PDQ Histo

PDQ
*
SERIES
ACKERMANN
PDQPHYSIOLOGY
BAKER, MURRAY
PDQBIOCHEMISTRY
CORMACK
PDQHISTOLOGY
DAVIDSON
PDQMEDICAL GENETICS
JOHNSON
PDQPHARMACOLOGY, 2/e
KERN
PDQHEMATOLOGY
McKIBBON
PDQ EVIDENCE-BASED PRINCIPLES AND PRACTICE
NORMAN, STREINER
PDQ STATISTICS, 3/e
SCIUBBA, REGEZI, ROGERS
PDQ ORAL DISEASE: DIAGNOSIS AND TREATMENT
STREINER, NORMAN
PDQ EPIDEMIOLOGY, 2/e
*
PDQ (Pretty Darned Quick)
PDQ
HISTOLOGY
DAVID H. CORMACK, PHD
Professor Emeritus
Division of Anatomy, Department of Surgery
University of Toronto
Toronto, Ontario
with illustrations by
Monique Guilderson, BSc., MSc., BMC
2003
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Preface
It is my sincere hope that this short laboratory guide will turn
out to be a handy study aid for those histology students who, because of
inexperience, are nervous about becoming overwhelmed by too much his-
tologic detail. In keeping with the approach taken by the other books in the
PDQ series, it deals only with basics, which it presents in a reasonably
rational manner. Because PDQ Histology focuses the readers attention on
the main structural features found in standard sections taken from the stu-
dents slide box and emphasizes the particular combination of histologic cri-
teria that characterizes each section, it represents a minireview of the dis-
tinctive microscopic features that permit histologic preparations of the
various tissues and organs of the body to be correctly identied. Included
on the accompanying CD-ROM are representative selections of review
questions, sample multiple-choice questions, and temporarily unidentied
test sections available for private assessment of the readers comprehension
and prociency at appropriate stages in the course of study.
Along with a good slide box, microscope, labeled relevant photomi-
crographs, and the timely assistance of expert slide interpreters, histology
students generally appreciate being provided with a fairly nitty-gritty
approach to slide identification. PDQ Histology is designed to help begin-
ners to make a productive start and then to keep them firmly on the rails.
To advance, students should, of course, endeavor to supplement their
knowledge base with further relevant information that they are receiving
from lectures, other textbooks, atlases, and recommended additional
sources. Before long, diligent students will find that they are able to com-
pile other reliable recognition features of their own. By learning how to
rationalize test situation responses, they will soon be able to avoid flagrant
guesswork and with sufficient practice will attain self-confidence in micro-
scopic identification of tissues.
David H. Cormack
January 2003
Contents
Preface, v
1 Introduction: Cells and Tissues, 1
2 Epithelial Tissue, 19
3 Connective Tissue, 35
4 Cartilage and Bone, 45
5 Blood and Myeloid and Lymphoid
Tissues, 61
6 Nervous Tissue, 83
7 Muscle, 99
8 Circulatory System, 109
9 Integumentary System, 119
10 Digestive System, 129
11 Respiratory System, 145
Contents vii
12 Urinary System, 155
13 Endocrine System, 169
14 Female Reproductive System, 185
15 Male Reproductive System, 201
Index, 214
PDQ
Histology
Introduction: Cells and
Tissues
Afundamental concept of body structure is that its interior
represents an intricate assembly of distinct body tissues. The constituent
cells of these tissues are so frail that they generally require the support of
extracellular matrix (ECM), but certain tissues have a minimal amount of
this component and others have an ECM that is represented by a uid. The
least conspicuous tissue component at a microscopic level is a body uid,
predominantly interstitial (tissue) uid and, less commonly, blood plasma
or lymph. A profusion of functional cell products passes into these body u-
ids from the tissues, ranging from secreted matrix constituents and plasma
proteins to hormones and various other signaling molecules.
Histology is essentially the study of the body tissues. Although there
are only four basic tissues, some have variant and special subtypes, as
shown in Table 11. Organs and body parts are all constructed from these
four basic tissues and their subtypes. Histology also addresses the question
of how tissue cells carry out body processes at the cellular level. By describ-
ing major body functions in terms of the relevant tissues and their dis-
tinctive cell structure, it serves as a structural foundation for allied and
applied health sciences such as physiology, biochemistry, hematology, and
pathology.
1
1
MICROSCOPES
The chief tool at the histologists disposal is the light microscope (LM).
Providing two-staged magnication (Figure 11), the LM has as its basis (1)
an objective lens assembly and (2) an ocular (eyepiece) lens assembly. For
an eye unaided by any magnifying device, the minimal distance between
two adjacent points in a specimen that is necessary for these to be distin-
guishable as separate entities (ie, retinal resolution) is 0.2 mm. Magnica-
tions up to 1,000, obtainable with the LM, enable small details up to
0.2 m apart to be discerned, but because 0.2 m is the LMs absolute limit
of resolution, any further photographic enlargement exceeding 1,400 dis-
closes no further details.
Another limitation of the LM is that incident light must pass through
the specimen viewed. Tissue slices (sections) or suitably thin alternative
preparations are therefore necessary. Also, appropriate histologic stains
(dyes) are required for most tissue components to be seen clearly, so the tis-
sue is routinely xed (killed and preserved with 4% aqueous formaldehyde
or some other protein coagulant) and processed further to permit its sec-
tioning and enhance its staining. The necessary harsh treatment can cause
unintentional distortion that may complicate the interpretation of LM
appearance. An atypical histologic appearance may indicate inadequate x-
ation or excessive shrinkage during tissue processing.
Further information about the LM and some hints on how to get it to
perform properly may be found in the accompanying CD-ROM.
2 PDQ HISTOLOGY
Table 11
Four Basic Tissues with Their Subtypes
Epithelial tissue Nervous tissue
Epithelial membranes
Epithelial glands Muscle tissue
Skeletal muscle
Connective tissue Cardiac muscle
Loose (areolar) tissue Smooth muscle
Dense ordinary (brous) tissue
Adipose (fat)
Cartilage
Bone
Blood cells
Myeloid tissue
Lymphoid tissue
When higher magnications and better resolution are required, an elec-
tron microscope (EM) is used instead. The transmission electron micro-
scope (TEM) operates in a manner similar to the LM but uses a beam of
electrons instead of incident light. The scanning electron microscope
(SEM) is somewhat different in design. When its narrow electron beam
tracks back and forth across a metallic or metal-coated surface, electrons
secondarily emitted or reected from the metal are collected. The resulting
signal is processed to produce a three-dimensional-looking television image
(scan) of the scanned surface.
The usual kind of electron micrograph found in histology texts and
atlases (see Figure 15) is obtained with a TEM, which can provide magni-
cations up to 50,000 and resolve details that are 1 nm apart. This greater
resolution is achieved by using an electron beam instead of light. Resolution
is proportional to wavelength, and the wavelength of an electron beam is
over 1,000 times shorter than that of light. Resolution in scanning electron
micrographs, which are encountered less commonly in histology courses, is
generally limited to about 15 nm by the coating thickness, but SEM resolu-
tions of 3 nm are technically possible.
Chapter 1 Introduction: Cells and Tissues 3
Image on retina
Intermediate image
in optical path
Section on stage
Eyepiece lenses
Objective lenses
Condenser lenses
Lamp
Figure 11 Optical path of the light microscope.
When EM sections of a tissue are viewed with the TEM following xa-
tion with glutaraldehyde and appropriate staining with heavy metals, elec-
trons pass unimpeded through areas where tissue components remain
unstained, whereas stained components scatter electrons out of the beam.
Through photographic reversal of the image on the viewing screen, electron-
dense components of the tissue accordingly appear black and electron-lucent
components appear white in the reversed prints (electron micrographs).
Although three-dimensional imaging of biologic surfaces is possible
with the SEM, routine photomicrographs and micrographs show tissues and
organs in section, so it is important to try to visualize the three-dimensional
structure of a living tissue or organ before it was xed, distorted by tissue
preparation, sectioned, and stained.
INTERPRETING TISSUE SECTIONS
When confronted with histologic sections, beginning students often ask what
they are supposed to see. A generalized answer is either individual cells or
aggregates of cells forming some characteristic arrangement with the ECM.
Indeed, many organs and tissues are recognizable histologically by their dis-
tinctive arrangement of cells and ECM. Some detailed mental reconstruction
is generally necessary before histologic sections can be visualized in three
dimensions. An initial general impression to seek is whether the composition
of the tissue or organ is predominantly cellular (Figure 12) or predomi-
nantly ECM (Figure 13). Another useful clue is whether the structure seems
solid, perforated with spaces, or hollow. Preliminary observation under scan-
ning power (30) or low power (100) generally provides helpful informa-
tion before details are sought. Lower magnications are often more inform-
ative than high ones, particularly for organ identication.
4 PDQ HISTOLOGY
Figure 13 Elastic cartilage tissue made up chiey of extracellular matrix.
Extracellular
matrix
Cells
Figure 12 Liver tissue made up chiey of cells.
CELLS
The appearance of cells in LM sections of liver may be seen in Figure 14. The
central spherical nucleus houses the cells complement of chromosomes that
contain gene-bearing deoxyribonucleic acid (DNA). The darkly stained struc-
ture toward the nuclear center, the nucleolus, is the site of production of ribo-
somal RNA (rRNA). The other darkly stained material in the nucleus, hete-
rochromatin (condensed chromatin), represents genetically inactive DNA that
is not being transcribed, that is, copied from DNA to RNA for protein synthe-
sis. The outer part of the cell, its cytoplasm, derives energy through cell metab-
olism and synthesizes proteins. Few details of its internal structure are seen in
routine histologic sections because the cytoplasmic organelles are so small, so
the EM, with its higher resolution, is generally employed instead. The perime-
ter of each cell is sometimes difficult to discern in LM sections (see Figure 14),
generally because it lies at the wrong angle for good visibility. The cell mem-
brane itself is not thick enough to be resolved in the LM.
6 PDQ HISTOLOGY
Figure 14 Light microscopic appearance of hepatocytes.
Nucleus
Nucleolus
Cell boundary
Cytoplasm
Figure 15 shows an example of cell and ECM appearance at the EM
level. The chief cytoplasmic organelles and their functions are summarized
in Table 12. For further information, see Web site links in the accompanying
CD-ROM.
Figure 15 Electron microscopic appearance of chondrocytes and their associated extracel-
lular matrix.
Lysosome Extracellular
matrix
Rough-surfaced
endoplasmic
reticulum
Heterochromatin Cell membrane
Nuclear envelope
8 PDQ HISTOLOGY
Table 12
Main Components of Cytoplasm
Component Structure Chief Functions
Cell membrane Fluid lipid bilayer with Regulates molecular exchanges
integral and peripheral with tissue uid, responds to
proteins and cell coat signals
on external surface
Cytosol Semiuid with minimal Contains soluble molecules and
internal support cytoplasmic organelles and
inclusions
Smooth-surfaced Branched tubular Synthesizes lipids and steroids,
endoplasmic system of smooth segregates calcium in muscle
reticulum intracellular membrane cells, metabolizes drugs and
glycogen in hepatocytes
Rough-surfaced Parallel at membranous Incorporates integral membrane
endoplasmic cisternae with bound proteins, segregates
reticulum ribosomes secretory and lysosomal
proteins in its lumen
Golgi apparatus Saucer-shaped stack Completes glycosylation of
of at smooth glycoproteins, modies and
membranous sorts secretory products,
saccules distributes secretory proteins
to secretory vesicles, sends
acid hydrolases to lysosomes
Secretory vesicles Spherical smooth Convey secretory proteins from
membranous saccules Golgi apparatus to cell
containing secretory surface
proteins
Lysosomes Small spherical smooth Segregate destructive enzymes
membranous saccules for general use in
containing acid intracellular digestion
hydrolases
Mitochondria Outer and inner limiting Produce ATP through oxidative
membranes, cristae, phosphorylation
internal mitochondrial
matrix
Ribosomes Electron-dense Provide sites where amino acids
ribonucleoprotein are assembled in the order
particles present as specied in mRNA sequences
subunits
Microtubules Unbranched tubular Provide support, guide
assemblies of tubulin intracellular transport, provide
subunits, 25 nm in ciliary and agellar motility
diameter, single and
assembled into
centrioles, cilia, and
agella
Continued
STAINS
A stain combination routinely used for histologic sections is hematoxylin and
eosin (H&E). Unless otherwise stated, the photomicrographs in this book
show sections that have been stained in this manner. Hematoxylin, a tree-
derived dye combined with Al
3+
ions, colors negatively charged components
blue or purple. Eosin is a negatively charged stain that colors positively charged
components pink or red. Basophilic components stain with hematoxylin (a
basic stain); acidophilic components stain with eosin (an acid stain). When a
blood stain is used, basophilic and acidophilic constituents stain equivalent
blue and pink colors. Periodic acidSchiff (PAS) staining generates aldehyde
groups in glycoproteins and polysaccharides and discloses these as magenta or
purple dye complexes.
Tissue components staining blue in H&E sections
contain macromolecules having a net negative charge, for example,
DNA, RNA (nucleic acids are rich in PO
4
3
groups);
have an affinity for hematoxylin because of its associated Al
3+
ions;
include nuclear chromatin, nucleoli, and cytoplasmic ribosomes.
Tissue components staining pink in H&E sections
contain macromolecules having a net positive charge in the staining pH
range of 5 to 6, for example, xed proteins with amino groups ionized
to NH
3
+
,
take up eosin because it has a negative charge, and
are in most cases proteins.
Tissue components staining magenta in PAS sections
may be secretory glycoproteins;
may be glycogen, a stored polysaccharide.
After H&E staining, why does the cell cytoplasm appear essentially pink
in Figure 16 and comparatively blue in Figure 17 and have areas of each
Table 12 (Continued)
Main Components of Cytoplasm
Component Structure Chief Functions
Microlaments Actin-containing 7 nm rods Interact with thick laments
(equivalent to in contractions, support
thin laments) microvilli
Thick laments Myosin-containing 1216 nm Interact with microlaments or
rods thin laments in contractions
Intermediate Heterogeneous class of Provide support, distribute
laments 10 nm rods stresses
ATP = adenosine triphosphate; mRNA = messenger ribonucleic acid.
color in Figure 18? What does this staining reveal about the particular
functions of each cell type?
The cytoplasm of muscle bers (see Figure 16)
has a high concentration of structural proteins (actin, myosin, etc) that
are moderately acidophilic, and
10 PDQ HISTOLOGY
Figure 17 Blue-stained cytoplasm in osteoblasts.
Osteoblasts
Figure 16 Pink-stained cytoplasm in skeletal muscle bers.
is functionally specialized for contraction.
The cytoplasm of osteoblasts (see Figure 17)
contains abundant ribosomes, and therefore basophilic RNA, bound to
its rough-surfaced endoplasmic reticulum (rER);
is functionally specialized for secretion (bone matrix constituents).
The cytoplasm of pancreatic acinar cells (see Figure 18)
contains abundant ribosomes, and therefore basophilic RNA, bound to
its rER;
contains abundant secretory vesicles with acidophilic secretory protein
content;
is functionally specialized for the synthesis and release of stored aci-
dophilic secretory proteins (digestive enzymes).
Alternative staining can provide supplementary information. For exam-
ple, the surface epithelium stained with H&E in Figure 19 exhibits round,
virtually colorless gaps containing some constituent with negligible affinity
for either stain. A PAS-stained section (Figure 110) discloses magenta-
colored mucus at these sites, indicating that the white gaps are goblet cells
Figure 18 Pink- and blue-stained cytoplasmic components in pancreatic acinar cells.
12 PDQ HISTOLOGY
Figure 19 Goblet cells in a hematoxylin and eosinstained section of the small intestine.
Figure 110 Goblet cells in a periodic acidSchiffstained section of the small intestine.
(glycoprotein-secreting surface epithelial cells). Special staining methods
are occasionally preferable, notably silver impregnation for nervous tissue
and immunohistochemical staining for specic proteins.
NUCLEAR APPEARANCE
Functional activity may be reected in the appearance of the nucleus as well
as the cytoplasm. Cells that are actively synthesizing proteins, for example,
often exhibit a large nucleolus (see Figure 14) because they require an
abundance of the rRNA exclusively transcribed there.
It is equally important to be able to recognize mitotic (dividing) cells
because the proportion and LM appearance of dividing cells can have diag-
nostic value. Mitotic gures may be spotted under low magnication (Fig-
ure 111). At higher magnification, the individual rod-shaped chromo-
somes may be discernible (Figure 112).
Figure 111 Mitotic gures present in crypt bases of the small intestine.
Mitotic gures
14 PDQ HISTOLOGY
Figure 112 Mitotic gure seen in more detail (crypt of small intestine).
Anaphase
The process of mitosis is summarized in Figure 113.
Prophase
Polar microtubules form
between centriole pairs
Chromosomes
become thread-like
Nuclear envelope
starts fragmenting
Kinetochore microtubules
form in mitotic spindle
Chromosomes (two
chromatids in each)
align in equatorial plane
Metaphase
Kinetochore microtubules
shorten
Spindle elongates due to
sliding between opposed
polar microtubules
Chromatids separate,
move toward poles
Anaphase
Nuclear envelope re-forms
Daughter chromosomes
(formerly the chromatids)
segregate
Equatorial region
constricts
Residual microtubules
form midbody
Telophase
Figure 113 The stages of mitosis.
Appropriate preparation and special staining of isolated mitotic cells
allow specific identification of the chromosomes they contain (Figure
114). An assembled chromosome map obtained from a representative cell
is called a persons karyotype.
Another important distinction that needs to be made is between mitotic
cells and dying cells. Sometimes both are present in the same eld of view.
All stages of mitosis except early prophase are recognizable by the presence
of dark-staining chromosomes and the absence of an intact nuclear enve-
lope (Figure 115). Dying or dead cells, on the other hand, exhibit terminal
disintegration of their nucleus (karyorrhexis), extreme condensation and
heavy (heterochromatic) staining of its chromatin (pkynosis), or unusually
pale heterochromatin staining (karolysis). Examples of these changes are
shown in Figure 116.
16 PDQ HISTOLOGY
Figure 114 Chromosomes isolated from a mitotic cell and stained to show their chromosome
bands.
Figure 116 Nuclear signs of cell death.
Pyknosis
Karyolysis
Karyorrhexis
Figure 115 Mitotic gures in dividing hepatoma cells.
Anaphase
Metaphase
Finally, a few cell types, for example, osteoclasts (Figure 117) and
skeletal muscle bers (see Figure 16), possess more than one nucleus. In
these cases, the multinucleate condition is a consequence of cell fusion.
Human erythrocytes (red blood cells) possess no nucleus because it is dis-
carded at a late precursor stage.
18 PDQ HISTOLOGY
Figure 117 Multiple nuclei in osteoclasts.
Osteoclasts
Epithelial Tissue
Body cavities are lined with extensive cellular sheets known as
epithelial membranes (epithelia). The body also has a protective epithelial
covering termed the epidermis. Epithelia are invaginated at certain sites,
extending into adjacent connective tissue as epithelial glands. Epithelial tis-
sue accordingly consists of epithelial membranes and glands.
EPITHELIAL MEMBRANES
The constituent cells of epithelial membranes are clearly polarized in struc-
ture. Their free (luminal or apical) surface borders on the lumen of a body
compartment or the exterior of the body. A well-defined extracellular
matrix layer known as a basement membrane (basal lamina) attaches their
basal surface to adjacent connective tissue. This special matrix layer is
described in Chapter 3, Connective Tissue. The diverse functional roles of
epithelia include protection, absorption, secretion, transcellular transport,
restriction of diffusion, and provision of sensory function. Progressive cell
renewal through mitosis in basal layers of epithelia is their chief means of
maintaining structural integrity. Because epithelial tissue is entirely avas-
cular, its cells typically obtain oxygen and nutrients by diffusion from cap-
illaries of adjacent loose connective tissue.
Subtypes
Epithelial membranes are characterized by the number of constituent cell
layers and cell shape at their free surface (Table 21).
Typical examples of simple epithelia as they appear in hematoxylin and
eosin (H&E) sections are shown in Figures 21, 22, and 23.
2
19
20 PDQ HISTOLOGY
Table 21
Epithelial Membranes
Simplea monolayer of cells
Squamousat shape
Cuboidalshort, roughly square in sections
Columnartall and narrow
Pseudostratiedall cells are attached to the basement membrane but some do not
reach the free surface, giving a false impression of stratication
Columnar
Ciliated columnarcommonly with goblet cells
Stratiedmore than one layer of cells
Squamouscuboidal basal cells, at surface cells
Keratinizedat keratinized cells on free surface
Nonkeratinizedat nonkeratinized cells on free surface
Cuboidalusually 2 layers, lines small ducts
Columnarusually 2 layers, lines larger ducts
Transitionalbulging surface cells become at if epithelium is stretched
Figure 21 Simple squamous epithelium.
Simple squamous
epithelium
Figure 22 Simple cuboidal epithelium.
Simple cuboidal epithelium
The pseudostratied epithelium lining the main airways (Figure 24)
possesses goblet cells and cilia that act together to trap and clear away any
inhaled suspended particles that settle. Cilia are hair-like motile cytoplas-
mic processes that project from the luminal surface of the epithelium. Using
the energy that they release from adenosine triphosphate, they continu-
ously sweep a thin coating of mucus across the luminal surface. Cilia are
characterized by nine peripheral pairs of microtubules (doublets that can
actively slide past each other) and two central single microtubules (Figure
25). The mucus coating that is swept along by the cilia is produced by gob-
let cells (Figure 26) and underlying mixed (seromucous) glands.
Chapter 2 Epithelial Tissue 21
Figure 23 Simple columnar epithelium.
Simple columnar epithelium
22 PDQ HISTOLOGY
Figure 24 Ciliated pseudostratied columnar epithelium with goblet cells.
Cilia
Goblet cell
Figure 25 Electron micrograph of cilia.
The chief stratied epithelia are shown in Figures 27, 28, and 29.
Cells approaching the surface of stratied squamous keratinizing epithe-
lium become a layer of evaporation-resistant soft keratin termed the stra-
tum corneum(Figure 27), whereas those of stratied squamous nonkera-
tinizing epithelium (Figure 28) remain nonkeratinized but still fairly
resistant to abrasion.
Figure 26 Electron micrograph of a goblet cell.
