) are fixed macrophages that line the liver sinusoids and
remove material from the blood. These macrophages have ingested a
dye that makes them stand out. Fixed macrophages may also be seen
in the spleen and lymph nodes. (X530) (b) Wandering macrophages
(MF) are found in the lung. (X630)
M
M
4-10a 4-10b
MAST CELLS Mast cells (M) are involved in the inflammatory response by degranulating and releasing histamine and other chemicals. (a) This is
loose areolar tissue with two mast cells shown. The lower mast cell has degranulated. (X630) (b) Several mast cells are visible in this tonsil
specimen. (X630)
41
4-11a 4-11b
UNILOCULAR ADIPOCYTES The cytoplasm and nucleus of unilocular adipocytes is pushed to the periphery and the single, large fat droplet occupies
the majority of the cell. (a) Unilocular adipocytes stained with Sudan Red, a fat-soluble stain that demonstrates the fat droplet. (X100) (b) Standard
slide preparation dissolves and removes the fat droplet and all that remains of the adipocytes is the membrane and some cytoplasm. (X100)
4-12
MULTILOCULAR ADIPOCYTES Only human embryos have multilocular adipocytes (although they
are found in other adult mammals). The fat is stored as several droplets rather than one big one.
(X250)
42
BV
PC
BV
4-14
PLASMA CELL AND NEUTROPHIL Plasma cells (PC) are lymphocytes that
are actively secreting antibodies. They are elongated cells with the nu-
cleus toward one end. The nucleus looks like a clock face because of
4-13a the chromatin distributed around its periphery. A pale region of cyto-
plasm near the nucleus is the site of a Golgi apparatus. A phagocytic
neutrophil (N) is also visible in the field. Its granular cytoplasm is not
very apparent, but the segmented nucleus is. Notice the blood vessels
(BV) in the field. (X630)
EF
CF
MF
MC
F
4-13b
LYMPHOCYTES The lamina propria of mucous membranes is often
infiltrated with lymphocytes, cells of the immune system that secrete
antibodies among other protective functions. They are small, round 4-15
cells with dark staining nuclei. (a) In this specimen from the small
LOOSE AREOLAR TISSUE A loose weave of collagen (CF) and elastic
intestine, the lymphocytes are quite dense. Some have even entered
fibers (EF) characterizes loose areolar tissue (LAT). LAT is found as a
the simple columnar epithelium (arrows). (X250) (b) This specimen is
supporting tissue and as filler between organs (fascia). Fibroblasts (F),
also from the small intestine, but the lymphocytes (L) in the lamina
macrophages (MF), and mast cells (MC) are visible in this specimen.
propria are not as dense. (X250)
(X320)
43
LAMINA PROPRIA Loose

fibrous connective tissue
comprises the lamina
propria of many epithelial
membranes. This specimen
is from the small intestine. BV
Notice the blood vessels
(BV) and lymphocytes (L) in L
the lamina propria. (X400)
4-16
BV
BV
C
C C
BV
4-17
DENSE IRREGULAR CONNECTIVE TISSUE The dermis of the skin is com-
posed of an irregular weave of tightly packed fiber bundles. Although 4-18
all fiber types are present, collagen bundles (C) are the most obvious.
Notice that the bundles are sectioned longitudinally, obliquely, and in FIBROUS CAPSULE Many organs, including the testis shown here, are
cross section, indicating their irregular arrangement. Most of the cells covered by a dense fibrous connective tissue capsule (C). Notice the
are fibroblasts. Blood vessels (BV) indicate this tissue is vascular. (X210) blood vessels (BV). (X50)
4-19a 4-19b
COLLAGENOUS DENSE REGULAR CONNECTIVE TISSUE (a and b) Tendons, shown here, and ligaments, are made of closely packed collagen bundles
all oriented in the same direction. Dense regular connective tissue is avascular. The cells are fibroblasts, and they are typically in rows. (X210)
44
4-20 4-21
ELASTIC DENSE REGULAR CONNECTIVE TISSUE This specimen is an WHITE FAT White fat is made of unilocular adipocytes and may be
elastic ligament from the neck. The waviness of the fibers is an found most anywhere loose connective tissue is found. This specimen
indication of their elasticity. (X250) is from a skeletal muscle, and the fat is filling in between the bundles
of muscle fibers. (X250)
RF RF
4-23
4-22
RETICULAR CONNECTIVE TISSUE Lymphoid organs and bone marrow
BROWN FAT Brown fat is made of multilocular adipocytes and in
have a reticular connective tissue framework. This lymph node specimen
humans, is primarily found in the embryo. (X250)
was prepared with a silver stain so the reticular fibers (RF) could be
seen. (X250)
Cartilage and Bone
C H A P T E R
5
Introduction to Skeletal Tissues and flexibility, respectively. Most of the time, collagen
Cartilage and bone are specialized connective tissues with a fibers are not easily seen in cartilage even though they are
firm to rigid matrix, suiting them for their skeletal and pro- abundant. A silver stain (or other specialized elastin stain)
tective functions. must be used to visualize elastic fibers since H&E does not
stain them.
Cartilage is avascular, but receives nourishment from
Basic Characteristics of Cartilage blood vessels in the perichondrium, the fibrous membrane
Cartilage is on joint surfaces, forms the framework of the that surrounds most cartilage (Figure 5-3). It is composed
nose, ears, respiratory tree and part of the rib cage, and is of a superficial fibrous layer and a deeper chondrogenic cell
located between vertebrae. It also comprises a majority of layer. In appositional growth, the chondrogenic cells differ-
the embryonic skeletal system before it is replaced by bone. entiate into chondroblasts that secrete new matrix on the
The semirigid matrix makes cartilage well-suited for these cartilage’s surface.
supporting functions.
Embryonically, cartilage begins developing from mes- Types of Cartilage
enchyme in chondrification centers. The mesenchymal cells There are three types of cartilage: hyaline cartilage, elastic
differentiate into chondroblasts, which begin secreting ma- cartilage, and fibrocartilage. Hyaline cartilage (Figures 5-1
trix. When they become surrounded by matrix, they are through 5-4) is the most abundant of the three cartilage
called chondrocytes and the space each chondrocyte occu- types. It is found in the nose and respiratory tree, on the
pies is called a lacuna (Figure 5-1). When a chondrocyte sternal ends of the ribs, and on articular surfaces. It also
grows and divides, it forms an isogenous group of cells in forms the cartilage model of the skeleton during develop-
the lacuna. These cells continue producing matrix and be- ment. The matrix of hyaline cartilage is glassy in appear-
come separated into different lacunae. The result is carti- ance (hence, “hyaline”) and has collagen fibers in it, though
lage growth from the interior of the matrix, a process they are not easily seen with the light microscope. Except in
known as interstitial growth. articular cartilage (Figure 5-5) a perichondrium is present.
The characteristic consistency of the matrix is due to Elastic cartilage (Figure 5-6) is found in the ears and
the grouped arrangement of proteoglycans and its degree of epiglottis. It is very similar to hyaline cartilage, except that
hydration. The matrix immediately around each lacuna is its matrix has numerous elastic fibers in addition to the col-
basophilic and is called territorial matrix; interterritorial lagen fibers. The chondrocytes also tend to be larger (at the
matrix is found between lacunae (Figure 5-2). Collagen and expense of the matrix) and more numerous than in hyaline
elastic fibers are present in the matrix and confer strength cartilage. A perichondrium is present.
45
46
Fibrocartilage (Figure 5-7) is found in the intervertebral but their relative amounts differ depending on the particu-
disks and the pubic symphysis. It has abundant collagen fibers lar bone and the specific part of the bone.
in the matrix and the chondrocytes tend to be seen in rows Compact bone has a dense bony matrix with canals
parallel to the fiber bundles. There is no perichondrium. that carry blood vessels. Volkmann’s canals penetrate from
the surface and Haversian canals run more or less parallel
Basic Characteristics of Bone— to the surface (Figure 5-14). The structural and functional
Matrix and Cells unit of compact bone is the Haversian system or osteon
Bone matrix is composed of about 25% organic (Type I (Figure 5-15). Each Haversian system consists of a central
collagen) and 65% inorganic (calcium hydroxyapatite crys- Haversian canal with its neurovascular bundle. The bony
tals) materials, with most of the remainder being water. The matrix is arranged in concentric rings around the canal
collagen provides some flexibility and tensile strength to the (concentric lamellae) with osteocytes occupying lacunae at
bone, whereas the inorganic materials confer hardness. their junctions. Osteocytes maintain contact with other os-
There are four major cells of bone tissue. These are osteo- teocytes by way of cellular processes that pass through tiny
progenitor cells, osteoblasts, osteocytes, and osteoclasts. canaliculi. In this fashion, nutrients and oxygen from the
Osteoprogenitor cells (Figure 5-8) are derived from blood vessel can diffuse from cell to cell outward toward
mesenchymal cells and have the ability to differentiate into the periphery of the Haversian system. A calcified cement
osteoblasts, and under some circumstances, into chondro- line encircles the outer margin of the Haversian system.
genic cells. They are elongated cells and are found associ- Spongy bone (Figure 5-16) is found in the interior of
ated with periosteum and endosteum (see below). bones. It is made of a delicate network of bony trabeculae
Osteoblasts (Figure 5-8) develop from osteoprogenitor that are arranged to strengthen the bone according to the
cells and are therefore also found lining external and inter- mechanical loads placed on it. Irregular arrangements of
nal bone surfaces, but they have more of a cuboidal shape. lamellae are present, but Haversian systems are not.
Osteoblasts are actively involved in secreting collagen-rich
osteoid, the precursor to true bone matrix during bone
growth, repair and remodeling. Osteoid is subsequently cal- Bone Growth
cified to make the bony matrix. There are two basic mechanisms of bone growth. In intra-
Osteocytes (Figure 5-8) are osteoblasts that have become membranous ossification, bone forms within a mesenchy-
surrounded by matrix. They occupy lacunae in the matrix mal membrane. In endochondral ossification, bone tissue
and are responsible for maintaining matrix composition. replaces a hyaline cartilage model. In both mechanisms,
Osteoclasts (Figures 5-8 and 5-9) are large, multinucle- bone is initially formed as spongy bone.
ate cells responsible for bone resorption; that is, they de- Intramembranous ossification (Figure 5-17) is responsi-
stroy bony matrix. They are often found associated with a ble for producing the flat bones of the skull. The process is
pit in the bone called a Howship’s lacuna. Osteoclasts are relatively straightforward.
active during the growth, repair, and remodeling processes. 1. Mesenchymal cells differentiate into osteoblasts at the
Their activities are balanced with those of osteoblasts so primary center of ossification and begin depositing
that (normally) bone mass remains constant. bony matrix to form trabeculae of spongy bone.
2. Ossification continues radially from the ossification
Membranes of Bone
center, like ripples spreading from dropping something
Bone surfaces are lined with one of two fibrous mem- in water.
branes: periosteum or endosteum. Periosteum (Figure 5-10)
3. Bone marrow develops in the spaces between trabeculae.
is the fibrous membrane that surrounds bone. Like its
counterpart in cartilage, periosteum has a superficial fi- 4. Periosteum and endosteum develop from the unossified
brous layer and a deeper cellular layer. The cellular layer mesenchyme membrane.
has osteoprogenitor cells that have the ability to become 5. The surfaces are remodeled to form compact bone. In
bone-forming cells. Periosteum is attached to bone by the skull, these layers are called inner and outer tables.
Sharpey’s fibers, collagen bundles that penetrate the bone The spongy bone between them is the diploë.
matrix (Figure 5-11). All internal surfaces are lined with a Endochondral ossification is involved in producing
reticular connective tissue called endosteum, which is also most bones of the skeleton, and is most easily understood
associated with osteoprogenitor cells (Figure 5-12). by studying appendicular long bones. While studying this
process, it is important to remember that the cartilagenous
Types of Bone Tissue model grows in length and diameter even as parts of it are
Bone tissue is of two types: spongy (cancellous) bone and being replaced by bone. (After all, the skeleton must be
compact bone (Figure 5-13). Both types are in all bones, functional and an appropriate size at all times during
C H A P T E R F I V E : C A R T I L A G E A N D B O N E
47
embryonic growth.) Endochondral ossification is outlined in 6. Growth in diameter occurs as periosteal bone is de-
Figure 5-18 and can be summarized by the following steps: posited on the surface while osteoclast activity removes
1. Hyaline cartilage occupies the zone of reserve cartilage bone from the marrow cavity side of the diaphysis (Fig-
and acts as a source of cartilage to undergo the process ure 5-22).
described below. 7. All bone tissue initially forms as spongy bone, so it
2. Normal chondrocytes multiply in the zone of prolifera- must be remodeled on the surface to become compact
tion (multiplication). bone. The process involves deposition of the lamellae
3. The chondrocytes enlarge (hypertrophy) in the zone of from outermost to innermost around the neurovascular
hypertrophy. bundle, which ends up in the remaining space, the
Haversian canal (Figure 5-23).
4. The cartilage matrix calcifies in the zone of calcifica-
tion. 8. At about the time of birth, secondary centers of ossifi-
cation appear in one or both ends (epiphyses) of the
5. Chondrocytes deteriorate and bone is deposited on the
long bone (Figure 5-24). These ossify the epiphyses
remaining fragments of calcified matrix in the zone of
leaving cartilage in two places: on the surface of the
ossification.
epiphysis as articular cartilage, and in the epiphyseal
6. Osteoclasts remove the newly formed bone to make the plate that joins the epiphysis with the diaphysis (Figure
marrow cavity in the zone of erosion. 5-25). The epiphyseal plate continues to produce new
Examine the photomicrographs in Figures 5-19 to 5-25 cartilage on its epiphyseal side at the same rate it is
as you read the details of this process. being replaced by bone on the diaphyseal side. In this
1. Endochondral ossification begins with a bone made of fashion, the plate remains the same thickness, but ap-
hyaline cartilage. A primary center of ossification (Fig- pears to grow away from the center of the bone. This
ure 5-19a) appears at the center of the shaft (diaphysis). results in longitudinal growth of the bone. At some time
This is evidenced by the chondrocytes of the region during development (typical for each bone), the epiphy-
multiplying and enlarging at the expense of the matrix. seal plate ossifies and longitudinal growth is completed.
What matrix is left becomes calcified and appears more
basophilic. Since the calcified matrix retards diffusion Synovial Articulations
of oxygen and nutrients, chondrocytes begin to deterio- Synovial joints (Figure 5-26) are characterized by having a
rate within the enlarged lacunae. synovial (joint) cavity. This makes them freely moveable.
2. Simultaneous with the activities of the primary ossifica- Lining the bones’ surfaces is articular cartilage. It is made
tion center, osteoblasts develop deep to the perichon- of hyaline cartilage (from the original cartilage model that
drium of the shaft and deposit a ring of periosteal bone remained unossified) without a perichondrium. Lining the
called a bony collar (Figure 5-19a). The bony collar cavity is a vascular and fibrous synovial membrane that se-
acts as a splint to support the bone as the internal carti- cretes a lubricating synovial fluid (Figure 5-27). Strengthen-
lage is eroded away. The perichondrium develops into a ing the joint from the outside is a fibrous joint capsule that
periosteum. is continuous with the periosteum of the articulating bones.
The capsule may contain thickenings that act as ligaments,
3. Blood vessels (periosteal buds) grow into the enlarged
or the ligaments may be separate from the capsule. Other
lacunae and form primary marrow spaces (Figures 5-19b
cartilages (generally fibrocartilage) may be associated with
and 5-19c). They bring osteoprogenitor cells into the
the interior of the joint. The menisci of the knee joint are an
region, some of which become osteoblasts that deposit
example.
osteoid and then bone on the remaining cartilagenous
fragments. Muscular Attachments to Bone
4. After a region has ossified, osteoclasts move in and re- Skeletal muscles either attach to bone directly or by way of
sorb the bone to form the marrow cavity (Figures 5-19d a tendon. In the first instance, the muscle fibers extend to
and 5-19e). This will not occur during endochondral the periosteum and the connective tissue components of the
ossification of short or irregular bones that lack a mar- muscle blend in with the connective tissue fibers of the pe-
row cavity. riosteum (Figure 5-28). The periosteum itself is attached to
5. The process of bone replacing cartilage spreads toward the bone by Sharpey’s fibers, and these are especially sub-
the ends of the bones (epiphyses), following the same stantial at the point of muscular attachment. If the muscle
sequence of events as occurred in the primary center: attaches by way of a tendon, then it is the tendon fibers
enlargement and multiplication of chondrocytes, calci- that mix with the periosteal fibers.
fication of matrix, deterioration of chondrocytes, and
deposition of bone (Figures 5-20 and 5-21).
48
Cb
TM
Cc
IM
5-1 5-2
THE CELLS OF CARTILAGE Chondroblasts (Cb), isogenous groups of CARTILAGE MATRIX Territorial matrix (TM) around the chondrocytes
chondrocytes (Cc), and some empty lacunae (L) (artifacts of prepa- stains darker than the interterritorial matrix (IM) between lacunae.
ration) are visible in this micrograph of hyaline cartilage. (X250) This specimen is hyaline cartilage. (X250)
5-3a 5-3b
PERICHONDRIUM The vascular fibrous membrane around cartilage is called perichondrium (P). (a) The pink layer is the perichondrium surrounding
hyaline cartilage. The distinction between the outer fibrous layer and inner chondrogenic layer is not sharp. (X125) (b) This hyaline cartilage speci-
men has a thicker perichondrium. The outer fibrous (F) and inner chondrogenic (C) layers are visible, but still are not sharply separated. (X125)
49
Cc
Cc
5-4a 5-4b
HYALINE CARTILAGE The most common form of cartilage is hyaline cartilage. Its matrix presents a smooth appearance even though collagen
fibers are present. Perichondrium (P) and chondrocytes (Cc) are visible in both specimens. Specimen (a) is from the trachea. (X65) Specimen (b) is
a section of the nasal septum. (X250)
AC
AC
AC
5-5a 5-5b
ARTICULAR CARTILAGE The hyaline cartilage covering joint surfaces lacks a perichondrium. (a) This bone has been removed from its joint during
preparation, but the articular cartilage (AC) on its joint surface is apparent. (X65) (b) This micrograph shows two articulating bones with their
articular cartilages. (X65)
50
Cc
P
5-6a
5-6b
ELASTIC CARTILAGE The framework of the

ear and epiglottis is made of elastic cartilage.
A silver stain was used to highlight the
elastic fibers in the top two specimens.
Cc Notice the perichondrium (P) and relatively
large chondrocytes (Cc). Specimen (a) is
from the external ear. (X210) Specimens (b)
and (c) are from the epiglottis (X100 and
5-6c X210, respectively).
51
CF
CF
Cc
Cc
5-7a 5-7b
FIBROCARTILAGE Intervertebral disks and the pubic symphysis are made of fibrocartilage. It is characterized by dense bundles of collagen fibers
(CF) in its matrix and the absence of a perichondrium. Often the chondrocytes (Cc) occur in lines, as in (b). Both specimens are from inter-
vertebral disks and were magnified. (X130)
Op
Oc
Oc
Ob
Ob
Ocl
Ocl
5-8a 5-8b
BONE CELLS These two micrographs show the four types of bone cells. Osteoprogenitor cells (Op) and osteoblasts (Ob) are found in rows on the
surface of bone, associated either with endosteum or periosteum. Osteoprogenitor cells are flatter than osteoblasts. Osteocytes (Oc) are cells of
mature bone and are encased by the bony matrix. Osteoclasts (Ocl) are large, multinucleate cells found on the surface of bone. Micrograph (a)
was magnified X250. Micrograph (b) was magnified X380.
52
Ob
Ocl
*
P
HL
HL
5-9 5-10
OSTEOCLASTS Bone is resorbed by multinucleate osteoclasts (Ocl). PERIOSTEUM Periosteum (P) is the fibrous membrane around bone (B).
They often erode a pit in the surface of the bone called a Howship’s It has an outer fibrous layer and an inner osteogenic layer with osteo-
lacuna (HL). (X380) progenitor or osteoblast (Ob) cells. In this specimen, the space (*) is an
artifact of preparation. (X250)
SF
B
E
5-11 5-12
SHARPEY’S FIBERS Periosteum (P) is anchored to the bone (B) by collagen ENDOSTEUM All inner surfaces of bone (B) are lined with a delicate
bundles called Sharpey’s fibers (SF). In this micrograph, they can be connective tissue called endosteum (E). In this specimen, the endo-
seen to penetrate the bony matrix (arrows). (X380) steum is associated with osteoprogenitor cells. (X250)
53
VC
HC
5-13 5-14
TYPES OF BONE TISSUE Dense, compact bone covers outer surfaces, THE CANALS OF COMPACT BONE The blood vessels of compact bone
whereas the interior of a bone is composed of more delicate spongy travel through canals. Volkmann’s canals (VC) carry vessels from the
bone tissue. Both types are found in all bones, but in different surface to the interior. Haversian canals (HC) transmit blood vessels
proportions. (Photograph by Gary D. Wisehart) parallel to the surface. In this decalcified bone specimen cut in cross
section, the Volkmann’s canals are cut longitudinally or obliquely,
whereas the Haversian canals are cut in cross section. Osteocytes are
barely visible as white specks in the matrix. (X100)
CL
Oc
Ca
L CL HC
Oc HC
Ca
5-15a 5-15b
COMPACT BONE IN CROSS SECTION (GROUND BONE PREPARATION) Due to the hardness of compact bone matrix, these specimens were not sectioned
with a microtome, but instead were ground down to their final thickness. The osteocytes and blood vessels often are absent from preparations
such as these. The central Haversian canals (HC), concentric lamellae (L), osteocytes (or at least their lacunae) (Oc), canaliculi (Ca), and cement
lines (CL) are visible in both specimens. (X100)
54
M
T
T Oc
T
T
Ob
5-16 5-17a
SPONGY BONE Trabeculae (T) of spongy bone contain osteocytes and
lamellae of bone, but these are not organized into Haversian systems.
M
(X50)
Ob
Oc
T
M
HC
T
Ob
5-17b
Oc
T
Ob M
M
T
5-17c
Oc
INTRAMEMBRANOUS OSSIFICATION Intramembranous ossi-
fication occurs within a mesenchymal membrane. Trabe-
culae of bone (T), osteoblasts (Ob), and osteocytes (Oc) are
identifiable, as is the membrane (M) in each specimen. (a) 5-17d
This is a fairly advanced specimen from the skull. The outer
and inner surfaces will be remodeled into compact bone called the inner and outer tables. The internal spongy bone will be called
diploë. (X50) (b) This membrane bone also has yet to produce the surface compact bone, but the trabeculae are well-formed. (X100)
(c) This specimen is from a section of the entire skull of an embryonic vertebrate. The membrane bone is orange-red and has the
characteristic osteoblasts on its surface and the trapped osteocytes within. The white space at the left is part of the nasal cavity, and the
purple curled structure is hyaline cartilage (HC) of a nasal concha. (X50) (d) This is a frontal section of an embryonic skull. The centers
of ossification and membrane are visible. Notice the osteoblasts on the surface that are partially surrounded by bone. The hyaline
cartilage of the nasal septum is the blue object at the center top. (X100)
55
Articular
cartilage
Epiphyseal
line
Secondary Epiphyseal
center of plate
ossification
Hyaline
cartilage
model
Marrow
Primary cavity
center of
ossification
bony collar
Regular cartilage
Replacement cartilage
Bone
Marrow
5-18
ENDOCHONDRAL OSSIFICATION IN A LONG BONE This complex process starts with a hyaline cartilage model of the bone. The cartilage is replaced
by bone tissue as the bone grows in size. See page 47 for details.
56
BC
Cc
Cc
MC
BC
5-19a 5-19b
P
P
BC
Cc
MC
HC
5-19d
5-19c ENDOCHONDRAL OSSIFICATION AT THE PRIMARY CENTER SHOWN IN
CROSS SECTION While they are from different bones, this collection of
images should allow the reader to piece together the dynamic process
of endochondral ossification. All images were magnified X50. (a) The
primary center of ossification has become active, as evidenced by the
enlarged chondrocytes (Cc) at the center and the pink bony collar (BC)
produced by subperiosteal osteoblasts. The periosteum (P) is also
visible. Notice that the hyaline cartilage at the periphery still looks like
typical hyaline cartilage. (b) The chondrocytes and matrix in the bone’s
center have begun to break down and osteoblasts and other marrow
cells are visible in the primitive marrow cavity (MC). (c) The marrow
cavity has gotten larger, the bony collar has gotten more complex and
the chondrocytes at the periphery have enlarged. That is, they’ve
reached the stage the interior chondrocytes were at in micrograph (a).
(d) In this micrograph, very little cartilage (HC) remains in the bone’s
center, having been replaced with spongy bone. The surface bone on
the left has begun remodeling into compact bone (note the circular
regions that will become Haversian systems—see arrows). (e) The
marrow cavity has been formed in this specimen through osteoclast
activity on the spongy bone recently produced.
5-19e
57
PO
PC
PO PB
PB Ob
B
CC
CC
B
E
O C H P R O C H P
5-20a 5-20b
ENDOCHONDRAL OSSIFICATION IN LONGITUDINAL SECTION The zones of endochondral ossification are visible in both specimens, which are oriented
so that the process is further along toward the left. Zones of reserve cartilage (R), proliferation (P), hypertrophy (H), calcification (C), ossification
(O), and erosion (E) are visible, as are the periosteum (PO), perichondrium (PC), and periosteal bone (PB). (a) Bone tissue is pink in this specimen
(see periosteal bone for the shade) as is calcified cartilage (which is somewhat paler). Notice how the spaces produced by deteriorated chondro-
cytes allow entry of osteoblasts (Ob) that deposit new bone (B) on the remaining slivers of calcified cartilage (CC). (X100) (b) The calcified
cartilage (CC) is noticeably darker than the normal matrix in this specimen. Bone (B) is a deep pink. (X100)
CC
CC
B
B
Ob
5-21b
5-21a Ob
B
DETAILS OF ENDOCHONDRAL OSSIFICATION (a) Chondrocytes enlarge CC
and multiply at the expense of matrix. When they die from lack of
oxygen in the calcified matrix (CC), there is little of the original
cartilage that remains. Bone (B) is deposited on the slivers of calcified
cartilage. Osteoblasts (Ob) have aligned themselves in rows on the
surface of the growing bone, a very distinctive feature of them. (X200)
(b) This image is of the zone of ossification. Bone tissue (B) is deep Ob
O
pink and calcified cartilage (CC) is lavender. Osteoblasts are visible in
their characteristic rows on the bone’s surface. (X200) (c) In this
specimen, bone is orange-red and calcified cartilage is light blue. The
blue just deep to the osteoblasts is osteoid (O), the precursor to true
bone matrix. (X320)
5-21c
58
Ob
P
Ob
5-22 5-23
PERIOSTEAL BONE While cartilage is being replaced by bone in the SURFACE REMODELING Bone tissue is produced as spongy bone. The
interior, osteoblasts (Ob) deep to the periosteum (P) deposit bone on compact bone on bone surfaces is the result of remodeling spongy bone.
the outer surface of the bone at the same rate that osteoclasts remove Developing Haversian systems (H) are visible in this specimen. Osteo-
bone from the marrow cavity. These coordinated activities result in blasts (Ob) deposit concentric lamellae in the open region until all that
circumferential growth of the bone. (X130) remains is the Haversian canal with its neurovascular bundle. (X250)
SC
SC
5-24a 5-24b
SECONDARY CENTER OF OSSIFICATION In long bones and some other bones, a secondary center (SC) of ossification opens in the epiphysis some
time after birth. The secondary center completely ossifies the epiphysis except for two regions that remain hyaline cartilage: the epiphyseal plate
that connects the epiphysis and diaphysis, and the articular cartilage. Both micrographs are X25 and show early activity in the secondary center.
59
EP
EP
5-25a 5-25b
EPIPHYSEAL PLATE After birth, longitudinal growth in long bones is due
to the activity of the epiphyseal plate (EP) of hyaline cartilage that joins
the epiphysis with the diaphysis. All the zones of cartilage being
replaced by bone are compressed into the epiphyseal plate; all are
there, but there may be some overlap. New cartilage is formed on the
epiphyseal side at the same rate as cartilage is replaced by bone on the
diaphyseal side of the plate. (a) New bone is forming on the lower side
of this epiphyseal plate, so the epiphysis (E) is being pushed upward in
this micrograph. (X50) (b) The epiphysis is toward the upper right in
this micrograph. Bone is red, reserve cartilage is light blue, and
calcified cartilage is lavender. (X100) (c) The epiphysis is toward the
right in this specimen. The stain used does not allow easy differ-
entiation of reserve cartilage, calcified cartilage, and bone. (X100)
EP
5-25c
SM
AC
FC
SYNOVIAL JOINT Synovial joints are freely movable. The bones

involved are held together with a fibrous articular capsule (FC).
The joint cavity is lined with a vascular synovial membrane (SM)
that secretes a lubricating synovial fluid. Also notice the articular
5-26 cartilage (AC) lacking a perichondrium. (X100)
60
SC
AC
SM
5-27
SYNOVIAL MEMBRANE The source of synovial fluid is the vascular and fibrous synovial membrane
(SM). Articular cartilages (AC) of the two bones and the synovial cavity (SC) are also visible.
(X250)
SF
B
P
SM
SF
5-28
MUSCULAR ATTACHMENT TO BONE In this specimen, the connective tissue of the skeletal muscle
(SM) blends with the periosteum (P), which is in turn attached to the bone (B) with Sharpey’s
fibers (SF). Notice that in this specimen, all tissues are pink, so identification relies on differing
textures and distribution of nuclei. (X125)
Blood and Bone Marrow
C H A P T E R
6
Introduction to Blood for 44% and 1%, respectively. The following are typical
Blood functions as a transport medium between organs spe- cell densities for each (Figure 6-2):
cialized for contacting and exchanging materials with the ◗ Red blood cells: 4,500,000 and 5,000,000 RBCs per
environment and the remainder of cells buried in the body. cubic millimeter of blood in females and males, respec-
It transports oxygen from the lungs to other body cells and tively
carbon dioxide from these same cells back to the lungs. It ◗ White blood cells: between 6,000 and 10,000 (average
distributes nutrients absorbed by the small intestines and 8,000) per cubic millimeter of blood
stored in the liver throughout the body, and it picks up ◗ Platelets: between 150,000 and 300,000—it is difficult
wastes and transports them to the kidneys for excretion. In to get an accurate count—per cubic millimeter of blood.
addition, it is involved in transporting cells of the body’s
Erythrocytes (Figure 6-3) are biconcave discs with a di-
defenses to sites where they are needed and distributing
ameter of about 7 to 8 µm. They develop from cells in bone
heat and hormones throughout the body.
marrow, lose most organelles (including their nucleus) dur-
Blood is a specialized connective tissue, with cells dis-
ing maturation, and are little more than bags of the red
persed in a fluid intercellular material called plasma. Since
oxygen-carrying pigment hemoglobin (plus a few soluble
some blood cells are not actually cells when functional, the
enzymes) when functional in the blood. Their shape in-
“cellular” portion is said to be made of formed elements.
creases surface area for oxygen exchange and makes them
pliable enough to fit through the smallest blood vessels (3
to 4 µm in diameter).
Leukocytes differ from erythrocytes in a number of
Formed Elements of Blood ways: they are larger, nucleated, and while found in the
When blood is spun in a centrifuge tube, the plasma sepa- blood during transport, usually function outside the blood.
rates from it and is found on top of the formed elements They are divided into two major groups: granulocytes and
(Figure 6-1). Comprising the majority of the formed ele- agranulocytes. Granulocytes have prominent cytoplasmic
ments is a red layer made up of erythrocytes (red blood granules whose staining properties form the basis for differ-
cells, RBCs). On top of the erythrocytes is the narrow buffy entiating the three major types (neutrophils, eosinophils, and
coat layer made of leukocytes (white blood cells, WBCs) basophils—see page 62). In addition, they have multilobed
and platelets (thrombocytes). The formed elements consti- nuclei (which leads to the other name for this group: poly-
tute 45% of the blood volume, with plasma making up the morphonuclear granulocytes) and are formed in the bone
other 55%. The erythrocyte and buffy coat layers account marrow. The characteristics of each granulocyte follows.
61
62
Neutrophils (Figure 6-4) are the most abundant WBC, Platelets (Figure 6-10) are cell fragments derived from
accounting for between 60% and 70% of all WBCs in megakaryocytes in bone marrow. They are involved in the
blood. They are slightly larger than RBCs (9 to 12 µm in di- clotting process.
ameter) and have a multilobed nucleus with the lobes often
joined by thin nuclear strands. Their cytoplasmic granules Composition of Bone Marrow
are unstained or are slightly lavender in standard blood Marrow is found in the marrow cavity of long bones and fills
smear preparations (Wright’s or Giemsa stains). Function- spaces between trabeculae in spongy bone of all bones (Fig-
ally, they are phagocytic cells. ure 6-11). It is a highly vascular tissue that contains numer-
Eosinophils (Figure 6-5) comprise less than 5% of all ous and connected blood sinusoids, a framework of reticular
WBCs in the blood. They are somewhat larger than neu- fibers, and cells of various types. Red bone marrow is found
trophils (10 to 14 µm in diameter) and have a two-lobed nu- in all fetal bones, but as the individual ages, its distribution is
cleus. Their cytoplasmic granules stain red in standard blood limited to ribs, sternum, vertebrae, and flat bones of the
smears. They are active in combating parasitic infections, skull. Its primary function is hemopoiesis (formation of
phagocytose antigen-antibody complexes, and are present blood cells—see below). Yellow bone marrow (Figure 6-12)
during allergic reactions, apparently to temper the response. replaces red bone marrow during aging, although it main-
Basophils (Figure 6-6) are rare in the blood, comprising tains the ability to revert to red marrow if the need for more
less than 1% of all WBCs. They are slightly larger than blood cells arises. It contains abundant fat cells.
RBCs (8 to10 µm in diameter) and have prominent, dark-
purple cytoplasmic granules that make seeing the S-shaped Postnatal Development of Blood Cells
nucleus difficult. They are involved in the inflammatory re- Hemopoiesis is the production of blood cells. According to
sponse and like mast cells, may be involved in hypersensi- current evidence, this process begins in the bone marrow
tivity reactions. with pluripotent stem cells that divide to give rise to five
Agranulocytes have unlobed nuclei and lack prominent different types of unipotent stem cells, each of which is re-
cytoplasmic granules. (All WBCs have lysosomes that appear sponsible for producing a particular type of blood cell.
as faint cytoplasmic granules.) Lymphocytes and monocytes Granulopoiesis, illustrated in Figure 6-13, is the pro-
are the two types of agranulocytes. duction of granulocytes. The unipotent stem cell in this se-
Lymphocytes (Figure 6-7) account for between 20 and ries is actually bipotent, since it also can progress through
25% of all WBCs in blood. They are slightly larger than the monocyte series. Each granulocyte type goes through
RBCs (8 to 10 µm in diameter) and have a round nucleus similar developmental stages, and all begin with undifferen-
that fills most of the cell, leaving only a thin ring of cyto- tiated myeloblasts followed by promyelocytes, which con-
plasm at the cell’s periphery. There are three functionally tains primary (nonspecific) cytoplasmic granules. When a
distinct, but morphologically indistinguishable, types of promyelocyte develops specific granules characteristic of
lymphocytes. These are B cells, T cells, and null cells. B each granulocyte it becomes a myelocyte. Metamyelocytes
cells differentiate into antibody-secreting plasma cells when are smaller and have an indented nucleus. Stab (band) cells
exposed to the proper antigen (Figure 6-8). They are respon- follow and are the last stage prior to a mature granulocyte.
sible for the humoral immune response. T cells do not secrete Stab cells are characterized by a U-shaped to slightly seg-
antibodies in response to antigen, but they do have antigen mented nucleus. (By convention, if the nuclear indentation
receptors in their membranes. Depending on the type of T is less than half the nuclear diameter, the cell is a metamye-
cell, it may initiate a cell mediated immune response as a cy- locyte. If more than half of the nuclear diameter, it is a stab
totoxic T cell, or it may regulate activity of other cells either cell.) Mature granulocytes have the segmented or lobed nu-
as a T-helper or T-suppressor cell. Null cells are so-named cleus and appear as they do in blood. Examples of develop-
because they lack the membrane markers that identify T and mental stages are shown in Figure 6-14.
B cells. Natural killer (NK) cells are null cells that kill foreign Monopoiesis is the production of monocytes (Figure
or infected cells without antigen-antibody interaction. 6-15). Monocytes develop from the same stem cell as gran-
Monocytes (Figure 6-9) comprise between 3% and 8% of ulocytes and pass through monoblast and promonocyte
all blood WBCs. They are about twice the size of RBCs (12 to (Figure 6-16) stages before the mature monocyte is formed.
15 µm in diameter). The cytoplasm is bluish-gray in stan- During development, the cells become smaller and the nu-
dard blood smears and the nucleus is often indented. Mono- clear indentation becomes more prominent. Both of these
cytes are the blood form of tissue macrophages. In addition to are rare in bone marrow smears.
phagocytic activity, some macrophages act as antigen- Lymphopoiesis is the production of lymphocytes (Figure
presenting cells (APCs) that phagocytose and digest foreign 6-17). Lymphoblasts and prolymphocytes are the develop-
material, then carry the antigen on their own surface to mental stages a lymphocyte passes through prior to matu-
“show” cells of the immune system and stimulate a response. rity. These are differentiated primarily by size, with the cells
C H A P T E R S I X : B L O O D A N D B O N E M A R R O W
63
becoming smaller with maturity, but they are rarely seen in erythroblast has a pinkish cytoplasm and a small, densely
normal bone marrow. A mature lymphocyte is shown in staining nucleus. The final developmental stage before the
Figure 6-16. cell is a mature RBC is the reticulocyte. It is anucleate and
Erythropoiesis is the process of RBC formation and the appears very similar to RBCs except its cytoplasm contains
sequence is shown in Figure 6-18. Examples of these cells a bluish network when stained with brilliant cresyl blue.
are shown in Figure 6-19. It begins with the proerythro- Thrombopoiesis (Figure 6-20) is the production of
blast, a large cell with a prominent nucleus, little basophilic platelets. Megakaryocytes are derived from a unipotent
cytoplasm, and no hemoglobin. The proerythroblast devel- stem cell, which undergoes several mitotic divisions as a
ops into a series of erythroblasts (normoblasts) that show a megakaryoblast without cytokinesis. The resulting mega-
progressive decrease in size, a loss of organelles (along with karyocyte is a large (up to 100 µm in diameter), polyploid
a loss of basophilic staining), and an increase in hemoglo- cell with a lobed nucleus. It produces platelets by cytoplas-
bin (along with increasing eosinophilia). The stages are ba- mic fragmentation. These cells are shown in Figures 6-21
sophilic erythroblast, polychromatophilic erythroblast, and and 6-22.
orthochromatophilic erythroblast. The orthochromatophilic
Plasma
Buffy coat
RBCs
6-1 6-2
HEMATOCRIT This capillary tube contains blood that has been BLOOD SMEAR Blood is usually examined as a smear on a slide stained with
centrifuged to separate the plasma (above) from the formed either Giemsa (as in this preparation) or Wright’s stain. RBCs outnumber
elements (below). A thin buffy coat (composed of white blood WBCs about 1000 to 1. Two WBCs are visible in this field (arrows). (X265)
cells and platelets) is visible at the RBC and plasma junction.
64
6-4a 6-4b
6-4c 6-4d
NEUTROPHILS Neutrophils are granulocytes with a pinkish to gray
cytoplasm. They are the most abundant WBC and are slightly larger
than RBCs. Mature neutrophils have a segmented nucleus (as in micro-
6-3 graphs a, b, and c). Immature neutrophils have an unsegmented nucleus
and are called band cells (as in micrograph d). About 30% of neutro-
RED BLOOD CELLS The biconcave disk shape of the RBCs is visible phils in blood samples from females demonstrate a “drumstick” pro-
in some cells in this Giemsa preparation. RBCs are very sensitive to truding from the nucleus, as in (c). This is the region of the inactive
osmotic conditions and their shape may change if the staining con- X chromosome. Micrograph (a) was stained with Giemsa stain; the
ditions are not isosmotic. (X660) others were prepared with Wright’s stain. All photos are X660.
6-5a 6-5b 6-6a 6-6b

