Lecture
THE NERVOUS
SYSTEM is the most complicated and highly organized of the various systems
which make up the human body. It is the mechanism concerned with the
correlation and integration of various bodily processes and the reactions and
adjustments of the organism to its environment. In addition the cerebral cortex
is concerned with conscious life. It may be divided into two parts, central
and peripheral.
The central nervous system consists of the encephalon or brain,
contained within the cranium, and the medulla spinalis or spinal cord
lodged
in the vertebral canal; the two portions are continuous with one another at the
level of the upper border of the atlas vertebra.
Neuron Theory
The nerve cell and its processes collectively constitute what is termed
a neuron, and Waldeyer formulated the theory that the nervous system is
built up of numerous neurons, “anatomically and genetically independent of one
another.” According to this theory (neuron theory) the processes of one
neuron only come into contact, and are never in direct continuity, with those
of other neurons; while impulses are transmitted from one nerve cell to another
through these points of contact, the synapses.
The synapse or synaptic membrane seems to allow nervous
impulses to pass in one direction only, namely, from the terminals of the
axis-cylinder to the dendrons. This theory is based on the following facts,
viz.: (1) embryonic nerve cells or neuroblasts are entirely distinct from one
another; (2) when nervous tissues are stained by the Golgi method no continuity
is seen even between neighboring neurons; and (3) when degenerative changes
occur in nervous tissue, either as the result of disease or experiment, they
never spread from one neuron to another, but are limited to the individual
neurons, or groups of neurons, primarily affected. It must, however, be added
that within the past few years the validity of the neuron theory has been
called in question by certain eminent histologists, who maintain that by the
employment of more delicate histological methods, minute fibrils can be
followed from one nerve cell into another. Their existence, however, in the
living is open to question. Mott and Marinesco made careful examinations of
living cells, using even the ultramicroscope and agree that neither Nissl
bodies nor neurofibrils are present in the living state.
For the present we may look upon the neurons as the units or structural
elements of the nervous system. All the neurons are present at birth which are
present in the adult, their division ceases before birth; they are not all
functionally active at birth, but gradually assume functional activity. There
is no indication of any regeneration after the destruction of the cell-body of
any individual neuron.
The peripheral nervous system consists of a series of nerves by
which the central nervous system is connected with the various tissues of the
body. For descriptive purposes these nerves may be arranged in two groups, cerebrospinal
and sympathetic, the arrangement, however, being an arbitrary one, since
the two groups are intimately connected and closely intermingled. Both the
cerebrospinal and sympathetic nerves have nuclei of origin (the somatic
efferent and sympathetic efferent) as well as nuclei of termination (somatic
afferent and sympathetic afferent) in the central nervous system. The
cerebrospinal nerves are forty-three in number on either side—twelve cranial,
attached to the brain, and thirty-one spinal, to the medulla spinalis.
They are associated with the functions of the special and general senses and
with the voluntary movements of the body. The sympathetic nerves transmit the
impulses which regulate the movements of the viscera, determine the caliber of
the bloodvessels, and control the phenomena of secretion. In relation with them
are two rows of central ganglia, situated one on either side of the
middle line in front of the vertebral column; these ganglia are intimately
connected with the medulla spinalis and the spinal nerves, and are also joined
to each other by vertical strands of nerve fibers so as to constitute a pair of
knotted cords, the sympathetic trunks, which reach from the base of the
skull to the coccyx. The sympathetic nerves issuing from the ganglia form three
great prevertebral plexuses which supply the thoracic, abdominal, and pelvic
viscera; in relation to the walls of these viscera intricate nerve plexuses and
numerous peripheral ganglia are found.
1. Structure of the Nervous System
The nervous tissues are composed of nerve cells and their various
processes, together with a supporting tissue called neuroglia, which,
however, is found only in the brain and medulla spinalis. Certain long
processes of the nerve cells are of special importance, and it is convenient to
consider them apart from the cells; they are known as nerve fibers.
To the naked eye a difference is obvious between certain portions of the
brain and medulla spinalis, viz., the gray substance and the white
substance. The gray substance is largely composed of nerve cells, while the
white substance contains only their long processes, the nerve fibers. It is in
the former that nervous impressions are received, stored, and transformed into
efferent impulses, and by the latter that they are conducted. Hence the gray
substance forms the essential constituent of all the ganglionic centers, both
those in the isolated ganglia and those aggregated in the brain and medulla
spinalis; while the white substance forms the bulk of the commissural portions
of the nerve centers and the peripheral nerves.
