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The continuity of the mesenchyme holds the body together, and

this characteristic is maintained through all the later development.
Through the pathways of the mesenchymal system every part of the body
is accessible from every other part. For this reason, the mesenchymal
tissue becomes the carrier of all the conduction systems: vessels, nerves,
and ducts. While some constituents of the piesenchyme (e.g., cartilage,
bone) remain in an indifferent stage and only later begin their tissuebuilding functions, one part differentiates early into embryonic
connective tissue (Fig. 49). This consists of stellate, branched, spindleshaped cells with relatively large nuclei, poor in chromatin. The
cytoplasmic processes anastomose so that the embryonic connective
tissue may be regarded as a reticular syncytium, the meshes of which are
filled by a soft, viscous, interstitial substance (intercellular or ground
From the embryonic connective tissue, which in embryos and
fetuses covers the whole body under the ectoderm and fills all the spaces
between organs, all other forms of connective tissue develop. In the
process the cells undergo changes, and fibrillar structures appear in the
intercellular substance. We distinguish collage-nous, elastic, and
reticular fibers.
Collagenous Fibers (Figs. 54, f *; 57,a). They consist of very fine
fibrils (0.3-0.5 n thick), which are smooth, unbranched, and usually
wavy. The fibrils are always, united by a mucoid cement into bundles
(fibers), which, in contrast to fibrils, may-branch. The collagenous fibers
usually unite to form strands and trabeculae. They are pale (i.e., only
slightly refractive), very slightly extensible, and possessed of great
tensile strength. (The Achilles tendon, for example, can withstand 6 kg.
per sq. mm.) When boiled, they yield glue, whence the term collagenous.
Their principal constituent is the protein collagen. In dilute acids and
alkalies they swell and gradually dissolve. They stain red with eosin and
van Gieson's stain, blue with Mallory's stain.
Elastic Fibers (Figs. 54,g*; 57,b,c). They may be very fine or very
thick (18 ). They are highly refractive (bright with dark borders). They
branch but do not form bundles. The ends are often curved like a
shepherd's crook. They are often mixed, singly or .as nets, with
collagenous fibers. They may be selectively stained brown with orcein,
dark blue with resorcin-fuchsin. They are elastic- i e they extensible and
return after stretching to their former length. Elastic fibers and th '
principal constituent, the protein elastin, are resistant to boiling, acids,
and alkalies
Reticular Fibers (Fig. 53). The name of these fibers is derived from
their presence in reticular connective tissue (see p. 36), and from the fact

that they branch and anastomose, as collagenous fibrils are not supposed
to do. They vary in thickness, but are usually very fine; they form no
bundles, but lattice-like networks They are more resistant to boiling,
acids, and alkalies than collagenous fibers but resemble the latter in their
reactions to Mallory's stain. According to Plenk, they are precollagenous
fibers. With Bielschow-sky's silver solution they stain black and are
therefore called argyrophil fibers.
OCCURRENCE. The reticular fibers are incorporated in the finer
structure of many organs. They are joined by the collagenous fibers,
with-which they may be continuous. Reticular fibers are found in
lymphatic tissue and in the interstitial tissue of various glands, e.g., in
endocrine glands. They are also distributed along the liver sinusoids and
the capillaries of the kidney. By the Bielschowsky method, certain
membranes that were formerly considered to be homo geneous have been
demonstrated to consist of a feltwork of argyrophil fibers, e.g., the
basement membrane of the epidermis and the glands, the membranelles
(sheaths) of smooth muscle fibers, the sarcolemma of striated muscle,
and the fat cell membrane.
The gelatinous fluid ground substance which fills the intercellular
spaces in embryonic connective tissue remains as a mucoid, amorphous,
totally displaceable mass in the other forms of connective tissue. It forms
membranes with the invested fibers in loose connective tissue and
constitutes the scanty cement that binds the fibrils into fibers. According
to the arrangement and nature of the intercellular substance or of the
cells, the following kinds of connective tissue are recognized:
1. Embryonic. Homogeneous ground substance predominates.
2. Reticular.
3. Fibrillar. Fibrillar intercellular substance predominates.
4. Elastic.
1. Embryonic Connective Tissue
This is described on p. 34. In mucous connective tissue (Fig. 51),
which represents the next stage of development from mesenchyme, the
fibrillar intercellular substance has not yet become predominant. It is
composed of a gelatinous, mucous ground substance in which are
embedded fine wavy collagenous fibers and stellate or rounded cells
connected by cytoplasmic processes.
OCCURRENCE. Mucous connective tissue is found in the umbilical
cord around the vessels, as Wharton's jelly, also in embryos in place of
the future perilymphatic spaces of the inner ear. The omasal papillae

