THE LYMPHATIC SYSTEM

AND IMMUNITY
22
T H E LY M P H AT I C S Y S T E M , D I S E A S E R E S I S T A N C E , A N D H O M E O S T A S I S
The lymphatic system contributes to homeostasis by draining interstitial fluid as well as providing
the mechanisms for defense against disease. •

Maintaining homeostasis in the body requires continual combat against harmful agents in our inter-
nal and external environment. Despite constant exposure to a variety of pathogens (PATH-ō-
jens), disease-producing microbes such as bacteria and viruses, most people remain healthy.
The body surface also endures cuts and bumps, exposure to ultraviolet rays in sunlight,
chemical toxins, and minor burns with an array of defensive ploys.
Immunity or resistance is the ability to ward off damage or disease through our de-
fenses. Vulnerability or lack of resistance is termed susceptibility. The two general types of
immunity are (1) innate and (2) adaptive. Innate (nonspecific) immunity refers to defenses
that are present at birth. They are always present and available to provide rapid responses to
protect us against disease. Innate immunity does not involve specific recognition of a mi-
crobe and acts against all microbes in the same way. In addition, innate immunity does not
have a memory component; that is, it cannot recall a previous contact with a foreign mole-
cule. Among the components of innate immunity are the first line of defense (the physical
and chemical barriers of the skin and mucous membranes) and the second line of defense
(antimicrobial substances, natural killer cells, phagocytes, inflammation, and fever). Innate
immune responses represent immunity’s early warning system and are designed to prevent
microbes from gaining access into the body and to help eliminate those that do gain access.
Adaptive (specific) immunity refers to defenses that involve specific recognition of a
microbe once it has breached the innate immunity defenses. Adaptive immunity is based on a
specific response to a specific microbe; that is, it adapts or adjusts to handle a specific microbe.
Unlike innate immunity, adaptive immunity is slower to respond, but it does have a memory com-
ponent. Adaptive immunity involves lymphocytes (a type of white blood cell) called T lymphocytes
(T cells) and B lymphocytes (B cells).
The body system responsible for adaptive immunity (and some aspects of innate immunity) is the
lymphatic system. This system is closely allied with the cardiovascular system, and it also functions
with the digestive system in the absorption of fatty foods. In this chapter, we will explore the mecha-
nisms that provide defenses against intruders and promote the repair of damaged body tissues.

