Organs Associated
C H A P T E R
with the Digestive
Tract
SALIVARY GLANDS 329
PANCREAS 332
LIVER 335
Hepatocytes & Hepatic Lobules 338
Structure & Function in the Liver 343
T he organs associated with the digestive tract include the
major salivary glands, the pancreas, the liver, and the
gallbladder. Products of these organs facilitate transport
and digestion of food within the gastrointestinal tract. e main
functions of the salivary glands are to moisten and lubricate
ingested food and the oral mucosa, to initiate the digestion of
carbohydrates and lipids with amylase and lipase, and to secrete
innate immune components such as lysozyme and lactoferrin.
e pancreas secretes digestive enzymes that act in the
small intestine and hormones important for the metabolism of
the absorbed nutrients. Bile, whose components are necessary
for digestion and absorption of fats, is made in the liver but
stored and concentrated in the gallbladder. e liver also plays
a major role in carbohydrate and protein metabolism, inacti-
vates many toxic substances and drugs, and synthesizes most
plasma proteins and factors necessary for blood coagulation.
SALIVARY GLANDS
Exocrine glands in the mouth produce saliva, which has diges-
tive, lubricating, and protective functions. With a normal pH
of 6.5-6.9, saliva also has an important bu ering function
and in some species is also important for evaporative cool-
ing. ere are three pairs of large salivary glands: the parotid,
submandibular, and sublingual glands (Figure 16–1), in
addition to the numerous minor or intrinsic salivary glands
located throughout most of the oral mucosa which secrete
about 10% of the total saliva volume.
MEDICAL APPLICATION
Inadequate saliva production, leading to dry mouth or xerosto-
mia, can be caused by various factors a ecting the major salivary
glands, such as mumps viral infection, radiation of the glands, or
the normal side e ect of drugs such as antihistamines.
BILIARY TRACT & GALLBLADDER 345
SUMMARY OF KEY POINTS 346
ASSESS YOUR KNOWLEDGE 348
A connective tissue capsule surrounds each major sali-
vary gland. e parenchyma of each consists of secretory units
on a branching duct system arranged in lobules, separated by
septa of connective tissue. e secretion of each gland is either
serous, seromucous, or mucous, depending on its content of
the glycoprotein mucin. Saliva from the parotids is serous
and watery. e submandibular and sublingual glands pro-
duce a seromucous secretion, while that of the minor glands
is mostly mucous. Saliva is modi ed by the cells of the duct
system draining the secretory units, with much Na+ and Cl−
reabsorbed while certain growth factors and digestive enzymes
are added.
ree epithelial cell types comprise the salivary secretory
units:
■ Serous cells are polarized protein-secreting cells, usu-
ally pyramidal in shape, with round nuclei, well-stained
RER, and apical secretory granules (Figures 16–2
through 16–4). Joined apically by tight and adherent
junctions, serous cells form a somewhat spherical unit
called an acinus (L. grape), with a very small central
lumen (Figure 16–2). Serous acinar cells secrete enzymes
and other proteins.
■ Mucous cells are somewhat more columnar in shape,
with more compressed basal nuclei (Figures 16–2 and
16–4). Mucous cells contain apical granules with hydro-
philic mucins that provide lubricating properties in
saliva but cause poor cell staining in routine preparations
(Figure 16–5). Mucous cells are most o en organized
as cylindrical tubules rather than acini. Mixed salivary
glands have tubuloacinar secretory units with both
serous and mucous secretion.
■ Myoepithelial cells, described in Chapter 4, are found
inside the basal lamina surrounding acini, tubules, and
the proximal ends of the duct system (Figures 16–2
and 16–4). ese small, attened cells extend several
329
330 CHAPTER 16 ■ Organs Associated with the Digestive Tract

FIGURE 16–1 Major salivary glands.

Parotid salivary gland

Parotid duct

There are three bilateral pairs of major sali-

vary glands, the parotid, submandibular,
Masseter muscle and sublingual glands, which together
produce about 90% of saliva. Their loca-
tions, relative sizes, and excretory ducts are
shown here. These glands plus microscopic
Mucosa (cut) minor salivary glands located throughout
Sublingual ducts the oral mucosa produce 0.75-1.50 L of
Submandibular duct saliva daily.
Sublingual salivary gland
Mylohyoid muscle (cut)
Submandibular salivary gland

FIGURE 16–2 Epithelial components of a submandibular gland lobule.

Myoepithelial cells Intercellular Myoepithelial cells

secretory canaliculi

Serous acinus

“Serous demilune”
Intercalated duct of a mixed acinus

Intercalated ducts

Striated ducts Basal laminae

Mucous tubule

The secretory portions are composed of pyramidal serous (violet)
and mucous (tan) cells. Serous acini consist of typical protein-
secreting cells with rounded nuclei, basal accumulation of RER,
and apical ends lled with secretory granules. The cells of mucous
tubules have attened, basal nuclei with condensed chromatin.
In the submandibular gland mixed tubuloacinar secretory units
also occur, combining short mucous tubules with distal clusters
of serous cells called “serous demilune.” The short intercalated
ducts are lined with low cuboidal epithelium. The striated ducts
consist of columnar cells with characteristics of ion-transporting
cells: basal membrane invaginations with mitochondrial accumu-
lations. Myoepithelial cells are shown around the serous acini.

