digestives system histology of embryology and cytology

1. 1. Digestive system. Oral cavity. Teeth.
2. Digestive system. Digestive tube.
3. Digestive glands. Gall bladder.
Presented by Dr. Mazhar Hussain
2020

2. The digestive system consists of the digestive tract—oral
cavity, esophagus, stomach, small and large intestines,
rectum, and anus—and its associated glands—salivary
glands, liver, and pancreas . Its function is to obtain from
ingested food the molecules necessary for the
maintenance, growth, and energy needs of the body.
Macromolecules such as proteins, fats, complex
carbohydrates, and nucleic acids are broken down into
small molecules that are more easily absorbed through
the lining of the digestive tract, mostly in the small
intestine. Water, vitamins, and minerals from ingested
food are also absorbed. In addition, the inner layer of the
digestive tract is a protective barrier between the content
of the tract's lumen and the internal milieu of the body.

3. The digestive system consists
of the tract from the mouth
(oral cavity) to the anus, as
well as the digestive glands
emptying into this tract,
primarily the salivary glands,
liver, and pancreas.
The first step in digestion occurs
in the mouth, where food is
moistened by saliva and ground
by the teeth into smaller pieces;
saliva also initiates the
breakdown of carbohydrates.
Digestion continues in the
stomach and small intestine,
where the food's basic
components (eg, amino acids,
monosaccharides, free fatty
acids) are absorbed. Water
absorption occurs in the large
intestine, causing undigested
material to become semisolid.

4. Major layers and organization of the
digestive tract.
• GENERAL STRUCTURE OF THE DIGESTIVE TRACT
The entire gastrointestinal tract has certain
common structural characteristics. It is a hollow tube
with a lumen of variable diameter and a wall made
up of four main layers: the mucosa, submucosa,
muscularis, and serosa. The structure of these
layers is summarized below and is illustrated for the
small intestine

5. Major layers and organization of the digestive tract.

6. The mucosa comprises an epithelial lining; an underlying lamina propria
of loose connective tissue rich in blood vessels, lymphatics,
lymphocytes and smooth muscle cells, sometimes also containing
glands; and a thin layer of smooth muscle called the muscularis mucosae
usually separating mucosa from submucosa. The mucosa is frequently
called a mucous membrane
The submucosa contains denser connective tissue with many blood and
lymph vessels and the submucosal plexus of autonomic nerves. It may
also contain glands and lymphoid tissue.
The thick muscularis is composed of smooth muscle cells that are spirally
oriented and divided into two sublayers. In the internal sublayer (closer to the
lumen), the orientation is generally circular; in the external sublayer, it is mostly
longitudinal. In the connective tissue between the muscle sublayers are blood and
lymph vessels, as well as another autonomic myenteric nerve plexus. This and the
submucosal plexus together comprise the local enteric nervous system of the
digestive tract, containing largely autonomic neurons functioning independently
of the central nervous system (CNS).
The serosa is a thin layer of loose connective tissue, rich in blood vessels,
lymphatics, and adipose tissue, with a simple squamous covering epithelium
(mesothelium)

7. The main functions of the digestive
tract's epithelial lining are to:
•Provide a selectively permeable barrier between the
contents of the tract and the tissues of the body,
•Facilitate the transport and digestion of food,
•Promote the absorption of the products of this
digestion,
•Produce hormones that affect the activity of the
digestive system,
•Produce mucus for lubrication and protection.

8. MEDICAL APPLICATION
• In certain diseases, such as Hirschsprung disease (congenital
megacolon) or Chagas disease (Trypanosoma cruzi infection),
the plexuses in the digestive tract are severely injured and
most of their neurons are destroyed. This results in
disturbances of digestive tract motility, with frequent
dilatations in some areas. The abundant innervation from the
autonomic nervous system that the digestive tract receives
provides an anatomic explanation of the widely observed
action of emotional stress on this tract.

9. ORAL CAVITY
The oral cavity is lined with stratified squamous epithelium, keratinized or
nonkeratinized, depending on the region. The keratin layer
protects the oral mucosa from damage during masticatory function and is best
developed on the gingiva (gum) and hard palate. The lamina propria in
these regions has many papillae and rests directly on bony tissue. Nonkeratinized
squamous epithelium covers the soft palate, lips, cheeks, and the floor
of the mouth. Surface cells are shed continuously and replaced by progeny of stem
cells in the basal epithelial layer. The lamina propria has papillae
similar to those in the dermis of the skin and is continuous with a submucosa
containing diffuse small salivary glands. The soft palate also has a core of
skeletal muscle and lymphoid nodules. In the lips, there is also striated muscle and a
transition from the oral nonkeratinized epithelium to the keratinized
epithelium of the skin

10. • Lip. Low-magnification micrograph of
a lip section showing one side
covered by typical oral mucosa (OM),
the opposite side covered by skin (S)
containing hair follicles (F) and
associated glands. Between the oral
portion of the lips and normal skin is
the vermilion (V), or the vermilion
zone, where epidermis is very thin,
lightly keratinized, and transparent to
blood in the rich microvasculature of
the underlying connective tissue.
Because this region lacks the glands
for oil and sweat, it is prone to
excessive dryness and chapping in
cold, dry weather. Internally, the lips
contain much-striated muscle (M)
and many minor salivary glands

11. Teeth
In the adult human there are normally 32 permanent teeth, arranged in two bilaterally
symmetric arches in the maxillary and mandibular bones . Each quadrant has eight
teeth: two incisors, one canine, two premolars, and three permanent molars. Twenty of
the permanent teeth are preceded by deciduous (baby) teeth which are shed; the others
are permanent molars with no deciduous precursors. Each tooth has a crown exposed
above the
gingiva, a constricted neck at the gum, and one or more roots below the gingiva that
hold the teeth in bony sockets called alveoli, one for each tooth

12. : The dentition of the permanent teeth is shown, as well as the approximate age at eruption for
each
tooth.