831
832 CHAPTER 22 • THE LYMPHATIC SYSTEM AND IMMUNITY
LYMPHATIC SYSTEM STRUCTURE
AND FUNCTION
 OBJECTIVES
• List the components and major functions of the lymphatic
system.
• Describe the organization of lymphatic vessels.
• Explain the formation and flow of lymph.
• Compare the structure and functions of the primary and
secondary lymphatic organs and tissues.
The lymphatic system (lim-FAT-ik) consists of a fluid called
lymph, vessels called lymphatic vessels that transport the lymph,
a number of structures and organs containing lymphatic tissue,
and red bone marrow, where stem cells develop into the various
types of blood cells, including lymphocytes (Figure 22.1). It
assists in circulating body fluids and helps defend the body
against disease-causing agents. As you will see shortly, most
components of blood plasma filter through blood capillary walls
to form interstitial fluid. After interstitial fluid passes into lym-
phatic vessels, it is called lymph (LIMF  clear fluid). The
major difference between interstitial fluid and lymph is location:
Interstitial fluid is found between cells, and lymph is located
within lymphatic vessels and lymphatic tissue.
Lymphatic tissue is a specialized form of reticular con-
nective tissue (see Table 4.4C on page 129) that contains large
numbers of lymphocytes. Recall from Chapter 19 that lympho-
cytes are agranular white blood cells (see page 699). Two types
of lymphocytes participate in adaptive immune responses: B
cells and T cells.
Functions of the Lymphatic System
The lymphatic system has three primary functions:
1. Drains excess interstitial fluid. Lymphatic vessels drain
excess interstitial fluid from tissue spaces and return it to the blood.
2. Transports dietary lipids. Lymphatic vessels transport lipids
and lipid-soluble vitamins (A, D, E, and K) absorbed by the gas-
trointestinal tract.
3. Carries out immune responses. Lymphatic tissue initiates
highly specific responses directed against particular microbes
or abnormal cells.
Lymphatic Vessels and Lymph Circulation
Lymphatic vessels begin as lymphatic capillaries. These tiny
vessels, which are located in the spaces between cells, are closed
at one end (Figure 22.2 on page 834). Just as blood capillaries
converge to form venules and then veins, lymphatic capillaries
unite to form larger lymphatic vessels (see Figure 22.1), which
resemble veins in structure but have thinner walls and more
valves. At intervals along the lymphatic vessels, lymph flows
through lymph nodes, encapsulated bean-shaped organs consist-
ing of masses of B cells and T cells. In the skin, lymphatic ves-
sels lie in the subcutaneous tissue and generally follow the same
route as veins; lymphatic vessels of the viscera generally follow
arteries, forming plexuses (networks) around them. Tissues that
lack lymphatic capillaries include avascular tissues (such as car-
tilage, the epidermis, and the cornea of the eye), the central ner-
vous system, portions of the spleen, and red bone marrow.
Lymphatic Capillaries
Lymphatic capillaries are slightly larger in diameter than blood
capillaries and have a unique one-way structure that permits
interstitial fluid to flow into them but not out. The ends of
endothelial cells that make up the wall of a lymphatic capillary
overlap (Figure 22.2b). When pressure is greater in the inter-
stitial fluid than in lymph, the cells separate slightly, like the
opening of a one-way swinging door, and interstitial fluid enters
the lymphatic capillary. When pressure is greater inside the lym-
phatic capillary, the cells adhere more closely, and lymph cannot
escape back into interstitial fluid. The pressure is relieved as
lymph moves further down the lymphatic capillary. Attached to
the lymphatic capillaries are anchoring filaments, which contain
elastic fibers. They extend out from the lymphatic capillary,
attaching lymphatic endothelial cells to surrounding tissues.
When excess interstitial fluid accumulates and causes tissue
swelling, the anchoring filaments are pulled, making the open-
ings between cells even larger so that more fluid can flow into
the lymphatic capillary.
In the small intestine, specialized lymphatic capillaries called
lacteals (LAK-tē-als; lact-  milky) carry dietary lipids into
lymphatic vessels and ultimately into the blood. The presence of
these lipids causes the lymph draining from the small intestine to
appear creamy white; such lymph is referred to as chyle (KĪ L 
juice). Elsewhere, lymph is a clear, pale-yellow fluid.
Lymph Trunks and Ducts
As you have already learned, lymph passes from lymphatic
capillaries into lymphatic vessels and then through lymph nodes.
As lymphatic vessels exit lymph nodes in a particular region of
the body, they unite to form lymph trunks. The principal trunks
are the lumbar, intestinal, bronchomediastinal, subclavian, and
jugular trunks (Figure 22.3 on page 835). The lumbar trunks
drain lymph from the lower limbs, the wall and viscera of the
pelvis, the kidneys, the adrenal glands, and the abdominal wall.
The intestinal trunk drains lymph from the stomach, intestines,
pancreas, spleen, and part of the liver. The bronchomediastinal
trunks drain lymph from the thoracic wall, lung, and heart. The
subclavian trunks drain the upper limbs. The jugular trunks
drain the head and neck.
Lymph passes from lymph trunks into two main channels, the
thoracic duct and the right lymphatic duct, and then drains into
venous blood. The thoracic (left lymphatic) duct is
about 38– 45 cm (15–18 in.) long and begins as a dilation called
the cisterna chyli (sis-TER-na KĪ -lē; cisterna  cavity or reser-
voir) anterior to the second lumbar vertebra. The thoracic duct is
the main duct for the return of lymph to blood. The cisterna
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Figure 22.1 Components of the lymphatic system.

The lymphatic system consists of lymph, lymphatic vessels, lymphatic tissues, and red bone marrow.