FIGURE 16–3 Parotid gland.
SD
A
A A
ID
a b
The large parotid gland consists entirely of serous acini with cells
producing amylase and other proteins for storage in secretory
granules.
(a) Micrograph of a parotid gland shows densely packed serous
acini (A) with ducts. Secretory granules of serous cells are clearly
contractile processes around the associated secretory
unit or duct and their activity is important for moving
secretory products into and through the ducts.
MEDICAL APPLICATION
Excessive saliva production, or sialorrhea, is associated with
the autonomic activity of nausea, in ammation within the
oral cavity, and rabies viral infection.
In the intralobular duct system, secretory acini and
tubules empty into short intercalated ducts, lined by cuboi-
dal epithelial cells, and several of these ducts join to form a
Salivary Glands 331
C H A P T E R
1 6
SD
Organs Associated with the Digestive Tract ■ Salivary Glands
A
CT
shown in this plastic section, as well as an intercalated duct (ID)
and striated duct (SD), both cut transversely. (X400; PT)
(b) Striations of a duct (SD) are better seen here, along with a
septum (CT) and numerous serous acini (A). The connective tissue
often includes adipocytes. (X200; H&E)
striated duct (Figure 16–2). e more columnar striated
duct cells have many infoldings of their basolateral mem-
brane, all aligned with numerous mitochondria that, by light
microscopy, appear as faint basal striations radiating toward
the nuclei (Figure 16–6). Striated ducts reabsorb Na+ ions
from the initial secretion and their folded cell membranes
present a large surface area with ion transporters, facilitat-
ing rapid ion transcytosis and making the secretion slightly
hypotonic.
Plasma cells in the connective tissue surrounding the
small intralobular ducts release IgA, which forms a complex
with the secretory component synthesized by the epithelial
cells of the serous acini and intralobular ducts. Transferred
into the saliva, the IgA complex released into the saliva pro-
vides defense against speci c pathogens in the oral cavity.
332 CHAPTER 16 ■ Organs Associated with the Digestive Tract
FIGURE 16–4 Ultrastructure of serous and
mucous cells.
M all saliva, are branched tubuloacinar glands, having pri-
L mixed units grouped serous cells occur distally on short
S In addition to α-amylase and proline-rich proteins,
My
A micrograph of a mixed acinus from a submandibular gland
shows both serous and mucous cells surrounding the small
lumen (L). Mucous cells (M) have large, hydrophilic granules like
those of goblet cells, while serous cells (S) have small, dense
granules. Small myoepithelial cells (My) extend contractile
processes around each acinus. (X2500)
(Used with permission from Dr John D. Harrison, King’s College
London Dental Institute, London, UK.)
Ducts from each lobule converge and drain into inter-
lobular excretory ducts with increasing size and thicker
connective tissue layers. e lining of these ducts is unusual,
combining various epithelial types, including simple cuboidal
or columnar, strati ed cuboidal or columnar, and pseudostrat-
i ed epithelia, distributed in no apparent pattern. ese atypi-
cal epithelia may re ect their composition of cells with many
diverse functions, including cells for ion reabsorption, cells
for secretion of mucin and other proteins, enteroendocrine
cells, and basal stem cells, all in highly branched ducts of small
diameter. Before emptying into the oral cavity, the main duct
of each gland is lined with nonkeratinized strati ed squamous
epithelium.
Vessels and nerves enter the large salivary glands at a
hilum and gradually branch into the lobules. A rich vascular
and nerve plexus surrounds the secretory and duct compo-
nents of each lobule. e capillaries surrounding the secretory
units provide uid important for saliva production, which is
stimulated by the autonomic nervous system. Parasympathetic
stimulation, usually elicited through the smell or taste of food,
provokes a copious watery secretion with relatively little
organic content. Sympathetic stimulation inhibits such secre-
tion and produces the potential for dry mouth o en associated
with anxiety.
Features speci c to each group of major salivary glands
include the following:
■ Parotid glands, located in each cheek near the ear, are
branched acinar glands with exclusively serous acini
(Figure 16–3). Serous cells of parotid glands secrete
abundant α-amylase that initiates hydrolysis of carbo-
hydrates and proline-rich proteins with antimicrobial
and other protective properties.
■ Submandibular glands, which produce two-thirds of
marily serous acini, but with many mixed tubuloacinar
secretory units (Figures 16–4 and 16–5a). Within the
mucous tubules and o en assume a crescent-shaped
arrangement called a serous demilune (Figure 16–5a).
serous cells of the submandibular gland secrete lyso-
zyme for hydrolysis of bacterial walls.
■ Sublingual glands, the smallest of the major glands, are
also considered branched tubuloacinar glands, but here
secretory tubules of mucous cells predominate and the
main product of the gland is mucus (Figure 16–5b). e
few serous cells present add amylase and lysozyme to the
secretion.
As described in Chapter 15, small, nonencapsulated sali-
vary glands are distributed throughout the oral mucosa and
submucosa with short ducts to the oral cavity. ese minor
salivary glands are usually mucous, except for the small serous
glands at the bases of circumvallate papillae. Plasma cells
releasing IgA are also common within the minor salivary
glands.
PANCREAS
e pancreas is a mixed exocrine-endocrine gland that pro-
duces both digestive enzymes and hormones. It is an elongated
retroperitoneal organ, with a large head near the duodenum
and more narrow body and tail regions that extend to the le
(Figure 16–7). e pancreas has a thin capsule of connec-
tive tissue, from which septa extend to cover the larger ves-
sels and ducts and to separate the parenchyma into lobules
(Figure 16–8). e secretory acini are surrounded by a basal
lamina that is supported only by a delicate sheath of reticu-
lar bers with a rich capillary network. Endocrine function of
the pancreas involves primarily smaller cells similar to entero-
endocrine cells located in variously sized clusters called the
pancreatic islets (islets of Langerhans). ese are described
with the endocrine organs in Chapter 20.
 Pancreas 333

FIGURE 16–5 Submandibular gland and sublingual gland.

C H A P T E R
A M
ID A

1 6
M

Organs Associated with the Digestive Tract ■ Pancreas

S S
A

M ID
M
S
ID
M M

M
SM
S

S A

SM
M M

a b

(a) The submandibular gland is a mixed serous and mucous gland
(serous cells predominate), and shows well-stained serous acini
(A) and “serous demilunes” (S) and pale-staining mucous cells
(M) grouped as tubules in this tubuloacinar gland. (The crescent-
shaped “serous demilunes” arise at least in part artifactually due
to disproportionate swelling of the adjacent mucous cells during
slide preparation.) Small intralobular ducts (ID) drain each lobule.
(X340; H&E)
(b) The sublingual gland is a mixed but largely mucous gland with
a tubuloacinar arrangement of poorly stained mucous cells (M).
Small intralobular ducts (ID) are seen in connective tissue, as well
as small fascicles of lingual striated muscle (SM). (X140; H&E)