EOSINOPHILS These granulocytes are relatively rare and are about BASOPHILS Basophils comprise only about 1% of all WBCs. They are
twice the size of RBCs. Their cytoplasmic granules stain red, and their slightly larger than RBCs and have dark purple cytoplasmic granules
nucleus usually has two lobes. Micrograph (a) was prepared with that obscure the nucleus. Both micrographs are X660. Micrograph (a)
Giemsa stain; (b) was prepared with Wright’s stain. Both were was prepared with Giemsa stain and (b) was prepared with Wright’s
magnified X660. stain.
65
6-7a 6-7b 6-7c

LYMPHOCYTES Lymphocytes are common in the blood, comprising up to 25% of all WBCs. Most are about the size of RBCs and have only a
thin halo of cytoplasm encircling their round nucleus. They belong to functional groups called B cells and T cells, which are morphologically
indistinguishable. Micrographs (a) and (b) are small lymphocytes and were prepared with Giemsa and Wright’s stains respectively. Some
lymphocytes are larger, as in micrograph (c). These are natural killer (NK) cells or some other type of “null” cell. A neutrophil is also seen in (c).
All micrographs are X660.
PLASMA CELLS B lymphocytes develop into plasma

cells when stimulated by the appropriate antigen.
Plasma cells secrete protective antibodies against
the antigen. These plasma cells (arrows) are
recognizable because of their elongated shape,
eccentric nucleus with “clock face” chromatin, and
a pale region near the nucleus (which is the site of
a Golgi apparatus). This specimen is from the
6-8 colon. (X660)
6-9a 6-9b 6-10a 6-10b

MONOCYTES Monocytes are the blood form of macrophages. They are PLATELETS Platelets (arrows) are cell fragments involved in clotting.
about twice the size of RBCs and have a round or indented nucleus. (a) Giemsa stain. (b) Wright’s stain. Both micrographs are X660.
Both micrographs are X660. (a) Giemsa stain. (b) Wright’s stain.
66
T S
6-11a 6-11b
RED MARROW The spaces between trabeculae of spongy bone (T) are often filled with red marrow. It consists of various developmental stages of
blood cells. (a) X50. (b) Blood sinusoids (S), marrow cells, and two megakaryocytes (arrows) are visible. (X250)
BC
6-12a 6-12b
YELLOW MARROW In most bones, red marrow is replaced by yellow marrow in the adult. The majority of cells are adipocytes. (a) X50.
(b) Adipocytes (A), sinusoids (S), and some remaining blood cells (BC) are seen. (X250)
67
nB nMM
nB
P
nM
P
6-14a 6-14b
nM
eM
nB
nB
eMM
6-14c 6-14d
6-13
GRANULOCYTE DEVELOPMENT
Through the promyelocyte stage, nS
the precursors of neutrophils, baso-
phils, and eosinophils are indistin-
guishable. Once myelocytes begin
accumulating granules specific to nB nMM
each granulocyte, each develop-
mental series can be traced separ-
ately. The overall pattern involves eS
accumulation of cytoplasmic
granules and progressive inden-
tation and segmentation of the
nucleus.
6-14e
BONE MARROW SMEARS ILLUSTRATING GRANU-
LOCYTE DEVELOPMENT These marrow smears nB
illustrate most of the cells in the granulocytic
developmental series. Some cells will be
encountered frequently (e.g., neutrophilic
cells) whereas others are much less common B
(e.g., basophils). (a through f X660) Key to
the symbols used: MB 4 myeloblast, P 4
promyelocyte, nB 4 neutrophilic band,
nM 4 neutrophilic myelocyte, nMM 4
neutrophilic metamyelocyte, nS = neutro-
philic segmented, eM 4 eosinophilic myelo- nB
cyte, eMM 4 eosinophilic metamyelocyte,
eS 4 eosinophilic segmented, B 4 basophilic
myelocyte.
6-14f
68
L
Monoblast
PM
PM
6-15 6-16 6-17

MONOCYTE DEVELOPMENT BONE MARROW SMEAR ILLUSTRATING MONOCYTE LYMPHOCYTE DEVELOPMENT
The unipotent stem cell is AND LYMPHOCYTE DEVELOPMENT The precursors Lymphocytes go through
the same cell that gives rise of lymphocytes and monocytes are relatively rare only two stages in develop-
to the granulocytic cells. in marrow smears. Shown here are a promono- ment in marrow. Unlike
Cells of monocyte develop- cyte (PM) and lymphocyte (L). Early stages of other blood cells, they
ment are difficult to find in lymphocyte development are often difficult to continue their development
marrow smears. tell apart. If there is doubt as to a cell’s identity, outside the marrow in
by convention it is just called a lymphocyte. lymphoid tissue throughout
(X660) the body.
6-18
ERYTHROCYTE DEVELOPMENT
The major changes in
erythrocyte development are
a decrease in size, loss of
organelles (most notably, the
nucleus) and an increase in
cytoplasmic hemoglobin.
oE
pE oE
pE
oE
proE
proE
6-19a 6-19b 6-19c

BONE MARROW SMEARS ILLUSTRATING ERYTHROCYTE DEVELOPMENT (a through c X660) Key to symbols used: proE 4 proerythroblast, pE 4
polychromatophilic erythroblast, oE 4 orthochromatophilic erythroblast. Note the oE extruding its nucleus in (b) and the dividing oE in (c).
Both are indicated with arrows.
69
6-20
THROMBOCYTES (PLATELET)
DEVELOPMENT Megakaryoblasts
6-21a
develop into megakaryocytes
by repeated mitotic divisions
without cytokinesis. Once formed,
megakaryocytes undergo fragmen-
tation to produce hundreds of
thrombocytes.
6-21b
BONE MARROW SMEARS ILLUSTRATING THROMBOCYTE DEVELOPMENT (a) Megakaryoblast with
a single, unlobed nucleus. (X400) (b) The characteristic lobed nucleus in this megakaryocyte
is visible. (X400)
70
6-22
SECTION OF BONE MARROW Because of its size, this megakaryocyte (arrow) stands out against the
marrow cells. (X265)
Muscle Tissue
C H A P T E R
7
Introduction to Muscle Tissue known as the sliding filament theory. In this model, the
Muscle tissue is a highly cellular and vascular tissue special- contractile unit is the region between two Z discs and is
ized for contraction. Contraction of muscle results in move- called a sarcomere. During contraction, the Z discs are
ment of a body part, propulsion of a substance, or in some brought closer together. As the actin and myosin filaments
instances, stopping movement of a substance. slide across one another, the I bands become narrower but
There are three basic types of muscle tissue: skeletal the A bands stay the same width.
muscle, cardiac muscle, and smooth (visceral) muscle. The Skeletal muscle tissue is combined with connective tis-
cells of all muscle tissues are long and thin, and are referred sues to form specific muscle organs, like the biceps brachii
to as muscle fibers. or gluteus maximus. The connective tissue components of
a muscle weave between the muscle fibers and bind the
whole organ together. Covering the entire muscle is epimy-
Characteristics of Skeletal Muscle sium (Figure 7-5), a dense irregular collagenous tissue layer.
Skeletal muscle (Figure 7-1) is associated with the bones of Smaller extensions of epimysium penetrate the muscle as
the skeleton and is responsible for movement of the body. It perimysium and surround several fibers forming a fascicle.
also may be found associated with the skin (as in the muscles Extending from the perimysium is a fine layer of reticular
of facial expression) and with some viscera (the proximal tissue and basal lamina called endomysium that surrounds
end of the esophagus, for example). In all cases, skeletal each individual fiber. These connective tissue layers are all
muscle is under voluntary control. continuous with the connective tissue of the muscle’s ten-
The fibers of skeletal muscle are long (up to 3 cm) with don, which in turn attaches to the bone’s periosteum (Fig-
a diameter between 10 and 100 µm. They are multinucle- ures 7-6 and 7-7). In addition, many of the connective
ate, with their nuclei pushed to the periphery of each fiber tissue fibers in the tendon are attached directly to the mem-
(Figure 7-2). With the light microscope, alternating dark branes of the muscle fibers. Thus, when a muscle contracts,
and light bands are visible, giving the fibers a striped or stri- it pulls on its connective tissue (including its tendon) which
ated appearance (Figure 7-3). The dark striations are called transmits the pull to the bone and moves it.
A bands, whereas the lighter ones are known as I bands. A Muscle fibers can be classified as one of three types
thin Z disc bisects each I band. With electron microscopy, it based on diameter, amount of myoglobin (structurally simi-
is seen that the striations are the product of the highly or- lar to hemoglobin and also acts in oxygen transport), rate of
ganized arrangement of actin and myosin filaments within contraction, and other features. The fiber types are red slow
each fiber (Figure 7-4). It is these filaments that form the con- twitch fibers, white fast twitch fibers, and intermediate fibers
tractile apparatus and cause fiber shortening by a mechanism (Figure 7-8). Red fibers are red because of their abundance of
71
72
myoglobin. They have a smaller fiber diameter, have a low has the innate ability to contract rhythmically, although the
glycogen content, and are capable of slower and weaker nervous system and hormones can modify contraction rate.
contractions than white fibers, but do not fatigue as easily. Cardiac muscle fibers are striated, branched, and unin-
White fibers have less myoglobin and more glycogen than ucleate (occasionally two nuclei are present), with the oval
red fibers. They also are larger, contract more rapidly and nucleus found near the fiber’s center (Figure 7-12). They are
with greater strength, but fatigue easily. Intermediate fibers less than 100 µm long and about 15 µm in diameter. Inter-
are intermediate between white and red fibers in these char- calated discs (Figure 7-13), found at the junctions between
acteristics. All muscles have all fiber types, but the propor- cells, are seen as dark transverse lines with the light micro-
tions differ depending on the primary role played by the scope. They physically bind the cells together so the force of
muscle. Red fibers are abundant in postural muscles, contraction is transmitted between the linked cells. They
whereas white fibers predominate in appendicular muscles. also promote the rapid spread of electrical action potentials
Each skeletal muscle fiber is supplied (innervated) by a from cell to cell.
somatic motor neuron. The structure formed between the Cardiac muscle is very vascular, with abundant capil-
muscle fiber and its somatic motor neuron is a motor end laries visible between fibers (Figure 7-14). An endomysium
plate (Figure 7-9). While not visible in light micrographs, of collagenous fibers is also present.
there is an actual gap between the membranes of the neu-
ron and muscle fiber called a synapse. When a nerve im- Characteristics of Smooth Muscle
pulse reaches the end of the neuron a neurotransmitter Smooth muscle (Figure 7-15) is found in the walls of or-
(acetylcholine) is released into the synapse, diffuses across gans, so it is also known as visceral muscle. It is under
it, and stimulates the muscle cell to contract. involuntary control and is capable of slow, sustained con-
Muscle spindles (Figure 7-10) are sensory organs in tractions. (Think of the uterus during childbirth!) Smooth
skeletal muscles that detect stretching. They are formed muscle fibers are long and tapered at both ends with a sin-
from modified muscle fibers and somatic sensory neurons. gle, central nucleus conforming to the cell’s shape (Figure
If stretched, they initiate a stretch reflex that results in 7-16). There are no striations.
contraction. Two layers of smooth muscle are often in the walls of
tubular organs, with the fibers of one layer running the
Characteristics of Cardiac Muscle length of the organ and the fibers of the other layer ori-
Cardiac muscle tissue (Figure 7-11) is found only in the ented around the organ (Figure 7-17). These longitudinal
heart where it forms the myocardium, the thickest layer of and circular layers are capable of producing a specialized
the heart wall. The tissue is under involuntary control and form of contraction in tubular organs called peristalsis.
7-1
SKELETAL MUSCLE Transverse Striations and many peripheral nuclei (N) characterize
skeletal muscle fibers. (X230)
C H A P T E R S E V E N : M U S C L E T I S S U E
73
7-2a 7-2b
SKELETAL MUSCLE IN VARIOUS SECTIONS These specimens from the tongue show skeletal muscle fibers sectioned in all planes. Even though the
striations are not visible in cross section, the peripheral nuclei are a useful feature for identifying this tissue. Both micrographs are X210.
7-4
THE SARCOMERE The orderly arrangement of actin and myosin fila-
I ments accounts for the striations in skeletal muscle. In addition to the
A Z discs and A and H bands, an M disc within the H band is visible.
The contractile unit of skeletal muscle is the sarcomere, the region
7-3 between Z discs. During contraction, the interdigitating filaments slide
across one another and the Z discs are brought closer together. As a
STRIATIONS In the light microscope striations are seen as thick dark result, I bands disappear. The I bands in this specimen are narrow
stripes (A bands) and faintly visible thin dark lines (Z discs) bisecting because the muscle is in a contacted state. (X22,000)
thick light stripes (I bands). Notice that the nuclei are not in the (Courtesy of UCSD Medical Center)
striated part of the cell. (X630)
74
En
Ep
F
Ep
7-5a 7-5b
CONNECTIVE TISSUE OF SKELETAL MUSCLES Muscle fibers in a skeletal muscle are held together by fibrous connective tissue. Epimysium (Ep)
surrounds the entire muscle, perimysium (P) surrounds a fascicle of fibers, and endomysium (En) surrounds each individual fiber. (a) A thick
epimysium is seen in this muscle. (X250) (b) The fibers in this specimen have separated during preparation, but this makes the three connective
tissue layers more visible and emphasizes their continuity. It also defines the fascicles (outline-F). (X250)
M T
7-6a 7-6b
MUSCULOTENDINOUS JUNCTION (a) In this micrograph, the end of the muscle (M) and its tendon (T) are seen. (X60) (b) In this higher magnifi-
cation of the boxed region in (a), the continuity of the muscle’s connective tissue components combining to form the tendon is visible. Also seen
are the connective tissue fibers bound to the muscle fibers’ membrane (arrows). (X250)
75
ST
FT
B
T
7-8
7-7 FAST AND SLOW TWITCH FIBERS Two fiber types are identifiable in
TENDINOUS ATTACHMENT TO BONE The collagenous fibers of the this micrograph stained for glycogen (PAS reaction), which is more
tendon (T) on the left blend with the periosteum and penetrate the abundant in the larger white fast twitch fibers (FT). The fibers lacking
bone (B) matrix to anchor the muscle firmly to the bone. (X60) glycogen tend to be the smaller, red slow twitch fibers (ST). (X250)
MEP
MS
SMN
MEP
7-9 7-10
MOTOR END PLATE The junction between a somatic motor neuron (SMN) and MUSCLE SPINDLES Muscle spindles (MS) are receptors that
its muscle fiber is a complex structure known as a motor end plate (MEP). Two provide sensory information about the amount of stretch
are seen here. Acetylcholine is the neurotransmitter that crosses the synaptic placed on a muscle. A motor end plate (MEP) is also seen.
gap. (X250) (X250)
76
7-11 7-12
CARDIAC MUSCLE Cardiac muscle is found only in the heart where it CARDIAC MUSCLE’S CENTRAL NUCLEI The central position of the nuclei
forms the myocardium. Its fibers are faintly striated (not visible in this is easily seen in this cross section. (X250)
micrograph), branched (arrows), and uninucleate. (X250)
En
C
ID
ID
7-13
7-14
INTERCALATED DISCS Iron-hematoxylin stain
accentuates the striations and intercalated discs CAPILLARIES OF CARDIAC MUSCLE Cardiac muscle is very vascular, as evidenced by the
(ID) of cardiac muscle fibers. The branching is numerous capillaries (C) seen in this micrograph. An endomysium (En), faint intercalated
also visible. (X630) discs (arrows), and branching fibers are also visible. (X250)
77
7-15a
SMOOTH MUSCLE Two specimens of smooth muscle are shown.
Note the absence of striations and the random distribution of
nuclei throughout the tissue. Smooth muscle is sometimes con-
fused with dense regular connective tissue, but in the latter,
nuclei are usually in rows (compare to Figure 4-19). Both
specimens were magnified X120.
7-15b
SMOOTH MUSCLE FIBERS Smooth muscle fibers are

elongated with tapered ends. Notice the single nucleus
(N) and absence of striations. (X120) 7-16
78
7-17a 7-17b
SMOOTH MUSCLE LAYERS In many organs, smooth muscle is found in longitudinal (L) and circular (C) layers, an arrangement that moves
materials through the organ by peristalsis. On slides, this allows viewing of smooth muscle in longitudinal and cross sections. (a) Small intestine
(X120). (b) Small intestine (X250).
Nervous Tissue and Organs
of the Nervous System
C H A P T E R
8
Introduction to Nervous Tissue 8-1). For a more detailed description of the role each part
fills in the body, the reader is referred to a physiology text-
Nervous tissue is specialized for coordinating and integrat-
book. Functionally, the nervous system can be divided into
ing the activities of body’s cells. It accomplishes this through
two parts: the afferent (sensory) nervous system, which
conduction of electrical nerve impulses and secretion of carries information toward the CNS or up the spinal cord
chemical neurotransmitters. toward the brain, and the efferent (motor) nervous system,
The organs of the nervous system are structurally clas- which carries information from the brain and away from
sified as either being part of the central nervous system the CNS. The afferent and efferent systems have somatic
(CNS), which includes the brain and spinal cord, or the pe- (relating to the skin and skeletal muscles) and visceral (re-
ripheral nervous system (PNS), which includes nerves, gan- lating to the internal organs) components. The visceral
glia and sensory receptors. motor system is also known as the autonomic nervous sys-
While it is beyond the scope of this book to provide a tem, which is structurally and functionally divided into a
detailed description of the nervous system’s functional or- sympathetic (thoracolumbar) division and a parasympa-
ganization, it is necessary to refer to its components (Figure thetic (craniosacral) division. The former prepares the body
Nervous System
Afferent (Sensory) System Efferent (Motor) System
Visceral Motor (Autonomic)
Visceral Somatic Somatic

Sensory Sensory Motor Parasympathetic Sympathetic
(Craniosacral) (Thoracolumbar)
Division Division
8-1
FUNCTIONAL ORGANIZATION OF THE NERVOUS SYSTEM
79
80
to contend with stressful and life threatening situations, Extending from the cell body are cytoplasmic processes
whereas the latter is involved in body maintenance during of two types. Dendrites receive impulses and conduct them
nonstressful times. to the cell body. They are usually short and highly branched.
Axons transmit the nerve impulse away from the cell body
Cells of Nervous Tissue to another neuron, a muscle, or a gland. Axons are up to a
meter in length and branch at the end. Depending on the
Nervous tissue is composed of two basic cell types:
type of neuron, there may be one to many dendrites, but
neurons, which are responsible for conduction and integra-
there is always a single axon. Any long, thin neuronal pro-
tion of nerve impulses, and neuroglia (glial cells), which
cess is called a nerve fiber, but usually the term applies to
support neurons (Figure 8-2). The region between cells, the
axons.
neuropil, is a three-dimensional mesh of cellular processes
Neurons can be functionally divided into three groups.
and blood vessels.
Sensory neurons carry impulses towards the CNS, whereas
Glial Cells motor neurons carry impulses away from the CNS. In-
terneurons conduct impulses between sensory and motor
Glial cells do not conduct nerve impulses. Instead, they sup-
neurons within the CNS.
port neurons and have many functions typically associated
There are three structural types of neurons. These are
with fibrous connective tissues. There are several types,
based on the shape of the cell body and the number and
some more easily identified than others. Microglia are
characteristics of the cytoplasmic processes. Multipolar
phagocytic cells of the mononuclear phagocytic system.
neurons (Figure 8-12) have many short dendrites and a sin-
They are the smallest of the glial cells and have an elon-
gle long axon arising from an irregularly shaped cell body.
gated dark staining nucleus with little cytoplasm. They are
The axon can be identified by the absence of Nissl sub-
difficult to identify with certainty in routine preparations
stance at its base, the axon hillock. Motor neurons and in-
(Figure 8-3). Astrocytes (Figure 8-4) are highly branched
terneurons are multipolar. Bipolar neurons have a single
cells that occupy most of the space between neurons in the
dendrite and a single axon emerging from opposite ends of
CNS. They may be associated with blood vessels and the
the elongated cell body (Figure 8-13). These are not found
pia mater (see page 81) by way of perivascular feet. In addi-
in many places of the body. Two examples are in the olfac-
tion to other poorly understood functions, they help form
tory and optic pathways. Unipolar (pseudounipolar) neu-
the blood-brain barrier. Protoplasmic and fibrous astro-
rons (Figure 8-14) are derived from bipolar neurons.
cytes are difficult to differentiate in light micrographs.
During development, the axon and dendrite fuse near the
Oligodendrocytes are responsible for producing myelin (see
cell body so that only a single process emerges from it. The
below) in the CNS (Figure 8-5). Each oligodendrocyte may
“dendrite” ends up being the longer of the two and is called
be associated with 50 or more different neurons. They are
the peripheral process. It carries the impulse from a sensory
small cells with a dark cytoplasm and nucleus. They are the
receptor to the cell body. The “axon” becomes the central
most abundant glial cells in white matter. Schwann cells are
process. It carries the impulse from the cell body to a neu-
responsible for producing myelin in the PNS (Figure 8-6).
ron in the CNS. Most sensory neurons are unipolar.
Ependymal cells line the ventricles and the central canal of
the spinal cord (Figure 8-7). They are cuboidal to columnar
in shape and join with the pia mater to form the choroid Myelination
plexus, a vascular structure responsible for production of Glial cells support nerve fibers. At the very least, fibers in
cerebrospinal fluid (CSF). Satellite cells (Figure 8-8) are as- the PNS are nestled into depressions of Schwann cells. These
sociated with and support neurons in ganglia (see below). fibers are said to be unmyelinated. Many fibers, however,
are covered with an insulating fatty material called myelin,
Neurons and are said to be myelinated.
Although neurons come in a variety of sizes and shapes, In the PNS, a fiber becomes myelinated when Schwann
they have many features in common (Figure 8-9). The por- cells wrap around it several times (Figures 8-15 and 8-16).
tion of the neuron containing the majority of cytoplasm With each wrap, the cytoplasm is squeezed into the remain-
and the nucleus is called the cell body or perikaryon. The der of the cell and the two layers of cytoplasmic membrane
nucleus often has a prominent nucleolus. The cytoplasm are pressed together to form the myelin. (Imagine squeezing
may show an abundance of neurofibrils (Figure 8-10) in it toothpaste from the base of the tube toward the opening.)
or have basophilic granules called Nissl bodies (Figure 8-11). The myelin sheath, then, is actually several double-layers of
The fibrils comprise the cytoskeleton and the Nissl bodies cytoplasmic membrane wrapped around the fiber. (Imagine
are regions of rough endoplasmic reticulum. A pale cyto- wrapping the empty part of the toothpaste tube around a
plasmic region may be visible near the nucleus. This is the stick several times.) Surrounding the myelin is the portion
site of a Golgi apparatus. of the Schwann cell containing the cytoplasm. This is the
C H A P T E R E I G H T : N E R V O U S T I S S U E A N D O R G A N S O F T H E N E R V O U S S Y S T E M
81
neurilemma. Occasionally, cytoplasm remaining in the lay- sensory neuron cell bodies surrounded by satellite cells.
ers of myelin is visible as Schmidt-Lanterman (clefts) lines Sympathetic (prevertebral) ganglia (Figure 8-25) are part of
(Figure 8-17). the sympathetic trunks on the ventral side of the vertebral
Nodes of Ranvier (Figure 8-18) are gaps between adja- column. They contain sympathetic postganglionic neuron
cent Schwann cells where the fiber’s membrane is exposed. cell bodies. The neurons are multipolar with lipofuchsin
Each myelinated segment is called an internode (Figure 8-19). granules (cellular debris) and lack a well-organized layer of
The activities associated with impulse production only satellite cells. Other ganglia are found in association with
occur at the nodes (rather than along the entire length of the viscera and contain autonomic neuron cell bodies (Fig-
the fiber) and thus transmission of the impulse is much ure 8-26).
more rapid in myelinated fibers than in unmyelinated fibers. A group of functionally related neurons in the CNS is
In the CNS, myelination is performed by oligodendro- called a nucleus. (Figure 8-27). By definition, nuclei are in
cytes. Each oligodendrocyte myelinates fifty or more differ- the gray matter, but they are less obvious than ganglia be-
ent fibers and, as such, does not surround each fiber with a cause they lack a connective tissue covering that forms an
neurilemma. Nodes of Ranvier and internodes are present outer boundary. There are many nuclei in the brain, espe-
in myelinated fibers of the CNS. cially in the thalamus and hypothalamus.
Gray and White Matter Other Neural Structures

Myelin is a fatty material, and has a whitish appearance to Meninges
it. In fresh preparations, dense collections of myelinated Connective tissue is absent from the interior of the CNS,
fibers are referred to as white matter, whereas unmyeli- but it is covered with three connective tissue layers referred
nated fibers and neuron cell bodies comprise gray matter to as meninges (Figure 8-28). The innermost, pia mater, is a
(Figure 8-20). These terms are applied to the CNS, but not delicate layer of collagen and elastic fibers with a basement
the PNS. membrane. The middle layer, the arachnoid, consists of
Nerves and Tracts fibrous strands that form a web-like layer continuous with
the pia matter. Blood vessels pass through the subarachnoid
Nerve fibers tend to travel together. In the PNS, collections
space, as does cerebrospinal fluid. The outermost layer is
of fibers are identifiable structures called nerves. Nerves are
made of dense connective tissue and is called the dura
either sensory (containing only sensory fibers) or mixed (a
mater. It may be visible in spinal cord preparations, but is
combination of motor and sensory fibers, both somatic and
usually absent from brain specimens since it contributes to
visceral.
the periosteum of the skull and is difficult to remove with
Each nerve is associated with collagenous connective
the brain.
tissue similar in arrangement to the connective tissues of
muscles (Figures 8-21 and 8-22). On the surface is epi- Choroid Plexus
neurium, a fibrous covering that wraps and supports the The ependymal cells of the brain’s ventricles associate with
nerve, separating it from the surrounding fascia. The peri- pia mater to form highly vascular infoldings called choroid
neurium is derived from the epineurium. It enters the nerve plexus (Figure 8-29). These are responsible for producing
and forms nerve fiber bundles called fascicles. Endoneurium cerebrospinal fluid, which bathes the CNS and acts as a
is a delicate connective tissue layer derived from per- shock absorber.
ineurium that surrounds each fiber and Schwann cell. This
Synapse Structure
arrangement weaves all the nerve fibers into a structurally
sound unit. A synapse is the intercellular junction between a neuron
In the CNS, fibers with a common origin and a com- and another cell. If the junction is formed with another
mon destination form tracts (Figure 8-23). Tracts do not neuron, it is called an axodendritic or axosomatic junction
have the connective tissue components of nerves and are (Figure 8-30). If it is formed with a skeletal muscle cell, it is
difficult to identify because there are no clear boundaries called a neuromuscular junction and the structure is called
between them. Tracts may either be ascending (sensory) or a motor end plate (Figure 8-31). The two cells involved in the
descending (motor), but are never mixed. synapse do not actually contact one another, but rather have
a space called the synaptic cleft separating them. When the
Ganglia and Nuclei electrical nerve impulse reaches the end of an axon, a chemi-
As with fibers, neuron cell bodies tend to be in groups. cal neurotransmitter is released by the presynaptic neuron
Neuron cell bodies in the PNS are found in structures called that diffuses across the synapse to bind with receptors on the
ganglia. Spinal (dorsal root) ganglia are located on either postsynaptic neuron. The consequence of neurotransmitter
side of the vertebral column in association with the dorsal binding to the postsynaptic neuron may be stimulation or in-
roots of spinal nerves (Figure 8-24). They contain unipolar hibition, depending on the neuron and the neurotransmitter.
82
Organs of the Central Nervous System of the pyramids. These fibers continue down into the cord
Spinal Cord to synapse with somatic motor neurons in the ventral gray
horns. Medullary gray matter includes various nuclei.
The spinal cord is continuous with the medulla oblongata
of the brain stem. It is formed as the medulla passes Cerebellum
through the foramen magnum at the base of the skull and The cerebellum is composed of gray matter around white
continues to its own termination even with the second lum- matter and has a prominently convoluted surface (Figure
bar vertebra. In cross section (Figures 8-32 and 8-33), the 8-37). The gray matter is the cerebellar cortex and consists
white matter surrounds gray matter, which is in the shape of a superficial molecular layer and a deeper granular
of the letter “H”. The gray matter is divided into three or layer. The molecular layer is composed mostly of unmyeli-
four regions, depending on the part of the spinal cord. The nated fibers and contains few neurons, whereas the granular
dorsal gray horns contain interneuron cell bodies and cell layer has numerous neurons. At the junction of the two lay-
bodies of second order sensory neurons that carry impulses ers is a row of distinctive neurons called Purkinje cells
to the thalamus. The dorsal horns usually extend to the sur- whose dendrites project into the molecular layer and whose
face of the cord. Ventral gray horns contain somatic motor axons project into the white matter of the cerebellar
neuron cell bodies. There is considerable white matter be- medulla (Figure 8-38). The cerebellum is involved in coor-
tween the ventral horns and the spinal cord’s surface. Lat- dinating and refining somatic motor activity. As such, it has
eral gray horns are found in spinal cord segments T1 connections to and from the motor cortex of the cerebrum,
through L2 and S2 through S4. The former contain sympa- and from proprioceptors in muscles, tendons and joints.
thetic preganglionic neuron cell bodies, whereas the latter
contain parasympathetic preganglionic neuron cell bodies.
Cerebral Cortex
Finally, the gray commissure primarily contains axons of
interneurons. In the middle of the gray commissure is the The cerebral cortex is the layer of gray matter on the sur-
central canal. It is small, lined with ependymal cells, and face of the cerebrum. The cortex is highly folded into sulci
carries cerebrospinal fluid. (depressions) and gyri (ridges), more commonly referred to
The white matter of the cord is divided into dorsal, lat- as convolutions. The cerebrum is the site of higher thought
eral, and ventral columns. The columns carry ascending processes and reasoning, as well as the location of neurons
and descending tracts of fibers, but the precise locations of responsible for interpreting sensory input to produce vari-
the tracts are not identifiable due to the absence of connec- ous sensations, initiating voluntary motor activity and stor-
tive tissue boundaries. ing memory.
The ventral median fissure is a deep indentation on the In most parts of the cerebral cortex, there are six identi-
ventral side of the cord along the midline. A shallower and fiable layers. These are listed below from superficial to deep
narrower dorsal median sulcus is on the dorsal side. (Figures 8-39 and 8-40).
Each spinal nerve is connected to the spinal cord by ◗ The molecular layer is the most superficial layer of the
two roots. These are the dorsal root and the ventral root cortex and is adjacent to the pia matter. It consists pri-
(Figure 8-34). Even though all spinal nerves are mixed, sen- marily of neuronal axons and dendrites and scattered
sory and motor fibers segregate in the roots. All sensory glial cells.
fibers entering the spinal cord follow the dorsal root and ◗ The outer granular layer consists of two neuron types:
have their cell bodies in the dorsal root ganglion, and all small pyramidal cells and stellate cells.
motor fibers leave the cord through the ventral root. ◗ The pyramidal cell layer is fairly thick and contains py-
Like other parts of the CNS, the spinal cord is covered ramidal cells of increasing size with increasing depth.
with meninges. The dura mater of the cord is continuous ◗ The inner granular layer is composed of stellate cells.
with the epineurium of the spinal nerve roots (Figure 8-35).
◗ The ganglionic layer has large pyramidal cells (Betz
Medulla Oblongata Cells), stellate cells, and cells of Martinotti.
The medulla oblongata (Figure 8-36) is the part of the ◗ The multiform cell layer is the deepest in the cortex. It
brain that joins the spinal cord. Its white matter is com- is characterized by a diverse grouping of neurons.
posed primarily of ascending and descending tracts. The White matter of the cerebrum is deep to the multiform
pyramidal tracts carry motor fibers arising from the motor cell layer of the cortex. It is composed of projection (both
cortex of the cerebrum and are visible as the raised pyramids ascending and descending), association (within a hemi-
on the ventral surface of the medulla. A majority of the fibers sphere), and commissural (connecting opposite hemispheres)
in these tracts cross to the opposite side at the decussation tracts.
83
Np
8-2 8-3
CELLS OF NERVOUS TISSUE Nervous tissue is made of two basic cell PROBABLE MICROGLIA Microglia (arrows) are phagocytic cells of neural
types: neurons (N), which are larger and conduct nerve impulses, and tissue, but are difficult to identify with a high degree of certainty in
smaller neuroglia (glial cells) (G), which perform a variety of functions. routine preparations. They are small cells with a dense and elongated
Neurons are larger and have a complex shape that generally cannot nucleus. (X320)
be fully appreciated in histology specimens because they have been
sectioned. The neuropil (Np) primarily consists of neuronal
cytoplasmic processes. (X210)
FIBROUS ASTROCYTES Perivascular feet of astrocytes

