1 Hist Sept 13 ReadingNerve - Tissue2020

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Learning objectives:

1. Describe the histological characteristics that are unique to neural tissue


2. Describe the cellular composition and general functions of neural tissue
3. Classify neurons based on their morphology and functions
4. Describe the microscopic structure of neurons and correlate these with their functions
5. Describe the different types of synapses and correlate their structure with their functions
6. Contrast fast versus slow axonal transport
7. Describe the microscopic structure of the glial cell types and correlate these with their
distribution and functions
8. Describe the development, structure, and functions of myelin
9. Compare myelinated and non-myelinated axons
10. Contrast the myelination of axons in the central nervous system (CNS) and peripheral
nervous system (PNS)
11. Predict functional outcomes that result from disorders of myelination (e.g., multiple
sclerosis)
12. Describe anterograde (Wallerian) and retrograde neuronal degeneration in response to
injury
13. Contrast the neuronal responses to injury in the CNS and PNS
14. Describe the microscopic structure and functions of the choroid plexus and the blood-
brain barrier

1. General definitions

A. Central Nervous System (CNS) vs. Peripheral Nervous System (PNS)


CNS includes brain and spinal cord; PNS is all outlying nervous tissue (nerve fibers and
ganglia).
Functionally, the PNS is divided into a sensory (afferent) component, which receives and
transmits impulses to the CNS for processing, and a motor (efferent) component, which
originates in the CNS and transmits impulses to effector organs throughout the body. The motor
component is further subdivided as follows:
1) In the somatic system, impulses originating in the CNS are transmitted directly, via a single
neuron, to skeletal muscles.
2) In the autonomic system, in contrast, impulses from the CNS first are transmitted to an
autonomic ganglion via one neuron; a second neuron originating in the autonomic ganglion then
transmits the impulses to smooth muscles, cardiac muscles, or glands.

B. Cell Types
1) Neurons (the parenchyma cells)
2) Glial cells (the stromal cells) – supportive and protective function

2. Development of the nervous system

The CNS develops from the primitive ectoderm. A simple epithelial disk, the neural plate,
rapidly rolls into a hollow cylinder, the neural tube. This process is known as neurulation.
Figure 9-1 Early stages of neural tube formation (Histology and Cell Biology, Kierszenbaum, Abraham L., M.D.,
Ph.D.; Tres, Laura L., M.D., Ph.D.)

Three cell sources of the CNS

The ectoderm germ cell layer gives rise to three major structures: (1) the surface ectoderm,
primarily the epidermis of the skin (including hair, nails, and sebaceous glands), lens and the
cornea of the eye, anterior pituitary, and tooth enamel; (2) the neural tube (brain and spinal
cord); and (3) the neural crest.

Cells of the neural crest migrate away from the neural tube and generate components of the
peripheral nervous system (Schwann cells and the sympathetic and parasympathetic nervous
system), the adrenal medulla, melanocytes of the skin, odontoblasts of the teeth, and neuroglial
cells.

3. Neurons

A. Components of a neuron
1) Soma or cell body
2) Dendrites
3) Axon
Figure 9-2 Components of a neuron (Histology and Cell Biology, Kierszenbaum, Abraham L., M.D., Ph.D.; Tres,
Laura L., M.D., Ph.D.)

B. Classification of Neurons
All neurons have a cell body and a variable number of processes, including at least one axon that
transmits impulses away from the cell body, and usually one or more incoming processes called
dendrites. Variations on this theme permit classification according to number and types of
processes.
Figure 9-3 Types of neurons: Bipolar, pseudounipolar, and multipolar neurons (Histology and Cell Biology,
Kierszenbaum, Abraham L., M.D., Ph.D.; Tres, Laura L., M.D., Ph.D.)
1) Bipolar
• A single axon and dendrite arise at opposite poles of the cell body.
• Found as sensory neurons in the ear/vestibular system, in the retina, and in the olfactory
mucosa.

