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Nu =
nucleus
Ax =
Axon
TEM of an
Unmyelinated
Axons
Nu = nucleus of an
Oligodendrocyte
• EM: demonstrates the myelin as a series of concentric layers of
Schwann cell plasma membrane
• mature myelin shows regular lamination with concentric, regular,
dense laminar or period lines of 30Ǻ (3μm) separated by light
intervals of about 100Ǻ (10μm); finer dense lines of 20Ǻ (2μm)
thickness called the intraperiod lines, dissect the inner cytoplasmic
surfaces of the Schwann cell plasma membrane & intraperiod lines
by appositions of the outer surfaces (“jelly roll” hypothesis)
TEM of a
Myelin
Sheath
Myelination of Axons by Oligodendrocytes
Sheath of Schwann (Neurolemma)
• composed of sheath of flattened cells termed the
neurolemmal sheath or neurolemma, forms a thin
sleeve around the myelin, which in turn, surrounds
the axon
• after passing through the white mater, a nerve
fiber leaves the CNS & acquires a thin, distinct
nucleated sheath of Schwann; as the nerve leaves
the CNS, it carries with it the glia cells to form the
sheath of Schwann
• in embryonic life, Schwann cells accompany
outgrowing axons & migrate from branch to
branch until they form complete neurolemmal
sheaths; this thin membrane contain flattened
nuclei that are alternately arranged along the
segments of the nerve fiber; a small Golgi complex
& a few mitochondria can be seen in the
attenuated cytoplasm of these cells of Schwann
• EM: reveals that myelin is actually part of the
Schwann cell being closely applied to the axon,
surrounding it & making several encircling turns
around it; the concentric lamellae of the plasma
membrane of the Schwann cell is called the myelin
sheath; the most superficial lamellae containing
the nucleus is called the neurilemmal sheath or
the Schwann sheath proper
• they are ectodermal in origin & are essential to
the vitality & function of peripheral nerve fibers;
they form myelin but are also necessary for
regeneration of axons; after division, an axon is
regenerated from the central stump, i.e. from the
end continuous with the nerve cell body; it then
grows peripherally to its termination along a
channel formed by Schwann cells; after injury,
these cells can also become phagocytic, removing
cellular debris
• they are not present in the CNS; their function is performed by
oligodendrocytes (oligodendroglia), a type of neuroglia
• myelin contains about 80% lipid, including cholesterol,
phospholipids, glycolipids, & about 20% protein; the high lipid
content explains the whiteness of peripheral nerves (e.g. the white
rami of the autonomic nerves as compared with the gray rami that
contain primarily unmyelinated fibers), as well as the distinction
between white and gray mater of the CNS
Classification of Nerve Fibers:
- may be classified as myelinated or non-myelinated
I. According to their coverings, they may be classified into 4 types for
purposes of microscopic study:
a. Medullated nerve fiber with Neurolemma
b. Medullated nerve fiber without Neurolemma
c. Non-medullated nerve fiber with Neurolemma
d. Non-medullated nerve fiber without Neurolemma
A. Medullated nerve fiber with Neurolemma
• contains both coverings of myelin sheath &
neurolemma or nucleated sheath of Schwann, so
that they are considered the biggest among the
other varieties
• it is made up of a central core of neurofibrils which
are the continuation of the neurofibrils originating
from the cell body; these fibrils are embedded in a
small amount of neuroplasm, called the axoplasm;
the bundle of fibrils constitutes the axis cylinder
called axon at its origin from the cell body; these
fibrils are composed of finer & parallel
neurofilaments; a few, thin mitochondria,
microtubules & slender ER are present in the
axoplasm; covering this axis cylinder is a delicate
membrane called the axolemma of Kuhne
• the myelin sheath covers the axis cylinder; the thickness of the
sheath determines size of the nerve fibers; there are regular
annular deficiencies or constrictions along the course of the fiber
termed the Nodes of Ranvier – these are the places where the
branches or collaterals of the nerve fiber are given off at right
angles (EM); these are the places where the Schwann cells meet &
where nutritive substances easily gain entrance into the axis
cylinder