MEDICAL APPLICATION
Pancreatic cancer, which is usually a carcinoma of duct
cells, can arise anywhere in the gland but occurs most often
in the head of the organ near the duodenum. The tumor is
usually asymptomatic until growth and metastasis are well
advanced, leading to the low rate of early detection and sub-
sequent high rate of mortality. Metastasis may be facilitated
by the relatively sparse connective tissue around the ducts
and vasculature of the pancreas.
e digestive enzymes are produced by cells of serous acini
in the larger exocrine portion of the pancreas (Figure 16–9a).
is somewhat resembles the parotid gland histologically,
although the pancreas lacks striated ducts and the parotid
glands lack islets of endocrine tissue. Each pancreatic aci-
nus consists of several serous cells surrounding a very small
lumen, without myoepithelial cells (Figure 16–9). e acinar
cells are polarized, with round basal nuclei, and numerous
zymogen granules apically, typical of protein-secreting cells
(Figure 16–10).
Each acinus is drained by a short intercalated duct of
simple squamous or low cuboidal epithelium. e initial cells
of these small ducts extend into the lumen of the acinus as
small pale-staining centroacinar cells that are unique to the
pancreas. Cells of the intercalated ducts secrete a large volume
of uid, rich in HCO3− (bicarbonate ions), which alkalinizes
and transports hydrolytic enzymes produced in the acini.
334 CHAPTER 16 ■ Organs Associated with the Digestive Tract
FIGURE 16–6 Striated ducts.
SD
a b
(a) A striated duct (SD) shows very faint striations in the basal half
of the columnar cells, which represent mitochondria located in
the folds of the lateral cell membrane. (X200; H&E)
(b) SEM indicates that the apical ends of the cells are joined
together near the small lumen (L), with interdigitating folds of cell
membrane best developed at the basal end (B). (X4000)
e intercalated ducts merge with intralobular ducts and
larger interlobular ducts, which have increasingly columnar
epithelia before joining the main pancreatic duct that runs the
length of the gland.
e exocrine pancreas secretes approximately 1.5 L of
alkaline pancreatic juice per day and delivers it directly into
the duodenum where the HCO3− ions neutralize the acidic
chyme entering there from the stomach and establish the pH
for optimal activity of the pancreatic enzymes. ese digestive
enzymes include several proteases, α-amylase, lipases, and
nucleases (DNAase and RNAase). e proteases are secreted
as inactive zymogens (trypsinogen, chymotrypsinogen,
proelastase, kallikreinogen, and procarboxipeptidases). Tryp-
sinogen is cleaved and activated by enteropeptidases in the
duodenum, generating trypsin that activates the other prote-
ases in a cascade. Pancreatic tissue is protected against autodi-
gestion by the following:
■ Restricting protease activation to the duodenum,
■ Trypsin inhibitor, which is copackaged in the secretory
granules with trypsinogen,
B
B
c
(c) SEM shows the bases (B) of several such cells with the basal
lamina removed, revealing the interlocking of folded membrane
between neighboring cells. Mitochondria within the folds supply
energy for rapid ion uptake from saliva. (X4000)
■ e higher pH in the acini and duct system due to
HCO3− secreted by the centroacinar and intercalated
duct cells, which helps keep all the enzymes inactive.
MEDICAL APPLICATION
In acute pancreatitis, the proenzymes may be activated and
digest pancreatic tissues, leading to very serious complica-
tions. Possible causes include infection, gallstones, alcohol-
ism, drugs, and trauma. Chronic pancreatitis can produce
progressive brosis and loss of pancreatic function.
Exocrine secretion in the pancreas is regulated mainly
through two polypeptide hormones produced by enteroendo-
crine cells of the small intestine:
■ Cholecystokinin (CCK) stimulates enzyme secretion by
the acinar cells.
■ Secretin promotes water and HCO3− secretion by the
duct cells.
 Liver 335

FIGURE 16–7 Pancreas and duodenum.

C H A P T E R
Body of pancreas

1 6
Main pancreatic duct
Common bile duct
Tail of
pancreas

Organs Associated with the Digestive Tract ■ Liver

Duodenum

Accessory
pancreatic duct Duodenojejunal
flexure
Hepatopancreatic
ampulla

Major duodenal
papilla Pancreatic
acini
Jejunum

Head of pancreas
(a) Duodenum and pancreas, anterior view

Pancreatic
Acinar cell islet

Pancreatic acinus
(b)

(a) The main regions of the pancreas are shown in relation to the (b) Micrographs show a pancreatic islet and several pancreatic
two pancreatic ducts and the duodenum. acini. (X75 and X200; H&E)

Autonomic (parasympathetic) nerve bers also stimulate LIVER

secretion from both acinar and duct cells.
MEDICAL APPLICATION
In the normal liver most dense connective tissue is found
only in the portal areas, surrounding the blood vessels and
bile ductule. In liver cirrhosis, which occurs late in chronic
liver disease, brosis and proliferation of broblasts and
hepatic stellate cells occur beyond the portal areas. The
excessive connective tissue may disrupt the normal hepatic
architecture and interfere with liver function.
e liver is the largest internal organ, in adults averaging
about 1.5 kg or 2% of the body weight. Located in the right
upper quadrant of the abdomen just below the diaphragm (see
Figure 15–1), the liver has major le and right lobes with two
smaller inferior lobes, most of which are covered by a thin cap-
sule and mesothelium of the visceral peritoneum. e capsule
thickens at the hilum (or porta hepatis) on the inferior side,
where the dual blood supply from the hepatic portal vein
and hepatic artery enters the organ and where the hepatic
vein, lymphatics, and common hepatic (bile) duct exit.
336 CHAPTER 16 ■ Organs Associated with the Digestive Tract

FIGURE 16–8 Pancreas.

A
A I
I
D
I
Low-power view of pancreas includes sev-
eral islets (I) surrounded by many serous
I acini (A). The larger intralobular ducts (D)
are lined by simple columnar epithelium.
The ducts and blood vessels (V) are located
in connective tissue, which also provides
V a thin capsule to the entire gland and thin
A septa separating the lobules of secretory
acini. (X20; H&E)

V D
D

FIGURE 16–9 Pancreatic acini.

Centroacinar cells

Basal lamina
A Intercalated duct
A

A
Zymogen granules Acinar cells
b

(a) Micrograph of exocrine pancreas shows the serous, enzyme-
producing cells arranged in small acini (A) with very small lumens.
Acini are surrounded by only small amounts of connective tissue
with broblasts (F). Each acinus is drained by an intercalated duct
with its initial cells, the centroacinar cells (arrow), inserted into the
acinar lumen. (X200; H&E)
(b) The diagram shows the arrangement of cells more clearly.
Under the in uence of secretin, the centroacinar and intercalated
duct cells secrete a copious HCO3−-rich uid that hydrates, ushes,
and alkalinizes the enzymatic secretion of the acini.