Neuroglia.—Neuroglia, the peculiar ground
substance in which are imbedded the true nervous constituents of the brain and
medulla spinalis, consists of cells and fibers. Some of the cells are stellate
in shape, with ill-defined cell body, and their fine processes become neuroglia
fibers, which extend radially and unbranched
among the nerve cells and fibers which they aid in supporting. Other
cells give off fibers which branch repeatedly. Some of the fibers start from
the epithelial cells lining the ventricles of the brain and central canal of
the medulla spinalis, and pass through the nervous tissue, branching repeatedly
to end in slight enlargements on the pia mater. Thus, neuroglia is evidently a
connective tissue in function but is not so in development; it is ectodermal in
origin, whereas all connective tissues are mesodermal.
Nerve Cells are largely aggregated in the gray
substance of the brain and medulla spinalis, but smaller collections of these
cells also form the swellings, called ganglia, seen on many nerves. These
latter are found chiefly upon the spinal and cranial nerve roots and in
connection with the sympathetic nerves.
The nerve cells vary in shape and size, and have one or more processes.
They may be divided for purposes of description into three groups, according to
the number of processes which they possess: (1) Unipolar cells, which
are found in the spinal ganglia; the single process, after a short course, divides
in a T-shaped manner. (2) Bipolar cells, also found in the spinal
ganglia when the cells are in an embryonic condition. They are best
demonstrated in the spinal ganglia of fish. Sometimes the processes come off
from opposite poles of the cell, and the cell then assumes a spindle shape; in
other cells both processes emerge at the same point. In some cases where two
fibers are apparently connected with a cell, one of the fibers is really
derived from an adjoining nerve cell and is passing to end in a ramification
around the ganglion cell, or, again, it may be coiled spirally around the nerve
process which is issuing from the cell. (3) Multipolar cells, which are
pyramidal or stellate in shape, and characterized by their large size and by
the numerous processes which issue from them. The processes are of two kinds:
one of them is termed the axis-cylinder process or axon because
it becomes the axis-cylinder of a nerve fiber. The others are termed the protoplasmic
processes or dendrons; they begin to divide and subdivide soon after
they emerge from the cell, and finally end in minute twigs and become lost
among the other elements of the nervous tissue.
The body of the nerve cell, known as the cyton, consists of a
finely fibrillated protoplasmic material, of a reddish or yellowishbrown color,
which occasionally presents patches of a deeper tint, caused by the aggregation
of pigment granules at one side of the nucleus, as in the substantia nigra and
locus cæruleus of the brain. The protoplasm also contains peculiar angular
granules, which stain deeply with basic dyes, such as methylene blue; these are
known as Nissl’s granules. They extend into the dendritic processes but
not into the axis-cylinder; the small clear area at the point of exit of the
axon in some cell types is termed the cone of origin. These granules
disappear (chromatolysis) during fatigue or after prolonged stimulation
of the nerve fibers connected with the cells. They are supposed to represent a
store of nervous energy, and in various mental diseases are deficient or
absent. The nucleus is, as a rule, a large, well-defined, spherical body, often
presenting an intranuclear network, and containing a well-marked nucleolus.
In addition to the protoplasmic network described above, each nerve cell
may be shown to have delicate neurofibrils running through its substance; these fibrils are continuous with the fibrils of
the axon, and are believed to convey nerve impulses. Golgi has also described
an extracellular network, which is probably a supporting structure.
Nerve Fibers.—Nerve fibers are found universally
in the peripheral nerves and in the white substance of the brain and medulla
spinalis. They are of two kinds—viz., medullated or white fibers,
and non-medullated or gray fibers.
The medullated fibers form the white part of the brain and
medulla spinalis, and also the greater part of every cranial and spinal nerve,
and give to these structures their opaque, white aspect. When perfectly fresh
they appear to be homogeneous; but soon after removal from the body each fiber
presents, when examined by transmitted light, a double outline or contour, as
if consisting of two parts . The central portion
is named the axis-cylinder; around this is a sheath of fatty material,
staining black with osmic acid, named the white substance of Schwann or medullary
sheath, which gives to the fiber its double contour, and the whole is
enclosed in a delicate membrane, the neurolemma, primitive sheath, or nucleated
sheath of Schwann.