TEM of a
Myelinated
Axons
Gaps = Nodes of
Ranvier
• the portion of the nerve fiber between 2
successive nodes of Ranvier is called internodal
segment which is longer when the nerve fiber is
thicker; its thickness is due to the increase of
myelin substance & the content of neurofibrils;
the segment appears shorter towards the end of a
nerve due to the diminution of neurofibrils; in the
region of the node, there are finger-like processes
of the neighboring cells of Schwann covering the
nodal area. The gap is important in the saltatory
transfer of action potential for stimulus
propagation along the nerve fiber
• Schmidt Lanterman’s Lines/Incisures of Schmidt –
several oblique & radially placed lines in the
myelin sheath within each internodal segment
(when stained with osmic acid); the portion
between 2 successive Schmidt incisures is called
myelin segment; on EM, these represents areas of
local separation of the spirally wrapped myelin
lamellae
• the myelin is covered externally by the nucleated
sheath of Schwann; this is a thin, nucleated
membrane whose nuclei appear alternately,
containing one for each internodal segment. At
the nodes of Ranvier, it dips & comes in contact
with the axis cylinder; this covering also
disappears like the myelin before the nerve fiber
ends so that all nerve endings are naked fibers
• there is a thin, delicate fibrous connective sheath outside
the neurolemma termed CT sheath of Key & Retzius or the
Sheath of Henle – this forms a part of the endoneurium
that carries the capillary blood vessels to the individual
nerve fibers
B. Medullated nerve fiber without Neurolemma
• found only in the white mater (CNS) corresponding to the
portions of axons & dendrons that traverse the white
mater after passing the gray mater
• it is composed of the axis cylinder surrounded by the
axolemma & which is covered by the myelin sheath
• the nodes of Ranvier are not prominent
• seen lying in rows between the myelinated fibers are the
interfascicular glia cells (oligodendrocytes), arranged in the
form of a membrane
Bundles of nerve
fibers (BNF)
surrounded by an
Epineurium (Epn)
Perineurium (arrow)
surrounds each
bundle
SEM of a nerve bundle
W
• White mater consists of nerve fibers, neuroglia & blood vessels; it
surrounds the gray mater & its white color is caused by the high
proportion of myelinated nerve fibers
Cerebellum
• Gray mater is located in the surface as a thin cortex overlying the
centrally placed white mater; there are also small collections of
nerve cell (nuclei) in the central parts of the cerebellum (nucleus
dentatus, emboliformis, globosi, & fastigii)
Low mag of the
Cerebellum or
Cerebellar
Cortex
(arrow)
Structure:
• the cerebellar cortex is uniform in all parts of the
cerebellum
• it has 3 layers (from the surface proceeding inwards):
1. Outer molecular layer – relatively few cells & few
myelinated fibers
- the superficial cells are stellate in shape with short, thin
dendrites & fine myelinated axons that run horizontally
- the large, deeper stellate cells situated in the vicinity of
the Purkinje cell bodies also are known as “basket cells”;
many of their numerous branching dendrites ascend to the
surface & the axons run parallel to the sagittal plane
ending in relation or around the bodies of the Purkinje
cells; a narrow horizontal plexus of myelinated fibers
composed of collaterals from the axons of Purkinje cell is
found only in its deepest portion
Cerebellum or Cerebellar Cortex
2. Middle Ganglionic layer – consists of a single row of large, flask-
shaped bodies with 2 processes (Purkinje cell); each cell gives off 2
or 3 main dendrites which enters the molecular layer forming a fan-
shaped dendritic arborization extending to the surface; the axon
rises from the part of the cell opposite the dendrites, acquires a
myelin sheath, passes through the granular layer & enters the
underlying white myelin to go to one of the deep cerebellar nuclei
or to some other parts of the cortex
High mag of the Purkinje Cells of the Cerebellum
The Dendritic Tree of Purkinje Cells
Cajal’s drawing of
Purkinje Cells
3. Granular layer – in ordinary stains, presents the
appearance of closely packed chromatic nuclei
resembling lymphocytes; irregular light spaces
constitute the so-called “islands” or “glomeruli”;