24 PDQ HISTOLOGY
Figure 27 Stratied squamous keratinizing epithelium.
Stratum corneum
Basal layer of epithelium
Figure 28 Stratied squamous nonkeratinizing epithelium.
Transitional epithelium (see Figure 29), conned to the urinary tract,
is unique in that, for unknown reasons, some of the large football-shaped
cells on its luminal border are multinucleated and a few are polyploid, that
is, they contain more than one diploid set of chromosomes.
Cell Junctions
Specialized sites known as cell junctions lie on the lateral and basal parts of
the epithelial cell membrane (Figure 210). Table 22 details their various
electron microscopic appearances and functionsessentially, cell-to-cell
sealing, cell attachment, and direct cell-to-cell communication.
Figure 29 Transitional epithelium.
Multinucleated cell
26 PDQ HISTOLOGY
Continuous tight
junction
Adhesion belt
Desmosome
Gap junction
Hemidesmosome
Basement
membrane
Desmosome Gap
Continuous tight
Adhesion belt
Figure 210 Epithelial cell junctions. A, Belt-shaped junctions on lateral surfaces. B, Circular
junctions on lateral and basal surfaces. C, Detailed structure of cell junctions.
A
B
C
Table 22
Principal Cell Junctions
Junction Structural Features Function
Continuous tight Circumferential band where By obstructing lateral
junction (zonula aligned integral membrane intercellular spaces,
occludens) proteins interlock and form these ridges limit
anastomosing ridges paracellular diffusion at
lateral cell borders
Adhesion belt Circumferential band where This junction maintains
(zonula adherens) linker glycoproteins extend strong cell-to-cell
into the intercellular gap, adherence at the site
strongly bonding contiguous where the attached
cell membranes; anchors marginal band
the marginal band of contracts
microlaments
Desmosome Circular area where strongly Tension in the attached
bonding linker glycoproteins intermediate laments
form an electron-dense line is resisted by the
along midline of the gap; rivet-like site of strong
electron-dense plaques cell-to-cell adherence
along desmosomes lateral
borders anchor intermediate
(keratin) laments
Hemidesmosome Circular area on basal surface, Tension in the attached
resembling half a desmosome, intermediate laments
where cell membrane, plaque, is resisted by the
and anchored intermediate rivet-like site of strong
laments lie apposed to anchorage to the ECM
basement membrane
Gap junction Circular area containing tiny By providing a route for
spool-like assemblies small molecules and
(connexons) of aligned ions to diffuse directly
interlocking integral from one cell to another,
membrane proteins; these channels permit
connexons bridge the metabolic coupling and
intercellular gap and in direct cell-to-cell
their open conguration communication
constitute walls of aqueous
channels
ECM = extracellular matrix.
EPITHELIAL GLANDS
Glands that release their secretion through a duct are classied as exocrine;
those that deliver it directly to the bloodstream are termed endocrine (see
Chapter 13, Endocrine System). The epithelial components of a gland (its
secretory units and ducts) are called its parenchyma; the connective tissue
components (loose connective tissue and brous capsule) are known as its
stroma. Exocrine gland classication is broadly based on the general struc-
ture and function of glands (Table 23).
28 PDQ HISTOLOGY
Table 23
Exocrine Epithelial Glands
Structural Classication
Simpleunbranched duct system
Tubular secretory unit is tubular and in some cases coiled
Acinar/alveolarsecretory unit is spherical/ask-shaped
Compoundbranched duct system
Tubular
Acinar/alveolar
Functional Classication
Serousproduces a watery secretion that generally contains some protein
Mucousproduces mucus
Seromucous (mixed)possesses separate serous and mucus-secreting cells
Othereg, sebaceous glands producing an oily secretion (sebum) from
disintegrated lipid-laden cells
In simple glands, the secretory portion is generally distinguishable from
the duct. In the example shown in Figure 211, a coiled simple tubular
sweat gland,
the pale-staining, low columnar epithelial cells are serous secretory cells,
the at nuclei closely apposed to their basal border are those of con-
tractile myoepithelial cells, and
the double layer of cuboidal epithelial cells with dark-staining nuclei are
duct cells.
Figure 211 Simple tubular gland (eccrine sweat gland).
Duct
Secretory portion
In compound glands (Figure 212), stromal elements such as intralobu-
lar loose connective tissue, brous interlobular septa, and the brous capsule
are also recognizable. Intralobular ducts (see Figure 214) are the narrow
duct branches lying within lobules. Interlobular ducts are the slightly wider
branches extending along brous septa that delineate lobules. These ducts
collect the secretion from intralobular ducts. Hence, the parenchymal organ-
ization of a compound gland may be likened to a dormant bush-like tree:
rounded buds = secretory units
small twigs = intralobular ducts
branches = interlobular ducts
trunk = main duct
Building on this analogy, a heavy fall of snow covering the tree and ll-
ing all its crevices = stromal connective tissue.
30 PDQ HISTOLOGY
Intralobular loose
connective tissue
Secretory unit
Fibrous interlobular
septum
Intralobular duct
Fibrous capsule
Interlobular duct
Figure 212 Organization of a compound gland.
Serous secretory units (Figures 213 and 214) are readily distinguish-
able from mucous secretory units (Figures 213 and 215) by the following
criteria:
Serous cells
large spherical nucleus
basophilic cytoplasm at base
acidophilic secretory granules
Mucus-secreting cells
small at dark-staining nucleus at base
essentially colorless in H&E sections because of their stored mucus
Figure 213 Mixed (seromucous) gland showing mucous and serous secretory units.
Mucous secretory units
Serous secretory units
Both types of secretory unit may be found in the same gland (see Figures
213 and 215). The serous and mucous secretory cells of these mixed (sero-
mucous) glands can lie in separate lobules (see Figure 213), a common
lobule (see Figure 215), or a common mixed secretory unit (Figure 216).
32 PDQ HISTOLOGY
Figure 214 Serous gland showing serous secretory units and duct.
Serous secretory units
Intralobular duct
Figure 215 Mixed (seromucous) gland showing mucous, serous, and mixed secretory units.
Serous secretory
unit
Mucous secretory unit
Serous cell in mixed
secretory unit
The serous and mucous cells of a mixed secretory unit release their secre-
tions independently into the same central lumen. Arranged as a cap on one
side of the mucous unit, the serous cells in such a unit appear in sections as
a serous demilune. Whether this is apparent as a crescent-shaped area
depends on the plane of section.
Figure 216 Mixed (seromucous) gland showing serous demilunes.
Intralobular duct Serous demilune
Connective Tissue
The three subtypes of connective tissue being considered in
this chapter are loose (areolar), dense ordinary (brous), and body fat (adi-
pose). In contrast to the extracellular matrix (ECM) that represents the
principal constituent of the rst two subtypes, the functionally important
constituent of adipose tissue is stored lipid.
INTERSTITIAL MATRIX
The ECM complex of connective tissue, known as the interstitial matrix, con-
tains collagen bers, elastic bers, and an amorphous component known as
ground substance. Hard to distinguish in hematoxylin and eosin (H&E)-
stained sections, these intercellular bers are recognizable in specially stained
tissue spreads (Figure 31). Representing strong bundles of nonbranching
collagen brils, collagen bers can be fairly acidophilic, for example, at sites
where they are abundant. They provide substantial tensile strength. Branched,
comparatively narrow, straight elastic bers provide elasticity. Incorporating
the protein elastin within a framework of unbranched microbrils made of
glycoproteins called brillins, elastic bers have little affinity for eosin and
require special stains (eg, orcein) for their presence to be evident. The nest
bundles of collagen brils (found, for example, in the liver and myeloid and
lymphoid tissues) branch owing to longitudinal bundle splitting. Demon-
strable with silver stains, these bundles are called reticular bers because they
constitute an intimate supporting network for cells. Whereas the coarse col-
lagen bers of connective tissue are assemblies of type I collagen, represent-
ing ~ 90% of the total collagen, reticular bers are assemblies of type III col-
lagen. Both collagen and elastic bers become assembled interstitially from
their respective secreted precursor proteins, procollagen and tropoelastin.
The ground substance of connective tissue, essentially colorless in H&E
sections, is a hydrated gel containing proteoglycans, the glycosaminoglycan
hyaluronic acid, and glycoproteins. This gel plays a key role in nutrient and
waste diffusion because it sequesters tissue uid in its aqueous channels.
3
35
LOOSE CONNECTIVE TISSUE
In suitably stained loose connective tissue prepared as a at spread (see Fig-
ure 31), it is possible to distinguish
capillaries (because it is a highly vascular tissue),
nuclei of the broblasts that secreted ECM macromolecules,
conspicuous mast cells full of histamine-containing secretory granules
wavy wide collagen bers made up of brils, and
straight narrow elastic bers.
It is important to realize that in such preparations, the entire thickness of
cells and bers is seen.
36 PDQ HISTOLOGY
Figure 31 Loose connective tissue spread (special stains).
Capillary Fibroblast nucleus
Mast cell
Elastic ber Collagen ber
The H&E-stained loose connective tissue shown adjacent to stratied
epithelium in Figure 32 looks dissimilar from that in Figure 31 because
it has been sectioned. This is how it is usually seen. Its recognizable charac-
teristics include the following:
abundant pink collagen bers
numerous capillaries extending into projecting papillae that supply the
thick epithelium with nutrients
Chapter 3 Connective Tissue 37
Figure 32 Loose connective tissue in section (lamina propria of the esophagus).
Loose connective tissue
Epithelium
Along the interface between loose connective tissue and other tissues
such as epithelium, the ECM is organized into a thin sheet of binding
matrix called a basement membrane. At this site, laminin, an adhesive gly-
coprotein with a binding receptor in the cell membrane, is intimately asso-
ciated with an interstitially assembled meshwork of type IV collagen. Other
interstitial glycoproteins and proteoglycans participate in binding the adja-
cent cells to the interstitial matrix. Basement membrane sites are periodic
acidSchiff positive because of this high glycoprotein content (Figure 33).
At the electron microscopic level, a cross-section of the basement mem-
brane appears as a thin fuzzy extracellular line of low electron density
(lamina densa) situated parallel and adjacent to the basal surface of the
attached cells.
CONNECTIVE TISSUE CELLS
Table 31 lists the main characteristics of the cell types found in loose con-
nective tissue. Because this tissue is involved in certain types of immune and
inammatory responses, lymphocytes and neutrophils may also be encoun-
tered (see Chapter 5, Blood and Myeloid and Lymphoid Tissues).
38 PDQ HISTOLOGY
Figure 33 Basement membranes of renal tubules.
Basement membrane
Table 31
Connective Tissue Cells
Cell Type Structural Features Chief Functions
Fibroblast Spindle-shaped, basophilic Secretes brous and
cytoplasm in secretory state amorphous interstitial
matrix constituents
Plasma cell Round with basophilic cytoplasm, Secretes immunoglobulin
pale Golgi region, eccentric
nucleus with condensed
peripheral chromatin
Macrophage Ovoid, eccentric kidney-shaped Phagocytoses
nucleus sometimes hidden by microorganisms, foreign
phagocytosed particles or particles, and cell debris;
hemosiderin (hemoglobin processes antigens;
breakdown pigment) secretes complement
proteins and growth
factors
Mast cell Ovoid, central ovoid nucleus Releases histamine and
sometimes hidden by granule other inammatory
content, large metachromatic mediators when triggered
secretory granules by local trauma or antigen
cross-linking of surface-
bound immunoglobulin E
Endothelial cell Squamous epithelial lining cell Regulates production of
of blood vessels and lymphatics tissue uid and uid
exudate, regulates blood-
tissue molecular
exchanges, releases
vasoactive mediators
(nitric oxide radical and
endothelin 1)
Pericyte Irregularly shaped with long Incompletely differentiated
cytoplasmic processes cell persisting as a
wrapped around capillaries potential source of new
and small venules broblasts and smooth
muscle cells
Adipocyte Large and round, with small Stores and mobilizes lipid
(fat cell) at nucleus at periphery of (long-term energy reserve)
large central lipid droplet
(which is extracted in tissue
processing)
Fibroblasts are most readily recognizable at connective tissue sites where
they are actively secreting ECM macromolecules (Figure 34). Plasma cells,
derived from antigen-activated B lymphocytes, are recognizable by their
round shape, characteristic eccentric clock-face nucleus, cytoplasmic
basophilia, and pale negative Golgi region (Figures 34 and 35).
40 PDQ HISTOLOGY
Figure 35 Plasma cells seen in more detail.
Clock-face nucleus
Golgi region
Figure 34 Fibroblasts and plasma cells in loose connective tissue.
Plasma cells
Fibroblasts
Macrophages are readily identiable if they contain phagocytosed par-
ticles, for example, a marker dye (Figure 36) or inhaled smoke-derived car-
bon particles. They are further characterized by having a kidney-shaped
nucleus.
DENSE ORDINARY CONNECTIVE TISSUE
Compared with loose connective tissue, dense ordinary (brous) connective
tissue is substantially less vascular and its collagen bundles are much coarser
(Figures 37 and 38). Scattered between the type I collagen bundles are
nuclei of inactive broblasts (brocytes). Because the form of dense ordinary
(brous) connective tissue shown here is an interweaving arrangement of
wavy collagen bundles extending in almost every direction, it is described as
irregular. Tendons and ligaments are characterized by the regular form of
this tissue, where orderly arrays of essentially parallel collagen bundles are
oriented like cable wires along the direction of tension.
Figure 36 Macrophages in loose connective tissue.
Macrophages
42 PDQ HISTOLOGY
Figure 37 Irregular dense ordinary connective tissue (reticular layer of dermis of thin skin).
Loose ordinary connective tissue
Dense ordinary connective tissue
Figure 38 Collagen bundles of irregular dense ordinary connective tissue.
ADIPOSE TISSUE
In loose connective tissue, adipocytes (fat cells) may be found singly or in
small groups. In adipose (fat) tissue, adipocytes represent the predominant
cell type (Figure 39). The white spaces created by lipid extraction give this
tissue a distinctive chicken wire appearance. Besides storing lipid for
potential energy production, adipose tissue pads parts of the body and lls
some crevices. The widely distributed kind of adipose tissue shown in Fig-
ure 39, called white fat, is the principal site of lipid metabolism. The less
extensive kind, called brown fat because it contains an even greater abun-
dance of mitochondrial cytochromes and blood capillaries, stores lipid as
multiple droplets and provides essential body heat, which is important in
newborn babies.
Figure 39 Adipocytes of white fat.
Cartilage and Bone
Each of the structurally specialized skeletal tissues cartilage and
bone is derived from mesenchyme, as are the other forms of connective tis-
sue. Although both tissues are made up of extracellular matrix and cells,
cartilage is totally avascular, whereas bone tissue is highly vascular. Another
key difference is that cartilage can grow from within (interstitial growth)
through internal production of more matrix as well as grow externally by
adding more matrix to its outer surface (appositional growth), whereas
bone grows entirely through apposition to its surface. Functionally relevant
differences also exist in the composition of the matrix.
CARTILAGE
Essentially, hyaline cartilage (Figure 41) is designed to resist compression.
Approximately 75% of its wet weight is tissue uid that provides a long-
distance diffusion route for nutrients and oxygen to reach the chondrocytes
embedded in its avascular matrix. Other functionally important con-
stituents of hyaline cartilage matrix are proteoglycan (chiefly present as
aggregates) and reinforcing brils composed of type II collagen. Chondro-
cytes occupy tiny spaces in the matrix (lacunae) that become visible because
the xed cells shrink. When the chondrocytes of growing cartilages divide,
some lacunae remain closely associated as cell nests.
The common type of cartilage is more fully described as hyaline (glass-
like). Articulating surfaces of bones are covered with a specialized form of
hyaline cartilage called articular cartilage (Figure 42). Besides hyaline car-
tilage, there are two other forms of cartilage, also avascular, known as elas-
tic cartilage (Figures 43 and 44) and fibrocartilage (Figure 45). The
distribution of these reinforced forms of cartilage is more limited.
4
45
46 PDQ HISTOLOGY
Figure 41 Hyaline cartilage.
Perichondrium
Cell nests
Matrix
Figure 42 Articular cartilage.
Articular cartilage
Subchondral bone
Typical features of hyaline cartilage (see Figure 41) include the following:
matrix slightly blue (or slightly pink)
peripheral brous perichondrium (irregular dense ordinary connective
tissue)
lacunae large and round, with shrunken chondrocytes, particularly in
its midregion
lacunae atter near its periphery
some lacunae associated as cell nests
blood vessels absent
Articular cartilage (see Figure 42) has the following characteristics:
matrix pink, reecting extent of type II collagen packing
no peripheral perichondrium (hence postnatal growth is entirely
interstitial)
lacunae near the articular surface are parallel and flat (sandwiched
between horizontal collagen brils)
in the midregion, some lacunae are arranged in longitudinal columns
(a feature in growing articular cartilages indicating chondrocyte
proliferation)
vascular subchondral bone supporting it appears more acidophilic
because it contains relatively more collagen
avascular
Chapter 4 Cartilage and Bone 47
The characteristics of elastic cartilage (see Figures 43 and 44; see also
Figure 13) are as follows:
matrix contains supplementary acidophilic elastic bers that give the
matrix additional elastic properties (these bers are more conspicuous
with an elastin stain, eg, in Figure 44)
resembles hyaline cartilage in other respects
found in the epiglottis and external ear
48 PDQ HISTOLOGY
Figure 43 Elastic cartilage (hematoxylin and eosin stain).
Figure 44 Elastic cartilage (special elastin stain).
Distinctive features of brocartilage (see Figure 45) are as follows:
matrix contains reinforcing pink-staining bundles of type I collagen
bers that give the matrix extra tensile strength
round lacunae, containing round chondrocytes, are typically arranged
in straight rows
matrix is slightly blue between lacunae
found in tendon insertions, intervertebral disks, menisci, and pubic
symphysis
BONE
Bone matrix is heavily calcied (hydroxyapatite constitutes 70% of its wet
weight), and type I collagen constitutes approximately 90% of its organic
content, making it highly resistant to compression, tension, bending, and
twisting. In contrast, cartilage matrix stays largely uncalcied except during
long bone development and growth. Other important constituents of bone
matrix are (1) proteins that induce bone formation and promote matrix
calcication and (2) tissue uid representing roughly 25% of its wet weight.
Because bone matrix is heavily calcied and has a comparatively low water
content, bone tissue requires short-range diffusion from blood capillaries;
hence, it is highly vascular. Numerous ne communicating channels called
canaliculi interconnect its lacunae (see Figures 416 and 417), conveying
tissue uid that facilitates such diffusion.
Figure 45 Fibrocartilage.
Bone Cells
Bone tissue forming in connective tissue membranes incorporates develop-
ing loose connective tissue that brings with it a capillary blood supply. In
this vascular environment, mesenchymal cells differentiate into osteoblasts
that start secreting the macromolecular constituents of bone matrix.
In intramembranously developing at bones (Figure 46), basophilic
osteoblasts cover the surface of acidophilic bony trabeculae (beams), laying
down new matrix on the surface (appositional growth). Bone consisting of
anastomosing trabeculae like this is called cancellous bone.
Osteoblasts show the cytoplasmic basophilia and pale negative Golgi
region characteristic of active protein secretion (Figure 47). Spindle-
shaped osteoprogenitor (osteogenic) cells are also present on the surface.
These progenitor cells persist as bipotential stem cells that give rise to
osteoblasts at well-vascularized sites but produce chondrocytes at avascular
sites. Osteocytes appear smaller and less rounded than osteoblasts and lie
within lacunae (see Figure 47).
50 PDQ HISTOLOGY
Figure 46 Cancellous bone developing in an intramembranous environment.
Bony trabecula Osteoblasts
Capillary
In addition to this bone-forming cell family, bone tissue contains large
multinucleated cells called osteoclasts that resorb bone matrix (Figures 48
and 49; see also Figure 117). Derived from blood monocytes, osteoclasts
are recognizable by their multiple nuclei, relative size, acidophilic cyto-
plasm, and resorptive ruffled border applied to a ragged matrix surface or
resorption bay (Howships lacuna). They decalcify and then partly digest the
adjacent matrix locally by secreting H
+
ions and lysosomal hydrolases into
a sealed compartment of extracellular space under this border.
Figure 47 Osteoblasts and osteocytes of cancellous bone.
Osteoprogenitor cell
Osteocyte
Endothelial cell of
blood vessel
Osteoblasts
52 PDQ HISTOLOGY
Figure 48 Osteoclasts resorbing bone matrix.
Osteoclasts
Figure 49 Osteoclast seen in more detail.
Bone matrix is subject to lifelong internal remodeling in which old or
weakened bone resorbed through osteoclast activity is immediately replaced
by new bone through osteoblast activity. Osteoporosis (ie, a decrease in the
bodys total bone mass) is a manifestation of inadequate compensation of
bone resorption by bone formation.
Growth of Long Bones
Development and postnatal growth of a long bone involves the progressive
bony replacement of a temporary cartilage model. This indirect process,
known as endochondral ossification, is more complicated than the
intramembranous ossication occurring directly in connective tissue mem-
branes (see Figure 46). It is a consequence of vascularization of the model
and progressive encroachment into the cartilage by bone cells that lay down
layers of bone matrix by apposition on cartilage remnants.
The following zones may be recognized in the cartilaginous growth
plate (epiphyseal plate) present at either end of a childs typical growing
long bone (Figure 410):
resting (reserve) cartilageanchors epiphyseal plate to bony epiphysis;
chondrocytes randomly distributed
proliferating cartilagereplaces chondrocytes lost during ossication;
chondrocytes arranged as longitudinal stacks
maturing cartilagehypertrophying (enlarging) chondrocytes initiate
calcication; chondrocytes large and pale staining
calcifying cartilagecalcium phosphate becomes deposited in matrix;
chondrocytes along diaphyseal border die; transverse partitions of lacu-
nae disintegrate
Bone becomes deposited by apposition on the longitudinal partitions of
calcied cartilage extending as remnants from the diaphyseal border.
