CHAPTER 4: EPITHELIAL TISSUE
SEE TABLE 4-1 ON PAGE 74 for the Main Characteristics of
FOUR BASIC TYPES OF TISSUE
1. Epithelial – composed of closely aggregated polyhedral
cells with strong adhesion to one another and attached
to a thin layer of ECM; cellular sheets that line the
cavities of organs and cover the body surface
Principal functions:
 Covering, lining and protecting surfaces (epidermis)
 Absorption (intestinal lining)
 Secretion (parenchymal cells of glands)
2. Connective - Abundant ECM produced by its cells
3. Nervous – long fine processes specialized to receive,
generate and transmit impulses
4. Muscular - elongated cells specialized for contraction
and movement
- All these contain cells and molecules of the
extracellular matrix (ECM), form different organs of
the body
- Most organs can be divided into parenchyma &
stroma
o PARENCHYMA which is composed of the
cells responsible for the organ’s specialized
functions
o STROMA cells which have a supporting role
in the organ except in the brain, spinal cord
the stroma is always connective tissue
CHARACTERISTICS AND FEATURES OF EPITHELIAL CELLS
- Shapes of cells: Columnar, cuboidal, squamous
- Tall cells have elongated nuclei and squamous cells
have flatted nuclei. Cuboidal or pyramidal cells have
spherical nuclei
- Nuclei allow one to determine the number of cell
layers in an epithelium
- Most epithelia rest on connective tissue that
contains the microvaculature bringing nutrients and
oxygen
Lamina Propria - connective tissue that underlies the
epithelia lining of organs of the digestive, respiratory and
urinary systems
Papillae
- area of contact between epithelium and connective
tissue may be increased by these irregularities at the
interface forming small evaginations
- occur most frequently in epithelial tissues subject to
friction
the four basic types of tissue
Epithelial cells generally show polarity, with organelles
and membrane proteins distributed unevenly within the
cell
 BASAL POLE – region of the cell contacting the
connective tissue
 APICAL POLE – opposite end, facing a space
BASEMENT MEMBRANES
- Subjacent connective tissue possessed by epithelial
cells at their basal surfaces
- Presence of glycoproteins
Using the TEM, it can be resolved into two structures:
 BASAL LAMINA/ ELECTRON DENSE LAYER
o 20-100 nm thick
o Nearest the basal poles
o Network of fine fibrils comprising the basal
lamina
 RETICULAR LAMINA
o More diffuse and fibrous than basal lamina
o Beneath basal lamina
The macromolecules of the basal lamina are secreted at
the basal poles of the epithelial cells and form three-
dimensional arrays
 LAMININ – large glycoproteins that self – assemble
as a lacelike network immediately below the cell’s
basal poles where they are held by transmembrane
integrins
 TYPE IV COLLAGEN – monomers of type IV collagen,
contain three polypeptide chains and self-assemble
further to form a feltlike layer
 The laminin and Type IV collagen are held together
by the adhesive glycoprotein entactin/nidogen and
by perlecan, a proteoglycan
These components are also produced by muscle cells,
adipocytes (fat-storing cells) and the cells supporting the
peripheral neurons
The more diffuse meshwork of reticular laminae contains
type III collagen and is bound to the basal laminae by
anchoring fibrils of type IV collagen
FUNCTIONS OF BASEMENT MEMBRANES
- Provide structural support and polarity to epithelial
cells
- Attach epithelia to underlying connective tissue
- Proteins of the layered meshwork help filter
substances entering the epithelium from below,
concentrate mitogenic growth factors and form a
scaffold for epithelial repair and regeneration
- Components help organize proteins in plasma
membrane
- Basement membrane proteins mediate many cell to
cell interactions involving epithelia and mark routes
for certain cell migrations along epithelia
INTERCELLULAR ADHESION AND OTHER JUNCTIONS
- Prominent in epithelial tissues
- Epithelial cells strongly adhere to neighboring cells
and basal laminae, epithelia subject to friction
SPECIALIZED INTERCELLULAR JUNCTIONS WITH FUNCTIONS:
 Tight or Occluding junctions (Zonulae occludens)
o seal between adjacent cells
o most apical of the junctions
o “zonula” – junction forms a band
completely encircling each cell
o Seal between the membrane is due to
interactions between the transmembrane
proteins claudin and occluding of each cell
o Clearly seen after cryofracture as band of
branching strands in the membrane around
each cell’s apical end
o Intercellular seal insures that molecules
crossing an epithelium
o These continuous zones around epithelial
cells prevent membrane proteins at the
apical cell surface from moving in the
membrane to the basal and lateral surfaces
o TWO MEMBRANE DOMAINS ARE
PRODUCED: Apical and Basolateral
o APICAL CELL MEMBRANES – part of the
luminal compartment of a tissue or organ
o BASOLATERAL DOMAINS – part of a basal
compartment that also encompasses the
underlying connective tissue
 Adherent or Anchoring junctions:
o sites of string cell adhesion
o The intercellular seal of this junctional type
ensures that molecules crossing an
epithelium in either direction do so by going
through the cells (trancellular path) rather
than between them (paracellular path)
 Zonula Adherens
o sites of string cell adhesion
o Encircles the epithelial cell, usually
immediately below zonula occludens
o Adherent junction, firmly anchoring cell to
its neighbors
o Cell adhesion here is mediated by
cadherins, transmembrane glycoproteins of
each cell that interact in the presence of Ca
2+
o At the cytoplasmic ends, cadherin bind
catenin that is linked to the zonula
adherens from part of the terminal web
 Desmosome/Macula Adherens
o Related adherent junction
o Resembles a single “spot- weld” and does
not form a belt around the cell
o Disc – shaped structures at the surface of
one cell that are matched with identical
structures at an adjacent cell surface
o Larger number of cadherin family called
desmoglein and desmocollin