contain a tissue of similar structure, as does the supporting lamella in the

comb and in the wattles of the chicken.
A related tissue in the adult is the pigmented connective tissue of
the eye (Fig. 50). It consists of branched connective tissue cells with
lobulated, often coarse, processes and yellow-brown to black pigment
granules. The cells anastomose. The nucleus is free of pigment and is
seen as a light spot under certain conditions. There are also bundles of
collagenous fibrils.
OCCURRENCE. The pigmented connective tissue is present in the
chorioid, ciliary body, iris, and sclera, and scattered in the skin.
2. Reticular Tissue
Reticular tissue presents a wide- or narrow-meshed network of
reticular cells united in a syncytium (Fig. 52). The arrangement
resembles that of mesenchymal cells. The reticular fibers (Fig. 53),
produced by the cells and occurring mostly in the cytoplasmic network,
serve to support and lend firmness to the syncytium. The fibers, which
pass through the cell processes, are usually entirely surrounded by
cytoplasm, but often appear to lie on the surface. The presence of elastic
and collagenous fibers in the reticulum has been reported.
One form of reticular tissue never occurs alone but always from its
first appearance contains in its meshes other mesenchymal derivatives,
the lymphocytes. These cells, which are not attached to the tissue, occur
at first sparsely, but later in such masses that they obscure the reticulum.
The latter can be seen only after removal of the lymphocytes by
brushing, shaking or selective staining. This mixed tissue of sessile
reticular cells, which produce the fiber network, and the loosely
embedded lymphocytes is called lymphatic tissue. 3 The reticular cells can
usually be recognized only by their nuclei, which are larger and stain
lighter than the lymphocyte nuclei.
OCCURRENCE. Reticular tissue forms the finer framework of
whole organs, such as lymph nodes, solitary and aggregated lymph
nodules, tonsils, spleen, and bone marrow; it is also found in the thymus,
intestinal mucosa, conjunctiva, and elsewhere. Where the reticular fibers
form a latticework around the end-pieces oi glands, muscle fibers, or
capillaries, the reticular cells lie on the outside of the lat tice.
The close relationship apparent in certain organs between reticular
and certain special endothelial cells, not only in cytoplasmic and fibrillar
connections but also in functional similarities, has led to the designation
reticulo-endothelial system. To this system belong the lining cells of the
sinuses of the lymph nodes; the sinusoids o) the spleen, bone marrow,

and liver; and the reticular cells of the splenic pulp and the other
lymphatic tissues (lymph nodes, tonsils, and lymph nodules). They are
intimately related and form the reticulo endothelial system in the narrow
sense. They are distinguished by their phagocytosis of erythrocytes,
foreign particles, bacteria, and protozoa, and by their storage of colloidal
substances (hemoglobin, lipoids, and dyes). They are concerned with
erythrocyte destruction, with iron, vitamin, and fat metabolism, with the
defense against noxious substances, and with the production of
antibodies. The activity of the cells of the reticulo-endothelial system is
shown in their morphological changes, enlargement, prolifera tion,
detachment, and metamorphosis to mobile macrophages (see p. 39).
Because of its great adaptability under the influence of functional
stimuli, it is less-to be regarded as a fixed anatomical concept than as an
especially labile, sensitive, physiological system. Vital staining by the
injection of acid colloidal dyes (e.g., lithium carmine) in the living body
has greatly furthered our knowledge of the work and structure of the
reticulo-endothelial system.
3. Fibrillar Connective Tissue
In fibrillar connective tissue the intercellular substance with its
collagerious fibrils predominates over the cells. The extremely fine
collagenous fibrils are held together by small amounts of amorphous
cement to form longitudinally striated fibers. The fibroblasts (Fig. 55) lie
on these bundles of fibrils (Fig. 57,a). The collagenous bundles are
always accompanied by elastic fibers (Fig. 57,6), which may predominate
in numbers, forming a special tissue. (For their characteristics see p. 34.)
The following kinds of cells occur in fibrillar connective tissue: The
fixed connective tissue cells, fibroblasts (Figs. 54,c*; 55} with their
elliptical, chromatin-poor nuclei are usually greatly flattened and
conform in shape to the course of the fibers. They may be curved or
sharply bent, sometimes with ragged, veil-like processes that connect the
fibroblasts with each other. They lie immediately upon the bundles of
fibrils and often form complete sheaths around them. The fibroblasts
produce the fibrous intercellular substance.
Histiocytes (Fig. 5 4,d*) are thin, often multiform cells which
resemble fibroblasts and are considered by some authors to be formed
from them as a result of irrita tion. Histiocytes, however, have sharper
outlines and stain better than do fibroblasts; they possess abundant
granular cytoplasmic inclusions, and their nuclei are richer in chromatin.
Above all, they have the power of phagocytosis, and for this reason they
are also called fixed macrophages. They store foreign substances, for
example, intravitally injected dyes (neutral red), in the form of granules.

