Chapter 7: CARTILAGE  Epiphyseal plates of long bones, where it makes

Junqueira’s Basic Histology: Text and Atlas possible longitudinal bone growth.

In the embryo:
 Hyaline cartilage forms the temporary skeleton
that is gradually replaced by bone.
Cartilage is a tough, durable form of supporting connective
tissue, characterized by an extracellular matrix (ECM) with
high concentrations of GAGs and proteoglycans, interacting
with collagen and elastic fibers.

Functions:

 Allows the tissue to bear mechanical stresses

without permanent distortion.
 Forms the framework supporting softer tissues
(respiratory tract, ears, and nose, cartilage).
 Provides cushioning and sliding regions within
skeletal joints and facilitates bone movements.
 Guides development and growth of long bones,
both before and after birth.
 The high content of bound water allows cartilage to
serve as a shock absorber, an important
functional role.

CHONDROCYTES (Gr. chondros, cartilage + kytos, cell)-

cells of cartilage that is embedded in the ECM. MEDICAL APPLICATION:
Lacunae - matrix cavities Osteoarthritis, a chronic condition that commonly
The physical properties of cartilage depend on electrostatic occurs during aging, involves the gradual loss or
changed physical properties of the hyaline cartilage that
bonds between type II collagen fibrils, hyaluronan, and the
sulfated GAGs on densely packed proteoglycans. lines the articular ends of bones in joints.

All types of cartilage lack vascular supplies and

chondrocytes receive nutrients by diffusion from capillaries Chondrocytes
in surrounding connective tissue (the perichondrium).
At the periphery: young chondrocytes or chondroblasts
Chondrocytes exhibit low metabolic activity. Cartilage also
have an elliptic shape, with the long axes parallel to the
lacks nerves.
surface.

Deeper: they are round and may appear in groups of up to

TYPES of Cartilage: eight cells that originate from mitotic divisions of a single
chondroblast and are called isogenous aggregates.
A. HYALINE CARTILAGE
As the chondrocytes become more active in secreting
Hyaline (Gr. hyalos, glass) cartilage, the most common of the collagens and other ECM components, the aggregated cells
three types, is homogeneous and semitransparent in the are pushed apart and occupy separate lacunae.
fresh state.
HC cells metabolize glucose mainly by anaerobic
In adults’ hyaline cartilage: glycolysis.
 Articular surfaces of movable joints,
 Walls of larger respiratory passages (nose, larynx, Nutrients from the blood diffuse to all the chondrocytes
trachea, bronchi), from the cartilage surface, with movements of water and
 Ventral ends of ribs, where they articulate with the solutes in the cartilage matrix promoted by intermittent
sternum tissue compression and decompression during body
movements. The limits of such diffusion define the

HISTOLOGY: CARTILAGE 1
maximum thickness of hyaline cartilage, which usually
exists as small, thin plates.

Perichondrium
C. FIBROCARTILAGE
A sheath of dense connective tissue that surrounds
cartilage. Essential for the growth and maintenance of Fibrocartilage takes various forms in different structures
cartilage. The outer region of the perichondrium consists but is essentially a mingling of hyaline cartilage and dense
largely of collagen type I fibers and fibroblasts, but an inner connective tissue.
layer adjoining the cartilage matrix also contains Found in:
mesenchymal stem cells which provide a source for new
chondroblasts that divide and differentiate into  Intervertebral discs, and act as lubricated cushions and
chondrocytes. Harbors the blood supply serving the shock absorbers preventing damage to adjacent
cartilage and a small neural component. vertebrae from abrasive forces or impacts.
 Attachments of certain ligaments,
 Pubic symphysis—all places where it serves as very
tough, yet cushioning support tissue for bone.

Chondrocytes of fibrocartilage occur singly and often in

aligned isogenous aggregates, producing type II collagen
and other ECM components. The relative scarcity of
proteoglycans overall makes fibrocartilage matrix more
acidophilic than that of hyaline or elastic cartilage. There is
no distinct surrounding perichondrium in
fibrocartilage.

Important features of the three major types of cartilage

are summarized in Table 7–1.

B. ELASTIC CARTILAGE

Elastic cartilage is essentially similar to hyaline cartilage

except that it contains an abundant network of elastic fibers
in addition to a meshwork of collagen type II fibrils, which
give fresh elastic cartilage a yellowish color.