The axis-cylinder is the essential part of the nerve fiber, and
is always present; the medullary sheath and the neurolemma are occasionally
absent, expecially at the origin and termination of the nerve fiber. The
axis-cylinder undergoes no interruption from its origin in the nerve center to
its peripheral termination, and must be regarded as a direct prolongation of a
nerve cell. It constitutes about one-half or one-third of the nerve fiber,
being greater in proportion in the fibers of the central organs than in those
of the nerves. It is quite transparent, and is therefore indistinguishable in a
perfectly fresh and natural state of the nerve. It is made up of exceedingly
fine fibrils, which stain darkly with gold chloride and at its termination may
be seen to break up into these fibrillæ. The fibrillæ have been termed the primitive
fibrillæ of Schultze. The axis-cylinder is said by some to be enveloped in
a special reticular sheath, which separates it from the medullary sheath, and
is composed of a substance called neurokeratin. The more common opinion
is that this network or reticulum is contained in the white matter of Schwann,
and by some it is believed to be produced by the action of the reagents
employed to show it.
The medullary sheath, or white matter of Schwann is
regarded as being a fatty matter in a fluid state, which insulates and protects
the essential part of the nerve—the axis-cylinder. It varies in thickness, in
some forming a layer of extreme thinness, so as to be scarcely distinguishable,
in others forming about one-half the nerve fiber. The variation in diameter of
the nerve fibers (from 2 to 16μ) depends
mainly upon the amount of the white substance, though the axis cylinder also
varies within certain limits. The medullary sheath undergoes interruptions in
its continuity at regular intervals, giving to the fiber the appearance of
constriction at these points: these are known as the nodes of Ranvier.
The portion of nerve fiber between two nodes is called an internodal
segment. The neurolemma or primitive sheath is not interrupted at the
nodes, but passes over them as a continuous membrane. If the fiber be treated
with silver nitrate the reagent penetrates the neurolemma at the nodes, and on
exposure to light reduction takes place, giving rise to the appearance of black
crosses, Ranvier’s crosses, on the axis-cylinder. There may also be seen
transverse lines beyond the nodes termed Frommann’s lines the
significance of these is not understood. In addition to these interruptions
oblique clefts may be seen in the medullary sheath, subdividing it into
irregular portions, which are termed medullary segments, or segments
of Lantermann, there is reason to believe that these clefts are
artificially produced in the preparation of the specimens. Medullated nerve
fibers, when examined in the fresh condition, frequently present a beaded or
varicose appearance: this is due to manipulation and pressure causing the oily
matter to collect into drops, and in consequence of the extreme delicacy of the
primitive sheath, even slight pressure will cause the transudation of the fatty
matter, which collects as drops of oil outside the membrane.
The neurolemma or primitive sheath presents the appearance
of a delicate, structureless membrane. Here and there beneath it, and situated
in depressions in the white matter of Schwann, are nuclei surrounded by a small
amount of protoplasm. The nuclei are oval and somewhat flattened, and bear a definite
relation to the nodes of Ranvier, one nucleus generally lying in the center of
each internode. The primitive sheath is not present in all medullated nerve
fibers, being absent in those fibers which are found in the brain and medulla
spinalis.
Wallerian Degeneration.—When nerve fibers are cut
across, the central ends of the fibers degenerate as far as the first node of
Ranvier; but the peripheral ends degenerate simultaneously throughout their
whole length. The axons break up into fragments and become surrounded by drops
of fatty substance which are formed from the breaking down of the medullary
sheath. The nuclei of the primitive sheath proliferate, and finally absorption
of the axons and fatty substance occurs. If the cut ends of the nerve be sutured
together regeneration of the nerve fibers takes place by the downgrowth of
axons from the central end of the nerve. At one time it was believed that the
regeneration was peripheral in origin, but this has been disproved, the
proliferated nuclei in the peripheral portions taking part merely in the
formation of the so-called scaffolding along which the new axons pass.
Non-medullated Fibers.—Most of the fibers of the
sympathetic system, and some of the cerebrospinal, consist of the gray
or gelatinous nerve fibers (fibers of Remak). Each of these consists of an axis-cylinder to
which nuclei are applied at intervals. These nuclei are believed to be in
connection with a delicate sheath corresponding with the neurolemma of the
medullated nerve fiber. In external appearance the non-medullated nerve fibers
are semitransparent and gray or yellowish gray. The individual fibers vary in
size, generally averaging about half the size of the medullated fibers.