Bone Subtypes
Cancellous bone forming in the metaphysis under the epiphyseal plate (Fig-
ure 411) shows the following features:
light-staining wide vascular soft tissue spaces representing > 50% of its
volume
lamellae (layers) of dark pink bone matrix deposited on pale blue rem-
nants of calcied cartilage
osteoblasts and osteoprogenitor cells on the bone surface
Adult cancellous bone (from iliac crest) is shown in Figure 412. The
pale tissue between the trabeculae is fat-storing (yellow) bone marrow. Can-
cellous bone (upper border of Figure 413) can give rise to dense (compact)
bone (lower third of Figure 413) through appositional thickening of its
trabeculae. The additional new lamellae diminish the former soft tissue
spaces to narrow haversian canals. Persisting cancellous bone nevertheless
provides necessary internal support in medullary cavities (see Figure 412).
54 PDQ HISTOLOGY
Figure 410 Zones of an epiphyseal plate.
Zones:
resting
proliferating
maturing
calcifying
Figure 412 Cancellous bone (iliac crest).
Figure 411 Cancellous bone developing endochondrally.
Dense (compact) bone (Figures 414 and 415) is characterized by the
following features:
lled-in soft tissue spaces representing only a small proportion of its
volume
cylindrical haversian systems (osteons), each having lamellae and osteo-
cytes arranged concentrically around a central haversian canal
blood vessels in longitudinal haversian canals and radial Volkmanns
canals
smooth inner and outer circumferential lamellae
external periosteummade up of an outer vascular dense ordinary con-
nective tissue and a deeper population of osteoprogenitor cells
internal endosteum represented by a single layer of osteoprogenitor
cells that lines the medullary cavity and haversian canals
56 PDQ HISTOLOGY
Figure 413 Dense bone forming from cancellous bone.
Periosteum
Outer circumferential
lamellae
Interstitial lamellae
Inner circumferential
lamellae
Haversian system (osteon)
Cancellous bone in
medullary cavity
Blood vessel in a
Volkmann's canal
Haversian vessel in
a haversian canal
Figure 414 Organization of dense bone.
Figure 415 Dense bone.
Several histologic features of dense bone, for example, haversian sys-
tems, bone canaliculi, and interstitial lamellae, are more recognizable in
ground sections (silver-stained transparent thin slices of undecalcied bone
matrix). Figure 416 shows some details of a haversian system. After parts
of a haversian system have been dismantled by osteoclasts during the course
of internal remodeling, the parts that are left behind constitute interstitial
lamellae (Figure 417).
58 PDQ HISTOLOGY
Figure 416 Haversian system of dense bone (ground section, silver stain).
Haversian canal
Osteocyte lacunae
interconnected by
canaliculi
Figure 417 Interstitial lamellae of dense bone (ground section, silver stain).
Lamellae of haversian
system
Interstitial lamellae
JOINT TISSUES
In sections of articulating synovial joints, it is possible to nd articular car-
tilage (see Figure 42), brocartilage at tendon or ligament insertions into
cartilage (see Figure 45), epiphyseal plates if the bones are still growing
(see Figure 410), and the lining synovial membrane of the joint.
The synovial membrane (synovium), not regarded as an epithelial
membrane because it is made up of connective tissue cells trapped between
abundant interstitial bers (both collagen and elastin), has three character-
istic appearances. The vascular area shown at left in Figure 418 is typical
of an areolar region. Secretory cells (synovocytes) near its surface produce
the viscous glycoprotein and hyaluronic acid present in synovial uid, the
lubricating uid of the joint. The dense ordinary connective tissue shown
at the right in Figure 418 is a brous region with coarse collagen bundles
that are highly resistant to friction. The central fold of tissue in Figure 419
is an adipose region with synovocytes near its surface and underlying
grouped adipocytes that collectively serve as a cushion.
Figure 418 Areolar and brous regions of synovium.
Areolar region Fibrous region
60 PDQ HISTOLOGY
Figure 419 Adipose region of synovium.
Articular cartilage
Adipose region of synovium
Blood and Myeloid and
Lymphoid Tissues
Peripheral blood consists of plasma (ie, serum along with
brinogen and clotting factors) and formed elements, which are commonly
known as blood cells even though erythrocytes have no nucleus and
platelets are only cytoplasmic fragments. The total blood volume is about
5 L, with blood cells occupying roughly 45% of this volume. Blood cells are
produced by myeloid tissue and lymphoid tissue, the two hematopoietic
(hemopoietic) tissues.
BLOOD CELLS
The chief characteristics of the various kinds of circulating blood cells are
outlined in Table 51. In addition to erythrocytes (red blood cells) and
platelets, ve different types of leukocytes (white blood cells) exist, each
having a characteristic appearance and distinctive functions. Erythrocytes
are more abundant than leukocytes in peripheral blood. The neutrophil is
the predominant type of leukocyte.
Individual blood cells are routinely observed in peripheral blood lms
stained with suitable blood stains.
5
61
62 PDQ HISTOLOGY
Table 51
Blood Cells
Morphology and Staining
Cell (Blood Stain) Chief Functions
Erythrocyte Pink disk, without nucleus Transports oxygen as
oxyhemoglobin
Platelet Small blue fragment with central Platelet aggregates stop
purple granules and no nuclear blood loss from damaged
component blood vessels
Leukocytes
Neutrophil Pale with small mauve granules; Phagocytoses and destroys
nucleus segmented (25 lobes) bacteria
Eosinophil Large red granules; nucleus bilobed Destroys parasitic worms,
regulates local allergic
inammation
Basophil Large blue granules; nucleus Releases histamine and other
bilobed or segmented inammatory mediators
when triggered by antigen
cross-linking of surface-
bound immunoglobulin E
(systemic allergic response)
Lymphocyte Diameter small or large; cytoplasm B and T cells mediating
blue, without granules; nucleus immune responses
spherical
Monocyte Diameter large; cytoplasm pale Phagocytoses debris;
blue, without granules; nucleus circulating precursor of
shaped like a kidney or horseshoe macrophages and
osteoclasts
Erythrocytes and Platelets
Distinctive features of erythrocytes (Figure 51) are as follows:
biconcave disks, diameter 7 m
middle third thinner and therefore paler
contain the oxygen-transporting protein hemoglobin
worn out and destroyed after 4 months in the circulation
Platelets (Figure 52) have the following characteristics:
biconvex disks, diameter 3 m, dispersed or aggregated in blood lms
irregular mass of purple granules at the center
derived through fragmentation of megakaryocytes
contain the vasoconstrictor serotonin and a potent growth factor
Chapter 5 Blood and Myeloid and Lymphoid Tissues 63
Figure 51 Erythrocytes.
Figure 52 Platelets.
Granulocytes
64 PDQ HISTOLOGY
Figure 53 Neutrophil.
Figure 54 Eosinophil.
Key features of neutrophils (Figure 53) include the following:
mauve granules, smaller and paler than those of eosinophils or basophils
distinctive segmented nucleus
presence at acutely inamed sites
short life span (~ 3 days)
In sections, mature neutrophils are easily recognizable by their charac-
teristic segmented nucleus with two to ve lobes. Tissue sites where signif-
icant numbers of neutrophils lie external to blood vessels are sites of acute
inammation.
Eosinophils (Figure 54) have the following characteristics:
conspicuous red granules and relatively large diameter
distinctive bilobed nucleus
found in loose connective tissue in local allergic responses
Distinctive features of basophils (Figure 55) are as follows:
large blue granules commonly obscuring bilobed or segmented nucleus
rarely encountered in blood lms (~ 1% of the total leukocyte count)
Nongranular Leukocytes
Small lymphocytes (Figure 56) are characterized by the following:
diameter close to that of erythrocytes
dark-staining spherical or slightly indented nucleus occupying most of
the cells volume
thin rim of blue cytoplasm
more T cells than B cells in peripheral blood
majority are long-lived T cells
recirculate from blood to lymph to blood, etc
found in (1) loose connective tissue at sites where antigen has entered
and (2) lymphoid tissues
Large lymphocytes (Figure 57) may be recognized by the following:
diameter approaching that of monocytes
similar in nuclear morphology and cytoplasmic color to the small lym-
phocyte but relatively more cytoplasm
66 PDQ HISTOLOGY
Figure 55 Basophil.
Figure 57 Large lymphocyte.
Figure 56 Small lymphocyte.
Typical monocytes (Figure 58) have the following:
a large diameter
a relatively large amount of pale blue cytoplasm
a large kidney-shaped nucleus
MYELOID TISSUE
All types of blood cells except T lymphocytes are produced in myeloid tis-
sue (red bone marrow). The intricate process of blood cell formation is
known as hematopoiesis. Hematopoietic cell populations include pluripo-
tential self-renewing hematopoietic stem cells, hematopoietic progenitor
cells, and blood cell precursors. Respective properties, potentialities, and
acronyms have been assigned to these stem cells and progenitors, based in
large part on the types of blood cells they produce in tissue culture. In each
lineage, precursor stages can nevertheless be recognized directly with a light
microscope. Erythroid precursors, granulocytic precursors, and megakary-
ocytes (precursor cells of platelets) are examples of hematopoietic precur-
sors that histology students may be asked to identify in myeloid tissue.
68 PDQ HISTOLOGY
Figure 58 Monocyte.
Erythroid Series
The erythroid precursors, with their various alternative names, are listed in
Table 52 following the sequence in which they are formed.
The erythroid precursors shown in Figures 59 to 513 are individually
identiable in stained lms of red bone marrow.
Table 52
Precursor Stages of Erythropoiesis
Proerythroblast (pronormoblast)
Basophilic erythroblast (basophilic normoblast)
Polychromatophilic erythroblast (early normoblast)
Normoblast (late normoblast; orthochromatophilic erythroblast)
Polychromatophilic erythrocyte (reticulocyte)
Erythrocyte (normocyte)
70 PDQ HISTOLOGY
Figure 511 Polychromatophilic erythroblast.
Figure 512 Normoblast showing extrusion
of its nucleus.
Figure 59 Proerythroblast.
Figure 510 Basophilic erythroblast.
Proerythroblasts (see Figure 59) are recognizable by the following:
large diameter
large spherical nucleus, sometimes with large nucleoli visible
abundant blue cytoplasm
Basophilic erythroblasts (see Figure 510) have the following characteristics:
diameter almost as large as that of proerythroblasts
spherical nucleus; chromatin now more condensed
blue cytoplasm
relatively abundant
Polychromatophilic erythroblasts (see Figure 511) are characterized by
the following:
slightly smaller diameter
spherical nucleus proportionately smaller; chromatin more condensed
and dark staining
bluish-pink cytoplasm (cytoplasmic ribonucleic acid [RNA] along with
newly synthesized hemoglobin)
nal dividing stage in the erythroid series
Normoblasts (see Figure 512) have the following characteristics:
small diameter
dark-staining spherical nucleus, highly condensed and pyknotic,
becomes extruded from the cell
bluish-pink cytoplasm (becoming increasingly pink)
At the polychromatophilic erythrocyte stage (see Figure 513),
the diameter is slightly larger than that of a mature erythrocyte;
the nucleus has been extruded;
compared with the color of mature erythrocytes, the cytoplasm still
looks a slightly bluish pink; and
the cell enters the circulation and after 2 days stains as a mature
erythrocyte.
The blue component of the characteristic cytoplasmic staining of erythroid
precursors is the RNA involved in globin synthesis. Its recognition is facilitated
by the use of special stains (new methylene blue or cresyl blue) that aggregate
and stain it, generally as a wreath-like network (Figure 514). Polychro-
matophilic erythrocytes stained this way are called reticulocytes. Cells lacking
such blue-stained particles represent mature erythrocytes (see Figure 514).
72 PDQ HISTOLOGY
Figure 513 Polychromatophilic erythrocyte
and mature erythrocyte.
Figure 514 Reticulocytes and mature
erythrocyte.
Mature erythrocyte Polychromatophilic
erythrocyte
Granulocytic Series
Table 53 lists the granulocytic precursors in the order of their formation.
The granulocytic precursors that are most readily identied in marrow
lms are those with recognizable specic granules that are mauve, red, or
blue. Neutrophil precursors, characterized by mauve granules, predomi-
nate. The earliest precursors, myeloblasts and promyelocytes, are large cells
with spherical nuclei, sometimes with discernible large nucleoli. Promyelo-
cytes also contain inconspicuous azurophilic (primary) granules. However,
because these two stages can have quite an ambiguous appearance, histol-
ogy students are not often required to recognize them.
Table 53
Precursor Stages of Granulopoiesis (Neutrophil Series)
Myeloblast
Promyelocyte
Myelocyte
Metamyelocyte
Band neutrophil
Neutrophil
74 PDQ HISTOLOGY
Figure 516 Metamyelocyte (neutrophilic).
Figure 517 Band neutrophil.
Figure 515 Myelocyte (neutrophilic).
The neutrophilic myelocyte (Figure 515), the last dividing cell in the
neutrophilic series, may be recognized by the following:
diameter similar to that of the basophilic erythroblast
spherical or slightly indented nucleus with rather condensed chromatin
mauve specific (secondary) granules in addition to preexisting fine
azurophilic (primary) granules
The nucleus becomes more kidney shaped and condensed at the
metamyelocyte stage (Figure 516).
The band neutrophil (Figure 517) is a slightly immature neutrophil
that is nevertheless able to enter the peripheral blood. The highly condensed
nucleus is shaped like a band or horseshoe before it begins to segment.
Changes in the proportion of immature neutrophils to mature neutrophils
in the peripheral blood can be diagnostically indicative.
76 PDQ HISTOLOGY
Figure 518 Bone marrow lm.
Erythroid precursors
Granulocytic precursors
Figure 520 Megakaryocyte in red bone marrow.
Megakaryocyte
Lipid droplet
Figure 519 Section of red bone marrow.
Megakaryocyte
Sinusoid
In bone marrow lms (Figure 518), it is often possible to nd ery-
throid precursors by looking for spherical nuclei and to nd granulocytic
precursors (which are more numerous) by spotting their content of gran-
ules and/or elongating nucleus. Histologic organization of the tissue, how-
ever, is totally disrupted. Sections of myeloid tissue (Figures 519 and 520)
reveal further structural details in the tissue:
wide blood channels termed sinusoids
white empty-looking spaces representing lipid droplets in large fat-
storing stromal cells
large megakaryocytes, some adjacent to sinusoids
an assortment of dividing and differentiating hematopoietic cells held
in a reticular ber meshwork
Megakaryocytes (see Figure 519) have the following characteristics:
massive size
large multilobed nucleus that becomes increasingly polyploid as the cell
matures
copious cytoplasm that fragments into platelets at maturity
LYMPHOID TISSUE
Lymphoid tissue is characterized by an abundance of lymphocytes. Dif-
fusely distributed below wet epithelial membranes and aggregated in the
tonsils, Peyers patches, and the appendix, it also constitutes a key compo-
nent of the lymphoid organs, which are the thymus, lymph nodes, and
spleen. Because all of these locations except the thymus represent sites
where immune responses to antigen occur, they are often described as the
secondary lymphoid organs and tissues. The thymus is considered one of
the primary lymphoid organs because it is the site where T cells differenti-
ate. The other primary lymphoid organ is bone marrow, the site where
mammalian B cells differentiate.
78 PDQ HISTOLOGY
Figure 521 Lingual tonsil.
Crypt
Lymphoid follicle
Mucous glands
Figure 522 Palatine tonsil.
Crypt
Lymphoid follicles
Tonsils are substantial aggregates of lymphoid follicles situated in the
pharynx (palatine tonsils), nasopharynx (pharyngeal tonsil), and tongue
(lingual tonsil).
The lingual tonsil (Figure 521) is recognized by the following:
conspicuous lymphoid follicles
overlying stratied squamous nonkeratinizing epithelium
underlying mucous glands
crypts kept clear of debris by secretion
The palatine tonsils (Figure 522) have the following characteristics:
large aggregates of lymphoid follicles
overlying stratied squamous nonkeratinizing epithelium
no underlying glands
deep crypts that commonly contain debris
more susceptible to infection
The palatine tonsils are the ones that are removed if they become chron-
ically infected.
80 PDQ HISTOLOGY
Figure 523 Lymph node.
Capsule Subcapsular sinus
Cortical lymphoid follicle
Medullary
sinus
Figure 524 Thymus.
Cortex
Medulla
Distinctive recognizable histologic features of lymph nodes (Figure
523) are the following:
afferent and efferent lymphatics
connective tissue capsule with subcapsular sinus
blue-staining cortex with lymphoid follicles and paracortical high
endothelial venules
antibody-secreting plasma cells present in medullary cords
pale lymph-containing medullary sinuses
The thymus (Figure 524) has the following characteristics:
incompletely lobulated appearance
dark blue-staining cortex packed with small lymphocytes (proliferating
and differentiating T cell progenitors)
cortical network of pale-staining thymic epithelial reticular cells (source
of thymic cytokines and hormones)
medullary pink spherical thymic (Hassalls) corpuscles
The spleen (Figure 525) has the following distinctive features:
connective tissue capsule with substantial inward projections (trabeculae)
blue areas (white pulp) containing abundant lymphocytes
close association between white pulp and splenic arterial blood vessels
(eg, the follicular arterioles found in its lymphoid follicles)
pink structural component (red pulp) with characteristic wide sinu-
soids and closely associated active macrophages
82 PDQ HISTOLOGY
Figure 525 Spleen.
Lymphoid follicle with
central arteriole
Red pulp
Sinusoid
Capsule Trabecula
Nervous Tissue
The nervous systemis made up of nervous tissue with a cer-
tain amount of associated connective tissue. The neurons (nerve cells) of
the central nervous system (CNS) are chiey supported by neuroglial cells
(glia), an ancillary population of heterogeneous cells that, in common with
neurons, are derived from neuroectoderm. Extending from the CNS is the
peripheral nervous system(PNS), consisting of the peripheral nerves, gan-
glia, and afferent and efferent nerve endings. A characteristic of the PNS is
that its nerve bers, neuronal cell bodies, and associated glia are intimately
supported and strengthened by substantial wrappings of connective tissue.
Three concentric external connective tissue membranes collectively known
as the meninges similarly invest and protect the brain and spinal cord.
CENTRAL NERVOUS SYSTEM
The brain and spinal cord are both made up of two dissimilar arrangements
of nervous tissue termed gray matter and white matter. Gray matter is an
intimate assortment of neuronal and glial cell bodies embedded in an
entwined mass of nerve bers (many without a myelin sheath) termed the
neuropil (see Figure 63). White matter is a more structured arrangement
of nerve bers, many with myelin sheaths, along with their associated glia.
Neuronal cell bodies, on the other hand, are absent from white matter. The
two different arrangements are more easily recognized in sections of spinal
cord and cerebellum than in sections of cerebral cortex. Myelin sheaths
invest and insulate fast-conducting axons. The axon is generally the neu-
rons longest nerve ber, and, in most cases, it propagates nerve impulses
away from the cell body (afferent neurons being an exception). Compared
with the axon, the dendrites are typically shorter, more branched, and
tapered, and their role is to receive nerve impulses. In hematoxylin and
eosin (H&E)-stained sections, it is rarely evident whether representative
unmyelinated nerve bers are axons or dendrites.
6
83
84 PDQ HISTOLOGY
Figure 61 Spinal cord.
Posterior horn
of gray matter
Subarachnoid
space
White
matter
Anterior horn of
gray matter
Dura
Figure 62 Gray matter and white matter of spinal cord.
Neuropil
Multipolar neurons in gray matter
Myelinated nerve bers
in white matter Pia
Spinal Cord
The spinal cord (Figure 61) can be recognized by a combination of
distinctive features:
central H-shaped area of gray matter (anterior and posterior horns with
interconnecting gray commissures)
surrounding white matter (funiculi containing descending and ascend-
ing longitudinal tracts)
anterior median ssure and posterior median sulcus
spinal nerve roots in the subarachnoid space
surrounding meninges (dura mater, arachnoid membrane, and pia
mater)
At higher magnication (Figure 62), it may be seen that
multipolar neurons are present in the gray matter,
delicate ingrowths of the pia mater (innermost meningeal membrane)
bring a blood supply,
shrinkage artifact is common (particularly between neuronal cell bod-
ies and the surrounding neuropil), and
white matter contains pale-looking myelin sheaths (myelin lipids are
extracted during processing) but lacks neuronal cell bodies.
Chapter 6 Nervous Tissue 85
86 PDQ HISTOLOGY
Figure 63 Multipolar neuron.
Axon
hillock
Nucleolus
in nucleus
Neuropil
Capillaries
Dendrite
Glial cell
nuclei
Figure 64 Electron micrograph of a chemical synapse.
Synaptic vesicles in
axon
Dendrite
Postsynaptic
thickening
Typical neurons (Figure 63) are characterized by the following features:
large diameter
asymmetric multipolar, pseudounipolar, or bipolar shape (multipolar
in this case)
intense cytoplasmic basophilia, typically patchy and indicative of
patches of rough-surfaced endoplasmic reticulum (Nissl bodies),
reecting active protein synthesis for maintenance of long axon
axon hillock (site where axon is attached) is generally not basophilic
dendrites with basophilic cytoplasm as in cell body
pale-staining central nucleus with central large nucleolus (owls eye
nucleus) that also reects active protein synthesis
Neuropil and blood capillaries may also be seen to advantage in this
photomicrograph.
The special sites on neurons where nerve impulses can be transmitted
from one cell to another are known as synapses. Chemical synapses (the
common type) release neurotransmitter into their synaptic cleft; electrical
synapses (uncommon) possess gap junctions that conduct impulses directly.
Typical features of an axodendritic directed chemical synapse seen in
the electron microscope (Figure 64) include synaptic vesicles containing
neurotransmitter and electron-dense material (variable in amount) on the
cytoplasmic surface of the postsynaptic membrane (postsynaptic thicken-
ing). In addition to chemical synapses, there are electrical synapses. These
are essentially gap junctions (see Figure 210, C) that permit direct con-
duction of nerve impulses.
88 PDQ HISTOLOGY
Figure 66 Cerebral cortex (silver impregnation).
Neuroglial
cell
Figure 65 Cerebral cortex (hematoxylin and eosin stain).
Pia Neuronal cell
body
Capillary
Brain
In H&E sections of cerebral cortex (Figure 65), the following may be
found:
supercial gray matter with neurons arranged as a series of indistinct
laminae (layers)
neuronal cell bodies, the largest of which are pyramid shaped (pyram-
idal cells)
widely scattered nuclei of neuroglia and surrounding neuropil
blood capillaries supplying the necessary oxygen and nutrients
Silver or gold impregnation methods (Golgi preparations) selectively
reveal representative neurons and neuroglia (Figure 66). With special
staining, the extensively interlacing long cytoplasmic processes of astro-
cytes, the predominant type of glial cell, may be seen (Figure 67).