13. : Diagram of a molar's internal structure is similar to that of all teeth, with an enamel-covered
crown, cementum-covered roots anchoring the tooth to alveolar bone of the jaw, and a slightly
constricted neck where the enamel and cementum coverings meet at the gingiva. A pulp cavity
extends into the neck and is filled with well-vascularized, well-innervated mesenchymal
connective tissue. Blood vessels and nerves enter the tooth through apical foramina at the
root tips.

14. THE CROWN
• The crown is covered by the extremely hard enamel and
the roots by a bone-like tissue called cementum. These
two coverings meet at the neck of the tooth. The bulk of a
tooth is composed of another calcified material, dentin,
which surrounds a soft connective tissue-filled space
known as the pulp cavity .The pulp cavity narrows in the
roots as the root canals, which extend to the tip of each
root, where an opening (apical foramen) permits the
entrance and exit of blood vessels, lymphatics, and nerves
of the pulp cavity. The periodontal ligaments are fibrous
connective tissue bundles of collagen fibers inserted into
both the cementum and alveolar bone, fixing the tooth
firmly in its bony socket (alveolus).

15. DENTIN
Dentin is a calcified tissue consisting of 70% calcium hydroxyapatite, making it
harder than bone. The organic matrix contains type I collagen fibers and
glycosaminoglycans secreted by odontoblasts, tall polarized cells that line the
tooth's internal pulp cavity . Mineralization of the predentin
matrix involves matrix vesicles in a process similar to that in osteoid . Long,
slender apical odontoblast processes lie within dentinal tubules which
penetrate the full thickness of the dentin, gradually becoming longer as the
dentin becomes thicker. Along their length the processes extend fine branches
into smaller lateral branches of the tubules Odontoblasts remain active in
predentin secretion into adult life,
gradually reducing the size of the pulp cavity.

16. Dentin and odontoblasts.
(a): Odontoblasts (O) are long polarized cells derived from mesenchyme of the developing
pulp cavity (PC). Odontoblasts are specialized for collagen and GAG synthesis and are bound
together by junctional complexes as a layer, with no basal lamina, so that a collagen-rich matrix
called predentin (P) is secreted only from their apical ends at the dentinal surface. Within
approximately one day of secretion predentin mineralizes to become dentin (D) as
hydroxyapatite crystals form in a process similar to that occurring in osteoid of developing
bones. In this process the collagen is masked and calcified matrix becomes
much more acidophilic and stains quite differently than that of predentin. When predentin
secretion begins an apical extension from each cell, the odontoblast process (OP), forms and is
surrounded by new matrix. As the dentin-predentin layer thickens, these processes lengthen.
When tooth formation is complete odontoblasts persist and their processes are maintained in
canals called dentinal tubules which run through the full thickness of the dentin. X400. Mallory
trichrome from M.F. Santos, Department of Histology and Embryology, Institute of Biomedical
Sciences, University of São Paulo, Brazil.)

18. . (b, c): Odontoblast processes can be silver-stained and shown to branch near the
junction of dentin with enamel (E) and along their length closer to
their source (c), with the lateral branches occupying smaller canaliculi within dentin.

19. (d): These odontoblast process (OP) connections to the odontoblasts (O),
shown with stained nuclei here, are important for the maintenance of dentin
in adult teeth. X400. Mallory trichrome, c and d
used, with permission, from M.F. Santos, Department of Histology and
Embryology, Institute of Biomedical Sciences, University of São Paulo, Brazil.)

20. Ultrastructure of dentinal tubule.
shows the calcification of dentin (D) at its border with not-yet- calcified predentin (P). An
odontoblast process (OP) with microtubules and a few
secretory vesicles occupies much of the space (S) in the dentinal tubule. A process extends from
each odontoblast and the tubules continue completely across
the dentin layer. X32,000.

21. (b): Cross-section of an odontoblast process (OP) near predentin (P) shows
its close association with an unmyelinated nerve fiber (N) extending there
from fibers in the pulp cavity. These nerves respond to various stimuli, such as
cold temperatures, reaching the nerve fibers through the dentinal tubules.

22. Teeth are sensitive to
stimuli such as cold,
heat, and acidic pH, all
of which can be
perceived as pain. Pulp
is highly innervated and
some unmyelinated
nerve fibers extend into
the dental tubules near
the pulp cavity. The
different stimuli can
affect fluid inside
dentinal tubules,
stimulating
these nerve fibers
located near
odontoblast processes.

23. Enamelis the hardest component of the human body, consisting of
nearly 98% hydroxyapatite and the rest organic material including at least
two unique proteins, amelogenin and enamelin, but no collagen. Other
ions, such as fluoride, can be incorporated or adsorbed by the
hydroxyapatite crystals; enamel containing fluorapatite is more resistant
to acidic dissolution caused by microorganisms, hence the addition of
fluoride to toothpaste and water
supplies.
Enamel consists of interlocking rods or columns, enamel rods (prisms), bound
together by other enamel. Each rod extends through the entire thickness of the
enamel layer; the precise arrangement of rods in groups is very important for
enamel's strength and mechanical properties.

24. Ameloblasts and enamel.
Ameloblasts (A) are tall polarized cells
whose apical ends initially contact
dentin (D). Ameloblasts are joined to
form a cell layer surrounded basally by
connective tissue (CT). As odontoblasts
secrete predentin, ameloblasts secrete
a matrix lacking collagens, but rich in a
few glycoproteins which quickly
initiate
calcium hydroxyapatite formation to
make enamel (E), the hardest material
in the body. Enamel forms a layer, but
consists of enamel rods or prisms,
solidly
fused together by more enamel. Each
enamel rod represents the product of
one ameloblast. No cellular processes
occur in enamel and the layer of
ameloblasts
surrounding the developing crown is
completely lost during tooth eruption.
Teeth that have been decalcified for
histological sectioning typically lose
their
enamel layer completely.