Structure of the Peripheral Nerves and Ganglia.—The
cerebrospinal nerves consist of numerous nerve fibers collected together
and enclosed in membranous sheaths. A small bundle of fibers, enclosed in a
tubular sheath, is called a funiculus; if the nerve is of small size, it
may consist only of a single funiculus; but if large, the funiculi are
collected together into larger bundles or fasciculi, which are bound
together in a common membranous investment. In structure the common membranous
investment, or sheath of the whole nerve (epineurium), as well as the
septa given off from it to separate the fasciculi, consist of connective
tissue, composed of white and yellow elastic fibers, the latter existing in
great abundance. The tubular sheath of the funiculi (perineurium) is a
fine, smooth, transparent membrane, which may be easily separated, in the form
of a tube, from the fibers it encloses; in structure it is made up of
connective tissue, which has a distinctly lamellar arrangement. The nerve
fibers are held together and supported within the funiculus by delicate
connective tissue, called the endoneurium. It is continuous with septa
which pass inward from the innermost layer of the perineurium, and shows a
ground substance in which are imbedded fine bundles of fibrous connective
tissue running for the most part longitudinally. It serves to support capillary
vessels, arranged so as to form a net-work with elongated meshes. The
cerebrospinal nerves consist almost exclusively of medullated nerve fibers,
only a very small proportion of non-medullated being present.
The bloodvessels supplying a nerve end in a minute capillary plexus, the
vessels composing which pierce the perineurium, and run, for the most part,
parallel with the fibers; they are connected together by short, transverse
vessels, forming narrow, oblong meshes, similar to the capillary system of
muscle. Fine non-medullated nerve fibers, vasomotor fibers, accompany
these capillary vessels, and break up into elementary fibrils, which form a
network around the vessels. Horsley has demonstrated certain medullated fibers
running in the epineurium and terminating in small spheroidal tactile
corpuscles or end bulbs of Krause. These nerve fibers, which
The nerve fibers, so far as is at present known, do not coalesce, but
pursue an uninterrupted course from the center to the periphery. In separating
a nerve, however, into its component funiculi, it may be seen that these do not
pursue a perfectly insulated course, but occasionally join at a very acute
angle with other funiculi proceeding in the same direction; from this, branches
are given off, to joint again in like manner with other funiculi. It must be
distinctly understood, however, that in these communications the individual
nerve fibers do not coalesce, but merely pass into the sheath of the adjacent
nerve, become intermixed with its nerve fibers, and again pass on to
intermingle with the nerve fibers in some adjoining funiculus.
Nerves, in their course, subdivide into branches, and these frequently
communicate with branches of a neighboring nerve. The communications which thus
take place form what is called a plexus. Sometimes a plexus is formed by
the primary branches of the trunks of the nerves—as the cervical, brachial,
lumbar, and sacral plexuses—and occasionally by the terminal funiculi, as in
the plexuses formed at the periphery of the body. In the formation of a plexus,
the component nerves divide, then join, and again subdivide in such a complex
manner that the individual funiculi become interlaced most intricate’y; so that
each branch leaving a plexus may contain filaments from all the primary nervous
trunks which form the plexus. In the formation also of smaller plexuses at the
periphery of the body there is a free interchange of the funiculi and primitive
fibers. In each case, however, the individual fibers remain separate and
distinct.
It is probable that through this interchange of fibers, every branch
passing off from a plexus has a more extensive connection with the spinal cord
than if it had proceeded to its distribution without forming connections with
other nerves. Consequently the parts supplied by these nerves have more extended
relations with the nervous centers; by this means, also, groups of muscles may
be associated for combined action.
The sympathetic nerves are constructed in the same manner as the
cerebrospinal nerves, but consist mainly of non-medullated fibers, collected in
funiculi and enclosed in sheaths of connective tissue. There is, however, in
these nerves a certain admixture of medullated fibers. The number of the latter
varies in different nerves, and may be estimated by the color of the nerve.
Those branches of the sympathetic, which present a well-marked gray color, are
composed chiefly of non-medullated nerve fibers, intermixed with a few
medullated fibers; while those of a white color contain many of the latter
fibers, and few of the former.
