The motor impulses for voluntary movement are mainly generated in the precentral gyrusof the

frontal lobe (primary motor cortex, Brodmann area 4) and in the adjacent cortical areas (first
motor neuron). They travel in the long fiber pathways (mainly the corticonuclear and corticospinal
tracts/pyramidal pathway), passing through thebrainstemand down thespinal cord to the anterior
horn, where they make synaptic contact with the second motor neuron—usually by way of one or
more intervening interneurons.

The nerve fibers emerging from area 4 and the adjacent cortical areas together make up
thepyramidal tract, which is the quickest and most direct connection between the primary motor
area and the motor neurons of the anterior horn. In addition, other cortical areas (especially the
premotor cortex, area 6) and subcortical nuclei (especially the basal ganglia, cf. p. 330) participate
in the neural control of movement. These areas form complex feedback loops with one another
and with the primary motor cortex and cerebellum; they exert an influence on the anterior horn
cells by way of several distinct fiber pathways in the spinal cord. Their function is mainly to
modulate movement and to regulate muscle tone.

Impulses generated in the second motor neurons of the motor cranial nerve nuclei and the
anterior horn of the spinal cord pass through theanterior roots, the nerve plexuses(in the cervical
and lumbosacral regions), and theperipheral nerveson their way to the skeletal muscles. The
impulses are conveyed to the muscle cells through themotor end platesof the neuromuscular
junction.

Lesions of the first motor neuron in the brain or spinal cord usually producespastic paresis, while
lesions of the second motor neuron in the anterior horn, anterior root, peripheral nerve, or motor
end plate usually produceflaccid paresis. Motor deficits rarely appear in isolation as the result of a
lesion of the nervous system; they are usually accompanied by sensory, autonomic, cognitive,
and/or neuropsychological deficits of various kinds, depending on the site and nature of the
causative lesion.

Central Components of the MotorSystemand Clinical Syndromes of Lesions Affecting Them

The central portion of the motor system for voluntary movement consists of theprimary motor
cortex (area 4)and theadjacent cortical areas(particularly the premotor cortex, area 6), and
thecorticobulbarandcorticospinal tractsto which these cortical areas give rise (Figs. 3.1and 3.2).

Motor Cortical Areas

Theprimary motor cortex(precentral gyrus, Fig. 3.1) is a band of cortical tissue that lies on the
opposite side of the central sulcus from the primary somatosensory cortex (in the postcentral
gyrus) and, like it, extends upward and past the superomedial edge of the hemisphere onto its
medial surface. The área representing the throat and larynx lies at the inferior end of the primary
motor cortex; above it, in sequence, are the areas representing the face, upper limbs, trunk, and
lower limbs (Fig. 3.2). This is the inverted “motor homunculus,” corresponding to the
“somatosensory homunculus” of the postcentral gyrus that was discussed in Chapter 2
(see Fig. 9.19, p. 374).

Motor neuronsare found not only in area 4 but also in the adjacent cortical areas. The fibers
mediating fine voluntary movements, however, originate mainly in the precentral gyrus. This is the
site of the characteristic, largepyramidal neurons (Betz cells), which lie in the fifth cellular layer of
the cortex and send their rapidly conducting, thickly myelinated axons (Fig. 3.3) into the pyramidal
tract. The pyramidal tract was once thought to be entirely composed of.
Betz cell axons, but it is now known that these account for only 3.4-4 % of its fibers. The largest
fiber contingent in fact originates from the smaller pyramidal and fusiform cells of Brodmann
areas 4 and 6. Axons derived from área 4 make up about 40 % of all pyramidal tract fibers; the
remainder come from other frontal areas, from areas 3, 2, and 1 of the parietal somatosensory
cortex (sensorimotor area), and from other areas in the parietal lobe (Fig. 3.1). The motor neurons
of area 4 subserve fine, voluntary movement of the contralateral half of the body; the pyramidal
tract is, accordingly, crossed (see Fig. 3.4). Direct electrical stimulation of area 4, as during a
neurosurgical procedure, generally induces contraction of an individual muscle, while stimulation
of area 6 induces more complex and extensive movements, e. g., of an entire upper or lower limb

Corticospinal Tract (Pyramidal Tract)

This tract originates in themotor cortexand travels through thecerebral white matter(corona
radiata), the posterior limb of theinternal capsule (where the fibers lie very close together), the
central portion of thecerebral peduncle(cruz cerebri), thepons, and the base (i.e., the anterior
portion) of themedulla, where the tract is externally evident as a slight protrusion called the
pyramid. The medullary pyramids (there is one on either side) give the tract its name. At the lower
end of the medulla, 80-85 % of the pyramidal fibers cross to the other side in the so –called
decussation of the pyramids. The fibers that do not cross here descend the spinal cord in the
ipsilateral anterior funiculus as theanterior corticospinal tract; they cross farther down (usually at
the level of the segment that they supply) through the anterior commissure of the spinal cord (cf.
Fig. 3.6). At cervical and thoracic levels, there are probably also a few fibers that remain uncrossed
and innervate ipsilateral motor neurons in the anterior horn, so that the nuchal and truncal
musculature receives a bilateral cortical innervation.