(arrows) attach to the basement membrane of capillaries
8-4 (C) to form the blood-brain barrier. (X210)
84
8-6
8-5 SCHWANN CELLS Myelin in the PNS is produced by Schwann cells. In
OLIGODENDROCYTES The most abundant glial cell of white matter is the this nerve cross section, the dark staining, crescent shaped Schwann
oligodendrocyte (arrow). These cells are responsible for producing cell nuclei (arrows) are visible in association with a few of the
myelin in the CNS. (X380) myelinated fibers. (X630)
E
SC
SC
8-7 8-8
EPENDYMAL CELLS The ventricular system of the brain is lined with an SATELLITE CELLS The neurons of some ganglia are supported by
epithelium-like layer of ependymal cells (E), as seen here. They also are satellite cells (SC). This specimen is from a dorsal root ganglion.
found as part of the choroid plexus, which is responsible for producing (X250)
cerebrospinal fluid. (X250)
85
Nu
CB
8-9 8-10
A TYPICAL NEURON Neurons are usually large, irregularly shaped cells. NEUROFIBRILS The cytoskeleton of neurofibrils (blue lines) is often
Most have a large nucleus (N) with a prominent nucleolus (Nu). Nu- visible in neurons. Notice the large nucleus with its nucleolus. This
merous cytoplasmic processes, called axons and dendrites, extend from specimen is a whole mount of neural tissue. (X250)
the cell body (CB) (perikaryon). The cytoskeleton is visible in this
multipolar neuron. (X250)
8-11a 8-11b
NISSL BODIES IN NEURONS With proper staining, Nissl bodies are visible in the cytoplasm of some neurons. Electron micrographic studies have
shown the Nissl substance to be rough endoplasmic reticulum. The Nissl substance is absent from the base of the axon, the axon hillock, making
it possible to identify the axon without seeing its entire length (arrows). Both specimens are multipolar neurons. (X380)
86
CB
8-13
BIPOLAR NEURONS OF THE RETINA Bipolar neurons have a single axon
and dendrite. Their cell bodies are in the layer labeled “CB” with the
8-12 axons (A) and dendrites (D) extending into the layers on either side.
They are found in the retina and a couple of other places in the body.
MULTIPOLAR NEURON This is a smear of neural tissue. Notice the (X100)
multiple cytoplasmic processes of this neuron and its cytoskeleton of
blue neurofibrils. The arrow indicates the axon. (X100)
UN
UNIPOLAR (PSEUDOUNIPOLAR) NEURONS

Unipolar neurons (UN) have a single
process (P) that divides into a peripheral
process, which comes from a sensory
receptor, and a central process, which leads
to the CNS. The cell body is spherical.
This specimen was taken from a dorsal
8-14 root ganglion. (X320)
87
Schwann cell cytoplasm
Axon
Schwann cell nucleus Mesaxon Neurilemma Myelin

sheath
8-15a
8-15c
Nu
M MYELIN SHEATH (a) Myelin is produced by compressed

F
cytoplasmic membrane of a Schwann cell as it wraps
around a nerve fiber. (b) This electron micrograph shows
the myelin sheath (My) as multiple wraps of the Schwann
cell membrane. The mesaxon (M), neurilemma (N),
My Schwann cell nucleus (Nu), and nerve fiber (F) are also
visible. (X15,000) (c) The layers of membrane are visible
in this higher magnification of myelin. (X18,750)
8-15b (Micrographs courtesy of UCSD Medical Center.)
88
M
F
S
8-16a 8-16b
MYELINATED FIBERS (a) In this nerve cross section, the myelin sheaths (M) surround nerve fibers (F), but are disrupted due to slide preparation.
Schwann cell nuclei (S) are also visible. (X630) (b) This nerve cross section was prepared with an osmium stain that stains fat black and thus
highlights the myelin. (X380)
NR
8-17
SCHMIDT-LANTERMAN CLEFTS During myelination, some cytoplasm is
trapped in the myelin sheath and is visible as Schmidt-Lanterman clefts.
A node of Ranvier (NR) is also visible. (X250)
8-18
NODES OF RANVIER Gaps in the myelin sheath between adjacent
Schwann cells are called nodes of Ranvier. The electrical activity
associated with impulse transmission only occurs at the nodes of a
myelinated fiber rather than its entire length, as in unmyelinated
fibers. This results in faster impulse conduction. The fiber is visible
as a gray line crossing the node. (X380)
89
INTERNODES Two nodes of Ranvier are indicated

with arrows; the region in between is known as
the internode. Since internodes are not involved
in impulse transmission and they constitute the
majority of the fiber, impulse transmission is more
rapid in myelinated fibers than in unmyelinated
8-19 ones. (X60)
W W
G G
8-20a 8-20b
GRAY AND WHITE MATTER In the CNS, regions of myelinated fibers have a whitish appearance and are known as white matter (W). The main
cells of white matter are oligodendrocytes. Gray matter (G) is composed of neuron cell bodies, glial cells and unmyelinated fibers (though a few
myelinated fibers may be found). Both specimens are from the spinal cord. (a) Silver stain, X50. (b) H&E stain, X50.
90
Ep
8-21a
CONNECTIVE TISSUE COMPONENTS OF A
NERVE Nerves are made of nerve fibers and
connective tissue. Each nerve is surrounded
by a connective tissue epineurium (Ep). The
perineurium (P) surrounds fascicles of fibers,
and the endoneurium (En) surrounds each
individual fiber. (a) This is a cross section
of a small nerve. (H&E stain, X25). (b) This
is an enlargement of the boxed area in (a). 8-21b
(X125) (c) This section of a nerve was
stained with Masson stain, which makes
connective tissue blue. The endoneurium
and part of the perineurium is visible. Also
seen are Schwann cells (S), myelin (My) and
fibers (F). (X1000)
P
En
S
My
8-21c
91
Fb
P
My
F
NR
NR
8-22a NR My
LONGITUDINAL SECTIONS OF A NERVE (a) The perineurium (P) En

with fibroblast nuclei (Fb) is obvious, but the endoneurium
is difficult to discern in this H&E preparation of a nerve cut
in longitudinal section. Myelin (My), nerve fibers (F) and
nodes of Ranvier (NR) are also visible. (X210) (b) This
longitudinal section of a nerve clearly illustrates the endo-
neurium (En). A blood vessel (BV) is also visible. Other
structures are labeled as in (a). (X320)
BV
8-22b
L A
TRACTS Collections of nerve fibers in the CNS are called
tracts and they comprise the white matter. While the general
location of each tract is known, their precise boundaries are
unclear due to the absence of connective tissue or any other
landmark around them. For instance, in this spinal cord
specimen the lateral (L) spinothalamic and anterior (A)
spinocerebellar tracts are adjacent, but there is no clear
8-23 boundary between them. (X20)
92
DRG
VR
8-24a
DORSAL ROOT GANGLION Neuron cell bodies in the PNS

are found in ganglia. Shown here are three examples of
dorsal root ganglia, that are located in the dorsal roots
of each spinal nerve and contain sensory neuron cell
bodies. (a) The dorsal root ganglion (DRG) is seen as an
expanded region of the dorsal root. The ventral root
(VR) is also visible. It contains motor fibers. (X25) (b
and c) The unipolar neuron cell bodies and satellite cells
surrounding them are seen in these micrographs. (X230)
8-24b
8-24c
93
8-25a 8-25b
AUTONOMIC GANGLIA (a) Sympathetic postganglionic neuron cell bodies are seen in this sympathetic ganglion. The neurons are multipolar and
satellite cells, while present, are fewer in number than in dorsal root ganglia. Also note the connective tissue between neurons. (X230) (b) In
addition to the features visible in (a), the neurons in this ganglion show brown lipofuchsin granules, which are thought to be breakdown products
from lysosome activity. They increase in number with age. (X320)
AP
8-27a
8-26
AUERBACH’S PLEXUS Some ganglia are embedded in other organs.
Neurons serving the musculature of the digestive tract are located
between muscle layers in the digestive organs and form Auerbach’s
plexus (AP). The submucosal plexus (Meissner’s plexus) is responsible
for regulating glandular activity. (X380)
NUCLEI IN THE CNS Groups of functionally related

neuron cell bodies in the CNS are called nuclei. The
olivary nucleus (O) of the medulla oblongata is seen
in (a). (X20) (b) Part of a nucleus in the brain is
shown in this micrograph. (X320) 8-27b
94
8-28a 8-28b
MENINGES (a)The three meningeal layers are visible in this spinal cord specimen. The outermost dura mater (D) has pulled away from the
arachnoid (A) during preparation. The pia mater (P) is visible as a blue line on the surface of the cord. (X65) (b) A blood vessel is visible in
the pia mater (P) of this cerebrum specimen. The other meningeal layers have been removed. (X130)
BV
BV
8-29a 8-29b
CHOROID PLEXUS The vascular choroid plexus is formed from pia mater and ependymal cells. (a) This micrograph shows choroid plexus from the
fourth ventricle. Ependymal cells (E) and blood vessels (BV) are visible. The cerebellum is in the lower right corner of the image. (X65) (b) This
specimen was magnified X250 and shows more detail than (a). Ependymal cells lining the ventricle are also visible, though some have pulled away
from the neural tissue during preparation (arrow).
95
SMN
MEP
8-31a
8-30
SYNAPSES The synapses between several axons and the cell body
(axosomatic) and dendrites (axodendritic) of a Purkinje cell in the
cerebellum are shown in this micrograph. (X380)
SMN
MEP
M
8-31b
MOTOR END PLATES Motor end plates (MEP) contain the synapse
between a somatic motor neuron (SMN) and a skeletal muscle fiber
(M). They are complex structures made from the membranes of both
cells. (a and b X250)
96
DC
DH
DMS
D
DC
DR CC
DH
DRG LC
LC C
VH
VH
VC
VR
VMF
VMF
VC
8-32a 8-32b
DC DH
DMS DMS
DC
LC
LH DH
C CC
LC
CC
VH
VH
VC
VMF
VMF
VC
8-32c 8-32d D
SPINAL CORD IN CROSS SECTION These four micrographs are of spinal cords from different specimens and regions. Specimens (a) and (c) are from
the thoracic region. Specimens (b) and (d) are from the cervical and lumbar regions, respectively. All were magnified X25. Key to symbols used:
DH 4 dorsal gray horn, VH 4 ventral gray horn, LH 4 lateral gray horn, DC 4 dorsal white column, LC 4 lateral white column, VC 4
ventral white column, C 4 central commissure, CC 4 central canal, DRG 4 dorsal root ganglion, DR 4 dorsal root, VR 4 ventral root, D 4
dura mater, VMF 4 ventral median fissure, and DMS 4 dorsal median sulcus.
97
E CC
CC
8-33a 8-33b
CENTRAL COMMISSURE AND CENTRAL CANAL OF SPINAL CORD (a) The central commissure is gray matter, but also contains fibers crossing from
one side of the cord to the other. The central canal (CC) is lined with ependymal cells (E) and contains cerebrospinal fluid. (X130) (b) The central
canal. (X250)
DRG
SC
D
E
EDR
DRG
VR
VR
8-34
SPINAL NERVE ROOTS Each spinal nerve is attached to its segment of E
8-35
the spinal cord by two roots. The dorsal root (DR) carries sensory
fibers entering the cord. The cell bodies of these sensory neurons are DURA MATER AND EPINEURIUM The dura mater (D) and epineurium (E)
located in the dorsal root ganglion (DRG). The ventral root (VR) form a continuous layer where the nerve roots join the spinal cord. The
carries motor fibers out of the cord. (X6) ventral root (VR), and dorsal root ganglion (DRG), and spinal cord
(SC) are shown. (X65)
98
ON ON
P P
ON P
8-36a 8-36b
MEDULLA OBLONGATA The medulla oblongata houses ascending and descending tracts, and several nuclei. (a) This is a panoramic view of the
medulla magnified X6. The fourth ventricle (V) is at the top. At the bottom are the pyramids (P), which are major descending tracts. The olivary
nucleus (ON) is also visible. (b) This is a higher magnification of the olivary nuclei and the pyramids. (X25)
M G
G
W
8-37a 8-37b
CEREBELLUM The cerebellum is involved in coordinating motor functions. Its surface is convoluted and it consists of gray matter around white
matter. The gray matter is the cerebellar cortex and is divided into a superficial molecular layer (M) and a deeper granular layer (G). The white
matter (W) carries fibers entering and leaving the cerebellum. (X25) (b) This micrograph was magnified X130 and shows the layers in the
cerebellum.
99
P P
8-38a
P P
8-38b
8-38c
PURKINJE CELLS The most distinctive cells of the cerebellum are the Purkinje cells (P). They are found at the
junction of the molecular (M) and granular (G) layers. They have highly branched dendrites that enter the
molecular layer. Their single axon penetrates the granular layer. (a) This micrograph illustrates the branched
dendrites of Purkinje cells. (X320) (b) The fibers in the molecular layer are visible in this specimen stained with a
silver stain. (X320) (c) Axosomatic and axodendritic synapses on the Purkinje cells are visible in this micrograph.
(X425)
100
II
D
III
IV D
D A
V
VI
8-40a
8-39
CEREBRAL CORTEX The cerebral cortex is a thin layer of gray
matter. In most parts of the cerebrum, it is divided into six
layers. From superficial to deep: (I) Molecular layer, (II) Outer
granular layer, (III) Pyramidal cell layer, (IV) Inner granular
layer, (V) Ganglionic layer, (VI) Multiform cell layer. (X50)
8-40b
CELLS OF THE CEREBRAL CORTEX (a) The most distinctive cells
of the cortex are the pyramidal cells. Their apex is pointed
toward the surface. The axon (A) emerges from the other end.
Dendrites (D) are found at the apex and at the corners of the
P P base. Pyramidal cells are found in Layers III and V, with the
larger ones being deeper in the cortex. (X320) (b) Two pyra-
midal cells are seen in this specimen stained with a silver stain.
(X530) (c) Fusiform cells (F) have a single axon emerging from
the side and dendrites coming off both ends. Pyramidal cells
(P) are also seen in this specimen. (X320)
F
8-40c
Special Sensory Organs
C H A P T E R
9
Introduction to the Senses in H&E stained specimens, but nucleus location provides a
Sensory receptors receive and respond to stimuli arising ei- high percentage basis for identification. The nuclei of basal
ther from within the body or from the external environ- cells are located in lower third of the epithelium, while nu-
ment. They are divided into receptors of the general senses clei of sustentacular cells are near the surface. The nuclei of
and the special senses. olfactory receptor cells are in the middle.
General sensory receptors are simple in construction The connective tissue deep to olfactory epithelium
and are widely distributed in the body. Examples include (lamina propria) contains Bowman’s glands. These serous
touch receptors (Meissner’s corpuscles) and pressure recep- glands produce a watery secretion that dissolves odorifer-
tors (Pacinian corpuscles) of the skin, which are covered in ous chemicals and makes them more able to stimulate the
Chapter 11. Other general sensory receptors are covered in receptors.
their respective chapters.
Taste Buds
The special senses include taste, smell, vision, hearing,
and balance and equilibrium. Their receptors are more Taste buds (Figure 9-2) are found in the epithelium of the
complex in construction and are more localized. They are tongue as well as other parts of the oral mucosa and house
the topic of this chapter. the chemoreceptors for the sensation of taste. They are
oval-shaped with an opening at the surface called the taste
Olfactory Receptors pore. Internally, lighter staining gustatory cells and darker
The receptors for olfaction are the simplest of the special sustentacular cells have long microvilli that project into the
senses. Olfactory epithelium covers the upper nasal cavity taste pore. While both cells are associated with neurons, the
and superior nasal conchae (Figure 9-1). It is a modified gustatory cells are considered to be the actual chemorecep-
PSCC that contains the olfactory receptor cells, which are tors. Electron micrographs have revealed basal cells, a third
bipolar neurons. The end of the dendrite is dilated to form cell type that may be the source of new cells.
an olfactory vesicle, whereas its axon passes through a
foramen of the cribriform plate to synapse with second The Eye
order neurons in the olfactory bulb. The eyeball is the site of photoreceptors responsible for vi-
Two other cells are located in the olfactory epithelium. sion. It is composed of three basic layers (Figure 9-3). From
Basal cells are small and located near the basement mem- external to internal, they are the fibrous tunic (tunica fibrosa),
brane. They are a source of new cells. Tall supporting (sus- the vascular tunic (tunica vasculosa), and the neural tunic
tentacular) cells provide physical and nutritive support to the (tunica nervosa). In addition, there are two cavities in the
receptor cells. The three cell types are difficult to differentiate eye. The anterior cavity is anterior to the lens and contains
101
102
the watery aqueous humor. The posterior cavity is poste- light intensity. The latter is the dilator pupillae muscle,
rior to the lens and occupies the greatest volume of the eye. which is responsible for its dilation when it is darker.
It contains the gel-like, refractive vitreous body. Aqueous humor is derived from capillaries in the ciliary
processes. It flows from the posterior chamber, through the
pupil, into the anterior chamber, and is returned to the
Fibrous Tunic blood by the canal of Schlemm. Aqueous humor supplies
The fibrous tunic is mostly composed of the sclera, with the the lens and cornea with oxygen and nutrients.
anterior one-sixth forming the cornea. The sclera is made of The lens (Figure 9-10) is a biconvex transparent disc.
densely arranged collagen fibers, which accounts for its white Its flexibility allows it to change shape (as a result of
appearance. It is an attachment site for the ocular muscles changes in tension brought about by contraction and relax-
and is continuous with the dura mater of the optic nerve. ation of the ciliary muscle) and bring images into focus on
The cornea (Figure 9-4) is transparent and avascular the retina. It consists of three layers. On the surface is a
with five distinct layers. On the anterior, the corneal epithe- transparent, collagenous lens capsule. The second layer is a
lium is a thin, nonkeratinized stratified squamous epithe- simple cuboidal subcapsular epithelium that lies deep to the
lium that is continuous with the conjunctival lining of the capsule on the anterior surface. Finally, up to 3000 elon-
inner eyelid. It has numerous nerve endings in it. Deep to gated and hexagonal lens fibers make up the greatest por-
the corneal epithelium is Bowman’s membrane, a thin basal tion of the lens. As they mature, they elongate along the
lamina made of collagen. The stroma is the thickest layer antero-posterior axis of the lens and lose their organelles.
and is composed of up to 250 lamellae of collagen fibers. The fibers’ refractive ability derives from the accumulated
The fibers in each lamella are parallel, but the fibers in ad- protein crystallin contained in them.
jacent lamellae are oriented in different directions. The canal
of Schlemm is found in the stroma at the sclero-corneal Neural Tunic
junction (Figure 9-5). Descemet’s membrane is a thick base- The photoreceptor cells of the retina are called rods and
ment membrane between the stroma and the corneal endo- cones. Their complex shape includes a nuclear region (cell
thelium. The posterior of the cornea is lined with a simple body), which contains the nucleus. Extending posteriorly
squamous corneal endothelium. Among other functions, the from the nuclear region are inner and outer segments. The
corneal endothelium keeps the stroma dehydrated to main- shape of the outer segment, which contains pigments, is the
tain optical qualities. basis for naming rods and cones. Extending anteriorly from
the nuclear region is a synaptic region.
Functionally, rods and cones differ in the amount of
Vascular Tunic
light necessary to stimulate them. Rods are stimulated in
Deep to the fibrous tunic is the vascular tunic. It is com- low light intensity and are responsible for monochromatic
posed of the choroid, ciliary body, and iris. The choroid (“black and white”) vision. Cones are only stimulated by
(Figure 9-6) is made of loose connective tissue. It is highly higher light intensities and are responsible for color vision.
vascular and contains an abundance of melanin, a black The retina (Figure 9-11) consists of ten layers. With their
pigment. complex shapes and their orientation perpendicular to the
The ciliary body (Figure 9-7) is a circular expansion of retina, different parts of rods and cones show up in several of
the vascular tunic even with the lens. Ciliary processes (Fig- the layers. The layers are listed below from outside to inside.
ure 9-8) extend from the ciliary body and produce aqueous
humor, which occupies the anterior cavity of the eye (see
◗ A pigmented epithelium abuts the choroid. It is com-
posed of short columnar cells that accumulate melanin.
below). Suspensory ligaments of the lens also extend from
This pigment absorbs light that has not been absorbed
the ciliary processes and insert on the lens. Smooth ciliary
by the rods and cones and prevents reflection back into
muscle within the ciliary body adjusts tension on the sus-
the eye. The pigment cells also surround the rod and
pensory ligaments and in turn on the lens, thus changing its
cone segments.
shape and allowing it to focus light on the retina.
The iris (Figure 9-9) is the colored part of the eye. It sep- ◗ The photoreceptive layer contains the rod and cone
arates the anterior chamber and the posterior chamber of the segments.
anterior cavity. The hole in its center is the pupil. On the pos- ◗ The thin acidophilic external limiting membrane is not
terior surface of the iris is a layer of melanocytes. The more a membrane at all. It is a region of tight junctions be-
abundant they are, the darker the eyes. Internally, there are tween Muller cells (large glial cells—see the next page)
smooth muscles arranged circularly and radially around the and the rods and cones.
pupil. The former is the sphincter pupillae muscle and is re- ◗ The outer nuclear layer consists of densely packed rod
sponsible for constriction of the pupil in conditions of high and cone cell bodies and their nuclei.
C H A P T E R N I N E : S P E C I A L S E N S O R Y O R G A N S
103
◗ The outer plexiform layer is a region of axodendritic epithelium. It opens into the nasopharynx via the auditory
synapses between the rods and cones and the next layer (Eustacian) tube. The ossicles are three small bones (mal-
of (bipolar) neurons. leus, incus, and stapes) that link the tympanic membrane to
◗ The inner nuclear layer contains cell bodies of bipolar the inner ear. Vibrations in the tympanic membrane due to
neurons, other neurons, and Muller cells. sound waves are amplified by the ossicles.
◗ The inner plexiform layer is a region of axodendritic Internal Ear
synapses between bipolar neurons and the neurons
The internal ear is made of small cavities and channels hol-
(ganglion cells) of the next layer.
lowed out of the temporal bone. Collectively, these con-
◗ The ganglion cell layer contains cell bodies of ganglion stitute the osseous (bony) labyrinth. The membranous
cells, whose axons form the optic tract. labyrinth is contained within and conforms (more or less)
◗ The optic nerve fiber layer is composed of ganglion cell in shape to the osseous labyrinth. Both are filled with fluid.
axons that converge at the optic disc and emerge as the The osseous labyrinth contains perilymph, whereas the
optic tract. (Since the neural tunic is an outgrowth of membranous labyrinth contains endolymph. There are
the brain, the optic nerves are more appropriately re- three regions in the internal ear—the vestibule, the semicir-
ferred to as “tracts.”) cular canals, and the cochlea.
◗ The innermost retinal layer is the inner limiting mem- The vestibule (Figure 9-15) is the middle portion of the
brane. It is the basal lamina of long glial cells called osseous labyrinth. Within it are the utricle and saccule.
Muller cells that extend from the inner limiting mem- Both are made of membranous labyrinth. They are lined
brane to the outer limiting membrane where they form with a simple cuboidal epithelium and each contains a mac-
tight junctions with the rods and cones. Muller cells ula (Figure 9-16). The maculae are oriented at right angles
physically and metabolically support rods and cones. to one another and are composed of receptor cells used in
Where the optic tract emerges from the eye, there is no maintenance of balance and equilibrium. The actual recep-
retina. This is the blind spot (Figure 9-12). A few millimeters tors are sensory hair cells that have stereocilia and a single
lateral and slightly inferior to it is a small pit in the retina cilium (kinocilium) projecting into a gelatinous layer of gly-
where visual acuity is greatest. This is the fovea centralis. coprotein. Otoliths made of calcium carbonate are also em-
bedded in the glycoprotein. Head movements result in
Eyelid and Conjunctiva bending of the stereocilia and stimulation of the hair cells.
The eyelid (Figure 9-13) derives structural support from a The complex sensory input from hair cell stimulation is
plate of fibroelastic tissue called the tarsus and muscle processed by the brain and used to determine the position
fibers from the orbicularis oculi and levator palpebrae (in of the head in space.
the upper eyelid) muscles. Anteriorly, it is covered with a Extending posteriorly from the vestibule are three semi-
thin skin. Posteriorly, it is covered by a stratified columnar circular canals arranged at right angles to one another.
epithelium with goblet cells that forms the palpebral con- Semicircular ducts made of membranous labyrinth are in
junctiva. The palpebral conjunctiva reflects and covers the each canal, both ends of which open into the utricle. The
exposed portion of the sclera as the bulbar conjunctiva. ampulla is a dilation at one end of each duct. It contains the
The mucous secretions lubricate the eyelids and eye surface. crista ampullaris (Figure 9-17) which has sensory hair cells
similar to those of the maculae. The cupula is a gelatinous
The Ear glycoprotein layer that overlies each crista. Body move-
The external ear consists of the pinna (auricle) and the ex- ments result in inertia in the endolymph, which pushes on
ternal auditory meatus. The pinna (Figure 9-14) is com- the cupulae and stimulates the hair cells. The complex pat-
posed of a framework of elastic cartilage covered with skin. tern of hair cell stimulation is interpreted by the brain as
It collects sound waves and funnels them through the exter- movement in a particular direction at a particular rate.
nal auditory meatus to the eardrum. The proximal part of The cochlea (Figure 9-15) contains the receptors for
the external auditory meatus is formed from elastic carti- hearing. It consists of a spiral canal extending anteriorly
lage, while the distal portion is bone. Both are lined with from the vestibule. The bony labyrinth is divided into two
skin containing glands that secrete cerumen (earwax). The separate canals by the membranous scala media, which is
fibrous tympanic membrane forms the partition between filled with endolymph. The scala vestibuli and scala tym-
the external and middle ear. pani are on either side of the scala media and contain peri-
lymph. Since the scala media does not extend the entire
Middle Ear length of the cochlea, the scala vestibuli and scala tympani
The middle ear cavity is a space within the temporal bone are continuous at the apex of the cochlea. The modiolus
and is lined with simple cuboidal to simple squamous forms the bony axis of the cochlea.
104
In cross section, the scala media looks triangular, with Vibration of a particular frequency (related to the wave-
the vestibular and basilar membranes separating it from the length of the sound that caused it) results in movement of a
scala vestibuli and scala tympani, respectively (Figure 9-18). particular part of the basilar membrane. This stimulates the
Resting on the basilar membrane is the organ of Corti (Fig- hair cells, which in turn stimulate the sensory bipolar neu-
ure 9-19), which contains hair cells and support cells. The rons that occupy the spiral ganglion (Figure 9-20). Their
hair cells are the actual sound receptors and have stereocilia axons form the auditory nerve. The auditory cortex inter-
embedded in the tectorial membrane that overlies them. prets the information as sound of a particular pitch.
OE
9-1
OLFACTORY EPITHELIUM Olfactory cells are the receptors for the sense 9-2a
of smell. They are located in the olfactory epithelium (OE), a modified
PSCC in the nasal cavity. It is difficult to identify the three cell types of
this epithelium in H&E stains, but nucleus location provides a means
of making a good guess. Basal cells have nuclei in the lower third,
nuclei of sustentacular cells are near the surface, and nuclei of
olfactory cells are in the middle. Bowman’s glands (B) and ducts are
visible in the lamina propria. (X210)
N P
G
S
V
V
F
M
9-2b
9-3 TASTE BUDS Chemoreceptors for the sensation of taste are located in
taste buds found in the epithelium lining the oral cavity. (a) Taste buds
LAYERS OF THE EYEBALL The three layers of the eyeball, from outside (arrows) on lingual papillae. (X130) (b) Detail of taste buds. Visible
in, are the fibrous tunic (F), vascular tunic (V), and neural tunic (N). are the taste pores (P), gustatory cells (G) with light staining nuclei,
An ocular muscle (M) is also visible in this specimen. (X50) and sustentacular cells (S) with darker nuclei. (X380)
105
C S
cS
C
S
E
9-4
CORNEA The transparent anterior portion of the fibrous tunic is the
cornea. There are five layers in the cornea. On the anterior surface
is the corneal epithelium (C), a nonkeratinized stratified squamous 9-5
epithelium. Bowman’s membrane (B) is a collagenous basal lamina
deep to the epithelium. The bulk of the cornea is the stroma (S), made CANAL OF SCHLEMM A venous sinus known as the canal of Schlemm
of highly organized collagen fibers. Lastly, Descemet’s membrane (D) (cS) is at the junction of the cornea (C) and sclera (S). It returns
and the simple squamous corneal endothelium (E) are posterior to the aqueous humor from the anterior chamber to the blood stream. (X60)
stroma. (X60)
CB
C
F
9-7
CILIARY BODY In the anterior of the eye, the vascular tunic is expanded
to form the ciliary body (CB). Also visible are the cornea (C), iris (I),
9-6 and lens (L). (X20)
CHOROID The vascular tunic is made up of the choroid (C), a pig-
mented layer between the fibrous (F) and neural (N) tunics. (X250)
106
CP
CB
S
9-8 *
CILIARY PROCESSES Ciliary processes (CP) are extensions
of the ciliary body (CB) that produce aqueous humor. The
space (*) is an artifact of preparation where the sclera (S) C
has separated from the ciliary body. (X100)
AC
I
P
PC
9-9a
AC
PM
PC
IRIS (a) The iris (I) divides the anterior cavity
into an anterior chamber (AC) and a posterior
chamber (PC). Also visible are the lens (L),
pupil (P), sclera (S), cornea (C), and the
posterior cavity with the vitreous body (V).
(X6) (b) This micrograph shows the iris in
detail. Visible is the layer of melanocytes (M)
on the posterior and the pupillary muscle (PM)
within. (X125) 9-9b
107
LC
SE
AC L PC
LF
9-10a 9-10b
LENS (a) The lens (L) divides the eye into anterior (AC) and posterior cavities (PC). In this micrograph, suspensory ligaments are not visible. The
iris also appears to lack a pupil, but this is due to the plane of the section. (X7) (b) In this higher magnification of the lens, the lens capsule (LC),
subcapsular epithelium (SE), and lens fibers (LF) are visible. (X250)
PC ILM
O
IP
R
IN
BS
OP S
ON
P
ELM
PE
O
C D
9-11
RETINA The posterior cavity (PC) is at the top of this micrograph and 9-12
the choroid (C) is at the bottom. In between is the retina. The ten
layers of the retina are labeled as follows: PE 4 pigmented epithelium, OPTIC NERVE Where the optic nerve (O) leaves the posterior of the eye,
P 4 photoreceptive layer (rod and cone segments), ELM 4 external there is no retina (R). This results in a blind spot (BS) in each eye.
limiting membrane, ON 4 outer nuclear layer, OP 4 outer plexiform Notice that the sclera (S) is continuous with the dura mater (D) of the
layer, IN 4 inner nuclear layer, IP 4 inner plexiform layer, G 4 optic nerve. (As an outgrowth of the brain, the optic nerve is better
ganglion cell layer, O 4 optic nerve fiber layer, ILM 4 inner lining referred to as the optic tract, so it is covered with dura mater, not
membrane. (X125) epineurium.)
108
G
E
S
T
S
9-13 C
EYELID The eyelid is covered with a highly folded thin skin (S) on its
anterior and a stratified columnar epithelium, the conjunctiva (C), 9-14
on its posterior. The skeletal muscle (M), fibrous tarsal plate (T), and
PINNA The external ear is covered with skin (S) which is tightly bound
tarsal (Meibomian) glands (G) are also visible. (X25)
to an internal framework of elastic cartilage (E). Numerous hair
follicles are also visible (arrows). (X65)
O
V G
SC
9-16
9-15
MACULA The vestibule of the inner ear houses the membranous utricle and
INTERNAL EAR This section of temporal bone shows much of saccule, each with a macula. This micrograph shows a macula with otoliths
the inner ear, a complex collection of cavities. The cochlea (C), (O) embedded in a glycoprotein (G) layer that covers the receptive hair cells
semicircular canals (SC), and vestibule (V). Also shown is the (H). The macula provides sensory information about head position. (X250)
auditory tube (A). (X8)
109
HC
M CA
9-17a 9-17b
AMPULLA OF A SEMICIRCULAR CANAL Each of the three semicircular canals has a dilated end called the ampulla. Within the ampulla is the crista
ampullaris, a receptor organ that provides sensory information about body movement. (a) The bony (B) and membranous (M) labyrinths of
the vestibule are labeled in this micrograph. Note that the membrane is unnaturally collapsed due to preparation of the specimen. The crista
ampullaris (CA) is also visible. (X50) (b) In this detail of the crista ampullaris, the hair cell layer (HC) is visible, but much of the cupula (the
glycoprotein layer covering the crista) is missing. (X100)
SV
SM
ST
9-18
DETAIL OF COCHLEA The membranous labyrinth is represented by the scala media (SM), which is
separated from the bony labyrinth by the vestibular membrane (V) and basilar membrane (B).
The bony labyrinth consists of the scala vestibuli (SV) and scala tympani (ST). The organ of
Corti (O) rests on the basilar membrane. (X100)
110
SM
TM
IH OH
OH
IH
P tC
P
ST BM
9-19
ORGAN OF CORTI The organ of Corti is the receptor for the sensation of hearing. The actual
receptors are hair cells, of which there are two types: an inner hair cell (IH) and outer hair cells
(OH). Other parts of the organ of Corti, such as pillar cells (P), other support cells, tunnel of
Corti (tC), basilar membrane (BM) and the tectorial membrane (TM) are also visible. The scala
media (SM) is above the organ of Corti and the scala tympani (ST) is below it. (X250)
SG
9-20
SPIRAL GANGLION The nerve cell bodies whose axons form the auditory nerve are located in
spiral ganglia (SG) within the modiolus. They transmit sensory impulses from the hair cells to
the cerebral cortex to produce the sensation of hearing. (X250)
Endocrine System
C H A P T E R
10
Introduction to the Endocrine System about the size of a garden pea. Its three parts have two dif-
The endocrine system is composed of endocrine glands and ferent embryological origins: the adenohypophysis (anterior
endocrine cells. The former are multicellular organs, and pituitary) and pars intermedia are derived from oral ecto-
the latter are cells dispersed within organs with more than derm, and the neurohypophysis (posterior pituitary), which
an endocrine function. is an outgrowth of the hypothalamus (Figure 10-1).
Endocrine glands are typically epithelial in origin and
are produced as a result of surface cells growing down into Adenohypophysis (Anterior Pituitary)
deeper tissues. Unlike exocrine glands that maintain their
Two main cell types are recognized in the adenohypophysis
connection with the surface (by way of their duct), the con-
using traditional staining methods. These are the easily
nection of endocrine glands is lost during development and
stained chromophils and the poorly staining chromophobes
the glands are “ductless.” Each gland produces one or more
(Figure 10-2). Chromophils are further differentiated into
peptide or lipid hormones, which are secreted into the
acidophils and basophils, depending on their affinity for
blood stream. For this reason, endocrine glands are very
acidic and basic stains, respectively. Immunohistochemical
vascular. The hormones have a physiological effect on spe-
techniques have resulted in categorizing these cells based on
cific targets, cells in organs elsewhere in the body. In this
their secretion. Somatotrophs and mammotrophs secrete
way, the endocrine system regulates activities in concert
growth hormone (GH) and prolactin, respectively, and are
with the nervous system.
acidophils. The basophilic corticotrophs, thyrotrophs, and
Endocrine glands are structurally diverse, but they do
gonadotrophs secrete adrenocorticotropic hormone (ACTH),
have some general features in common. The secretory cells
thyroid stimulating hormone (TSH), and follicle stimulating
are referred to as parenchyma. There is also often a connec-
hormone (FSH) and luteinizing hormone (LH), respectively.
tive tissue capsule and, of course, the numerous capillaries
Chromophobes are thought to be chromophils that
into which hormones are secreted.
have discharged their secretory granules (i.e., degranulated).
Endocrine glands covered in this chapter are the pitu-
itary gland, thyroid gland, parathyroid glands, adrenal
glands, pineal gland, pancreas, testes, and ovaries. Neurohypophysis (Posterior Pituitary)
The neurohypophysis consists of unmyelinated fibers of
Pituitary Gland (Hypophysis) neurosecretory cells whose cell bodies are located in the
The pituitary gland is located inferior to the hypothalamus hypothalamus (Figure 10-3). Glial-like pituicytes support
and occupies the sella turcica of the sphenoid bone. It is the fibers.
111
112
Pars Intermedia and Pars Tuberalis its cells stain more intensely than the spongiocytes. These
The pars intermedia consists of basophilic cells located be- cells secrete small amounts of sex hormones.
tween the anterior and posterior pituitary and is visible in The adrenal medulla (Figure 10-7) is essentially a gan-
some specimens. Small cysts with acidophilic colloidal ma- glion of the sympathetic nervous system. Its cells are the de-
terial characterize it and produce melanocyte stimulating velopmental and functional equivalents of sympathetic
hormone. postganglionic neurons. They secrete the hormones epi-
The pars tuberalis is found associated with the in- nephrine and norepinephrine that act along with direct
fundibular stalk of the pituitary. Its function is not known. sympathetic innervation to prepare the body for dealing
with stressful situations. The secretory cells are basophilic
Thyroid Gland and have large nuclei.
The thyroid gland is found anterior to the larynx in the
neck. It is composed of two lobes and is surrounded by a Pineal Gland
thin capsule. Connective tissue septa divide the thyroid The pineal gland is a small outgrowth from the roof of the
gland into lobules. third ventricle (Figure 10-8). Modified neurons called
Unlike most endocrine glands, the thyroid does not pinealocytes secrete the hormones melatonin and serotonin,
store its secretion intracellularly. Rather, the inactive hor- and are characterized by round nuclei with prominent nu-
mones are stored in the lumina of thyroid follicles (Figure cleoli. Interstitial cells, which are structurally and function-
10-4). Each follicle is formed by a simple cuboidal epithe- ally similar to astrocytes, are found surrounding clusters of
lium that produces the hormones T3 and T4, which are re- pinealocytes. Pineal sand (corpora arenacea), made of cal-
sponsible for regulating metabolic rate. In addition, the cium phosphate and calcium carbonate, becomes more
much less abundant clear (C) cells produce the hormone abundant with age, but is of unknown function.
calcitonin, which regulates blood calcium levels.
Pancreas
Parathyroid Glands The pancreas has endocrine and exocrine components.
The parathyroid glands are located on the posterior of the Islets of Langerhans comprise the endocrine portion (Figure
thyroid gland. Each is enclosed by a fibrous capsule that 10-9). The majority of cells (up to 70%) in an islet are the
sends septa into the gland and divides it into lobules (Figure insulin secreting b cells. The glucagon secreting a cells are
10-5). Parathyroid hormone is involved in regulating blood toward the periphery of the islet and comprise about 20%
calcium and phosphate levels. of the cells. These hormones decrease and increase blood
The parenchyma is composed of hormone secreting sugar levels, respectively. The remaining cells are of three
chief cells and oxyphil cells of unknown function. Chief types, defined by their secretions: somatostatin, gastrin, and
cells are small with round, centrally positioned nuclei and pancreatic polypeptide. Typical histological stains do not
pale, eosinophilic cytoplasm. Oxyphils are larger with an distinguish between the five different cells that manufacture
abundance of eosinophilic cytoplasm. and secrete each chemical, but they can be differentiated
with immunocytochemical stains specific for their secretions.
Adrenal Glands
The adrenal glands are located superior to each kidney. Testes
They consist of a fibrous capsule surrounding the paren- The testes have a cytogenic function (sperm production)
chyma, which is divided into an outer cortex and an inner and an endocrine function. The interstitial cells of Leydig
medulla (Figure 10-6a). are located in the vascular connective tissue between semi-
The adrenal cortex is divided into three layers. The zona niferous tubules of the testes (Figure 10-10). They are
glomerulosa (Figure 10-6b) is a thin layer deep to the cap- responsible for secretion of testosterone, the male sex hor-
sule. Its cells are in rounded clusters and have dense staining mone. The cells have a round nucleus with one or two
nuclei. They secrete the mineralocorticoid hormone aldos- eccentric nucleoli and lipids in the cytoplasm.
trerone, which increases blood pressure by increasing sodium
reabsorption in the kidney tubules. The zona fasciculata (Fig- Ovaries
ure 10-6c) is deep to the zona glomerulosa and is the thick- Like the testes, ovaries have a cytogenic and an endocrine
est of the three cortical layers. Its cells are arranged in rows function. As ovarian follicles develop (Figure 10-11), the
and are called spongiocytes (Figure 10-6d) due to their ap- ovum undergoes meiosis and the surrounding follicle cells
pearance from the abundant cytoplasmic lipids. The main proliferate. The thecal cells of developing ovarian follicles
hormone is cortisol, which raises blood glucose levels among produce estrogen, one of the female sex hormones. After
other metabolic effects. The zona reticularis (Figure 10-6e) ovulation, the ovarian follicle undergoes changes and devel-
is the innermost cortical layer. The nuclei and cytoplasm of ops into a corpus luteum (Figure 10-12). The granulosa
C H A P T E R T E N : E N D O C R I N E S Y S T E M
113
lutein cells secrete progesterone and some estrogen. If preg- degenerates and the placenta assumes the role of hormone
nancy occurs, the corpus luteum continues growing and se- production. If pregnancy does not occur, the corpus luteum
creting hormones for the first trimester, after which it degenerates approximately within two weeks of ovulation.
PT
C
AP
PP
PI
10-1
PITUITARY GLAND The drastically different appearances of the anterior 10-2
pituitary (AP) and posterior pituitary (PP) are consistent with their ANTERIOR PITUITARY GLAND The secretory cells of the anterior pituitary
different embryonic origins. The anterior pituitary develops from oral gland can be differentiated with H&E stain into chromophils, which
ectoderm, whereas the posterior pituitary grows from the hypothalamus. stain intensely, and chromophobes (C), which don’t. The former can
Also visible are the infundibular stalk (I), pars tuberalis (PT), and pars further be differentiated into reddish acidophils (A) and purplish
intermedia (PI). (X7) basophils (B). Further identification requires immunohistochemical
staining for the specific hormones. Note the numerous blood vessels
surrounding the secretory cells. (X380)
POSTERIOR PITUITARY The bulk of the posterior

