Nervous Tissue

I. Location & Description

The structures that make up the nervous system include:

• brain, cranial nerves and their branches
• the spinal cord, spinal nerves and their branches
• ganglia, enteric plexuses, and sensory receptors

Nervous tissue consists of two types of cells:

• Neurons are cell processes (axons & dendrites) that extend from the nucleus-containing cell body. They
provide most of the unique functions of the nervous system, such as sensing, thinking, remembering,
controlling muscle activity, and regulating glandular secretions.
• On the other hand, the non-irritable supporting cells, neuroglia, nourish, and protect the neurons and
maintain homeostasis in the interstitial fluid that bathes them.

II. Neuron

A. PARTS OF A NEURON

There are three parts to a neuron: a cell body and two types of cellular projections. The cell body is called the
neuron cell body, or soma (body); as with any other type of cell, the cell body’s nucleus is the source of information
for protein synthesis. One type of cellular projection is called a dendrite (tree), referring to its branching organization.
The other type of cellular projection is called the axon (axis), referring to the straight alignment and uniform diameter
of most axons. Axons are also called nerve fibers.

• The cell body, also known as the perikaryon or soma, contains a nucleus surrounded by cytoplasm that
includes typical cellular organelles such as lysosomes, mitochondria, and a Golgi complex. Most neurons
 have a spherical, unusually large, euchromatic (pale-staining) nucleus with a prominent nucleolus. Bi-
nuclear neurons are seen in sympathetic and sensory ganglia. The chromatin is finely dispersed, reflecting
the intense synthetic activity of these cells. Neuronal cell bodies also contain free ribosomes and prominent
clusters of rough endoplasmic reticulum, termed Nissl bodies. The ribosomes are the sites of protein
synthesis. Newly synthesized proteins produced by Nissl bodies are used to replace cellular components as
material for growth of neurons and to regenerate damaged axons in the PNS. When appropriate stains are
used, RER and free ribosomes appear under the light microscope as basophilic granular areas called Nissl bodies -
their number varies according to neuronal type and functional state, in large neurons such as motor neurons they are
particularly abundant. The Golgi complex is located only in the cell body and consists of multiple parallel arrays of
smooth endoplasmic reticulum (SER) arranged around the periphery of the nucleus. Mitochondria are scattered
throughout the cytoplasm of the perikaryon.

• Dendrites are short, often highly branched cytoplasmic extensions that are tapered from their bases at the
neuron cell body to their tips. They form many synapses and are the principle signal reception and processing sites
on neurons. Most neurons have numerous dendrites, which considerably increase the receptive area of the cell. The
arborization of the dendrites allows one neuron to receive and integrate a great number of axon terminals from other
neurons. It has been established that up to 200,000 axonal terminations establish functional contact with the dendrites
of a Purkinje cell of the cerebellum. That number maybe even higher in other neurons.

Unlike axons, which maintain a constant diameter from one end to another, dendrites become thinner as they
subdivide into branches. The cytoplasmic composition of the dendrite base, close to the neuron body, is
similar to that of the perikaryon but is devoid of Golgi complexes.

Many dendrite surfaces have small extensions, called dendritic spines, where axons of other neurons form
synapses with the dendrites. Dendrites receive input from other neurons’ axons and from the environment.
When stimulated, they generate small electric currents, which are conducted to the neuron cell body.
Dendrite spines are the first processing locale for synaptic signals arriving on a neuron. The processing
apparatus is contained in a complex of proteins attached to the cytosolic surface of the postsynaptic
membrane, which is visible under the electron microscope and received the name postsynaptic membrane
long before its function was disclosed. Dendritic spines participate in the plastic changes that underline
adaptation, learning, and memory. They are dynamic structures with a morphological plasticity based on the
cytoskeletal protein actin, which is related to the development of synapses and their functional adaptation in
adults.

• An axon is a cylindrical process that varies in length and diameter according to the type of the neuron. Most
neurons have only one axon; a very few have no axon at all. Axons are usually very long processes,
although some neurons have short axons. For example, axons of the spinal cord motor neurons that
innervate the foot muscles may be up to 100 cm (about 40 inches) in length.