the granule cells are small, multipolar with short 3
or 4 dendrites which arborized in peculiar claw-
like endings; the unmyelinated axons ascend to
the molecular & run transversely to the dendritic
expansion of the Purkinje cells
- scattered throughout the layer are the Golgi type
II cells or cells of Gehucten which have vesicular
nuclei & chromophilic bodies; their branching
dendrites enter the molecular layer, while their
terminal fibrils come in contact with the dendritic
terminals of the terminals of the granule cell
• it has the terminations of the mossy & climbing
fibers passing into the cerebellar cortex from the
white mater of the brain stem & spinal cord; the
mossy fibers are thick & synapse in cells of the
granular layer, while climbing fibers pass through
the granular layer to terminate on Purkinje cells
Cerebral Cortex
• Gray mater is located on the surface as cerebral
cortex
• White mater underneath the gray cortex
(medullary substance) is composed of bundles of
myelinated fibers passing in all directions; these
fibers are supported by neuroglia & are
functionally composed of 3 main groups:
association fibers, projection fibers, &
commissural fibers
The Cerebral Hemispheres
• a striking feature of cortical structure is its
lamination; in sections stained by Nissl method,
cell bodies are not uniformly distributed, but are
arranged in superimposed horizontal layers
• proceeding from the surface of the cortex towards
the white or medullary substance, these layers
include:
I. Molecule/plexiform layer
it contains relatively few cells of 2 types:
a. cells with horizontal axons (horizontal cells of Cajal)
b. Golgi type II cells
within this are found the terminal dendritic
ramifications of the pyramidal & fusiform cells
from the deeper layers & the axonal endings of
the cells of Martinotti
Cerebrum or Cerebral Cortex
M – molecular; G - granular; P – pyramidal layers
M
G
P
II. External Granular layer
consists of numerous closely packed small cells of
triangular or pyramidal shape whose apical
dendrites terminate in the molecular layer; their
axons descend to the deepest cortical layers where
many terminate; some enter the white mater as
association fibers
III. External Pyramidal layer
composed of typical well-formed pyramidal
neurons
there are 2 sublayers:
a. superficial layer of medium-sized pyramids
b. deeper layer of large ones
their apical dendrites go to the first layer, while
most of their axons enter the white mater as
association or commissural fibers
in the most superficial part of the layer, there are a
great number of horizontal myelinated fibers
constituting the band of Kaes-Bechterew
IV. Internal Granular layer
composed chiefly of closely packed stellate cells;
these are very small with short axons ramifying
within the layer; other larger cells have
descending axons terminating in deeper layers, or
may enter the white substance
the whole layer is permeated by a dense
horizontal plexus of myelinated fibers forming
external band Baillarger, which are considered to
be mainly the terminal ramifications of the
thalamo-cortical fibers
V. Internal Pyramidal/Ganglion layer
consists principally of medium-sized & large pyramidal neurons
(Giant pyramidal cells of Betz), which constitutes the origin of the
pyramidal tract – the corticospinal & corticobulbar tracts
they are intermingled with granule cells & the cells of Martinotti
their axons enter the white mater chiefly as projection fibers; the
rich horizontal fiber plexus in the deeper portion of this layer
constitutes the internal band of Baillarger
VI. Multiform/Fusiform cell layer
contains predominantly spindle-shaped cells whose long axis are
perpendicular to the cortical surface; it contains also granule,
Martinotti, & stellate cells
it may be subdivided into:
a. an upper sublayer of more densely packed, larger cells
b. a lower sublayer in which the smaller cells are loosely arranged
the whole layer is pervaded by fiber bundles that enter or leave the
medullary substance
Cortical Cells & Fibers:
• the principal types of cells found in the cerebral
cortex are:
1. Pyramidal cells – the most characteristic of the
cortex, having the form of a pyramidal or triangle
whose upper pointed end is continued toward the
brain surface as the apical dendrite
- there are also a number of more or less
