Tissues Introduction and Epithelium 1

MD1Histology: Fall 2012

Tissues

An organized aggregation of cells that function in a collective manner is called a tissue. Body tissues can be classified into four basic types according to their function and structure:
1. Epithelial tissue covers body surfaces and lines hollow organs, body cavities, and ducts. It also forms glands. It is composed of closely aggregated cells with very little extracellular substance.
2. Connective tissue protects and supports the body and its organs. Various types of connective tissue bind organs together, store energy reserves as fat, and help provide immunity to disease-causing organisms. It is characterized by abundance of extracellular material produced by its cells.

Tissues
3. Muscle tissue generates the physical force needed to make body structures move. It is composed of elongated cells that have the specialized function of contraction. 4. Nervous tissue detects changes in a variety of conditions inside and outside the body and responds by generating nerve impulses that help maintain homeostasis. It is composed of cells with elongated processes that receive, generate, and transmit nerve impulses.

Tissues

All tissues of the body develop from three primary germ layer

ectoderm mesoderm endoderm (the first tissues that form in a human embryo)

All three primary germ layers contribute to epithelial tissues. Mesoderm gives rise to all connective tissues and most muscle tissues. Ectoderm develops into nervous tissue.

Epithelial tissue

Main characteristics:

Epithelia line and cover all body surfaces except the articular cartilage, the enamel of the tooth. Most epithelial cells renew continuously by mitosis Epithelia lack a direct blood (avascular) and lymph supply. Nutrients are delivered by diffusion. Epithelia have no or very little extracellular substance. The cohesive nature of the epithelia is maintained by cell adhesion molecules and junctional complexes. Epithelia are anchored to a basal lamina. The basal lamina and connective tissue components cooperate to form basement membrane. Epithelia have structural and functional polarity.

Form and Shape

The form and dimensions of epithelial cells range from columnar to cuboidal to low squamous cells. The epithelial cell nuclei have distinctive shape, varying from spherical to elongated to elliptical corresponding to the shape of the cell. Nuclear form is used as a criteria to determine the arrangement of the cells in layers, a primary criterion for classification of epithelial tissues

Epithelial tissue

Functions:

protection, covering and lining of surfaces (e.g., skin, intestines) absorption (e.g., intestines) secretion (e.g., glands) gas exchange (e.g., lungs) sensation (e.g., gustative and olfactory neuroepithelium) contractility (e.g., myoepithelial cells).

Because epithelial cells line all external and internal surfaces of the body, everything that enters or leaves the body must cross an epithelial sheet.

Epithelial polarity

The various surfaces of epithelial cells often differ in structure and have specialized functions.

The apical (free) surface. Apical surfaces may contain cilia or microvilli. The lateral surfaces of an epithelial cell face the adjacent cells on either side. The basal surface of an epithelial cell is opposite the apical surface and adheres to extracellular materials. The basement membrane is a thin extracellular layer that commonly consists of two layers, the basal lamina and reticular lamina. The basal lamina is closer to the epithelial cells and is secreted by them

1. The Basal domain and cell to extracellular matrix adhesion: Basal lamina & Basement membrane
Epithelial cells are separated from the connective tissue by a sheet of extracellular material called the basal lamina. In EM, it appears as a dense layer, 20100 nm thick, consisting of a delicate network of very fine fibrils (lamina densa). Basal laminae may have an electron-lucent layer on one or both sides of the lamina densa, called lamina rara or lamina lucida. Between cell layers without intervening connective tissue, such as in lung alveoli and in the renal glomerulus, the basal lamina is thicker as a result of fusion of the basal laminae of each epithelial cell layer. The main components of basal laminae are type IV collagen, the glycoproteins laminin and entactin, and proteoglycans.

Basal lamina
Basal laminae are attached to the underlying connective tissues by anchoring fibrils formed by type VII collagen These components are secreted by epithelial, muscle, adipose, and Schwann cells. In some instances, reticular fibers are closely associated with the basal lamina, forming the reticular lamina. Connective tissue cells produce the reticular fibers.

Basal lamina
Basal laminae have many functions. It

provide support to the cells provide a barrier that limits or regulates the exchange of macromolecules between connective tissue and cells of other tissues.

is able to influence cell polarity, regulate cell proliferation and differentiation by binding with growth factors, influence cell metabolism, and serve as pathways for cell migration.
contain the information necessary for certain cellto-cell interactions, such as the reinnervation of denervated muscle cells.

Basal Infoldings

Many cells that transport fluid have infoldings at the basal surface. They significantly increase the surface area of the basal cell domain allowing for more transport proteins and channels to be present. These basal surface modifications are prominent in cells that participate in active transport of molecules e.g. in proximal and distal tubules of the kidney and in certain ducts of the salivary glands. Mitochondria are typically concentrated at this basal site to provide energy. The mitochondria are usually oriented vertically within the folds

2. The lateral domain and its specializations:

Intercellular Junctions

Several membrane-associated structures contribute to cohesion and communication between cells. The lateral membrane of epithelial cells has specialized intercellular junction that serve as site of adhesions and as a seal to prevent the passes of materials through the intercellular membrane. These junctions are distributed from the apex to the base of the cells.

Intercellular junctions

Zonula occludens

Tight junctions or Zonula occludens junctions: form the primary intercellular difusion barrier between adjacent cells.