All axons originate from a short pyramid-shaped region, the axon hillock that usually arises from the
perikaryon. The plasma membrane of the axon is called axolemma (axon + Gr. eilema, sheath); its contents
are known as axoplasm. In neurons that give rise to a myelinated axon, the portion of the axon between the
axon hillock and the point at which the myelination begins is called the initial segment. This is the site at
which various excitatory and inhibitory stimuli impinging on the neuron are algebraically summed, resulting
in the decision to propagate - or not to propagate - an action potential, or nerve impulse. It is known that
several types of ion channels are located in the initial segment and that these channels are important in
generating the change of electric potential that constitutes the action potential. In contrast to dendrites,
 axons have a constant diameter and do not branch profusely. Occasionally, the axon, shortly after its
departure from the perikaryon, gives rise to a branch that returns to the area of the perikaryon. Along the
length of an axon, side branches called axon collaterals may branch off, typically at a right angle to the
axon. The axon and its collaterals end by dividing into many fine processes called axon terminals
(telodendria).

Axoplasm possesses mitochondria, microtubules, neurofilaments, and some cisternae of SER. The absence
of polyribosomes and RER emphasizes the dependence of the axon on the perikaryon for its maintenance.
If the axon is severed, its peripheral parts degenerate and die.

Along the axon a bi-directional transports of molecules and organelles occurs. Anterograde
flow continuously delivers macromolecules and organelles, synthesized in the perikaryon, to the axon
terminals. It occurs at three distinct speeds - a slow stream (a few millimiters per day) transport proteins and
actin fillaments; a flow of intermediate speed transports mitochondria, and a fast stream (100 more rapid)
transports substances contained in vesicles that are needed at the axon terminals during
neurotransmission. Simultaneously, a retrograde flow in the opposite direction transports several
molecules, including material taken up by endocytosis (including viruses and toxins), to the perikaryon. This
process is used to study the pathways of neurons; peroxidase or another marker is injected in region with
axon terminals, and its destribution is followed after a certain period of time.

B. OTHER STRUCTURES (as seen in the pic)

Cytoplasmic extensions of the Schwann cells in the PNS and of the oligodendrocytes in the CNS surround axons to
form either myelinated or unmyelinated axons. Myelin sheath protects and electrically insulates axons from one
another. In addition, action potentials travel along myelinated axons more rapidly than along unmyelinated axons. In
myelinated axons, the extensions from Schwann cells or oligodendrocytes repeatedly wrap around a segment of an
axon to form a series of tightly wrapped membranes rich in phospholipids, with little cytoplasm sandwiched between
the membrane layers. The tightly wrapped membranes constitute the myelin sheath and give myelinated axons a
white appearance because of the high lipid concentration. The myelin sheath is not continuous but is interrupted
every 0.3–1.5 mm. At these locations are slight constrictions where the myelin sheaths of adjacent cells dip toward
the axon but do not cover it, leaving a bare area 2–3 µm in length. These interruptions in the myelin sheath are the
nodes of Ranvier.

C. BASIC TYPES OF NEURONS

Based on the size and shape of their processes, most neurons can be placed in one of the following categories: multipolar
neurons have more than two cell processes, one being the axon and the others dendrites; bipolar neurons have an axon and
one dendrite; and pseudounipolar neurons have a single process that emerges from the perikaryon and shortly afterwards
forms a T-shape, with one branch extending to the peripheral endings of the dendrites and the other towards the CNS. In
pseudounipolar neurons, stimuli that are picked up by the dendrites travel directly to the axon terminals without passing through
the perikaryon. During the maturation process of pseudounipolar neurons, the central (axon) and the peripheral (dendrite) fibers
fuse, becoming one single fiber. In these neurons, the cell body does not seem to be involved in the conduction of the impulses,
although it does synthesize many molecules, including neurotransmitters that migrate to the peripheral fibers.
Most neurons of the body are multipolar. Bipolar neurons are found in the cochlear and vestibular ganglia, retina, and the
olfactory mucosa. Pseudounipolar neurons are found in the spinal ganglia. They are also found in most of the cranial ganglia.