They are considered to be "resting wandering cells," which may resume

their wanderings upon suitable stimulation. They are found principally in
the subcutaneous tissue and mesentery, where they are especially
numerous near the small blood vessels. They have great significance in
the defense activities of the body.
Lymphocytes continually pass from the blood vessels into the
connective tissue and thence into, or through, the epithelia.
Coarsely granular acidophil leukocytes (granulocytes, p. 91) wander
out of the blood vessels. They are especially numerous in the intestinal
Mast cells (Fig. 54,4*) resemble the coarsely granular, basophil
leukocytes of the blood (oval cell body, rounded nucleus, basophilic
water-soluble granules), but are not identical with them. They are found
only in loose connective tissue, often in rows along the vessels. They
have been associated with the production of heparin.
Plasma cells (Fig. 56) are round or polyhedral cells with an
eccentric nucleus and a larger amount of cytoplasm than the
lymphocytes, which they otherwise resemble. In the fixed and stained
nucleus the chromatin is arranged about the periphery like the spokes of
a wheel. The cytoplasm has a mottled appearance and is basophilic,
except for an unstained area at one side of the nucleus. De generating
plasma cells contain spherical acidophil bodies in the cytoplasm. Plasma
cells and lymphocytes are probably derived from a common stem cell,
and the immature forms of both have been credited with the production
of antibodies.
Fibrillar connective tissue may be subdivided into loose,
membranous, regular fibrous, and irregular dense connective tissue.
a. Loose Connective Tissue
Loose connective tissue (Figs. 54*, 57) has no definite outward
form. It is a displaceable substance which forms the gliding, mobile
layers of the body. The collagenous fibers and trabeculae (Fig. 57,a) run
a wavy course, cross, and interweave. Loose connective tissue is always
traversed in all directions by abundant elastic fibers (b,c). All the cells
described on pp. 38-39 are included in its structure. In addition, fat cells
(p. 44) occur in varying numbers. It invades the organs as interstitial
tissue and serves as a filling and connecting mass between neighboring
organs and body parts, as well as between the larger divisions (lobes) of
the organs. As inter- and intraparenchymatous connective tissue, it forms
the binding substance in muscles, nerves, and glands. The
interparenchymatous connective tissue of the organs invades them from
their connective tissue capsules and separates the divisions of the

parenchyma (bundles and lobules), enclosing each in a more or less firm

sheath. The delicate framework of bundles of fibrils inclosing the tissue ;
elements of the organs is the intraparenchymatous connective tissue.
The interstitial tissue is a continuous framework, the trabeculae of
which are formed of collagenous fibers and carry the vessels and nerves.
The trabeculae cross in all directions and form a network. The
amorphous ground substance combines with the fiber nets to form
membranes enclosing tissue spaces (tissue lacunae) with small amounts
of tissue fluid. When air is introduced (e.g., into the subcutis), these
small clefts are distended to vesicular spaces (emphysema). This occurs
inwounds. In the same way, blood stasis can cause the accumulation of
fluid in the tissue spaces (hydrops, edema, or anasarca). This
characteristic gave rise to the old terms areolar tissue or cellular tissue;
in the latter term cell has its original meaning of any small space
surrounded by walls. (Compare with cellulitis.) The looseness of the
tissue depends upon the size of the tissue spaces.
In addition to the mechanical function (maintenance of form),
which is an at tribute of the fibrous components, loose connective tissue
also has important metabolic functions to perform. The exchange of
metabolites between the capillaries and the various cells of the organs
takes place through the connective tissue covering the capillaries. The
tissue fluid which mediates this exchange is a component of the
intercellular substance of the connective tissue. The connective tissue is
also important to the water economy of the body. The water in the
organism is continually passing back and forth between the blood and the
tissues (intercellular substance). The cells of the connective tissue
(histiocytes and wandering cells), with the blood, are concerned with the
defense against noxious substances in the organs. This is one of the most
important functions of connective tissue, which re acts rapidly to irritants.
Finally, the loose connective tissue plays a role in regener ation. All the
connective tissues repair defects with the products of the fibro-blasts.
Strong mechanical demands on the connective tissue cause a reduction in
the cells other than fibroblasts and a greater development of the
intercellular structures.
b. Membranous Connective Tissue
In membranous connective tissue the bundles of fibrils of different
sizes crossing each other in various directions, are spread flat. They are
accompanied by a close network of elastic fibers. The cellular elements
are the same as ia loose connective tissue. The lamina propria of the
serous membranes (p. 102) is composed of membranous connective tissue
in which the elastic nets form a layer on both sides of the fibrous sheet.