25. Ameloblasts and
enamel.
:Micrograph of a thin
preparation of a tooth
prepared by grinding. Fine
tubules can be observed in
the dentin (D) and rods
aligned the same way can be
faintly observed (arrows) in
the enamel (E). The more
prominent lines that cross
enamel diagonally represent
incremental growth lines
produced as the enamel
matrix was secreted cyclically
by the ameloblast layer

26. Enamel
• Enamel is produced by cells of ectodermal
origin, whereas most of the other structures
of teeth derive from mesodermal and neural
crest cells. Together these cells produce a
series of structures around the developing
oral cavity, the enamel organs, each of which
forms one tooth

27. Tooth formation
Tooth formation begins in the sixth week of human development
when ectodermal epithelium lining the oral cavity begins to grow
into the underlying mesenchyme of the developing jaws. At a series
of sites corresponding to each future tooth, these epithelial cells
proliferate extensively and become organized as enamel organs,
each shaped rather like a wine glass with its stem initially still
attached to the oral lining. Ameloblasts form from the innermost
layer of cells in the enamel organ. Mesenchymal cells inside the
concave portion of the enamel organ include neural crest cells
which differentiate as the layer of odontoblasts with their apical
ends in contact with the apical ends of the ameloblasts.

28. Tooth formation
: When production of dentin and enamel
has begun, the enamel organ
appears as shown in this micrograph. The
ameloblast layer (A) is separated from the
outer enamel epithelium (OEE) by a thick
intervening region rich in GAGs
but having fewer, widely separated cells.
Surrounding the enamel organ is
mesenchyme, some parts of which begin
to undergo intramembranous bone
formation (B) and form the jaws. Inside
the cavity of each enamel organ,
mesenchymal cells comprise the dental
papilla (DP), in which the outermost cells
are
the layer of odontoblasts (O) facing the
ameloblasts. These two cell layers begin
to move apart as the odontoblasts begin
to produce the layer of predentin
(PD). Contact with dentin induces each
ameloblast to begin secretion of a rod or
prism of enamel matrix. More slowly
calcifying interprismatic enamel fuses all
the enamel rods into a very strong, solid
mass.

29. Tooth
formation
: Detail of an enamel organ at a later
stage showing the layers of predentin
(PD) and dentin (D) and a layer of
enamel (E), along with the organized cell
layers that produced this material.
Odontoblasts (O) are in contact with the
very cellular mesenchyme of the dental
papilla (DP) which will become the pulp
cavity. Ameloblasts (A) are prominent in
the now much thinner enamel organ,
which is very close to developing bone
(B). Details of these cell layers are
presented further. Enamel formation
continues until shortly before tooth
eruption; formation of dentin continues
after eruptionuntil the tooth is fully
formed. Odontoblasts persist around the
pulp cavity, with processes penetrating
the dental layer, producing factors to help
maintain dentin. Mesenchymal cells
immediately around the enamel organ
differentiate into the cells of cementum
and other periodontal tissues.

30. Tooth pulp consists of connective tissue
resembling mesenchyme. Its main components are
the layer of odontoblasts, many fibroblasts, thin
collagen
fibrils, and ground substance. Pulp is a highly
innervated and vascularized tissue. Blood vessels
and myelinated nerve fibers enter the apical
foramen and divide into numerous branches. Some
nerve fibers lose their myelin sheaths and extend
into the dentinal tubules. Pulp fibers are
sensitive to pain.

31. PERIODONTIUM
The periodontium comprises the structures responsible for maintaining the teeth in
the maxillary and mandibular bones. It consists of the cementum,
periodontal ligament, alveolar bone, and gingiva.
Cementum covers the dentin of the root and is similar in composition to bone,
although osteons and blood vessels are absent. It is thicker in the apical
region around the root, where there are cementocytes, cells resembling
osteocytes, in lacunae. Unlike osteocytes, however, cementocytes do not
communicate via canaliculi and their nourishment comes from external tissues. Like
bone, cementum is labile and reacts to the stresses to which it is
subjected by resorbing old tissue or producing new tissue. Continuous production
of cementum in the apex compensates for the physiologic wear of the
teeth and maintains close contact between the roots of the teeth and their sockets.

32. MEDICAL APPLICATION
In comparison with bone, the cementum has
lower metabolic activity because it is not
irrigated by blood vessels. This feature allows the
movement of teeth within alveolar bone by
orthodontic appliances without significant root
resorption.

33. The periodontal ligament is connective tissue 150 to 350 m thick
with collagen fiber bundles connecting the cementum and the
alveolar bone of the tooth socket. It permits limited movement
of the tooth within the socket and the fibers are organized to
support the pressures exerted during mastication. This avoids
transmission of pressure directly to the bone which would cause
localized bone resorption. Unlike typical ligaments, it is highly
cellular and has a rich supply of blood vessels and nerves, giving
the periodontal ligament supportive, protective, sensory, and
nutritive functions.
Collagen of the periodontal ligament has an unusually high
turnover rate (as demonstrated by autoradiography) and a high
content of soluble collagens,
with the space between its fibers filled with glycosaminoglycans
(GAGs).

34. Dental Pulp.
The periphery of the dental pulp
contains the organized odontoblasts
(O) contacting the surrounding dentin
(D). Centrally the pulp consists of
delicate, highly
cellular connective tissue resembling
undifferentiated mesenchyme but
with many thin-walled venules (V) and
capillaries. Pulp has reticulin fibers and
other
fine collagen fibers, with much ground
substance. Nerves fibers are also
present. The blood and nerve supplies
enter the pulp cavity via the apical
foramen at
the apex of the roots.