The cerebrospinal and sympathetic nerve fibers convey various
impressions. The sensory nerves, called also centripetal or afferent
nerves, transmit to the nervous centers impressions made upon the
peripheral extremities of the nerves, and in this way the mind, through the medium
of the brain, becomes conscious of external objects. The centrifugal or efferent
nerves transmit impressions from the nervous centers to the parts to which
the nerves are distributed, these impressions either exciting muscular
contraction or influencing the processes of nutrition, growth, and secretion.
Origins and Terminations of Nerves.—By the
expression “the terminations of nerve fibers” is signified their connections
with the nerve centers and with the parts they supply. The former are sometimes
called their origins or central terminations; the latter their peripheral
terminations.
Origins of Nerves.—The origin in some cases is
single—that is to say, the whole nerve emerges from the nervous center by a
single root; in other instances the nerve arises by two or more roots which
come off from different parts of the nerve center, sometimes widely apart from
each other, and it often happens, when a nerve arises in this way by two roots,
that the functions of these two roots are different; as, for example, in the
spinal nerves, each of which arises by two roots, the anterior of which is
motor, and the posterior sensory. The point where the nerve root or roots
emerge from the surface of the nervous center is named the superficial
or apparent origin, but the fibers of the nerve can be traced for a
certain distance into the substance of the nervous center to some portion of
the gray matter, which constitutes the deep or real origin of the
nerve. The centrifugal or efferent nerve fibers originate in the nerve cells of
the gray substance, the axis-cylinder processes of these cells being prolonged
to form the fibers. In the case of the centripetal or afferent nerves the
fibers grow inward either from nerve cells in the organs of special sense, e.
g., the retina, or from nerve cells in the ganglia. Having entered the
nerve center they branch and send their ultimate twigs among the cells,
without, however, uniting with them.
Peripheral Terminations of Nerves.—Nerve fibers
terminate peripherally in various ways, and these may be conveniently studied
in the sensory and motor nerves respectively. The terminations of the sensory
nerves are dealt with in the section on Sense Organs.
Motor nerves can be traced into either unstriped
or striped muscular fibers. In the unstriped or involuntary muscles
the nerves are derived from the sympathetic, and are composed mainly of
non-medullated fibers. Near their terminations they divide into numerous
branches, which communicate and form intimate plexuses. At the junction of the
branches small triangular nuclear bodies (ganglion cells) are situated. From
these plexuses minute branches are given off which divide and break up into the
ultimate fibrillæ of which the nerves are composed. These fibrillæ course
between the involuntary muscle cells, and, according to Elischer, terminate on
the surfaces of the cells, opposite the nuclei, in minute swellings.
In the striped or voluntary muscle the nerves supplying
the muscular fibers are derived from the cerebrospinal nerves, and are composed
mainly of medullated fibers. The nerve, after entering the sheath of the
muscle, breaks up into fibers or bundles of fibers, which form plexuses, and
gradually divide until, as a rule, a single nerve fiber enters a single
muscular fiber. Sometimes, however, if the muscular fiber be long, more than
one nerve fiber enters it. Within the muscular fiber the nerve terminates in a
special expansion, called by Kühne, who first accurately described it, a motor
end-plate. The nerve fiber, on approaching the muscular fiber, suddenly
loses its medullary sheath, the neurolemma becomes continuous with the
sarcolemma of the muscle, and only the axis-cylinder enters the muscular fiber.
There it at once spreads out, ramifying like the roots of a tree, immediately
beneath the sarcolemma, and becomes imbedded in a layer of granular matter,
containing a number of clear, oblong nuclei, the whole constituting an
end-plate from which the contractile wave of the muscular fiber is said to start.
Ganglia are small aggregations of nerve cells. They are
found on the posterior roots of the spinal nerves; on the sensory roots of the
trigeminal, facial, glossopharyngeal, and vagus nerves, and on the acoustic
nerves. They are also found in connection with the sympathetic nerves. On
section they are seen to consist of a reddish-gray substance, traversed by
numerous white nerve fibers; they vary considerably in form and size; the
largest are found in the cavity of the abdomen; the smallest, not visible to
the naked eye, exist in considerable numbers upon the nerves distributed to the
different viscera. Each ganglion is invested by a smooth and firm, closely
adhering, membranous envelope, consisting of dense areolar tissue; this sheath
is continuous with the perineurium of the nerves, and sends numerous processes
into the interior to support the bloodvessels supplying the substance of the
ganglion.