pituitary is composed of unmyelinated fibers
from neurons in the hypothalamus. These are
10-3 supported by glial-like pituicytes (P). (X250)
114
10-4a 10-4b
10-4c 10-4d
THYROID GLAND (a) The follicular nature of the thyroid gland is apparent in this micrograph. (X65) (b) This is a higher magnification of the
boxed region in (a). Each follicle is made of simple cuboidal cells that secrete inactive thyroid hormones into its lumen. (X250) (c) Larger and
paler staining clear cells (C) are found between follicles. They secrete calcitonin, a hormone involved in regulating blood calcium levels. (X250)
(d) This is a higher magnification of clear cells. Notice the thyroid follicles surrounding them. (X380)
115
10-5a 10-5b
PARATHYROID GLAND (a) The parathyroid glands are physically
associated with the thyroid gland. In this micrograph, the thyroid
gland is on the left and the parathyroid is on the right. The thin
fibrous capsule (C) and a septum (S) are also visible. (X130) (b) The
parathyroid secretory cells, chief cells, are organized into clusters
C
or strands. They are small with round nuclei and a pale cytoplasm.
(X130) (c) Oxyphil cells (O) are found in clusters and are larger
than chief cells (C). Their number increases with age. (X380)
10-5c
116
ZG
ZF
ZG
ZR
ZF
10-6b
10-6a
ZF
10-6d
ZF
10-6c
ADRENAL CORTEX (a) The zona glomerulosa (ZG), zona fasciculata
(ZF), and zona reticularis (ZR), which comprise the adrenal cortex,
and the adrenal medulla (M) are visible in this micrograph. (X65)
(b) The zona glomerulosa cells are in rounded clusters deep to the
fibrous capsule (C). These cells secrete aldosterone. The zona ZR
fasciculata is also visible. (X250) (c) The zona fasciculata cells are
arranged in columns. The cells are called spongiocytes. (X130) (d)
Spongiocytes (S) have a pulpy appearance due to cytoplasmic lipids.
They secrete cortisol. (X130) (e) The thin zona reticularis stains
more intensely than the neighboring zona fasciculata. These cells
secrete small amounts of sex hormones. (X130)
10-6e
117
ZR
10-7 10-8
ADRENAL MEDULLA The cells of the adrenal medulla (M) are modified PINEAL GLAND The pinealocytes (P) are modified neurons with a round
neurons that secrete epinephrine and norepinephrine. It is found at the nuclei and obvious nucleoli. They are arranged in clusters associated
core of the adrenal gland. Note the zona reticularis (ZR) surrounding with numerous capillaries (C). Also present are glial cells (G). (X250)
the adrenal medulla. (X130)
PA
PA
I b
I
10-9a 10-9b
PANCREAS (a) Islets of Langerhans (I) comprise the endocrine portion of the pancreas. Surrounding the islets are the exocrine pancreatic acini
(PA). (X135) (b) Immunocytochemical stains allow recognition of five cell types in the islet, but H&E stains (used in this specimen) do not, so
all cells look alike. Insulin secreting b cells (b) are the most abundant cells of the islet and tend to be located in the center. The a cells (a), which
secrete glucagon, tend to be found near the periphery. (X250)
118
ST ST
ST
I
I
*
10-10a ST
10-10b
I
TESTICULAR INTERSTITIAL CELLS OF LEYDIG (a) The interstitial
cells (I) are found between seminiferous tubules (ST). (X65)
(b) Interstitial cells (I) secrete testosterone and can be identified
based on their location between seminiferous tubules and their
obvious, eccentric nucleoli. The space (*) between the interstitial
cells and the seminiferous tubules is an artifact of preparation.
ST (X250) (c) This is another example of interstitial cells. These
would have to be identified based on their position between
I seminiferous tubules, since nuclear staining is not very good.
(X250)
10-10c G
C
OVARY THECAL AND GRANULOSA CELLS

During development, thecal (T) and
granulosa (G) cells in the ovarian follicle
have an endocrine function and secrete
estrogen. Note the abundance of capillaries
(C). (X250) 10-11
119
CL
CL
10-12a 10-12b
CORPUS LUTEUM After ovulation, the follicle cells remaining in the
ovary develop into a corpus luteum (CL), an endocrine gland respon-
sible for secreting progesterone and some estrogen. (a) This is a
corpus luteum after ovulation. If pregnancy does not occur, it will
degenerate. (X130) (b) If pregnancy does occur, the corpus luteum
continues enlarging and secreting hormones. In fact, it can become
GL almost the size of the ovary. (X25) (c) The actual secretory cells are
the large, pale staining granulosa lutein cells (GL). (X250)
10-12c
Integumentary System
C H A P T E R
11
Introduction to the Integument The main epidermal cells are called keratinocytes. The
The integumentary system consists of the skin and its ap- basal cells are the healthiest due to their proximity to the
pendages, such as hair, nails, and various glands. It func- dermal capillaries, and they are the ones that undergo mito-
tions as a covering for the entire body. In this role, it sis. As they do, they push preceding generations of cells
protects against mechanical damage and presents a barrier toward the surface and down the oxygen concentration
to penetration by chemicals and infectious agents. It also gradient. Eventually, the keratinocytes occupy a position
is involved in sensation, thermoregulation (through sweat where they can’t get enough oxygen to satisfy their meta-
glands and regulating blood flow), and vitamin D synthesis. bolic needs and they die. This process involves degeneration
It is the largest organ in the body. The average thick- of the nucleus and accumulation of the protein keratin, and
ness of the skin is about 1 to 2 mm, but it tends to be is reflected in the layers seen in epidermis.
thicker on dorsal surfaces than on ventral surfaces (the The epidermis of thick skin presents five distinct layers
palms of the hands and soles of the feet are exceptions to (Figure 11-3), whereas in thinner skin only three are gener-
this). ally visible. From deep to superficial (which coincides with
the stages of development), these are the stratum basale,
Layers of the Integument stratum spinosum, stratum granulosum, stratum lucidum,
The skin is composed of two layers: the superficial epider- and stratum corneum. The stratum basale is the deepest
mis and the deeper dermis (Figure 11-1). Deep to the der- layer of the epidermis. It is composed of healthy (due to
mis is the hypodermis, also known as superficial fascia. their proximity to the dermal capillaries) cuboidal to low
Where it is replaced with adipose tissue, it is known as sub- columnar cells that undergo mitosis. The stratum spinosum
cutaneous fat. While the hypodermis is not part of the in- is the thickest layer. It is characterized by the “prickly” ap-
tegument, it will be covered in this chapter since it is visible pearance of its cells due to the desmosomes that attach ad-
on most skin slides and some epidermal appendages pene- jacent cells. The deeper cells continue mitotic activity, but
trate it. this ability is lost in the more superficial layers. The stratum
granulosum is characterized by cells containing dark stain-
Epidermis ing keratohyalin granules, an indication that they have
The epidermis is derived from ectoderm and is composed of begun to die. It is usually about five cells thick. Secretion of
keratinized stratified squamous epithelium. Its thickness a lipid material by these cells makes the epidermis water-
ranges from 0.1 mm (over most of the body) to 1.4 mm (on proof. The stratum lucidum is characterized by cells that
the soles of the feet). Figure 11-2 illustrates thick and thin have lost their nuclei and have accumulated keratin. It is
skin. only seen in thick skin. The most superficial layer of the
121
122
epidermis is the stratum corneum. It consists of dead, anu- Hair Follicles

cleate cells that have accumulated abundant keratin. As The hair follicle (Figure 11-9) is an angular downgrowth of
these cells approach the surface, they lose their intercellular the epidermis into the dermis or hypodermis. The base of the
attachments (desmosomes) and are sloughed off. follicle is dilated to form the hair root, which is penetrated by
Besides keratinocytes, other cells are seen in the epider- a dermal papilla that houses capillaries. Together, the root
mis. Langerhans cells (Figure 11-3c) are a component of the and papilla form the hair bulb. The follicle is surrounded by
immune system and function as antigen presenting cells. a thick basement membrane called the glassy membrane,
They are characterized by a dark staining nucleus with pale which itself is wrapped in dense connective tissue.
cytoplasm and numerous cytoplasmic extensions (hence The hair is made of keratinized cells. It is produced by
their other name, “dendritic cells”). They are found in many dividing cells at the base of the follicle called the hair ma-
places, but are most abundant in the stratum spinosum. trix, which is the functional equivalent of the stratum
Melanocytes (Figure 11-4) are pale staining cells found in basale of the epidermis. Thus, the hair grows from its base,
the stratum basale. They produce the brown/black pigment not its tip. Cells become keratinized in a mechanism similar
melanin in membrane-bound structures known as melano- to that seen in the epidermis. The portion of the hair above
somes that are deposited in the cytoplasm of keratinocytes the skin’s surface is called the hair shaft. Frequently, the
of the stratum spinosum. The melanin accumulates near the hair falls out of the follicle during slide preparation, so all
nucleus on the side closest to the surface where it protects that is seen is the follicle and associated structures.
the nucleus by absorbing ultraviolet radiation. Figure 11-4 Two types of hairs are present in humans. Vellus hairs
shows pigmented skin. Merkel cells are found in the stra- are short, fine, and microscopic. These comprise the ma-
tum basale of fingertips and oral mucosa, and have in- jority of hairs on the human body. Terminal hairs are larger
dented nuclei. They are thought to act as mechanoreceptors. and coarser. These are the hairs everyone recognizes as
hairs.
Dermis In cross section, the hair and follicle present several lay-
The dermis is deep to the epidermis and is the “true skin.” ers (Figure 11-10).
The surface in contact with the epidermis is highly folded ◗ The medulla is at the center of the hair. It is most obvi-
into elongated dermal ridges or conical dermal papillae ous in the root; in the shaft the cells are cornified or
(Figure 11-5). These interlocking surfaces anchor the epi- completely absent.
dermis to the dermis and resist separation of the two layers ◗ The cortex makes up the bulk of the hair. In the root,
when subjected to shearing forces. Dermal ridges are seen the cortex is made of cuboidal cells, but in the shaft the
on the surface of the palm and sole as fingerprints. cells are flattened and keratinized. Pigment granules
The portion of the dermis in contact with the epidermis and air spaces between the cortical cells produce hair
and comprising the dermal papillae is a looser and finer color.
connective tissue and forms the papillary layer. The papil-
◗ The cuticle of the hair is made of hard keratin and sur-
lae contain capillary loops and tactile receptors called
rounds the cortex.
Meissner’s corpuscles (Figure 11-6). The majority of the
dermis is composed of a vascular, dense irregular connec- ◗ The internal root sheath extends from the base of the
tive. This is the reticular layer (Figure 11-5). Fibroblasts are follicle to where the sebaceous gland enters. It consists
the most common cell. Large blood vessels, nerves, and epi- of three layers. The cuticle of the internal root sheath is
dermal appendages are present in the reticular layer. Pres- similar in construction to the cuticle of the hair and its
sure receptors called Pacinian corpuscles may also be seen cells interdigitate with it. Huxley’s layer consists of a
(Figure 11-7). couple of layers of flattened cells. Henle’s layer is a sin-
gle row of flattened rectangular cells.
Hypodermis ◗ The external root sheath is composed of several cell
The hypodermis consists of loose connective tissue and layers. The single layer of cells in contact with the con-
anchors the skin to the underlying tissues without binding nective tissue sheath is columnar; the remaining cells
it too tightly. This allows free movement of underlying are polygonal in shape. These cells are derived from the
muscles without pulling on the skin. The loose connective outermost part of the matrix and are separated from
tissue may be replaced by fat (Figure 11-8). Hair follicles, the connective tissue sheath by the glassy membrane.
Pacinian corpuscles, and sweat glands may be seen.
Sebaceous Glands
Appendages of the Skin Hair follicles are associated with branched acinar sebaceous
The epidermis gives rise to a variety of appendages. These glands (Figures 11-9a and 11-11). Gland cells near the folli-
are nails, hair follicles, and sweat glands. cle disintegrate and release the oily substance sebum into
C H A P T E R E L E V E N : I N T E G U M E N T A R Y S Y S T E M
123
the follicle, which moisturizes and lubricates the hair. Divi- distal sides of the fingers and toes. The main part of the nail
sion of cells at the base of the gland replaces these cells. Be- is the nail plate that ends distally as the free edge, the part
cause sebaceous glands are thicker than their respective that gets trimmed. At the proximal end is the nail root that
follicle, these glands are often seen in specimens without lies beneath a fold of skin. The nail rests on an epidermal
their follicle. layer (corresponding to the stratum basale and stratum
spinosum) called the nail bed. At the proximal end, the nail
Arrector Pili Muscles
bed is thickened to form the nail matrix. It is responsible
Arrector pili muscles (Figures 11-9a and 11-12) are smooth for producing the nail in a process similar to hair produc-
muscles that insert on the follicle’s connective tissue sheath. tion. The stratum corneum folds over the proximal end of
They are positioned in the obtuse angle formed by the folli- the nail plate as the eponychium and under the free edge as
cle and the skin surface. Their contraction straightens the hyponychium.
hair, which in other mammals causes the fur to form a
thicker insulating layer, but in humans only pro-
duces “goose bumps.” The sebaceous gland is
found in the angle formed between the arrector
pili and the follicle. E
Sudoriferous (Sweat) Glands

Sudoriferous or sweat glands are downgrowths
of the epidermis. They are simple, coiled tubu-
lar glands of two types. Merocrine sweat glands
(Figures 11-13a and 11-13b) are more numer-
ous and produce a watery secretion as a stress
response or a cooling response. The coiled se-
cretory portion is made of simple cuboidal ep-
ithelium and the lumen is small. The relatively
straight duct leading to the surface is made of
two cell layers. Apocrine sweat glands (Figure
11-13d) are found in the axillary and groin re- D
gions. The lumen of the secretory portion is
wider than in merocrine glands and the secre-
tion is more viscous.
Nails
Nails (Figure 11-14) are made of keratinized
cells that form a hard plate on the dorsal and
THE INTEGUMENT The integument consists

of the superficial epidermis (E), a keratinized
stratified squamous epithelium, and the deeper
dermis (D), a dense irregular connective tissue.
The hypodermis (H) is deep to the dermis and
is adipose tissue in this specimen. (X65) 11-1
124
11-2a 11-2b
THICK AND THIN SKIN (a) This specimen is from the sole of the foot and is representative of thick skin. Note the thick keratinized layer (K) on the
surface. (X65) (b) Thin skin has a much thinner keratinized layer that appears shredded. (X130)
SC
SL
SG
EPIDERMAL LAYERS The epidermis of thick skin presents five
SS layers, each representing a stage in the life of a keratinocyte.
From deep to superficial, these are the stratum basale (SB),
stratum spinosum (SS), stratum granulosum (SG), stratum
lucidum (SL), and stratum corneum (SC). (a) X130 (b) X265
SB (c) The stratum spinosum is characterized by cells with spiny
intercellular junctions. Note the Langerhans (dendritic) cell
(L). (X660)
11-3a
L
SC
SL
SG
SS
SB
11-3b 11-3c
125
11-4a
11-4b
PIGMENTED SKIN Skin accumulates a brown to
black pigment called melanin. This pigment
accounts for the color of darker skinned races,
and the tanning in lighter skinned races. (a) This
is pigmented thin skin. Note the abundance of
melanin in the basal layers of the epidermis.
(X265) (b) This thin skin specimen is pigmented,
but not as much as (a). Note the melanocytes (M).
(X265) (c) Melanin is produced by melanocytes.
These are large, pale staining cells found in the
basal layers of the epidermis. Also note the spiny
cells of the stratum spinosum. (X660)
11-4c
126
DP
DP
11-5a
DERMIS (a) Dermal papillae (DP) project into
the epidermis. The irregular margin between
the two layers increases surface area and limits
separation due to shearing forces. (X130) (b)
This micrograph shows capillaries (C) in the
dermal papillae. (X260) (c) The papillary layer
(P) of the dermis is in contact with the epidermis
and is made of a loose connective tissue. The C
deeper reticular layer (R) is thicker and much
coarser. The difference in texture is obvious in C
this microgram. (X130)
11-5b
11-5c
127
MC
MEISSNER’S CORPUSCLES
Meissner’s corpuscles
(MC) are receptors for
light touch found in
dermal papillae. (X210) 11-6
11-7
PACINIAN CORPUSCLES These pressure
receptors (P) look like a sliced onion in
section and are found in the dermis and D
hypodermis. (X130)
S
HYPODERMIS Deep to the dermis
(D) is a layer of loose connective
tissue or adipose tissue (A) that S
loosely binds the integument to
the underlying structures. It allows
free movement of muscles without A
pulling on the skin. This micro- A
graph shows adipose tissue and
two sweat glands (S) in the hypo-
dermis. (X130) 11-8
128
ERS
SG
AP
HB
11-9b
Ma
11-9a
HAIR FOLLICLE Hair follicles are epidermal invaginations into the dermis.
They form an angle with the surface and rarely are seen in their entirety
due to the sections being so thin. (a) This shows a fairly complete hair GM
follicle, but pieces of other follicles are also visible (arrows). Sebaceous
glands (SG) and an arrector pili muscle (AP) can also be seen. (X50)
(b) The deepest part of the follicle is dilated and forms the hair bulb
(HB). Also visible are the cortex (C), and external root sheath (ERS).
(X110) (c) Each follicle has a dermal papilla (DP) that has a capillary DP
and a nerve. It is separated from the hair root cells by a basement
membrane (B), which is continuous with the glassy membrane (GM)
on the outside of the follicle. Note the large amount of pigment in the
hair matrix cells (Ma). (X400)
11-9c
GM
Cu
CTS
ERS
IRS
IRS
ERS
C
C IRS
GM
ERS
CTS
GM
11-10a CTS 11-10b
HAIRS IN CROSS SECTION (a) This cross section of two hair follicles was made at the level of the hypodermis. Key to the symbols used: C 4
cortex, Cu 4 cuticle, IRS 4 internal root sheath, ERS 4 external root sheath, GM 4 glassy membrane, CTS 4 connective tissue sheath.
(X130) (b) This section was made higher in the dermis. Symbols used are the same as in (a). (X260)
129
SG
11-11a
SEBACEOUS GLANDS Cells of sebaceous glands (SG)
are large and often have a foamy appearance. Be-
cause they are larger than their associated follicle (F),
sebaceous glands are often seen with no follicle in the
section. (a) The sebaceous gland on the left is seen to
empty into the follicle (arrow). The ducts of the others F
are out of the plane of section. (X130) (b) Shown here
is a follicle cut in oblique section. The sebaceous SG
glands show no connection with it. (X130) (c) In this
cross section, the size of sebaceous glands relative to
their follicle is seen. (X250)
SG
11-11b
SG
SG
SG
11-11c
130
AP
CTS
11-12a
AP
SG
11-12b
ARRECTOR PILI MUSCLES Contraction of an arrector pili muscle causes the hair to straighten. (a) This
micrograph shows an arrector pili (AP) muscle attachment to the connective tissue sheath (CTS) of
a follicle cut in oblique section through the dermis (D). (X130) (b) It is very difficult to find a
perfectly sectioned hair follicle, so one has to make do with what is available. In this micrograph, no
follicle is present, but the sebaceous gland (SG) and arrector pili muscle are seen in the dermis.
(X130)
131
D
D
11-13a 11-13b
11-13c 11-13d
SWEAT GLANDS Sweat glands are of two types—merocrine and apocrine. (a and b) Here are two merocrine sweat glands in the hypodermis. The
secretory portion (S) is highly coiled, has large pale staining cells, and a small lumen. Myoepithelial cells (M) are associated with the secretory
portion. The duct (D) is lined with two layers of darker staining cells. (X260) (c) This micrograph shows a sweat gland duct as it passes through
the stratum corneum. (X210) (d) Apocrine sweat glands are found in the axillary and genital regions. The secretory portion is coiled and tubular
with a large lumen. The ducts empty into hair follicles and are similar in construction to the ducts of merocrine sweat glands. (X210)
132
E
NR
NP
NB
NM
11-14a
NP
NB
NR
NM
DP
11-14b
NAILS (a) This micrograph is of a fetal finger cut in sagittal section. Notice the continuity of the
stratum corneum with the eponychium and hyponychium. (X25) (b) This micrograph shows greater
detail of the nail root. (X65) Key to symbols used: NB 4 nail bed, NM 4 nail matrix, NP 4 nail
plate, NR 4 nail root, E 4 eponychium, H 4 hyponychium, and DP 4 distal phalanx.
Cardiovascular System
C H A P T E R
12
Introduction to the Cardiovascular System (intima) interna, tunica media, and tunica (adventitia) ex-
The cardiovascular system consists of blood pumped by the terna. They are best demonstrated in arteries (Figure 12-1).
heart through the blood vessels to all parts of the body. Its The tunica interna is a simple squamous endothelium
function is to carry materials from organs that exchange supported by a thin layer of connective tissue. In some ves-
with the environment to and from cells buried deep in the sels (arteries), an internal elastic (lamina) membrane is also
body. Oxygen is picked up in the lungs and distributed present at the junction with the tunica media. The tunica
throughout the body, whereas carbon dioxide is picked up media is composed of varying amounts of smooth muscle
from the cells and delivered to the lungs where it is exhaled. and elastic connective tissue oriented circularly around the
Food absorbed by the digestive tract is distributed through- vessel. In larger arteries, there may be an external elastic
out the body, and wastes are picked up from cells and re- lamina or membrane at the junction with the tunica ex-
moved from the blood by the kidneys. terna. The tunica externa is composed of fibrous connective
The human circulatory system consists of two closed- tissue that often blends with surrounding connective tis-
loop paths: the pulmonary circuit and the systemic circuit. sues, making the outer limit of the vessel difficult to see.
In the adult, oxygen-poor blood is returned to the right If the vessel’s walls are thicker than the effective dis-
atrium by the superior and inferior venae cavae. From here, tance for diffusion of oxygen and nutrients, the wall may
it moves to the right ventricle and is pumped out into the be penetrated with small blood vessels called vasa vasorum.
pulmonary trunk where it is then sent to the lungs to pick They are especially common in larger veins due to the lower
up oxygen. It is returned to the left atrium by the pul- oxygen concentration in systemic venous blood.
monary veins. This completes the pulmonary circuit. From
Types of Blood Vessels
the left atrium, the now oxygen-rich blood is sent to the left
ventricle and is then pumped out the aorta to be distributed Blood vessels are of three main types: arteries, veins, and
to the capillaries of the entire body. Blood in the capillaries capillaries. Each has a distinctive structure appropriate to its
loses its oxygen to the surrounding tissues, picks up CO2 function. Arteries, veins, and nerves typically travel together
and is sent back to the right atrium. This completes the sys- to form a neurovascular bundle (Figure 12-2).
temic circuit. Arteries carry blood away from the heart at high pres-
sure. As such, their walls are thicker than their correspon-
Basic Blood Vessel Structure ding vein. The two arteries that exit the heart are the aorta
Blood vessels are composed of as many as three layers or and pulmonary trunk. These branch into smaller and smaller
tunics that are modified according to the vessel’s function. arteries and eventually lead to the capillaries of the systemic
From innermost tunic to outermost these are the tunica circuit and pulmonary circuit, respectively. As arteries get
133
134
smaller, the proportion of elastic tissue decreases and the they carry blood at low pressure and against the pull of
smooth muscle increases (relative to the wall’s thickness). gravity, many veins are supplied with endothelial folds that
As viewed under the microscope, arteries typically have a form valves. As viewed under the microscope, veins often
circular lumen, distinct layers, and the tunica media is the are collapsed and have an irregularly shaped lumen, the
thickest of the three (Figure 12-3). layers are indistinct and the thickest layer is the tunica ex-
Elastic arteries (Figure 12-3) are the largest and include terna (compare with the artery in Figure 12-2).
the pulmonary trunk and aorta, and their major branches. The smallest veins are called venules (Figure 12-5).
The tunica media is dominated by elastic tissue in the form Venules are structurally similar to capillaries, but are larger
of fenestrated membranes, which alternate with smooth in diameter. As they emerge from the capillary bed, pericytes
muscle. Internal and external elastic membranes are pres- are present, but these are replaced with smooth muscle in
ent. The tunica adventitia is very thin and often has vasa the tunica media as the venules get larger. High endothelial
vasorum. venules are found in certain lymphatic organs and have a
Muscular arteries (Figure 12-4) are larger than 0.5 mm cuboidal endothelium.
in diameter and include most of the named arteries. The Medium veins are less than 1 cm in diameter (Figure
internal elastic membrane is wavy in appearance and the 12-7). Their tunica intima is composed of endothelium and
tunica media is composed of up to 40 layers of smooth connective tissue, but there is no internal elastic membrane.
muscle cells. An external elastic membrane may be present The tunica media consists of smooth muscle, and the tunica
and the tunica adventitia has collagen, and elastic and smooth adventitia has collagen and elastic fibers, and smooth muscle.
muscle fibers oriented longitudinally along the vessel. Valves formed by pockets of tunica intima are present in
Arterioles (Figure 12-5) are the smallest arteries, with a medium veins that carry blood against gravity (Figure 12-8).
diameter of 30 to 200 µm. The tunica media is composed If blood flows the wrong way, the pockets fill with blood
of one to a few layers of smooth muscle. An internal elastic and the lumen is closed.
membrane is present in larger arterioles but is absent in Large veins (Figure 12-9) have a thicker tunica intima
smaller ones. The tunica adventitia is approximately the due to more connective tissue. The tunica media is gener-
same size as the tunica media. The final arteriole before the ally absent, though smooth muscle cells are present in veins
capillary bed is called a metarteriole. Blood flow into the carrying blood against gravity. The tunica adventitia is the
capillary bed is regulated by precapillary sphincters, circu- thickest layer and is composed of elastic and collagen
larly arranged smooth muscle fibers around the capillaries fibers. Vasa vasorum are also present.
emerging from the metarteriole. An arteriovenous anasto-
mosis directly connects the arteriole and venule and allows The Heart
blood to bypass the capillary bed when the sphincters are The heart is a hollow, muscular organ. It consists of four
closed. chambers: the right atrium and ventricle, and the left
Capillaries are the smallest blood vessels. The have a atrium and ventricle. The superior vena cava brings blood
diameter of 4 to 10 µm and consist only of endothelium from the head and upper limbs to the right atrium, whereas
and basal lamina. Contractile pericytes may be present on the inferior vena cava does the same for the lower limbs
capillaries (and some venules). They regulate blood flow. and abdomen. The pulmonary trunk emerges from the
Based on electron micrographs, three types of capillaries right ventricle. Pulmonary veins bring blood from the lungs
are recognized (Figure 12-6). Continuous capillaries have a to the left atrium and the aorta carries blood out of the left
complete endothelial lining. They are the most common ventricle.
and are found in muscle, nervous, and connective tissues, Valves in the heart ensure blood travels in the correct
as well as the lungs. In fenestrated capillaries, membrane- direction. Between each atrium and its ventricle is an atri-
covered pores 60 to 80 nm in diameter are present. These oventricular valve. The tricuspid valve is between the right
are found in endocrine glands, lamina propria of the small atrium and ventricle and consists of three flaps. The bicus-
intestine, and kidneys (where there is no membrane cover- pid (mitral) valve has two flaps and separates the left
ing the pores). Sinusoidal capillaries are larger than other atrium and ventricle. At the base of the aorta and pul-
capillaries and the surrounding cells determine their shape. monary trunk is a semilunar valve made of endothelial
Their discontinuous endothelium promotes exchange. They pockets similar in construction to the valves of veins. These
are found in bone marrow, the liver, spleen and lymph valves prevent backflow into the ventricle during ventricu-
nodes, as well as other places. lar relaxation (diastole).
Veins carry blood toward the heart under low pressure. The heart wall consists of three layers. The endocardium
Their walls are thinner than the corresponding artery. Veins is comparable to the tunica intima of blood vessels (Figure
begin at the capillaries and end at the heart as either the in- 12-10). It consists of endothelium plus a loose connective
ferior or superior vena cava or the pulmonary veins. Because tissue. The endocardium folds inward and is reinforced with
C H A P T E R T W E L V E : C A R D I O V A S C U L A R S Y S T E M
135
connective tissue to form the flaps of the bicuspid and tri- over the atria causing their contraction. It also reaches the
cuspid valves. The myocardium (Figure 12-11) is the middle atrioventricular (AV) node located in the myocardium near
and thickest layer of the heart wall. It is composed of car- the tricuspid valve. When stimulated, the AV node sends its
diac muscle and is highly vascular. Specialized cardiac muscle own signal via the bundle of His down the interventricular
fibers called Purkinje fibers are found in the subendocardial septum to the heart’s apex. From there, the signal spreads
region. Purkinje fibers are part of the heart’s conducting upwards through the ventricular myocardium along spe-
system (see below) and transmit impulses from the atrio- cialized cardiac muscle fibers called Purkinje fibers, causing
ventricular node to the apex of the heart. Epicardium is on contraction (Figure 12-13). Purkinje fibers are located deep
the outer surface of the heart (Figure 12-12). It is also to the endocardium and are distinctive due to their large
called visceral pericardium and is made of a simple squa- size, lack of striations and pale cytoplasm. They are some-
mous mesothelium plus underlying fibrous connective tissue, times binucleate.
which sometimes accumulates fat. It also is the layer in The cardiac skeleton is made of dense connective tissue.
which the coronary vessels travel. It separates the atria from the ventricles and is also found at
Cardiac muscle cells have an intrinsic contraction rate, the bases of the pulmonary trunk and aorta. It is an attach-
but the sinoatrial (SA) node, located at the junction of the ment site for cardiac muscle and prevents transmission of
superior vena cava and the right atrium, regulates that rate. impulses from the atria to the ventricles except via the bundle
The SA node and the following structures constitute the of His.
conduction system of the heart. The SA node’s signal spreads
TE
TM
TI
12-1
BLOOD VESSEL STRUCTURE The innermost layer in a blood vessel, the tunica interna (TI), is composed
of a simple squamous endothelium (arrow) and underlying connective tissue. It is the only layer
found in all blood vessels. Depending on the vessel, the tunica media (TM) is made of fibrous and/or
elastic connective tissue and smooth muscle. It is absent in capillaries. The tunica externa (TE) is on
the outside of vessels other than capillaries. (X250)
136
A
L
V
V
12-2a
NEUROVASCULAR BUNDLE Typically arteries (A), veins (V), and nerves
(N) are found together in the body. Collectively, they are known as N
a neurovascular bundle. (a) Shown in this micrograph is a fairly large
neurovascular bundle. Notice the round lumen and thicker wall layers
in the artery. Corresponding layers in the walls of the artery and vein
stain the same. (X60) (b) This is a much smaller neurovascular bundle
and includes a small lymph vessel (L). (X120)
12-2b
TI
TM
AORTA In this specimen of the aorta,

