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Connective tissue is the medium which surrounds and supports the other
tissues of the body. The term stroma is used to describe the
connective tissue component of an organ, and the term parenchyma
refers to the actual functioning component of the organ, e.g., the secretory
epithelium of glands. Most connective tissue (C.T.) arises from the
embryonic C.T. or mesenchyme which is derived from the mesoderm.
Blood is a type of connective tissue, but will be discussed separately.
A. Functions of Connective Tissue
1. Mechanical support and protection of soft tissues
2. Physiological support - a. Serves as a pathway for vessels
and nerves b. Tissue fluids of C.T. act as a diffusion medium
for exchange of metabolites between tissues and vessels
3. Storage of energy in the form of fat
4. Defense against infection
5. Repair of injuries
B. Characteristics of C.T.
All types of C.T. are characterized by:
1. The presence of cells, fibers, and ground substance in varying combinations
and amounts.
2. A relative sparsity of cells and an abundance of the secreted,
non-living, intercellular matrix of fibers and ground substance.
C. Components of C.T.
The nature and function of the various types of C.T. depend upon the
admixture of cells, fibers and ground substance.
The fibers and ground substance are referred to collectively as the
matrix.
1. Cells found in connective tissue
A variety of cells appear in C.T. Some, like fibroblasts or adipose
cells, are indigenous to the C.T. and engage in production of the matrix
or in intracellular storage. Many of the other cell forms originate
in the blood, which is itself a specialized form of C.T., and infiltrate
C.T. proper in response to infection or injury.
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a. Fibroblasts - This most common and characteristic C.
T. cell type secretes the fibers and ground substance of connective
tissue. The fibroblast has a large, flattened, euchromatic, ovoid
nucleus, a fusiform shape, and long, tapering cytoplasmic processes
often indistinct in outline. When not actively engaged in synthesis,
these cells may be called fibrocytes.
b. Adipose cells (fat cells) - Specialized for storage
of lipid. The solubilization of the large central lipid droplet
during paraffin embedding leaves only the nucleus and a thin rim
of cytoplasm, giving the cells a "signet ring" appearance.
c. Mast cells - Commonly occur along blood vessels, but
do not stain distinctly with H&E. The cells are large, ovoid and
densely packed with metachromatic granules containing heparin,
serotonin, and histamine. The granules usually obscure the nucleus
unless, as readily happens, degranulation occurs.
d. Macrophages (histiocytes) - These are phagocytic cells
derived from blood monocytes. They may be fixed in position or
may migrate into the C.T. after tissue injury or infection. Their
nucleus is smaller and darker staining that that of the fibroblast,
and is ovoid or somewhat indented. The cell is irregular in shape,
but has a distinct outline. Ingested particles may be visible
in the cytoplasm.
e. Plasma cells - These cells synthesize antibody molecules
which circulate in the blood stream. Plasma cells are commonly
found in the C. T. of the respiratory, gastrointestinal and genitourinary
systems. These cells are oval and have an abundant, more-or-less
basophilic cytoplasm. The nucleus is eccentrically placed and
is small and dense, with clumps of heterochromatin dispersed in
a cart-wheel pattern around the periphery ("clock face" nucleus).
In H&E preparations, the cells typically have an clear perinuclear
halo marking the position of the pale-staining Golgi complex.
f. Lymphocytes - These blood cells are normally found
in C.T. in small numbers but produce dense tissue infiltrations
during chronic inflammatory states. The cell is usually small
(7-8 mm in diameter) with a relatively large nucleus and scanty
cytoplasm. In H&E preparations they appear as dark purple staining
spheres scattered in the C.T.
g. Neutrophils and Eosinophils - Blood cells which infiltrate
C.T. in large numbers only during tissue inflammation. The neutrophil
is characterized by a multilobed nucleus. The eosinophil contains
reddish-orange granules and has a horseshoe-shaped (bilobed) nucleus.
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2. Fibers found in Connective Tissue
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a. Collagen Fibers - The most common C.T. fiber; these
are long, non-branching and straight (or slightly wavy in the
absence of tension). They do not impregnate with silver (non-argyrophilic)
or stain strongly with PAS. The fibers consist of bundles of fine
collagen fibrils, composed of repeating tropocollagen molecules.
The partially overlapping, 280 nm long, tropocollagen units impart
to the fibrils a 64nm periodicity visible at the EM level.
b. Reticular Fibers - Fine collagen fibrils bound to
glycoprotein components. They form delicate supporting networks
in adipose and lymphoid tissue and along the basement membranes
where C.T. lies in contact with epithelial cells or muscle fibers.
Reticular fibers are not detectable with H&E, but stain strongly
with PAS* and are intensely argyrophilic (have an affinity for
silver staining). The fibrils have the typical 64 nm periodicity
and may develop into larger diameter collagen fibers during maturation.
c. Elastic Fibers - Do not stain with H&E and are best
demonstrated by either orcein or Weigert's resourcin-fuchsin stain.
They are long, thin, branching threads, exhibiting no periodicity
at the EM level. The fibers are composed of tiny peripheral microfibrils
and a central core of elastin, a protein which contains two unusual
amino acids, desmosine and isodesmosine.
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3. Ground Substance
Amorphous material containing proteoglycans which consist of 5% protein
and 95% carbohydrate as polymers of disaccharide units (glycosaminoglycans).
Ground substance serves as a diffusion medium in the spaces around the
cells and fibers. It plays a major role in determining the physical
nature of a C.T. In C.T. Proper, it is relatively free flowing, while
in Supportive C.T., such as bone or cartilage, it is rigid. The ground
substance may stain weakly due to its high water content and because
of partial solubilization of certain glycosaminoglycan components during
fixation and embedding.
*PAS - Periodic acid - Schiff reaction
stains complex carbohydrates a deep red (magenta), e.g. mucin in goblet
cells, basement membrane in the glomerulus, the brush border of kidney
tubules or glycogen deposits are all PAS-positive.
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