Figure 67 Fibrous astrocytes (special stain).
90 PDQ HISTOLOGY
Figure 68 Cerebellum.
Molecular layer Granular layer
White matter
Figure 69 Purkinje cell of the cerebellum.
Granular layer Molecular layer
Purkinje cell
The cerebellar cortex (Figure 68) may be recognized by the following
characteristics:
deeply folded appearance (ssures and sulci)
pink outer cortical layer of gray matter (molecular layer)
blue inner cortical layer of gray matter (granular layer)
ask-shaped neuronal cell bodies (Purkinje cells) scattered along the
interface between the outer and inner cortical layers
pink central medullary core of white matter
Characteristic features of Purkinje cells (Figure 69) include the
following:
large diameter
cytoplasmic basophilia (Nissl bodies) and nuclear morphology both
typical of neurons
extensively branching dendritic tree extending far into the molecular
layer
PERIPHERAL NERVOUS SYSTEM
The PNS is more strongly reinforced by abundant connective tissue. Fur-
thermore, under the right conditions, peripheral nerves show considerable
potential for nerve ber regeneration when they become damaged. Small
peripheral nerves may often be found in adventitial connective tissue of
organs or structures. Sensory cranial ganglia and spinal ganglia (see Figure
613) have a slightly different histologic appearance from autonomic gan-
glia (see Figure 614), but the difference is subtle.
92 PDQ HISTOLOGY
Axon
Schwann cell
and myelin sheath
Fascicle
Epineurium
Perineurium
Endoneurium
Figure 610 Organization of a peripheral nerve.
Figure 611 Peripheral nerve (cross-section).
Perineurium surrounding
fascicle
Myelinated nerve bers
Peripheral Nerves
A moderate-sized peripheral nerve has the structural organization shown in
Figure 610. Its outermost brous connective tissue covering, not repre-
sented in small nerves, is termed its epineurium. Internal to this, brous
perineurium lls the crevices between nerve ber bundles and surrounds
each bundle as a sheath. Inside the bundle, loose connective tissue known
as endoneurium lies between the individual nerve bers.
Typical features of peripheral nerves cut in cross-section (Figure 611)
or longitudinal section (Figure 612) are as follows:
nerve ber bundles (fascicles), each bundle surrounded with perineurium
nerve bers, with or without white myelin sheaths (compare Figures
611 and 612)
endoneurium often difficult to discern
epineurium not represented except in large nerves
nuclei of Schwann cells, occasionally with nuclei of endoneurial broblasts
Figure 612 Fascicle of peripheral nerve (longitudinal section).
Perineurium surrounding
fascicle
Schwann cell nuclei
94 PDQ HISTOLOGY
Figure 614 Autonomic ganglion (parasympathetic ganglion in pancreas).
Nuclei of capsule cells
Cell body of ganglion cell
Unmyelinated
nerve bers
Figure 613 Spinal ganglion.
Cell body of ganglion cell
Nuclei of capsule cells
Spinal Ganglia
A section of spinal (posterior root or dorsal root) ganglion (Figure 613) is
recognizable by the following:
pseudounipolar ganglion cells with large spherical cell bodies
ganglion cell nucleus central to the cell body, conspicuous large nucle-
olus and cytoplasmic basophilia (Nissl bodies) indicative of active pro-
tein synthesis
each ganglion cell body enclosed by a layer of comparatively small cap-
sule cells (neuroglial cells)
AUTONOMIC NERVOUS SYSTEM
Viscera, exocrine glands, blood vessels, and smooth muscles have their func-
tional activities regulated automatically by way of their autonomic inner-
vation. Sympathetic and parasympathetic ganglia of the autonomic nervous
system (Figure 614) are both characterized by the following features:
large multipolar ganglion cells, cell bodies more irregular in shape because
of multiple dendrites but otherwise resembling spinal ganglion cells
ganglion cell nucleus more commonly eccentric in position
the population of small capsule cells associated with each ganglion cell
appears discontinuous rather than as a layer because of the neurons
multipolar shape
96 PDQ HISTOLOGY
Neural retina Retinal pigment
epithelium
Figure 615 Position of the retina. Box indicates the site and orientation of Figures 616
and 617.
Inner nuclear
layer
Outer plexiform layer
Nuclei and cell bodies
of photoreceptors
External limiting membrane
Outer and inner segments
of photoreceptors
Retinal pigment epithelium
(simple cuboidal; outermost)
Cone
Rod
Neuroglial cells
Figure 616 Photoreceptive part of the retina.
OPTIC RETINA
The functional importance of the retina (Figures 615, 616, and 617a
toluidine bluestained semithin section) is that it is uniquely provided with
photoreceptors. Some essential features by which retinal sections may be
identied are as follows:
inner multilayered neural retina (photoreceptors, other retinal neurons,
optic nerve bers)
outer single-layered retinal pigment epithelium (RPE) (a simple
cuboidal melanin-containing epithelium)
outer and inner segments of photoreceptors (rods and cones) adjacent
to RPE
a band of photoreceptor nuclei adjacent to the layer containing the
inner segments
a wide band containing nuclei of other retinal neurons and tall colum-
nar supporting neuroglia
inner layers (not shown) containing fewer nuclei and more nerve bers
optic nerve bers forming the supercial retinal layer that lines most of
the vitreous cavity of the eye
Figure 617 Outer part of the retina.
Muscle
Muscles are described histologically as consisting of mus-
cle cells associated with moderate to large amounts of connective tissue.
Each of the three muscle typesskeletal, cardiac, and smoothhas dis-
tinctive features. When seen with a microscope, skeletal muscle (responsible
for voluntary movements) and cardiac muscle (involuntary heart muscle)
show striations. Smooth muscle (involuntary muscle of viscera and
contractile vessels) lacks striations.
Detailed descriptions of muscle cells commonly use terms that would
benefit from some initial explanation. In the case of skeletal or smooth
muscle, the term muscle ber is often used as a synonym for muscle cell. In
cardiac muscle, however, it means a row of individual cells joined end to
end. Myobrils, present in skeletal and cardiac muscle but not smooth mus-
cle, are thread-like organized arrangements of thick and thin laments. Fil-
aments of all three classes (thick, thin, and intermediate) are present in all
three types of muscle cells. The underlying structural basis for all termi-
nology is summarized in Figure 71.
SKELETAL MUSCLE
As in peripheral nerves, skeletal muscle bers are arranged in bundles (fas-
cicles), with brous perimysiumsurrounding each bundle (see Figures 71
and 72). The muscle is enclosed as a whole by brous epimysium, and del-
icate endomysium lies between the muscle bers.
7
99
100 PDQ HISTOLOGY
Thin and thick
filaments
Myofibril
Skeletal muscle
fiber
Epimysium
Perimysium
Fascicle
Endomysium
Endomysium
Figure 71 Organization of a skeletal muscle.
Chapter 7 Muscle 101
Figure 72 Skeletal muscle at low magnication.
Skeletal muscle bers
in fascicle
Endomysium Perimysium
102 PDQ HISTOLOGY
Figure 73 Skeletal muscle bers in cross-section.
Nucleus of capillary
endothelial cell
Myobrils
Capillary
endothelial cell
Skeletal muscle ber
containing myobrils
Nuclei of
muscle bers
The robust perimysial sheaths of skeletal muscle fascicles are recogniz-
able in cross-sections of skeletal muscle, particularly at very low magnica-
tions (see Figure 72). With more magnication, it is evident that skeletal
muscle bers are multinucleated (as a result of fusion of myoblasts, their
precursor cells), and their nuclei lie in the peripheral cytoplasm (Figure
73). The interior portion of the muscle bers appears closely packed with
myobrils (the breaks between them are xation artifact). The distinctive
multinucleated appearance of skeletal muscle bers becomes more appar-
ent in longitudinal sections (Figure 74).
Striations
Under optimal viewing conditions (or after special staining), transverse
striations may also be seen on the myobrils: dark-staining A bands and
light-staining I bands. These bands seem to extend across the entire muscle
fiber because striations of myofibrils are aligned transversely across the
ber. Scattered broblast nuclei or capillary endothelial nuclei (see Figure
73) can sometimes be found in the pale-staining endomysium separating
the muscle bers.
Figure 74 Skeletal muscle bers in longitudinal section, showing striations.
Nuclei of a skeletal
muscle ber
Striations
Sarcomeres
A distinctive arrangement of thick and thin laments known as the sar-
comere (Figure 75) is repeated along myobrils in the form of an extended
linear series. The sarcomere is considered the basic contractile module of
skeletal and cardiac muscle. The thin laments, which contain actin, tro-
ponin, and tropomyosin, are anchored to Z lines made of -actinin. The
thick laments, which contain myosin, are centrally interconnected in the
transverse plane by a narrow M line containing myomesin. A bands contain
both thick and thin laments. I bands do not include any thick laments. H
bands do not include any thin laments. During contraction, the H band
disappears and the I bands shorten because the thin laments are pulled far-
ther into the A band, and this decreases the distance between the Z lines at
the ends of the sarcomere.
CARDIAC MUSCLE
Although cardiac muscle resembles skeletal muscle in certain respects, car-
diac muscle is involuntary and has fibers made up of individual cardiac
muscle cells, each with one or two central nuclei. Distinctive structures
called intercalated disks attach these muscle cells to each other, forming
bers that show a certain amount of branching. Hence, two key histologic
features of cardiac muscle bers are their limited anastomosis and their dis-
tinctive assembly pattern incorporating separate cardiac muscle cells with
intercalated disks between them (Figure 76). A delicate endomysium occu-
pies the spaces between cardiac muscle bers, but epimysium and perimy-
sium, characteristic of skeletal muscle, are unrepresented. Unlike skeletal
muscle, which can regenerate new muscle bers from satellite (myosatellite)
cells situated at the periphery of its muscle bers, cardiac muscle is charac-
terized by a negligible inherent potential for regeneration. If it dies, it is
replaced by brous scar tissue.
104 PDQ HISTOLOGY
I A I
H Z Z
Relaxed
Contracted
M
Figure 75 Sarcomere, showing its bands and the arrangement of thin and thick laments.
Intercalated disks (see Figure 76) are not evident at the light micro-
scopic level unless special stains are used. Electron microscopy reveals that
they are specialized for attachment and electrical conduction. They traverse
the muscle ber in a step-like manner at the level of Z lines. Their distinc-
tive features are as follows:
extensive interdigitation of cell borders
exclusive site of gap junctions that allow free ion movement (direct elec-
trical conduction)
transverse adhering junctions (two types) that anchor (1) thin laments
and (2) intermediate laments
Desmosome-like
junction
Fascia adherens
junction
Gap
junction
Intermediate
filaments
Thick
filament
Thin
filament
Z line
Figure 76 Part of an intercalated disk, showing the component junctions and associated
laments.
106 PDQ HISTOLOGY
Figure 77 Cardiac muscle in cross-section.
Central nuclei
Endomysial
capillary
Peripheral myobrils
in muscle bers
Figure 78 Cardiac muscle in longitudinal section.
Branched muscle bers
Endomysial capillaries
Central nuclei of muscle bers
Cross-sections of cardiac muscle fibers (Figure 77) have these
characteristics:
a large diameter
a central nucleus
peripheral myobrils
highly vascular associated endomysium
In longitudinal sections (Figure 78), muscle ber branching may also
be seen, together with striations at higher magnications. Intercalated disks
are not usually identifiable at this magnification in hematoxylin and
eosinstained sections.
SMOOTH MUSCLE
The appearance of smooth muscle bers cut in cross- and longitudinal sec-
tion may be seen in Figure 79. Smooth muscle bers (commonly called
smooth muscle cells) may be recognized by the following:
small diameter compared with other types of muscle ber
arranged as muscle coats, layers, or bundles
surrounding sheaths of loose connective tissue
central nucleus
absence of the discrete myobrils and aligned striations that character-
ize skeletal and cardiac muscle
distinctive pleating of the nucleus in contracted and partly contracted
muscle bers (compare Figures 710 and 711)
Other features of smooth muscle include gap junctional coupling (basis
of excitation spread through direct conduction, as in cardiac muscle) and
postnatal potential for mitosis and supplemental derivation from pericytes.
Figure 79 Smooth muscle in cross-section (left) and longitudinal section (right).
108 PDQ HISTOLOGY
Figure 711 Smooth muscle bers in the contracted state.
Pleated nuclei
Figure 710 Smooth muscle bers in the relaxed state.
Circulatory System
Circulatory system is a broadly inclusive term for (1) the
heart and the various types of blood vessels characterizing the blood circu-
latory systemand (2) the lymph-collecting vessels of the lymphatic system.
Components of the circulatory system are conventionally described as being
made up of three concentric layers known as tunicaethe tunica intima,
tunica media, and tunica adventitiabut the borders between these layers
are hard to distinguish in certain cases. In general, the tunica intima is a
loose connective tissue layer with endothelium, a characteristic simple
squamous lining epithelium, anchored to it by underlying basement mem-
brane. The intimal lining layer of the heart is known as the endocardium,
and heart valves are sheet-like inward extensions of this layer. Deep in the
endocardium lie specialized impulse-conducting cardiac muscle fibers
(Purkinje bers) belonging to the impulse-conducting system of the heart.
In arterial vessels, the intima is characterized by an additional substantial
layer of fenestrated elastin called the internal elastic lamina. The tunica
media of vessels and myocardium of the heart, the middle layer of their
walls, consist primarily of muscle cellssmooth muscle in most vessels and
cardiac muscle in the heart. The total amount of muscle present, however,
is consistent with the components function: abundant in the heart, meager
in venules, and unrepresented in capillaries. The outermost region of the
wall, the tunica adventitia, is a layer of loose connective tissue that in the
great veins also contains substantial bundles of smooth muscle. The tunica
adventitiaparticularly that of veinsand also the outer region of the
media of some vessels are nourished by vasa vasorum (small nutritive ves-
sels). The equivalent cardiac layer is the epicardium, consisting of brous
connective tissue and the overlying visceral layer of serous pericardium.
Lymphatic vessels share this wall structure, but the layers of the wall are
harder to distinguish. The component vessels of the blood circulatory and
lymphatic systems are listed in Table 81. In light microscopic sections,
metarterioles resemble arterioles and lymphatic capillaries may be difficult
to recognize and distinguish with condence from blood capillaries.
The main features of blood vessel wall structure are shown in Figure 81.
8
109
110 PDQ HISTOLOGY
Table 81
Blood Circulatory and Lymphatic Vessels
Blood circulatory system
Elastic arteries (aorta)
Distributing (muscular) arteries
Arterioles and metarterioles
Capillaries
Venules
Small- and medium-sized veins
Large collecting veins (vena cava)
Lymphatic system
Lymphatics and lymphatic ducts
Lymphatic capillaries
Aorta Distributing artery Arteriole
Vena cava Vein Venule
(large)
Smooth muscle
(media)
Smooth
muscle
(media)
Smooth muscle
in adventitia
Smooth muscle
(media)
Subendothelial layer
(intima)
Internal
elastic
lamina Smooth muscle
(media)
Elastin
(laminae, fibers)
External
elastic
lamina
Figure 81 Comparative composition of the walls of blood vessels.
ELASTIC ARTERIES
Distinctive wall features of the aorta, which is an elastic artery (Figure 82;
see also Figure 81), are as follows:
marked overall thickness
substantial content of elastin (maintains diastolic blood pressure)
uniformly laminated appearance of the very wide media (rows of dark
pink smooth muscle cells separated by pale pink elastic laminae)
relatively thick intima compared with intima of a muscular artery
(endothelium with substantial subendothelial layer containing elastin,
smooth muscle cells, and some broblasts)
somewhat indistinct border between intima and media
even more indistinct internal and external laminae
vasa vasorum in adventitia, penetrating outer part of media
together with muscular (distributing) arteries, a site of development of
atherosclerotic plaques
Chapter 8 Circulatory System 111
Figure 82 Aorta.
Intima
Media
Adventitia
112 PDQ HISTOLOGY

Figure 83 Distributing artery and vein (hematoxylin and eosin stain).
Figure 84 Distributing artery (toluidine blue stain).
Artery Vein
Internal elastic lamina
Smooth muscle cells
Endothelium
DISTRIBUTING ARTERIES
Also known as muscular arteries, distributing arteries (Figures 83 and 84;
see also Figure 81) may be recognized by the following:
substantial wall thickness compared with vein but thinner than aortic
wall
very thin intima compared with aorta (endothelium and underlying
internal elastic lamina)
wavy pale-staining internal elastic lamina (demonstrated to advantage
by toluidine blue staining; see Figure 84)
wide media containing dark pink smooth muscle cells circularly
arranged (regulates blood ow)
distinct border between muscular media and substantial connective tis-
sue adventitia
VEINS
Features of the accompanying veins (see Figures 81 and 83) include the
following:
relatively thin walls (lower venous pressure)
media generally thinner, less distinct, and less muscular (except in
supercial leg veins)
adventitia fairly thick relative to other layers
internal elastic lamina unrecognizable
intimal ap valves, rarely seen in sections
degenerative changes in their wall components can lead to dilatation
(varicose veins)
The inferior vena cava (see Figure 81) has some additional character-
istics:
wide lumen
fairly thin media compared with adventitia
thick contractile adventitia containing longitudinal bundles of smooth
muscle
114 PDQ HISTOLOGY
Figure 85 Arteriole.
Figure 86 Arteriole, venules, and capillaries.
Smooth muscle cells
Endothelium
Venules
Capillaries
Arteriole
SMALL BLOOD VESSELS
The features by which arterioles (Figures 85 and 86; see also Figure 81)
may be recognized include the following:
small overall diameter (< 100 m)
thick walled relative to lumen (ie, wall thickness almost comparable
with luminal diameter)
distinct media made up of only one or two layers of circularly arranged
smooth muscle cells
internal elastic lamina generally discernible except in the smallest arte-
rioles (see Figure 86)
indistinct border between media and adjacent thin adventitia
Venules may often be found in the vicinity of their companion arteri-
oles (see Figure 86). They are characterized by the following:
small diameter generally between that of arterioles and capillaries
very thin walls
barely recognizable smooth muscle cells in media (unrepresented in
small venules)
allowing leukocytes and fluid exudate to escape at sites of acute
inflammation
Blood capillaries are distinguishable from arterioles and venules by
their small luminal diameter (see Figure 86). However, few details are dis-
cernible without the electron microscope.
116 PDQ HISTOLOGY
Figure 88 Lymphatic.
Figure 87 Electron micrograph of a capillary.
Pericyte
Erythrocyte
Endothelial
cell
Typical blood capillaries (Figure 87) have the following characteristics:
small diameter (810 m, only slightly larger than an erythrocyte; see
Figure 87)
very thin endothelial lining (continuous or fenestrated)
associated pericytes
no media
LYMPHATIC VESSELS
Lymphatics (Figure 88) may be recognized through any combination of
the following:
uniform pink staining of the lymph protein in their lumen
presence of lymphocytes (without other blood cells) in their lumen
relatively wide lumen (appropriately scaled down in lymphatic capillaries)
very thin walls, occasionally showing recognizable smooth muscle cells
in larger lymphatics
poorly organized or skimpy appearance of walls
proximity to lymphoid organs (eg, lymph nodes)
Lymphatic capillaries differ from blood capillaries in the following
respects:
slightly wider lumen
blind ending
incomplete endothelial basement membrane
basal border of endothelium anchored to adjacent collagen bers
pericytes absent
Integumentary System
The integumentary systemconsists of the skin and vari-
ous types of skin appendages (Table 91). Each kind of skin is constructed
from two tissue components: epidermis, a stratied squamous keratinizing
epithelium derived from ectoderm, and dermis, its connective tissue layer
derived from mesoderm. The term hypodermis denotes the subcutaneous
adipose layer found immediately under the dermis. The skin-associated
glands, hairs, and nails (ie, all skin appendages) are entirely epidermal
derivatives.
In thick skin, which covers the palms of the hands, soles of the feet, and
the exor surface of digits, epidermis is almost half as thick as the dermis,
whereas in the thin skin covering the remainder of the body, epidermal
thickness is only about one-fth of the dermal thickness (Figures 91 and
92). The terminology is based on the thickness of the epidermis, not the
total thickness of the skin. Unless understood, this can be quite confusing
because thin skin has a greater total thickness than thick skin! One reason
9
119
Table 91
Integumentary System
Skin
Epidermis
Dermis
Epidermal skin appendages
Hairs
Nails
Sebaceous glands
Sweat glands
Eccrine type
Apocrine type
120 PDQ HISTOLOGY
Reticular layer of dermis
Sweat gland
Subcutaneous
fat
Pacinian
corpuscle
Blood vessel
Stratum corneum
Stratum lucidum
Stratum granulosum
Stratum spinosum
Stratum
germinativum
Papillary layer
of dermis
A
B
Figure 91 A, Organization of thick skin. B, Details of the epidermis.
Hair
Papillary layer
of dermis
Sebaceous gland
Reticular layer
of dermis
Arrector pili muscle
Sweat gland
Root sheath of
hair follicle
Connective tissue
papilla
Subcutaneous fat
Figure 92 Organization of thin skin.
for making a distinction between two types of skin is that certain structural
differences exist, notably the presence of friction ridges and protective
thickened stratum corneum in thick skin and the presence of hair follicles
and sebaceous glands in thin skin.
Because epidermis is an avascular epithelium, it must receive nutrients
from capillaries in the underlying papillary layer of dermis. Progressive dis-
placement of its keratinocytes toward the surface isolates them from this
source, so they die and ake off. The turnover time for keratinocytes to
reach the skin surface from the stratum germinativum is almost 1 month.
The deep parts of certain skin appendages (namely, the growing region
of hair follicles and the secretory portion of most sweat glands) extend
down from the dermis into the subcutaneous tissue (see Figures 92 and
97). Encapsulated pressure-sensitive sensory nerve endings resembling
onions (pacinian corpuscles) may also be found in this adipose layer, par-
ticularly in sections of thick skin (Figure 93).