the fenestrated elastic membranes are
visible in the tunica media (TM) as
black lines. They account for the
elasticity of this large artery. The
tunica externa (TE) and part of the TE
tunica interna (TI) are also visible.
(X65) 12-3
137
TM
TM
IEM
IEM
12-4a 12-4b
MUSCULAR ARTERY Medium sized arteries are also known as muscular arteries because of the abundance of smooth muscle fibers in the tunica
media (TM). Notice in both specimens that the tunica media is the thickest layer, a feature typical of arteries in general. (a) In this specimen, the
internal elastic membrane (IEM) is obvious as a dark squiggly line. (X130) (b) The internal elastic membrane is still very apparent in this
micrograph even though it did not stain a color that makes it stand out. Also note the vein (V) at the right. (X130)
A TM
V
A V
A
M
V L
12-5a 12-5b
ARTERIOLE AND VENULE (a) Arterioles (A) are less than 200 µm in diameter. In this specimen, the endothelial lining and internal elastic membrane
are visible and the tunica media (TM) is about three muscle cells thick. Notice that the tunica adventitia blends in with the surrounding connective
tissue. Two smaller arterioles are also present, each with a single layer of smooth muscle. To the right and below the largest arteriole are two
smaller venules (V). Note their thin walls. (X265) (b) In this specimen, the nuclei of smooth muscle cells (M) in the arterioles are larger and
lighter than the endothelial cells’ nuclei (E). Notice that the tunica media of the smallest arterioles consists of one layer of smooth muscle. The
wall of the venule is extremely thin, but the lumen is relatively large. There is also a small lymphatic vessel (L) visible in the field. (X660)
138
12-6a 12-6b
CAPILLARIES Capillaries are the smallest of blood vessels and are the
site of material exchange between the blood and tissues. They can be
recognized by the wall consisting only of endothelium and the red
blood cells often lining up in single file within the lumen. There are
three types of capillaries recognized based on structures visible with
the electron microscope. While they can’t be differentiated using light
micrographs, following are examples of each type. In each case, an
arrow indicates the capillaries. (a) This lung specimen provides
examples of continuous capillaries, the most abundant of the three.
(X660) (b) Glomerular capillaries in the kidneys are an example
of the fenestrated type. (X660) (c) This liver specimen provides an
example of sinusoidal capillaries. These have an endothelium, but it is
not complete. The cells around it determine the overall shape of the
capillary. Notice that these are larger than the other capillary types as
evidenced by the number of RBCs that fit across the diameter. (X265)
12-6c
139
SM
TE
TM
TI
A
EF
12-7a 12-7b
VEINS Veins are typically thin-walled and the tunica externa is the thickest layer. (a) This micrograph of a vein clearly shows the layers, which
stain the same as the comparable layers in the artery (A). Notice the nuclei of the endothelial cells (E) in the tunica interna (TI), the smooth
muscle (SM) of the tunica media (TM), and elastic fibers (EF) in the tunica externa (TE). (X125) (b) The layers are not as distinct in this
micrograph of a small vein. Note the arteriole (arrow). (X260)
12-8a 12-8b
VALVES IN VEINS Veins carrying blood against gravity often have pocket-shaped valves (V) made of infoldings of tunica interna. (a) X125 (b) X260.
140
TI
TE
12-9
VENA CAVA Large veins have a thick tunica interna (TI) and tunica externa (TE). The tunica media is
absent or not well-defined. (X125)
12-10
ENDOCARDIUM The inside of the heart is lined with endocardium composed of an endothelium and
loose connective tissue. In this micrograph, the endocardium (E) and myocardium (M) are visible.
(X65)
141
ID
ID
ID
12-11a 12-11b
MYOCARDIUM The thickest layer in the heart’s wall is the myocardium. Shown here are two micrographs of cardiac
muscle prepared with different stains. In (a), the striations and intercalated discs (ID) are easily seen. (X380) In (b),
striations are difficult to see and the intercalated discs are faint, but the branching fibers are very obvious. Notice the
numerous capillaries (continuous type) between the fibers. (X250)
12-12
EPICARDIUM The outer surface of the heart is lined with serous membrane known as epicardium (E) or
visceral pericardium. Notice the adipocytes (A) and myocardium (M). (X130)
142
PF
12-13a
PF
12-13b
PURKINJE FIBERS Purkinje fibers (PF) are part of the heart’s conducting system and are responsible
for carrying the impulse from the apex upward across the ventricles. They are located between the
endocardium (E) and myocardium (M). Purkinje fibers are modified cardiac muscle cells, but are
larger and lighter staining. Both micrographs are X130.
Lymphatic System
C H A P T E R
13
Introduction to the Lymphatic System B cells possess an antigenic marker that identifies them.
The primary role of the lymphatic system is that of immu- They are produced in the bone marrow and also become
nity, of allowing the body to differentiate between “self” immunocompetent there. When stimulated, the appropriate
and “nonself.” Nonself chemicals or cells are broadly re- B cell multiplies and differentiates into plasma cells and
ferred to as antigens. (Immunologists define antigens more memory B cells. Plasma cells actively secrete antibodies (hu-
precisely than that, but this definition will serve our pur- moral immunity), whereas memory cells remain dormant
poses.) The body’s response to an antigen is either humoral, until subsequent contact with the same antigen. Plasma
in which antibodies are secreted into tissue fluids, or cellu- cells are elongated and have nuclear chromatin resembling
lar, in which the defense is provided directly by the immune a clock face. A light region near the nucleus is the site of the
cells. In either case, the response is specific for each antigen. Golgi apparatus. Memory B cells are indistinguishable from
The lymphatic system is also involved in recovery and other lymphocytes in routine histological preparations.
transport of tissue fluid, as well as absorption of fats in the T cells are produced in the bone marrow but become
small intestine. immunocompetent in the thymus. They are identified by a
T cell antigenic marker and are involved in cell mediated re-
sponses. When stimulated, T cells proliferate and differen-
Cells of the Immune System tiate into one of several functional groups. T helper (TH)
There are several types of immune cells, but they are mostly cells assist other T cells or B cells in their immune response.
difficult to differentiate with the light microscope. The fol- Cytotoxic T (Tc) cells are active in killing foreign cells or vi-
lowing relies on identification by sophisticated techniques rally infected cells. Memory T (TM) cells perform the same
that are beyond the scope of an introductory histology function as their B cell counterparts. These T cells are indis-
course, as is an exhaustive description of their functions. For tinguishable from other lymphocytes in routine histological
more information, the reader is referred to an immunology preparations.
text. Natural killer (NK) cells are large lymphocytes that
Lymphocytes (Figure 13-1) constitute approximately lack the T and B cell antigenic markers, and as such belong
20–25% of all leukocytes seen in the blood. They are typi- to the population of null cells. NK cells kill cells coated
cally about the size of erythrocytes, but larger lymphocytes with antibodies in a process called antibody-dependent cell-
are also seen. Lymphocytes are categorized into one of three mediated cytotoxicity.
functional groups: B cells, T cells, and NK cells. When func- Another category of cells involved in the immune re-
tional, they are not found in the blood, but rather in lym- sponse are antigen presenting cells (APCs), which are in-
phoid organs or as aggregations in extravascular tissues. volved in processing an antigen and “showing” it to the
143
144
appropriate immune cell to begin the immune response. macrophages, populate them. Antigens in tissue fluid enter
Macrophages (Figure 4-9) and dendritic (Langerhans) cells the lymph and are removed by these cells before they can
(Figure 11-3c) of the skin are examples. get into the blood.
Lymph nodes are bean-shaped and are covered with a
Organs of the Lymphatic System dense connective tissue capsule. Afferent lymph vessels carry
Lymphatic tissue is found in encapsulated lymphoid organs, lymph to the node and enter along the node’s convex sur-
such as lymph nodes, the thymus, and the spleen, and also face. The indentation is the hilus and is the point of entry
as unencapsulated clusters in the walls of other organs such and exit for blood vessels, as well as exit of the efferent
as those of the digestive and respiratory tracts. lymph vessel (Figure 13-7).
Internally, the node is divided by extensions of the cap-
Thymus
sule called trabeculae (Figure 13-8). Connecting the afferent
The thymus (Figure 13-2) is found in the mediastinum, the vessels with the efferent vessel is a series of channels called
region of the thoracic cavity between the lungs. It is most sinuses, each named according to its location. The subcap-
highly developed at puberty, then is replaced with adipose tis- sular sinus is on the periphery of the node, and the cortical
sue as the individual ages. A dense connective tissue capsule and medullary sinuses are in the node’s cortex and medulla,
covers the thymus, and connective tissue trabeculae or septa respectively. As lymph passes through the sinuses, it con-
arising from it divide the organ’s two lobes into lobules. The tacts the immune cells inhabiting the node.
outer portion is the cortex and the inner part is the medulla. Most lymphocytes enter the node through the artery,
The thymic cortex (Figure 13-3) stains darker and more with the remainder entering through the afferent vessels. A
basophilic than the medulla, and is occupied by numerous framework of reticular fibers coursing through the node
T lymphocytes (thymocytes) that have migrated from the supports them. In the cortex, lymphocytes are found in
bone marrow. They proliferate and as they mature, they dense, spherical clusters called lymph follicles or lymph
move toward the medulla. They become immunocompetent nodules (Figure 13-9). Primary follicles are uniform in ap-
at the periphery of the cortex. Epithelial reticular cells sepa- pearance and are mostly composed of small, inactive B
rate the cortex from the blood vessels in the trabeculae and cells. Secondary follicles have a light staining germinal center
form the blood-thymus barrier that prevents antigens from composed of proliferating B cells surrounded by a darker
contacting thymocytes. mantle zone populated by inactive B cells. A lighter mar-
The thymic medulla (Figure 13-4) is eosinophilic and ginal zone may be seen around the mantle zone. T lympho-
has fewer lymphocytes and more epithelial reticular cells cytes dominate the deepest portion of the cortex called the
than the cortex. From the medulla, mature T cells enter paracortex. Follicles are not found in the paracortex. Lym-
blood or lymphatic vessels and spread throughout the body phocytes of the medulla are organized into elongated cords
to populate other lymphatic organs. Thymic (Hassall’s) cor- (Figure 13-10). The cells are primarily plasma cells and B
puscles are the most distinctive features of the medulla. lymphocytes.
They are concentrically arranged keratinized epithelial The mantle zone of a secondary follicle is often wider
reticular cells. Their function is unknown, but they proba- on the side near the capsule. Likewise, the germinal center
bly are produced by a degenerative process. shows polarity, with the medullary side being darker (the
Lymph Nodes and Lymph Vessels dark zone) than the side facing the capsule (the light zone).
Lymphatic vessels (Figure 13-5) return tissue fluid to the It is in the germinal center that B cells divide and produce
blood vascular system. The lymph vasculature begins as plasma cell precursors and memory B cells. The memory B
blind capillaries consisting of a simple squamous endothe- cells enter circulation and take up residence in other lym-
lium. The capillaries are tributaries to small lymph vessels, phatic tissues whereas the plasma cell precursors move to
which continue to converge and get larger. Ultimately, the medullary cords and finish their differentiation.
lymph vessels empty into the right lymphatic duct and the
thoracic duct, which drain lymph into the right and left Spleen
subclavian veins, respectively. On slides, RBCs are not seen The spleen is a lymphoid organ that occupies the left upper
in lymph vessels, but lymphocytes may be present. quadrant of the abdominal cavity. It is responsible for im-
In addition to their pattern of convergence, lymph ves- mune responses to blood antigens as well as phagocytosis
sels also resemble veins in being thin-walled (but thinner) of worn out RBCs and other particulate matter.
and having valves. The tunics are indistinct in most lymph The capsule (Figure 13-11) is composed of fibrous con-
vessels, but the tunica media of the lymphatic ducts has lon- nective tissue and smooth muscle. Connective tissue tra-
gitudinal and circular layers of smooth muscle in it. beculae penetrate the spleen from the capsule and reticular
Lymph nodes (Figure 13-6) are located periodically fibers form a framework in which the lymphocytes are
along the length of lymph vessels. B and T cells, as well as suspended.
C H A P T E R T H I R T E E N : L Y M P H A T I C S Y S T E M
145
Spleen tissue is divided into red pulp and white pulp. of lymph follicles, mostly with germinal centers. The strati-
Red pulp (Figure 13-12) is the blood vascular component fied squamous epithelium of the oral mucosa covers them
and includes blood sinuses and the intervening splenic cords and penetrates downward as tonsilar crypts. Dendritic cells
composed of macrophages, lymphocytes, and blood cells. in the epithelium act as APCs. Connective tissue trabeculae
The sinuses are made of endothelium and are enveloped in are present and there is a deep connective tissue capsule,
reticular fibers. White pulp (Figure 13-13) is composed of but the tonsils are not surrounded by it. The pharyngeal
lymphocyte aggregates. T cells form periarterial lymphatic tonsil (Figure 13-16) is located in the nasopharynx. It is
sheaths (PALS) around the central arteries. The other type covered with PSCC and some stratified squamous epithe-
of white pulp is made primarily of B cells in lymph follicles lium. There are no crypts, but the mucosa is folded into
with germinal centers and mantle zones. A lighter marginal pleats. The lingual tonsils (Figure 13-17) are at the base of
zone of active macrophages is found at the periphery of the the dorsum of the tongue. They are covered with stratified
follicle. squamous epithelium. Each has a single crypt.
Gut Associated Lymphatic Tissue (GALT) is composed
Other Unencapsulated Lymphatic Tissue
of lymph follicles in the walls of digestive organs (Figure
Lymphatic tissue is found scattered in various organs. In 13-18). They are most abundant in the Peyer’s Patches of
some cases, the lymphocytes are just dispersed in other con- the ileum. GALT is drained by efferent lymph vessels, but
nective tissues. In other cases they are organized into lymph has no afferent vessels. Instead, special epithelial cells called
follicles. M cells overlie the follicles and transfer antigens to macro-
Scattered lymphocytes in the lamina propria and epi- phages in the follicles which in turn present them to T cells.
thelium of various organs constitutes the Mucosa Associated GALT is also abundant in the colon and the vermiform ap-
Lymphatic Tissue (MALT). T cells are the primary occupants pendix, a tubular extension of the cecum.
of MALT, but some B cells are also present (Figure 13-14). Bronchus Associated Lymphatic Tissue (BALT) is com-
Tonsils are probably the most well known examples of mon at the branch points of the respiratory tree, where M
MALT. They are small aggregations of unencapsulated lym- cells replace the respiratory epithelium (Figure 13-19). B
phatic tissue located in the oropharynx and nasopharynx. cells are the most abundant cells, but APCs and T cells are
The two palatine tonsils (Figure 13-15) are located at the also present.
junction of the oral cavity and the oropharynx. They consist
13-1a 13-1b
LYMPHOCYTES (a) Lymphocytes are the second most abundant leukocyte in blood. (X660) (b) Lymphocytes are found in blood, but they are
usually functional outside the blood stream. Shown here is a spherical aggregation of lymphocytes in the wall of the digestive tract. Each tiny
purple dot is a lymphocyte. (X260)
146
THYMUS The thymus is covered with a
connective tissue capsule (C). Trabeculae (T)
are projections of the capsule into the thymus
T that divide it into lobules. The cortex (Cx)
and medulla (M) are clearly seen. (X50)
Cx
13-2
C
T
Cx
13-3a
E
THYMIC CORTEX The outer region of the thymus
is called the cortex. (a) In this micrograph, the
capsule (C) and trabeculae (T) are visible. The
more basophilic cortex (Cx) stands out against
the more eosinophilic medulla (M). (X60)
(b) Most of the cortical cells are thymocytes
(T lymphocytes). Other cells of the cortex are
macrophages and epithelial cells (E—these are
difficult to see in normal preparations because
of the numerous thymocytes). T cells that react
with “self” die and are removed by the
macrophages. (X660)
13-3b
147
13-4a 13-4b
THYMIC MEDULLA Epithelial cells (E) involved in the positive and negative selection process of T cells are more visible in the medulla. They are
rounded and light staining. Keratinized clumps of dying epithelial cells form Hassall’s corpuscles (H), a distinctive feature of the thymus. (a)
Epithelial cells and an early Hassall’s corpuscle are seen in this micrograph. (X660) (b) A more advanced Hassall’s corpuscle is shown in this
micrograph. Epithelial cells are also visible. (X660)
V
V
V
V V
13-5a 13-5b
LYMPH VESSELS Like veins, lymph vessels have valves, but they have thinner walls. (a) In this whole mount, the valve (V) and thin wall are
apparent. (X60) (b) This is a section through a couple of lymph vessels with valves. (X65)
148
P C
F
Cx
Co Cx
H M
Co
P
F
H
13-6a 13-6b
LYMPH NODE Each lymph node is covered by a connective tissue capsule (C). Lymphatic tissue comprising the cortex (Cx) is arranged in spherical
follicles (F) or in irregular masses in the paracortex (P), whereas it is organized into cords (Co) in the medulla (M). Sinuses run between masses
of lymphatic tissue. Blood vessels and efferent lymph vessels connect at the hilus (H). (a and b) Both micrographs are panoramic views of lymph
nodes. (X22)
L
V
13-7
EFFERENT LYMPH VESSELS In this micrograph, efferent lymph vessels emerging from the hilus are
visible. Note the valve (V), the lymphocytes (L), and the absence of RBCs in the vessel. (X130)
149
LF
SS
CS
T SS
13-8a C
LF CS LF
13-8b
CS
LYMPH NODE TRABECULAE AND SINUSES (a,

SS
b, and c) The interior of the lymph node is
penetrated by connective tissue trabeculae
(T) arising from the capsule (C). The sub-
capsular sinus (SS) and cortical sinuses (CS)
LF are visible in places, but are obscured in
others by lymphocytes from the lymph
13-8c follicle (LF). (a is X115, b and c are X130.)
150
MnZ
LZ
DZ
GC
LF
13-9a 13-9b
MrZ CC
MnZ
MF
GC
CC
13-9c 13-9d
LYMPH NODE CORTEX (a) The cortex of a lymph node is composed of spherical aggregations of B lymphocytes called lymph follicles (LF). Deep
to the follicles is the paracortex (P), occupied mainly by T cells. The medulla (M) is also visible. (X65) (b) Shown in this field are secondary
follicles demonstrating the lighter germinal center (GC) and darker mantle zone (MnZ). Notice that the mantle zone is thicker on the side of the
capsule and that the germinal center has a dark zone (DZ) toward the medulla and gets lighter (LZ) toward the capsule. Secondary follicles have
multiplying B cells in the germinal center. (X130) (c) This is a secondary follicle. In addition to the germinal center and mantle zone, the lighter
marginal zone (MrZ) is visible. (X265) (d) B cells multiply and differentiate in the germinal center, beginning in the darker portion at the
medullary end and finishing at the lighter capsular end. During development, they pass through centroblast and centrocyte stages before they
become plasma cells. Centroblasts are large cells with a round nucleus. These are mitotically active and produce the centrocytes. Centrocytes
(CC) are recognized by their irregular nuclear membranes. Plasma cells and memory B cells are located in the light region before they migrate to
the medulla. Tingible body macrophages (MF) are also seen. These cells phagocytose B cells that have failed to differentiate properly in response
to the antigen. (X660)
151
MC T
C
MS
RP
WP
MC
MS
13-10
LYMPH NODE MEDULLA The lymphocytes of the medulla (mostly
plasma cells) are arranged into cords (MC) and are separated by
medullary sinuses (MS). The reticular connective tissue framework 13-11a
is visible (arrows), as are trabeculae (T).
T C
RP
WP
13-11b
SPLEEN Both micrographs show a panoramic view of the spleen with its
13-12 capsule (C) and trabeculae (T). Red pulp (RP), made of blood sinuses, and
white pulp (WP), made of lymphocytes, are also visible. (a) X20 (b) X25
SPLENIC RED PULP The portion of splenic parenchyma com-
posed of blood sinuses (S), cords (C) of lymphocytes and
macrophages is called red pulp due to its appearance in fresh
specimens. Blood in the spleen eventually finds its way to blind
capillaries surrounded by macrophages. The RBCs must pass
by this macrophage layer in order to enter the red pulp. Then,
they must pass through slits in the basement membrane of the
sinuses, a task likely to break old, brittle RBCs. This contact
with macrophages and difficult passage into the sinuses is the
mechanism for removing RBCs from circulation.
152
GC
PALS
A
MnZ
MrZ
13-13a 13-13b
SPLENIC WHITE PULP White pulp is found in two forms. Masses of T cells form periarterial lymphatic sheaths (PALS) around arterioles (A), as
in (a). B cells form follicles with a germinal center (GC), mantle zone (MnZ) and marginal zone (MrZ). These are illustrated in (b). Both micro-
graphs are X130.
L L
13-14a 13-14b
DIFFUSE LYMPHATIC TISSUE Lymphocytes are often found in the connective tissue layer of a mucous membrane and are referred to as MALT. They
are characterized by small, dark staining nuclei. (a) Numerous lymphocytes (L) are visible in this preparation of small intestine mucosa. (X250)
(b) This is another small intestine specimen. Lymphocytes are present in the connective tissue, but are also seen in the epithelium. (X380)
153
Cr
SS F
F
SS
F IF
IF
Cr
13-15a 13-15b
PALATINE TONSILS The palatine tonsils are covered by stratified squamous epithelium (SS) that projects down into tonsilar crypts (Cr). The bulk
of the tonsil is composed of follicles with germinal centers (F), with interfollicular (IF) regions composed of T cells making up the remainder. A
fibrous capsule (C) marks the lower margin of the tonsil. (a) X8 (b) X25.
Ep
Ep
P
C
F Ca
13-16
PHARYNGEAL TONSIL The pharyngeal tonsil is similar in construction to 13-17
the palatine tonsils, but has a thinner capsule, is covered with PSCC
(Ep), and has shallower depressions called pleats (P). Lymph follicles LINGUAL TONSIL Each lingual tonsil has a single crypt (C), a thin
(F) are abundant. The white region indicated by the arrow is an capsule at its base (Ca), and is covered with stratified squamous
artifact of preparation. (X50) epithelium (Ep). Follicles (F), some with germinal centers, are also
present. (X7)
154
13-18a 13-18b
GUT ASSOCIATED LYMPHATIC TISSUE (GALT) Lymph follicles are present
in the mucosa of many digestive organs. (a) Here is a single follicle in
the small intestine’s mucosa. Diffuse lymphatic tissue is also present.
(X110) (b) Aggregates of lymph follicles in the ileum are known as
Peyer’s patches. They are found in the submucosa. (X25) (c) The
vermiform appendix is found on the cecum of the large intestine. It
has abundant lymphatic tissue in it, as this cross section illustrates.
(X20)
13-18c
BRONCHUS ASSOCIATED LYMPHATIC TISSUE

(BALT) Lymph follicles are also found
associated with the respiratory tree,
especially at branch points. (X110) 13-19
Digestive System
C H A P T E R
14
Introduction to the Digestive System secretion is viscous due to the mucus in it. Occasionally,
The digestive system is involved in the ingestion, mechanical serous and mucous acini are found together, with the serous
and chemical digestion, and absorption of food. It consists cells forming a cap over the mucous cells, a structure re-
of the organs of the oral cavity (teeth and tongue), digestive ferred to as a serous demilune.
tube (esophagus, stomach, small intestine, and large intes- Acini empty into ducts made of secretory cells called
tine) and accessory glands (salivary glands, liver, gall blad- intercalated ducts, which lead to larger striated ducts. These
der, and pancreas). are lined with a simple columnar epithelium whose nuclei
tend to be toward the free surface and whose cytoplasm ap-
Organs of the Oral Cavity pears striated due to numerous folds of the basal mem-
The oral cavity (Figures 14-1 and 14-2) is lined with a strati- brane. Striated ducts secrete lysozyme and antibodies
fied squamous epithelium. It is nonkeratinized on the inside (immunoglobin A).
of the lips, cheeks, and hard and soft palates. It is partially Each acinus is associated with a flattened, contractile
keratinized over the gingivae (gums) and some papillae of myoepithelial cell with processes that wrap around the aci-
the tongue. nus. Upon contraction, myoepithelial cells assist in moving
Chemical digestion of starch begins in the oral cavity, the secretion into the duct.
but the main function is the mechanical digestion of food as There are three main salivary glands. The parotid glands
it is broken up by the teeth and mixed with saliva to form a (Figure 14-4) are located anterior to each ear and their ducts
bolus. empty into the oral cavity near the second maxillary molar.
A connective tissue capsule sends septa into the gland and
Salivary Glands divides it into lobules. Most acini are the serous type. Adi-
Salivary glands (Figure 14-3) produce a secretion called pocytes become more abundant with age.
saliva. Saliva is about 99% water, with mucus and the en- The sublingual glands (Figure 14-5) are located in the
zymes amylase and lysozyme comprising the remaining 1%. floor of the oral cavity. They are composed primarily of
The secretory cells are arranged into acini of two types: mucous acini, some with serous demilunes. Adipocytes may
serous acini and mucous acini. Serous acini produce a watery also be seen, especially in adults as they increase in number
secretion containing enzymes and antibodies. They are typi- with age.
cally basophilic and have a granular cytoplasm (consistent The submandibular glands (Figure 14-6) contain both
with their function of protein synthesis.) Mucous acini gen- serous and mucous acini, some with serous demilunes. The
erally appear pale with a smooth cytoplasm and the cells’ relatively long striated ducts make them abundant in sec-
nuclei pushed toward the basement membrane. Their tioned specimens.
155
156
Teeth buds. In addition, it is responsible for mixing chewed food
There are typically 20 deciduous teeth that are replaced by with saliva to form a bolus. It is also involved in speech.
32 permanent teeth in the adult human. There are 16 per- The surface of the tongue is covered by papillae of
manent teeth in each dental arch and the pattern in the three types. Filiform papillae are the most numerous and
mandibular and maxillary arches is the same, with two in- are hair-like in appearance. Their epithelium is keratinized.
cisors, one canine, two premolars, and three molars on Fungiform papillae are globular with a nonkeratinized
each side (Figure 14-7). stratified squamous epithelium and vascular connective tis-
Each tooth (Figure 14-8) consists of a crown that pro- sue within, which makes them appear reddish. Six to 14
jects above the gingivae or gums. The crown is covered with circumvallate papillae are located in a row in the region
enamel, the hardest material in the body (95% calcium known as the sulcus terminalis, about two-thirds of the
salts and 5% organic substances). Below the gumline, one way back on the tongue. Each is surrounded by a cleft into
or more roots project into alveoli (sockets) in the bone. The which serous von Ebner’s glands drain. Most taste buds are
roots are covered with cementum, a bone-like material that located in the mucosa of these papillae.
lacks the Haversian systems and blood vessels of bone. The The bulk of the tongue is skeletal muscle arranged in
fibrous periodontal ligament (periodontal membrane) con- horizontal, transverse, and longitudinal bands. These are
nects the cementum with the alveolar bone and acts as its responsible for producing the complex movements of the
periosteum. The neck of the tooth is a constriction at the tongue. Filling in between the muscle and epithelial layers is
junction of the crown and roots. Internally, the tooth con- a loose connective tissue lamina propria with occasional
sists of bone-like dentin (70% calcium salts) and a pulp cav- mucous and serous salivary glands.
ity filled with the vascular loose connective tissue called Taste buds (Figures 9-2 and 14-10f) are light staining
dental pulp. The gingiva is a mucous membrane that covers oval objects found in the epithelium of the oral cavity and
the bone tissue of the maxilla and mandible up to the tooth’s the tongue. At the surface is the taste pore. Both the lighter
enamel. It has a keratinized stratified squamous epithelium. staining gustatory cells and darker sustentacular cells are
Deciduous teeth develop from an invagination of thick- associated with neurons, but the gustatory cells are consid-
ened embryonic oral epithelium called the dental lamina in ered to be the actual chemoreceptors.
each jaw (Figure 14-9). The dental lamina dilates and forms
an enamel organ for each tooth, then eventually degener-
ates. Deep to the enamel organ, mesenchyme begins to Organs of the Digestive Tube
form the dental papilla. Concurrently, the enamel organ be- The digestive tube consists of the esophagus, stomach,
comes cap-shaped as it grows around the papilla. Tall small intestine, and large intestine. The wall is divided into
columnar enamel-producing ameloblasts begin to develop four main layers that show modification from one organ to
along the concave side of the enamel organ next to the another depending on function (Figure 14-11). From the
papilla, then dentin producing odontoblasts form between lumen outward, these layers are the mucosa, submucosa,
the ameloblasts and the dental papilla. Further growth of muscularis externa, and serosa or adventitia.
the enamel organ results in a bell-shaped structure that de- The mucosa is composed of either a stratified squamous
termines the form of the particular tooth’s crown. Connec- epithelium or a simple columnar epithelium. Deep to it is a
tive tissue around the enamel organ differentiates into the loose connective tissue layer called the lamina propria. In
dental follicle, the precursor of the periodontal ligament. It some organs, the lamina propria is displaced by glandular
also forms the cementum of the root. epithelium; in others it is occupied by numerous lympho-
Odontoblasts secrete dentin between the odontoblast cytes. Together, the epithelium and lamina propria form the
and ameloblast layers. As they do so, they migrate inward. mucous membrane of the gut. Finally, a thin layer of
A long, thin cytoplasmic process of each odontoblast re- smooth muscle (skeletal muscle in the proximal esophagus)
mains in the dentin as the odontoblastic process, visible in forms the muscularis mucosae.
light micrographs as a dentinal tubule. Subsequently, amelo- Deep to the mucosa is the submucosa, a loose connec-
blasts deposit enamel next to the dentin. Thus, the odonto- tive tissue that houses blood vessels, nerves, and lymphat-
blast and ameloblast layers become separated by the dentin ics, as well as glands derived from the surface epithelium in
and enamel each has secreted. some organs.
The muscularis externa is composed of smooth muscle
Tongue in most organs, but skeletal muscle is seen in the proximal
The tongue (Figure 14-10) is a muscular organ covered esophagus and anal canal. It is composed of an inner circu-
with mucous membrane consisting of a stratified squamous lar layer and an outer longitudinal layer, the coordinated
epithelium and lamina propria. It possesses general sensory contraction of which results in the propulsive movement
receptors as well as chemoreceptors for taste called taste called peristalsis.
C H A P T E R F O U R T E E N : D I G E S T I V E S Y S T E M
157
If the organ is in the peritoneal cavity, the outer surface chemical digestion of proteins and further mechanical di-
is lined with a serous membrane called visceral peritoneum, gestion of the food to make chyme.
and this constitutes the serosa. It is composed of a simple The stomach is divided into four regions. The esophagus
squamous mesothelium and a deeper fibrous connective tis- enters at the cardia. Superior to this is the fundus. The main
sue. In most parts of the gut, the serosal layer joins itself on portion is called the body or corpus, and the end joining the
one side of the organ and forms the double-layered mesen- small intestine is the pyloris.
tery that suspends the organ and carries blood vessels to The mucosa of the esophagus changes abruptly as it
and from it. If the organ is outside the peritoneal cavity, it is joins the cardia (Figure 14-14). At the esophageal-cardiac
covered with a fibrous adventitia that blends in with sur- junction, the stratified squamous becomes a simple columnar
rounding connective tissues. epithelium, and the lamina propria becomes occupied by
In addition to these basic layers, neural structures re- gastric glands (see below). All other layers are continuous
sponsible for regulating glandular secretion and muscular across the junction.
contraction are present. Meissner’s plexus is found in the The mucosa of the stomach (Figure 14-15) has depres-
submucosa and supplies the muscularis mucosae and mu- sions called gastric pits. Leading from each pit is an isthmus
cosal glands. The myenteric plexus (of Auerbach) is located which gives rise to tubular gastric glands. With the light
between the two muscular layers of the muscularis externa. microscope, three cell types are identifiable. These are the
These are aggregations of parasympathetic postganglionic chief cells, mucous cells, and parietal cells.
neurons that supply the muscle. Mucous cells are found on the luminal surface and line
the pits. They are also in the necks of the glands. These mu-
Esophagus cous cells can be differentiated from the surface mucous
The esophagus (Figure 14-12) is a muscular tube found in cells using the electron microscope. The mucus produced
the mediastinum of the thoracic cavity leading from the coats the stomach lining and protects it from the low pH
laryngopharynx to the stomach. Peristaltic waves convey and enzymes of gastric juice.
the bolus of food to the stomach. A physiological sphincter Parietal cells are eosinophilic and appear reddish with
normally prevents food from being regurgitated from the H&E stains. They secrete hydrochloric acid and intrinsic
stomach into the esophagus. factor (necessary for absorption of Vitamin B12). They are
The relaxed esophageal mucosa is highly folded, but it most abundant in the isthmus region, but may be found
flattens as the esophagus distends to accommodate the throughout the gland.
swallowed bolus. The epithelium is a nonkeratinized strati- Chief (zymogenic) cells are distributed throughout the
fied squamous. There is a thin lamina propria and discontin- gland, but are most abundant near the base. These cells are
uous muscularis mucosae. basophilic and their granular cytoplasm appears purplish with
The submucosa is composed of an elastic connective H&E stains. They secrete the precursor to the enzyme pepsin.
tissue (to accommodate swallowing) and contains serous There is regional variation in the gastric mucosa (Figure
esophageal glands. 14-16). Deep pits and an abundance of mucous cells in the
Since swallowing is a voluntary action, the proximal glands characterize the cardiac region. The fundus and
one-third of the esophagus contains skeletal muscle in the body, which comprise the majority of the stomach, have
muscularis externa and muscularis mucosae. By the distal shallow pits and typical gastric glands composed of mu-
one-third of the esophagus, the muscle layers are made ex- cous, parietal, and chief cells. The pyloric region has deep
clusively of smooth muscle. The muscularis externa is thick pits penetrating to about half the thickness of the mucosa.
and has well-defined layers. Mucus cells are abundant in the glands.
All but the distal centimeter of the esophagus is located The stomach’s submucosa is not remarkable, but the
in the mediastinum and as such is covered by a fibrous ad- muscularis externa has an additional innermost oblique
ventitia. The terminal part of the esophagus, which has layer. At the pyloric-duodenal junction, the circular layer
passed through the diaphragm to join the stomach, is lined thickens to form the pyloric sphincter. The outer surface is
with a serosa. covered with a serosa.
Stomach Small Intestine