o On the cytoplasmic side of eac cell
membrane, these proteins inset into a
dense attachment plaque of anchoring
proteins (plakoglobin and desmoplakin)
that bind intermediate filaments rather
than actin filaments
o Cable – like filaments of cytokeratin
mediate filaments of the cytoskeleton are
very strong, desmosomes provide firm
adhesion among the cells. In nonepithelial
cells, the intermediate filaments attached
to desmosomes are composed of other
proteins, such as desmin or vimentin
 Gap Junctions
o channels for communication between
adjacent cells
o mediate communication rather than
adhesion or occlusion between cells
o functionally important in nearly all
mammalian tissues
o cryofracture shows rgar gap junction consist
of aggregated transmembrane protein
complexes that form circular patches in the
plasma membrane
o gap junction proteins are called connexins
form hexameric connexons each of which
has a central hydrophilic pore about 1.5 nm
in diameter
 o When two cells attach, connexins in the
adjacent cell membrane move laterally and
align to form connexons between the two
cells, with each junction having dozens or
hundreds of aligned connexon pairs
o Gap junctions permit intercellular exchange
of molecules with small less than 1.5 nm of
diameters
o Cyclic nucleotides and ions move rapidly
through gap junctions, allowing cells in
many tissues to act in a coordinated
manner rather than as independent units
o The basal domain of an epithelial cell
attaches to the subjacent basal lamina by
junctions called Hemidesmosomes which
can be observed by TEM
o Hemidesmosomes are half- desmosome
contain integrins rather than cadherins
o The transmembrane integrin proteins bind
the extracellular macromolecules laminin
and collagen type IV
SPECIALIZATIONS OF THE APICAL CELL SURFACE
MICROVILLI
- Specialized for absorption, the apical surfaces
present an array of projections
- In cells such as the lining of the small intestine,
apical surfaces are densely covered with uniform
microvilli visible as a brush or striated border on
these cells
- Average is about 1 microgram long and 0.1
microgram wide but with hundreds or thousands
present on the end of each absorptive cell, the total
surface area can increase by 20-30 fold
- Glycocalyx covering intestinal microvilli is thick and
includes enxymes for digestion of certain
macromolecules
- Each microvillus contains many bundles of actin
filaments capped and cross- linked to each other and
to the surrounding plasma membrane by many
different actin – binding proteins
- Microfilament arrays are dynamic and undergo
various myosin- based movements which help
maintain optimal conditions for absorption via
numerous channels, receptors, and other proteins in
the plasmalemma
- The actin filaments insert into the terminal web of
similar filaments at the base of the microvilli
STEREOCILIA
- Much less common type of apical process, restricted
to absorptive epithelial cells lining the epididymis
and the proximal part of the ductus deferens in the
male reproductive system
- Increase the cell’s surface area, facilitating
absorption
- Resemble microvilli in containing arrays of actin
filaments and various actin – binding proteins, with
smilar diameters and with similar connections to the
cell’s terminal web
- Much longer and less motile than microvilli and may
show distal branching
CILIA
- Long projecting structures, larger than microvilli
- Contain internal arrays of microtubules
- Most if not all cell types have at least one cilium of
variable length, called primary cilium which is not
motile but is enriched with receptors and signal
transduction complexes for detection of light, odors ,
motion and flow of liquid past the cells. Primary cilia
are important in the embryo
- Motile cilia are found only in epithelia, where they
are abundant on the apical domains of many
cuboidal or columnar cells
- Typically 5-10 micrograms long and 0.2 micrograms
in diameter, which is longer and wider than a typical
microvillus
- Each cilium has a core structure consisting of nine
peripheral microtubular doublets in which a few
tubulin protofilaments are shared
- The nine doublets form an array around two central
microtubules; 9+2 assembly of microtubules is called
an axoneme
- Kinesin and Dynein move along the peripheral
microtubules for transport of molecular components
into these structures
- Axonemes are continuous with basal bodies which
are apical cytoplasmic structures just below the cell
membrane
- Basal bodies are similar to centrioles having triplets
of microtubules and dynamic tubulin protofilaments
forming rootlets anchoring the entire structure to
the cytoskeleton
- Ciliary motion occurs to successive changes in the
axoneme
- Complexes with axonemal dyneins bound to a
microtubule in each doublet extend as “arms”
extended to the next doublet
 - With energy from ATP (adenosine triphosphate),
dynein powered sliding of adjacent doublets relative
to each other bends the axoneme and a rapid series
of these sliding movements produces the beating
motion of epithelial cilia
- Flagellum has similar structure with cilia
TYPES OF EPITHELIA
- Two main groups: covering (or lining) epithelia and
secretory(glandular) epithelia
1. Covering or Lining Epithelia
- Organized into one or more layers that cover the
external surface or line the cavities of an organ
 SIMPLE EPITHELIA - contain one cell layer
o Squamous – thin cells
o Cuboidal – cell width and thickness roughly
similar
o Columnar – cells taller than they are wide
 STRATIFIED EPITHELIA – two or more layers
o Most stratified epithelia are classified
according to cell shape of the superficial
layers: squamous, cuboidal, columnar
The very thin surface cells of stratified squamous epithelia
can be “keratinized” (filled with keratin intermediate
filaments) or “nonkeratinized” (with relatively sparse
amounts of keratin
Stratified squamous keratinized epithelium
- found mainly in the epidermis of the skin where it
prevents dehydration of tissue