THICK SKIN AND THIN SKIN CHARACTERISTICS
The following generalizations apply to thick skin (see Figure 91):
ve distinguishable epidermal layers (germinativum, spinosum, granu-
losum, lucidum, corneum)
thick stratum corneum
dermal papillae arranged on double (secondary) dermal ridges under
friction ridges
no hair follicles or sebaceous glands
eccrine sweat glands opening onto friction ridges
General features of thin skin (see Figure 9-2) include the following:
four distinguishable epidermal layers (germinativum, spinosum, gran-
ulosum, corneum)
thin stratum corneum
dermal papillae randomly arranged
hair follicles with associated ask-shaped sebaceous glands
eccrine and apocrine sweat glands (apocrine glands, limited to pubic,
perineal, and axillary areas and breasts, open into hair follicles)
nail plates in nail grooves
Chapter 9 Integumentary System 121
122 PDQ HISTOLOGY
Figure 94 Thick skin seen in more detail.
Corneum
Granulosum
Spinosum
Papillary layer
of dermis
Duct of sweat gland
Reticular layer
of dermis
Sweat gland
Figure 93 Thick skin at low magnication.
Corneum
Sweat gland
Adipose tissue
Pacinian corpuscle
Seen under low power, sections of thick skin (see Figure 93) are rec-
ognizable by the following:
thick stratum corneum
total depth (epidermis + dermis down as far as subcutaneous fat)
smaller than in thin skin
hair follicles and sebaceous glands absent (note that thorough low-
power searching of the entire section is mandatory for certainty about
this)
pacinian corpuscles also present in this particular case (compatible with
its being thick skin but does not rule out thin skin)
Increased magnications of thick skin (Figure 94) disclose the following:
epidermal layers (notably, spinosum, granulosum, and thick corneum)
vascular dermal papillae (part of the papillary layer of dermis)
comparatively coarse collagen bundles indicative of the reticular layer of
dermis (beneath the papillary layer)
eccrine sweat glands, with their duct portion extending up through the
dermis and their secretory portion situated in the subcutaneous adipose
tissue
124 PDQ HISTOLOGY
Figure 95 Thin skin at low magnication.
Figure 96 Epidermis of thin skin seen in more detail.
Corneum
Granulosum
Spinosum
Papillary layer
of dermis
Reticular layer
of dermis
Stratum granulosum
Melanin in stratum germinativum
Sections of thin skin (Figures 95 and 96) typically show the following:
relatively thin epidermis
recognizable germinativum, spinosum, and corneum; granulosum
usually somewhat indistinct, and lucidum is not represented
relatively thin stratum corneum
melanin mostly confined to deep layers of the epidermis in light-
skinned races (variable in amount; synthesized by neural crestderived
epidermal melanocytes and acquired by keratinocytes through
phagocytosis)
In histology tests, students are generally expected to state whether the
skin being identied is thick or thin.
HAIR FOLLICLES
Although sections of thin skin may happen to include isolated small por-
tions of hair follicles and/or their associated sebaceous glands, this is often
not the case. These particular skin appendages are more easily observed in
sections of the scalp, where the hair follicles are more tightly packed and
there are numerous oblique or longitudinal cuts to look at.
The scalp is an area of thin skin uniquely characterized by a dense pop-
ulation of hair follicles (Figure 97). Readily identiable features include
the following:
hair follicles containing growing hairs, extending down into subcuta-
neous tissue
follicle-associated, flask-shaped sebaceous glands (produce sebum
through disintegration of their lipid-laden cells)
arrector pili muscles (smooth muscles that give sebaceous glands a
squeeze and make hairs stand up on end)
representative parts of hairs and hair follicles, composite drawings of
which are shown in Figure 98
It is seldom expected that students will memorize every microscopic
detail of these structures. Recognition of the following components could
be considered challenging enough:
connective tissue papilla (nourishes the growth zone)
matrix region of the hair (the growth zone)
cortex and medulla of hair (medulla unrepresented in some types of hair)
internal and external root sheaths
connective tissue sheath (external to root sheath)
126 PDQ HISTOLOGY
Figure 97 Scalp.
Sebaceous
glands
Arrector
pili muscles
Papilla of
hair follicle
Adipose
tissue
Blood
vessel
Sebaceous
gland
Arrector pili
muscle
Connective tissue
sheath
External root sheath
Internal root sheath
Stratum
germinativum
Hair matrix
Connective
tissue papilla
Trichohyalin
granules
Cortex
of hair
Medulla
of hair
A. B.
Figure 98 Organization of a hair follicle. A, Longitudinal section. B, Cross-section at the level
indicated in A.
SWEAT GLANDS
Two types of sweat gland exist, termed eccrine and apocrine.
Eccrine Type
The widely distributed, ordinary kind of sweat gland (eccrine sweat gland;
Figure 99), represented in both thick and thin types of skin, shows the fol-
lowing features:
simple tubular gland
irregularly coiled secretory portion
helically winding straighter duct portion
embedded in loose connective tissue
secretory portion with single layer of pale-staining low columnar serous
secretory cells
duct portion with double layer of cuboidal epithelial cells, characterized
by dark-staining nuclei
Apocrine Type
Largely resembling eccrine sweat glands in their histologic appearance,
apocrine sweat glands open into hair follicles instead of opening directly
onto the skin surface. Also, their secretory portion is rather large in com-
parison with their duct portion. The postpubertal secretory product of
apocrine sweat glands can undergo bacterial degradation, giving rise to
body odors. Sweat glands of the apocrine type are found chiefly in the
armpits and the pubic and perineal areas.
128 PDQ HISTOLOGY
Figure 99 Eccrine sweat gland.
Secretory portion Duct
Digestive System
The digestive system, represented by the tubular gastroin-
testinal tract (gut) and its various accessory digestive glands (Table 101),
is endoderm derived. The duct portions of its many accessory glands open
directly into the gut lumen. An obvious general histologic feature of the
tract is its regional specialization for nutrient processing, digestion, and
absorption. Even though constituent regions share a common wall structure
based on four concentric layers (Figure 101), additional features indicat-
ing specic functions allow each part to be recognized individually. Thus,
absorptive villi (nger-like projections) are found in the small intestine but
not elsewhere. Lymphoid tissue is more conspicuous in the ileum (Peyers
patches) and appendix than it is elsewhere. A mixed population of simple
columnar absorptive epithelial cells and mucus-secreting goblet cells char-
acterizes the intestine but not the esophagus or stomach.
Special arrangements are needed for continuous renewal of the lining
epithelial cells because they are subject to abrasion and exposure to acid or
damaging enzymes. In the intestine, the epithelial stem cells are protected
by being situated deep in crypts extending below the luminal surface. Gas-
tric epithelial cells are renewed every 4 to 6 days and villous epithelial cells
are renewed every 5 to 6 days. Rapid epithelial turnover and secretion of
protective mucus by gastric epithelium, goblet cells, and mucous glands are
key mechanisms in maintaining an uninterrupted epithelial barrier between
gut contents and the internal environment of the body.
10
129
Table 101
Digestive System
Oral cavity Ileum
Esophagus Appendix
Stomach Colon
Fundus Accessory glands
Pylorus Salivary glands
Duodenum Pancreas
Jejunum Liver
130 PDQ HISTOLOGY
Serosa or
adventitia
Outer layer
(longitudinal)
Inner layer
(circular)
Submucosa with
submucosal glands
Muscularis mucosae
(circular)
Lamina propria
Epithelium
Mucosa
Muscularis
externa
Figure 101 Layers of the gastrointestinal tract.
Figure 102 Esophagus.
Epithelium
Muscularis externa
Muscularis mucosae
Lamina propria
Submucosa
WALL ORGANIZATION
The four layers of the gut wall (see Figure 101) are as follows:
mucosa made up of epithelium(lining and glandular invaginations, eg,
gastric pits, crypts of Lieberkhn), lamina propria (loose connective
tissue extending into villi), and muscularis mucosae (smooth muscle
regulating size and independent movement of luminal folds)
submucosa (loose connective tissue extending into rugae and plicae;
contains medium-sized blood vessels and submucosal autonomic nerve
plexus; site of mucous glands in the duodenum and esophagus)
muscularis externa (major smooth muscle layers involved in peristalsis;
inner circular and outer longitudinal layers; contains myenteric auto-
nomic nerve plexus)
serosa or adventitia (covering of simple squamous visceral peritoneum =
serosa; connective tissue attachment, eg, to abdominal wall = adventitia)
ESOPHAGUS
Sections of esophagus (Figure 102) may be identified by the following
criteria:
four wall layers evident
stratified squamous nonkeratinizing epithelial lining, resistant to
abrasion
muscularis mucosae comparatively substantial in the lower part
thick muscularis externa made of skeletal muscle in the upper third,
smooth muscle in the lower third, and both types of muscle in the mid-
dle third
adventitia instead of a serosa
Chapter 10 Digestive System 131
132 PDQ HISTOLOGY
Figure 103 Fundic region of the stomach.
Gastric pit
Mucus-secreting simple
columnar epithelium
Chief cells Parietal cells
Figure 104 Pyloric region of the stomach.
Gastric pit Mucus-secreting simple
columnar epithelium
Pyloric gland Lamina propria
Muscularis mucosae
STOMACH
Two representative parts of the stomach are shown here: the body of the
stomach (designated the fundic region) and the pyloric region.
The fundic region (Figure 103) may be recognized by the following:
pale-staining mucus-secreting simple columnar surface epithelium
(without goblet cells)
the same epithelium dipping down into the lamina propria as gastric
pits (foveolae)
long fundic glands extending down through the lamina propria from
the pits, their bases reaching down to the substantial muscularis
mucosae
fundic glands containing pink parietal cells (which produce hydrochlo-
ric acid and secrete intrinsic factor, a glycoprotein required for vitamin
B
12
absorption), purple chief cells (which secrete pepsinogen and
lipase), enteroendocrine cells (which secrete hormones), and mucus-
secreting cells
large mucosal folds with submucosal cores (rugae)
Some additional features by which the pyloric region (Figure 104) may
be identied:
mucus-secreting pyloric glands, resembling mucus-secreting surface
epithelium (demonstrated here to advantage by periodic acidSchiff
staining, which colors glycoproteins magenta)
muscularis externa thickened at the pyloric sphincter
134 PDQ HISTOLOGY
Villus
Surface
epithelium
Crypt
Crypt
Lamina
propria
Muscularis
mucosae
Figure 105 Organization of a villus and crypts.
Figure 106 Crypt of the small intestine.
Goblet cells Absorptive cells
Enteroendocrine
cell
Paneth cells
SMALL INTESTINE
In the small intestine, the total absorptive area is increased by folds, villi,
and microvilli on the absorptive luminal surface of epithelial cells. Situated
between villi projecting into the lumen are crypts, straight simple tubular
mucosal glands invaginated into the lamina propria (Figure 105). Without
a clear understanding that in contrast to villi, which are tissue extensions
surrounded by luminal space, crypts are invaginations with a central space
and surrounding tissue, it can be difficult to tell which is which in sections.
In general, isolated proles surrounded by space (the tissue islands seen
on the luminal border in Figure 107) are villi, and those extending into
solid tissue are crypts.
In the lower part of crypts (Figure 106) it is possible to find the
following:
basal epithelial stem cells called crypt base columnar cells
undifferentiated dividing epithelial cells
mucus-secreting goblet cells
Paneth cells, characterized by large acidophilic secretory granules that
contain an antibacterial enzyme (lysozyme)
enteroendocrine cells of many kinds that secrete a variety of gastroin-
testinal hormones
136 PDQ HISTOLOGY
Figure 108 Ileum.
Villi
Crypts
Submucosal
lymphoid follices
Muscularis
externa
Figure 107 Duodenum.
Villi
Muscularis
mucosae
Submucosal
mucous glands
Muscularis
externa
Distinctive features of the duodenum (Figure 107) are the following:
relatively wide villi (seen in various planes of section) with crypts in the
lamina propria
conspicuous large mucus-secreting (Brunners) glands in submucosa,
generally extending through muscularis mucosae into lamina propria
(major source of protective alkaline mucus)
The ileum (Figure 108) is recognizable by the following:
narrower villi compared with duodenum
absence of large submucosal glands
conspicuous large aggregates of conuent lymphoid follicles (Peyers
patches) in submucosa, extending through muscularis mucosae into
lamina propria and raising surface epithelium between villi
The jejunumhas the villi, crypts, and basic wall structure typical of the
small intestine but lacks conspicuous submucosal glands and lymphoid
follicles.
138 PDQ HISTOLOGY
Figure 1010 Appendix.
Crypts Muscularis mucosae
Submucosal
lymphoid follices
Muscularis externa
Figure 109 Large intestine.
Crypt
Lamina propria
Muscularis
mucosae
LARGE INTESTINE
Distinguishing characteristics of the large intestine in general (Figure 109)
include the following:
absence of villi (often difficult to establish rmly unless a large area can
be searched for evidence of the tissue islands that would indicate villi)
higher proportion of goblet cells relative to columnar absorptive epithe-
lial cells
relatively long straight crypts containing numerous goblet cells along
with a few enteroendocrine cells
Sections of the appendix (Figure 1010) are additionally characterized by
the following:
relatively small irregularly shaped lumen
conspicuous confluent masses of lymphoid follicles in submucosa,
extending into lamina propria (enlarged appearance in children)
SEROUS ACCESSORY GLANDS
A few microscopic similarities exist between the parotid (a salivary gland of
the serous type) and the pancreas (another example of a compound serous
accessory digestive gland).
140 PDQ HISTOLOGY
Figure 1012 Pancreas.
Centro-acinar cell
Intralobular duct
Islet
Figure 1011 Parotid.
Intralobular
ducts
Serous
secretory units
The parotid (Figure 1011) shows the following features:
thick brous capsule, substantial septa, and obvious lobular organization
relatively numerous intralobular ducts, lined with simple cuboidal or
simple columnar epithelium
typical serous secretory units (acini)
In contrast, the pancreas (Figures 1012 and 1013) has the following:
a imsy investing sheath of loose connective tissue
relatively thin and delicate septa
large main and accessory ducts, both supported by substantial amounts
of brous connective tissue
relatively few intralobular ducts (mostly with simple cuboidal or even
lower epithelium; see Figure 1013)
central pale-staining centroacinar cells (proximal duct cells) included in
some of its serous acini, depending on their plane of section
irregularly shaped islets of Langerhans (pancreatic islets, the diffuse
endocrine component of this gland)
Figure 1013 Intralobular duct of the pancreas.
Acinus Intralobular duct
142 PDQ HISTOLOGY
Figure 1015 Central part of a liver lobule.
Sinusoids
Central vein Hepatocytes
Figure 1014 Liver.
Central vein
Portal area
LIVER
Histologic sections of the liver, the largest compound gland of all, appear
characteristically sponge-like. At low magnification (Figure 1014), it is
possible to distinguish the following:
portal areas containing a wide portal vein, hepatic artery or arteriole,
bile duct (with characteristic simple cuboidal epithelium), and com-
monly also a lymphatic
wide central veins
roughly hexagonal liver lobules, with portal areas indicating their
periphery and central veins indicating their middle
irregularly anastomosing rows of hepatocytes arranged in a radial pat-
tern that converges on the central vein
Further details seen with additional magnification (Figure 1015)
include the following:
anastomosing radiating rows of hepatocytes (representing cross-sections
of widely perforated plates having a thickness of a single hepatocyte)
thin-walled venous sinusoids (ie, pale-looking blood channels between
these rows that convey the blood delivered by the hepatic artery and
portal vein of a portal area toward a central vein)
cytologic details of hepatocytes
The liver acinus, an essentially functional concept, is not strictly recog-
nizable in routine histologic sections. However, its central axis is repre-
sented by an arteriolar branch of the hepatic artery, along with an accom-
panying venular branch of the portal vein, extending along a border of a
lobule from a portal area (Figure 1016). Its periphery is partly indicated by
the central veins on either side of this axis. Visualized in three dimensions,
the liver acinus resembles a hard-boiled egg lying on its side. Pursuing this
analogy, the highest concentrations of blood-borne nutrients and oxygen
reach the yolk area (termed zone 1). The lowest nutrient and oxygen levels
are found in the outer part of the egg white (termed zone 3).
144 PDQ HISTOLOGY
Central vein
Hepatic artery Portal vein
Portal
area
1 2 3
Figure 1016 Organization of a liver acinus.
Respiratory System
Extending posteriorly from the nasal cavities and
nasopharynx and descending into the thorax are the constituent parts of the
respiratory system (Table 111). This system is conventionally subdivided
into (1) its conducting portion, which extends as far as the terminal bronchi-
oles, and (2) its respiratory portion, which continues to the alveoli. A dichoto-
mously branching system of airways, the tracheobronchial tree, cleans and
conducts the incoming air and also provides the exit route. Most of the lining
layer is ciliated pseudostratied columnar epithelium with goblet cells, which
is specialized to produce and propel a particle-trapping coating of mucus. The
respiratory portion of the system lying distal to this represents the essential
site of gas exchange. Beyond respiratory bronchioles, its structural compo-
nents (alveolar ducts, alveolar sacs, and alveoli) represent air spaces separated
by intervening walls. A functionally important component of the tracheo-
bronchial tree, visceral pleura, and interalveolar walls is elastin, the passive
recoil of which is the chief factor expelling air in expiration. Besides provid-
ing a simple squamous mesothelial (serosal) gliding surface with underlying
(subserosal) dense broelastic tissue, the visceral pleura contains a plexus of
pulmonary lymphatics draining, by way of interlobular septal lymphatics,
toward the hilum of the lung. Another functionally important constituent of
the respiratory portion is its extensive population of alveolar macrophages.
When these phagocytes engulf inhaled particles settling beyond the mucus
coating of the airways, they subsequently become trapped in the mucus layer
and are swept along with it toward the pharynx.
11
145
Table 111
Respiratory System
Nasal cavities and nasopharynx Bronchioles
Larynx and epiglottis Alveolar ducts and alveoli
Trachea Pleura
Bronchi
146 PDQ HISTOLOGY
Figure 111 Trachea.
Pseudostratied
epithelium
Mixed submucosal
glands
Cartilage
Fibrous perichondrium
Figure 112 Bronchus.
Cartilage
Mixed submucosal
gland
TRACHEA AND BRONCHI
Sections of the trachea (Figure 111) may be recognized by the following:
C-shaped hyaline cartilages (keeping this major airway open when the
neck is bent)
ciliated pseudostratied columnar epithelium with goblet cells (respon-
sible for maintaining ascending mucus coating)
mixed submucosal glands (providing supplementary watery mucus)
smooth (trachealis) muscle (interconnecting posterior ends of C-shaped
cartilages)
dense broelastic perichondrium (extending as cartilage interconnections)
Sections of the intrapulmonary bronchi (Figure 112) appear fairly
similar to those of the trachea, except that their
lumen is smaller than that of the trachea,
supporting cartilages are more irregular in shape,
mixed submucosal glands are smaller than those of the trachea, and
adventitia is attached to the surrounding lung tissue.
Crisscrossing smooth muscle bundles lie in the submucosal connective
tissue between the epithelium and the cartilage, but because they follow hel-
ical courses, they are not always easy to recognize.
Chapter 11 Respiratory System 147
148 PDQ HISTOLOGY
Figure 113 Large bronchiole.
Smooth muscle
Ciliated pseudostratied
columnar epithelium
Figure 114 Small bronchiole.
Simple cuboidal epithelium
BRONCHIOLES
Larger bronchioles such as a preterminal bronchiole (Figure 113) have the
following characteristics:
luminal diameter < 1 mm
smooth muscle arranged as helical bundles crisscrossing deep to the
elastic lamina propria (excessive tonus of this muscle can lead to expi-
ratory difficulties in asthmatic patients)
ciliated pseudostratied columnar epithelium with goblet cells (con-
tinuation of the mucociliary escalator)
adventitia attached to surrounding lung tissue (bronchiolar walls pulled
outward by tension in interalveolar walls when alveoli inate)
exible walls unsupported by cartilages
submucosal glands are absent
Additional features of the smaller bronchioles (Figure 114) are as follows:
decreased total wall thickness and smaller luminal diameter
smooth muscle even less conspicuous
simple cuboidal epithelium, with some cells nonciliated
no goblet cells
150 PDQ HISTOLOGY
Figure 116 Respiratory bronchiole leading into alveolar ducts.
Respiratory
bronchiole
Alveoli Alveolar sac
Alveolar ducts
Figure 115 Respiratory bronchiole.
Alveolar duct Alveolar sac
Respiratory bronchiole
RESPIRATORY BRONCHIOLES AND ALVEOLAR AIR SPACES
Respiratory bronchioles represent the last few orders of branching of the
tracheobronchial tree. It is usually easier to recognize them from their dis-
tal position in the dichotomously branching air passages (Figure 115) than
from the small numbers of alveoli extending from their walls. Alveolar ducts
(Figure 116; see also Figure 115) are essentially cylindrical air spaces
extending distally from the ends of the nal order of respiratory bronchi-
oles. The more distal alveolar sacs represent compound air spaces that are
smaller than alveolar ducts yet larger than alveoli. Their perimeters consist
of bubble-like alveoli that bulge out from the sac (see Figures 115 and
116). The ultimate structural and functional unit of gas exchange, the alve-
olus, is recognizable in lung sections as the smallest white-looking intra-
pulmonary air space. Most of the very cellular tissue lying between air
spaces in this section represents juxtaposed walls of collapsed alveoli
because it is a section of collapsed lung.
152 PDQ HISTOLOGY
Figure 117 Alveolar sacs.
Alveolar sac
Alveolus
Capillary in
interalveolar wall
Figure 118 Cell types in the interalveolar wall.
Alveolar macrophage
Type I
pneumocyte
Erythrocytes in
alveolar capillary
Type II
pneumocyte
Additional histologic features of the respiratory portion appearing to
advantage in sections of expanded lung (Figures 117 and 118) include the
following:
thin interalveolar walls (septa)
pulmonary blood capillaries (lymphatics present in the bronchovascu-
lar connective tissue but not the interalveolar walls)
at squamous epithelial cells (type I pneumocytes) lining the alveoli,
recognizable by their at nuclei
rounded secretory epithelial cells (type II pneumocytes) secreting
pulmonary surfactant, recognizable by their slightly foamy-looking
cytoplasm
rounded alveolar macrophages, recognizable by their content of phago-
cytosed particles (particularly in smokers lungs) but generally hard to
distinguish from type II pneumocytes in light microscopic sections
unless their position is supercial
elastic bers (demonstrable in interalveolar walls when a special elastin
stain is used)
Interalveolar walls are essentially sandwich-like in construction (Figure
119):
type I pneumocytes, with occasional interposed type II pneumocytes,
constitute the slices of bread;
a continuous basement membrane underlying these two types of
epithelial cells represents the butter;
a flat plexus of anastomosing pulmonary capillaries (ie, rolled-up
endothelial cells with their surrounding basement membrane) repre-
sents a major part of the lling;
an extensible supportive framework of interstitial fibers (elastic and
reticular) represents another part of the lling;
motile alveolar macrophages, scattered over the alveolar surfaces, cor-
respond to crumbs adhering to the at outer surfaces of the sandwich.