35. The gingiva is a mucous membrane firmly bound to the periosteum of the maxillary and
mandibular bones . It is composed of stratified
squamous epithelium and lamina propria with numerous connective tissue papillae. A
specialized part of this epithelium, named junctional epithelium, is
bound to the tooth enamel by means of a cuticle resembling a thick basal lamina. The
epithelial cells are attached to this cuticle by numerous
hemidesmosomes. Between the enamel and the epithelium is the gingival sulcus, a
groove up to 3 mm deep surrounding the neck
The alveolar bone is in immediate contact with the periodontal ligament, which serves as its
periosteum. It is primary (immature) bone, with the
collagen fibers not arranged in the typical lamellar pattern of adult bone. Many of the collagen fiber
bundles of the periodontal ligament penetrate this
bone and bind it to the cementum . The bone closest to the roots of the teeth forms the socket. Vessels
run through the alveolar bone and
penetrate the periodontal ligament along the root, with some vessels and nerves entering the pulp at
the apical foramen of each root.

36. The periodontium of
each tooth consists of the
cementum, periodontal
ligament, alveolar bone, and
gingiva. (a): Micrograph of
decalcified tooth showing
the gingiva. The free gingiva (FG)
is against the dentin (D), with
little of the gingival sulcus
apparent. Gingiva has many
layers of stratified epithelial cells
covering the connective tissue of
the lamina propria (LP). The
connective tissue is continuous
with that of the periosteum (P)
covering the alveolar bone (B)
and
with the periodontal ligament
(PL).

37. Periodontium.
(b): Micrograph shows the
periodontal ligament (L) with
its many blood vessels (V)
and insertions into the
alveolar bone (B). This
ligament serves as the
periosteum of the alveolar in
tooth sockets and is also
continuous with developing
layers of cementum (C) that
covers the dentin. Cementum
forms a thin layer
of bone-like material secreted
by large, elongated cells
called cementoblasts.

38. Periodontium.
(c): Micrograph
shows the
continuity of
collagen fibers in
alveolar
bone (B) with the
bundles in the
periodontal
ligament (L). X200.
Picrosirius in
polarized light.

39. ESOPHAGUS
The part of the gastrointestinal tract called the esophagus is a muscular tube whose function is
to transport food from the mouth to the stomach. It is
lined by nonkeratinized stratified squamous epithelium with stem cells scattered throughout the
basal layer . In general, the esophagus has
the same major layers as the rest of the digestive tract. In the submucosa are groups of small
mucus-secreting glands, the esophageal glands,
secretions of which facilitate the transport of foodstuffs and protect the mucosa. In the lamina
propria of the region near the stomach are groups of glands,
the esophageal cardiac glands, which also secrete mucus.

40. ESOPHAGUS
(a): Longitudinal section of
esophagus shows mucosa
consisting of
nonkeratinized stratified
squamous epithelium (SS),
lamina propria (LP), and
smooth
muscles of the muscularis
mucosae (MM). Beneath
the mucosa is the
submucosa containing
esophageal mucous glands
(GL) which empty via ducts
(D) onto
the luminal surface.

41. ESOPHAGUS
(b): Transverse section
showing the muscularis
halfway along the esophagus
reveals a combination of
skeletal muscle (right)
and smooth muscle fibers
(left) in the outer layer, which
are cut both longitudinally
and transversely here. This
transition from muscles under
voluntary control
to the type controlled
autonomically is important in
the swallowing mechanism.

42. STOMACH
The stomach, like the small intestine, is a mixed exocrine-endocrine organ that digests food and
secretes hormones. It is a dilated segment of the
digestive tract whose main functions are to continue the digestion of carbohydrates initiated in
the mouth, add an acidic fluid to the ingested food,
transform it by muscular activity into a viscous mass (chyme), and promote the initial digestion of
proteins with the enzyme pepsin. It also produces a
gastric lipase that digests triglycerides . Gross inspection reveals four regions: cardia, fundus,
body, and pylorus . The fundus and body
are identical in microscopic structure so that only three histologically distinct regions are
recognized. The mucosa and submucosa of the empty stomach
have longitudinally directed folds known as rugae, which flatten when the stomach is filled with
food. The wall in all regions of the stomach is made up of
all four major layers

43. REGIONS OF STOMACH

44. Wall of the stomach
with rugae.
Wall of the stomach with rugae.
Low magnification micrograph of
the stomach wall at the fundus
shows the relative thickness of the
four major layers: the mucosa (M),
the submucosa (SM),
the muscularis externa (ME), and
the serosa (S). Two rugae (folds) cut
transversely and consisting of
mucosa and submucosa are
included. The mucosa is
packed with branched tubular
glands penetrating the full thickness
of the lamina propria so that this
sublayer cannot be distinguished at
this magnification.
The muscularis mucosae (arrows),
immediately beneath the basal ends
of the gastric glands, is shown. The
submucosa is largely loose
connective tissue, with
blood vessels (V) and lymphatics

45. Mucosa
Changing abruptly at the esophago-gastric junction, the mucosa of the stomach consists of a
simple columnar surface epithelium that invaginates into
the lamina propria, forming gastric pitS. Emptying into the gastric pits are branched, tubular
glands characteristic of the
stomach region (cardiac, gastric, and pyloric). Stem cells for the entire epithelial lining of the
stomach are located in the upper regions of these glands
near the gastric pits. The vascularized lamina propria that surrounds and supports these pits
and glands contains smooth muscle fibers and lymphoid
cells. Separating the mucosa from the underlying submucosa is a layer of smooth muscle, the
muscularis mucosae

46. Esophagogastric
junction
At the junction of the esophagus
(E) and the cardiac region of the
stomach (C) there is an abrupt
change in the mucosa from
stratified squamous epithelium to
simple columnar epithelium
invaginating as gastric pits (GP).
The mucosa contains many
mucus-secreting esophageal
cardiac glands (ECG), whose
function is
supplemented by mucous cardiac
glands (CG) opening into the
superficial gastric pits. Strands of
muscularis mucosae (arrow)
separate the mucosa and
submucosa (SM).