FIGURE 16–10 Pancreatic acinar cell ultrastructure.
L
S
C nucleus (N) surrounded by cytoplasm
G
N
RER
e main digestive function of the liver is production
of bile, a complex substance required for the emulsi cation,
hydrolysis, and uptake of fats in the duodenum. e liver is
also the major interface between the digestive system and the
blood, as the organ in which nutrients absorbed in the small
intestine are processed before distribution throughout the
body. About 75% of the blood entering the liver is nutrient-
rich (but O2-poor) blood from the portal vein arising from the
stomach, intestines, and spleen; the other 25% comes from the
hepatic artery and supplies the organ’s O2.
Hepatocytes (Gr. hepar, liver), the key cells of this organ,
are among the most functionally diverse cells of the body.
Liver 337
C H A P T E R
1 6
Organs Associated with the Digestive Tract ■ Liver
TEM of a pancreatic acinar cell shows its
pyramidal shape and the round, basal
packed with cisternae of rough ER (RER).
The Golgi apparatus (G) is situated at the
apical side of the nucleus and is associ-
ated with condensing vacuoles (C) and
numerous secretory granules (S) with
zymogen. The small lumen (L) of the aci-
nus contains proteins recently released
from the cell by exocytosis. Exocytosis of
digestive enzymes from secretory gran-
ules is promoted by CCK, released by
enteroendocrine cells of the duodenum
when food enters that region from the
stomach. (X8000)
In addition to an exocrine function in the secretion of bile
components, hepatocytes and other liver cells process the con-
tents of blood, with many speci c functions:
■ Synthesis and endocrine secretion into the blood of the
major plasma proteins, including albumins, brinogen,
apolipoproteins, transferrin, and many others
■ Conversion of amino acids into glucose
(gluconeogenesis);
■ Breakdown (detoxi cation) and conjugation of
ingested toxins, including many drugs;
■ Amino acid deamination, producing urea removed
from blood in kidneys;
338 CHAPTER 16 ■ Organs Associated with the Digestive Tract

■ Storage of glucose in glycogen granules and triglycerides
in small lipid droplets;
■ Storage of vitamin A (in hepatic stellate cells) and
other fat-soluble vitamins;
■ Removal of e ete erythrocytes (by specialized macrophages,
or Kup er cells);
■ Storage of iron in complexes with the protein ferritin.
Hepatocytes & Hepatic Lobules
e liver’s unique histologic organization and microvascu-
lature allow hepatocytes to perform their diverse metabolic,
exocrine, and endocrine functions. Hepatocytes are large
cuboidal or polyhedral epithelial cells, with large, round cen-
tral nuclei and eosinophilic cytoplasm rich in mitochondria.
e cells are frequently binucleated and about 50% of them
are polyploid, with two to eight times the normal chromosome
number.
e liver parenchyma is organized as thousands of small
(~0.7 × 2 mm) hepatic lobules in which hepatocytes form
hundreds of irregular plates arranged radially around a small
central vein (Figures 16–11 through 16–13). e hepa-
tocyte plates are supported by a delicate stroma of reticulin
bers (Figure 16–13b). Peripherally each lobule has three to

FIGURE 16–11 Liver.

Hepatic sinusoid

Central vein
Hepatocytes Hepatic Bile canaliculi
lobule
Kupffer cell
Central vein
Hepatic sinusoid
Bile canaliculi
Hepatocyte

Portal triad
Branch of
(a) Hepatic lobules bile duct
Branch of
hepatic portal vein
Branch of
hepatic artery
C

PV
H (b) Hepatocytes and sinusoids

L
B

HA

(c) Portal triad and hepatic lobule

The liver, a large organ in the upper right quadrant of the abdo-
men, immediately below the diaphragm, is composed of thou-
sands of polygonal structures called hepatic lobules, which are
the basic functional units of the organ.
(a) Diagram showing a small central vein in the center of a
hepatic lobule and several sets of blood vessels at its periph-
ery. The peripheral vessels are grouped in connective tissue
of the portal tracts and include a branch of the portal vein, a
branch of the hepatic artery, and a branch of the bile duct (the
portal triad).
(b) Both blood vessels in this triad branch as sinusoids, which run
between plates of hepatocytes and drain into the central vein.
(c) Micrograph of a lobule shows the central vein (C), plates of
hepatocytes (H), and in an adjacent portal area a small lymphatic
(L) and components of the portal triad: a portal venule (PV),
hepatic arteriole (HA), and bile ductule (B). (X220; H&E)

FIGURE 16–12 Hepatic lobule.
A
C
A
V
D
a b
Cut transversely, hepatic lobules are polygonal units show-
ing plates of epithelial cells called hepatocytes radiating
from a central venule (C). (a) Hepatic lobules of some mam-
mals, such as the pig, are delimited on all sides by connective
tissue.
six portal areas with more brous connective tissue, each of
which contains three interlobular structures that comprise the
portal triad (Figures 16–11 and 16–13d):
■ A venule branch of the portal vein, with blood rich in
nutrients but low in O2.
■ An arteriole branch of the hepatic artery, which
supplies O2.
■ One or two small bile ductules of cuboidal epithelium,
branches of the bile conducting system.
Most of the peripheral portal areas also contain lymphat-
ics and nerve bers and in some species (eg, pigs) extend thin
sheets of brous connective tissue completely around the lob-
ules, making individual lobules easier to distinguish than in
humans (Figure 16–12b).
Between all of the anastomosing plates of hepatocytes
of a hepatic lobule are important vascular sinusoids, which
emerge from the peripheral branches of the portal vein and
hepatic artery and converge on the lobule’s central vein (Fig-
ures 16–11 through 16–13c). e venous and arterial blood
mixes in these irregular hepatic sinusoids. e anastomosing
sinusoids have thin, discontinuous linings of fenestrated endo-
thelial cells surrounded by sparse basal lamina and reticular
bers. e discontinuities and fenestrations allow plasma to
Liver 339
C H A P T E R
1 6
Organs Associated with the Digestive Tract ■ Liver
C
D
A D
V
(b) In humans these lobules have much less connective tissue and
their boundaries are more di cult to distinguish. In both cases
peripheral connective tissue of portal areas contains the portal
triad: small bile ductules (D), venule (V) branches of the portal vein,
and arteriole (A) branches of the hepatic artery. (Both X150; H&E)
ll a narrow perisinusoidal space (or space of Disse) and
directly bathe the many irregular microvilli projecting from
the hepatocytes into this space (Figure 16–14). is direct
contact between hepatocytes and plasma facilitates most key
hepatocyte functions involving uptake and release of nutri-
ents, proteins, and potential toxins.
Two other functionally important cells are found with the
sinusoids of hepatic lobules:
■ Numerous specialized stellate macrophages, usually
called Kup er cells, are found within the sinusoid lin-
ing (Figure 16–15). ese cells recognize and phagocy-
tose aged erythrocytes, freeing heme and iron for reuse
or storage in ferritin complexes. Kup er cells are also
antigen-presenting cells and remove any bacteria or debris
present in the portal blood.
■ In the perisinusoidal space are hepatic stellate cells (or
Ito cells) with small lipid droplets, which store vitamin A
and other fat-soluble vitamins (Figure 16–15b). ese
mesenchymal cells, which are di cult to see in routine
preparations, also produce extracellular matrix (ECM)
components (becoming myo broblasts a er liver
injury) and cytokines that help regulate Kup er cell
activity.
340 CHAPTER 16 ■ Organs Associated with the Digestive Tract