The majority of pyramidal tract fibers cross in the decussation of the pyramids, then descend the
spinal cord in the contralateral lateral funiculus as the lateral corticospinal tract.This tract shrinks
in cross-sectional area as it travels down the cord, because some of its fibers terminate in each
segment along the way. About 90 % of all pyramidal tract fibers end in synapses onto
interneurons, which then transmit the motor impulses onward to the largeαmotor neurons of the
anterior horn, as well as to the smallerγmotor neurons (Fig. 3.4).

Corticonuclear (Corticobulbar) Tract

Some of the fibers of the pyramidal tract branch off from the main mass of the tract as it passes
through the midbrain and then take a more dorsal course toward the motor cranial nerve nuclei
(Figs. 3.4and 4.54, p. 212). The fibers supplying these brainstem nuclei are partly crossed and
partly uncrossed (for further details, cf. Chapter 4, section 4.4 “Cranial Nerves”). The nuclei
receiving pyramidal tract input are the ones that mediate voluntary movements of the cranial
musculature through cranial nerves V (the trigeminal nerve), VII (the facial nerve), IX, X, and XI (the
glossopharyngeal, vagus, and accessory nerves), and XII (the hypoglossal nerve).

Corticomesencephalic tract.There is also a contingent of fibers traveling together with the

corticonuclear tract that arises, not in areas 4 and 6, but rather in area 8, the frontal eye field (Figs.
3.1and 3.4). The impulses in these fibers mediate conjugate eye movements (p. 46), which are a
complex motor process

Because of its special origin and function, the pathway originating in the frontal eye fields has a
separate name (the corticomesencephalic tract), though most authors consider it a part of the
corticonuclear tract.

The corticomesencephalic tract runs in tandem with the pyramidal tract (just rostral to it, in the
posterior limb of the internal capsule) and then heads dorsally toward the nuclei of the cranial
nerves that mediate eye movements, i.e., cranial nerves III, IV, and VI (the oculomotor, trochlear,
and abducens nerves). Area 8 innervates the eye muscles exclusively in synergistic fashion, rather
than individually. Stimulation of area 8 induces conjugate gaze deviation to the opposite side. The
fibers of the corticomesencephalic tract do not terminate directly onto the motor neurons of
cranial nerve nuclei III, IV, and VI; the anatomical situation here is complicated and incompletely
understood, and is discussed further in Chapter 4 (p. 46 ff).

Other Central Components of the Motor System

A number of central pathways beside the pyramidal tract play major roles in the control of motor
function (Fig. 3.5). One important group of fibers (thecorticopontocerebellar tract) conveys
information from the cerebral cortex to the cerebellum, whose output in turn modulates planned
movements (cf. Chapter 5 “Cerebellum”). Other fibers travel from the cortex to thebasal ganglia
(mainly the corpus striatum = caudate nucleus and putamen), thesubstantia nigra, the
brainstemreticular formation, and other nuclei (e. g., in the midbrain tectum). In each of these
structures, the impulses are processed and conveyed onward, via interneurons, to efferent tracts
that project to the motor neurons of the anterior horn—the tectospinal, rubrospinal,
reticulospinal, vestibulospinal, and other tracts (Fig. 3.6). These tracts enable the cerebellum, basal
ganglia, and brainstem motor nuclei to influence motor function in the spinal cord. (For further
details, see Chapter 4 “Brainstem,” and Chapter 8 “Basal Ganglia.”)