47. Digestive tube.
3. Digestive glands. Gall bladder.

48. Gastric pits
and glands.
(a): SEM of the
stomach lining
cleared of its
mucus layer
reveals closely
placed gastric pits
(P) surrounded by
polygonal apical
ends of surface
mucous cells.

49. Gastric pits
and glands.
(b): Micrograph of the same
lining shows that these
surface mucous cells are part
of a simple columnar
epithelium continuous with
the lining of the pits
(P). Each pit extends into the
lamina propria and then
branches into several tubular
glands. These glands branch
further, coil slightly, and fill
most of the
volume of the mucosa.
Around the glands, which
contain other cells besides
columnar cells, a small
amount of connective tissue
comprising the lamina propria
is also seen.

50. REGIONAL DIFFERENCES IN THE
STOMACH MUCOSA
The cardia is a narrow circular region, only 1.5–3 cm in
width, at the transition between the esophagus and the
stomach . The pylorus is
the funnel-shaped region opening into the small
intestine. The mucosa of these two stomach regions
contains tubular glands, usually branched, with coiled
secretory portions called cardial glands and pyloric
glands . The pits leading to these glands are longer in
the pylorus. In both regions
the glands secrete abundant mucus, as well as lysozyme,
an enzyme that attacks bacterial walls.

51. Salivary Glands
These are the parotid, submandibular and sublingual glands, and
numerous small glands situated in the mucous membrane of the
lips (labial glands), cheeks (buccal glands), tongue (lingual glands),
and palate (palatine glands). Some salivary gland tissues may be
seen in the palatine and pharyngeal tonsils. The secretions of
these glands help to keep the mouth moist, and provide a
protective and lubricant coat of mucous. Some enzymes (amylase,
lysozyme), and immunoglobulin IgA are also present in the
secretions.

53. Salivary Glands
Salivary glands are compound tubulo-alveolar glands (racemose glands). Their
secretory elements (also referred to as end pieces or as the portio terminalis)
may be rounded (acini), pear shaped (alveoli), tubular, or a mixture of these
(tubulo-acinar, tubulo-alveolar). The secretory elements lead into a series of
ducts through which their secretions are poured into the oral cavity. In
sections through salivary glands we see a large number of closely packed
acini (or alveoli) with ducts scattered between them. These elements are
supported by connective tissue that also divides the glands into lobules, and
forms capsules around them. Blood vessels, lymphatics and nerves run in the
connective tissue that may at places contain some adipose tissue
The cells lining the alveoli of salivary glands are usually described either as serous
or mucous. In sections stained with haematoxylin and eosin serous cells stain
darkly (because of the presence of zymogen granules: the colour varies from pink
to dark purple). They have rounded nuclei that lie towards the base. In contrast
mucous cells stain very lightly and, therefore, appear empty. The cells are in fact
almost completely filled in by a mucoid material that stains very poorly Fig.
14.14). This material pushes the nuclei towards the basement membrane. The
nuclei are flattened. Further details of the structure of mucous and serous cells
are considered below

54. An alveolus is typically made up entirely of serous cells or of
mucous cells. However, in some cases mucous alveoli are covered
(on one or more sides) by groups of serous cells that are arranged
in the form of crescents or demilunes .In the parotid gland the
alveoli are almost entirely serous, only an occasional mucous
alveolus being present . In the submandibular gland some alveoli
are serous and some are mucous, the latter being frequently
capped by serous crescents . The sublingual glands are made up
predominantly of mucous alveoli, but a few serous demilunes may
be presen.

55. A secretory unit, or gland, with only one type of cell (serous or
mucous) is said to be homocrine. If it contains more than one
variety of cells it is said to be heterocrine. From what has been
said above it will be clear that all the three major salivary glands
are heterocrine.
Secretions produced in alveoli pass along a system of ducts,
different parts of which have differing structure. The smallest
ducts are called intercalated ducts. These are lined by cuboidal
or flattened cells.
Intercalated ducts open into striated ducts lined by columnar
cells. They are so called because the basal parts of the cells
show vertical striations. Striated ducts open into excretory ducts
that are lined by simple columnar epithelium.

56. Some Additional Details
1. Serous cells are usually arranged in the form of rounded acini. As a result each cell
is roughly pyramidal having a broad base (towards the basement membrane) and a
narrow apex (towards the lumen). Some microvilli and pinocytotic vesicles are seen at
the apex of the cell. The lumen of the acinus often extends for some distance
between adjacent cells: these extensions are called intercellular secretory canaliculi.
Deep to these canaliculi the cell membranes of adjoining cells are united by tight
junctions. Deep to these junctions, the lateral cell margins show folds that
interdigitate with those of adjoining cells. The apical cytoplasm contains secretory
granules that are small, homogeneous, and electron dense. The cytoplasm also
contains a prominent Golgi complex and abundant rough endoplasmic reticulum, both
features indicating considerable synthetic activity. Mitochondria, lysosomes, and
microfilaments are also present.