In structure all ganglia are essentially similar, consisting of the same
structural elements—viz., nerve cells and nerve fibers. Each nerve cell has a
nucleated sheath which is continuous with the neurolemma of the nerve fiber
with which the cell is connected. The nerve cells in the ganglia of the spinal
nerves are pyriform in shape, and have each a single process. A short distance
from the cell and while still within the ganglion this process divides in a
T-shaped manner, one limb of the cross-bar turning into the medulla spinalis,
the other limb passing outward to the periphery. In the sympathetic ganglia the
nerve cells are multipolar and each has one axis-cylinder process and several
dendrons; the axon emerges from the ganglion as a non-medullated nerve fiber.
Similar cells are found in the ganglia connected with the trigeminal nerve, and
these ganglia are therefore regarded as the cranial portions of the sympathetic
system. The sympathetic nervous system includes those portions of the nervous
mechanism in which a medullated nerve fiber from the central system passes to a
ganglion, sympathetic or peripheral, from which fibers, usually non-medullated,
are distributed to such structures, e. g., bloodvessels, as are not
under voluntary control. The spinal and sympathetic ganglia differ somewhat in
the size and disposition of the cells and in the number of nerve fibers
entering and leaving them. In the spinal ganglia the nerve cells are much
larger and for the most part collected in groups near the periphery, while the
fibers, which are mostly medullated, traverse the central portion of the
ganglion; whereas in the sympathetic ganglia the cells are smaller and
distributed in irregular groups throughout the whole ganglion; the fibers also
are irregularly scattered; some of the entering ones are medullated, while many
of those leaving the ganglion are non-medullated.
Fasciculi, tracts or fiber systems are groups
of axons having homologous origin and homologous distribution (as regards their
collaterals, subdivisions and terminals) and are often named in accordance with
their origin and termination, the name of the nucleus or the location of the
cell body from which the axon or fiber arises preceding that of the nucleus or
location of its termination. A given topographical area seldom represents a
pure tract, as in most cases fibers of different systems are mixed.
The Spinal Cord or Medulla Spinalis
The medulla
spinalis or spinal cord forms the elongated, nearly cylindrical,
part of the central nervous system which occupies the upper two-thirds of the
vertebral canal. Its average length in the male is about
The position of the medulla spinalis varies with the movements of the
vertebral column, its lower extremity being drawn slightly upward when the
column is flexed. It also varies at different periods of life; up to the third
month of fetal life the medulla spinalis is as long as the vertebral canal, but
from this stage onward the vertebral column elongates more rapidly than the
medulla spinalis, so that by the end of the fifth month the medulla spinalis
terminates at the base of the sacrum, and at birth about the third lumbar
vertebra.
The medulla spinalis does not fill the part of the vertebral canal in
which it lies; it is ensheathed by three protective membranes, separated from
each other by two concentric spaces. The three membranes are named from without
inward, the dura mater, the arachnoid, and the pia mater.
The dura mater is a strong, fibrous membrane which forms a wide, tubular
sheath; this sheath extends below the termination of the medulla spinalis and
ends in a pointed cul-de-sac at the level of the lower border of the second
sacral vertebra. The dura mater is separated from the wall of the vertebral
canal by the epidural cavity, which contains a quantity of loose areolar
tissue and a plexus of veins; between the dura mater and the subjacent
arachnoid is a capillary interval, the subdural cavity, which contains a
small quantity of fluid, probably of the nature of lymph. The arachnoid
is a thin, transparent sheath, separated from the pia mater by a comparatively
wide interval, the subarachnoid cavity, which is filled with cerebrospinal
fluid. The pia mater closely invests the medulla spinalis and sends
delicate septa into its substance; a narrow band, the ligamentum
denticulatum, extends along each of its lateral surfaces and is attached by
a series of pointed processes to the inner surface of the dura mater.
Thirty-one pairs of spinal nerves spring from the medulla spinalis, each
nerve having an anterior or ventral, and a posterior or dorsal root, the latter
being distinguished by the presence of an oval swelling, the spinal
ganglion, which contains numerous nerve cells. Each root consists of
several bundles of nerve fibers, and at its attachment extends for some
distance along the side of the medulla spinalis. The pairs of spinal nerves are
grouped as follows: cervical 8, thoracic 12, lumbar 5, sacral 5, coccygeal 1,
and, for convenience of description, the medulla spinalis is divided into
cervical, thoracic, lumbar and sacral regions, corresponding with the
attachments of the different groups of nerves.