- Cells form many layers, with less differentiated
cuboidal cells near the underlying connective tissue
- Keratinazation makes cells more irregular in shape
and flatten as they accumulate keratin in the process
and are moved progressively to the skin surface
Stratified Squamous nonkeratinized epithelium
- Lines wet cavities (mouth, esophagus and vagina)
where water loss is not a problem
Stratified cuboidal and Stratified columnar epithelium
- Both relatively rare
- Stratified cuboidal epithelium is restricted to
excretory ducts of salivary and sweat glands
- Stratified columnar epithelium can be found in the
conjunctiva lining the eyelids, where it is both
protective and mucus secreting
Transitional epithelium or eurothelium
- Lines much of the urinary tract, extending from the
kidneys to the proximal part of the urethra
- Characterized by a superficial layer of large, dome-
like cells sometime called umbrella cells
- Specialized to protect underlying tissues from the
hypertonic and potentially cytotoxic effects of urine
Pseudostratified columnar epithelium
- Tall, irregular cells are attached to the basement
membrane but their nuclei are at different levels and
not all cells extend to see the free surface, giving a
stratified appearance
- Lining of the upper respiratory tract, where cells are
heavily ciliated
2. Secretory Epithelia & Glands
- Glands function mainly to produce and secrete
various macromolecules in epithelia
- Products to be secreted are generally stored in the
cells within small membrane – bound vesicles called
secretory granules
- Secretory epithelial cells may synthesize, store and
release proteins in the pancreas, lipids (adrenal,
sebaceous glands) or complex carbohydrates such as
salivary glands
- Epithelia of mammary glands secrete all three
substances
- Sweat glands are a little synthetic activity and
secrete mostly water and electrolytes transferred
from the blood
- Scattered secretory cells sometimes called
unicellular glands are common in simple cuboidal,
simple columnar, and pseudostratified epithelia of
many organs
- An example is the goblet cell abundant in the lining
of the small intestine
- Glands develop from covering epithelia during fetal
life by means of cell proliferation and growth into
the underlying connective tissue, followed by further
differentiation
- Exocrine glands retain their connection with the
surface epithelium, connection forming the tubular
ducts lined with epithelium by which secreted
material leaves the gland
 o Epithelia of exocrine glands are organized
as a continuous system composed of many
small secretory portions and ducts that
transport the secretion out of the gland
- Endocrine glands lose the connection to their
original epithelium and therefore lack ducts
- Stroma of connective tissue support both exocrine
and endocrine secretory units
- A layer of connective tissue also encloses the gland
as its capsule surrounds the larger ducts and forms
partitions or septa that separate the gland into
lobules each containing secretory units connected to
a small part of the duct system
KEYPOINTS
 Glands can be simple (ducts not branched) or
compound (ducts with two or more branches)
 Secretory portions can be tubular (either short or
long and coiled) or acinar (rounded and saclike);
either type of secretory unity may be branched even
if the duct is not branched
 Compound glands can have branching ducts and can
have multiple tubular, acinar, or tubuloacinar
secretory portions
3 BASIC MECHANISMS FOR RELEASING PRODUCTS OF
EPITHELIAL CELLS
1. Merocrine secretion
- Most common; typical exocytosis of proteins or
glycoproteins from membrane bound vesicles
2. Holocrine Secretion
- Cells accumulate product as they mature and
undergo terminal cell differentiation culminating in
complete cell disruption with release of the product
and cell debris into the gland’s lumen. Best seen in
sebaceous glands
3. Apocrine Secretion
- Product at cell’s apical ends
- Product released together with a bit of cytoplasm
and plasma membrane
- Mechanism by which droplets of lipid are secreted in
the mammary gland
Exocrine glands with merocrine secretion can be serous
or mucous
 SEROUS CELLS
o synthesize proteins that are mostly
nonglycosylated, such as digestive enzymes
o well developed RER and Golgi complexes
and are filled apically with secretory
granules in different stages of maturation
o stain intensely with basophilic and
acidophilic stains
o Acini of the pancreas and parotid salivary
glands are composed of serous cells
 MUCOUS CELLS
o Such as goblet cells have RER and golgi
complexes and are filled apically with
secretory granules
o Contain heavily glycosylated proteins called
mucins
o When mucins are released from the cell,
they become hydrated and form mucus
o Hydrophilic mucins are washed from cells
during routine histological preparations,
causing mucinogen granules to stain poorly
with eosin
o Sufficient oligosaccharides usually remain,
however, to allow mucous cells to be
stained by the periodic acid-schiff (PAS)
method
 Some salivary glands are mixed seromucous glands
with both serous acini and mucous tubules. The
product of such gland is a mixture of digestive
enzymes and watery mucus
Epithelia of many exocrine glands: sweat, lachrymal,
salivary and mammary glands contain contractile
myoepithelial cells located inside the basal lamina
Long processes of these cells embrance an acinus as an
octopus might embrace a rounded boulder. Along ducts,
they are more longitudinally arranged. Connected to
each other and to the other epithelial cells by both gap
junctions and desmosomes, myoepithelial cells are rich in
actin filaments and myosins. Their contractions serve to
help propel secretory products into and up the duct
system
Endocrine signaling involves hormone transport in the
blood to target cells throughout the body with other
endocrine glands
 The signaling is then termed as paracrine if close to the
hormone- secreting cell or autocrine if secreting the cell
by itself