Palatine tonsil
Submandibular node Left internal jugular vein
Cervical node
Left subclavian vein
Right internal jugular vein
Right lymphatic duct
Right subclavian vein
Thoracic duct
Thymus

Axillary node
Lymphatic vessel

Thoracic duct
Spleen

Cisterna chyli (b) Areas drained by

right lymphatic and
Aggregated lymphatic thoracic ducts
Intestinal node follicle (Peyer’s patch)
Small intestine Area drained by
Large intestine right lymphatic duct
Iliac node Area drained by
Appendix thoracic duct

Red bone marrow Inguinal node

Functions
Lymphatic vessel
1. Drains excess interstitial fluid.
2. Transports dietary lipids from the gastrointestinal tract to the blood.
3. Protects against invasion through immune responses.

Anterior view of principal components of lymphatic system

? What tissue contains stem cells that develop into lymphocytes?

833
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834 CHAPTER 22 • THE LYMPHATIC SYSTEM AND IMMUNITY
Figure 22.2 Lymphatic capillaries.
Lymphatic capillaries are found throughout the body except in avascular tissues, the central nervous system,
portions of the spleen, and bone marrow.
Blood
capillary
Venule
Tissue cell Arteriole
Blood Blood
Lymph
Interstitial fluid Lymphatic
capillary
(a) Relationship of lymphatic capillaries
to tissue cells and blood capillaries
? Is lymph more similar to blood plasma or to interstitial fluid? Why?
chyli receives lymph from the right and left lumbar trunks and
from the intestinal trunk. In the neck, the thoracic duct also
receives lymph from the left jugular, left subclavian, and left
bronchomediastinal trunks. Therefore, the thoracic duct re-
ceives lymph from the left side of the head, neck, and chest,
the left upper limb, and the entire body inferior to the ribs
(see Figure 22.1b). The thoracic duct in turn drains lymph into
venous blood at the junction of the left internal jugular and left
subclavian veins.
The right lymphatic duct (Figure 22.3) is about 1.2 cm (0.5
in.) long and receives lymph from the right jugular, right subcla-
vian, and right bronchomediastinal trunks. Thus, the right lym-
phatic duct receives lymph from the upper right side of the body
(see Figure 22.1b). From the right lymphatic duct, lymph drains
into venous blood at the junction of the right internal jugular and
right subclavian veins.
Formation and Flow of Lymph
Most components of blood plasma filter freely through the capil-
lary walls to form interstitial fluid, but more fluid filters out
of blood capillaries than returns to them by reabsorption (see
Figure 21.7 on page 771). The excess filtered fluid—about
3 liters per day—drains into lymphatic vessels and becomes
lymph. Because most plasma proteins are too large to leave
blood vessels, interstitial fluid contains only a small amount of
protein. Proteins that do leave blood plasma cannot return to the
blood by diffusion because the concentration gradient (high level
of proteins inside blood capillaries, low level outside) opposes
Lymph
Endothelium
of lymphatic
capillary
Tissue cell
Interstitial fluid
Anchoring
filament
Opening
(b) Details of a lymphatic capillary
such movement. The proteins can, however, move readily
through the more permeable lymphatic capillaries into lymph.
Thus, an important function of lymphatic vessels is to return the
lost plasma proteins to the bloodstream.
Like veins, lymphatic vessels contain valves, which ensure
the one-way movement of lymph. As noted previously, lymph
drains into venous blood through the right lymphatic duct and
the thoracic duct at the junction of the internal jugular and sub-
clavian veins (Figure 22.3). Thus, the sequence of fluid flow
is blood capillaries (blood) : interstitial spaces (interstitial
fluid) : lymphatic capillaries (lymph) : lymphatic vessels
(lymph) : lymphatic ducts (lymph) : junction of the internal
jugular and subclavian veins (blood). Figure 22.4 on page 837
illustrates this sequence, along with the relationship of the lym-
phatic and cardiovascular systems.
The same two “pumps” that aid the return of venous blood to
the heart maintain the flow of lymph.
1. Skeletal muscle pump. The “milking action” of skeletal
muscle contractions (see Figure 21.9 on page 774) compresses
lymphatic vessels (as well as veins) and forces lymph toward the
junction of the internal jugular and subclavian veins.
2. Respiratory pump. Lymph flow is also maintained by
pressure changes that occur during inhalation (breathing in).
Lymph flows from the abdominal region, where the pressure is
higher, toward the thoracic region, where it is lower. When the
pressures reverse during exhalation (breathing out), the valves
prevent backflow of lymph. In addition, when a lymphatic vessel
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LYMPHATIC SYSTEM STRUCTURE AND FUNCTION 835