The connective tissue of the greater omentum (Fig. 58) is-a

variation in which the fiber bundles surround large or small holes in a
lacelike structure. Flat fibroblasts lie on the bundles, which are
accompanied by elastic fibers and may be cov ered by very delicate
membranes containing circular reticular fibers. Vessels run in the thicker
trabeculae. Omentum and serous membranes are covered by simple
squamous epithelium (mesothelium).
A further variation is found in the arachnoid, where the fiber
bundles do not run in one plane but in all directions. The surface of the
trabeculae is covered by an incomplete layer of flat cells.
c. Regular Fibrous Connective Tissue
Regular (tense, stretched) fibrous connective tissue is developed
under conditions of mechanical stress of the fibrous intercellular
substance. It is distinguished by its tensile strength and by the regular
arrangement of its fiber bundles. It is usually poor in elastic tissue.
Fibroblasts of shapes characteristic of the various or gans are almost the
only cells present (Figs. 55,d; 59,b; 60,6).
OCCURRENCE. It forms the ligaments, tendons, aponeuroses, and
certain deep fasciae.
The tendons (Figs. 59, 60), like the ligaments, are composed of
parallel, tautly stretched bundles of fibrils, the primary tendon bundles
(tendon fibers). Between them lie the rows of fibroblasts or tendon cells.
These are flat, polygonal, stellate, or fusiform, usually have winglike
processes, and are marked by many pressure grooves and ridges from the
embedded fibers. The nuclei are elongated and often rqsemble those of
smooth muscle cells. The tendon cells lie in anastomosing tissue spaces,
which they almost fill (Fig. 60,4). A number of primary bundles unite to
form secondary and these to form tertiary bundles. The tendons are
covered superficially by loose connective tissue (peritenonium
externum), except where the tendon sheaths occur. From the
peritenonium externum, processes (peritenonium in ternum) invade the
tendon and clothe the bundles as interfascicular tissue. The latter
contains elastic fibers, fat cells, and occasionally cartilaginous
The blood vessels form a well-developed vascular net in the
peritenonium, but they are lacking inside the bundles. The lymph vessels
are distributed similarly. The tendon fibers are richly supplied with
sensory nerve fibers, which end in ten don spindles and terminal
corpuscles. The functional demand on the tendons has a profound
influence on the nature and the course of their fibers.