horizontally running basal dendrites that spring
from the cell body & arborize in the vicinity of the
cell; the axon emerges from the base of the cell &
descends towards the medullary substance to
enter the white mater as a projection or
association fiber
Pyramidal Cells of the Cerebral Cortex
Silver stained Pyramidal Cells of the Cerebral Cortex
Cajal Pyramidal Cell Drawing (circa 1900) and replica
- they contain a large vesicular nucleus with prominent
Nissl bodies; they are classified usually as small, medium,
and large neurons
- the giant pyramidal cells of Betz may be more than 100
micra in height & are found in the motor area of the pre-
central gyrus
2. Stellate/Granule cells – are small with a polygonal or
triangle shape
- have dark-staining nuclei & scanty cytoplasm, varying in
size from 4 – 8 micra
- these cells have a number of dendrites passing in all
directions, and a short axon that ramifies close to the cell
body
- some resemble pyramidal cells in that they have no apical
dendrite that extends to the surface, these are known as
stellate, pyramidal cells or star pyramids
- they are found throughout all layers of the cortex but are
especially numerous in layer IV
3. Spindle/Fusiform cells – found mainly in the
deepest layer, with their long axis usually vertical
to the surface
4. Horizontal cells of Cajal – small fusiform or pear-
shaped cells found in the most superficial layer;
their long axons run horizontally & arborize within
the layer
5. Cells of Martinotti – are small, triangular or
polygonal cells where axons are directed toward
the surface
- they are present in practically all cell layers &
extend variable distance; some arborize in the
same layer, while others send collateral to a
number of layers
• fibers in the cerebral cortex are disposed radially &
tangentially; they include the axons of pyramidal,
fusiform & stellate cells which leave the cortex as
projection or association fibers and the entering
afferent projection & association fibers
terminating within the cortex; ascending axons of
the cells of Martinotti, likewise, have a vertical
course
• the tangential fibers, running horizontal to the
surface, are composed principally of the terminal
branches of the afferent projection & association
fibers, the axons of the horizontal & granule cells,
& the terminal branches of collaterals from the
pyramidal & fusiform cells
Meninges
Cerebellum
(arrow)
• it is composed of loose CT of the pia mater,
covered by a simple cuboidal or low columnar
epithelium of neural tube origin; the epithelial
cells possess numerous irregular microvilli & there
are junctional complexes near their free ends; the
capillaries beneath the epithelium are unlike those
found elsewhere in the brain, they have thin walls
& fenestrations or pores closed by thin
diaphragms; the endothelial cells are held
together by tight junctions; these are the site of
“blood-CSF barrier”
• its CT is quite cellular, containing many
macrophages; they engulf intravenously injected
supravital dyes (trypan blue)
• its main function is to secrete CSF by the epithelial
cells covering the plexuses
The Choroid
Plexus
A capillary tuft
surrounded by
Ependymal Cells
The Choroid Plexus – Capillaries surrounded by
Ependymal Cells (arrow)
Cerebrospinal Fluid
• it is the fluid elaborated by the choroid plexuses
that completely fills the ventricles, the central
canal of the spinal cord, the subarachnoid space,
& the perivascular space
• it is important for the metabolism of the CNS,
protecting the nervous system from concussions &
mechanical injuries; it bathes & suspends the CNS,
thus cushioning the nervous tissue against trauma
• its amount is variable, estimated at 80 – 150ml; it
is slightly viscous, has a low specific gravity (1.004
– 1.008), contains traces of proteins, small
quantities of inorganic salt & dextrose, and a few
lymphocytes; it resembles the aqueous humor of
the eye more closely than it does any other fluid
of the body
• its sources are primarily the blood vessels of the
choroid plexus, the pia mater and the brain
substance; it is constantly renewed, circulating
slowly through the brain ventricles, and through
the meshes of the subarachnoid space
*ventricular fluid normally flows from the lateral
ventricles of the cerebral hemispheres, through
the foramina of Monroe 3rd ventricle