More flexible than hyaline cartilage,

Found in:

 Auricle of the ear,

 the walls of the external auditory canals,
 the auditory (Eustachian) tubes,
 the epiglottis
 Upper respiratory tract.

Elastic cartilage in these locations includes a

perichondrium similar to that of most hyaline cartilage.
Throughout elastic cartilage the cells resemble those of
hyaline cartilage both physiologically and structurally.

HISTOLOGY: CARTILAGE 2
CARTILAGE FORMATION, GROWTH, & REPAIR

All cartilage forms from embryonic mesenchyme in the

process of chondrogenesis.

The first indication of cell differentiation is the rounding up

of the mesenchymal cells, which retract their extensions,
multiply rapidly, and become more densely packed
together.

In general the terms “chondroblasts” and

“chondrocytes” respectively refer to the cartilage cells
during and after the period of rapid proliferation.

At both stages the cells have basophilic cytoplasm rich in

RER for collagen synthesis. Production of the ECM encloses
the cells in their lacunae and then gradually separates
chondroblasts from one another.

During embryonic development, the cartilage

differentiation takes place primarily from the center
outward; therefore the more central cells have the
characteristics of chondrocytes, whereas the peripheral
cells are typical chondroblasts. The superficial mesenchyme
develops as the perichondrium.

Once formed, the cartilage tissue enlarges both by

interstitial growth, involving mitotic division of preexisting
chondrocytes, and by appositional growth, which involves
chondroblast differentiation from progenitor.

Fibrocartilage varies histologically in different structures,

but is always essentially a mixture of hyaline cartilage
and dense connective tissue.

Appositional growth of cartilage is more important during

postnatal development, although as described in Chapter 8,
interstitial growth in cartilaginous regions within long
bones is important in increasing the length of these
structures. In articular cartilage, cells and matrix near the
articulating surface are gradually worn away and must be
replaced from within, because there is no perichondrium to
add cells by appositional growth.

Except in young children, damaged cartilage undergoes

slow and often incomplete repair, primarily dependent on
cells in the perichondrium which invade the injured
area and produce new cartilage. In damaged areas the
perichondrium produces a scar of dense connective tissue
instead of forming new cartilage. The poor capacity of
cartilage for repair or regeneration is due in part to its
avascularity and low metabolic rate.

HISTOLOGY: CARTILAGE 3
 ■ Fibrocartilage contains varying combinations of hyaline
cartilage in small amounts of dense connective tissue.

■ Histologically it consists of small chondrocytes in a

hyaline matrix, usually layered with larger areas of bundled
type I collagen with scattered fibroblasts.

■ Fibrocartilage provides very tough, strong support at

tendon insertions and in intervertebral discs and certain
Cartilage SUMMARY OF KEY POINTS other joints. Cartilage Formation, Growth, & Repair
■ Cartilage is a tough, resilient type of connective tissue that ■ All forms of cartilage form from embryonic mesenchyme.
structurally supports certain soft tissues, notably in the
respiratory tract, and provides cushioned, low-friction ■ Cartilaginous structures grow by mitosis of existing
surfaces in joints. chondroblasts in lacunae (interstitial growth) or formation
of new chondroblasts peripherally from progenitor cells in
■ Cells of cartilage, chondrocytes, make up a small the perichondrium (appositional growth).
percentage of the tissue’s mass, which is mainly a flexible
mass of extracellular matrix (ECM). ■ Repair or replacement of injured cartilage is very slow
and ineffective, due in part to the tissue’s avascularity and
■ Chondrocytes are embedded within lacunae surrounded low metabolic rate.
by the ECM.

■ Cartilage ECM typically includes collagen as well as

abundant proteoglycans, notably aggrecan, which bind a
large amount of water.

■ Cartilage always lacks blood vessels, lymphatics, and Cartilage ASSESS YOUR KNOWLEDGE
nerves, but it is usually surrounded by a dense connective
tissue perichondrium that is vascularized.