Figure 22.3 Routes for drainage of lymph from lymph trunks into the thoracic and right lymphatic ducts.
All lymph returns to the bloodstream through the thoracic (left) lymphatic duct and right lymphatic duct.

Right internal jugular vein Left internal jugular vein

RIGHT LEFT JUGULAR TRUNK
JUGULAR
LEFT SUBCLAVIAN TRUNK
TRUNK

RIGHT THORACIC (LEFT LYMPHATIC) DUCT

SUBCLAVIAN
TRUNK Left subclavian vein
Left brachiocephalic vein
Right subclavian First rib
vein Accessory hemiazygos vein

LEFT BRONCHO-
RIGHT LYMPHATIC MEDIASTINAL TRUNK
DUCT
Right brachiocephalic THORACIC (LEFT
vein LYMPHATIC) DUCT
RIGHT BRONCHO-
MEDIASTINAL TRUNK
Hemiazygos vein
Superior vena cava

Rib

Intercostal
muscle

Azygos vein

LEFT LUMBAR
CISTERNA CHYLI TRUNK
RIGHT LUMBAR
TRUNK
Inferior vena cava INTESTINAL
TRUNK

(a) Overall anterior view

Right jugular trunk

Right subclavian trunk Left jugular trunk

Left subclavian trunk

Thoracic (left lymphatic) duct

Right lymphatic duct

Right bronchomediastinal trunk

Left bronchomediastinal trunk

(b) Detailed anterior view

? Which lymphatic vessels empty into the cisterna chyli, and which duct receives lymph from the cisterna chyli?
836 CHAPTER 22 • THE LYMPHATIC SYSTEM AND IMMUNITY

Figure 22.4 Schematic diagram showing the relationship of the lymphatic system to the cardiovascular system.
The arrows indicate the direction of flow of lymph and blood.
The sequence of fluid flow is blood capillaries (blood) : interstitial spaces (interstitial fluid) : lymphatic
capillaries (lymph) : lymphatic vessels (lymph) : lymphatic ducts (lymph) : junction of the internal
jugular and subclavian veins (blood).
SYSTEMIC CIRCULATION PULMONARY CIRCULATION

Lymph node
Lymphatic capillaries

Lymphatic
duct
Subclavian vein Pulmonary blood
capillaries

Veins
Lymphatic vessel

Valve

Heart Arteries

Lymph node
Systemic blood capillaries

Lymphatic capillaries

? Does inhalation promote or hinder the flow of lymph?