FIGURE 16–13 Hepatic lobule microvasculature.

H
S

S
a b

S
H PV
S

HA BD
S
c S d

(a) Hepatocytes (H) are polygonal epithelial cells, which form
branching, irregular plates separated by venous sinusoids (S).
(H&E X400)
(b) Reticulin (collagen type III) bers (R) running along the plates
of hepatocytes (H), supporting these and the intervening sinu-
soids. Most connective tissue in the liver is found in the septa and
portal tracts. (X400; Silver)
(c) With plates of hepatocytes (H) appearing to radiate from it, the
central vein (C) of the lobule has more collagen than the smaller
sinusoids (S) that drain into it from all directions (arrows). (X200;
Mallory trichrome)
(d) Peripheral portal areas contain more connective tissue and
are the sites of the portal triad: a portal venule (PV), an arteriole
branching o the hepatic artery (HA), and one or two bile duct-
ules (BD). (X200; H&E)

e endothelium of the central vein in the middle of each
hepatic lobule is supported by a very thin layer of brous con-
nective tissue (Figure 16–13c). Central venules from each lob-
ule converge into larger veins, which eventually form two or
more large hepatic veins that empty into the inferior vena
cava.
Blood always ows from the periphery to the center of
each hepatic lobule. Consequently, oxygen and metabolites, as
well as all other toxic or nontoxic substances absorbed in the
intestines, reach the lobule’s peripheral cells rst and then the
more central cells. is direction of blood ow partly explains
why the properties and function of the periportal hepatocytes

FIGURE 16–14 Ultrastructure of hepatocytes, perisinusoidal space, and bile canaliculi.
M
G
S PS
E
RER
a H
(a) TEM of hepatocytes shows small bile canaliculi (BC) between
tight junctions (TJ) joining two cells. A hepatocyte nucleus (H) is
in the lower right corner, surrounded by small tubular vesicles of
smooth ER (SER), much rough ER (RER), many mitochondria (M),
small electron-dense glycogen granules, and Golgi complexes
(G). Between the hepatocytes and the fenestrated endothelial cell
(E) of the sinusoid (S) is the very small perisinusoidal space (PS)
almost lled with microvilli. (X9500)
(Figure 16–14a, used with permission from Douglas L. Schmucker,
Department of Anatomy, University of California, San Francisco, CA.)
(b) SEM of the luminal surface of the endothelium lining a hepatic
sinusoid shows grouped fenestrations (F). At the border are seen
di er from those of the centrolobular cells. Hepatocytes near
the portal areas can rely on aerobic metabolism and are o en
more active in protein synthesis, while the more central cells
are exposed to lower concentrations of nutrients and oxy-
gen and are more involved with detoxi cation and glycogen
metabolism.
While the sinusoidal (basolateral) domains of hepa-
tocytes process nutrients and other blood components and
Liver 341
C H A P T E R
H
E
1 6
PS
BC
Organs Associated with the Digestive Tract ■ Liver
F
TJ
M E
H
b
PS
SER
BC H
M
c
cut edges of endothelial cells (E) in this discontinuous sinusoid
and hepatocytes (H). Between these two cells is the thin perisinu-
soidal space (PS), into which project microvilli from the hepato-
cytes surface. (X6500)
(Figure 16–14b, used with permission from Eddie Wisse, Electron
Microscopy Unit, Department of Pathology, University of Maastricht,
Maastricht, the Netherlands.)
(c) SEM of hepatocytes (H) broken apart from one another reveals
the length of a bile canaliculus (BC) along the cell’s surface. Such
canaliculi run between the cells of the hepatocyte plates in the
hepatic lobules and carry bile toward the portal areas where the
canaliculi join cuboidal bile ductules. (X8000)
secrete the plasma proteins, the smaller apical surfaces of the
hepatocytes form bile canaliculi and are involved in exo-
crine secretion of bile (Figures 16–14 and 16–16). Within the
hepatic plates hepatocytes adhere rmly with desmosomes
and junctional complexes. e apical surfaces of two adherent
hepatocytes are grooved and juxtaposed to form the canalicu-
lus, sealed by tight junctions, into which bile components are
secreted (Figure 16–14). ese canaliculi are elongated spaces
342 CHAPTER 16 ■ Organs Associated with the Digestive Tract
FIGURE 16–15 Hepatic sinusoids.
K PS
K
a b
In the endothelial lining of the hepatic sinusoids are numerous
specialized stellate macrophages or Kup er cells, which detect
and phagocytose e ete erythrocytes.
(a) Kup er cells (K) are seen as black cells in a liver lobule from a
rat injected with particulate India ink. (X200; H&E)
(b) In a plastic section, Kup er cells (K) are seen in the sinusoid (S)
between two groups of hepatocytes (H). They are larger than the
(total length > 1 km) with lumens only 0.5-1μm in diameter
with large surface areas due to the many short microvilli from
the constituent hepatocytes (Figures 16–14 and 16–16).
e bile canaliculi form a complex anastomosing network
of channels through the hepatocyte plates that end near the
portal tracts (Figures 16–11b and 16–17). e bile ow there-
fore progresses in a direction opposite to that of the blood, that
is, from the center of the lobule to its periphery. Bile canaliculi
are the smallest branches of the biliary tree or bile conducting
system. ey empty into bile canals of Hering (Figure 16–17)
composed of cuboidal epithelial cells called cholangiocytes.
e short bile canals quickly merge in the portal areas with
the bile ductules lined by cuboidal or columnar cholangio-
cytes and with a distinct connective tissue sheath. Bile duct-
ules gradually merge, enlarge, and form right and le hepatic
ducts leaving the liver.
Into the canaliculi hepatocytes continuously secrete bile, a
mixture of bile acids (organic acids such as cholic acid), bile
salts (the deprotonated forms of bile acids), electrolytes, fatty
acids, phospholipids, cholesterol, and bilirubin. Some bile
components are synthesized in hepatocyte SER, but most are
taken up from the perisinusoidal space; all are quickly secreted
HS
E
H
H
S
attened endothelial cells (E). Between the endothelium and the
hepatocytes is a very thin space called the perisinusoidal space
(PS) of Disse, in which are located small hepatic stellate cells (HS),
or Ito cells, which maintain the very sparse ECM of this compart-
ment and also store vitamin A in small lipid droplets. These cells
are numerous but are di cult to demonstrate in routine histologic
preparations. (X750; PT)
into the bile canaliculi (Figure 16–16). Bile acids/salts have an
important function in emulsifying the lipids in the duodenum,
promoting their digestion and absorption.
Bilirubin is a pigmented breakdown product of heme that
is released from splenic macrophages primarily, but also from
Kup er cells, and carried to hepatocytes bound to albumen.
Released into the duodenum with bile, bilirubin is converted
by intestinal bacteria into other pigmented products, some of
which are absorbed in the intestinal mucosa to be processed
and excreted again in the liver or excreted into urine by the
kidneys. ese bilirubin-related compounds give feces and
urine their characteristic colors.
MEDICAL APPLICATION
The brosis characteristic of cirrhosis produces connective
tissue that can ll the perisinusoidal space and interfere with
metabolic exchange between the hepatocytes and the sinu-
soids. Blockage of hepatocyte secretion into the blood can
result in clotting disorders, hypoalbuminemia, and other
medical problems.
 Liver 343