2) Pseudounipolar
• Only one cell process arises from the cell body & then divides into 2 branches.
Developmentally, begin as bipolar neurons. Functionally, one process carries impulses to
the cell, while the other carries it away. Therefore, although it appears unipolar,
developmentally and functionally it is bipolar, giving rise to its name.
• Found in peripheral sensory ganglia, such as dorsal root ganglia, and in cranial ganglia.

3) Multipolar
• MOST COMMON
• One axon, multiple dendrites
• Found in brain, spinal cord, and peripheral ganglia

C. Cell Body: neurons have a characteristic cytology reflecting a high metabolic activity.
This activity is reflected in their histological appearance.

1) Also called the soma or perikaryon and consists of the nucleus and cytoplasm.

2) Nucleus
• Single large nucleus, usually centrally placed. Moves to periphery if cell is injured. Fine
chromatin.
• Large, prominent nucleolus (“owl-eye” nucleus).
• Reflect a high degree of transcriptional activity.

3) Cytoplasm
• Well-developed rough ER and polyribosomes. Appear as clumps with H&E stains (Nissl
substance). Especially prominent in large active neurons such as the motor neurons of
the spinal cord. Present in the cell body
• There is also a well-developed Golgi to produce secretory products, and large numbers of
mitochondria to supply the high energy requirements. Lysosomes are numerous because
of a high turnover of cell membrane and other cell components, and residual bodies
containing lipofuscin are often prominent, particularly in the elderly.
• May contain pigment such as melanin (neurons of the substantia nigra).
• Abundant neurofilaments (intermediate filaments) help form the cytoskeleton of the
perikaryon.

D. Dendrites:
• Numerous, short processes that become attenuated and branch with distance from the
perikaryon. Provide receptor sites for up to 200,000 axon terminals from other neurons,
and carry impulses to the cell body.
• Ultrastructurally, contain Nissl bodies, mitochondria, neurofilaments, microtubules, and
other features of the perikaryon, but no Golgi complex.

E. Axons:
• Multipolar neurons have a single axon for transmission of impulses away from the
perikaryon. Although collateral branches may arise a short distance from the body, the
axon is otherwise unbranched until it terminates; it is constant in diameter throughout
its length.
• Arises from the axon hillock. This contains no Golgi complex or Nissl substance.
• The axon cytoplasm, or axoplasm, contains many longitudinally oriented microtubules
and neurofilaments, mitochondria, and smooth ER. It never contains Nissl substance or
Golgi.
• Surrounded by a plasma membrane called the axolemma. A protective layer of glial
cells further surrounds this.
• Bidirectional transport:
i. Macromolecules and organelles made in the perikaryon are transported to the axon
terminals by anterograde flow.
ii. Retrograde flow transports molecules (viruses and toxins included) to the cell body.
iii. Major proteins involved are dynein and kinesin.

Figure 9-4 Axonal transport (Histology and Cell Biology, Kierszenbaum, Abraham L., M.D., Ph.D.; Tres, Laura L.,
M.D., Ph.D.)

F. Synaptic terminals and synapses:


The synaptic terminal is specialized for the transmission of a chemical message in response to
an action potential. The synapse is the junction between the presynaptic terminal of an axon
and a postsynaptic membrane receptor surface, generally a dendrite.
Figure 9-5 Synaptic transmission (Histology and Cell Biology, Kierszenbaum, Abraham L., M.D., Ph.D.; Tres,
Laura L., M.D., Ph.D.)

Synapses are classified by their location on the postsynaptic neuron as follows:


1. Axospinous synapses are axon terminals facing a dendritic spine.
2. Axodendritic synapses are axon terminals on the shaft of a dendrite.
3. Axosomatic synapses are axon terminals on the soma of a neuron.
4. Axoaxonic synapses are axon terminals ending on axon terminals.
Figure 9-6 Types of synapses (Histology and Cell Biology, Kierszenbaum, Abraham L., M.D., Ph.D.; Tres, Laura
L., M.D., Ph.D.)