Enlargements.—The medulla spinalis is not quite
cylindrical, being slightly flattened from before backward; it also presents
two swellings or enlargements, an upper or cervical, and a lower or lumbar.
The cervical enlargement is the more pronounced, and corresponds
with the attachments of the large nerves which supply the upper limbs. It
extends from about the third cervical to the second thoracic vertebra, its
maximum circumference (about
The lumbar enlargement gives attachment to the nerves which
supply the lower limbs. It commences about the level of the ninth thoracic
vertebra, and reaches its maximum circumference, of about
The Anterior Median Fissure (fissura mediana anterior) has
an average depth of about
The Posterior Median Sulcus (sulcus medianus posterior) is
very shallow; from it a septum of neuroglia reaches rather more than half-way
into the substance of the medulla spinalis; this septum varies in depth from 4
to
On either side of the posterior median sulcus, and at a short distance
from it, the posterior nerve roots are attached along a vertical furrow named
the posterolateral sulcus. The portion of the medulla spinalis which
lies between this and the posterior median sulcus is named the posterior
funiculus. In the cervical and upper thoracic regions this funiculus
presents a longitudinal furrow, the postero-intermediate sulcus; this
marks the position of a septum which extends into the posterior funiculus and
subdivides it into two fasciculi—a medial, named the fasciculus gracilis
(tract of Goll); and a lateral, the fasciculus cuneatus (tract
of Burdach). The portion of the medulla spinalis
which lies in front of the posterolateral sulcus is termed the antero-lateral
region. The anterior nerve roots, unlike the posterior, are not attached in
linear series, and their position of exit is not marked by a sulcus. They arise
by separate bundles which spring from the anterior column of gray substance
and, passing forward through the white substance, emerge over an area of some
slight width. The most lateral of these bundles is generally taken as a
dividing line which separates the antero-lateral region into two parts, viz.,
an anterior funiculus, between the anterior median fissure and the most
lateral of the anterior nerve roots; and a lateral funiculus, between
the exit of these roots and the postero-lateral sulcus. In the upper part of
the cervical region a series of nerve roots passes outward through the lateral
funiculus of the medulla spinalis; these unite to form the spinal portion of
the accessory nerve, which runs upward and enters the cranial cavity through
the foramen magnum.
Gray Substance (substantia grisea centralis).—The
gray substance consists of two symmetrical portions, one in each half of the
medulla spinalis: these are joined across the middle line by a transverse
commissure of gray substance, through which runs a minute canal, the central
canal, just visible to the naked eye. In a transverse section each half of
the gray substance is shaped like a comma or crescent, the concavity of which
is directed laterally; and these, together with the intervening gray
commissure, present the appearance of the letter H. An imaginary coronal plane
through the central canal serves to divide each crescent into an anterior
or ventral, and a posterior or dorsal column.
The Anterior Column (columna anterior; anterior cornu),
directed forward, is broad and of a rounded or quadrangular shape. Its
posterior part is termed the base, and its anterior part the head, but
these are not differentiated from each other by any well-defined constriction.
It is separated from the surface of the medulla spinalis by a layer of white
substance which is traversed by the bundles of the anterior nerve roots. In the
thoracic region, the postero-lateral part of the anterior column projects
lateralward as a triangular field, which is named the lateral column (columna
lateralis; lateral cornu).
The Posterior Column (columna posterior; posterior cornu)
is long and slender, and is directed backward and lateralward: it reaches
almost as far as the posterolateral sulcus, from which it is separated by a
thin layer of white substance, the tract of Lissauer. It consists of a base,
directly continuous with the base of the anterior horn, and a neck or
slightly constricted portion, which is succeeded by an oval or fusiform area,
termed the head, of which the apex approaches the posterolateral
sulcus. The apex is capped by a V-shaped or crescentic mass of translucent,
gelatinous neuroglia, termed the substantia gelatinosa of Rolando, which
contains both neuroglia cells, and small nerve cells. Between the anterior and
posterior columns the gray substance extends as a series of processes into the
lateral funiculus, to form a net-work called the formatio reticularis.