distends, the smooth muscle in its wall contracts, which helps
move lymph from one segment of the vessel to the next.
Lymphatic Organs and Tissues
The widely distributed lymphatic organs and tissues are clas-
sified into two groups based on their functions. Primary
lymphatic organs are the sites where stem cells divide and
become immunocompetent, that is, capable of mounting an im-
mune response. The primary lymphatic organs are the red bone
marrow (in flat bones and the epiphyses of long bones of adults)
and the thymus. Pluripotent stem cells in red bone marrow give
rise to mature, immunocompetent B cells and to pre-T cells. The
pre-T cells in turn migrate to the thymus, where they become
immunocompetent T cells. The secondary lymphatic organs
and tissues are the sites where most immune responses occur.
They include lymph nodes, the spleen, and lymphatic nodules
(follicles). The thymus, lymph nodes, and spleen are considered
organs because each is surrounded by a connective tissue cap-
sule; lymphatic nodules, in contrast, are not considered organs
because they lack a capsule.
Thymus
The thymus is a bilobed organ located in the mediastinum
between the sternum and the aorta (Figure 22.5a). An envelop-
ing layer of connective tissue holds the two lobes closely
together, but a connective tissue capsule separates the two.
Extensions of the capsule, called trabeculae (tra-BEK-ū-lē 
little beams), penetrate inward and divide each lobe into lobules
(Figure 22.5b).
Each thymic lobule consists of a deeply staining outer cortex
and a lighter-staining central medulla (Figure 22.5b). The cortex
is composed of large numbers of T cells and scattered dendritic
cells, epithelial cells, and macrophages. Immature T cells (pre-T
cells) migrate from red bone marrow to the cortex of the thymus,
where they proliferate and begin to mature. Dendritic cells
(dendr-  a tree), which are derived from monocytes, and so
named because they have long, branched projections that resem-
ble the dendrites of a neuron, assist the maturation process. As
you will see shortly, dendritic cells in other parts of the body,
such as lymph nodes, play another key role in immune re-
sponses. Each of the specialized epithelial cells in the cortex has
several long processes that surround and serve as a framework
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LYMPHATIC SYSTEM STRUCTURE AND FUNCTION 837

Figure 22.5 Thymus. (See Tortora, A Photographic Atlas of the Human Body, Second Edition, Figure 7.2a.)
The bilobed thymus is largest at puberty and then atrophies with age.

Blood vessels

Capsule
Lobule:
Cortex
Thymic (Hassall’s)
corpuscle
Thyroid Medulla
Brachiocephalic
gland veins Trabecula
Trachea
Right common Thymus
carotid artery

Superior Parietal
vena cava pericardium
LM 30x

(b) Thymic lobules

Right Left
lung lung
T cell

Thymic (Hassall’s)
corpuscle

Epithelial cell

Diaphragm

(a) Thymus of adolescent LM 385x

(c) Details of the thymic medulla

? Which type of lymphocytes mature in the thymus?

for as many as 50 T cells. These epithelial cells help “educate”
the pre-T cells in a process known as positive selection (see
Figure 22.22). Additionally, they produce thymic hormones that
are thought to aid in the maturation of T cells. Only about 2% of
developing T cells survive in the cortex. The remaining cells die
via apoptosis (programmed cell death). Thymic macrophages
help clear out the debris of dead and dying cells. The surviving
T cells enter the medulla.
The medulla consists of widely scattered, more mature
T cells, epithelial cells, dendritic cells, and macrophages (Figure
22.5c). Some of the epithelial cells become arranged into
concentric layers of flat cells that degenerate and become filled
with keratohyalin granules and keratin. These clusters are called
thymic (Hassall’s) corpuscles. Although their role is uncertain,
they may serve as sites of T cell death in the medulla. T cells
that leave the thymus via the blood migrate to lymph nodes, the
spleen, and other lymphatic tissues where they colonize parts of
these organs and tissues.
In infants, the thymus is large, with a mass of about 70 g (2.3 oz).
After puberty, adipose and areolar connective tissue begin to re-
place the thymic tissue. By the time a person reaches maturity,
the gland has atrophied considerably, and in old age it may
weigh only 3 g (0.1 oz). Before the thymus atrophies, it popu-
lates the secondary lymphatic organs and tissues with T cells.
However, some T cells continue to proliferate in the thymus
throughout an individual’s lifetime.
Lymph Nodes
Located along lymphatic vessels are about 600 bean-shaped
lymph nodes. They are scattered throughout the body, both
superficially and deep, and usually occur in groups (see Figure
22.1). Large groups of lymph nodes are present near the mam-
mary glands and in the axillae and groin.
Lymph nodes are 1–25 mm (0.04–1 in.) long and, like the
thymus, are covered by a capsule of dense connective tissue that
extends into the node (Figure 22.6). The capsular extensions,
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838 CHAPTER 22 • THE LYMPHATIC SYSTEM AND IMMUNITY

Figure 22.6 Structure of a lymph node. Arrows indicate the direction of lymph flow through a lymph node.
Lymph nodes are present throughout the body, usually clustered in groups.