154 PDQ HISTOLOGY
Basement
membrane
Type I
pneumocyte
Type II
pneumocyte
Alveolar
macrophage
Endothelial lining
cell of capillary
Interstitial
fibers
Figure 119 Organization of the interalveolar wall.
Urinary System
Represented by the kidneys, ureters, urinary bladder, and
urethra, the urinary system is responsible for producing and storing urine.
Internal organization is most complex in the kidneys; the other components
are all characterized by a central lumen and an appropriate wall structure.
KIDNEYS
The kidneys are largely composed of structural and functional units termed
nephrons that form urinary filtrate continuously from circulating blood
plasma and then concentrate, process, and deliver it to the collecting duct
system. The histologic basis for distinguishing the renal cortex from the
renal medulla is the route taken by the various parts of the nephron and the
collecting tubules and ducts into which it empties. The renal cortex repre-
sents the unique site of the renal corpuscle (the proximal ltration unit),
convoluted segments of the nephron (its proximal and distal convoluted
tubules), and medullary rays (ray-like cortical areas containing straight seg-
ments of several nephrons along with cortical collecting tubules). The renal
medulla is characterized by its content of straight segments of nephrons
(thick- and thin-walled parts of loops of Henle) and associated straight
medullary collecting tubules that merge and drain into main collecting
ducts opening onto the renal papillae at the hilum of the kidney. Each con-
ical medullary pyramid, with its covering cap of cortex, represents a kidney
lobe. The human kidney is regarded as multilobar because it contains up to
18 lobes. Between kidney lobes, renal cortex extends down into the renal
medulla as renal columns. A convenient way to substantiate the position of
the corticomedullary boundary is to look for arcuate vessels fanning out
distally from the interlobar blood vessels at this level like the ribs of an
umbrella.
12
155
156 PDQ HISTOLOGY
Figure 121 Renal cortex.
Interlobular blood vessels
Cortical labyrinth
Renal corpuscles
Figure 122 Lobule of renal cortex.
Interlobular blood vessels
Medullary ray
Some distinctive histologic features of the renal cortex (Figure 121) are
as follows:
renal corpuscles scattered throughout the cortex
cortical labyrinth (multiple cuts through proximal and distal convo-
luted tubules)
interlobular blood vessels, afferent and efferent arterioles
medullary rays (predominantly longitudinal cuts through straight
tubules resembling the medullary tubules)
brous external capsule may be included in the section, immediately
adjacent to outer cortex (not shown)
The following additional features characterize the cortical lobule (Figure
122):
the lateral borders of the lobule are indicated by the positions of the
interlobular blood vessels (interlobular arteries or veins);
the central core of the lobule is made up of longitudinal or oblique cuts
through parallel straight tubules, that is, it is a medullary ray.
Although cortical lobules are recognizable in favorable planes of sec-
tion, they are not always easy to discern.
Chapter 12 Urinary System 157
158 PDQ HISTOLOGY
Renal corpuscle
Proximal convoluted
tubule
Distal convoluted
tubule
Collecting
duct
Site of juxtaglomerular
apparatus
Afferent arteriole
Glomerulus
Loop of Henle
Figure 124 Parts of the nephron with associated collecting tubules.
Figure 123 Renal medulla.
Sections of the renal medulla (Figure 123) typically contain longitu-
dinal or oblique cuts of
thick- and thin-walled parts of loops of Henle,
medullary collecting tubules and papillary ducts (main collecting ducts),
long straight capillaries known as vasa recta, and
traces of loose connective tissue (comprising the site of an interstitial
osmotic gradient).
Each nephron is made up of a renal corpuscle, proximal convoluted
tubule, loop of Henle, and distal convoluted tubule (Figure 124). The com-
ponent segments of the nephron have distinctive histologic appearances
and are readily distinguishable where they have been cut in cross-section
(see Figure 129).
160 PDQ HISTOLOGY
Figure 126 Renal corpuscle seen in more detail.
Capsular space
Podocyte
Glomerular
capillary
Glomerular
basement membrane
Parietal (capsular)
epithelium
Proximal convoluted
tubule (tubular pole)
Vascular pole
Figure 125 Renal corpuscles.
Renal corpuscle
Arterioles
Renal Corpuscle
The blood being ltered by renal corpuscles comes from their afferent arte-
riole and leaves by way of their efferent arteriole (Figure 125). It is seldom
possible to tell whether a given arteriole lying in close association with a
renal corpuscle is afferent or efferent. Even though the glomerulus-
associated end of the afferent arteriole is characterized by the presence of
juxtaglomerular (JG) cells in its media (see Figure 127), these periodic
acidSchiff (PAS)-positive secretory cells are present in small numbers and
therefore difficult to nd.
Distinctive features by which renal corpuscles (Figure 126) may be
recognized are as follows:
parietal (capsular) epitheliuma simple squamous epithelium with a
supporting basement membrane that appears to advantage in PAS-
stained sections
visceral (glomerular) epitheliumrepresented by scattered cell bodies
and nuclei of podocytes (the pedicels or foot processes of which are
attached to the glomerular basement membrane)
central tuft of glomerular capillaries
glomerular basement membrane (the renal ltration barrier compo-
nent that minimizes exit of plasma macromolecules), also appearing to
advantage in PAS-stained sections
capsular space into which urinary ltrate passes
vascular pole (representing the joint attachment site of the afferent and
efferent arterioles)
tubular pole (representing the proximal end of the proximal convoluted
tubule)
162 PDQ HISTOLOGY
Figure 128 Medullary ray.
Medullary ray
Efferent
arteriole
Afferent
arteriole
JG cells
Lacis cells
Macula
densa
Blood from
glomerulus
Distal
convoluted
tubule
Blood to
glomerulus
Urinary filtrate to
collecting tubule
Figure 127 Organization of the juxtaglomerular apparatus (see Figure 124 for its orientation).
Juxtaglomerular Apparatus
JG cells are the principal effector component of the juxtaglomerular appa-
ratus (Figure 127), the major features of which are as follows:
its juxtaglomerular positionlocated where the distal convoluted
tubule lies in apposition to the afferent and efferent arterioles at the vas-
cular pole of the renal corpuscle
granule-containing JG cells (modied smooth muscle cells) in media of
afferent arteriole, which respond to decreased stretch of the vessel wall
by releasing renin, the enzyme that produces angiotensin from its pre-
cursor; angiotensin raises blood pressure
apposed macula densa (densely nucleated spot in the epithelial wall of
the distal convoluted tubule), which detects falls in the sodium and
chloride contents of urinary filtrate and triggers renin release by JG
cells
lacis (extraglomerular mesangial) cells, interposed between macula
densa cells and JG cells and presumed to be involved in cell-to-cell
signaling
Renal Tubules
The following are readily recognizable characteristics of a medullary ray
(Figure 128):
flanked by a combination of cortical labyrinth and renal corpuscles
(indicative of renal cortex)
straight thick-walled parts of loops of Henle (segments of supercial
nephrons)
straight cortical collecting tubules (recognizable by the fact that the
lateral margins of their cuboidal epithelial lining cells are relatively
distinct)
164 PDQ HISTOLOGY
Figure 1210 Proximal and distal convoluted tubules.
Proximal convoluted tubule
Distal convoluted tubule
Proximal convoluted
Distal convoluted
Thin-walled loop of Henle
Collecting tubule
Figure 129 Renal tubules in cross-section.
In routine histologic sections, the straight tubules characterizing
medullary rays and the medulla are seldom totally represented as longitu-
dinal sections. The various kidney tubules, however, are also individually
recognizable in cross- or oblique section (Figure 129). Cortical proximal
and distal convoluted tubules are commonly cut in these two planes.
Proximal and distal convoluted tubules (Figure 1210) are typically dis-
tinguishable by the following criteria:
proximal convoluted tubule longer than distal convoluted tubule
(therefore seen with greater frequency in sections)
epithelial lining cells of the proximal convoluted tubule larger, pinker,
and wider than those of the distal convoluted tubule
epithelial lining cells of the proximal convoluted tubule characterized
by a distinct striated luminal border (abundant absorptive microvilli
with an associated thick PAS-positive cell coat), whereas those of the
distal convoluted tubule lack such a border
166 PDQ HISTOLOGY
Figure 1211 Ureter.
Smooth muscle Lamina propria
Transitional epithelium
Figure 1212 Urinary bladder.
Lamina propria
Transitional
epithelium
Smooth
muscle
bundles
URETERS
Sections of a ureter (Figure 1211) typically show the following features:
tubular organization
distinctive stellate lumen
deeply folded transitional epithelial lining (exclusive to the urinary
tract)
fibroelastic lamina propria and adventitia (with resulting luminal
folding)
muscular coat made up of inner longitudinal and outer circular layers
of smooth muscle (reverse arrangement of that found in the digestive
tract)
URINARY BLADDER
The urinary bladder (Figure 1212) may generally be recognized by the
following:
wall structure less distinctly organized (sac-like, not tubular)
transitional lining epithelium (indicative of urinary tract)
comparatively shallow mucosal folds
broelastic lamina propria and adventitia
smooth muscle layers, the respective orientations of which are hard to
distinguish
Endocrine System
The glands generally included in histology courses as repre-
sentative of the endocrine system are listed in Table 131 (the ovaries are
described in Chapter 14, Female Reproductive System, and the testes in
Chapter 15, Male Reproductive System). In contrast to epithelial glands of
the exocrine type, the endocrine glands are not provided with any ducts.
Instead, their target cell-specific secretory products (hormones) pass
directly into capillaries, enabling these products to reach their target cells by
way of the bloodstream. Each hormone (or its precursor) is synthesized
continuously. Many hormones then become stored in the cytoplasm until
their release is suitably triggered. Thyroid and steroid hormones, however,
are lipid soluble; hence, they diffuse out of secretory cells as soon as they are
made. Whereas the water-soluble hormones bind to specific hormone
receptors on the surface of their target cells, thyroid and steroid hormones
diffuse into cells and bind to specic intracellular receptor proteins in their
target cells. The complexes thus formed specically promote the transcrip-
tion of appropriate genes. The secretory cells of most endocrine glands are
arranged as anastomosing cords, columns, or clumps lying adjacent to wide
fenestrated capillaries and supported by a brous capsule and connective
tissue trabeculae.
13
169
Table 131
Endocrine Organs
Pituitary Thyroid
Anterior lobe Parathyroids
Posterior lobe Pancreatic islets
Adrenal Ovaries
Cortex Testes
Medulla
170 PDQ HISTOLOGY
Anterior lobe
Infundibular stalk
Posterior lobe
Pars intermedia
Figure 131 Parts of the pituitary.
Figure 132 Pituitary seen under scanning power.
Anterior lobe
Posterior lobe
PITUITARY
The pituitary (hypophysis) is a relatively small endocrine gland, roughly
ovoid in shape, that is connected to the hypothalamus by its infundibular
stalk (Figure 131). Functionally and histologically, the anterior and posterior
lobes of the pituitary are dissimilar. Situated between these lobes is the pars
intermedia, poorly represented in the human pituitary and hard to recognize
except for a few colloid-lled cysts with negligible functional signicance. The
anterior and posterior lobes may both be represented in vertical or horizon-
tal planes of section. Secretory epithelial cells of the anterior lobe, a deriva-
tive of the oral ectoderm, secrete the hormones listed in Table 132. Hypo-
thalamic hormones regulating the release of these hormones are included in
this table. The posterior lobe (pars nervosa), derived from a downgrowth of
the brain (the oor of the third ventricle), is entirely composed of nervous tis-
sue. It is the exclusive site of release of two hypothalamic peptide hormones:
oxytocin and vasopressin (antidiuretic hormone [ADH]).
In a horizontal section of the pituitary (Figure 132), the anterior larger
lobe, with its brightly stained secretory epithelial cells, is readily distin-
guishable from the pale-staining nervous tissue that characterizes the pos-
terior lobe.
Chapter 13 Endocrine System 171
Table 132
Anterior Pituitary Hormones
Release-Regulating Hormones
Hormones (Hypothalamic)
Adrenocorticotropic Corticotropin releasing
Follicle stimulating Gonadotropin releasing
Growth Growth hormone releasing
Growth hormone inhibiting
(somatostatin)
Luteinizing Gonadotropin releasing
Melanocyte stimulating None established
Prolactin Prolactin releasing
Prolactin inhibiting (dopamine)
Thyroid stimulating Thyrotropin releasing
172 PDQ HISTOLOGY
Figure 133 Anterior and posterior pituitary.
Posterior lobe
Anterior lobe
Figure 134 Anterior pituitary.
Acidophils Fenestrated capillary
Basophils Chromophobes
Juxtaposition of a mass of nervous tissue (representing the posterior lobe)
and a large mixed population of brightly stained secretory epithelial cells, some-
what haphazard in arrangement and without ducts (representing the anterior
lobe), is reliably indicative of the pituitary gland (Figure 133). The pars inter-
media may or may not be discernible, depending mainly on the plane of section.
The posterior lobe contains the following:
hypothalamohypophyseal tracts of pale-staining unmyelinated axons
belonging to neurosecretory neurons situated in the supraoptic and
paraventricular nuclei of the hypothalamus (the major respective
sources of ADH and oxytocin)
small pale-staining neuroglia (pituicytes) represented by their dark-
staining nuclei
capillaries of the fenestrated type, into which ADH and oxytocin are
released from axon terminals of the neurosecretory neurons
Characteristic features of the anterior pituitary, suitably stained (eg,
with Gomoris stain) to show its acidophils and basophils and viewed at
higher magnication (Figure 134), are as follows:
groups of chromophils of two main kindsred-staining acidophils and
blue-staining basophils
groups of chromophobesunstained, smaller, quiescent, or degranu-
lated secretory cells
no ducts
abundant wide capillaries of the fenestrated type
The various pituitary acidophils and basophils, together with the hor-
mones they secrete, are listed in Table 133.
Table 133
Anterior Pituitary Hormone Sources
Cells and Cytoplasmic Staining Secreted Hormones
Somatotrophs (acidophil) Growth hormone
Mammotrophs (acidophil) Prolactin
Mammosomatotrophs (acidophil) Prolaction and growth hormone
Corticotrophs (basophil) Adrenocorticotropic hormone (and
putative source of melanocyte-
stimulating hormone)
Corticothyrotrophs (basophil) Adrenocorticotropic hormone and
thyroid-stimulating hormone
Corticogonadotrophs (basophil) Adrenocorticotropic hormone,
follicle-stimulating hormone, and
luteinizing hormone
Thyrotrophs (basophil) Thyroid-stimulating hormone
Gonadotrophs (basophil) Follicle-stimulating hormone and
luteinizing hormone
174 PDQ HISTOLOGY
Capsule
Zona
glomerulosa
Zona
fasciculata
Fenestrated
capillary
Zona
reticularis
Medullary
chromaffin
cells
Figure 135 Organization of an adrenal gland.
Figure 136 Adrenal seen under scanning power.
Zona reticularis
Capsule Medulla
ADRENALS
The adrenals (suprarenals) are distinctively shaped endocrine glands
attached to the superomedial borders of the kidneys. Each adrenal gland has
a mesoderm-derived cortex with a neural crestderived medulla that is
functionally separate. Compared with the medulla, the cortex is more elab-
orate in organization (Figure 135).
The following steroid hormones are produced by the adrenal cortex
(adrenocorticosteroids):
aldosterone, secreted by the cells in the glomerulosa (outermost zone of
the adrenal cortex)
cortisol and weak androgens, secreted jointly by the cells in the fascicu-
lata and reticularis (middle and innermost zones of the adrenal cortex)
The following catecholamine hormones are produced by the adrenal
medulla:
epinephrine, secreted by many of the medullary chromaffin cells
norepinephrine, secreted by the remainder of the medullary chromaf-
n cells
Features generally permitting sections of the adrenals to be recognized
under scanning power (Figure 136) include the following:
characteristically folded, essentially triangular outline (indicating the
overall shape of the capsule)
bright-staining zona reticularis, demarcating the border between the
cortex and the medulla
paler-staining zona fasciculata, external to zona reticularis
relatively uniform-looking pale pink medulla with large medullary
veins that commonly have the appearance of wide empty spaces
176 PDQ HISTOLOGY
Figure 137 Adrenal cortex.
Zona fasciculata Zona glomerulosa Capsule
Zona reticularis Medulla
Figure 138 Adrenal cortex seen in more detail.
Capsule
Glomerulosa
Fasciculata
Reticularis
Medulla
Working inward, further features of adrenal sections that may be dis-
tinguished at higher magnication (Figures 137 and 138) are as follows:
broelastic thick external capsule
narrow outer zone containing spherical clusters of small purple secre-
tory cells, with closely associated fenestrated capillariesthe zona
glomerulosa of the adrenal cortex
wide middle zone containing radial columns, approximately one cell
thick, of larger pink secretory cells, with closely associated long straight
fenestrated capillariesthe zona fasciculata of the adrenal cortex
narrow inner zone containing anastomosing cords of small, pale purple
secretory cells arranged as a network, with closely associated fenestrated
capillariesthe zona reticularis of the adrenal cortex
central mass of slightly larger, relatively pale-staining, round cells that
include chromaffin cells of two types (either epinephrine secreting or
norepinephrine secreting) and occasional basophilic sympathetic gan-
glion cells, together with wide branches of the medullary vein and fen-
estrated capillariesthe adrenal medulla
178 PDQ HISTOLOGY
Figure 1310 Thyroid (periodic acidSchiff stain).
Figure 139 Thyroid (hematoxylin and eosin stain).
THYROID
The bilobed thyroid gland, lying anterior and lateral to the trachea just infe-
rior to the larynx, incorporates two independent populations of hormone-
secreting cells. The thyroid follicular epithelial cells, derived from endo-
derm, produce thyroid hormone (thyroxine plus triiodothyronine). The
parafollicular (C) cells, derived from neural crest, produce calcitonin.
A characteristic histologic feature of the thyroid gland is the thyroid
follicle, a cyst-like spherical structure lined by simple cuboidal thyroid
follicular epithelial cells and filled with extracellular thyroglobulin, the
macromolecular stored precursor of thyroid hormone (Figure 139).
Thyroglobulin is demonstrated to advantage by periodic acidSchiff staining
(Figure 1310) because it is a glycoprotein. On release through exocytosis
from the epithelial lining cells of the follicle, the secreted thyroglobulin
becomes iodinated extracellularly. It is subsequently ingested by these cells
and submitted to intracellular proteolysis, releasing thyroid hormone.
An outer fascial sheath surrounds the external connective tissue capsule
of the thyroid. The minimal connective tissue stroma is profusely supplied
with fenestrated capillaries.
180 PDQ HISTOLOGY
Figure 1311 Thyroid follicle.
Follicular
epithelial cell
Figure 1312 Parafollicular cells of the thyroid.
Follicular
epithelial cell
Parafollicular
cell
Seen here at higher magnication, thyroid follicular epithelial cells consti-
tute a simple cuboidal epithelium bordering on the stored thyroglobulin (Fig-
ure 1311). In a less active thyroid, these cells can appear low columnar (Fig-
ure 1312). In the course of tissue processing, the thyroid tends to undergo
uneven shrinkage, accounting for the wide split extending from the bottom left
to the upper right in Figure 1311 and the majority of empty-looking spaces
and thyroglobulin bubbles commonly seen in thyroid sections.
The slightly larger round cells with central nuclei that lie between fol-
licles (see Figure 1312) are parafollicular (C) cells. These cells lie internal
to the follicular basement membrane but do not border on the thyroglobulin
(Figure 1313). They release calcitonin into adjacent fenestrated capillaries.
Basement
membrane
Thyroid
follicle
Parafollicular
cells
Figure 1313 Position of parafollicular cells in the thyroid.
182 PDQ HISTOLOGY
Figure 1315 Pancreatic islet.
Acinus Islet
Figure 1314 Parathyroid.
Chief cells
Oxyphil cells
PARATHYROIDS
Distinctive features of the parathyroids (Figure 1314), the small endo-
derm-derived endocrine glands lying on the posterior surface of the thyroid
that secrete parathyroid hormone, are as follows:
large areas of relatively small round chief cells with an intensely stained small
central nucleus, representing the cells that secrete parathyroid hormone
smaller groups of larger, pinker cells with a fairly similar nucleus
(oxyphil cells) that rst appear at puberty and are assumed to represent
retired or nonsecreting chief cells
numerous fenestrated capillaries
no ducts
PANCREATIC ISLETS
Pancreatic islets (islets of Langerhans), representing the diffuse endoderm-
derived endocrine component of the pancreas (Figure 1315), may be rec-
ognized by the following:
proximity to serous acini (a readily recognizable pancreatic exocrine
component)
relatively pale staining of the cytoplasm of their constituent cells in
hematoxylin and eosinstained sections
irregular shape, size, and distribution of the islets
A () cells producing glucagon can be distinguished from B () cells
producing insulin by specially devised staining methods (eg, Gomori). Also
present are D () cells secreting somatostatin, F cells secreting pancreatic
polypeptide, and fenestrated capillaries.
Female Reproductive System
This chapter deals with the parts of the female reproductive
system listed in Table 141 and describes how the histologic appearance of
some of them is affected by physiologic variation in the ovarian steroid hor-
mone levels. From the age of puberty until menopause, the ovaries consti-
tute the exclusive site of cyclic maturation of ovarian follicles, monthly
release of the secondary oocyte formed at the time of ovulation, and phasic
secretion of ovarian hormones. The secondary oocyte liberated in the course
of ovulation enters a uterine tube, where it may become fertilized. Peristaltic
waves of contraction, produced by smooth muscle of the tubal wall, convey
the liberated germ cell to the uterine body. The blastocyst developing from
a fertilized ovum generally implants in the endometrium (uterine mucosa)
at some posterior site. Estrogen and progesterone, the ovarian steroid hor-
mones, are essential for appropriate growth and functional activity of the
endometrium, myometrium (uterine smooth muscle), cervical mucous
glands, and breast and also affect the cytologic appearance of the vagina.