The stomach is a dilation of the digestive tube that becomes The small intestine is approximately 3 meters in length. It
further distended as food is stored in it. Internally, the mu- begins at the duodenum (25 cm long) and continues as the
cosa of an empty stomach is thrown into nonpermanent jejunum (1 m long), and ends with the ileum (2 m long).
longitudinal folds called rugae (Figure 14-13). As the stom- Each segment has characteristic features (see next page). It
ach fills, the rugae flatten to accommodate the increased is anchored to the dorsal body wall by a fan-shaped mesen-
volume. In addition to storage, the stomach is involved in tery that supplies blood vessels, nerves, and lymphatics.
158
Chemical digestion of chyme continues in the small in- Large Intestine
testine as it is mixed with enzymes in pancreatic juice and The large intestine is divided into a cecum, ascending colon,
secretions of the intestinal mucosa. It is also the primary transverse colon, descending colon, and rectum. Its primary
site of nutrient absorption into the blood. Consistent with function is to absorb water from the undigested, unabsorbed
its absorptive function, the mucosa is modified to increase material entering from the small intestine. The large intes-
surface area (to an amazing 600m2!). On a gross level, plicae tine begins at the ileocecal junction and the ileocecal valve
circulares (Figure 14-17) are permanent mucosal folds that prevents backflow of material into the small intestine.
are visible to the naked eye. Extending from the mucosal Nonpermanent folds of the colon’s mucosa may be pres-
surface are the finger-like villi that are just barely visible to ent, but there are no villi (Figure 14-22). The epithelium is a
the naked eye. These give the internal surface an appear- simple columnar with abundant goblet cells. Straight crypts
ance of velvet. Lastly, each epithelial cell has microscopic of Lieberkühn extend down to the muscularis mucosae.
microvilli extending from its surface, visible with the light These are involved in absorption of water and production
microscope as a brush border. of new cells by mitosis. Lymphatic follicles and diffuse lym-
The intestinal mucosa (Figure 14-18) is lined by a simple phatic tissue may be seen in the lamina propria.
columnar epithelium with a brush border and mucous se- The muscularis mucosae is usually prominent. The
creting goblet cells. Villi project out of the mucosa and in- inner circular muscle layer is unremarkable, but the outer
crease its surface area. Between villi are intervillous spaces. longitudinal layer is modified to form three bands called
Intestinal crypts (of Lieberkühn) are depressions into the taenia coli muscles (Figure 14-23). The surface is covered
mucosa between the villi. The space within a crypt is very with serosa, except for parts of the ascending and descend-
small compared to the intervillous space. Further, the crypts ing colon.
often appear as ovals lined with epithelium that are discon- The rectum is the straight, terminal portion of the large
nected with the surface due to the plane of section. Crypts intestine. Its mucosa resembles the rest of the large intestine,
produce an alkaline fluid and act as a source of new epithe- but the crypts are shallower and the goblet cells are more
lial cells to replace old, worn out ones. Paneth cells, which abundant. At the rectoanal junction (Figure 14-24), the
produce antimicrobial substances, may be seen deep in the simple columnar epithelium abruptly changes back to a
crypts. They possess eosinophilic granules near their apices. stratified squamous. Thickening of the circular muscularis
The lamina propria within each villus has a capillary externa forms the internal anal sphincter. A ring of skeletal
bed, a lymphatic lacteal (capillary), and an abundance of muscle outside the muscularis externa forms the external
scattered lymphocytes. anal sphincter.
Brünner’s glands in the submucosa characterize the The appendix (Figure 13-18c) is a short tubular struc-
duodenum (Figure 14-19). They empty their alkaline, mu- ture arising from the cecum. Its mucosa resembles the rest
coid secretion into the duodenal lumen via the crypts where of the large intestine, but there is an abundance of lympho-
it counteracts the acidity of the chyme entering from the cytes in the lamina propria and the submucosa.
stomach. The cells have a typical mucous-secreting cell
look. That is, they are pale staining with a flattened nucleus Glands of the Digestive Tract
near the base. The liver, pancreas, and gall bladder are derived as out-
Other than villus shape (that is difficult to determine in growths of the embryonic digestive tube and they retain
most preparations), the jejunum has no microscopic fea- their connection with the gut’s lumen in the postnatal
tures that differentiate it from the duodenum and ileum individual.
(Figure 14-20). In most instances, it must be identified by
the absence of the features seen in the other parts. Liver
The ileum (Figure 14-21) is characterized by Peyer’s The liver is located inferior to the diaphragm in the upper
patches, aggregations of several dozen lymph follicles in the right abdominal quadrant and is the largest internal organ of
submucosa on the side of the intestine opposite the mesen- the body. It is divided into four lobes, which are further di-
tery. Whereas isolated lymph follicles are visible in all parts vided into microscopic lobules (Figures 14-25 and 14-26).
of the small intestine, only the ileum has them in this great Traditionally, lobules have been considered the structural and
a quantity. functional units of the liver, but other interpretations also
The small intestine’s muscularis externa is smooth have merit. This being a histology book and not a physiology
muscle and the circular and longitudinal layers are well- book, the traditional view will be presented.
defined. A serosa lines the outer surface, except for the pos- The hepatic artery and the hepatic portal vein provide
terior of the duodenum, which is retroperitoneal. two sources of blood to the liver. The former supplies
159
oxygen-rich blood, whereas the latter brings nutrient-rich with here. Pancreatic juice is alkaline, contains over 20 en-
blood from the digestive organs to be processed by the zymes, and is delivered to the duodenum by the pancreatic
hepatocytes, the cells of the liver. duct.
The liver performs a variety of functions, including de- The pancreas (Figures 14-30 and 14-31) has a thin fi-
toxification of chemicals, phagocytosis of worn out RBCs, brous capsule that sends septa into the gland to divide it
synthesis of plasma proteins and plasma lipoproteins, glu- into lobules. The exocrine component is composed of nu-
cose metabolism (starch storage and gluconeogenesis), stor- merous secretory acini. The cells are roughly triangular and
age of certain vitamins, and production of bile. have a granular cytoplasm typical of zymogenic (enzyme se-
All liver functions are performed by the hepatocytes, creting) cells. That is, they have a basophilic cytoplasm due
and most of them require close contact with the blood. As to the extensive rough endoplasmic reticulum. They also
such, the hepatocytes are arranged in plates or cords of cells have eosinophilic zymogen granules at their apex. There are
separated by large capillaries called sinusoids. The cords are no myoepithelial cells associated with the acini.
usually a single cell thick, and the sinusoids are lined with Each acinus empties its pancreatic juice into intercalated
endothelial cells and phagocytic Kupffer cells. Blood from ducts made of centroacinar cells, which are unique in begin-
branches of the hepatic artery and hepatic portal vein enters ning within the acinus. Intercalated ducts empty into intra-
the sinusoids at several points along the periphery of the lobar ducts and then into interlobar ducts in the gland’s
lobule. As blood flows by the hepatocytes, they remove and septa. Ultimately, pancreatic juice is delivered to the duode-
deposit whatever nutrients are appropriate. The blood is num by the main and accessory pancreatic ducts. The main
drained from the lobule by a central vein, which ultimately pancreatic duct joins the common bile duct in the wall of
leads to the hepatic vein. the duodenum as the ampulla of Vater (Figure 14-32).
Small intercellular channels called bile canaliculi (Figure
14-27) carry bile produced by the hepatocytes. The canali-
culi only exist as separations between cells; they are not HF
tubes apart from the cells. Bile flows through the canaliculi SG
toward the periphery of the lobule and empties into a bile

duct. Bile ducts converge and ultimately lead to hepatic
ducts that emerge from the liver.
The system of bile ducts and the branches of the he-
patic artery and hepatic portal vein travel together within
the liver and constitute a hepatic (portal) triad (Figure 14- SM SS
28). Note that flow of materials within the triad’s compo-
nents is in opposite directions; blood is coming in and bile
is flowing out.
Gall Bladder
The gall bladder (Figure 14-29) is a small, blind sac found
inferior to the liver. It receives bile from the common HF
hepatic duct via the cystic duct. Once in the gall bladder,
bile is concentrated and stored until it is needed. Contrac-
tion of the gall bladder ejects bile back out the cystic duct,
into the common bile duct and into the duodenum. Bile is
alkaline and assists in neutralizing the acidic chyme entering
the duodenum from the stomach. It also is important in
lipid digestion as it emulsify fats.
The mucosa is folded into rugae. A simple columnar epi-
thelium that is active in absorbing water and electrolytes to
concentrate the bile lines it. There is no muscularis mucosae
or submucosa. The muscularis is made of indistinct layers. 14-1
Pancreas LIP Shown is the posterior of the lip. The oral mucosa is lined
with nonkeratinized stratified squamous (SS). Also present are
The pancreas has both endocrine (Chapter 10) and ex- accessory salivary glands (SG) and skeletal muscle (SM). Barely
ocrine components. It is the latter that we are concerned visible at the left are portions of two hair follicles (HF). (X25)
160
HP
ME
SD
SS
SG S
14-2
SOFT PALATE The soft palate is lined with a nonkeratinized stratified
squamous epithelium (SS). Also visible are accessory salivary glands 14-3a
(SG) and the maxillary bone of the hard palate (HP). (X25)
StD
S
StD
StD A
14-3b
BASIC COMPONENTS OF A SALIVARY GLAND Salivary glands have secre-
S tory mucous acini (M) and serous acini (S). The cells of mucous acini
tend to be light staining with the nuclei pushed toward the basement
membrane. They secrete mucus. Cells of serous acini tend to be darker
staining (basophilic) and granular, an appearance typical of enzyme
secreting cells. On occasion, serous demilunes (SD) form over a
mucous acinus. Myoepithelial cells (ME) wrap around the acini and
assist in moving the secretion out the ducts. Striated ducts (StD) are
14-4 lined with a simple columnar epithelium. The nuclei are toward the
PAROTID GLAND The parotid gland is made predominantly of dark apical edge and the basal cytoplasm is vertically striated. (a) This
staining serous acini (S). A few adipocytes (A) and a couple of striated specimen is from the sublingual gland. (X265) (b) This specimen is
ducts (StD) are also visible. (X265) from the submandibular gland. (X400)
161
ME
M StD S
ME
A
SD
BV
14-5a 14-6a
M
S
StD
D
M
Sp
SD Sp
14-6b
14-5b
SUBLINGUAL GLAND The sublingual gland consists mostly of mucous
acini (M), some with serous demilunes (SD). (a) In addition to the
S
acini, myoepithelial cells (ME) and adipocytes (A) are visible in this
specimen, as is a blood vessel (BV). (X230) (b) A duct (D) is seen in a
connective tissue septum (Sp) in this specimen. (X265)
M D
SUBMANDIBULAR GLAND The submandibular

gland is a tubuloacinar gland, with the serous
components (S) outnumbering the mucous
components (M). The mucous cells are often
in a tubular arrangement. Striated ducts (StD),
myoepithelial cells (ME) and connective tissue StD
septa (Sp) are also present. (a) In this specimen,
the tubular arrangement of mucous cells is
clearly seen (arrow). (X265) (b) The lobed
nature of the submandibular gland is visible Sp
in this preparation. (X210) (c) A major duct
is visible in this specimen (D). It is lined with
a stratified columnar epithelium. (X230) 14-6c
162
M2
M1
P2
P1
LI C
MI
14-7
THE ADULT DENTITION Shown is a radiograph of maxillary (upper) and mandibular (lower) dental arches. Starting from the
center of each arch are the medial (MI) and lateral incisors (LI), the canine (C), the first (P1) and second premolars (P2), and the
first (M1), second (M2), and third molars. Note: the third molars of each dental arch are the wisdom teeth and they have been
removed from this patient. (Radiograph courtesy of Paul C. Howard, D.M.D.)
P
GP
D C PM B
GP
G
14-8a 14-8b
TOOTH (a) In this section of a tooth, dentin (D), enamel (E), pulp (P), gingivae (G), gingival pocket (GP), and alveolar bone (B) are visible.
(X25) (b) This is a high power magnification of the tooth root. The periodontal membrane (PM) made of collagen fibers (Sharpey’s fibers)
binds the tooth to the bony alveolus (B). Cementum (C) and dentin (D) are also visible. (X380)
163
OE
EO
B
DP
14-9a 14-9b
EO
D
*
O O
DT
14-9c 14-9d
EO
Os
*
A
DT
D *
O
14-9e 14-9f
TOOTH DEVELOPMENT (a) The dental lamina has dilated to form a bud, the first evidence of an enamel organ. (X130) (b) The cap stage is char-
acterized by the formation of a dental papilla, which becomes surrounded by the developing enamel organ. (X110) (c) In this much later stage,
the enamel organ is fully formed. (X65) (d) This micrograph shows dentin formation. (X250) (e) This micrograph shows the ameloblast layer
enamel. It also shows dentin formation. (X365) (f) This is a cross section of a developing tooth. (X60) Key to symbols used: DL 4 dental lamina,
B 4 bud, OE 4 oral epithelium, DP 4 dental papilla, D 4 dentin, E 4 enamel, EO 4 enamel organ, O 4 odontoblasts, DT 4 dentinal
tubules, A 4 ameloblasts, Os 4 bone, * 4 artifactual space.
164
P
P
SG
SM
14-10a 14-10b
CvP
TB
SG
VE
VE
14-10c 14-10d
TP
T C
V GC
14-10e 14-10f
TONGUE (a) In this panoramic view of the tongue, lingual papillae (P) are seen as projections from the surface. Skeletal muscle (SM), adipose
tissue (A), and accessory salivary glands (SG) are also visible. (X20) (b) Filiform papillae are the most numerous of papillae and are lined with a
keratinized stratified squamous epithelium. (X50) (c) These fungiform papillae are lined with a nonkeratinized stratified squamous epithelium.
(X65) (d) Circumvallate papillae (CvP) have the greatest density of taste buds (TB) in their nonkeratinized stratified squamous epithelium. Serous
von Ebner’s glands (VE) are seen in the lamina propria and empty into the cleft (C) around the papilla. (X25) (e) Skeletal muscle in the tongue is
arranged vertically (V), longitudinally (L), and transversely (T), all of which are seen in this specimen. (X60) (f) These taste buds (TB) are in
fungiform papillae. A taste pore (TP) and gustatory cells (GC) are visible. (X250)
165
Mesentery
Vein
Artery
Nerve
Epithelium
Lamina propria Mucosa
Muscularis mucosae
Submucosa
Inner circular layer
Muscularis
Outer longitudinal layer externa
Connective tissue
Serosa
Mesothelium
Lumen Submucosal
gland
14-11
BASIC PLAN OF THE DIGESTIVE TUBE All parts of the digestive tube are built on this basic plan, with
modifications in each organ appropriate to their different functions.
166
EG
M
A ME SM M
L
LP
SM
MM
ME
14-12a
A
14-12b
M
LP
MM
M
SM
LP
ME MM
14-12c
EG
ESOPHAGUS (a) In this panoramic view, one can observe the collapsed
lumen (L) of the esophagus when empty. (X15) (b) This specimen is
from the upper third of the esophagus as evidenced by skeletal muscle
in the muscularis externa. Also note the esophageal glands (EG) in the
submucosa and the incomplete muscularis mucosae. (X60) (c) Note the SM
thick nonkeratinized stratified squamous epithelium and the thick, but
still incomplete muscularis mucosae. The skeletal muscle in the muscu-
laris externa tells us that the specimen is from the upper third of the 14-12d
esophagus. (X65) (d) Esophageal glands (EG) secrete lubricating mucus
for the inner lining of the esophagus. Note the duct of one gland pass-
ing to the surface. (X55) Key to symbols used: M 4 mucosa, SM 4
submucosa, ME 4 muscularis externa, A 4 adventitia, LP 4 lamina
propria, MM 4 muscularis mucosae.
167
ME
MM
14-14
ESOPHAGEAL-CARDIAC JUNCTION The stratified squamous epithelium
14-13 stops abruptly (arrow) and becomes a simple columnar epithelium
GASTRIC RUGAE The stomach’s mucosa presents nonpermanent where the esophagus (E) joins the cardiac stomach (S). All other layers
longitudinal folds called rugae. These flatten as the stomach fills with are continuous between the two organs. (X50)
food. One ruga is shown in this micrograph. Notice that the
muscularis mucosa (MM) is folded, indicating the whole mucosa
is folded, but the muscularis externa (ME) is not. (X20)
GP
GP
LP GG I
GG
14-15b
MM
14-15a
GASTRIC MUCOSA The stomach’s mucosa is lined with a simple CT
columnar epithelium. Depressions called gastric pits (GP) lead to PC
gastric glands (GG) which extend to the muscularis mucosae (MM)
and occupy most of the lamina propria (LP). (a) This micrograph
shows the entire thickness of the gastric mucosa (X50). (b) Shown
here are gastric pits and the beginnings of gastric glands. Note the BV
stomach’s surface is lined with mucus cells, which are also found
down in the pits, and the isthmus (I) and neck of the glands. (X265)
(c) This micrograph shows that gastric glands extend down to the
muscularis mucosae. Enzyme secreting chief cells (CC) and HCl MM CC
secreting parietal cells (PC) are indicated. Note the blood vessel (BV)
and how little connective tissue (CT) there is between glands.
(X265) 14-15c
168
GP
LN
14-16a
STOMACH REGIONS The mucosa of each part of the GP
stomach has a characteristic appearance. (a) In the
cardiac stomach the gastric pits (GP) are deep and
the cardiac glands secrete mostly mucus. Note the
lymphatic nodule (LN). (X65) (b) The body and
fundus of the stomach have shallow pits and typical
gastric glands with chief (CC) and parietal cells (PC).
(X115) (c) Deep pits extending down about half the
PC
mucosa’s thickness characterize the pyloric region.
The gastric glands (GG) secrete mucus. (X130)
CC
14-16b
GP
GG
14-16c
169
INCREASING SURFACE AREA IN THE SMALL
INTESTINE The mucosa of the small intestine
has several modifications to increase surface
area for absorption. (a) Plicae circulares (PC)
are permanent mucosal folds. As with rugae,
note that the muscularis mucosae (MM) is
folded, but the muscularis externa (ME) is
not. (X25) (b) Villi (V) extend from the muco-
sal surface. In this micrograph, the villi are
projecting from the surface of a plica. Note
that the muscularis mucosae does not follow
the contours of the villi. (X25) (c) Shown here PC
are the adjacent sides of two villi. The simple
columnar epithelium covering villi is modified
with microvilli, which in light micrographs is
seen as a brush border (BB). Also note the
goblet cells (GC) and lymphocytes (L). (X660)
MM
ME
14-17a
BB
L
L
V GC
V
MM
ME
14-17b 14-17c
170
INTESTINAL MUCOSA The intestinal
mucosa has villi (V) projecting from it
and crypts of Lieberkühn (CL) projecting
into it. The intervillous spaces (IV) are
GC much larger than the space within the
IV crypts, providing a clue as to where the
SC intestinal surface actually is. Simple
V columnar cells (SC), goblet cells (GC),
and Paneth cells (P) are present in the
epithelium. (a) The level of the intestinal
surface is indicated with a line at the
right of the field. Villi project up from
the surface and crypts project below the
surface. Note the dark nuclei of lympho-
LP cytes in the lamina propria (LP), the
muscularis mucosae (MM) and the
CL submucosa (SM). (X130) (b) This speci-
men has had its blood vessels injected
with a red dye. Note the capillaries in the
lamina propria of the villi. (X130) (c) This
micrograph shows smooth muscle fibers
P (MF) and a capillary (C) in the lamina
MM
propria. A clear brush border (BB) is also
visible. (X400)
SM
14-18a
MF
BB
C
14-18b
14-18c
171
DUODENUM The duo-
denum is identifiable
by mucus secreting
Brünner’s glands (BG) in
the submucosa (SM).
(a) In this micrograph,
the Brünner’s glands are
not very dense. One
gland is seen emptying
into a crypt (arrow).
(X65) (b) The Brünner’s
glands are much denser
in this specimen. Several
glands are seen emptying
into the crypts (arrows).
BG
(X65)
SM
BG
SM
14-19a 14-19b
PC
CL
LP
LF
MM
SM
PP
14-20
JEJUNUM The jejunum has no diagnostic feature of its own, so 14-21
tentative identification may be made based on the absence of
Brünner’s glands and Peyer’s patches. Note the lymphatic follicle ILEUM The ileum has distinctive aggregations of lymphatic follicles called
(LF) in the submucosa (SM) and lamina propria (LP). Villi (V), Peyer’s patches (PP) in the submucosa and sometimes extending into the
crypts (CL), and the muscularis mucosae (MM) are also visible. lamina propria. Note the plica circulares (PC). (X25)
(X65)
172
GC
CL L
LP
LP
CL
MM MM
SM
SM
14-22a 14-22b
COLON MUCOSA The colon’s mucosa is characterized by long, straight crypts of Lieberkühn (CL) that penetrate the lamina propria (LP) to the
muscularis mucosae (MM). The submucosa (SM) is also visible. Note the abundant goblet cells (GC) in the epithelium and the lymphocytes (L)
in the lamina propria. Micrograph (a) is X65; micrograph (b) is X130.
M M
SM
SM
IC
TC
TC
IC
14-23a 14-23b
TAENIA COLI MUSCLES The longitudinal layer of muscle in the colon is thicker in three strips called taenia coli muscles (TC). As evidenced by
these micrographs, the taenia coli are not exceptionally thick for a longitudinal muscle layer. It is the thin regions between them that make them
prominent. Also visible are the mucosa (M), submucosa (SM), and inner circular (IC) layer of muscle. Both micrographs are X25.
173
RECTOANAL JUNCTION Compared to
the colon, the rectum has shorter crypts CL
(CL) with more goblet cells. At the
IS
rectoanal junction (arrow), the simple
columnar epithelium abruptly changes
to a nonkeratinized stratified squamous
epithelium. The internal anal sphincter
(IS) is a thickening of the inner circular
muscle layer. (X65)
14-24
HT
HT
HC
CV
HC CV
S
S
14-25a 14-25b
LIVER LOBULES The liver is divided into lobules, which are not very distinct in humans. These specimens are from other mammals to illustrate
lobule structure. At the center of the lobule is the central vein (CV). Radiating outward like spokes on a wheel are cords (HC) of hepatocytes,
between which are blood sinusoids (S). Hepatic triads (HT) composed of branches of the hepatic artery, hepatic portal vein, and bile duct are
present at corners of the lobules. (a) X60, (b) X65.
174
S
CV
CV
14-26a 14-26b
CV
CV
S
E
S
H
H
E K
14-26c 14-26d
CELLS OF THE LOBULE Hepatocytes (H) are arranged into cords that are
separated by the sinusoids (S), which are lined with endothelial cells (E).
Sinusoids drain into the central vein (CV). Phagocytic Kupffer cells (K)
are also found in the sinusoids. (a and b) Note the continuity of the
sinusoids and central vein and the reddish purple staining glycogen
in the hepatocytes in (a). Both micrographs are X265. (c) Note the
dark staining nuclei of the endothelial cells, which form an incomplete
barrier between the sinusoids and the hepatocytes. (X400) (d) Kupffer
cells in this preparation have phagocytosed black particles and are
easily distinguished from the endothelial cells. (X400).
BC
BILE CANALICULI Bile production is among the many

functions performed by hepatocytes. Bile canaliculi
(BC) are tiny channels formed between hepatocytes
that lead to a branch of the bile duct in the hepatic
triad. These bile canaliculi have been injected with a
black dye to make them more visible. Notice that they
are only found over hepatocytes, not in the sinusoids.
Normally, blood and bile don’t mix. (X265) 14-27
175
BD BD
PV HA
PV
HA
14-28a 14-28b
HEPATIC (PORTAL) TRIAD Branches of the hepatic artery (HA), hepatic portal vein (PV), and bile duct (BD) travel together throughout the liver
and form a hepatic triad. (a) Notice that the triad components are not unusual in appearance. That is, the artery is smaller and has a thicker wall
than the vein, and the bile duct is lined with a simple cuboidal epithelium. (X265) (b) This specimen was sectioned just above a branch point, so
each component is represented more than once. (X265)
SM Mu
SM
A Mu
14-29a 14-29b
GALL BLADDER The mucosa (M) of the gall bladder is lined with a simple columnar epithelium and is folded longitudinally. The submucosa (SM)
is a loose connective tissue. Deep to it is the muscularis (Mu), which is made of smooth muscle arranged into indistinct longitudinal, circular and
oblique layers. A thick adventitia (A) covers the outer surface. Micrograph (a) and (b) were magnified X25 and X65, respectively.
176
I
A D
I
C
14-30a
D
A
D
14-31
PANCREAS DETAIL This micrograph illustrates pancreatic acini
(A), an islet of Langerhans (I), and a small interlobar duct (D).
Note the eosinophilic secretory granules at the apices of the
exocrine cells. Two centroacinar cells (C), which are lining
cells of the intercalated duct, are also visible. (X265)
14-30b
PANCREAS The pancreas has an endocrine
component and an exocrine component. The
endocrine islets of Langerhans (I) secrete the
hormones insulin and glucagon. The exocrine
pancreatic acini (A) secrete pancreatic juice,
an alkaline fluid rich in digestive enzymes.
Interlobar ducts (D) are also present and are
recognizable by the connective tissue covering.
(a) Note the lymphatic tissue in the interlobar
duct at the right. (X115) (b) The light-staining
region at the bottom is a portion of a large B
interlobar duct. (X115) Also see Figure 10-9.
V
P
AMPULLA OF VATER The common

bile duct and the main pancreatic
duct join within the duodenal wall
to form the ampulla of Vater (V),
which delivers pancreatic juice and
bile to the duodenum. Also visible in
this micrograph are the pancreas (P)
—note the lobes—and Brünner’s
glands (B). (X25) 14-32
Respiratory System
C H A P T E R
15
Introduction to the Respiratory System is lined with stratified squamous epithelium and thick hairs,
The respiratory system is responsible for gas exchange be- called vibrissae. These act as a coarse filter.
tween the environment and lung capillaries. Anatomically, it Most of the nasal cavity is lined by respiratory epithe-
is divided into the upper respiratory tract including the nasal lium, a pseudostratified ciliated columnar epithelium (PSCC)
with goblet cells (Figures 9-1 and 15-2). Deep to the epithe-
cavity, pharynx, and larynx, and a lower respiratory tract
lium is a very vascular lamina propria that also contains
composed of the trachea, bronchial tree, and lungs. Besides
serous and mucous glands. Secreted mucus maintains a
conducting air, other important functions of the upper respi-
moist environment that humidifies the air. It also traps inert
ratory tract include warming and moistening the air to pre-
particles and microorganisms in inspired air, and the cilia
pare it for gas exchange, filtering it to prevent infection and
sweep the mucus to the throat where it is swallowed. The
the blockage of smaller airways, olfaction, and vocalization.
blood vessels of the lamina propria are responsible for pro-
Functionally, the respiratory system is divided into a
viding the heat to warm air.
conducting portion, responsible for transmitting air to the
In the upper nasal cavity and superior nasal conchae,
lungs, and a respiratory portion, in which respiratory gas respiratory epithelium is replaced with olfactory epithelium
exchange occurs. The actual sites of gas exchange are called (Figures 9-1 and 15-2). It differs from typical respiratory
alveoli, microscopic sacs lined with simple squamous epi- epithelium in its thickness and lack of mucous cells. Bipolar
thelium and surrounded by capillaries. neurons act as olfactory receptor cells. They are chemore-
The abundant alveoli occupy the majority of lung vol- ceptors. Short basal cells and taller sustentacular cells are
ume, so a discussion of lung histology is primarily a discus- also present, although they are difficult to differentiate in
sion of alveolar anatomy. However, on a gross level, each standard H&E preparations. Bowman’s glands in the lam-
lung is divided into lobes (three in the right lung, two in the ina propria produce a watery secretion that dissolves chem-
left), which in turn are divided into segments (ten in the icals and makes them more able to stimulate the receptors.
right lung, 8 in the left). The lung’s surface is covered with
serous membrane called visceral pleura (Figure 15-1). Larynx
The larynx is positioned between the pharynx and the tra-
Nasal Cavity chea. It is made of a cartilagenous (mostly hyaline) frame-
The nasal cavity is formed from a framework of bone and work lined with mucous membrane consisting of PSCC and
cartilage with a vascular mucous membrane covering. It is a lamina propria. On its anterior and superior aspect is the
divided by the nasal septum into right and left cavities. Just epiglottis (Figure 15-3), which is made of a leaf-shaped elas-
inside the external naris of each side is the vestibule, which tic cartilage covered with stratified squamous epithelium
177
178
on its superior part. It closes the opening to the larynx to ◗ There is an increase (relative to the size of the tube) in
prevent food from “going down the wrong pipe” when the amount of smooth muscle and elastic tissue.
swallowing. ◗ The number of glands and goblet cells decreases.
Within the larynx are two pairs of mucosal folds. The The bronchi (Figure 15-8) resemble the trachea, but the
true vocal folds (Figure 15-4) are involved in phonation cartilage is plate-like, with the plates being found on all
and are the superior pair. The false vocal folds are inferior. sides (that is, there is no open side as with the C-shaped
The vocal ligaments are deep to the stratified squamous rings). There are smooth muscle and elastic fibers present,
epithelium of the true folds. Muscular action increases and as are glands and lymph follicles, which are especially seen
decreases tension on the vocal folds to alter the pitch of the at branch points of the airways. The epithelium is PSCC
sound produced. with goblet cells that gets progressively shorter.
Bronchioles (Figures 15-9 and 15-10) are about 1 mm
Trachea
in diameter or less, but they are easily identified due to their
The trachea (Figure 15-5) is a tubular organ anterior to the absence of cartilage in the wall. The epithelium is generally
esophagus. It begins at the junction with the larynx and a simple ciliated columnar in larger bronchioles that tran-
continues to its bifurcation into the two primary bronchi. C- sitions into a simple ciliated cuboidal, and then to a simple
shaped hyaline cartilage rings provide support, while fibrous nonciliated cuboidal in the smallest bronchioles. Goblet
connective tissue filling between them provides flexibility. cells are only seen in the largest of bronchioles. Clara cells
Histologically, the trachea is composed of a mucosa, are also seen in bronchiolar epithelium and are identified
submucosa, and adventitia, and its structure establishes the by their dome-shaped apical surface. They provide a source
basic plan for the rest of the respiratory tree (Figure 15-6). of new cells and produce surfactant that reduces surface
The mucosa is composed of a typical respiratory epithe- tension in the lungs (see page 179). Glands are absent from
lium—a tall PSCC with goblet cells—and a fibrous lamina the lamina propria, but lymphoid tissue is often seen. Spiral
propria, often containing lymph follicles. An elastic lamina layers of smooth muscle occupy a majority of the wall.
separates the lamina propria from the submucosa, which is The conducting portion of the bronchial tree ends with
made of dense irregular connective tissue. Various serous the terminal bronchioles. These are less than 0.5 mm in
and mucous glands may be found in the submucosa. The diameter. The epithelium is a simple cuboidal, some ciliated,
adventitia is a fibroelastic connective tissue that houses the with Clara cells. Only a couple of muscle layers are present.
cartilagenous rings. The trachealis muscle (Figure 15-7), a Terminal bronchioles divide to produce respiratory
band of smooth muscle, occupies the open posterior of the bronchioles, which represent the beginning of the respira-
C-shaped rings. tory portion of the bronchial tree (Figures 15-9 and 15-11).
These histologically resemble terminal bronchioles but are
Bronchial Tree identifiable by the presence of alveoli in their walls. It is in
The trachea branches into two primary bronchi, each of the distal portion of respiratory bronchioles that cuboidal
which goes to a lung. Once in the lung, primary bronchi cells no longer have cilia.
divide to form secondary (lobar) bronchi. (From the sec- As one progresses through the respiratory bronchioles,
ondary bronchi on, all branches are intrapulmonary.) There the alveoli become more and more numerous. Eventually,
are three secondary bronchi supplying the three lobes of the wall is exclusively alveoli and the bronchiole has be-
the right lung and two supplying the two lobes of the left come an alveolar duct (Figures 15-9 and 15-12). A smooth
lung. Secondary bronchi divide to form tertiary (segmen- muscle cell is positioned around the opening of each alveo-
tal) bronchi, each of which supplies a distinct lung segment. lus giving the appearance of knobs at the entrance of the
Further branching produces, in order, bronchioles, terminal alveolus to the alveolar duct. At the end of the alveolar duct
bronchioles, respiratory bronchioles, alveolar ducts, and are a few clusters of alveoli called alveolar sacs, which have
alveolar sacs. a common opening called the atrium.
The general trends seen in the bronchial tree are as Alveoli
follows:
The diameter of an alveolus is about 200 µm, and between
◗ The diameter of the tubes decreases. the two lungs there are about 300 million of them with a
◗ The epithelium as a whole gets shorter and the cells get total surface area of approximately 140 m2. The extensive
flatter, going from a tall PSCC in the trachea to a simple surface area and the thin alveolar wall make them efficient
squamous in the alveoli. in gas exchange with the blood. It also accounts for the
◗ There is a decrease in the size of the cartilage, going spongy consistency of the lungs on a gross level.
from C-shaped rings in the trachea, to more plate-like There are three cell types found associated with alveoli
in the bronchi, to absent in the bronchioles. (Figure 15-13). These are the type I pneumocytes, type II
C H A P T E R F I F T E E N : R E S P I R A T O R Y S Y S T E M
179
pneumocytes, and alveolar macrophages (dust cells). The pores join adjacent alveoli through the septum and allow
majority of alveolar surface is covered by the simple squa- pressure equalization within the various lung compart-
mous type I pneumocytes, although they comprise only ments. The actual blood-air barrier is composed of the type
about 5% of the alveolar cells. They are thin, even by squa- I pneumocytes and capillary endothelium, plus their fused
mous standards, having a thickness of as little as 80 nm. basal laminae. It is through these layers oxygen and carbon
Their nuclei are infrequently seen due to the cells’ extensive dioxide must diffuse during gas exchange between the air
cytoplasm. and blood.
Type II pneumocytes (great alveolar cells) are cuboidal
with a rounded apical surface and a central nucleus. They
secrete surfactant that reduces surface tension and prevents
the alveoli from collapsing during expiration.
Alveolar macrophages are often seen in the alveolar
wall or lumen. They are derived from blood monocytes and
OE
phagocytose small particles that have slipped past the other
defenses.
The interalveolar septum is composed of the epithelium RE
of two alveoli separated by numerous capillaries supported