■ There are three major forms of cartilage: (1) hyaline 1. The molecular basis for the shock absorbing properties of
cartilage, (2) elastic cartilage, and (3) fibrocartilage. Hyaline cartilage involves which of the following?
Cartilage a. Electrostatic interaction of proteoglycans with type IV
collagen
■ The ECM of hyaline cartilage is homogenous and glassy, b. Ability of glycosaminoglycans to bind anions
rich in fibrils of type II collagen and aggrecan complexes c. Noncovalent binding of glycosaminoglycans to protein
with bound water. cores
■ The ECM has less collagen and more proteoglycan d. Sialic acid residues in the glycoproteins
immediately around the lacunae, producing slight staining e. Hydration of glycosaminoglycans
differences in this territorial matrix.
2. What distinguishes cartilage from most other connective
■ Chondrocytes occur singly or in small, mitotically derived tissues?
isogenous groups. a. Its extracellular matrix is rich in collagen.
b. Its predominant cell type is a mesenchymal derivative. c.
■ Perichondrium is usually present, but not at the hyaline
Its predominant cell type secretes both fibers and
cartilage of articular surfaces or the epiphyses of growing
proteoglycans.
long bones. Elastic Cartilage
d. It lacks blood vessels.
■ Elastic cartilage generally resembles hyaline cartilage in e. It functions in mechanical support.
its chondrocytes and major ECM components, but its matrix
includes abundant elastic fibers, visible with special stains, 3. Which feature is typical of elastic cartilage?
which increase the tissue’s flexibility. a. Primary skeletal tissue in the fetus
b. No identifiable perichondrium
■ Elastic cartilage provides flexible support for the external c. Found in intervertebral discs
ear as well as certain structures of the middle ear and d. Most widely distributed cartilage type in the body
larynx; it is always surrounded by perichondrium. e. Collagen is mainly type II
Fibrocartilage

HISTOLOGY: CARTILAGE 4
4. Which area in cartilage is relatively collagen-poor and nasal cartilage resulting in a “saddle nose” deformity.
proteoglycan-rich? Erosions in the enamel of her front teeth are noted. The
a. Fibrocartilage breakdown of the nasal cartilage releases collagen fibers
b. Territorial matrix primarily of which type?
c. Epiphyseal plate a. Type I
d. Interterritorial matrix b. Type II
e. Perichondrium c. Type III
d. Type IV
e. Type VII

5. What is the source of the mesenchymal progenitor cells 10. A 66-year-old man who suffered from severe
activated for the repair of hyaline cartilage of accident- osteoarthritis is referred to an orthopedic surgeon for
damaged costal cartilages? replacement of his right knee. He had been actively involved
a. Perichondrium in both high school and intercollegiate football and had
b. Adjacent loose connective tissue continued running until about the age of 45 as a form of
c. Bone of the adjacent rib(s) and sternum relaxation and exercise. With the patient’s permission the
d. Chondrocytes of the injured cartilage removed joint is used by investigators performing a
e. Stem cells circulating with blood proteomic analysis of different joint tissues. The meniscus
was found to contain almost exclusively type I collagen and
6. How does articular cartilage differ from most other aggrecan was undetectable. What is the most likely
hyaline cartilage? explanation for this result?
a. It undergoes mainly appositional growth.
b. It contains isogenous groups of chondrocytes. a. The meniscus normally consists of dense regular
c. It lacks a perichondrium. connective tissue, which contains primarily type I collagen.
d. Its matrix contains aggrecan. b. The meniscus normally consists of fibrocartilage, which
e. It is derived from embryonic mesenchyme. contains only type I collagen.
c. The meniscus had undergone repeated rounds of repair
due to wear-and-tear during which its hyaline cartilage
7. Which step occurs first in chondrogenesis? component was replaced by dense connective tissue.
a. Appositional growth d. Osteoarthritic injury in the knee resulted in the
b. Conversion of chondroblasts to chondrocytes chondrocytes of the meniscus switching from expression of
c. Formation of mesenchymal condensations genes for type II collagen to type I collagen.
d. Interstitial growth e. Elastic cartilage is normally replaced by fibrocartilage
e. Secretion of collagen-rich and proteoglycan-rich matrix during aging and this process can be accelerated by
exercise.
8. Osteoarthritis is characterized by the progressive erosion
of articular cartilage. The matrix metalloproteinases
involved in this erosion primarily act on which matrix Answers: 1e, 2d, 3e, 4b, 5a, 6c, 7c, 8d, 9b, 10c
component?
a. Aggrecan
b. Link proteins
c. Network-forming collagen
d. Fibril-forming collagen
e. Chondronectin

9. A 28-year-old woman visits the family medicine clinic

complaining of loss of the sense of smell, nosebleeds,
problems with swallowing, and hoarseness. She admits to
“casual, social use” of cocaine on a regular basis since her
sophomore year of college. A complete examination of her
nose with a speculum and otoscope shows severe rhinitis
(inflammation). There is also perforation and collapse of the

HISTOLOGY: CARTILAGE 5