Clinical significance: Axonal transport of rabies virus

We emphasize once more the bidirectional transport of cargos (including synaptic


vesicles and mitochondria) along the axon:
1. Kinesin-mediated anterograde axonal transport of neurotransmitters, from the cell body
toward the axon terminal and the plus end of microtubules.
2. Cytoplasmic dynein-mediated retrograde axonal transport of growth factors and
recycling of axon terminal components from the axon terminal to the cell body and the minus
end of microtubules.

As you recall, kinesin and dynein motor proteins have a globular motor domain in their heavy
subunits that bind to microtubules and hydrolyses adenosine triphosphate (ATP) to propel cargos
along the microtubule tracks. Cargos are attached to kinesin by the light subunit of the motor
protein. Dynactin is a protein complex involved in the attachment of the cargo to dynein.

There are two types of axonal transport:


1. Fast axonal transport, responsible for the movement of vesicles and mitochondria.
2. Slow axonal transport, which is responsible for driving cytoplasmic proteins and
cytoskeletal proteins for the assembly of microtubules and neurofilaments.
Axonal transport is important in the pathogenesis of neurologic infectious diseases. For example,
the rabies virus introduced by the bite of a rabid animal replicates in the muscle tissue from as
little as 2 to 16 weeks or longer.
After binding to the acetylcholine receptor, the viral particles are mobilized by retrograde
axonal transport to the cell body of neurons supplying the affected muscle.
The rabies virus continues to replicate within infected neurons and after the shedding of the
virions by budding, they are internalized by the terminals of adjacent neurons.
Further dissemination of the rabies virus occurs in the CNS. From the CNS, the rabies virus is
transported by anterograde axonal transport by the peripheral nerves to the salivary glands.
The virus enters the saliva to be transmitted by the bite.
Painful spasm of the throat muscles on swallowing accounts for hydrophobia (aversion to
swallowing water).
The retrograde axonal transport to the CNS of tetanus toxin, a protease produced by the
vegetative spore form of Clostridium tetani bacteria after entering at a wound site, blocks the
release of inhibitory mediators at spinal synapses. Spasm contraction of the jaw muscles (known
as trismus), exaggerated reflexes, and respiratory failure are characteristic clinical findings.

4. Glial cells:
Glial cells are more numerous than neurons and retain the capacity to proliferate. Most brain
tumors, benign or malignant, are of glial origin.
When the CNS is injured, glial cells mobilize, clean up the debris, and seal off the local area,
leaving behind a “glial scar” (gliosis), which interferes with neuronal regeneration.
Unlike neurons, glial cells do not propagate action potentials and their processes do not receive
or transmit electrical signals.
The function of glial cells is to provide neurons with structural support and maintain local
conditions for neuronal function.

A. Astrocyte – CNS only


1) Structure
• Large, oblong nucleus with stellate long processes
• Cytoplasm can’t be seen with H&E
• Contains abundant glycogen and unique intermediate filaments called glial fibrillary acid
protein (GFAP)
2) Function – gives structural support to neural tissues by binding neurons to capillaries and to the
pia mater. This is accomplished by expansion of their distal processes to form “end feet”.
Responsible for transfer of metabolic products from blood to neurons. Also proliferate
following injury to the brain to form scar tissue.
3) Fibrous astrocytes (white matter) vs. protoplasmic astrocytes (gray matter)
Figure 9-7 Astrocytes (Histology and Cell Biology, Kierszenbaum, Abraham L., M.D., Ph.D.; Tres, Laura L., M.D.,
Ph.D.)

B. Oligodendrocyte – CNS only


1) Structure – smaller than the astrocyte, with small, round dense nucleus and clear cytoplasm.
Does not contain glycogen or GFAP.
2) Function – responsible for myelination in the CNS. One cell can myelinate several axons.

C. Schwann Cell – PNS only


1) Structure – small, dark, flat nucleus.
2) Function – responsible for myelination in the PNS. Each cell can myelinate only one axon,
however each axon can be myelinated by several Schwann cells! Cytoplasm can also cover
axons without myelinating them.