FIGURE 16–16 Hepatocyte ultrastructure and major functions.

C H A P T E R
1

1 6
Lipid
2
Bile

Organs Associated with the Digestive Tract ■ Liver

RER canaliculus A diagram of hepatocyte cytoplasmic organization,
SER with major functions localized. (1) RER is primarily
Tight engaged in synthesis of plasma proteins for release
Lysosomes (occluding) into the perisinusoidal space. (2) Potentially toxic com-
junctions
pounds, bilirubin (bound to albumin) and bile acids
Golgi Golgi are taken up from the perisinusoidal space, processed
SER Desmosome by enzymes in the tubulovesicular system of the SER,
Mitochondria and secreted into the bile canaliculi. (3) Glucose is
taken up from the perisinusoidal space and stored in
glycogen granules, with the process reversed when
glucose is needed.
RER
Glycogen Microvilli

Perisinusoidal space
Endothelium

Fenestration Reticular fibers

Structure & Function in the Liver

FIGURE 16–17 Bile ductules.
As mentioned previously, hepatocytes are highly versatile cells
Bile canaliculi with diverse functions that are re ected in their structure
(Figure 16–16). Abundant rough ER is focused on synthesis
of plasma proteins and causes cytoplasmic basophilia, which
is o en more pronounced in hepatocytes near the portal areas
(Figure 16–12). Abundant smooth ER, distributed more
evenly throughout the cytoplasm, contains the enzyme sys-
tems for the biotransformation or detoxi cation of substances
in blood, which are then usually excreted with bile. ese
include enzymes responsible for oxidation, methylation, and
Bile ductule conjugation of steroids, barbiturates, antihistamines, anticon-
vulsants, and other drugs. Under some conditions prolonged
presence of drugs can lead to increased amounts of SER in
hepatocytes, thus improving the liver’s detoxi cation capac-
ity. Other SER enzymes (glucuronosyl transferases) conjugate
bilirubin to glucuronate, rendering it more soluble and facili-
tating its excretion in bile.
Hepatocytes Bile canals Cholangiocytes Glycogen granules and small lipid droplets in hepa-
of Hering tocytes, and very small electron-dense ferritin complexes
(hemosiderin) primarily in the Kup er cells, respectively
Near the periphery of each hepatic lobule, many bile canaliculi mediate temporary storage of glucose, triglycerides, and iron.
join with the much larger bile canals of Hering, which are lined Hepatocyte peroxisomes are also abundant and impor-
by cuboidal epithelial cells called cholangiocytes. These canals tant for oxidation of excess fatty acids, catalase-mediated
soon join the bile ductules in the portal areas and drain into
the biliary tree. breakdown of the hydrogen peroxide generated by fatty acid
oxidation (by means of catalase activity), and conversion of
344 CHAPTER 16 ■ Organs Associated with the Digestive Tract
excess purines to uric acid. Many Golgi complexes are also
present, involved in synthesis of both plasma proteins and bile
components. e numerous mitochondria provide energy for
all these activities (Figure 16–16).
MEDICAL APPLICATION
Fatty liver disease is a reversible condition in which large
lipid droplets containing triglycerides accumulate abnormally
in hepatocytes via the process called steatosis. This disorder
has multiple causes, but it occurs most commonly in indi-
viduals with alcoholism or obesity. Accumulation of fat in
hepatocytes may produce a progressive in ammation of the
liver, or hepatitis, in this case called steatohepatitis.
FIGURE 16–18 Concepts of structure-function relationships in liver.
(a) Classic Hepatic Lobule
Drains blood from the portal
vein and the hepatic artery to
the hepatic or the central vein
Hepatic arteriole
Bile duct
Portal vein
Central
(or hepatic)
venule
Studies of liver microanatomy, physiology, and pathology have
given rise to three related ways to view the liver’s organization,
which emphasize di erent aspects of hepatocyte activity.
(a) The classic lobule concept o ers a basic understanding of the
structure-function relationship in liver organization and empha-
sizes the endocrine function of hepatocytes as blood ows past
them toward the central vein.
(b) The portal lobule emphasizes the hepatocytes’ exocrine
function and the ow of bile from regions of three classic lobules
toward the bile duct in the portal triad at the center here. The area
drained by each bile duct is roughly triangular.
(c) The hepatic acinus concept emphasizes the di erent oxygen
and nutrient contents of blood at di erent distances along the
e di erent categories of hepatocyte functions—
including secretion of proteins into blood, the exocrine secre-
tion of bile, and the removal of diverse small compounds from
blood—have led to three ways of considering liver lobule
structure, which are summarized in Figure 16–18.
■ e classic hepatic lobule (Figure 16–18a), with blood
owing past hepatocytes from the portal areas to a cen-
tral venule, emphasizes the endocrine function of the
structure producing factors for uptake by plasma.
■ e concept of portal lobules of hepatocytes is more
useful when considering the exocrine function of these
cells, that is, bile secretion. e portal area has the bile
ductule at the center, and bile, moving in the opposite
direction as the blood, ows toward it from all the sur-
rounding hepatocytes. e tissue draining bile into
(b) Portal Lobule (c) Hepatic Acinus
Drains bile from Supplies oxygenated
hepatocytes to the blood to hepatocytes
bile duct
Central vein
Zone III
least
oxygenated
Zone II
Zone I
most
oxygenated
sinusoids, with blood from each portal area supplying cells in
two or more classic lobules. Major activity of each hepatocyte is
determined by its location along the oxygen/nutrient gradient:
periportal cells of zone I get the most oxygen and nutrients and
show metabolic activity generally di erent from the pericentral
hepatocytes of zone III, exposed to the lowest oxygen and nutri-
ent concentrations. Many pathologic changes in the liver are best
understood from the point of view of liver acini.
(Used with permission from Boron WF, Boulpaep EL. Medical
Physiology: A Cellular and Molecular Approach. Philadelphia, PA:
Saunders Elsevier, 2005.)