Outer Cortex

Cells of inner cortex Cells around germinal center Cells in germinal center

T cells B cells B cells Follicular Macrophages

Dendritic dendritic
cells cells

Cells of medulla
Subcapsular sinus
Reticular fiber
Trabecula

B cells Plasma Macrophages Trabecular sinus

cells Outer cortex:
Germinal center
in secondary lymphatic
nodule
Afferent lymphatic
Cells around germinal
vessel
center
Inner cortex
Medulla
Medullary sinus
Valve
Reticular fiber

Efferent lymphatic
vessels

Valve

Hilum

Route of lymph flow

through a lymph node:
Afferent lymphatic vessel

Subcapsular sinus

Trabecular sinus
Capsule
Afferent
lymphatic vessel Medullary sinus

Efferent lymphatic vessel

(a) Partially sectioned lymph node
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Capsule
Subcapsular sinus
Trabecula
Trabecular sinus
Outer cortex
Germinal center in
secondary lymphatic
nodule
Inner cortex
Medullary sinus
Medulla
LM 55x
(b) Portion of a lymph node
? What happens to foreign substances in lymph that enter a lymph node?
called trabeculae, divide the node into compartments, provide
support, and provide a route for blood vessels into the interior
of a node. Internal to the capsule is a supporting network of
reticular fibers and fibroblasts. The capsule, trabeculae, reticular
fibers, and fibroblasts constitute the stroma (supporting connec-
tive tissue) of a lymph node.
The parenchyma (functioning part) of a lymph node is
divided into a superficial cortex and a deep medulla. The cortex
consists of an outer cortex and an inner cortex. Within the outer
cortex are egg-shaped aggregates of B cells called lymphatic
nodules (follicles). A lymphatic nodule consisting chiefly of B
cells is called a primary lymphatic nodule. Most lymphatic nod-
ules in the outer cortex are secondary lymphatic nodules (Figure
22.6), which form in response to an antigenic challenge and are
sites of plasma cell and memory B cell formation. After B cells
in a primary lymphatic nodule recognize an antigen, the primary
lymphatic nodule develops into a secondary lymphatic nodule.
The center of a secondary lymphatic nodule contains a region of
light-staining cells called a germinal center. In the germinal cen-
ter are B cells, follicular dendritic cells (a special type of den-
dritic cell), and macrophages. When follicular dendritic cells
“present” an antigen (described later in the chapter), B cells pro-
liferate and develop into antibody-producing plasma cells or de-
velop into memory B cells. Memory B cells persist after an ini-
tial immune response and “remember” having encountered a
specific antigen. B cells that do not develop properly undergo
apoptosis (programmed cell death) and are destroyed by
macrophages. The region of a secondary lymphatic nodule sur-
rounding the germinal center is composed of dense accumula-
tions of B cells that have migrated away from their site of origin
within the nodule.
The inner cortex does not contain lymphatic nodules. It con-
sists mainly of T cells and dendritic cells that enter a lymph
node from other tissues. The dendritic cells present antigens to
LYMPHATIC SYSTEM STRUCTURE AND FUNCTION 839
Nerve
Skeletal muscle
Lymph node
Lymphatic vessel
(c) Anterior view of an inguinal lymph node
T cells, causing their proliferation. The newly formed T cells
then migrate from the lymph node to areas of the body where
there is antigenic activity.
The medulla of a lymph node contains B cells, antibody
producing plasma cells that have migrated out of the cortex into
the medulla, and macrophages. The various cells are embedded
in a network of reticular fibers and reticular cells.
As you have already learned, lymph flows through a node in
one direction only (Figure 22.6a). It enters through afferent
lymphatic vessels (afferent  to carry toward), which penetrate
the convex surface of the node at several points. The afferent
vessels contain valves that open toward the center of the
node, directing the lymph inward. Within the node, lymph enters
sinuses, a series of irregular channels that contain branching
reticular fibers, lymphocytes, and macrophages. From the affer-
ent lymphatic vessels, lymph flows into the subcapsular sinus,
immediately beneath the capsule. From here the lymph flows
through trabecular sinuses, which extend through the cortex
parallel to the trabeculae, and into medullary sinuses, which
extend through the medulla. The medullary sinuses drain into
one or two efferent lymphatic vessels (efferent  to carry
away), which are wider and fewer in number than afferent
vessels. They contain valves that open away from the center of
the lymph node to convey lymph, antibodies secreted by plasma
cells, and activated T cells out of the node. Efferent lymphatic
vessels emerge from one side of the lymph node at a slight
depression called a hilum (HĪ -lum). Blood vessels also enter
and leave the node at the hilum.
Lymph nodes function as a type of filter. As lymph enters one
end of a lymph node, foreign substances are trapped by the retic-
ular fibers within the sinuses of the lymph node. Then
macrophages destroy some foreign substances by phagocytosis
while lymphocytes destroy others by immune responses. The
filtered lymph then leaves the other end of the lymph node.
840 CHAPTER 22 • THE LYMPHATIC SYSTEM AND IMMUNITY