Oligodendrocytes and Schwann cells: Myelinization


Figure 9-8 Oligodendrocytes and nodes of Ranvier in the CNS and PNS (Histology and Cell Biology,
Kierszenbaum, Abraham L., M.D., Ph.D.; Tres, Laura L., M.D., Ph.D.)
Figure 9-9 Myelination (Histology and Cell Biology, Kierszenbaum, Abraham L., M.D., Ph.D.; Tres, Laura L., M.D.,
Ph.D.)

Demyelinating diseases:

he integrity of myelin, but not the axon, is disturbed in demyelinating diseases affecting
the survival of oligodendrocytes or the integrity of the myelin sheath.

The myelin of the CNS is the target for attack by the immune system in multiple sclerosis, which
is of unknown cause.
Myelin is vital for the CNS to function effectively, and its destruction in multiple sclerosis results
in severe functional deficits, such as paralysis, loss of sensation and/or loss of coordination. The
nature of the deficit depends on the area of the CNS affected.
Figure 9-10 Multiple Sclerosis. (Stevens & Lowe’s Human Histology, Lowe, James S., BMedSci, BMBS, DM,
FRCPath; Anderson, Peter G., DVM, PhD)
Low-power micrograph of cerebral cortex from a patient with multiple sclerosis (MS). Tissue is stained with Luxol
fast blue to demonstrate myelin in the white matter (W) while the grey matter (G) contains far less myelin. Two pale
non-staining areas, called MS plaques (P), demonstrate focal areas of myelin loss.

D. Microglia – CNS only


1) Structure – smaller than other cells in the CNS with dense, elongated nucleus.
2) Function – not usually seen in normal brain tissue. The phagocytic cell of the CNS and involved
in inflammation and repair. These cells are the primary site of HIV-1 infection in the CNS.

E. Ependymal Cells – CNS only


1) Structure – cuboidal epithelial cells lining the ventricles and central canal of the spinal cord.
2) Function – form barrier between brain and CSF. Also found in the choroid plexus (see below).

F. Satellite Cell – PNS only


1) Structure – small, dark nuclei.
2) Function – Cluster around and provide support for neuron cell bodies in peripheral ganglia.
Figure 9-11 Support Cells of the CNS. (Stevens & Lowe’s Human Histology, Lowe, James S., BMedSci, BMBS,
DM, FRCPath; Anderson, Peter G., DVM, PhD)
The support cells of the CNS are called ‘glia’ and have several roles. Astrocytes not only support neurons and their
synapses but also extend foot processes around capillaries to maintain a blood–brain barrier. At the surface of the
brain, astrocytes line a basement membrane and form the glia limitans, which surrounds the whole CNS.
Oligodendrocytes myelinate the axons of the nerve cells, and a vast network of antigen-sensing microglial cells is
present throughout the CNS. Phagocytic macrophages, which also have an immune defence role, reside in the
perivascular space, outside the CNS substance. The ependymal cells form an epithelial-like sheet which, unlike
other epithelia, does not lie on a basement membrane. This sheet lines the fluid-filled ventricular cavities of the
brain and the central canal of the spinal cord.

5. Central Nervous System (CNS)

A. Distribution of white matter and gray matter


1) Gray matter contains neurons, dendrites, the initial portion of unmyelinated axons, and glial
cells. White matter contains myelinated axons, a few unmyelinated fibers, glial cells, but NO
neurons. The non-neuron component is often collectively referred to as neuropil.
2) Brain–gray matter forms the cerebral and cerebellar cortex; white matter lies beneath.
Aggregates of neurons (known as nuclei) may form islands within the white matter.
Figure 9-12 White versus gray matter.
A cross section of H&E-stained spinal cord shows the transition between white matter (left region) and gray matter
(right). The gray matter has many glial cells (G), neuronal cell bodies (N), and neuropil; white matter also contains
glia (G) but consists mainly of axons (A) whose myelin sheaths were lost during preparation, leaving the round
empty spaces shown. Each such space surrounds a dark-stained spot that is a small section of the axon. (X400)