57. 2. Mucous cells are usually arranged in the form of tubular secretory elements.
Cresents present in relation to them are located at the ends of the tubules. The cells lining
mucous cells tend to be columnar rather than pyramidal. Their secretory granules are large
and ill defined. Rough endoplasmic reticulum and Golgi complex are similar to those in
serous cells, but microvilli, foldings of plasma membrane, and intercellular canaliculi are
not usually seen.
3. From the point of view of ultrastructure many cells of salivary glands are intermediate
between serous and mucous cells. They are referred to as seromucous cells. Most of the
cells identified as serous with light microscopy in the parotid and submandibular glands
are really seromucous. The secretions of all types of salivary secretory cells contain
protein-carbohydrate complexes. Their concentration is lowest in cases of serous cells, very
high in mucous cells, and with widely differing concentrations in seromucous cells.
4. We have seen that in the submandibular glands mucous acini are often capped by
serous demilunes. The serous cells of a demilune drain into the lumen of the acinus
through fine canaliculi passing through the intervals between mucous cells

58. Innervation of Salivary Glands
Secretion by salivary glands is under hormonal as well as neural control. A local hormone
plasmakinin formed by secretory cells influences vasodilation. Salivary glands are
innervated by autonomic nerves, both parasympathetic (cholinergic) and sympathetic
(adrenergic). Parasympathetic nerves travel to secretory elements along ducts, while
sympathetic nerves travel along arteries. Synaptic contacts between nerve terminals and
effector cells form neuro-effector junctions
Two types of junction, epilemmal and hypolemmal, are described. At epilemmal junctions
the nerve terminal is separated from the secretory or effector cell by the basal lamina. At
hypolemmal junctions the nerve terminal pierces the basal lamina and comes into direct
contact with the effector cell. Nerve impulses reaching one effector cell spread to others
through intercellular contacts. Classically, salivary secretion has been attributed to
parasympathetic stimulation. While this is true, it is believed that sympathetic nerves can
also excite secretion either directly, or by vasodilation. Autonomic nerves not only
stimulate secretion, but also appear to determine its viscosity and other characteristics.
Autonomic nerve terminals are also seen on myoepithelial cells and on cells lining the ducts
of salivary glands. The latter probably influence reabsorption of sodium by cells lining the
ducts. Salivary glands are sensitive to pain, and must therefore have a sensory innervation
as well.

59. 5. Myoepithelial cells are present in relation to alveoli and intercalated ducts of salivary glands. They
may also be seen in relation to larger ducts (intralobular and extralobular). These cells lie between
the epithelial cells and their basement membrane. The myoepithelial cells located on alveoli are often
branched (stellate) and may form ‘baskets’ around the alveoli. Those located on the ducts are
fusiform and run longitudinally along them. With the EM myoepithelial cells are seen to contain the
usual organelles. In addition they have conspicuous filaments that resemble myofilaments of smooth
muscle cells. These filaments are numerous in processes arising from the cell. Cilia are present on
some myoepithelial cells. It has been suggested that the cilia may subserve a sensory or
chemoreceptor function. Myoepithelial cells are contractile, their contraction helping to squeeze out
secretion from alveoli. The cells receive an autonomic innervation (details of which are controversial).
6. The cells lining the striated ducts show an interesting ultrastructure. Their basal
striations are seen to be due to the presence of numerous deep infoldings of the basal
parts of the cell membranes. Numerous elongated mitochondria are present in the
intervals between the folds. Similar cells are also present scattered in the epithelium
of the excretory ducts. These cells are believed to play a role in regulating the water
and electrolyte content of saliva to make it hypotonic. Immunoglobulin A, produced
by plasma cells lying subjacent to the epithelium, passes into saliva through the cells
lining the striated ducts.

60. Pyloric
glands.
The pyloric region of the stomach
has deep gastric pits (P) leading to
short, coiled pyloric glands (G) in
the lamina propria. Cardial glands
are rather similar
histologically and functionally.
Cells of these glands secrete
mucus and lysozyme primarily,
with a few G cells also present.
The glands and pits are
surrounded
by cells of the lamina propria (LP),
connective tissue also containing
lymphatics and MALT.
Immediately beneath the glands
is the smooth muscle layer of the
muscularis mucos

61. In the fundus and body, the mucosa's lamina propria is filled
with branched, tubular gastricglands, three to seven of which
open into the bottom of
each gastric pit. Each gastric gland has an isthmus, a neck, and a
base; the distribution of epithelial cells in the glands is not
uniform . The isthmus, near the gastric pit, contains
differentiating mucous cells that migrate and replace surface
mucous cells, a few undifferentiated stem
cells, and a few parietal (oxyntic) cells; the neck of the glands
consists of stem cells, mucous neck cells (different from the
isthmus mucous cells), and
parietal cells; the base of the glands contains parietal cells and
chief (zymogenic) cells. Various enteroendocrine cells are
dispersed in the
neck and the base of the glands.

62. These cells of the gastric glands
provide key stomach functions.
Important properties of each are as
follows:
•Mucous neck cells are present in clusters or as single
cells between parietal cells in the necks of gastric
glands . They are irregular in shape, with the nucleus
at the base of the cell and the secretory granules near
the apical surface. Their mucus secretion is less
alkaline and quite different from that of the surface
epithelial mucous cells.

63. These cells of the gastric glands provide
key stomach functions. Important
properties of each are as follows:
Parietal cells are present mainly in the upper half of gastric glands, with fewer in the
base. They are large rounded or pyramidal cells, each with one central spherical
nucleus and cytoplasm that is intensely eosinophilic due to the high density of
mitochondria. A striking feature of the active secreting cell seen in the electron
microscope is a deep, circular invagination of the apical plasma membrane, forming
an intracellular canaliculus. Parietal cells secrete both hydrochloric acid (HCl) and
intrinsic factor, a glycoprotein required for uptake of vitamin B12 in the small
intestine. Carbonic anhydrase produces H2CO3 which dissociates in the cytoplasm
into H+ and HCO3 . The active cell also releases K+ and Cl– and the Cl– ions combine
with H+ to form HCl. The abundant mitochondria provide energy for the ion pumps
located mainly in the extensive cell membrane of the microvilli projecting into the
canaliculi.
Secretory activity of parietal cells is stimulated both through cholinergic nerve
endings (parasympathetic stimulation) and by histamine and a
polypeptide called gastrin, both secreted by local enteroendocrine cells.