By limiting the movement of water and other molecules through the intercellular space, they maintain physiochemical separation of the tissue compartments.
In EM, plasma membranes of adjoining cells come in close contact to seal off the intercellular space. The local fusion of transmembrane proteins of adjoining cell form the junction. Three major groups of proteins found in zonula occludens are occludin, claudins and junctional adhesion molecules (JAM). The cytoplasmic portion of these transmembrane proteins contain zonula occludin proteins ZO-1, ZO-2 and ZO-3. A number of bacteria, viruses and parasite products attack zonula occludens junction or associated molecules to gain entry to the body.

Zonula occludens

Tight Junctions or Zonula occludens

Zonula adherens

It encircles the cell and provides for the adhesion of one cell to its neighbor. Numerous actin filaments are inserted into electron-dense plaques of material on the cytoplasmic surfaces of the junctional membranes. There is gap of 15 -20 nm between the adjoining membranes. It is composed of the transmembrane cell adhesion molecule E-cadherin. On the cytoplasmic side the tail of Ecadherin is bound to catenin complex.

Zonula adherens

Macula adherens or desmosomes

The desmosome is a complex disk-shaped structure at the surface of one cell that is matched with an identical structure at the surface of the adjacent cell. The cell membranes in this region are very straight and are frequently somewhat farther apart (>30 nm) than the usual 20 nm. On the cytosolic side of the membrane of each cell and separated from it by a short distance is a circular plaque of material called an attachment plaque, made up of at least 12 different proteins.

In epithelial cells, groups of intermediate cytokeratin filaments are inserted into the attachment plaque or make hairpin turns and return to the cytoplasm.
Because intermediate filaments of the cytoskeleton are very strong, desmosomes provide a firm adhesion among the cells. In nonepithelial cells, the intermediate filaments attached to desmosomes are made not of cytokeratin but of other proteins, such as desmin or vimentin. Proteins of the cadherin family participate in the adhesion provided by desmosomes.

Desmosome

Hemidesmosomes

In the contact zone between certain epithelial cells and the basal lamina, hemidesmosomes are observed. These structures take the form of half a desmosome and bind the epithelial cell to the subjacent basal lamina. However, in desmosomes the attachment plaques contain mainly cadherins, whereas in hemidesmosomes the plaques are made of integrins, a family of transmembrane proteins that is a receptor site for the extracellular macromolecules laminin and type IV collagen.

Gap junction or nexus

Gap or communicating junctions (also called nexus) can occur almost anywhere along the lateral membranes of epithelial cells.

Gap junctions are found in nearly all mammalian tissues, except in skeletal muscle.
They are seen, in conventional transmission electron micrographs, as a close (2-nm) apposition of adjacent cell membranes. The individual unit of the gap junction is called a connexon. Each connexon is formed by six gap junction proteins called connexins. which join together leaving a hydrophilic pore about 1.5 nm in diameter in the center. Connexons of adjacent cells are aligned to form a hydrophilic channel between the two cells. Signaling molecules such as some hormones, cyclic AMP and GMP, and ions can move through gap junctions, causing the cells in many tissues to act in a coordinated manner.

Gap Junction

3. Apical domain: Specializations of the apical cell surface

In many epithelial cells

the apical surface exhibits special structural surface modifications to carry out specific functions. In addition, the apical surface may contain specific enzymes, ion channels and carrier proteins.

The structural surface modification include:

Microvilli: cytoplasmic processes that extend from the surface. Stereocilia: microvilli of large size Cilia: motile cytoplasmic processes

Microvilli
Microvilli are fingerlike extensions measuring about 1 m high and 0.08 m wide. They are found mainly on the free cell surface. Hundreds of microvilli are found in absorptive cells, such as the lining epithelium of the small intestine and the cells of the proximal renal tubule The complex of microvilli and glycocalyx may be seen with the light microscope and is called the brush or striated border.

Microvilli
Microvilli contain a core of actin filaments, which are anchored to villin at the tip of microvillous and extend down to apical cytoplasm where they interact with a horizontal network of actin filaments, the terminal web. The actin filament are cross linked at 10 nm interval by actin bundling proteins, fascin and fimbrin. This provides support and rigidity to the microvilli. Myosin I is also associated with the actin filaments of the core. Terminal web formed by horizontal actin filaments at the base provide support to the actin filaments of the core.

Stereocilia

Stereocilia are long, nonmotile extensions of cells of the epididymis and ductus deferens that are actually long and branched microvilli and should not be confused with true cilia. They are also found in sensory (hair) cells in inner ear.

Stereocilia increase the cell surface area, facilitating the movement of molecules into and out of the cell.

Stereocilia

Stereocilia also have core of actin filaments that are cross linked by fimbrin. In stereocilia, ezrin, a plasma membrane associated molecule anchors actin to plasma membrane. -actinin cross bridges between actin filaments are also present. There is no villin present in the stereocilia.

Cilia

Cilia are motile structure capable of moving fluid and particles along epithelial surfaces.

Cilia are cylindrical, 510 m long and 0.2 m in diameter.

They are surrounded by the cell membrane and contain a central pair of isolated microtubules surrounded by nine pairs of microtubules.

The two microtubules of the peripheral pairs are joined to each other.
Cilia are inserted into basal bodies, which are small cylindrical structures at the apical pole just below the cell membrane.

Basal bodies have a structure similar to that of the centrioles

Cilia