In the folded cerebral cortex, neuroscientists recognize six layers of neurons with different sizes
and shapes. The most conspicuous of these cells are the efferent pyramidal neurons. Neurons of
the cerebral cortex function in the integration of sensory information and the initiation of
voluntary motor responses.
Figure 9-13 Cerebral cortex.
(a) Important neurons of the cerebrum are the pyramidal neurons (P), which are arranged vertically and interspersed
with numerous smaller glial cells, mostly astrocytes, in the eosinophilic neuropil. (X200; H&E)
(b) From the apical ends of pyramidal neurons (P), long dendrites extend in the direction of the cortical surface,
which can be best seen in thick silver-stained sections in which only a few other protoplasmic astrocytes (A) cells
are seen. (X200; Silver)

The sharply folded cerebellar cortex coordinates muscular activity throughout the body and is
organized with three layers:
• A thick outer molecular layer has much neuropil and scattered neuronal cell bodies.
• A thin middle layer consists only of very large neurons called Purkinje cells. These are
conspicuous even in H&E-stained sections, and their dendrites extend throughout the
molecular layer as a branching basket of nerve fibers.
• A thick inner granular layer contains various very small, densely packed neurons
(including granule cells, with diameters of only 4-5 μm) and little neuropil.
Figure 9-14 Cerebellum.
(a) The cerebellar cortex is convoluted with many distinctive small folds, each supported at its center by tracts of
white matter in the cerebellar medulla (M). Each fold has distinct molecular layers (ML) and granular layers (GL).
(X6; Cresyl violet)
(b) Higher magnification shows that the granular layer (GL) immediately surrounding the medulla (M) is densely
packed with several different types of very small rounded neuronal cell bodies. The outer molecular layer (ML)
consists of neuropil with fewer, much more scattered small neurons. At the interface of these two regions a layer of
large Purkinje neuron (P) perikarya can be seen. (X20; H&E)
(c) A single intervening layer contains the very large cell bodies of unique Purkinje neurons (P), whose axons pass
through the granular layer (GL) to join tracts in the medulla and whose multiple branching dendrites ramify
throughout the molecular layer (ML). Dendrites are not seen well with H&E staining. (X40; H&E)
(d) With appropriate silver staining dendrites from each large Purkinje cell (P) are shown to have hundreds of small
branches, each covered with hundreds of dendritic spines. Axons from the small neurons of the granular layer are
unmyelinated and run together into the molecular layer where they form synapses with the dendritic spines of
Purkinje cells. (X40; Silver)

3) Spinal cord–white matter is peripheral; gray matter forms a central “H”-shaped structure
containing the central canal.

The two anterior projections of this gray matter, the anterior horns, contain cell bodies of very
large motor neurons whose axons make up the ventral roots of spinal nerves. The two posterior
horns contain interneurons which receive sensory fibers from neurons in the spinal (dorsal root)
ganglia. Near the middle of the cord the gray matter surrounds a small central canal, which
develops from the lumen of the neural tube, is continuous with the ventricles of the brain, is lined
by ependymal cells, and contains CSF.

Figure 9-15 Spinal cord.


(a) The gray matter contains abundant astrocytes and large neuronal cell bodies, especially those of motor neurons
in the ventral horns.
(b) The white matter surrounds the gray matter and contains primarily oligodendrocytes and tracts of myelinated
axons running along the length of the cord. (Center X5, a, b X100; All silver-stained)
(c) With H&E staining the large motor neurons (N) of the ventral horns show large nuclei, prominent nucleoli, and
cytoplasm rich in Nissl substance, all of which indicate extensive protein synthesis to maintain the axons of these
cells that extend great distances.
(d) In the white commissure ventral to the central canal, tracts (T) run lengthwise along the cord, seen here in cross
section with empty myelin sheaths surrounding axons, as well as small tracts running from one side of the cord to
the other. (Both X200; H&E)

B. Meninges
1) Dura mater – dense irregular connective tissue. There is an epidural space above and a subdural space
below.
2) Arachnoid – simple squamous epithelium that is avascular. Sends down fibrous trabeculae into the
subarachnoid space to connect the arachnoid with the pia mater. Subarachnoid space is filled with CSF
and communicates with the ventricles of the brain; it also contains blood vessels (subarachnoid hemorrhage).
• Arachnoid villi perforate through the dura mater, allowing CSF to flow into the venous sinuses.
3) Pia mater – loose connective tissue, very vascular. Partially follows blood vessels that enter the brain,
resulting in a perivascular space around these vessels lined by pia mater.
4) Blood-Brain Barrier – functional barrier composed of endothelial cells with tight junctions, their
basement membranes, and astrocyte foot processes (see details in “astrocyte”).