by elastic and collagen fibers. Tiny openings called alveolar
LP
SS CT
VP
15-2
NASAL EPITHELIUM Respiratory epithelium (RE) is a PSCC with goblet
cells. It lines most of the nasal cavity as well as the trachea and
bronchi. In the superior portion of the nasal cavity, the typical
respiratory epithelium is replaced by olfactory epithelium (OE),
identifiable by its thickness and lack of goblet cells. An arrow marks
A the junction of the two epithelia. The vascular lamina propria (LP) is
deep to the epithelium. (X210)
LP
G SS
15-1
P
VISCERAL PLEURA The outer surface of the lungs is covered with a
serous membrane known as visceral pleura (VP). It consists of a simple
squamous mesothelium (SS) and underlying connective tissue (CT).
Pulmonary alveoli (A) are seen deep to the pleura. (X250) EC
EPIGLOTTIS The epiglottis projects over the larynx and prevents

food from going down it during swallowing. The superior and
anterior portion is lined with nonkeratinized stratified squamous
(SS), as shown here. Mucous and serous glands (G) may be seen in
the lamina propria (LP). Internally, there is a leaf-shaped piece of
elastic cartilage (EC) surrounded by a perichondrium (P). (X20) 15-3
180
A CR
Es
SS E
V VL
VF
VM
15-4 15-5a
VOCAL FOLD The true vocal folds (VF) in the larynx are responsible
for sound production. Each (one is shown here) is lined with non- GC
keratinized stratified squamous epithelium (SS) and has the vocal
ligament (VL) and vocalis muscle (VM) within. The space above is
E
the ventricle (V). (X50)
LP
G
CR
15-5b
E
TRACHEA The tracheal mucosa is a PSCC (E) with a loose
connective tissue lamina propria (LP). Tracheal glands (G)
P may be present in the submucosa. C-shaped cartilagenous
rings (CR) made of hyaline cartilage and covered by a
perichondrium (P) provide structural support and are found
in the fibrous adventitia (A). The rings are positioned with
their open side toward the esophagus (Es). Their ends are
joined by the trachealis muscle (T). (a) The entire trachea is
CR shown in this micrograph. The esophagus is toward the left.
(X20) (b) In this micrograph, the light staining goblet cells
(GC) in the PSCC are just barely visible. (X50) (c) Mucous
secreting tracheal glands are seen in the submucosa of this
specimen. The space indicated by the arrow is a preparation
artifact. (X125)
P
15-5c
181
GC
BM
GC
15-6a
BM
TRACHEAL MUCOSA The PSCC with goblet cells (GC)
is characteristic of respiratory epithelium. Note the
thick basement membrane (BM) in both micrographs.
(a) In this specimen, there is very little lamina propria
and submucosa between the mucosa and the hyaline
cartilage. In fact, if it weren’t for the fact that they
are separated, it would be difficult to distinguish the
perichondrium (P) from the submucosa (S). (b) A G
tracheal gland (G) is visible in this micrograph. Both
micrographs are X380.
15-6b
CR
TRACHEALIS MUSCLE The trachealis muscle (T) is

smooth muscle that joins the ends of the C-shaped
rings in the trachea. The adventitia (A) and hyaline
15-7 cartilage ring (CR) are also visible. (X65)
182
SM
PV
L
E
SM
C
15-8a
E
15-8b
BRONCHI Bronchi are the first generations of
branches off the trachea and have a similar
construction. The plate-like cartilages (C)
and thinner epithelium (E) are the main
differences. (a) This specimen has only a
single cartilage at the plane of section and it
E does not encircle the bronchus—it is plate-
like. Note the smooth muscle (SM) and the
C
lymphatic tissue (L) in the submucosa. Also
SM seen is a branch of the pulmonary vein (PV).
As a side note, this section was made near a
point of bifurcation, as evidenced by the
indentations of the bronchial wall into the
lumen (arrows). (X50) (b) More plates of
cartilage are seen in this specimen, indicat-
ing it is from a larger bronchus than in (a).
(X50) (c) Smooth muscle and one piece of
cartilage are visible in this specimen. (X250)
15-8c
183
Bronchiole
Branch of pulmonary
vein
Lymphatic vessel
Pulmonary artery
Interlobular septum
Terminal bronchiole
Smooth muscle
Respiratory bronchiole
Capillary beds
Alveolar duct
Alveolar sac
Pleura Alveoli
15-9
RESPIRATORY TREE The final few branches of the respiratory tree are shown in this illustration. Terminal bronchioles represent the end of the
conducting portion of the respiratory tree. Beyond these, all components belong to the respiratory portion because they have alveoli and are
involved in gas exchange.
184
PA
PA
SM
15-10a
C
CC
LP
SM
15-10b
L BRONCHIOLES Bronchioles are only found within
the lung and maintain the general appearance of
the bronchi, only smaller. The absence of cartilage
distinguishes them from bronchi. The epithelium
ranges from low PSCC to simple ciliated columnar.
Smooth muscle is present. (a) The mucosa of these
SM bronchioles is folded, a result of the smooth muscle
(SM) in the wall and the absence of cartilage. Notice
the lung tissue around the bronchioles and the two
branches of the pulmonary artery (PA). (X65) (b)
L This bronchiole is lined with a simple ciliated
PA columnar (CC) epithelium lacking goblet cells. A
few dome-shaped and nonciliated Clara cells (C) are
present. Notice the thin lamina propria (LP) and
small amount of smooth muscle in the wall (SM).
(X250) (c) Lymphatic tissue (L) and smooth muscle
(SM) are seen in this bronchiole. Based on the PSCC
epithelium, it can be concluded that this bronchiole
is larger than the specimen in (b) even though only a
portion of it is shown. Notice the small branch of
15-10c the pulmonary artery (PA). (X230)
185
A
A
SM
C
C
C
A
A
A
15-11a 15-11b
RESPIRATORY BRONCHIOLES Respiratory bronchioles resemble terminal bronchioles in structure except for the alveoli (A) in their walls. Clara
cells (C) are present in the simple cuboidal epithelium. Only a few of the epithelial cells are ciliated. A thin layer of smooth muscle (SM) is also
visible. Both micrographs are X210.
RB
AS
RB AD
AS AD
AS
AD
AS
AS
15-12a 15-12b
ALVEOLAR DUCTS Each respiratory bronchiole (RB)
leads to an alveolar duct (AD), a passage made of
alveoli (A) that terminates in alveolar sacs (AS). (a) In
this micrograph, a respiratory bronchiole (RB) is seen AD
to lead into an alveolar duct and two (in this plane of
section) alveolar sacs. (X50) (b) This micrograph
shows a respiratory bronchiole branching into two
alveolar ducts. (X110) (c) Here is another example AS
of an alveolar duct and alveolar sacs. It takes some
searching of lung specimens to find examples cut in
the proper plane to illustrate the branching, so don’t
be discouraged if you don’t see one right away.
(X110)
AS
15-12c
186
TII
IS
MF
15-13a
ALVEOLI Each alveolus is lined by a simple squamous
epithelium that is difficult to see clearly in standard
preparations. More easily seen are the interalveolar
septa (IS) formed from the epithelia of neighboring
alveoli, plus capillaries (C) and a small amount of TII
elastic connective tissue between them. The alveolar
epithelium is composed of simple squamous Type I
pneumocytes (I) and the cuboidal Type II pneumo-
cytes (II). Alveolar macrophages (MF) may be seen
in the interalveolar septa or free in the alveolar IS
I
spaces. All micrographs are X630.
C
IS
15-13b
MF
I
TII
IS
C
15-13c
Urinary System
C H A P T E R
16
Introduction Uriniferous tubules constitute the functional units of
The organs of the urinary system are the two kidneys and the kidneys. Each tubule is composed of a nephron and a
ureters, the urinary bladder, and urethra. Its function is to collecting tubule, though the latter are shared by many
maintain fluid and electrolyte homeostasis by filtering the nephrons. Collecting tubules converge to form the ducts of
blood to remove wastes and excess ions, and conserve Bellini.
water and ions that are in short supply. It also is involved in There are approximately 1.3 million nephrons in each
maintaining blood pressure by adjusting blood volume. kidney. Each nephron consists of a renal corpuscle (Bow-
man’s capsule and glomerulus—see below), proximal con-
Kidneys voluted tubule, loop of Henle, and distal convoluted tubule
The kidneys are paired organs that occupy the retroperi- (Figure 16-1). Cortical nephrons are located in the cortex,
toneal region just inferior to the diaphragm on the dorsal with their loops of Henle barely penetrating the medulla.
body wall. They are kidney bean in shape (!) and the inden- Juxtamedullary nephrons are located near the junction of
tation, called a hilus, faces medially. Renal blood vessels, the cortex and medulla. They have long loops of Henle that
lymphatics, and the ureter enter at the hilus. The renal penetrate deep into the medulla.
pelvis is a dilation of the ureter’s proximal end. It branches The oval renal corpuscles (Figure 16-2) are the site of
to form major and minor calyces that connect with the blood filtration. They are located in the cortex and are quite
apices of the pyramids (see below). A fibrous capsule covers complex in structure. Each is composed of a capillary tuft
the kidney’s surface. called a glomerulus, which is surrounded by a Bowman’s
Internally, the kidney presents a renal cortex and renal capsule. The glomerulus is made of an endothelium with
medulla. In section, the medulla appears as six to 12 trian- fenestrations up to 100 nm in diameter. A three-layered basal
gular and striated regions called renal pyramids, which ac- lamina is deep to the endothelial cells and contributes to the
curately describes their three-dimensional shape. The apex filtration mechanism.
of each pyramid projects into a minor calyx as a renal During development, the glomerulus pushes into the di-
papilla and has approximately twenty openings called ducts lated and spherical Bowman’s capsule at the proximal end
of Bellini. This forms the area cribrosa. The base of each of the nephron, not unlike a fist pushing into a deflated
pyramid is adjacent to the cortex. The cortex has a granu- playground ball and collapsing it upon itself. The outer
lar appearance and occupies the outer portion of the kid- layer of Bowman’s capsule, the parietal layer, is a simple
ney, forming the cortical arch. Thin extensions of medullary squamous epithelium. The inner layer, which is in contact
tissue into the cortical arch are called medullary rays. Corti- with the glomerular capillaries, is called the visceral layer
cal columns are found between the pyramids. and is composed of cells called podocytes (Figure 16-3).
187
188
Podocyte processes wrap around the glomerular capillaries, (Figure 16-7). The distal tubule passes between its own af-
but leave small gaps about 25 nm in width called filtration ferent and efferent arterioles. This portion is called the mac-
slits. ula densa and is characterized by narrower cells. The pars
Blood enters the glomerulus through the afferent arteri- convoluta is tortuous but shorter than its proximal counter-
ole and leaves through the efferent arteriole, both of which part (and therefore less abundant in sections of renal cor-
penetrate the renal corpuscle at its vascular pole (Figure tex). The cells are paler staining with round, apical nuclei
16-4). The afferent arteriole is larger than the efferent arteri- and prominent nucleoli, and the lumen is typically open.
ole, resulting in resistance to blood flow. The pressure pro- The distal tubule is impermeable to water and urea, which
duced by this size difference forces fluid and molecules contributes to the mechanism of urine concentration. In ad-
smaller than a molecular weight of 65,0000 daltons from dition, a sodium-potassium pump reclaims sodium from
the blood through the filtration slits and basal lamina, and the urine in response to the hormone aldosterone from the
into the interior of Bowman’s capsule, Bowman’s space. adrenal cortex, and excess hydrogen and potassium ions
The fluid is called glomerular ultrafiltrate. In healthy kid- are pumped into the lumen.
neys, blood cells and large molecules are absent from the The juxtaglomerular (JG) apparatus (Figure 16-8) is
ultrafiltrate. formed from the macula densa, juxtaglomerular (JG) cells
Intraglomerular mesangial cells are associated with the in the tunica media of the afferent arteriole (and sometimes
glomerulus and are phagocytic in function. Their primary the efferent arteriole) and the extraglomerular mesangial
role is to remove molecules trapped on the basal lamina cells. The basal lamina is absent from the macula densa
that would impede filtration if they weren’t removed. They cells, so they are in direct contact with the JG cells. The JG
are also contractile and adjust blood flow through the apparatus is involved in maintaining blood pressure via two
glomerulus. Extraglomerular mesangial cells are located at separate mechanisms. First, if the macula densa cells detect
the vascular pole. a low sodium concentration, they cause dilation of the af-
Emerging from the urinary pole of the renal corpuscle ferent arteriole, which increases intraglomerular pressure.
is the proximal tubule, whose lumen is continuous with Second, they stimulate the JG cells to release renin into the
Bowman’s space (Figure 16-5). The epithelium is simple blood. Renin activates the angiotensin system, which causes
cuboidal with a brush border and a prominent basal lam- vasoconstriction and increases blood pressure. It also stim-
ina, demonstrable with a PAS stain. In most paraffin prepa- ulates the adrenal cortex to release aldosterone, resulting in
rations, the lumen is closed. Juxtamedullary nephrons have greater sodium resorption and an increase in blood volume,
two parts to the proximal tubule: The tortuous proximal which also increases blood pressure.
convoluted tubule (pars convoluta) in the cortex, and the The distal convoluted tubules of several nephrons
straight descending thick limb of Henle’s loop (pars recta) empty into a collecting tubule, which is not considered part
in medullary rays. Up to 80% of the sodium, chloride, and of the nephron. Cortical collecting tubules have a simple
water is reabsorbed in the proximal tubule and returned cuboidal epithelium and are located in the medullary rays
to the blood, as are all the glucose and amino acids in the (Figure 16-9). They are involved in acid-base homeostasis
ultrafiltrate. through secretion of hydrogen ions. Cortical collecting
The descending thin segment of Henle’s loop is a con- tubules join to form medullary collecting tubules, which
tinuation of the thick segment (Figure 16-6). It passes into also have a simple cuboidal epithelium but are located in
the medulla where it makes a hairpin turn called Henle’s the pyramids. These join and form papillary collecting
loop, and continues back to the cortex as the ascending tubules (ducts of Bellini) (Figure 16-10) that empty into a
thin segment of Henle’s loop. All parts are lined with a sim- minor calyx at the renal papilla. All collecting tubules are
ple squamous epithelium. The thin limbs resemble capillar- impermeable to water, but their permeability is increased in
ies (which are also present—see page 189) except for their the presence of antidiuretic hormone (ADH); the more
thicker epithelial cells with less dense staining nuclei and ADH, the more water leaves the tubule, and the more con-
absence of red blood cells within. Henle’s loop is involved centrated the urine becomes.
in a countercurrent mechanism for concentration of the
urine; it establishes the range of osmolarities for final urine Renal Circulation and the Renal Interstitium
concentration. This is accomplished, in part, by the relative Normally, gross circulation is not in the domain of histol-
impermeability to water by the ascending limb. ogy, but often the following blood vessels are identifiable in
The ascending limb of Henle’s loop continues as the kidney sections. Blood enters the kidney through the renal
distal tubule. It is composed of an ascending thick limb artery at the hilus. The renal artery branches into segmental
of Henle’s loop (pars recta), the macula densa, and the dis- arteries, lobar arteries, interlobar arteries (found next to the
tal convoluted tubule (pars convoluta). A simple cuboidal pyramids), arcuate arteries (found along the base of pyra-
epithelium lines the distal tubule, which is in the cortex mids), and interlobular arteries (which go into the cortical
C H A P T E R S I X T E E N : U R I N A R Y S Y S T E M
189
arch between medullary rays). Several afferent arterioles serosa. The mucosa consists of a transitional epithelium
branch off the interlobular arteries, empty into the glomeru- and a lamina propria made of a superficial dense connective
lus, which is drained by the efferent arteriole. Peritubular tissue and a deeper loose connective tissue. The epithelium
capillaries (for cortical nephrons) and vasa recta (for juxta- is an osmotic barrier, preventing water from the bladder’s
glomerular nephrons) arise from the efferent arteriole and wall entering the hyperosmotic urine. In the empty bladder,
form a plexus around the convoluted tubules and loop of the mucosa demonstrates nonpermanent folds called rugae
Henle, respectively (Figure 16-11). These extensive capil- (except at the trigone). These flatten as the bladder fills.
lary networks pick up materials that have left the renal Three loosely organized layers of smooth muscle form
tubule as a result of active transport, osmosis, or diffusion. the muscularis. These correspond to the layers seen in the
Thus, useful materials that left the blood as ultrafiltrate are distal ureter. That is, an inner longitudinal layer, middle cir-
returned to the blood. cular layer, and an outer longitudinal layer, but these are
Arcuate veins drain the capillaries. The veins leading often difficult to identify. The circular layer is thicker at the
out of the kidney follow the arteries coming in, and have internal urethral orifice where it forms the internal urinary
the same names (with the exception of lobar and segmental sphincter. A dense irregular connective tissue comprises the
veins, which are absent). adventitia, which covers most of the organ. The superior
All blood vessels in the kidney are found in the renal in- aspect of the bladder is covered with a serosa, the parietal
terstitium, the region between the tubules. It is composed of peritoneum.
a small amount of fibrous connective tissue with fibrob-
lasts, macrophages, and interstitial cells. Urethra
The urethra carries urine from the bladder to the external
Calyces and Renal Pelvis environment. As the urethra penetrates the urogenital dia-
Each renal papilla projects into a minor calyx. Minor phragm, a ring of skeletal muscle acts as the voluntary
calyces combine to form three or four major calyces, which external urinary sphincter.
in turn empty into the renal pelvis. All are lined with a tran- The female urethra (Figure 16-14) is about 5 cm long
sitional epithelium and lamina propria. A thin smooth and 6 mm in diameter, and the external urethral orifice
muscle layer is deep to the lamina propria and is respon- opens into the vestibule of the vagina. The mucosa presents
sible for moving urine toward the ureters. longitudinal folds and is mostly lined with nonkeratinized
stratified squamous epithelium with occasional patches of
Ureters PSCC. Glands of Littré, found in the lamina propria, se-
The ureters are 25 to 30 cm long and 4 mm in diameter. crete mucus to lubricate the mucosa. An inner longitudinal
They pass along the dorsal body wall and enter the pos- layer and an outer circular layer of smooth muscle com-
terior and inferior aspect of the urinary bladder. Each is prise the muscularis. The muscularis is surrounded by
composed of three layers: mucosa, muscularis, and adventitia skeletal muscle at the external urinary sphincter.
(Figure 16-12). The male urethra (Figure 16-15) is longer than the fe-
The mucosa is made of a transitional epithelium and a male urethra, being up to 20 cm in length. It is divided into
dense irregular connective tissue in the lamina propria. three segments based on location. These are the prostatic
Deep to this is the muscularis made of two smooth muscle urethra, membranous urethra, and spongy (penile) urethra.
layers: an inner longitudinal layer and an outer circular The prostatic urethra, lined with transitional epithe-
layer. Note that the arrangement is just the opposite of the lium, passes through the prostate gland. The microscopic
muscularis layers in the digestive tube, but the function is prostatic ducts and the two ejaculatory ducts empty into it.
the same; that is, peristalsis. In the distal third of the ureter, The membranous urethra passes through the urogenital di-
an additional layer of longitudinal muscle is added on the aphragm. Its epithelium is stratified columnar with occa-
surface of the circular layer. The outermost layer is the sional regions of PSCC. This segment is the site of the
fibrous adventitia, which blends with the renal capsule and external urinary sphincter. The spongy (penile) urethra
the adventitia of the urinary bladder. passes through the corpus spongiosum of the penis and
leads to the external urethral orifice at the end of the glans
Urinary Bladder penis. Most of its length is lined with stratified columnar or
The urinary bladder stores urine until micturition. A PSCC epithelium with stratified squamous at the orifice.
smooth, triangular region called the trigone is formed As with the female, numerous glands of Littré are pres-
where the two ureters enter and the urethra leaves. ent in the lamina propria of all segments.
There are three layers in the urinary bladder wall (Figure
16-13). These are the mucosa, muscularis, and adventitia or
190
Capillaries of Bowman’s capsule

glomerulus
Renal capsule
Distal
convoluted
tubule
Distal convoluted tubule

Efferent
arteriole
Artery
Afferent Renal
Vein corpuscle
arteriole
Proximal Collecting duct

convoluted
Capillary tubule
bed
Proximal Collecting
convoluted Thick tubules
tubule ascending
limb
Thin
descending
limb Papillary
Ascending duct
limb
Descending limb
Loop of
Henle
16-1
THE NEPHRON Shown is an artist’s rendition of a juxtamedullary nephron.
BC
C BC
BS
BS
M
G
P
C RENAL CORPUSCLE Renal corpuscles consist of
a glomerulus (G) surrounded by a Bowman’s
capsule. The parietal layer of Bowman’s
capsule (BC) is a simple squamous epithelium
and lines Bowman’s space (BS). The glomerular
capillaries (C), podocytes (P), and mesangial
cells (M) can be difficult to sort out in some
BC preparations, but are visible in this specimen.
16-2 (X530)
191
Pe P
Pe
RBC E
BL
F
16-3b
ELECTRON MICROGRAPH OF A GLOMERULUS (a) Visible in this

micrograph is a podocyte (P) with pedicels (Pe) on the endothelium
(E) of a glomerular capillary. An RBC and a mesangial cell (M) are
also visible. (X3700) (b) This is an enlargement of the boxed area in
(a). The gaps between pedicels (Pe) and the capillary fenestrations (F)
are visible, as is the three-layered basal lamina (BL). (X18,500)
(Courtesy of UCSD Medical Center.)
16-3a
AA VP
EA GC
BC
BS BS
16-4a 16-4b
AFFERENT AND EFFERENT ARTERIOLES (a) In this micrograph, the blood vessels have been injected with a red dye. Notice the afferent arteriole
(AA) has a larger diameter than the efferent arteriole (EA) emerging from the glomerulus (G). Bowman’s space (BS) is also visible though the
details of Bowman’s capsule are not. (X380) (b) In standard sections, the vascular pole (VP) of the renal corpuscle is often visible, but it is
difficult to distinguish between the afferent and efferent arterioles. Glomerular capillaries (GC) with blood cells and the parietal layer of
Bowman’s capsule (BC) are visible, however. (X680)
192
DCT
DCT
PCT
BS M BM
PCT
C
16-5b
M
C
DCT
16-5a
PROXIMAL CONVOLUTED TUBULE (a) The lumen of a proximal
convoluted tubule (PCT) is continuous with Bowman’s space
(BS). Proximal tubules have a simple cuboidal epithelium with
microvilli (M), barely discernable in this specimen. All the
tubules in this field are proximal convoluted tubules. Note
the abundant capillaries (C) surrounding the tubules. (X660)
(b) In this PAS preparation, the basement membranes (BM) M
are prominently shown, as are the microvilli lining the proxi-
mal tubules. Most tubules in the cortex are proximal tubules,
but a couple of distal convoluted tubules (DCT) are also identi- C
fiable in this specimen by their open lumen and lack of micro-
villi. (X260) (c) The microvilli of proximal convoluted tubules
are seen in this micrograph, as are capillaries and a distal con-
voluted tubule. (X400)
16-5c
CT CT
CT
H
H
VR H
H
CT
16-6a 16-6b
RENAL MEDULLA The medulla has thick and thin segments of the ascending and descending limbs of Henle’s loop as well as collecting ducts.
(a) In this micrograph, thin segments of Henle’s loop (H) and collecting tubules (CT) are visible, as are vasa recta (VR) with blood cells in them.
(X660) (b) In this section of the medulla, thick segments of Henle’s loops are more abundant than in (a). Collecting tubules can be differentiated
from other tubules by their pale cytoplasm and prominent lateral cell membranes. Capillaries (C) and an occasional thin segment of Henle’s loop
(arrow) are also visible. (X265)
193
PCT PCT
C
J
DCT
MD
PCT
DCT
PCT
PCT
PCT
16-8a
MD
16-7
J
DISTAL CONVOLUTED TUBULE Distal convoluted tubules (DCT) are
shorter than proximal tubules (PCT), and so are seen less frequently
in sections of renal cortex. Their lumen is more open and they lack
microvilli. (X260)
CT
MR
C 16-8b
JUXTAGLOMERULAR APPARATUS The distal convoluted tubule contacts the
afferent (and sometimes the efferent) arteriole at the vascular pole of its
renal corpuscle to form the juxtaglomerular apparatus. The arteriole
contributes juxtaglomerular cells (J), whereas the distal tubule cells
form the macula densa (MD). The macula densa cells are smaller with
darker staining nuclei than the other cells of the distal tubule. (a) Note
RC MR the abundance of proximal tubules (PCT) and the relative scarcity of
distal tubules in this specimen. Also notice the capillary (C). (X365) (b)
The macula densa and juxtaglomerular cells are clearly evident in this
micrograph. Note that the distal tubule of the JG apparatus is the only
RC one in the field. (X610)
MEDULLARY RAY Medullary tissue is found

in the subcapsular cortex as medullary rays
(MR). They are easily distinguished from
the cortex (C) by the absence of renal
corpuscles (RC). The rays contain cortical
16-9 collecting tubules (CT). (X65)
194
MD
P
MC
16-10
RENAL PAPILLA The medullary collecting tubules 16-11a
converge at the renal papilla to form papillary
collecting tubules (ducts of Bellini). The papilla
projects into the minor calyx (MC). (X110)
VR
VR
16-11b
VR
BLOOD VESSELS ASSOCIATED WITH THE NEPHRON