Aponeuroses are flat, sheetlike tendinous structures in which groups

of parallel fibers sometimes cross at right angles.
Lamellar connective tissue is seen in the fibrillar intercellular
substance of lamel-~ lar bone, where the parallel fibers of one lamella
take a different direction from those of the neighboring lamellae. The
perineurium and the lamina propria of the cornea have a similar
d. Irregular Dense Connective Tissue
Irregular dense connective tissue forms the remaining fibrous
membranes, which are characterized by a feltwork of interwoven fibers
and trabeculae. Examples are fasciae, periosteum, perichondrium,
connective tissue capsules of many organs (tunicae albugineae and
tunicae fibrosae), sheaths of the blood vessels and nerves, synovial
membranes, bursae, sclera, and corium. When the fasciae serve as mus cle
covering, they are rich in elastic fibers; when they act as tendons, they
have an aponeurotic structure.
The tendon sheaths consist of two cylinders, which are continuous
with each other at the ends of the sheath. Between them is a space filled
with synovial fluid. The wall of the outer cylinder is a tough fibrous coat
joined by loose connective tissue to the synovial membrane lining the
space. The synovial membrane bears flat cells of connective tissue origin
and vascular villi. This membrane is reflected over the tendon, fuses with
it, and forms the inner cylinder of the tendon sheath.
In the subtendinous bursae, the outer layer of the wall, which is
attached to the surrounding tissues, is composed of dense connective
tissue. The inner surface is lined by a synovial membrane. In the places
where the flat lining cells are lack ing, there is usually compact
connective tissue mixed with cartilage cells. The sub cutaneous bursae
have walls of thickened fibrillar connective tissue, from which trabeculae
extend across the cavity. (For the structure of joint cap sules, see p. 56.)
4. Elastic Tissue
Elastic fibers, unlike collagenous fibrils, do not form bundles; they
often branch (Fig. 61,V); and in addition to woven textures they form
characteristic fine or coarse nets by anastomosis (Fig. 62). At the
intersections where the fibers fuse, there are often weblike expansions.
OCCURRENCE. As thick elastic fibers, it is the main constituent of
certain ligaments (ligamentum nuchae, ligamenta arcualia, or flava, liga
menta vocaha, and ligamentum suspensorium penis). In these structures

the fibers are stretched parallel and are held together by loose connective
tissue (Fig 64) The elastic ligaments are yellow. In general, a pronounced
yellowish color in a supporting tissue indicates a high elastin content.
Elastic tissue also appears as plastic networks and as fenestrated or
continuous elastic membranes (lamellae), &g; m the large arteries.
Elastic tissue plays a large part in the structure of the lungs and other
organs. It also occurs in the form of nodules, as in the epiglottis of the
Adipose tissue is composed mostly of fat cells (Fig. 65). These are
spherical, 60-50 M in diameter. The cytoplasm is displaced by a large fat
globule, so that it forms only a thin peripheral layer. In the region of the
elliptical, parietal, often flattened nucleus there is a somewhat thicker
layer of cytoplasm (signet ring form, Fig. 65, a). In addition, each fat
cell possesses a fine membrane of reticular fibers.
In fresh preparations the nucleus is usually invisible. In stained
sections many nuclei show a space that represents a fat-filled cavity (Fig.
65,*). Fat stains black 'c acid, red with Sudan III and scarlet red. The fat
is extracted by the fat solvents embedding, so that only the empty cells
are seen in paraffin and celloidin sections. In the dead animal, tufted
margarin crystals are often found in e fat cells (Fig. 65,b).
Fat cells may occur singly or in groups anywhere in the connective
tissue. When they are aggregated in large masses like bunches of grapes,
they are the main constituents of adipose tissue, which is rich in nerves
and vessels (Figs. 65, 66). This tissue is regularly encountered in certain
regions of the body (the fat organs). The cell groups are of various sizes;
the microscopic ones are called primary lobules and, when avascular, fat
islands. In large deposits of fat, a number of primarv lobules are united
by loose connective tissue to form secondary lobules and the latter may
form larger tertiary lobules, and so on.
In severe starvation the fat in the cells is reduced to small droplets
There re mains in the flattened cells a pale cytoplasmic mass mixed with
mucoid fluid (serous fat cells).
OCCURRENCE. Adipose tissue serves as a supporting substance
that is elastic under pressure, and as a protective, insulating, contourbuilding, filling, and structural material. In well-nourished animals, it is
especially abundant in the subcutis (panniculus adiposus) and in the
mesentery, also on and around very mobile or sensitive organs and those
which must be protected from chilling. Fat plays an im portant structural
role in and around joints and in the foot pads and digital cushions of