each portal area duct is roughly triangular in shape,
with the central veins of three classic lobules at its
angles (Figure 16–18b).
■ e hepatic acinus, a third way of viewing liver cells,
emphasizes the nature of the blood supply to the hepa-
tocytes and the oxygen gradient from the hepatic artery
branch to the central vein. In a liver acinus hepatocytes
make up an irregular oval or diamond-shaped area
extending from two portal triads to the two closest cen-
tral veins (Figure 16–18c). Periportal hepatocytes nearest
the hepatic arteriole, comprising zone I in the acinus,
get the most oxygen and nutrients and can most readily
carry out functions requiring oxidative metabolism such
as protein synthesis. Hepatocytes in zone III, near the
central vein, get the least oxygen and nutrients. ey are
the preferential sites of glycolysis, lipid formation, and
drug biotransformations and are the rst hepatocytes to
undergo fatty accumulation and ischemic necrosis. In
the intervening zone II, hepatocytes have an intermedi-
ate range of metabolic functions between those in zones
I and III. e major activities in any given hepatocyte
result from the cell adapting to the microenvironment
produced by the contents of the blood to which it is
exposed.
MEDICAL APPLICATION
An important function of hepatocyte SER is the conjugation
of hydrophobic (water-insoluble), yellow bilirubin by glucuro-
nosyl transferases to form water-soluble, nontoxic bilirubin
glucuronide, which is excreted into the bile canaliculi. When
bilirubin glucuronide is not formed or excreted properly, vari-
ous diseases characterized by jaundice can result.
A frequent cause of jaundice in newborns is an underde-
veloped state of the hepatocyte SER (neonatal hyperbilirubi-
nemia). A treatment in these cases is exposure to blue light
from ordinary uorescent tubes, which transforms unconju-
gated bilirubin into a water-soluble photoisomer that can be
excreted by the kidneys.
Unlike the salivary glands and pancreas, the liver has
a strong capacity for regeneration despite its normal slow
rate of cell renewal. Hepatocyte loss from the action of toxic
substances triggers mitosis in the remaining healthy hepa-
tocytes in a process of compensatory hyperplasia that
maintains the original tissue mass. Surgical removal of a
liver portion produces a similar response in the hepatocytes
of the remaining lobe(s). e regenerated liver tissue is usu-
ally well organized, with the typical lobular arrangement, and
replaces the functions of the destroyed tissue. is regenera-
tive capacity is important clinically because one major liver
lobe can sometimes be donated by a living relative for surgi-
cal transplant and full liver function restored in both donor
and recipient.
Biliary Tract & Gallbladder 345
Besides proliferation of existing hepatocytes, a role for
liver stem cells in regeneration has been shown in some
C H A P T E R
experimental models. Such cells, o en called oval cells, are
present among cholangiocytes of the bile canals near portal
areas and produce progenitor cells for both hepatocytes and
cholangiocytes.
1 6
MEDICAL APPLICATION
Most malignant tumors of the liver derive from hepatocytes
Organs Associated with the Digestive Tract ■ Biliary Tract & Gallbladder
or cholangiocytes of the hepatic ducts. The pathogenesis of
liver carcinoma is associated with a variety of acquired disor-
ders, such as chronic viral hepatitis (B or C) and cirrhosis.
BILIARY TRACT & GALLBLADDER
e bile produced by the hepatocytes ows through the bile
canaliculi, bile ductules, and bile ducts. ese structures
gradually merge, forming a converging network that ulti-
mately forms the common hepatic duct, which joins the
cystic duct from the gallbladder and continues to the duode-
num as the common bile duct (Figure 16–19).
e hepatic, cystic, and common bile ducts are lined with
a mucous membrane having a simple columnar epithelium of
cholangiocytes. e lamina propria and submucosa are rela-
tively thin, with mucous glands in some areas of the cystic
duct, and surrounded by a thin muscularis. is muscle layer
becomes thicker near the duodenum and nally, in the duode-
nal papilla, forms a sphincter that regulates bile ow into the
small bowel.
e gallbladder is a hollow, pear-shaped organ (Figure 16–19)
attached to the lower surface of the liver, capable of storing
30-50 mL of bile that is concentrated during storage. e wall
of the gallbladder consists of a mucosa composed of simple
columnar epithelium and lamina propria, a thin muscularis
with bundles of muscle bers oriented in several directions,
and an external adventitia or serosa (Figure 16–20a). e
mucosa has numerous folds that are particularly evident when
the gallbladder is empty.
e lining epithelial cells of the gallbladder have promi-
nent mitochondria, microvilli, and large intercellular spaces,
all indicative of cells actively transporting water, in this case
for concentrating bile (Figure 16–20b). e mechanism for
this includes activity of Na+ pumps in the basolateral mem-
branes, followed by passive movement of water from the bile.
To move stored bile into the duodenum, contraction of the
gallbladder muscularis is induced by cholecystokinin (CCK)
released from enteroendocrine cells of the small intestine.
Release of CCK is, in turn, stimulated by the presence of
ingested fats in the small intestine. Gallbladder removal due
to obstruction or chronic in ammation leads to the direct
ow of bile from liver to gut, with few major consequences
on digestion.
346 CHAPTER 16 ■ Organs Associated with the Digestive Tract
FIGURE 16–19 Biliary tract and gallbladder.
Left and right hepatic ducts
Common hepatic duct
Cystic duct
Stored bile
Gallbladder
Minor
duodenal papilla
Hepatopancreatic
ampulla with
hepatopancreatic
sphincter
Major duodenal
papilla
Duodenum
Bile leaves the liver in the left and right hepatic ducts, which
merge to form the common hepatic duct, which connects to the
cystic duct serving the gallbladder. The latter two ducts merge to
form a common bile duct. 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.
MEDICAL APPLICATION
Reabsorption of water from bile in the gallbladder is involved
in the formation of gallstones in the lumen of the gallblad-
der or biliary ducts, a condition called cholelithiasis.
This disorder usually originates with bile that already
contains excessive amounts of normal bile components.
Supersaturation of cholesterol in bile can lead to the
Organs Associated with the Digestive Tract
Salivary Glands
■ Salivary glands have secretory units of either protein-secreting
serous cells, usually organized in round or oval acini, or of mucin-
secreting mucous cells in elongated tubules.
■ Parotid glands have only serous acini; sublingual glands are
mixed but have primarily mucous tubules, some with serous
demilunes; submandibular glands are also mixed but have
mainly serous acini.
■ Salivary secretory units are drained by simple cuboidal intercalated
ducts, which merge as simple columnar striated ducts, which
merge further as the larger interlobular or excretory ducts.
1 Left and right hepatic ducts merge
to form a common hepatic duct.
2 Common hepatic and cystic ducts
merge to form a common bile duct.
Common bile duct
Accessory
pancreatic duct
Main pancreatic duct
3 Main pancreatic duct merges with common
bile duct at the hepatopancreatic ampulla,
which extends into the duodenum.
4 Bile and pancreatic juices enter
duodenum at the major duodenal papilla.
The main pancreatic duct merges with the common bile duct
at the hepatopancreatic ampulla, which enters the wall of the
duodenum at a major papilla (of Vater); the accessory pancreatic
duct enters the duodenum at a minor papilla. Bile and pancreatic
juices are mixed before release into the duodenal lumen.
formation of cholesterol stones, the most common form.
Brown or black pigment stones can form when bile contains
excessive amounts of unconjugated bilirubin, which can
result from chronic hemolysis associated with disorders such
as sickle cell anemia. Gallstones can lead to biliary obstruc-
tion or more commonly to in ammation in acute or chronic
cholecystitis.
SUMMARY OF KEY POINTS
■ Cells of striated ducts have mitochondria-lined, basolateral mem-
brane folds specialized for electrolyte reabsorption from the secre-
tion; excretory ducts are unusual in having strati ed cuboidal or
columnar cells.
Pancreas
■ Pancreatic islets of endocrine cells are embedded in exocrine serous
acinar tissue, which comprises most of the pancreas and in which
the cells secrete hydrolytic digestive enzymes for delivery to the
duodenum.