64. Lamina Propria through Serosa
The lamina propria of the small intestine is composed of loose
connective tissue with blood and lymph vessels, nerve fibers,
and smooth muscle cells. The
lamina propria penetrates the core of each intestinal villus,
bringing with it microvasculature, lymphatics, and. Smooth
muscle fibers inside the villi are responsible for their rhythmic
movements, which are important for efficient absorption. The
muscularis mucosae also
produces local movements of the villi and plicae circulares.

65. Microvasculature, lymphatics, and muscle in villi

66. The proximal part of the duodenum has, primarily in its
submucosa but extending into the mucosa, large clusters of
branched tubular mucous glands, the
duodenal (or Brunner) glands, with small excretory ducts
opening among the intestinal crypts . The product of the
glands is distinctly
alkaline (pH 8.1–9.3), which neutralizes chyme entering the
duodenum from the pylorus, protecting the mucous membrane
and bringing the intestinal
contents to the optimum pH for pancreatic enzyme action. In
the ileum both the lamina propria and submucosa contain the
lymphoid nodule aggregates known as Peyer patches, an
important component of the MAL

67. Duodenal
(Brunner) glands.
Concentrated mainly in the upper
duodenum are large masses of
compound branched mucous glands,
the duodenal glands (DG), with many
lobules that
occupy much of the submucosa and
may extend above the muscularis
mucosae (MM) into the mucosa. Many
small excretory ducts (D) extend from
these
lobules through the lamina propria
and empty into the lumen among the
small intestinal crypts (IC). Alkaline
mucus from duodenal glands
neutralizes the pH
of material entering the duodenum
and supplements the mucus from
goblet cells in lubricating and
protecting the lining of the small
intestine.

68. LARGE INTESTINE
The large intestine or bowel consists of a mucosal membrane with no
folds except in its distal (rectal) portion and no villi . The mucosa is
penetrated throughout its area by tubular intestinal glands lined by
goblet and absorptive cells, with a small number of enteroendocrine cells
. The absorptive cells or colonocytes are columnar and have short,
irregular microvilli.Stem cells for the epithelium of
the large bowel are located in the bottom third of each gland. The large
intestine is well suited to its main functions: absorption of water,
formation of the
fecal mass from undigestible material, and production of mucus that
lubricates the intestinal surface.

70. (a):
Anterior view of the large intestine with the proximal end exposed shows the ileocecal valve
at its attachment to the ileum, along with the blind sac called the
cecum and its extension, the appendix. The mucosa has shallow plicae but no villi. The
muscularis has two layers, but the outer longitudinal layer consists only
of three distinct bundles of muscle fibers called taeniae coli (ribbons of the colon). These
bands cause the colon wall to form a series of sacs called haustra.
The serosa of the colon is continuous with that of the supporting mesenteries and displays a
series of suspended masses of adipose tissue called omental
appendages.

71. (b): At the distal end of the
rectum, the anal canal, the
mucosa and submucosa are
highly vascularized, with
venous sinuses, and are
folded as a series of
longitudinal anal folds with
intervening anal sinuses. Fecal
material accumulates in the
rectum is eliminated by
muscular contraction,
including action of an
internal anal sphincter of
smooth (involuntary) muscle
and an external sphincter of
striated (voluntary) muscle.

72. Wall of the large intestine.

73. a): Diagram showing the wall of the large intestine composed of the
four typical layers.
(b): The mucosa is occupied mostly by tubular intestinal glands
extending as deep as the muscularis mucosae and by lamina propria rich
in MALT. The submucosa is well vascularized. The muscularis has a typical
inner
circular layer, but the outer longitudinal muscle is only present in three
equally spaced bands, the taeniae col

74. The Anal Canal
The anal canal is about 4 cm long. The upper 3 cm
are lined by mucous membrane, and the lower 1 cm
by skin. The area lined by mucous membrane can be
further divided into an upper part (15 mm) and a
lower part (15 mm).
The mucous membrane of the upper 15 mm of the canal is lined by columnar epithelium. The
mucous membrane of this part shows six to twelve longitudinal folds that are called the anal
columns. The lower ends of the anal columns are united to each other by short transverse
folds called the anal valves. The anal valves together form a transverse line that runs all round
the anal canal: this is the pectinate line. The mucous membrane of the next 15 mm of the
rectum is lined by non-keratinised stratified squamous epithelium. This region does not have
anal columns. The mucosa has a bluish
appearance because of the presence of a dense venous plexus between it and the muscle
coat. This region is called the pecten or transitional zone. The lower limit of the pecten forms
the white line (of Hilton). The lowest 8 to 10 mm of the anal canal are lined by true skin in
which hair follicles, sebaceous glands and sweat glands are present.

75. Above each anal valve there is a depression called the anal sinus. Atypical
(apocrine) sweat glands open into each sinus. They are called the anal (or
circumanal) glands. The anal canal is surrounded by circular and longitudinal
layers of muscle continuous with those of the rectum. The circular muscle is
thickened to form the internal anal sphincter. Outside the layer of smooth
muscle, there is the external anal sphincter that is made up of striated muscle.
For further details of the anal musculature see a book on gross anatomy.
Prominent venous plexuses are present in the submucosa of the anal canal. The
internal haemorrhoidal plexus lies above the level of the pectinate line, while
the external haemorrhoidal plexus lies near the lower end of the canal.