animals. Fat is formed in the involution of the thymus. It is absent in the

cranium, eyelids, scrotum, and skin of the penis and clitoris. It occurs in
very small amounts in the ear, nose, and lips. In lean animals fat is found
only in the orbits, around the kidneys, in the mesen tery, bone marrow,
and vertebral canal, along the nerves and arteries, and in the lyssa of the
tissue arises from the middle germ layer through the mesen-chyme (p.
34). A part of the mesenchymal cells are to be regarded as producers of
the collagenous fibrils and are therefore called fibroblasts. The fibrils
probably form by association and parallel orientation of long molecules
of protein (p. 8) secreted by the cells. There is a question as to whether
this takes place within the cytoplasm or in the intercellular substance. In
areas of great concentration of actively fibril-producing cells, extensive
organs of fibrillar connective tissue are developed. In these formed
connective tissues, e.g., tendons, the division into primary, secondary,
and tertiary fiber bundles is always apparent. When there are only a few
scattered fibroblasts of moderate activity, the more delicate membranes,
or large or small irregular formations of loose connective tissue are
It is probable that the elastic fibers are also produced by fibroblasts.
The development of elastic fibers may prevail over the production of
collagenous fibrils, so that gradually the collagenous fibers and the cells
come to form the minor, almost unnoticeable portion of the whole
structure, as in the ligamentum nuchae. The clastic fibers appear
homogeneous from the beginning.
Adipose tissue develops very late in ontogenesis, for the fetus must
reach a high stage of development before it can store reserve material
for nutritional purposes, previous to that, all available nutrients are used
in the formation of the organs. In restricted areas, where the fat bodies
are to form, the mesenchyme in the vicin ity of the blood vessels takes on
a reticular character. The individual cells store fat, Which first appears as
single droplets. These coalesce finally to one large globule (a (a
univacuolar fat cell, Fig. 67). The cytoplasm forms a thin layer around
the fat globule, which presses the nucleus to the periphery. Fat storage in
the cells of reticular tissue is particularly evident in the transformation of
red to yellow bone marrow. Fat cells may also come from fibroblasts.
In poultry, there is much yellow, plurivacuolar adipose tissue, in
which many fat droplets lie around the central nucleus, as in embryonic
fat cells. The "hibernating gland" on the necks of rodents is also of this
type of adipose tissue, but is brown in color.

Pigmented connective tissue cells are also derived from the

undifferentiated mesenchymal cells. It is assumed that there is a division
of labor among the mesenchymal cells which separate from the parietal
layer of the mesoderm, some becoming fibroblasts with elaboration of
intercellular substance, some becoming fat cells, and others becoming
cells which produce intracellular pigment granules.
Cartilage is composed of an abundant intercellular substance and
the encapsulated cartilage cells. The intercellular substance, which is
firm but flexible, is composed of a homogeneous mass and fibrillar
elements. It may have a hyaline appearance, in which case the fibrils are
masked, or it may contain visible elastic Bar collagenous fibers, bundles,
or networks. Therefore we distinguish hyaline car tilage, elastic
cartilage, and fibrocartilage.
1. Hyaline Cartilage
The cartilage cells fill completely the cavities they occupy in the
intercellular substance (Figs. 6870). They may be spherical, oval, halfspherical, or flattened into a lens shape. They are usually surrounded by
a closely applied capsule (Figs.68:69b), which, however, is sometimes
invisible. The capsule is a differentiation of the intercellular substance
produced by the cells. The cells have a soft, delicate cytoplasm, which
contains granular or threadlike mitochondria and inclusions of glycogen,
fat, and, rarely, pigment. The nucleus is spherical, contains nucleoli and
may occasionally be double. The Golgi apparatus is well developed. The
cells shrink easily on fixation, become wrinkled, and no longer fill their
The daughter cells derived from one mother cell are often
associated in isogenous groubs (Fig. 69,e). After new ground*substance
appears between the daughter cells, their common origin may still be
recognized by their proximity, by the fact that their apposed surfaces are
flattened, or by a common capsule around the individual capsules. Near
the free surface of the cartilage, the cells are often flattened and lie
parallel to the surface in layers. Farther within the cartilage the cells are
more rounded, and in the deepest portions they are often arranged in
rows perpendicular to the surface (Fig. 70).
The intercellular substance (Figs. 6B,c; 69,f) is very firm and
elastic, has a bluish-white translucent appearance, and seems to be
homogeneous throughout like ground glass. However, after maceration
with 10 percent NaCl, KMnO 4 , baryta water, lime water, or trypsin, a
fibrillar structure becomes visible. The intercellular substance is