VR
(a) Peritubular capillaries (P) are found in the
interstitium around proximal and distal
convoluted tubules. Endothelial cell nuclei (E),
and the macula densa (MD) and juxtaglomerular
cells (J) of the JG apparatus are also seen in this
specimen. (X580) (b) The vasa recta (VR) of the
medulla near the cortex are often in bundles and
are surrounded by thick segments of Henle’s
loop (H). (X265) (c) The vasa recta near the
cortex are seen longitudinally in this injected
specimen. (X65) 16-11c
195
A
OL
C
IL
LP
M T
LP
T
IL
C
16-12a
16-12b
T
LP
IL
16-12c 16-12d
URETER The ureter wall consists of a mucosa (M) made of transitional epithelium (T) and a dense connective tissue lamina propria (LP). Inner
longitudinal (IL) and outer circular (C) layers of smooth muscle comprise the muscularis. On the outer surface is a fibrous adventitia (A). (a) In
the distal third of the ureter, an outermost longitudinal (OL) layer of smooth muscle is found. (X50) (b) This specimen was magnified X100. (c)
This specimen was magnified X190. (d) The transitional epithelium lining the ureter is clearly seen in this longitudinal section. (X320)
T
LP
R M
LP
T R M
16-13a 16-13b
URINARY BLADDER Transitional epithelium (T) and a lamina propria (LP) of superficial dense connective tissue and deeper loose connective tissue
are in the mucosa of the urinary bladder. The mucosa has nonpermanent folds called rugae (R) that allow the bladder to accommodate filling
with urine. The muscularis (M) has three indistinct layers of smooth muscle. On the outer surface is a fibrous adventitia or a serosa, but they are
not shown in these micrographs. Both micrographs are X20.
196
LP
16-14a 16-14b
FEMALE URETHRA The female urethra is lined by a transitional epithelium (T) in the proximal segment and a nonkeratinized stratified squamous
in the distal part. The mucosa is folded due to a highly elastic lamina propria (LP). Thin walled veins (V) form erectile tissue similar to the corpus
spongiosum of the male. Inner longitudinal and outer circular smooth muscle layers comprise the muscularis (M). (a) In this panoramic view, the
longitudinal mucosal folds and the venous sinuses are prominent. (X100) (b) The transitional epithelium indicates that this section is from the
proximal urethra. (X250)
E
E
LP
LP
16-15a 16-15b
MALE URETHRA The male urethra differs in structure in its three segments. The epithelium of the prostatic urethra is transitional, but most of its
length is stratified columnar (E) or PSCC. Shown in these micrographs is the longest portion, the spongy urethra. A fibroelastic lamina propria
(LP) and blood sinusoids (S) of the corpus spongiosum surround it. (a) X65 (b) X250
Reproductive Systems
C H A P T E R
17
Introduction to the Reproductive Systems layer and a visceral layer in contact the testicular surface.
The male and female reproductive systems are responsible The space between is lubricated by serous fluid. Deep to the
for producing gametes (sex cells), a process known by the tunica vaginalis is a tough, thick fibrous covering called the
general term gametogenesis. Ova are the female gametes tunica albuginea (Figure 17-1). Deep to the tunica albuginea
and are produced by oogenesis, whereas the male gamete, is a layer of vascular loose connective tissue called the tu-
sperm, is produced by spermatogenesis. The organs respon- nica vasculosa. Fibrous extensions of the tunica albuginea
sible for gametogenesis are called gonads—ovaries in the fe- form septa and divide each testis into about 250 lobules.
male and testes in the male. In addition to gametogenesis, Each testicular lobule contains up to four seminiferous
the gonads also produce sex hormones, which are necessary tubules, which are responsible for spermatogenesis. The
for reproduction as well as establishment of morphological seminiferous tubules empty into the rete testis, a network of
sex in the male and secondary sex characteristics in both tubules that lead to the ductuli efferentes, which carry sperm
males and females. Lastly, the reproductive systems have to the epididymis located on the posterior of the testis.
organs of copulation—the penis of the male and the vagina The interior of the testis is composed of seminiferous
of the female. tubules and the testosterone producing interstitial cells (of
Leydig) that occupy the vascular connective tissue (derived
Male Reproductive System from the tunica vasculosa) between the seminferous tubules.
The organs of the male reproductive system include the Each seminiferous tubule is up to 70 cm long and is
testes, epididymis, ductus deferens, ejaculatory duct, and composed of a thin connective tissue covering called the tu-
urethra. Accessory glands include the prostate gland, seminal nica propria and the relatively thick germinal (seminifer-
vesicles, and the bulbourethral glands. ous) epithelium with its basal lamina. Contractile myoid
cells are present in the tunica propria of some animals, but
Testes not humans. Several layers of spermatogenic cells in various
The testes are the primary sex organs or gonads of the stages of meiosis, which reduces the diploid chromosome
male. They develop in the abdominal cavity, then descend number to the haploid number, and the taller Sertoli cells
into the scrotum about the 28th week. In the adult, they are comprise the germinal epithelium.
about 4 cm long and 2.5 cm in diameter. Spermatogenic cells are the most abundant cells in the
Each testis is almost completely covered by a double germinal epithelium (Figure 17-2). Type A spermatogonial
layer of peritoneum called the tunica vaginalis obtained cells are located near the basal lamina and undergo mitotic
from the abdominal cavity during its descent into the scro- divisions to produce more type A cells as well as type
tum. As with all serous membranes, there is an outer parietal B spermatogonia. The latter divide to produce primary
197
198
spermatocytes, which migrate closer to the lumen. These at the tail. Long microvilli, called stereocilia (a misnomer),
are identifiable by their abundant cytoplasm and condensed project from the tallest cells and are involved in reabsorp-
chromatin in the nucleus. Primary spermatocytes undergo tion of fluid. Smooth muscle increases from a single circular
the first meiotic division to produce secondary spermato- layer at the head to three layers in the tail resembling what
cytes, which are difficult to find because they immediately is seen in the ductus deferens.
undergo the second meiotic division and produce sper-
Ductus Deferens and Ejaculatory Duct
matids with dark staining nuclei. The haploid spermatids
then undergo spermiogenesis, the conversion into a mature The ductus (vas) deferens (Figure 17-7) is a muscular tube
sperm cell, and are easily identified by their small and that carries sperm from the epididymis to the urethra. Its
pointed nuclei and their position along the luminal surface. terminal portion, the ampulla, is dilated and joins with
Mature sperm cells (Figure 17-3) are composed of a the seminal vesicle to form the ejaculatory duct. The mu-
head, midpiece, and tail. The head contains the haploid nu- cosa is folded longitudinally and is lined with a pseudos-
cleus and is capped by the acrosomal vesicle. It is derived tratified columnar epithelium resting on a thin lamina
from Golgi apparatus and contains hydrolytic enzymes propria. The thick muscularis is composed of an inner lon-
used in fertilization. The midpiece contains abundant mito- gitudinal, a middle circular, and an outer longitudinal layer
chondria responsible for producing ATP necessary to oper- of smooth muscle. These produce peristaltic contractions
ate the flagellum, which constitutes the tail. during ejaculation.
Sertoli cells are columnar and have a basal nucleus with Urethra
dispersed chromatin and a single nucleolus. They are re- The male urethra (Figures 16-15 and 17-10) is between 15
sponsible for nourishing the sperm cells during their devel- and 20 cm long. Three segments are identifiable based on
opment in the seminiferous tubule’s wall. Spermatogenesis location. These are the prostatic urethra, membranous ure-
requires follicle stimulating hormone (FSH) from the anterior thra, and spongy (penile) urethra. The prostatic urethra
pituitary gland. Sertoli cells, which respond by producing passes through the prostate gland and is lined with transi-
and secreting an androgen binding protein into the lumen, tional epithelium. The microscopic prostatic ducts and the
are thought to be its target. two ejaculatory ducts empty into it. The membranous ure-
A round nucleus with dispersed chromatin and one or thra, lined with stratified columnar epithelium, passes
two nucleoli, and an eosinophilic cytoplasm with lipid vac- through the urogenital diaphragm. This segment is the site
uoles characterize the interstitial cells (Figure 17-4). They of the external urinary sphincter. The spongy (penile) ure-
occupy the region between seminiferous tubules and are thra is found within the corpus spongiosum of the penis. It
surrounded by numerous capillaries, into which they se- is lined with stratified columnar or PSCC epithelium with
crete the male sex hormone testosterone. It is necessary for stratified squamous at the external urethral orifice.
embryonic development into a male, development of sec-
ondary sex characteristics, and normal sperm development. Seminal Vesicle
Interstitial cell activity is under the control of luteinizing The seminal vesicles (Figure 17-8) produce a secretion rich
hormone (LH) from the anterior pituitary. in fructose, prostaglandins, and other materials. The mu-
cosa is highly folded and is lined with a pseudostratified ep-
Rete Testis, Ductulus Efferens, and Epididymis ithelium. The secretory cells are tall and have a foamy
appearance due to the lipid contents. A muscularis consist-
The seminiferous tubules empty into a network of tubules
ing of an inner circular layer and an outer longitudinal
called the rete testis (Figure 17-5). These are lined with a
layer of smooth muscle creates peristaltic contractions dur-
simple cuboidal epithelium whose cells have microvilli and
ing ejaculation and forces the secretion into the ejaculatory
a single cilium. Contraction of surrounding myoid cells and
duct.
ciliary action move sperm along the tubule.
Fifteen to twenty ductuli efferentes receive sperm from Prostate Gland
the rete testis and deliver them to the epididymis where they The prostate gland (Figure 17-9) is about 4 cm long and 3
mature and become motile. The efferent ductules are lined cm wide and is found inferior to the urinary bladder. It sur-
with simple ciliated columnar or simple nonciliated rounds the proximal (prostatic) urethra and also surrounds
cuboidal epithelium. A thin smooth muscle layer is also the ejaculatory ducts. Its secretion is rich in many chemi-
present. cals, including citric acid, lipids, and proteolytic enzymes.
An epididymis (Figure 17-6) is found on the posterior A vascular dense irregular connective tissue capsule
of each testis. It consists of a head (superior), body and tail containing smooth muscle cells surrounds the prostate and
(inferior) and is the site of sperm storage and final matura- penetrates into the gland as the stroma. Up to 50 tubulo-
tion, which includes developing motility. It is lined with a alveolar glands are arranged into concentric layers around
pseudostratified epithelium that is thicker at the head than the urethra. The mucosal glands form the layer next to the
C H A P T E R S E V E N T E E N : R E P R O D U C T I V E S Y S T E M S
199
urethra. These are surrounded by the submucosal glands full maturity over a woman’s reproductive years at a rate of
and the main glands, which comprise the thickest layer. A one per month.
simple to pseudostratified columnar epithelium lines them. Beginning with puberty and approximately every 28
Pancreatic concretions (corpora amylacea) are often found days thereafter until menopause, and under the influence of
inside the glands. They are made of glycoprotein and are of Follicle Stimulating Hormone (FSH) from the anterior pitu-
unknown function, but they do increase with age. itary, primordial follicles continue their development. At
this stage of development, the primordial follicles consist of
Penis
a single layer of flat cells surrounding the primary oocyte.
The penis is a cylindrical organ that functions in urine elim- The primary oocyte has a large, eccentric nucleus with one
ination and as the male copulatory organ. It is covered with nucleolus and is approximately 25 µm in diameter. Devel-
skin, which is hairless at the distal end, but has coarse hairs opment continues as the primary oocyte enlarges to about
proximally. A loose hypodermis is also present.
100 µm and the follicle cells, now called granulosa cells
Internally, three cylindrical bodies of erectile tissue run
(that form the zona granulosa), become stratified to form a
its length (Figure 17-10). These are the two dorsal corpora
multilaminar primary follicle. The zona pellucida, an amor-
cavernosa and the ventral corpus spongiosum, which sur-
phous glycoprotein layer, forms between the primary
rounds the spongy urethra. A fibrous connective tissue layer
oocyte and the granulosa cells. In addition, the stroma near
called the tunica albuginea surrounds each cavernous body.
the follicle forms an inner vascular and cellular layer called
The erectile tissue is composed of vascular spaces lined with
the theca interna, and an outer fibrous layer called the
endothelium and separated by connective tissue trabeculae.
theca externa. The theca interna and granulosa cells pro-
These spaces fill with blood during erection.
duce the hormone estrogen.
Semen Development of a primary follicle into a secondary fol-
Semen contains only about 5% sperm cells by volume. The licle involves continued proliferation of the granulosa cells
seminal vesicles (70%), prostate gland (25%), and bulbo- and the formation of small intercellular spaces that coalesce
urethral glands (minimal) contribute the remainder of semi- to form the antrum. Follicular fluid (liquor folliculi) accu-
nal fluid. mulates in the antrum. Some granulosa cells remain around
the oocyte and form the cumulus oophorus. Further devel-
opment results in the layer of cells next to the zona pellu-
Female Reproductive System cida retracting from the oocyte but remaining attached via
Female reproductive organs include the ovaries, uterus and thin cytoplasmic threads. This single layer of cells is called
fallopian tubes, vagina, and breasts with mammary glands. the corona radiata.
In addition, the placenta and umbilical cord are organs of Of the many secondary follicles that form each month,
pregnancy. only a few continue development into a mature Graafian
follicle, which attains a size of 2.5 cm and bulges from the
Ovary ovary’s surface. The primary oocyte and the corona radiata
Ovaries are the female gonads (Figure 17-11). Like the detach and float free in the follicular fluid until ovulation.
testes, the ovaries have a gametogenic and an endocrine The primary oocyte completes the first meiotic division and
function. A simple cuboidal epithelium called germinal epi- forms a secondary oocyte and the first polar body. Most of
thelium and derived from peritoneum covers each ovary. the cytoplasm ends up in the secondary oocyte; the polar
Deep to it is the thin tunica albuginea, made of dense irreg- body serves its purpose by carrying the second haploid nu-
ular connective tissue. The bulk of the ovary is divided into cleus produced by meiosis I. Then, the secondary oocyte be-
a cellular cortex and a loose connective tissue medulla, al- gins the second meiotic division.
though the boundary between them is not sharp. Ovulation is a complex process, but basically is associ-
Within the ovarian cortex (Figure 17-12) are develop- ated with a spike of Luteinizing Hormone (LH) from the an-
ing ovarian follicles surrounded by a connective tissue terior pituitary gland, which leads to two events. These are
stroma. The embryonic ovarian cortex is populated by an increased internal pressure due to follicular fluid accumu-
oogonial cells from the yolk sac. Of the 5 to 7 million origi- lation, and the weakening and rupture of the follicle’s granu-
nal oogonial cells, only about 1 million become surrounded losa cells near the ovarian surface. The secondary oocyte
by follicle cells to form primordial follicles and survive to enters the infundibulum of the fallopian tubes (see next
the female child’s birth. The cells derived from oogonia are page) and begins its journey toward the uterus and, perhaps
called primary oocytes and they are suspended in prophase fertilization. If fertilization does not occur within about 24
of meiosis I until they continue development just prior to hours after ovulation, the secondary oocyte is phagocytosed.
ovulation. Only about 400,000 primordial follicles survive Meanwhile, under the influence of luteinizing hormone
to the beginning of puberty. Of these, only about 450 reach (LH) the granulosa cells remaining in the ovary form a corpus
200
hemorrhagicum, which develops into a corpus luteum (Fig- the superior portion of the body. The fundus is located
ure 17-13). The majority of corpus luteum cells is derived superior to the entry of the fallopian tubes. The cervix is
from the granulosa cells. These granulosa-lutein cells are the cylindrical portion that projects into the vagina.
large and pale staining, and secrete progesterone and estro- The uterine wall is composed of three major layers (Fig-
gens. Cells derived from the theca interna cells are at the ure 17-17). These are the endometrium, myometrium, and
periphery and are smaller, fewer in number, and darker perimetrium. The endometrium is the uterine mucosa and is
staining than the granulosa lutein cells. They are called the site of blastocyst (the multicellular derivative of the fer-
theca-lutein cells and also produce progesterone and estro- tilized ovum after about one week of development) implan-
gens. Failing fertilization, this corpus luteum of menstrua- tation during pregnancy. It is lined with a simple columnar
tion dies and becomes replaced with fibroblasts to form the epithelium overlying a vascular stroma. Glycogen-secreting
corpus albicans (Figure 17-14). simple tubular uterine glands are the most distinctive fea-
If fertilization does occur, the corpus luteum continues ture of the uterine mucosa. During the proliferative (follicu-
enlarging to a size of 5 cm and forms a corpus luteum of lar) phase of the menstrual cycle, these glands elongate and
pregnancy (Figure 17-15), which supplies estrogens and become coiled under the influence of ovarian estrogen. Co-
progesterone necessary to maintain the pregnancy. The pla- incident with this, the stroma gets thicker and more vascu-
centa assumes this function after about 3 months, but the lar. Glycogen secretion and the vascularity of the stroma
corpus luteum continues to contribute for several months both serve to support the implanted blastocyst until the pla-
into the pregnancy. centa forms. The secretory (luteal) phase begins after ovula-
The ovarian medulla (Figure 17-11a) consists of a tion. During this phase the endometrium is maintained by
loose, fibrous connective tissue with large blood vessels, luteinizing hormone from the corpus luteum. If no fertili-
lymphatics, and nerves. The main cells are fibroblasts and zation and implantation occur, progesterone levels decrease
the epithelioid hilus cells that produce androgens. as a result of the corpus luteum dying and the endometrium
Fallopian Tube is shed during menstruation.
The cyclic growth, death and repair of the endometrium
The fallopian (uterine) tubes are lateral extensions from the
have resulted in the recognition of three layers in it (Figure
superior portion of the uterus. There are four segments to
17-18). The stratum basalis is next to the myometrium and
each. These are the infundibulum, ampulla, isthmus, and
is relatively unchanged during the cycle. It serves as the
intramural region. The infundibulum is the funnel-shaped,
source of new endometrium after menstruation. The thick-
open end that receives the ovulated secondary oocyte from
est layer, characterized by a spongy looking stroma is the
the ovary. It has fingerlike projections called fimbriae ex-
stratum spongiosum. On the surface is the thin stratum
tending from its margin. The ampulla is the longest portion
compactum, characterized by a denser stroma. Together,
of the fallopian tubes and is the usual site of fertilization.
the stratum spongiosum and compactum undergo the most
Where the ampulla becomes more constricted, it forms the
change during the menstrual cycle and are collectively re-
isthmus, which leads to the intramural region within the
ferred to as the stratum functionalis.
uterine wall.
Straight arteries and spiral arteries supply the endo-
There are three layers in the fallopian tubes (Figure
metrium. The former provide blood to the stratum basalis,
7-16). These are the mucosa, muscularis, and serosa. The
whereas the latter supply the stratum functionalis. The spiral
mucosa forms longitudinal folds, which are branched in the
arteries constrict in the absence of progesterone, which
ampulla, and is lined with a simple columnar epithelium.
makes the stratum functionalis ischemic, resulting in its
Ciliated cells sweep the oocyte and fluid formed by noncili-
death.
ated secretory (peg) cells toward the uterus. The secretion
also nourishes the sperm and assists in their final matura- The myometrium (Figure 17-19) is composed of smooth
tion. Deep to the epithelium is an unremarkable lamina muscle in three poorly defined layers. There are inner and
propria made of a loose connective tissue. Movement of the outer longitudinal layers with a thick circular layer be-
oocyte is also assisted by peristaltic waves produced by an tween. The outer surface of the fundus and posterior body
inner circular and an outer longitudinal layer of smooth is covered with a serosa (parietal peritoneum). The remain-
muscle. The whole fallopian tube is covered with perito- der is covered with a fibrous adventitia.
neum, which forms the serosa. The cervix (Figure 17-20) is divided into an endocervi-
cal canal and an ectocervix, which protrudes into the
Uterus vagina. The endometrium of the endocervical canal is lined
The uterus is a pear-shaped organ, 7 cm by 4 cm, located in by a mucous secreting simple columnar epithelium, which
the pelvic cavity. There are three regions of the uterus. also lines depressions called endocervical glands. The ecto-
These are the body, fundus, and cervix. The uterine body is cervix resembles the vagina in that it is lined with a strati-
the major portion of the organ. The fallopian tubes enter fied squamous epithelium. The myometrium is mostly
201
replaced with elastic connective tissue as an adaptation to mixing of maternal and fetal blood—no blood cells cross the
the stretching necessary during childbirth. Lymphocytes placental barrier (see next column).
may be seen in this layer near the surface. The placenta is attached to the endometrium by a struc-
ture known as the decidua basalis, derived from the endo-
Vagina metrial stratum functionalis. Beginning with the third month,
The vagina (Figure 17-21) is the female copulatory organ chorionic villi anchor the developing embryo to the decidua
and serves as the birth canal. It is about 9 cm long and is basalis. In the mature placenta, the villi are branched and
lined with a mucosa composed of nonkeratinized stratified are not attached. Thus, free villi outnumber anchoring villi.
squamous epithelium and a dense, elastic lamina propria. Both types are covered with a cellular layer composed of
The mucosa folds when relaxed and the lumen is closed. syncytial trophoblasts, whose nuclei are seen to form clus-
There are no vaginal glands; cervical mucus is primarily re- ters called syncytial knots. The core of each villus contains
sponsible for its lubrication. Deep to the mucosa is a mus- fetal capillaries (and sometimes larger vessels), and embry-
cularis composed of an inner circular and an outer onic mesoderm. Open spaces in the decidua called lacunae
longitudinal layer of smooth muscle, though they are not or intervillous spaces surround the villi and contain mater-
well-defined and the fibers mix. Deep to the muscularis is a nal blood. Thus, the villi are bathed in maternal blood, but
fibrous adventitia, which blends in with the adventitial layers the syncytial trophoblasts and connective tissue form the
of the urinary bladder and rectum. placental barrier and prevent mixing with fetal blood.
Umbilical Cord
Breast
The umbilical cord (Figure 17-24) is derived from the early
The breasts contain mammary glands and are responsible connecting stalk that attaches the embryo to the tropho-
for producing milk (Figure 17-22). Mammary glands are blast layer. When fully formed, it is covered by amniotic
present in both sexes, but under the influence of estrogen, membrane and is filled with a ground substance called
progesterone and prolactin (from the anterior pituitary Wharton’s jelly, which contains mesenchymal cells. The
gland) in the female, the glands develop further. The 15 to two umbilical arteries, which carry oxygen-poor blood to
25 lobes in each breast are made of independent compound the chorionic villi, and the single umbilical vein, which
tubuloacinar glands. A lactiferous duct drains the gland to brings oxygen-rich blood to the fetus, are also present.
a lactiferous sinus, which leads to the
nipple. The glands are surrounded by
adipose tissue and are separated by
fibrous septa. The skin covering the
nipple has sebaceous glands not associ-
TA
ated with hair follicles.
During pregnancy, the mammary
glands branch and grow into alveoli,
BV
composed of a simple columnar ep-
ithelium and surrounded by myoep- IC
ithelial cells. Initially they produce a
viscous, protein-rich substance called
colostrum; within a few days after the ST
child is born, they produce true milk. BV
Placenta IC
The placenta (Figure 17-23) is a com-
plex organ that connects the embryo/
fetus with the mother’s uterus. It is
constructed from embryonic and ma-
ternal tissues. The fully formed pla-
centa serves as the fetus’ kidneys, lungs,
and intestines. That is, it absorbs oxy- 17-1
gen and nutrients from maternal blood,
TESTIS In this micrograph, the fibrous tunica albuginea (TA) of the testis and a portion of a
and transfers metabolic wastes into it. lobule are visible. Also seen are seminiferous tubules (ST), intestinal cells (IC), and a couple of
Under normal circumstances, there is no blood vessels (BV). (X65)
202
M
PS
M
Sd
TP
TP
Sz
A
Sd
B
PS A
B
SC
Sz
17-2a 17-2b
SEMINIFEROUS TUBULES Stages of spermatogenesis and spermiogenesis are visible in these micrographs. Type A spermatogonia (A) are identifiable
by their basal position in the tubule and a round nucleus with condensed chromatin and a nuclear vacuole. Type B spermatogonia (B) lack the
vacuole and have dispersed chromatin with a prominent central nucleolus. Primary spermatocytes (PS) are located closer to the lumen. They are
large cells with clumped chromatin. Secondary spermatocytes divide quickly after their production and are rarely seen in histological prepara-
tions. Spermatids (Sd) are small cells with round, dark nuclei. Spermatozoa (Sz) have pointed nuclei. Sertoli cells (SC) are located near the base
of the tubule and are characterized by an ovoid nucleus with a prominent nucleolus. In addition to cells of the spermatogenic series, note the
fibromuscular tunica propria (TP) surrounding this tubule. Several myoid cells (M) are visible. Micrographs (a) and (b) were magnified X400
and X265, respectively.
T M
H
17-3
SPERM CELLS The head (H), midpiece (M), and tail (T) are visible in these
sperm cells. Also visible in some is the acrosome (A). (X660)
203
IC
Sz
BV
ST
CT
PS
IC
17-4a 17-4b
INTERSTITIAL CELLS OF LEYDIG Interstitial cells (IC) are found in the vascular connective tissue (CT) located between seminiferous tubules (ST)
and constitute the main endocrine portion of the testis. (a) The interstitial cells are dark in this micrograph, but the nuclei with prominent
nucleolus are discernible. Note the blood vessels (BV) in the connective tissue. (X230) (b) In this micrograph, the interstitial cells and their lipid
vacuoles, characteristic of steroid secreting cells, are easily seen. Note also the spermatogenic cells and myoid cells of the seminiferous tubule.
Symbols used are the same as in Figure 17-2. (X580)
ST
CT
17-5
RETE TESTIS Seminiferous tubules empty into the rete testis, a network of tubules lined with
simple cuboidal epithelium (E) having microvilli. Each cell also has a single cilium. A vascular
connective tissue (CT) and myoid cells surround the tubules. A seminiferous tubule (ST) is also
seen. (X265)
204
E
S
E
LP
IL
17-7a OL
17-6a
E
M
E
M
S
LP
17-7b
17-6b
EPIDIDYMIS Pseudostratified epithelium (E) lines the ducts of the epididymis.
The tallest cells have long microvilli (stereocilia) (S), which aid in fluid ab-
sorption. The amount of smooth muscle (M) in the ducts’ walls increases
toward the ductus deferens. (a) X130 (b) X380
E
DUCTUS DEFERENS The longitudinally folded mucosa of
the ductus (vas) deferens consists of a pseudostratified
epithelium (E) and connective tissue lamina propria (LP).
The very muscular wall is composed of inner (IL) and LP
outer longitudinal (OL) smooth muscle layers, with a
circular (C) layer between. (a) X110 (b) X110 (c) X215 17-7c
205
S
G
17-8a
17-9a
CA
E
LP
G
L
17-8b
SEMINAL VESICLE Each seminal vesicle joins a ductus deferens and
empties its fructose-rich secretion into it. The mucosa is lined with a
pseudostratified epithelium (E). These cells have lipid droplets in them,
which often results in a foamy appearance. Lipofuchsin granules are 17-9b
also sometimes seen. The muscularis consists of an inner circular (C)
and an outer longitudinal (L) layer. (a) X25 (b) X110
SC
PROSTATE GLAND Shown here are three micrographs of CA

the prostate gland. Branched tubulo-acinar glands (G)
predominate and are lined with basal cells (B) and surface
secretory cells (SC). A fibromuscular stroma (S) fills the
space between glands. Corpora amylacea (CA), made of
glycoprotein, are found in the glands and increase with
age. (a) A portion of the capsule (C) is visible in this
micrograph. (X25) (b) The branching is apparent in this
gland. (X120) (c) Corpora amylacea become calcified
with age to form prostatic concretions. (X240) 17-9c
206
S
V
H
CC
TA
TA
CS
TA
U
17-10a 17-10b
TA
17-10c 17-10d
PENIS Externally, the penis is covered with skin (S) and a loose hypodermis (H). Internally, three cylinders of erectile tissue are present. These
are the two dorsal corpora cavernosa (CC) and the ventral corpus spongiosum (CS) around the urethra (U). Each is surrounded by a fibrous
connective tissue layer called the tunica albuginea (TA) and contains endothelium lined venous sinuses (V). (a) This micrograph is a transverse
section of half a penis. (X7) (b) The venous sinuses and the tunica albuginea of the corpus cavernosum are seen in this micrograph. (X25)
(c) The urethra is visible in the center of the corpus spongiosum. Note the tunica albuginea is thinner than in the corpora cavernosa. (X25)
(d) The spongy (penile) urethra is lined with a stratified or pseudostratified columnar epithelium (E). Mucous para-urethral glands (glands of
Littré) are also visible (G). (X55)
207
GE
TA
S
C
S
F
17-11a
S S
17-11b
THE OVARY The ovary is covered with a germinal epithelium (GE), which is actually peritoneum,
overlying a connective tissue tunica albuginea (TA). The bulk of the ovary is filled with connective
tissue stroma (S), which is divided into a cortex (C) and a medulla (M). The cortical region is
characterized by ovarian follicles (F), which are absent in the medulla. (a) This is a section of cat
ovary. (X65) (b) Shown in this micrograph is the ovarian cortex with follicles in various stages
of development. (X130)
208
GE
TE
TA
TI
PF GC
ZP
PO
S
CL
17-12a 17-12b
GC ZP
CO
A
GC
TI
TI
TE
17-12c 17-12d TE
MG
MG
PO CR
MG
TI TE
A CO
A
PO
17-12e 17-12f
OVARIAN FOLLICLE DEVELOPMENT For reference purposes, be advised that the ovum stays the same size in all stages of development. (a) A layer
of primordial follicles (PF) is found deep to the tunica albuginea (TA) and germinal epithelium (GE) of the ovary. A single layer of flat cells
surrounds each primordial follicle. A corpus luteum (CL) and the stroma (S) are also visible. (X220) (b) In this multilaminar primary follicle,
there are several layers of granulosa cells (GC) separated from the primary oocyte (PO) by the amorphous zona pellucida (ZP). A vascular and
cellular theca interna (TI) and the fibrous theca externa (TE) have begun to develop in the stroma. (X220) (c) The antrum (A) has begun to de-
velop in this secondary follicle and is filled with liquor folliculi, the diagnostic feature of a secondary follicle. Notice the more advanced state of
the theca interna (TI) and theca externa (TE). (X210) (d) In this secondary follicle, the cumulus oophorus (CO) is starting to develop. (X190) (e)
This micrograph shows a mature Graafian follicle. At this point, the primary oocyte is about to complete the first meiotic division. Note the well-
developed antrum and membrana granulosa (MG). (X50) (f) In this micrograph, the primary oocyte and cumulus oophorus are starting to detach
from the membrana granulosa in preparation for ovulation. The cells next to the primary oocyte form the corona radiata (CR). The theca interna
and theca externa are also visible. (X95)
209
CL
GL
CL
17-13a
17-13b
CORPUS LUTEUM The corpus luteum (CL) develops from the mem-
brana granulosa, which forms the granulosa-lutein (GL) cells, and
the theca interna, which contributes theca-lutein (TL) cells. Granulosa-
BV lutein cells are large and pale staining. The less abundant theca-lutein
cells are smaller and darker staining than the granulosa cells. They
TL tend to be located near the periphery of the corpus luteum. (a) In this
BV panoramic view, the large size of the corpus luteum is apparent. A
second corpus luteum is visible in the lower left of the field. (X20) (b)
The large granulosa-lutein cells are visible in this micrograph. (X210)
(c) Blood vessels (BV) and a few theca-lutein cells are visible in this
corpus luteum. (X210)
17-13c
GL
CA
TL
17-15
CORPUS LUTEUM OF PREGNANCY The corpus luteum of pregnancy
17-14
continues to grow and develop. It remains functional for the first
CORPUS ALBICANS After it has served its purpose, the corpus luteum is few months of pregnancy. In this micrograph, a portion of the corpus
invaded by fibroblasts, the luteal cells undergo autolysis, and a corpus luteum is shown. Granulosa-lutein cells (GL) and theca-lutein cells
albicans (CA) is formed. (X20) (TL) are visible. (X50)
210
MS
MS
MU
MU
17-16a
17-16b
MS
MU
V
LP
V
17-16c 17-16d
V LP
E
17-16e 17-16f
FALLOPIAN TUBES The Fallopian tubes are muscular structures extending laterally from the uterus. Their wall is made of a folded mucosa (MU),
a muscularis (MS) consisting of poorly defined inner circular and outer longitudinal smooth muscle layers, and a serosa made of parietal peritoneum.
Micrograph (a) is from the infundibulum, (b) is from the ampulla, and (c) is from the isthmus. These show that the mucosal folding is most intricate
in the ampulla. (a) and (b) are X20; (c) is X110. Micrographs (d) and (e) are of fimbriae, the fingerlike projections extending from the infundi-
bulum that assist in capturing the ovum at ovulation. The ciliated epithelium (E) sweeps into the Fallopian tube. Note the abundant, large veins
(V) in the lamina propria (LP). Micrograph (d) is X50; (e) is X230. Micrograph (f) illustrates the epithelium made of ciliated cells (C) and
nonciliated peg cells (P). (X400)
211
UTERUS This micrograph shows the
uterine wall. Uterine glands (UG)
penetrate the lamina propria (LP)
LP of the endometrium (E). The thick
myometrium (M) with its poorly
organized smooth muscle layers is
UG also seen. The perimetrium, made
of serous membrane, is not visible
E
in this preparation. (X25)
17-17 UG
SC
LP
SS
M
SF
17-18b
SB
LP
17-18a
E
ENDOMETRIUM The endometrium is lined by a simple colum-
nar epithelium (E) supported by a dense fibrous lamina propria
(LP). Two endometrial layers are recognized. These are the
basal stratum basalis (SB) and the more superficial stratum
functionalis (SF), itself made of the stratum compactum (SC)
UG LP
and the stratum spongiosum (SS). During the menstrual cycle,
the endometrium grows thicker, becomes more vascular, and
the glands become coiled. (a) This micrograph shows the
endometrium in the proliferative (follicular) phase of the
menstrual cycle. Note the few, straight glands (UG). (X20) (b)
The glands are more coiled in this micrograph of the endo-
metrium during the secretory (luteal) phase. (X25) (c) A coiled
gland is shown in this micrograph. Note the cellular lamina
propria and the dark staining lymphocytes (L). (X230) 17-18c
212
BV
BV
L C L
17-19
MYOMETRIUM The smooth muscle of the myometrium is arranged into poorly defined inner and
outer longitudinal layers (L), separated by a vascular middle circular layer (C). This micrograph
shows muscle fibers in both orientations as well as a few blood vessels (BV) and connective tissue
(CT). (X130)
SS
SC
EG
CT
17-20
CERVIX The endocervical canal is lined by a simple columnar epithelium (SC), which secretes
mucus, but it changes to a nonkeratinized stratified squamous epithelium (SS) where it projects
into the vagina as the ectocervix. Endocervical glands (EG) are also visible. Deep to the epithelium
is a dense elastic connective tissue layer (CT) with a few smooth muscle cells. (X130)
213
SS
SS
LP
LP
17-21a 17-21b
VAGINA The vagina consists of a mucosa, muscularis, and an adventitia. The mucosa is made of a nonkeratinized stratified squamous epithelium
(SS) and is supported by a thick, fibrous, and elastic lamina propria (LP). Longitudinal and circular smooth muscle fibers comprise the muscularis
(M), but the layers are not well-defined. An outer fibrous adventitia (A) blends with surrounding structures. (a) X25 (b) X130
CT
G
D
CT
G
17-22a 17-22b
BREAST The mammary glands are compound tubulo-acinar glands occupying the approximately 20 lobes in each breast. (a) This micrograph
is of inactive mammary glands (G). A duct (D), interlobar connective tissue (CT), and adipose tissue (A) are also visible. Note the absence of
secretion in the glands. (X110) (b) The branching glands (arrows) are apparent in this active mammary gland. Note the secretion in the glands
and the interlobar connective tissue. (X110)
214
T
V
IV
CT
V
IV
BV
IV
IV
17-23a 17-23b
PLACENTA (a) This micrograph is a panoramic
view of the placenta. Large and small chorionic
villi (V) and intervillous spaces (IV) are visible.
V Note the branching of the smallest villi. (X20)
V IV
(b) In this cross section of a villus, the trophoblast
layer (T), connective tissue (CT), and fetal blood
vessels (BV) are seen. Maternal blood is found in
SK the intervillous space (IV). (X555) (c) Clusters of
syncytial trophoblast nuclei form clusters called
syncytial knots (SK). (X245)
SK
17-23c
UMBILICAL CORD This umbilical W

cord has been cut in cross section.
The two umbilical arteries (A) and
the single umbilical vein (V) are
surrounded by mesenchymal tissue
called Wharton’s jelly (W). The A
outer surface is covered with
amniotic membrane (A). (X8) 17-24
Further Reading
Ballard, Marguerite. Atlas of Blood Cell Morphology. Centers for Disease Control.
Atlanta, Georgia, 1985.
Berman, Irwin. Color Atlas of Basic Histology, 3rd Ed. McGraw-Hill/Appleton &
Lange. Stamford, Connecticut, 2003.
Campbell, Neil A. and Jane B. Reece, Biology, 6th Ed. Benjamin Cummings. San
Francisco, CA, 2002.
Diggs, L.W., Dorothy Sturm, and Ann Bell. The Morphology of Human Blood Cells,
5th Ed. Abbott Laboratories. Abbott Park, Illinois, 1985.
Eroschenko, Victor P. DiFiore’s Atlas of Histology: With Functional Correlations,

9th Ed. Lippincott, Williams & Wilkins. Baltimore, Maryland, 2000.
Gartner, Leslie P. and James L. Hiatt. Color Textbook of Histology, 2nd Ed. W.B.
Saunders Company. Philadelphia, Pennsylvania, 2001.
Gartner, Leslie P. and James L. Hiatt. Color Atlas of Histology, 3rd Ed. Lippincott
Williams & Wilkins. Baltimore, Maryland, 2000.
Junqueira, L. Carlos, José Carneiro, and Robert O. Kelley. Basic Histology, 10th Ed.
McGraw-Hill/Appleton & Lange, 2002.
Kerr, Jeffrey B. Atlas of Functional Histology. Mosby. London, UK, 1998.
Ward’s Natural Science 2002 Biology Catalog. Table of microbiological stains.

Rochester, NY.
Young, Barbara and John W. Heath. Wheater’s Functional Histology A Text

and Colour Atlas, 4th Ed. Churchill Livingstone, Harcourt Publishers Limited.
Edinburgh, UK, 2000.
215

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A Photographic Atlas

Michael J. Leboffe, D.A.

Morton Publishing Company

Copyright © 2003 by Morton Publishing Company

All rights reserved. No part of this publication may be reproduced,

Printed in the United States of America

Chapter 1 Introduction 1 Epithelial Terminology 23

Chapter 6 Blood and Bone Marrow 61 Chapter 12 Cardiovascular System 133

Histological Techniques—Microscopy Strategies for Studying Histology

◗ TABLE 1-1 Commonly Used Histological Stains

AZURE B DNA is stained blue-green, RNA is stained purple.

DAFANO SILVER METHOD Golgi material stains black.

FEULGEN STAIN DNA is stained reddish violet.

OSMIUM Stains lipid a dark brown color.

Stage clip On/off switch

Coarse focus knob

Fine focus knob

LIGHT MICROSCOPE The light microscope uses Mechanical stage

LIGHT MICROGRAPH A photograph of a microscopic image is a

ELECTRON MICROSCOPE The

SECTIONS OF A TUBE Study this

T cell structure and ultrastructure before embarking

CELLS AS VIEWED WITH THE LIGHT

CELL MEMBRANE AS SEEN

EUCHROMATIN AND HETEROCHROMATIN In this

NUCLEAR ENVELOPE AND PORES In this transmission electron

RER AFFECTS THE APPEARANCE OF THE CYTOPLASM

MITOCHONDRIA Mitochondria (M) are the

2-34a 2-34b 2-34c

2-34d 2-34e 2-34f

MITOSIS IN LYMPHATIC TISSUE

AN EPITHELIAL MEMBRANE All epithelial membranes

Cilia on luminal surface

Basal layer Basement

Simple tubular Simple branched tubular Simple coiled tubular

SIMPLE TUBULAR GLANDS OF THE COLON

Functions of Fibrous Connective Tissues Extracellular Matrix of Adult Connective Tissues

connective tissue has an abundance of elastic fiber bundles

Reticular Connective Tissue

ELASTIC FIBERS IN ELASTIC TISSUE This specimen

LAMINA PROPRIA Loose

ELASTIC CARTILAGE The framework of the

SYNOVIAL JOINT Synovial joints are freely movable. The bones

6-5a 6-5b 6-6a 6-6b

6-7a 6-7b 6-7c

PLASMA CELLS B lymphocytes develop into plasma

6-9a 6-9b 6-10a 6-10b

6-15 6-16 6-17

6-19a 6-19b 6-19c

SMOOTH MUSCLE FIBERS Smooth muscle fibers are

Afferent (Sensory) System Efferent (Motor) System

Visceral Motor (Autonomic)

Visceral Somatic Somatic

Gray and White Matter Other Neural Structures

FIBROUS ASTROCYTES Perivascular feet of astrocytes

UNIPOLAR (PSEUDOUNIPOLAR) NEURONS

Schwann cell cytoplasm

Schwann cell nucleus Mesaxon Neurilemma Myelin

M MYELIN SHEATH (a) Myelin is produced by compressed

INTERNODES Two nodes of Ranvier are indicated