77. Mucosa of the
large intestine
(colon).
(a): Transverse section of
the colon shows the
muscularis externa (ME),
including a taenia coli cut
transversely in the lower
part of the figure, the
submucosa
(S), the mucosa (M) filled
with tubular intestinal
glands. Some of these
glands are cut
longitudinally, but most
seen here are cut
transversely.

78. Mucosa of the large
intestine (colon).
(b):
Transversely cut glands are
seen to consist of simple
columnar epithelium
surrounded a tubular
lumen (asterisk) and
embedded in lamina
propria (LP) with
many free lymphocytes.
Lymphocytes can also be
seen penetrating the
epithelium (arrow).

79. Mucosa of the large
intestine (colon).
(c): Longitudinal
section of one gland
stained for
glycoproteins shows
mucus in the lumen
and two major cell
types in the
epithelium: goblet
cells (G) and other
columnar cells
specialized for water
absorption.
X400. PAS.

80. Mucosa of the large
intestine (colon).
(d): TEM micrograph of
the absorptive cells, called
colonocytes, reveals short
microvilli at their apical
ends, prominent Golgi
complexes above the
nuclei, and dilated
intercellular spaces with
interdigitating leaflets of
cell membrane (L), a sign
of active water transport.
The absorption of water is
passive,
following the active
transport of sodium from
the basolateral surfaces of
the epithelial cells

81. Near the beginning of the large intestine, the appendix is an
evagination of the cecum. It is characterized by a relatively
small and irregular lumen,
shorter and less dense tubular glands, and no taeniae coli.
Although it has no function in digestion, the appendix is a
significant component of the MALT,
with abundant lymphoid follicles in its wall

82. Appendix.
A blind evagination off
the cecum, the
appendix, has a very
small lumen, fewer
glands in its mucosa,
and no taeniae coli.
The laminar propria
and submucosa
are generally filled
with lymphocytes and
lymphoid follicles,
making the appendix a
significant part of the
MALT

83. MEDICAL APPLICATION
Because the appendix is a closed sac and its contents are
relatively static, it can easily become a site of
inflammation (appendicitis). With the
small lumen and relatively thin wall of the appendix,
inflammation and the growth of lymphoid follicles in the
wall can produce swelling that can
lead to bursting of the appendix. Severe appendicitis is a
medical emergency since a burst appendix will produce
infection of the peritoneal cavity.

84. In the anal region, the mucous membrane forms a series of longitudinal folds, the anal
columns. About two cm above the anal opening,
at the recto-anal junction, the lining of the mucosa is replaced by stratified squamous
epithelium. In this region, the lamina propria
contains a plexus of large veins that, when excessively dilated and varicose, can produce
hemorrhoids.

85. Mucosa of the recto-anal
junction.
The simple columnar
epithelium with tubular
glands that lines the
rectum (left side) changes
abruptly to stratified
squamous epithelium in
the anal canal (right
side), as seen in this
longitudinal section. The
connective tissue of the
lamina propria is seen to
contain many free
lymphocytes. X40. H&E.

86. Biliary tract and gall bladder.
Bile leaves the liver in the left and right hepatic ducts, which merge (1) to form the
common hepatic duct, which connects to the cystic duct serving the gall bladder.
The latter two ducts merge (2) to form a common bile duct. The main pancreatic duct
merges with the common bile duct at the hepatopancreatic ampulla (3) which enters the
wall of the duodenum. Bile and pancreatic juices together are secreted from the major
duodenal papilla (of Vater) into the duodenal lumen (4). All these
ducts carrying bile are lined by cuboidal or low columnar cells called cholangiocytes,
similar to those of the small bile ductules in the liver.
The hepatic, cystic, and common bile ducts are lined with a mucous membrane having a
simple columnar epithelium of cholangiocytes. The lamina propria and
submucosa are relatively thin, with mucous glands in some areas of the cystic duct, and
surrounded by a thin muscularis. This muscle layer becomes thicker near
the duodenum and finally, in the portion within the duodenal wall, forms a sphincter that
regulates bile flow.

87. The gallbladder is a small organ that stores
bile. It is attached to your digestive system by
a system of hollow ducts called the biliary
tree. The gallbladder sits in an indenture
underneath the right lobe of the liver.

89. The hepatic, cystic, and common bile ducts are lined with a mucous membrane having a simple
columnar epithelium of cholangiocytes. The lamina propria and
submucosa are relatively thin, with mucous glands in some areas of the cystic duct, and
surrounded by a thin muscularis. This muscle layer becomes thicker near
the duodenum and finally, in the portion within the duodenal wall, forms a sphincter that
regulates bile flow.
MEDICAL APPLICATION
Most malignant tumors of the liver derive from hepatocytes or
cholangiocytes of the hepatic ducts. The pathogenesis of liver
carcinoma is associated with a
variety of acquired disorders, such as chronic viral hepatitis (B or C)
and cirrhosis. In the exocrine pancreas, most tumors also arise from
duct epithelial
cells; the mortality rate from pancreatic tumors is high.

90. Gallbladder.
The gallbladder is a saclike structure
that stores and concentrates bile, and
releases it into the duodenum after a
meal
(a): Its wall consists largely of a highly
folded
mucosa, with a simple columnar
epithelium (arrows) overlying a typical
lamina propria (LP); a muscularis (M)
with bundles of muscle fibers oriented
in all directions to
facilitate emptying of the organ; an
external adventitia (A) where it is
against the liver and a serosa where it
is exposed

91. (b): TEM of the epithelium
shows
cells specialized for water
uptake across apical
microvilli (MV) and release
into the intercellular
spaces (arrows) along the
folded basolateral cell
membranes. Abundant
mitochondria provide the
energy for this pumping
process. Scattered apical
secretory granules (G)
contain mucus. X5600.