composed of a feltwork of fine collagenous fibrils permeated by a

mucoid cement. Because the fibrils and the interfibrillar ground
substance have the same refractive index, the fibrils are invisible,
masked. Hyaline cartilage contains a great deal of water (costal cartilage
of calf, 75 percent), and shrinks on dry ing to a horny translucent mass.
With prolonged boiling, cartilage yields chondrin (cartilage glue), which
contains among other things chondroitin-sulfuric acid, the cause of the
basophilia of the ground substance (which stains dark blue with
hematoxylin). The intercellular substance immediately surrounding the
capsules has a different staining reaction, forming a halo or cell zone
which often includes a whole group of cells (Fig. 69,*).
In many cartilages, e.g., costal cartilages, there are areas in the
hyaline intercellular substance where one can see fibers without the. use
of reagents. They often radiate from one point. Such areas present an
asbestos-like appearance to the naked eye. In old age, cartilage often
loses its elasticity through calcification.
With the exception of the articular cartilages, the hyaline cartilages
are surrounded by a dense connective tissue membrane which blends with
the. cartilage. This perichondrium (Fig. 7Q,a) plays an important role in
the growth and regeneration of cartilage. Because the cartilage itself is
avascular, it is nourished by diffusion from the perichondrium. The
development of blood vessels in the cartilage is al ways a sign of
beginning calcification or ossification. This process begins early in the
costal cartilagesat six months in the dog, forming a core of spongy
bone by two years of age.
OCCURRENCE. Hyaline cartilage is found in articular cartilages;
in the thyroid, cricoid, and the base of the arytenoid laryngeal cartilages;
in the cartilage of trachea afid bronchi; in the cartilages of the nose; and
in the primordia of most of the bones during development.
2. Elastic Cartilage
Elastic cartilage (Fig. 71) is characterized by a network of fine or
coarse elastic fibers in the. otherwise hyaline intercellular substance. The
fiber networks vary in density in different cartilages. The fibers
gradually become fewer and thinner as they extend into the
perichondrium. The cartilage cells are spherical or flattened, often occur
in groups, and are surrounded by a distinct capsule. Elastic cartilage is
more flexible than hyaline. It contains very few collagenous fibers and is
more or less yellow in color.
OCCURRENCE. This type occurs in the epiglottis, the corniculate
and cuneiform cartilages of the larynx, the auricular cartilages, the

cartilage of the third eyelid, part of the cartilage of the Eustachian tube,
and part of the arytenoid cartilage.
3. Fibrocartilage
In fibrocartilage the intercellular substance contains bundles of
collagenous Sbers, which sometimes run parallel, sometimes form an
irregular texture. The cells lie in thick capsules between the fibers,
singly or in groups or rows. Fibrocartilage often merges gradually into
connective tissue or hyaline cartilage.
OCCURRENCE. This is found in articular discs and menisci,
intervertebral discs, marginal parts of the articular cartilages and
synchon-droses, accessory cartilage of the patella, and cartilaginous parts
of the tendons and tendon sheaths.
DEVELOPMENT. Cartilage can develop directly from indifferent
mesenchymal cells (chondroblasts) or by the transformation of another
tissue, e.g., perichondrium. Embryonic cartilage arises from a syncytium
of cells lying close together without cell boun daries. The boundaries
become distinct with the secretion of a capsule around each cell. As
growth continues, the outer layer of the capsule changes to hyaline
intercellular substance, which at first forms only a thin-walled
framework around the cells (cellular or precartilage) but increases in
amount as it pushes the cells farther apart. Car tilage growth may be
interstitial by cell division followed by the formation of new
intercellular substance or it may be appositional by the transformation of
the deepest layer of perichondrium into cartilage.
Fibrocartilage shows collagenous bundles as early as the cellular
cartilage stage. The fibers of elastic cartilage develop on the surface of
the cell rows and are at first very fine. They exhibit an independent
growth, becoming thicker and splitting off new fibers.
Other Forms of Supporting Tissue. In addition to cartilage and
bone, there are other, relatively soft, tissues that have a supporting
function. One example is adipose tissue, composed of elastic cellular
sacs filled with fat. Similar to fat cells are the cells of the vesicular
supporting tissue in the notochord (chordoid type). They are large saclike
cells with a strong membrane, parietal nucleus, and a fluid, nonfatty
content, which presses the cytoplasm to the wall. The cell fluid is under
strong pressure (turgor) and, because of the close contact of the cells,
lends firmness and resistance to the tissue. The cells are held together by
an elastic and collagenous notochordal sheath to form a flexible
supporting rod.

Chondroid tissue is composed of glassy, translucent, elastic cell

bodies, which maintain a constant form. The nucleus is spherical.
Between the cells are collagenous fibers and a thin interstitial substance
arranged like a honeycomb. In contrast to cartilage cells, chondroid cells
do not shrink. The tissue has a cartilaginous consistency. It occurs mostly
in lower animals, but is also found in birds and mammals, e.g., in tendon
attachments, sesamoid bones of the cat, and tuber calcanei of t calf. It is
closely related to fibrocartilage.