The neural tube caudal to the fourth pair of somites develops into the spinal

cord (Figs. 17-4 and 17-5). The lateral walls of the neural tube thicken,
gradually reducing the size of the neural canal until only a minute central
canal of the spinal cord is present at 9 to 10 weeks (see Fig. 17-5C). Initially,
the wall of the neural tube is composed of a thick, pseudostratified, columnar
neuroepithelium (see Fig. 17-5D). These neuroepithelial cells constitute the
ventricular zone (ependymal layer), which gives rise to all neurons and
macroglial cells (macroglia) in the spinal cord (Figs. 17-5E and 17-6).
Macroglial cells are the larger members of the neuroglial family of cells, which
includes astrocytes and oligodendrocytes. Soon a marginal zone composed of
the outer parts of the neuroepithelial cells becomes recognizable (see Fig. 175E). This zone gradually becomes the white matter (substance) of the spinal
cord as axons grow into it from nerve cell bodies in the spinal cord, spinal
ganglia, and brain. Some dividing neuroepithelial cells in the ventricular zone
differentiate into primordial neurons-neuroblasts. These embryonic cells form
an intermediate zone (mantle layer) between the ventricular and marginal
zones. Neuroblasts become neurons as they develop cytoplasmic processes (see
Fig. 17-6).
Microglial cells (microglia), which are scattered throughout
the gray and white matter, are small cells that are derived from
mesenchymal cells (see Fig. 17-6). Microglial cells invade the
CNS rather late in the fetal period after it has been penetrated
by blood vessels. Microglia originate in the bone marrow and
are part of the mononuclear phagocytic cell population.
The primordial supporting cells of the central nervous system-glioblasts
(spongioblasts)-differentiate from neuroepithelial cells, mainly after neuroblast
formation has ceased. The glioblasts migrate from the ventricular zone into the
intermediate and marginal zones. Some glioblasts become astroblasts and later
astrocytes, whereas others become oligodendroblasts and eventually
oligodendrocytes (see Fig. 17-6). When the neuroepithelial cells cease
producing neuroblasts and glioblasts, they differentiate into ependymal cells,
which form the ependyma (ependymal epithelium) lining the central canal of
the spinal cord. Sonic hedgehog signaling controls the proliferation, survival,
and patterning of neuroepithelial progenitor cells by regulating Gli transcription
factors (see Fig. 17-2).
Proliferation and differentiation of neuroepithelial cells in the developing spinal
cord produce thick walls and thin roof- and floor-plates (see Fig. 17-5B).
Differential thickening of the lateral walls of the spinal cord soon produces a
shallow longitudinal groove on each side-the sulcus limitans (Figs. 17-5B and
17-7). This groove separates the dorsal part, the alar plate (lamina), from the

ventral part, the basal plate (lamina). The alar and basal plates produce
longitudinal bulges extending through most of the length of the developing
spinal cord. This regional separation is of fundamental importance because the
alar and basal plates are later associated with afferent and efferent functions,
respectively.
Cell bodies in the alar plates form the dorsal gray columns that extend the
length of the spinal cord. In transverse sections of the cord, these columns are
the dorsal gray horns. Neurons in these columns constitute afferent nuclei, and
groups of these nuclei form the dorsal gray columns. As the alar plates enlarge,
the dorsal median septum forms. Cell bodies in the basal plates form the ventral
and lateral gray columns. In transverse sections of the spinal cord, these
columns are the ventral gray horns and lateral gray horns, respectively (see
Fig. 5-7C). Axons of ventral horn cells grow out of the spinal cord and form the
ventral roots of the spinal nerves. As the basal plates enlarge, they bulge
ventrally on each side of the median plane. As this occurs, the ventral median
septum forms, and a deep longitudinal groove-the ventral median fissuredevelops on the ventral surface of the spinal cord.

Positional Changes of the Spinal Cord

The spinal cord in the embryo extends the entire length of the
vertebral canal (see Fig. 17-10A). The spinal nerves pass through the
intervertebral foramina opposite their levels of origin. Because the
vertebral column and dura mater grow more rapidly than the spinal
cord, this positional relationship to the spinal nerves does not persist.
The caudal end of the spinal cord gradually comes to lie at relatively
higher levels. In a 6-month-old fetus, it lies at the level of the first
sacral vertebra (see Fig. 17-10B). The spinal cord in the newborn
terminates at the level of the second or third lumbar vertebra (see Fig.
17-10C). The spinal cord in the adult usually terminates at the
inferior border of the first lumbar vertebra (see Fig. 17-10D). This is
an average level because the caudal end of the spinal cord may be as
superior as the 12th thoracic vertebra or as inferior as the third lumbar
vertebra. As a result, the spinal nerve roots, especially those of the
lumbar and sacral segments, run obliquely from the spinal cord to the
corresponding level of the vertebral column. The nerve roots inferior
to the end of the cord-the medullary cone (Latin [L]. conus
medullaris)-form a bundle of spinal nerve roots, the cauda equina (L.
horse's tail). Although the dura mater and arachnoid mater usually end
at S2 vertebra in adults, the pia mater does not. Distal to the caudal
end of the spinal cord, the pia mater forms a long fibrous thread, the
terminal filum (L. filum terminale), which indicates the original level
of the caudal end of the embryonic spinal cord (see Fig. 17-10C). This
thread extends from the medullary cone and attaches to the
periosteum of the first coccygeal vertebra.
Myelination of Nerve Fibers
Myelin sheaths surrounding nerve fibers within the spinal cord begin
to form during the late fetal period and continue to form during the
first postnatal year. Myelin basic proteins, a family of related
polypeptide isoforms, are essential in myelination. In general, fiber
tracts become myelinated at approximately the time they become
functional. Motor roots are myelinated before sensory roots. The
myelin sheaths surrounding nerve fibers within the spinal cord are
formed by oligodendrocytes. The plasma membranes of these cells
wrap around the axon, forming a number of layers (Fig. 17-11F to H).

The myelin sheaths around the axons of peripheral nerve fibers are
formed by the plasma membranes of neurolemma cells (Schwann
cells), which are analogous to oligodendrocytes. These neuroglial
cells are derived from neural crest cells that migrate peripherally and
wrap themselves around the axons of somatic motor neurons and
preganglionic autonomic motor neurons as they pass out of the central
nervous system (see Figs. 17-8 and 17-11A to E). These cells also
wrap themselves around both the central and peripheral processes of
somatic and visceral sensory neurons, as well as around the axons of
postsynaptic autonomic motor neurons. Beginning at approximately
20 weeks, peripheral nerve fibers have a whitish appearance, resulting
from the deposition of myelin.

The developing human keith l moore