FIGURE 16–20 Gallbladder.
LP
LP
M
A
a b
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 bers ori-
ented in all directions to facilitate emptying of the organ; and an
external adventitia (A) where it is against the liver and a serosa
where it is exposed. (X60; H&E)
■ Each pancreatic acinar cell is pyramidal, with secretory (zymo-
gen) granules in the narrow apical end and Golgi complexes,
much rough ER, and a large nucleus at the basal end.
■ Intercalated ducts draining pancreatic acini, including their initial
centroacinar cells that insert into the acinar lumen, secrete bicar-
bonate ions (HCO3−) to neutralize chyme entering the duodenum
from the stomach.
Liver
■ Liver hepatocytes are large epithelial cells with large central nuclei
(polyploid and o en binucleated), much smooth and rough ER,
and many small Golgi complexes.
■ Hepatocytes have many functions, including endocrine (plasma
protein secretion), exocrine (bile secretion), glucose stor-
age (glycogen granules), and detoxi cation (using SER and
peroxisomes).
■ In the liver, hepatocytes are organized into irregular plates to form
polygonal hepatic lobules in which the hepatocyte plates radiate
toward a small central vein.
■ Each hepatic lobule is surrounded by sparse connective tissue that
is more abundant in the portal areas at the corners.
Biliary Tract & Gallbladder 347
C H A P T E R
G MV
1 6
Organs Associated with the Digestive Tract ■ Biliary Tract & Gallbladder
(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.
From these spaces water is quickly removed by capillaries in the
lamina propria. Abundant mitochondria provide the energy for
this pumping process. Scattered apical secretory granules (G) con-
tain mucus. (X5600)
■ Portal areas or tracts contain a small lymphatic and the portal triad:
a portal venule branch from the portal vein, a hepatic arteriole
branch of the hepatic artery, and a bile ductule branch of the biliary
tree.
■ In the lobules the portal venule and hepatic arteriole both branch
into irregular sinusoids between the hepatic plates where the nutri-
ent-rich and O2-rich blood mixes, ows past hepatocytes, and drains
to the central vein.
■ e endothelium of the hepatic sinusoids is discontinuous and
fenestrated; between it and the hepatocytes is the perisinusoidal
space (of Disse) where exchange occurs between the hepatocytes
and blood plasma.
■ e sinusoidal endothelium includes many specialized stellate
macrophages or Kup er cells, which recognize and remove e ete
erythrocytes, releasing iron and bilirubin for uptake by hepatocytes.
■ Also present in the perisinusoidal spaces are hepatic stellate cells
(or Ito cells) containing many small lipid droplets for storage of
vitamin A and other fat-soluble vitamins.
■ Between adherent hepatocytes in the hepatic plates are grooves called
bile canaliculi, sealed by tight junctions, into which hepatocytes
secrete water and bile components, including bilirubin and bile acids.