Lateral and medial motor tracts in the spinal cord.The motor tracts in the spinal cord are
anatomically and functionally segregated into two groups: alateral group, comprising the
corticospinal and rubrospinal tracts, and a medial group, comprising the reticulospinal,
vestibulospinal, and tectospinal tracts (Kuypers, 1985). The lateral tracts mainly project to the
distal musculature (especially in the upper limbs) and also make short propriospinal connections.
They are primarily responsible for voluntary movements of the forearms and hands, i.e., for
precise, highly differentiated, fine motor control. The medial tracts, in contrast, innervate motor
neurons lying more medially in the anterior horn and make relatively long propriospinal
connections. They are primarily responsible for movements of the trunk and lower limbs (stance
and gait)
67

Peripheral Components of the Motor System and Clinical Syndromes of Lesions Affecting Them

The peripheral portion of the motor system comprises the motor cranial nerve nuclei of the
brainstem, the motor anterior horn cells of the spinal cord, the anterior roots, the cervical and
lumbosacral nerve plexuses, the peripheral nerves, and the motor end plates in skeletal muscle.

Anterior horn cells (αandγmotor neurons).The fibers not only of the piramidal tract but also of the
nonpyramidal descending pathways (the reticulospinal, tectospinal, vestibulospinal, and
rubrospinal tracts, among others), as well as afferent fibers from the posterior roots, terminate on
the cell bodies or dendrites of the larger and smallerαmotor neurons. Fibers of all of these types
also make synaptic contact with the smallγmotor neurons, partly directly, and partly through
intervening interneurons and the association and commissural neurons of the intrinsic neuronal
apparatus of the spinal cord (Fig. 3.6). Some of these synapses are excitatory, others inhibitory.
The thin, unmyelinated

neurites of theγmotor neurons innervate the intrafusal muscle fibers. In contrast to the
pseudounipolar neurons of the spinal ganglia, the anterior horn cells are multipolar. Their
dendrites receive synaptic contact from a wide variety of afferent and efferent systems (Fig. 3.6).

The functional groups and nuclear columns of neurons in the anterior horn are not separated from
one another by anatomically discernible borders (cf. Fig. 2.5b, p. 25). In the cervical spinal cord,
the motor neurons for the upper limbs lie in the lateral portion of the gray matter of the anterior
horn; those for the truncal muscles lie in its medial portion. The same somatotopic principle
applies in the lumbar spinal cord, where the lower limbs are represented laterally, the trunk
medially.

Inhibition of anterior horn cells by Renshaw cells. Among the various types of interneurons of the
anterior horn, the Renshaw cells deserve special mention (Fig. 2.11, p. 34). These small cells
receive synaptic contact from colateral axons of the largeαmotor neurons. Their axons then
project back onto the anterior horn cells and inhibit their activity. Renshaw inhibition is an
example of a spinal negative feedback loop that stabilizes the activity of motor neurons.

Anterior roots.The neurites of the motor neurons exit the anterior aspect of the spinal cord as
rootlets (fila radicularia) and join together, forming the anterior roots. Each anterior root joins the
corresponding posterior root just distal to the dorsal root ganglion to form a spinal nerve, which
then exits the spinal canal through the intervertebral foramen.

Peripheral nerve and motor end plate.There is one pair of spinal nerves for each segment of the
body. The spinal nerves contain afferent somatosensory fibers, efferent somatic motor fibers,
efferent autonomic fibers from the latera horns of the spinal gray matter, and afferent autonomic
fibers (cf. p. 22). At cervical and lumbosacral levels, the spinal nerves join to form the nerve
plexuses, which, in turn, give rise to the peripheral nerves that innervate the musculature of the
neck and limbs (Figs. 3.31,3.32, and 3.34).
The thick, myelinated, rapidly conducting neurites of the largeαmotor neurons are calledα1 fibers
(Fig. 2.11, p. 34). These fibers travel to the working musculature, where they divide into a highly
variable number of branches that terminate on muscle fibers. Synaptic impulse transmission
occurs at the neuromuscular junctions (motor end plates).

Motor unit. An anterior horn cell, its neurites, and the muscle fibers it innervates are collectively
termed a motor unit (Sherrington). Each motor unit constitutes the final common pathway for
movement-related impulses arriving at the anterior horn cell from higher levels: its activity is
influenced by impulses in a wide variety of motor tracts that originate in different areas of the
brain, as well as by impulses derived from intrasegmental and intersegmental réflex neurons of
the spinal cord. All of these movement-related impulses are integrated in the motor unit, and the
result of this integration is transmitted to the muscle fibers.

Muscles participating in finely differentiated movements are supplied by a large number of

anterior horn cells, each of which innervates only a few (5-20) muscle fibers; such muscles are thus
composed ofsmall motor units. In contrast, large muscles that contract in relatively
undifferentiated fashion, such as the gluteal muscles, are supplied by relatively few anterior horn
cells, each of which innervates 100-500 muscle fibers (large motor units).