Figure 9-16 Diagram of the skull and the layers of the meninges covering the brain. (Textbook of Histology, Leslie
P. Gartner, PhD).

C. Choroid Plexus
1) In roof of 3rd & 4th ventricles and walls of lateral ventricles. Composed of “fibrovascular” core
of loose connective tissue and blood vessels covered by specialized ependymal cells. These cells
contain microvilli and cilia on their apical surface.

2) Main function is to produce cerebrospinal fluid. CSF passes from ventricles to subarachnoid
space and then enters the venous circulation via the arachnoid villi.
Figure 9-17 Choroid plexus.
The choroid plexus consists of ependyma and vascularized pia mater and projects many thin folds from certain walls
of the ventricles.
(a) Section of the bilateral choroid plexus (CP) projecting into the fourth ventricle (V) near the cerebellum. (X12;
Kluver-Barrera stain)
(b) At higher magnification each fold of choroid plexus is seen to be well-vascularized with large capillaries (C) and
covered by a continuous layer of cuboidal ependymal cells (E). (X150)
(c) The choroid plexus is specialized for transport of water and ions across the capillary endothelium and ependymal
layer and the elaboration of these as CSF.

6. Peripheral Nervous System (PNS)


A. Nerve Fibers -- an axon or collection of axons, plus any surrounding sheaths of
ectodermal origin. The presence or absence of a sheath, and the nature of the sheath
(cytoplasm vs. myelin) are used to further characterize nerve fibers.

1) Unmyelinated Fibers
a. Small axons in the PNS are embedded in clefts of the cytoplasm of Schwann cells.
Each Schwann cell can sheathe (but not myelinate) a dozen axons.
b. In the CNS, the majority of axons are unmyelinated. The axons are NOT sheathed by
any type of cell.

2) Myelinated Fibers
Axons that become myelinated are generally larger than 1 μM in diameter. Myelin is
deposited just beyond the axon hillock and continues to near the region of termination of
the axon.
a. Myelin is laid down by Schwann cells. The membranes of the Schwann cell wrap around
the axon several times and fuse to form a myelin sheath. The sheath is interrupted by gaps
called nodes of Ranvier which represent the spaces between adjacent Schwann cells along
the length of the axon. A Schwann cell can only myelinate a single axon; however, each
axon is myelinated by several Schwann cells.
b. In the CNS, myelin is laid down by oligodendrocytes. Each cell can myelinate several
axons.