OVARIES
The ovarian cortex is characterized by its distinctive content of ovarian
follicles representing various stages of maturation, embedded in a swirly-
looking stroma. Just under the simple cuboidal covering epithelium lies a
relatively brous layer termed the tunica albuginea. The ovarian medulla
contains similar broblast-like stromal cells and is the site of substantial
ovarian blood vessels, some of which are fairly convoluted.
14
185
Table 141
Female Reproductive System
Ovaries Vagina
Uterus Breast
Uterine tubes (fallopian tubes)
186 PDQ HISTOLOGY
Corpus
luteum
Primary
follicle
Primordial
follicle
Early secondary
follicle
Corpus albicans
Atretic follicle
Secondary
follicle
Theca
Mature follicle
Simple
cuboidal
epithelium
Cortex
Medulla
Figure 141 Maturation stages of ovarian follicles.
Figure 142 Primary ovarian follicles.
The histologic appearance of the ovaries depends on both the species
represented and the current menstrual phase at the time of xation. Com-
posite drawings showing the maturation stages of ovarian follicles (Figure
141) are based on all of their possible histologic appearances. Only a rep-
resentative selection is found in any given section. Furthermore, mature
follicles may appear surprisingly large after they accumulate follicular uid,
and several follicles may be found to be approaching full maturity if, as is
often the case, the ovary was obtained from some animal that produces lit-
ters rather than single offspring. All stages except the corpus luteum are
generally present during the proliferative (estrogenic, follicular) phase. The
corpus luteum appears as an added feature in the secretory (progestational,
progravid) phase and, in particular, in the rst trimester of a pregnancy,
when it becomes very large. Follicles may degenerate (undergo atresia) at
any stage in their maturation. Atretic and ruptured (ovulated) follicles
become replaced by a long-lasting brous scar called a corpus albicans.
The early stages of ovarian follicular maturation resulting from ovarian
stimulation by follicle-stimulating hormone (FSH) (Figure 142) are clas-
sied according to the histologic appearance of the ovarian follicular (gran-
ulosa) cells surrounding the primary oocyte:
primordial folliclessingle layer of low cuboidal (almost squamous)
unstimulated follicular cells
primary folliclestwo or more layers of cuboidal to columnar follicu-
lar cells but no antrum
secondary folliclesmany layers of columnar follicular cells with
antrum containing follicular uid
mature (tertiary, graafian) folliclecyst-like structure with primary
oocyte attached to follicular wall at cumulus oophorus; surrounding
stroma constitutes theca
Chapter 14 Female Reproductive System 187
188 PDQ HISTOLOGY
Figure 144 Mature ovarian follicles.
Theca Antrum
Cumulus oophorus Zona pellucida
Figure 143 Atretic ovarian follicles.
Microscopic signs of follicular atresia (Figure 143) include the following:
loss, disorganized arrangement, or morphologic deterioration of the
follicular cells
degenerative changes of the primary oocyte, for example, loss of its
zona pellucida (external glycoprotein covering), loss of nuclear staining,
indications of nuclear disruption, invasion of the follicle by leukocytes,
such as neutrophils
presence of broblasts but not scar tissue inside the follicle
Mature follicles (Figure 144) show the following features:
substantial size and spherical shape
sizable antrum lled with follicular uid
numerous follicular epithelial (granulosa) cells (producing estrogen)
vascular theca interna (releases the androgen substrate used in estrogen
production)
brous theca externa
cumulus oophorus containing primary oocyte
substantial pink zona pellucida present on oocyte surface
190 PDQ HISTOLOGY
Figure 146 Corpus luteum.
Theca lutein cells
Granulosa lutein cells
Figure 145 Corpus albicans.
Corpus albicans
A corpus albicans (Figure 145) may generally be recognized by its
essentially round to ovoid shape and rather irregular periphery merg-
ing with the stroma,
pink hyalinized collagen content, and
residual broblast nuclei (also showing that scarring has occurred).
Histologic features of a corpus luteum (Figure 146) include the
following:
during the rst part of the secretory phase of a menstrual cycle: a diam-
eter similar to or smaller than that of a ripening ovarian follicle
during the rst trimester of a pregnancy: a large diameter (generally
peaking at about half the ovarian volume)
relatively pale-staining granulosa lutein cells (producing both estrogen
and progesterone)
slightly smaller and pinker theca lutein cells (producing additional
progesterone and the androgen substrate used in estrogen production)
connective tissue septa, extending into its interior from the theca and
giving it a somewhat lobular appearance
persisting remnants of the intrafollicular blood clot formed at ovulation
192 PDQ HISTOLOGY
Figure 148 Uterine body.
Myometrium
Endometrium
Early proliferative Late proliferative Late secretory
Spiral
arteries
Endometrial
glands
Figure 147 Endometrium at representative phases of the menstrual cycle.
UTERUS, ENDOMETRIUM, AND MENSTRUAL CYCLE
The histologic appearance of the endometriumreects cyclical variation in
the circulating levels of estrogen and progesterone (Figure 147). Following
4 to 5 days of partial endometrial loss (the menstrual phase of the cycle),
rising estrogen levels from FSH-stimulated follicular granulosa cells pro-
mote mitosis and endometrial regeneration (proliferative, estrogenic, or
follicular phase). After ovulation, rising progesterone and estrogen levels
from luteinizing hormone and FSH-stimulated granulosa lutein cells pro-
mote hypertrophy and secretory activity of endometrial glands in prepara-
tion for implantation (secretory, progestational, or progravid phase). Sub-
sequent fall of the progesterone and estrogen levels leads to ischemic
necrosis of the endometrium (ischemic phase).
Sections of the uterine body (Figure 148) may be recognized by the
following:
hollow pear shape, with wide central lumen
substantial mucosa termed the endometrium, characterized by simple
tubular glands that appear straight at this menstrual phase
thick multilayered muscular wall termed the myometrium (uterine
smooth muscle)
thin external serosa
194 PDQ HISTOLOGY
Figure 1410 Secretory endometrium.
Sacculated
endometrial glands
Figure 149 Proliferative endometrium.
Straight
endometrial glands
Endometrium observed fairly late in the proliferative phase (Figure
149) may be recognized by the following:
simple columnar lining epithelium
predominantly straight or slightly tortuous long endometrial glands
deeply invaginated into a relatively cellular lamina propria
a few mitotic gures in the glandular epithelium or lamina propria
underlying smooth muscle (myometrium)
Endometrium that is approaching the end of the secretory phase (Fig-
ure 1410) typically shows the following additional features:
hypertrophy and sacculation of the endometrial glands (giving grazing
sections a ladder-like appearance)
pale-stained glandular secretory cells with a ragged luminal border
indicative of glycogen secretion
empty-looking spaces in the lamina propria, representing sites of inter-
cellular accumulation of tissue uid (edema spaces)
elongated spiral arteries in the lamina propria
196 PDQ HISTOLOGY
Figure 1411 Uterine tube.
Lamina propria
Ciliated simple
columnar epithelium
Smooth muscle
Figure 1412 Vagina.
Stratied squamous
nonkeratinizing epithelium
Lamina propria
UTERINE TUBE
Sections through the ampullary region of a uterine tube (Figure 1411)
may be recognized by the following:
elaborately folded mucosa characterized by substantial primary, sec-
ondary, and tertiary folds
loose ordinary connective tissue (lamina propria) extending into each
fold
simple columnar lining epithelium, predominantly ciliated, with scat-
tered groups of nonciliated secretory cells
fairly substantial muscular wall consisting of an inner circular layer and
an outer longitudinal layer of smooth muscle
external serosal covering
Care should be taken not to misidentify a section of this part of a uter-
ine tube as a section of seminal vesicle bearing a supercial resemblance to
it (see Figure 159).
VAGINA
Histologic features of the vagina (Figure 1412) are as follows:
substantial stratied squamous nonkeratinizing lining epithelium char-
acterized by pale staining owing to its content of stored glycogen and
lipid (estrogen effect)
thick, vascular broelastic lamina propria containing numerous small
veins and venules
smooth muscle coat consisting of inner circular and outer longitudinal
bundles
brous adventitia
198 PDQ HISTOLOGY
Figure 1414 Lobule of resting breast.
Intralobular connective tissue
Interlobular connective tissue
Figure 1413 Resting breast.
Intralobular connective
tissue
Interlobular connective tissue
Intralobular ducts
BREAST
Breast tissue that is not currently lactating is commonly described as rest-
ing breast. At low magnication (Figure 1413), resting breast is recogniz-
able by the following criteria:
small groups of ducts showing a somewhat scattered distribution indica-
tive of spherical lobules (permitting their distinction from sweat glands)
a relatively brous kind of connective tissue around the lobules that has
a variable content of adipocytes (not shown)
Additional features conrmed at higher magnication (Figure 1414)
are as follows:
epithelial lining of some larger ducts consists of two layers of cuboidal
cells; smaller ducts lined with a single layer
no secretory component (unlike sweat glands or lactating breast)
intralobular connective tissuerelatively cellular; interlobular connec-
tive tissuerelatively brous
Distinctive histologic features are acquired by the lactating breast
(Figure 1415) in the second half of a pregnancy and lasting until the end
of breast-feeding:
distended, rounded lobules separated by septa of brous interlobular
connective tissue
numerous pale-staining secretory alveoli variably lled with extracellu-
lar secretion (colostrum until about the end of the rst postnatal week
and then milk)
In the rst half of a pregnancy, high estrogen levels induce a prolifera-
tive response in the ductal epithelium, bringing about further development
of the ducts and formation of the secretory alveoli. In the second half, ris-
ing additional levels of progesterone, acting in concert with prolactin and a
number of other hormones, ensure full breast differentiation and elicit
secretory activity of the alveoli.
200 PDQ HISTOLOGY
Figure 1415 Lactating breast.
Fibrous interlobular
connective tissue
Intralobular duct Secretory
alveoli
Male Reproductive System
The parts of the male reproductive system considered in this
chapter are listed in Table 151. Beginning at the age of puberty, spermato-
zoa are produced from spermatogonial stem cells on an ongoing basis as
synchronously developing clones. This process of spermatogenesis occurs in
the seminiferous epithelium of the seminiferous tubules of each testis. The
other important function of the testis is testosterone secretion by its inter-
stitial (Leydig) cells. From the seminiferous tubules, spermatozoa pass into
a ductus epididymis, a storage compartment where spermatozoa begin to
mature and become motile. On each side, a long muscular-walled tube, the
ductus deferens, conveys motile spermatozoa through the inguinal canal
and pelvic cavity to its site of junction with the proximal end of a seminal
vesicle, an accessory gland that contributes nutritive fluid secretion to
semen. The prostate, the other major accessory gland, similarly contributes
prostatic uid to semen. The anatomic position of the prostate, surround-
ing the urethra at the base of the urinary bladder, is potentially of clinical
importance because constrictive prostatic enlargement (benign or malig-
nant) is a common condition in aging men. On each side, the ejaculatory
duct leading from the junction between a ductus deferens and a seminal
vesicle traverses the prostate inferiorly and opens into the prostatic urethra.
Distally, the spongy part of the urethra terminates at the glans penis.
15
201
Table 151
Male Reproductive System
Testes
Epididymides
Ducti deferentes
Seminal vesicles
Prostate
Penis
202 PDQ HISTOLOGY
Ductus
deferens
Rete
testis
Tunica
albuginea
Seminiferous
tubule
Ductuli
efferentes
Ductus
epididymis
Figure 151 Organization of the testis and epididymis.
Figure 152 Testis.
Mesothelium
Seminiferous
tubules
Tunica
albuginea
TESTES
Each testis (Figure 151) is an ovoid structure enclosed by a capsule-like
tunica albuginea. Fibrous septa extend inward and subdivide the interior
incompletely into lobules, each of which contains up to four tightly packed
seminiferous tubules. The two ends of each looped seminiferous tubule
open through the rete testis and ductuli efferentes of the mediastinum testis
into the long convoluted ductus epididymis extending from the head of the
epididymis to its tail.
In sections of testis observed at low magnification, the thick tunica
albuginea and numerous component seminiferous tubules are readily iden-
tiable (Figure 152). The external surface is covered with squamous peri-
toneal mesothelium (tunica vaginalis testis). Scattered small groups of
interstitial (Leydig) cells may be discerned in the connective tissue stroma
lying between the tubules.
Chapter 15 Male Reproductive System 203
204 PDQ HISTOLOGY
Figure 154 Seminiferous tubule.
Sertoli cell
Spermatozoa Spermatid Spermatogonium
Basement
membrane
Loose connective
tissue
Primary
spermatocyte
Sertoli cell
Myoid cell
Spermatid
Spermatogonium
Spermatozoon
Figure 153 Seminiferous tubule.
Seminiferous Tubules
The wall of a seminiferous tubule (Figure 153) is made up of (1) a single
layer of tall columnar epithelial cells called Sertoli cells and (2) a diverse
population of quiescent, dividing, and differentiating germ cell progenitors
representing the various stages of spermatogenesis. Occupying pockets in
the sides of Sertoli cells, the progenitor cells are so abundant that the Ser-
toli cells appear to be widely separated from each other in sections. At some
level, however, all of these constitutive epithelial cells stay laterally inter-
connected by continuous tight junctions that seal off the adluminal differ-
entiation compartment from the basal stem cell compartment. Adjacent to
the basement membrane at the basal end of the Sertoli cells is an investing
layer of loose connective tissue containing contractile at myoid cells.
Seminiferous tubules observed at an appropriate magnication (Figure
154) commonly show various combinations of the following characteristics:
smooth, round perimeter (basement membrane with associated con-
nective tissue and myoid cells)
fairly large nuclei, with an ovoid or almost pyramidal shape and radial
orientation, representing Sertoli cells (which respond to follicle-
stimulating hormone by secreting androgen-binding protein)
fairly substantial rounded cells, with a spherical-to-ovoid central
nucleus, that lie in the basal stem cell compartment immediately inter-
nal to the basement membrane of the seminiferous tubule (spermato-
gonia = spermatogenic stem cells)
round dividing cells containing condensed chromosomes (predomi-
nantly spermatogonia and primary spermatocytes)
spermatids that are transforming into spermatozoa as they approach
the luminal border
formed spermatozoa awaiting release from the luminal border of Ser-
toli cells
206 PDQ HISTOLOGY
Figure 156 Interstitial (Leydig) cells of testis.
Interstitial (Leydig) cells
1.
2.
3.
4.
Golgi
apparatus
Mitochondrion
Centriole
Forming
acrosome
Forming
flagellum
Head
cap
Residual
cytoplasm
(discarded)
Mitochondrial
sheath
Flagellum
Figure 155 Principal stages of spermiogenesis.
Spermiogenesis
Based on electron microscopic observations, key stages in spermiogenesis
(spermatozoal production from the preceding stage, spermatids) are as
follows:
morphologic transformation of haploid spermatids with no further cell
division (Figure 155)
approximation of the Golgi apparatus to the anterior pole of the nucleus
spreading of the acrosomal vesicle (a Golgi saccule) over the anterior
pole of the nucleus, formation of the acrosome and head cap (site of
penetration enzymes)
caudal migration of the centriole pair
nuclear elongation and further chromatin condensation
agellar growth from the distal centriole (which then disappears)
mitochondrial aggregation and helical rearrangement in proximal
axoneme as the mitochondrial sheath (middle piece)
Interstitial (Leydig) Cells
Testicular interstitial (Leydig) cells (Figure 156) possess the following
characteristics:
interstitial position (small groups in stroma between seminiferous
tubules)
relatively large diameter and central round-to-ovoid, pale-staining
nucleus
pale cytoplasmic staining reecting cholesterol content (stored steroid
precursor)
proximity to blood capillaries and lymphatic capillaries
respond to luteinizing hormone by secreting testosterone
208 PDQ HISTOLOGY
Figure 157 Ductus epididymis.
Figure 158 Ductus deferens.
Lamina propria
Pseudostratied
columnar epithelium Smooth muscle
EFFERENT TESTICULAR DUCTS
Distinctive features of the ductus epididymis (Figure 157) are as follows:
stored spermatozoa in lumen
pseudostratified columnar epithelium with stereocilia (immotile
groups of long microvilli)
surrounding thin circular layer of smooth muscle
supportive connective tissue stroma
A section of a ductus deferens, also widely known as a vas deferens (Fig-
ure 158), is recognizable by the following:
small corrugated lumen with low longitudinal mucosal folds
pseudostratied columnar epithelium with stereocilia
broelastic lamina propria
remarkably substantial muscular wall consisting of a longitudinal inner
layer, circular middle layer, and longitudinal outer layer of smooth mus-
cle (propels through peristalsis during emission)
connective tissue adventitia
210 PDQ HISTOLOGY
Figure 159 Seminal vesicle.
Figure 1510 Prostate.
Fibromuscular stroma
SEMINAL VESICLES
The histologic characteristics of a seminal vesicle (Figure 159) are as follows:
multiple cuts through a single tortuous tubular secretory diverticulum
that is folded into a fairly compact mass
wide, uid-lled lumen containing an elaborate arrangement of thin
folds of broelastic lamina propria covered with secretory epithelium
(different in appearance from the folds found in the ampulla of a uter-
ine tube; see Figure 1411)
dark-staining pseudostratied columnar or simple columnar secretory
epithelium
thick muscular wall consisting of inner circular and outer longitudinal
smooth muscle
connective tissue adventitia binding tubular convolutions together
PROSTATE
Distinctive histologic features of the prostate (Figure 1510) are as follows:
many lobules (compound tubuloalveolar gland)
characteristic extensive folding of the secretory epithelium
purple-staining tall columnar secretory epithelium (pseudostratied or
simple)
broelastic lamina propria
212 PDQ HISTOLOGY
Figure 1511 Secretory unit of prostate.
Secretory unit Calcied concretion
Secretory
epithelium
Stromal smooth
muscle cells
Figure 1512 Penis.
Corpora cavernosa Tunica albuginea
Spongy urethra
Corpus spongiosum
distinctive bromuscular capsule and stroma: smooth muscle cells
deep pink, collagen berslighter pink (Figure 1511)
calcied concretions found in the lumen of some secretory units (vari-
able in occurrence and more common in the prostate of aging men)
PENIS
Sections of the shaft of the penis (Figure 1512) may be readily recognized
by the following criteria:
essentially ovoid outline, thin skin at periphery
paired corpora cavernosa, each enclosed by its fibrous sheath
(tunica albuginea)
ventral spongy urethra, lined with stratied (or pseudostratied)
columnar epithelium and surrounded by the corpus spongiosum
with its enclosing tunica albuginea
Entries with f following a page
number indicate gures; entries
with t following a page number
indicate tables.
A bands, 103, 104, 104f
A cells, of pancreatic islets, 183
Acidophilic staining, 9, 10f
Acidophils, of pituitary, 172f,
173, 173t
Acinus, of liver, 144, 144f
Acrosome, 206f, 207
Adhesion belt, 26f, 27t
Adipocytes, 39t
Adipose tissue, 43, 43f
Adrenals, 174f, 175, 176f, 177
Adrenocorticosteroids, 175
Adrenocorticotropic hormone,
171t, 173t
Adventitia
of blood vessels, 109, 110f
of gut, 130f, 131
Afferent arteriole, of renal cor-
puscle, 158f, 161, 162f, 163
Aldosterone, 175
Alveolar ducts, 150f, 151
Alveolar macrophages, 145, 152f,
153, 154f
Alveolar sacs, 150f, 151, 152f
Alveoli, of lungs, 150f, 151, 152f
Anaphase, 14f, 15f, 17f
Androgen-binding protein, 205
Angiotensin, 163
Antidiuretic hormone, 171
Aorta, 110f, 111, 111f
Apocrine sweat glands, 121, 128
Appendix, 138f, 139
Appositional growth, of cartilage
and bone, 45, 50
Arachnoid membrane, 85
Arector pili muscle, 126, 126f,
127f
Arteries
distributing, 110f, 112f, 113
elastic, 110, 111, 111f
muscular, 113
Arterioles, 110f, 114f, 115
afferent, of renal corpuscles,
158f, 161, 162f, 163
efferent, of renal corpuscles,
161, 162f
Articular cartilage, 45, 46f, 47,
60f
Astrocytes, 89, 89f
Atherosclerotic plaques, 111
Atresia, of ovarian follicles, 187,
188f, 189
Autonomic ganglia, 94f, 95
Autonomic nervous system, 95
Axon, 83, 92f
Band neutrophil, 73t, 74f
Basement membrane, 38, 38f
glomerular, 160f, 161
Basic tissues, 1, 2t
Basophilic erythroblasts, 69t,
70f, 71
Basophilic normoblast, 69t
Basophilic staining, 9, 10f
Basophils
214
Index
of blood, 62t, 65, 66f
of pituitary, 172f, 173, 173t
B cells, 66, 77
of pancreatic islets, 183
Bladder, urinary, 166f, 167
Blood cells, 61-68. See also under
individual types
B lymphocytes, 66, 77
Bone
cancellous, 50, 50f, 51f, 54, 55f
dense (compact), 56-58, 56f-58f
growth of, 53
Bone marrow, 68, 76f, 77
Bone matrix, 49
Brain, 89
Breast
lactating, 200, 200f
resting, 198f, 199
Bronchi, 146f, 147
Bronchioles, 148f, 149, 150f, 151
Brown fat, 43
Brunners glands, 137
Calcitonin, 179, 181
Canal, haversian, 56, 57f, 58f
Canaliculi, 49, 58, 58f
Cancellous bone, 50, 50f, 51f, 54,
55f
Capillaries, 114f, 116f, 117
Capsule cells, of ganglia, 94f, 95
Cardiac muscle, 104-107, 106f
Cartilage, 45-49, 46f
articular, 45, 46f, 47, 60f
elastic, 5f, 48, 48f
bro-, 49, 49f
hyaline, 45-47, 46f
Cartilage matrix, 45, 46f, 47
Catecholamine hormones, 175
C cells, 181
Cell, microscopic appearance of,
6, 6f
Cell junctions, 25, 26f, 27t
Cell nests, 45, 46f
Cells. See under individual
names
Central nervous system, 83-91
Central veins, 142f, 143, 144f
Centroacinar cells, 140f, 141
Cerebellar cortex, 90f, 91
Cerebral cortex, 88f, 89
Chief cells
of fundic glands, 132f, 133
of parathyroid, 182f, 183
Chondrocytes, 7f, 45
Chromatids, 15f
Chromophils, 173
Chromophobes, 173
Chromosomes, 15f, 16, 16f
Cilia, 21, 22f
Circulatory system, 109-117
Collagen
bers, 35, 36, 36f
types, 35, 38, 45
Collecting ducts, of kidney, 158f,
159, 164f
Compact bone. See Dense bone
Compound glands, 28t, 30, 30f
Cones, retinal, 96f, 97
Connective tissue, 35-42, 36f, 37f,
42f
dense ordinary, 41, 42f
loose, 36-41, 36f, 37f
Convoluted tubules, of kidney,
158f, 164f, 165
Corneum, of epidermis, 24f,
120f, 122f, 124f
Corpus albicans, 186f, 187, 190f,
191
Corpus luteum, 186f, 190f, 191
Corpuscles
Hassalls (thymic), 81
renal, 155, 156f, 158f, 160f, 161
Corticogonadotrophs, 173t
Corticothyrotrophs, 173t
Corticotrophs, 173t
Cortisol, 175
Index 215
Crypts, intestinal, 134f, 135, 136f,
138f
Cytoplasm, main components
of, 8t-9t. See also Table 1-3 on
CD-ROM
Cytosol, 8t
D cells, of pancreatic islets, 183
Demilune, serous, 33, 33f
Dendrites, 83, 86f, 87
Dense bone, 56-58, 56f-58f
Dense ordinary connective
tissue, 41, 42f
Dermal papillae, 121
Dermis, 119, 120f, 122f, 124f
Desmosome, 26f, 27t
Digestive system, 129-144
Distal convoluted tubule, 158f,
164f, 165
Distributing arteries, 110f, 112f,
113
Dorsal root ganglia, 95
Ducts
alveolar, 150f, 151
collecting of kidney, 158f, 159,
164f
intralobular and interlobular,
30, 30f
papillary, 159
Ductus deferens, 208f, 209
Ductus epididymis, 208f, 209
Duodenum, 136f, 137
Dura mater, 84f, 85
Eccrine sweat glands, 121, 128,
128f
Efferent arteriole, of renal
corpuscle, 161, 162f
Elastic arteries, 110, 111, 111f
Elastic cartilage, 5f, 48, 48f
Elastic bers, 35, 36f
Elastin, in lungs, 145, 153
Electron micrographs, 3, 4. See
also 7f, 22f, 23f, 116f
Electron microscope, 3
Endocardium, 109
Endochondral ossication, 53,
55f
Endocrine glands, 169-183, 169t.