III. Neuroglia
Neuroglia (glue) or glia make up about half the volume of the CNS. Their name derives from the idea of early
histologists that they were the “glue” that held nervous tissue together. Generally, neuroglia are smaller than neurons,
and they are 5 to 50 times more numerous.They are derived from neuroectoderm (macroglia: astrocytes,
oligodendrocytes, ependyma) or from bone marrow (microglia). Nerve tissue has a very small amount of extracellular
matrix, and glial cells furnish a microenvironment suitable for neuronal activity. Glial cells have important structural
and metabolic interactions with neurons and their dendritic and axonal processes. They also have a primary role in a
wide range of normal functions and reactions to injury, including inflammation, repair, fluid balance, and energy
metabolism. In contrast to neurons, glia do not generate or propagate action potentials, and they can multiply and
divide in the mature nervous system.
The size and the shape of the nucleus helps in the light microscopic distinction of one glial cell type from another, as
their cytoplasmic processes are often not apparent on H&E preparations and can be demonstrated only with the use
of metalic impregnation, immunohistochemical, or electron microscopic methods.
Of the six types of neuroglia, four—astrocytes, oligodendrocytes, microglia, and ependymal cells—are found only in
the CNS. The remaining two types—Schwann cells and satellite cells—are present in the PNS.
• Astrocytes are star-shaped cells with multiple radiating processes. Astrocytes are the most numerous glial
cells and exhibit an exceptional morphological and functional diversity. Astrocytes have bundles of
intermediate filaments made of glial fibrillary acidic protein (GFAP) that reinforces their structure. The
filaments are either aggregated in fascicles (protoplasmic astrocytes) or dispersed diffusely throughout the
cytoplasm (fibrous astrocytes). Astrocyte foot processes cover the surfaces of neurons, blood vessels,
and the pia mater membrane of the brain and spinal cord. The astrocytes provide structural support and
play a role in regulating what substances from the blood reach the neurons. In addition to their supporting
function, astrocytes participate in controlling the ionic and chemical environment of neurons.
• Oligodendrocytes (Gr. oligos, small, + dendro, branch, + kytos, cell) form the myelin sheaths around axons in the
CNS. They have a denser, more homogeneous chromatin in a rounder and smaller nucleus (8 µm) than astrocytes do.
Oligodendrocytes also have a small number of cytoplasmic processes each of which forms a broad, flat structure after
emanating from the cell body. An individual oligodendrocyte straddles a bundle of axons, and each of its processes
wraps itself around the axon of a different neuron in the bundle. The processes form concentric circles of fatty tissue
that make up a segment of that neuron's myelin sheath; in between each segment of myelin is an unmyelinated section
of the axon called the Node of Ranvier. Myelin has a composition of 80% lipids and 20% protein, and performs a
number of crucial functions in the nervous system. It insulates nerve fibers by reducing the leakage of ions through the
cell membrane, and increases the velocity at which action potentials are conducted along the nerve fiber, by enabling
them to "jump" from one unmyelinated region to the next, in a process called saltatory conduction. Polio, multiple
 sclerosis, and other demyelinating diseases are caused by the breakdown of the myelin sheath in various regions of
the nervous system, and can result in blindness and paralysis
• Ciliated ependymal cells lining the ventricles of the brain and the central canal of the spinal cord help
move cerebrospinal fluid. Ependymal cells on the surface of the choroid plexus secrete cerebrospinal fluid.
• Microglia (Gr. micros, small, + glia) are small elongated cells with short irregular processes. They can be recognized
in routine hematoxylin and eosin preparations by their dense elongated nuclei, which contrast with the spherical of
other glial cells. Microglia are phagocytic cells that represent the mononuclear phagocytic system in the nerve tissue
and are derived from precursor cells in the bone marrow. They are involved with inflammation and repair in the adult
CNS, and they produce and release neutral proteases and oxidative radicals. When activated, microglia retract their
processes and assume the morphological characteristics of macrophages, becoming phagocytic and acting as
antigen-presenting cells. Microglia secrets a number of immunoregulatory cytokines and dispose of unwanted cellular
debris caused by CNS lesions.
• Schwann cells form the myelin sheath of an axon within the PNS. In this sense, Schwann cells are the
peripheral nervous system's analogues of the central nervous system oligodendrocytes. However, unlike
oligodendrocytes, each myelinating Schwann cell provides insulation to only one axon. This arrangement
permits saltatory conduction of action potentials with repropagation at the Nodes of Ranvier, the gaps
between myelinated segments. In this way, myelination greatly increases speed of conduction and saves
energy. Interestingly, they are called neurilemmocytes because they form neurilemma as they wrap around
the axons of neurons. The neurilemma is a layer of Schwann cell wrapped around the axon; it is a coil of
Schwann cell including two layers of plasma membrane and the cytoplasm between them. Schwann cells
are sometimes called neurilemmocytes for this reason. If the Schwann cell has produced a myelin sheath
around the nerve fiber, then the neurilemma is only the outermost coil of the cell, surrounding the myelin; the
term doesn’t include the layers of myelin.
• Satellite cells are small, flattened cells found in the ganglia of the peripheral nervous system (ganglion =
collection of cell bodies). Researchers have yet to determine the specific functions of satellite cells, but it is
generally assumed that they help regulate and stabilize the environment around ganglion cell bodies.