• CLINICAL CONNECTION Metastasis Through Spleen

Lymphatic Vessels
Metastasis (me-TAS-ta-sis; meta-  beyond; stasis  to stand), the
spread of a disease from one part of the body to another, can occur via
lymphatic vessels. All malignant tumors eventually metastasize. Cancer
cells may travel in the blood or lymph and establish new tumors where
they lodge. When metastasis occurs via lymphatic vessels, secondary
tumor sites can be predicted according to the direction of lymph flow
from the primary tumor site. Cancerous lymph nodes feel enlarged, firm,
nontender, and fixed to underlying structures. By contrast, most lymph
nodes that are enlarged due to an infection are softer, tender, and
movable. •
The oval spleen is the largest single mass of lymphatic tissue
in the body, measuring about 12 cm (5 in.) in length (Figure 22.7a).
It is located in the left hypochondriac region between the
stomach and diaphragm. The superior surface of the spleen is
smooth and convex and conforms to the concave surface of the
diaphragm. Neighboring organs make indentations in the vis-
ceral surface of the spleen—the gastric impression (stomach),
the renal impression (left kidney), and the colic impression (left
colic flexure of large intestine). Like lymph nodes, the spleen
has a hilum. Through it pass the splenic artery, splenic vein, and
efferent lymphatic vessels.
A capsule of dense connective tissue surrounds the spleen and
is covered in turn by a serous membrane, the visceral

Figure 22.7 Structure of the spleen. (See Tortora, A Photographic Atlas of the Human Body, Second Edition, Figure 7.3a.)
The spleen is the largest single mass of lymphatic tissue in the body.

SUPERIOR

Splenic vein
Splenic artery
Colic impression
Gastric impression
Hilum
POSTERIOR ANTERIOR
Renal impression

(a) Visceral surface

Splenic artery Capsule

Splenic vein

White pulp

Red pulp:
Venous sinus Red pulp
Splenic cord
Central artery
White pulp
Trabecula
Central artery
Capsule

(b) Internal structure

Trabecula
LM 25x
(c) Portion of the spleen

? After birth, what are the main functions of the spleen?


