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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
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.
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
ox.
B. ADIPOSE TISSUE
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
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.