See also under individual names
Endocrine system, 169-183
Endometrium, 192f, 193, 194f, 195
Endomysium, 99, 100f, 101f
Endoneurium, 92f, 93
Endoplasmic reticulum, 8t, 11
Endosteum, 56
Endothelial cells, 39t, 116f
Endothelium, 109, 112f, 114f
Enteroendocrine cells, 134f, 135
Eosin, 9
Eosinophils, 62t, 64f, 65
Epicardium, 109
Epidermis, 119, 120f, 121, 124f
Epididymis, 202f
Epimysium, 99, 100f
Epinephrine, 175, 177
Epineurium, 92f, 93
Epiphyseal plate, 53, 54f
Epithelial glands, 28-33, 28t,
29f-33f
Epithelial membranes, 19-25, 20t.
See also under individual types
Epithelium
pseudostratied, 20t, 21, 22f
retinal pigment, 96f, 97
simple, 20t, 20f, 21f
stratied, 20t, 23, 24f, 25f
stratied squamous, 23, 24f
transitional, 25, 25f
Erythroblasts
basophilic, 69t, 70f, 71
polychromatophilic, 69t, 70f, 71
Erythrocytes, 62t, 63, 63f
polychromatophilic, 69t, 72, 72f
216 PDQ HISTOLOGY
Erythroid (erythropoietic)
precursors, 69-72, 9t, 70f, 72f,
76f
Esophagus, 130f, 131
Estrogen, 185, 189, 191, 193, 200
Estrogenic phase, of menstrual
cycle, 192f, 193, 194f, 195
Exocrine glands, 28, 28t, 29f-33f
Extracellular matrix, 35, 38
Fallopian tubes. See Uterine tubes
Fasciculata, of adrenal cortex,
174f, 175, 176f, 177
Fat cell, 39t
Fat tissue, brown and white, 43
F cells, of pancreatic islets, 183
Female reproductive system,
185-200
Fibers
collagen, 35, 36, 36f
elastic, 35, 36f
Purkinje, 109
reticular, 35, 77
Fibrillins, 35
Fibroblasts, 39t, 40, 40f
Fibrocartilage, 49, 49f
Filaments
cytoplasmic, 9t
of muscle cells, 99, 100f, 104,
104f
Fixation, 2
Follicle-stimulating hormone,
187, 205
Follicles
hair, 125, 126, 126f, 127f
ovarian, 186f, 187, 188f, 189
primary, secondary, and
tertiary, 187
primordial, 186f, 187
Follicular atresia, 187, 188f, 189
Follicular epithelial cells, of
thyroid, 179, 180f, 181
Follicular phase, of menstrual
cycle, 192f, 193, 194f, 195
Foveolae, 133
Fundic region, of stomach, 132f,
133
Ganglia
autonomic, 94f, 95
spinal or posterior (dorsal)
root, 94f, 95
Ganglion cells, 94f, 95
Gap junction, 26f, 27t, 87, 105,
105f
Gastric pits, 132f, 133
Gastrointestinal tract, 129-139
Germinativum, of epidermis,
120f, 121, 124f
Gland(s)
adrenal, 174f, 175, 176f, 177
Brunners, 137
compound, 28t, 30, 30f
digestive accessory, 129t, 139
endocrine, 169-183, 169t. See
also under individual names
epithelial, 28-33, 28t, 29f-33f
exocrine, 28, 28t, 29f-33f
fundic, 132f, 133
mixed (seromucous), 32-33,
32f, 33f
parathyroid, 182f, 183
parotid, 140f, 141
pituitary, 170f, 171, 172f, 173
pyloric, 132f, 133
sebaceous, 126, 126f, 127
simple, 28t, 29, 29f
sweat, 121, 128, 128f
thyroid, 178f, 179, 180f, 181
Glia, 83, 86f, 88f
Glomerular basement membrane,
160f, 161
Glomerulosa, of adrenal cortex,
174f, 175, 176f, 177
Index 217
Glucagon, 183
Goblet cells, 21, 22f, 23f
Golgi apparatus, 8t, 40f
Gonadotrophs, 173t
Graafian follicles, 187
Granulocytes, 64. See also under
individual types
Granulocytic (granulopoietic)
precursors, 73, 73t, 76f
Granulosa cells, 189
Granulosa lutein cells, 190f, 191
Granulosum, of epidermis, 120f,
121, 122f, 124f
Gray matter, 83, 84f
Ground substance, of connective
tissue, 35
Growth hormone, 171t, 173t
Gut, 129-139
Hair follicles, 125, 126, 126f, 127f
Hassalls corpuscles, 81
Haversian canal, 56, 57f, 58f
Haversian system, 56-58, 57f,
58f
H bands, 104, 104f
Head cap, 206f, 207
Hematopoiesis, 68
Hematoxylin, 9
Hemidesmosome, 26f, 27t
Henle, loop of, 158f, 159, 163, 164f
Histamine, 36, 39t
Hormone
adrenocorticotropic, 171t, 173t
antidiuretic, 171
follicle-stimulating, 187, 205
gastrointestinal, 133, 135
growth, 171t, 173t
luteinizing, 193, 207
parathyroid, 183
release-regulating
(hypothalamic), 171, 171t
thyroid, 179
thyroid-stimulating, 171t, 173t
Hormone receptors, 169
Howships lacunae, 51
Hyaline cartilage, 45-47, 46f
Hydrochloric acid, 133
Hydroxyapatite, 49
Hypodermis, 119
Hypophysis. See Pituitary
Hypothalamic release-regulating
hormones, 171, 171t
Hypothalamohypophysial tracts,
173
Hypothalamus, 173
I bands, 103, 104, 104f
Ileum, 136f, 137
Inferior vena cava, 110f, 113
Insulin, 183
Integumentary system, 119-128
Interalveolar walls, 152f, 153-154,
154f
Intercalated disks, 104, 105, 105f
Interlobular ducts, 30, 30f
Intermediate laments, 9t, 27t,
105, 105f
Internal elastic lamina, 109, 110f,
112f
Interstitial cells, of testis, 203,
206f, 207
Interstitial growth, of cartilage,
45
Interstitial lamellae, 57f, 58, 58f
Interstitial matrix, 35
Intestine
large, 138f, 139
small, 135-137
Intima, of blood vessels, 109,
110f
Intralobular ducts, 30, 30f, 32f
Intramembranous ossication,
50, 50f
Intrinsic factor, 133
218 PDQ HISTOLOGY
Iodination, of thyroglobulin, 179
Ischemic phase, of menstrual
cycle, 193
Islets, pancreatic (of Langerhans),
140f, 141, 182f, 183
Jejunum, 137
JG cells, 161, 162f, 163
Junctions
cell, 25, 26f, 27t
gap, 26f, 27t
tight, 26f, 27t
Juxtaglomerular apparatus, 158f,
162f, 163
Juxtaglomerular cells, 161, 162f,
163
Karyolysis, 16, 17f
Karyorrhexis, 16, 17f
Karyotype, 16
Keratin, 23
Keratinocytes, 121
Kidneys, 155-165
lobule of, 156f, 157
Lacis cells, 163
Lactation. See Breast, lactating
Lacunae, 45, 49
Howships, 51
Lamina densa, 38
Lamina propria, of gut, 130f, 131
Laminin, 38
Langerhans, islets of, 140f, 141,
182f, 183
Large intestine, 138f, 139
Leukocytes, 62t. See also under
individual types
Leydig cells, 203, 206f, 207
Ligaments, 41
Light microscope, 2. See also in
Appendix on CD-ROM
Lingual tonsil, 78f, 79
Liver, 142f, 143, 144
acinus, 144, 144f
lobules of, 142f, 143
Loop of Henle, 158f, 159, 163, 164f
Loose connective tissue, 36-41,
36f, 37f
Lucidum, of epidermis, 120f, 121
Luteinizing hormone, 193, 207
Lymph nodes, 80f, 81
Lymphatic capillaries, 117
Lymphatics, 116f, 117
Lymphocytes, 62t, 66, 67f
Lymphoid organs, primary and
secondary, 77
Lymphoid tissue, 77-82
Lysosomes, 8t
Lysozyme, 135
Macrophages, 39t, 41, 41f
alveolar, 145, 152f, 153, 154f
Macula densa, 162f, 163
Male reproductive system,
201-213
Mammosomatotrophs, 173t
Mammotrophs, 173t
Mast cells, 36f, 39t
Matrix
of bone, 49
of cartilage, 45, 46f, 47
extracellular, 35, 38
interstitial, 35
Media, of blood vessels, 109, 110f
Medullary rays, 155, 156f, 157,
162f, 163
Megakaryocytes, 76f, 77
Melanin, 124f, 125
Melanocytes, 125
Membrane
arachnoid, 85
basement, 38, 38f
epithelial, 19-25, 20t. See also
under individual types
Index 219
synovial, 59, 59f, 60f
Meninges, 83, 85
Menstrual phase, 193
Mesangial cells, 163
Metamyelocytes, 73t, 74f
Metaphase, 15f
Microlaments, 9t
Microscope
electron, 3
light, 2. See also in Appendix
on CD-ROM
Microtubules, 8t, 15f
Mitochondria, 8t
Mitosis, 15f
Mitotic gures, 13, 13f, 14f, 17f
Mixed glands, 32-33, 32f, 33f
M line, 104, 104f
Monocytes, 62t, 68, 68f
Mucosa, of gut, 130f, 131
Mucous secretory units, 31, 31f,
32f
Muscle
cardiac, 104-107, 106f
skeletal, 99-104, 100f-103f
smooth, 107, 107f, 108f
Muscular arteries, 113. See also
Distributing arteries
Muscularis externa, 130f, 131
Muscularis mucosae, 130f, 131
Myelin sheath, 83, 85, 92f, 93
Myeloblasts, 73t
Myelocytes, 73t, 74f
Myeloid tissue, 68-77, 76f
Myobrils, 99, 100f, 102f, 103
Myoid cells, 204f, 205
Myosatellite cells, 104
Nephrons, 155, 158f, 159
Nerves, 91, 92f, 93, 93f
Nervous tissue, 83-95
Neuroglial cells, 83, 86f, 88f
Neurons, 83, 86f, 87
Neuropil, 83, 86f
Neutrophilic myelocytes, 74f, 75
Neutrophils, 62t, 64f, 65
band, 73t, 74f
Nissl bodies, 87, 91, 95
Norepinephrine, 175, 177
Normoblasts, 69t, 70f, 71
types of, 69t
Normocyte, 69t
Nucleolus, 6, 6f, 13
Nucleus, 13
Oocyte
primary, 187, 189
secondary, 185
Ossication
endochondral, 53, 55f
intramembranous, 50, 50f
Osteoblasts, 10f, 50, 51f
Osteoclasts, 18, 18f, 51, 52f
Osteocytes, 50, 51f
Osteon, 56, 57f
Osteoporosis, 53
Osteoprogenitor (osteogenic)
cells, 50, 51f
Ovarian follicles, 186f, 187, 188f,
189
Ovaries, 185-191
Oviduct. See Uterine tube
Oxyphil cells, 182f, 183
Oxytocin, 171
Pacinian corpuscles, 120f, 121,
122f
Palatine tonsils, 78f, 79
Pancreas, 140f, 141, 182f, 183
Pancreatic islets, 140f, 141, 182f,
183
Pancreatic polypeptide, 183
Paneth cells, 134f, 135
Papilla
dermal, 121
of hair, 126, 126f, 127f
Papillary ducts, 159
220 PDQ HISTOLOGY
Parafollicular cells, 179, 180f, 181
Parathyroid hormone, 183
Parathyroids, 182f, 183
Parenchyma, 30
Parietal cells, 132f, 133
Parotid gland, 140f, 141
Pars intermedia and pars ner-
vosa, of pituitary, 171
Pedicels, 161
Penis, 212f, 213
Pepsinogen, 133
Pericytes, 39t, 117
Perimysium, 99, 100f, 101f, 103
Perineurium, 92f, 93
Periodic acid-Schiff staining, 9,
12f
Periosteum, 56, 57f
Peripheral nerves, 91, 92f, 93, 93f
Peripheral nervous system, 83,
91-95
Peyers patches, 137
Photoreceptors, 96f, 97
Pia mater, 84f, 85
Pituicytes, 173
Pituitary, 170f, 171, 172f, 173
Pituitary hormones, 171t, 173,
173t
Plasma, 61
Plasma cells, 39t, 40, 40f
Platelets, 62t, 63, 63f
Pleura, 145
Pneumocytes, 152f, 153, 154, 154f
Podocytes, 161
Polychromatophilic
erythroblasts, 69t, 70f, 71
Polychromatophilic erythrocytes,
69t, 72, 72f
Portal areas, 142f, 143, 144f
Posterior root ganglia, 95
Primary follicles, 186f, 187
Primary oocyte, 187, 189
Primordial follicles, 186f, 187
Proerythroblasts, 69t, 70f, 71
Progestational (progravid) phase,
of menstrual cycle, 187, 193
Progesterone, 185, 191, 193, 200
Prolactin, 171t, 173t, 200
Proliferative phase, of menstrual
cycle, 192f, 193, 194f, 195
Promyelocytes, 73t
Pronormoblast, 69t
Prophase, 15f
Prostate, 210f, 211, 212f, 213
Proximal convoluted tubule,
158f, 164f, 165
Pseudostratied epithelium, 20t,
21, 22f
Purkinje cells, 90f, 91
Purkinje bers, 109
Pyknosis, 16, 17f
Pyloric region, of stomach, 132f,
133
Red pulp, of spleen, 82, 82f
Remodeling, of bone, 53
Renal columns, 155
Renal corpuscles, 155, 156f,
158f, 160f, 161
Renal cortex, 155, 156f, 157
Renal medulla, 155, 158f, 159,
165
Renin, 163
Resolution, 2, 3
Resorption, of bone, 51-53, 52f
Resorption bays, 51
Respiratory bronchioles, 150f,
151
Respiratory system, 145-154
Rete testis, 202f, 203
Reticular bers, 35, 77
Reticularis, of adrenal cortex,
174f, 175, 176f, 177
Reticulocytes, 72, 72f
Retina, 96f, 97
Retinal pigment epithelium, 96f,
97
Index 221
222 PDQ HISTOLOGY
Ribosomes, 8t, 11
Rods, retinal, 96f, 97
Root sheath, 126, 127f
Rugae, 133
Sarcomeres, 104, 104f
Satellite cells, 104
Scalp, 126, 126f
Scanning electron microscope, 3
Schwann cells, 92f, 93
Sebaceous glands, 126, 126f, 127f
Secondary follicles, 186f, 187
Secondary oocyte, 185
Secretory phase, of menstrual
cycle, 187, 192f, 193, 194f
Secretory units
mucous, 31, 31f, 32f
serous, 31, 31f, 32f
types of, 31-33, 31f-33f
Seminal vesicles, 210f, 211
Seminiferous tubules, 204f, 205
Seromucous glands, 32, 32f, 33f
Serosa, of gut, 130f, 131
Serotonin, 63
Serous demilune, 33, 33f
Serous secretory units, 31, 31f,
32f
Sertoli cells, 204f, 205
Serum, 61
Sheath(s)
of hair follicle, 126, 127f
mitochondrial, 206f, 207
myelin, 83, 85, 92f, 93
Simple epithelium, 20t, 20f, 21f
Simple glands, 28t, 29, 29f
Sinusoids
of bone marrow, 76f, 77
Sinusoids
of liver, 142f, 143
of spleen, 82, 82f
Skeletal muscle, 99-104, 100f-103f
Skin
appendages, 119t
thick, 119, 120f, 121, 122f, 123
thin, 119, 120f, 124f, 125
Small intestine, 135-137
Smooth muscle, 107, 107f, 108f
Somatostatin, 171t, 183
Somatotrophs, 173t
Spermatids, 204f, 205, 207
Spermatocytes, 204f, 205
Spermatogonia, 204f, 205
Spermatozoa, 204f, 205, 206f, 207
Spermiogenesis, 207
Spinal cord, 84f, 85
Spinal ganglia, 94f, 95
Spinosum, of epidermis, 120f,
121, 122f, 124f
Spleen, 82, 82f
Staining
acidophilic, 9, 10f
basophilic, 9, 10f
periodic acid-Schiff, 9, 12f
Stains, 9
Stereocilia, 209
Stomach, 132f, 133
Strata, of epidermis, 120f, 121
Stratied epithelium, 20t, 23, 24f,
25f
Stratied squamous epithelium,
23, 24f
Striations, of muscle bers, 103,
103f
Stroma, 30
Subarachnoid space, 84f, 85
Submucosa, of gut, 130f, 131
Suprarenals. See Adrenals
Sweat glands, 121, 128, 128f
Synapses, 86f, 87
Synovial joints, 59
Synovial membrane (synovium),
59, 59f, 60f
Synovocytes, 59
System
circulatory, 109-117
digestive, 129-144
endocrine, 169-183
female reproductive, 185-200
haversian, 56-58, 57f, 58f
integumentary, 119-128
male reproductive, 201-213
respiratory, 145-154
urinary, 155-167
T cells, 66, 77
Telophase, 15f
Tendons, 41
Tertiary follicles, 187
Testes, 202f, 203
Testosterone, 207
Theca interna and externa, 188f,
189
Theca lutein cells, 190f, 191
Thick laments, 104, 104f, 105f
Thick skin, 119, 120f, 121, 122f,
123
Thin laments, 104, 104f, 105f
Thin skin, 119, 120f, 124f, 125
Thymus, 80f, 81
Thyroglobulin, 179, 181
Thyroid gland, 178f, 179, 180f,
181
Thyroid hormone, 179
Thyroid-stimulating hormone,
171t, 173t
Thyrotrophs, 173t
Thyroxine, 179
Tight junction, 26f, 27t
Tissue
adipose (fat), 43, 43f
basic, 1, 2t
connective, 35-42, 36f, 37f, 42f
dense ordinary connective, 41,
42f
epithelial, 19-33
joint, 59
loose connective, 36-41, 36f,
37f
lymphoid, 77-82
myeloid, 68-77, 76f
nervous, 83-95
T lymphocytes, 66, 77
Tonsils, 78f, 79
Trabeculae, of bone, 50, 50f
Trachea, 146f, 147
Tracheobronchial tree, 145
Transitional epithelium, 25, 25f
Transmission electron
microscope, 3
Tubular pole, of renal corpuscle,
160f, 161
Tubules
cortical collecting, 163, 164f
proximal and distal
convoluted, 158f, 164f, 165
renal, 163, 164f
seminiferous, 204f, 205
Tunica adventitia, 109, 110f
Tunica albuginea
of ovary, 185
of testis, 202f, 203
Tunica intima, 109, 110f
Tunica media, 109, 110f
Ureters, 166f, 167
Urinary bladder, 166f, 167
Urinary system, 155-167
Uterine tube, 196f, 197
Uterus, 192f, 193
Vagina, 196f, 197
Varicose veins, 113
Vas deferens, 209
Vasa recta, 159
Vasa vasorum, 109
Vascular pole, of renal corpuscle,
160f, 161
Vasopressin, 171
Veins, 110f, 112f, 113
central, 142f, 143, 144f
Vena cava, 110f, 113
Venules, 110f, 114f, 115
Villi, 129, 134f, 135, 136f
Index 223
White fat, 43
White matter, 83, 84f
White pulp, of spleen, 82
Z line, 104, 104f, 105f
Zona pellucida, 188f, 189
Zones, of adrenal cortex, 174f,
175, 176f, 177
Zonula adherens, 26f, 27t
Zonula occludens, 26f, 27t
224 PDQ HISTOLOGY

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112 PDQ HISTOLOGY

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