peritoneum. Trabeculae extend inward from the capsule. The
capsule plus trabeculae, reticular fibers, and fibroblasts consti-
tute the stroma of the spleen; the parenchyma of the spleen
consists of two different kinds of tissue called white pulp and
red pulp (Figure 22.7b, c). White pulp is lymphatic tissue,
consisting mostly of lymphocytes and macrophages arranged
around branches of the splenic artery called central arteries.
The red pulp consists of blood-filled venous sinuses and cords
of splenic tissue called splenic (Billroth’s) cords. Splenic cords
consist of red blood cells, macrophages, lymphocytes, plasma
cells, and granulocytes. Veins are closely associated with the
red pulp.
Blood flowing into the spleen through the splenic artery
enters the central arteries of the white pulp. Within the white
pulp, B cells and T cells carry out immune functions, similar to
lymph nodes, while spleen macrophages destroy blood-borne
pathogens by phagocytosis. Within the red pulp, the spleen
performs three functions related to blood cells: (1) removal by
macrophages of ruptured, worn out, or defective blood cells and
platelets; (2) storage of platelets, up to one-third of the body’s
supply; and (3) production of blood cells (hemopoiesis) during
fetal life.
• CLINI C AL CONNECTION Ruptured Spleen
The spleen is the organ most often damaged in cases of abdominal
trauma. Severe blows over the inferior left chest or superior abdomen
can fracture the protecting ribs. Such crushing injury may result in a
ruptured spleen, which causes significant hemorrhage and shock.
Prompt removal of the spleen, called a splenectomy, is needed to pre-
vent death due to bleeding. Other structures, particularly red bone mar-
row and the liver, can take over some functions normally carried out by
the spleen. Immune functions, however, decrease in the absence of a
spleen. The spleen’s absence also places the patient at higher risk for
sepsis (a blood infection) due to loss of the filtering and phagocytic
functions of the spleen. To reduce the risk of sepsis, patients who have
undergone a splenectomy take prophylactic (preventive) antibiotics be-
fore any invasive procedures. •
Lymphatic Nodules
Lymphatic nodules (follicles) are egg-shaped masses of lym-
phatic tissue that are not surrounded by a capsule. Because
they are scattered throughout the lamina propria (connective
tissue) of mucous membranes lining the gastrointestinal, urinary,
and reproductive tracts and the respiratory airways, lymphatic
nodules in these areas are also referred to as mucosa-associated
lymphatic tissue (MALT).
Although many lymphatic nodules are small and solitary,
some occur in multiple large aggregations in specific parts of the
body. Among these are the tonsils in the pharyngeal region and
the aggregated lymphatic follicles (Peyer’s patches) in the ileum
of the small intestine. Aggregations of lymphatic nodules also
occur in the appendix. Usually there are five tonsils, which form
a ring at the junction of the oral cavity and oropharynx and at the
junction of the nasal cavity and nasopharynx (see Figure 23.2b).
The tonsils are strategically positioned to participate in immune
DEVELOPMENT OF LYMPHATIC TISSUES 841
responses against inhaled or ingested foreign substances. The
single pharyngeal tonsil (fa-RIN-jē-al) or adenoid is embedded
in the posterior wall of the nasopharynx. The two palatine
tonsils (PAL-a-tı̄n) lie at the posterior region of the oral cavity,
one on either side; these are the tonsils commonly removed in
a tonsillectomy. The paired lingual tonsils (LIN-gwal), located
at the base of the tongue, may also require removal during a
tonsillectomy.
 CHECKPOINT
1. How are interstitial fluid and lymph similar, and how do
they differ?
2. How do lymphatic vessels differ in structure from veins?
3. Diagram the route of lymph circulation.
4. What is the role of the thymus in immunity?
5. What functions do lymph nodes, the spleen, and the
tonsils serve?
DEVELOPMENT OF
LYMPHATIC TISSUES
 OBJECTIVE
• Describe the development of lymphatic tissues.
Lymphatic tissues begin to develop by the end of the fifth
week of embryonic life. Lymphatic vessels develop from lymph
sacs that arise from developing veins, which are derived
from mesoderm.
The first lymph sacs to appear are the paired jugular
lymph sacs at the junction of the internal jugular and subcla-
vian veins (Figure 22.8). From the jugular lymph sacs, lym-
phatic capillary plexuses spread to the thorax, upper limbs,
Figure 22.8 Development of lymphatic tissues.
Lymphatic tissues are derived from mesoderm.
Jugular
lymph sac Subclavian
vein
Thoracic
duct
Inferior
Cisterna vena cava
chyli
Posterior
lymph sac
? When do lymphatic tissues begin to develop?