TRANSPORT ACROSS EPITHELIA

SODIUM POTASSIUM PUMP – allows cells to maintain the
required low intracellular sodium concentration (5-15
mmol/L vs ~ 140 mmol/L in extracellular fluid)
(SEE FIGURE 4-27 ON PAGE 96)

Trancellular transport
 Specialize in the transfer of ions and water in either
direction across the epithelium. Apical tight
junctions prevent paracellular diffusion or backflow
between the cells
 Kidney tubules are sites of ion and water transport,
maintaining the body’s balance of salts and water
 Cells of the proximal renal tubules specialized
structurally for transcellular transport
 Apical surface is freely permeable to Na + and the
basolateral membranes have sodium pumps for the
active extrusion of Na + into the interstitial fluid
outside the tubules
 Osmotic and electrical balance is maintained by the
passive transfer of Cl- ions and water into the cell
 Basal membrane of these cells is elaborately folded,
with mitochondria located between folds to supply
ATP for Na+/K+ pumps
 Lateral membrane between cells further increase
the surface area for transport

 Thin cells have few organelles other than pinocytotic

vesicles which cross the thin cells in both directions
and release their contents on the opposite side by
exocytosis. This process is called trancytosis and also
occurs between apical and basolateral membranes
domains in cells of the simple

RENEWAL OF EPITHELIAL CELLS

- Renewed continuously by mitotic activity and stem
cell population
- Fast in tissues such as intestinal epithelium, which is
replaced every week, or slow, as in the large glands
- In stratified epithelial tissues, stem cells and mitosis
occur only within the basal layer in contact with the
basal lamina
- Epithelia are normally capable of rapid repair and
replacement of apoptotic or damaged cells