B. Nerves = groups or bundles of nerve fibers. Covered by connective tissue.


1) Epineurium–the outer layer of dense connective tissue. It surrounds several nerve bundles.
Contains blood vessels and lymphatics.
2) Perineurium–connective tissue and flattened epithelial cells that surrounds each bundle
(fascicle) of nerve fibers. Cells are joined by tight junctions to provide a barrier to passage
of most macromolecules (blood-nerve barrier).
3) Endoneurium–thin layer of reticular fibers that surround individual nerve fibers.
Figure 9-18 Peripheral nerve connective tissue: Epi-, peri-, and endoneurium.
(a) The diagram shows the relationship among these three connective tissue layers in large peripheral nerves. The
epineurium (E) consists of a dense superficial region and a looser deep region that contains the larger blood vessels.
(Reproduced, with permission, from McKinley M, O'Loughlin VD. Human Anatomy. 2nd ed. New York, NY:
McGraw-Hill; 2008).
(b) The micrograph shows a small vein (V) and artery (A) in the deep epineurium (E). Nerve fibers (N) are bundled
in fascicles. Each fascicle is surrounded by the perineurium (P), consisting of a few layers of unusual squamous
fibroblastic cells that are all joined at the peripheries by tight junctions. The resulting blood-nerve barrier helps
regulate the microenvironment inside the fascicle. Axons and Schwann cells are in turn surrounded by a thin layer of
endoneurium. (X140; H&E) (Reproduced, with permission, from Berman I. Color Atlas of Basic Histology. 3rd ed.
New York, NY: McGraw-Hill; 2003).
(c) As shown here and in the diagram, septa (S) of connective tissue often extend from the perineurium into larger
fascicles. The endoneurium (En) and lamellar nature of the perineurium (P) are also shown at this magnification,
along with some adjacent epineurium (E). (X200; PT)
(d) SEM of transverse sections of a large peripheral nerve showing several fascicles, each surrounded by
perineurium and packed with endoneurium around the individual myelin sheaths. Each fascicle contains at least one
capillary. Endothelial cells of these capillaries are tightly joined as part of the blood-nerve barrier and regulate the
kinds of plasma substances released to the endoneurium. Larger blood vessels course through the deep epineurium
that fills the space around the perineurium and fascicles. (X450)
Figure 9-19 Peripheral nerve ultrastructure.
This low-magnification TEM shows a fibroblast (F) surrounded by collagen (C) in the epineurium (E) and three
layers of flattened cells in the perineurium (P) that form another part of the blood-nerve barrier. Inside the
perineurium the endoneurium (En) is rich in reticulin fibers (RF) that surround all Schwann cells. Nuclei of two
Schwann cells (SC) of myelinated axons (A) are visible as well as many unmyelinated axons (UM) within Schwann
cells. (X1200)
Figure 9-20 Small nerves.
Small nerves can be seen in sections from most organs.
(a) In cross section an isolated, resin-embedded nerve is seen to have a thin perineurium (P), one capillary (C), and
many large axons (A) associated with Schwann cells (S). A few nuclei of fibroblasts can be seen in the endoneurium
between the myelinated fibers. (X400; PT)
(b) In longitudinal sections the flattened nuclei of endoneurial fibroblasts (F) and more oval nuclei of Schwann cells
(S) can be distinguished. Nerve fibers are held rather loosely in the endoneurium and in low-magnification
longitudinal section are seen to be wavy rather than straight. This indicates a slackness of fibers within the nerve,
which allows nerves to stretch slightly during body movements with no potentially damaging tension on the fibers.
(X200; H&E) (Reproduced, with permission, from Berman I. Color Atlas of Basic Histology. 3rd ed. New York,
NY: McGraw-Hill; 2003).
(c) In sections of mesentery and other tissues, a highly wavy or tortuous disposition of a single small nerve (N) will
be seen as multiple oblique or transverse pieces as the nerve enters and leaves the area in the section. (X200; H&E)
(d) Often, a section of small nerve will have some fibers cut transversely and others cut obliquely within the same
fascicle, again suggesting the relatively unrestrained nature of the fibers within the endoneurium (E) and
perineurium (P). (X300; H&E) (Reproduced, with permission, from Berman I. Color Atlas of Basic Histology. 3rd
ed. New York, NY: McGraw-Hill; 2003).

C. Ganglia – encapsulated groups of neurons and associated glial cells and connective tissue.
1) Sensory ganglia – receive afferent impulses and transmit to the CNS. Neurons are pseudounipolar and
surrounded by satellite cells. Entire group surrounded by a connective tissue capsule.
2) Autonomic (motor) ganglia – multipolar neurons typically found in walls of organs (GI tract). They do
have a few satellite cells associated with them, but do not have a connective tissue capsule.
Figure 9-21 Sensory and sympathetic ganglia (Stevens & Lowe’s Human Histology, Lowe, James S., BMedSci,
BMBS, DM, FRCPath; Anderson, Peter G., DVM, PhD)
7. Nerve degeneration

Damage to the neuron and its axon leads to axonal degeneration (wallerian degeneration).

Figure 9-22 Degeneration and regeneration of a peripheral nerve (Stevens & Lowe’s Human Histology, Lowe,
James S., BMedSci, BMBS, DM, FRCPath; Anderson, Peter G., DVM, PhD)

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