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A major source of confusion about the pyramidal tract stems from the traditional

view, formulated at the turn of the 20th century, that it originates entirely from the
large motor cells of Betz in the fifth layer of the precentral convolution (the primary
motor cortex, or area 4 of Brodmann) (Figs. 3-3 and 22-1). However, there are only
some 25,000 to 35,000 Betz cells, whereas the medullary pyramid contains about 1
million axons (Lassek). Thus the pyramidal tract contains many fibers that arise from
cortical neurons other than Betz cells, particularly in Brodmann areas 4 and 6 (the
frontal cortex immediately rostral to area 4, including the medial portion of the
superior frontal gyrus, i.e., the supplementary motor area); in the primary
somatosensory cortex (Brodmann areas 3, 1, and 2); and in the superior parietal lobule
(areas 5 and 7). Data concerning the origin of the pyramidal tract in humans are
scanty, but in the monkey, by counting the pyramidal axons that remained after
cortical excisions and long survival periods, Russell and DeMyer found that 40
percent of the descending axons arose in the parietal lobe, 31 percent in motor area 4,
and the remaining 29 percent in premotor area 6. Studies of retrograde transport of
tracer substance in the monkey have confirmed these findings.

Fibers from the motor and premotor cortices (Brodmann areas 4 and 6, Fig. 22-1),
supplementary motor cortex, and portions of parietal cortex (areas 1, 3, 5, and 7)
converge in the corona radiata and descend through the posterior limb of the internal
capsule, basis pedunculi, basis pontis, and medulla. As the corticospinal tracts
descend in the cerebrum and brainstem, they send collaterals to the striatum,
thalamus, red nucleus, cerebellum, and reticular formations. Accompanying the
corticospinal tracts in the brainstem are the corticobulbar tracts, which are distributed
to motor nuclei of the cranial nerves ipsilaterally and contralaterally (Fig. 3-2). It has
been possible to trace the direct projection of axons of cortical neurons to the
trigeminal, facial, ambiguus, and hypoglossal nuclei (Iwatsubo et al). No axons were
seen to terminate directly in the oculomotor, trochlear, abducens, or vagal nuclei.
Insofar as the corticobulbar and corticospinal fibers have a similar origin and the
motor nuclei of the brainstem are the homologues of the motor neurons of the spinal
cord, the term upper motor neurons may suitably be applied to both these systems of

The corticospinal tracts decussate at the lower end of the medulla, although some of
their fibers may cross above this level. The proportion of crossed and uncrossed fibers
varies to some extent from one person to another. Most textbooks state that 75 to 80
percent of the fibers cross and that the remaining fibers descend ipsilaterally, mostly
in the uncrossed ventral corticospinal tract. In exceptional cases, these tracts cross
completely; equally rarely, they remain uncrossed. These variations are probably of
functional significance in determining the amount of neurologic deficit that results
from a unilateral lesion such as capsular infarction. A few well-studied cases are
found, such as the one described by Terakawa and colleagues, of acute stroke of the
cerebral hemisphere causing hemiplegia on the same side. Also, Yakovlev found 3
instances of completely uncrossed pyramids among 130 autopsies of mentally
retarded neonates but considering the maldevelopment of these brains, the finding is
not surprising (personal communication).

Beyond their decussation, the corticospinal pathways descend as well-defined bundles
in the anterior and posterolateral columns of white matter (funiculi) of the spinal cord
(Fig. 3-2). The course of the noncorticospinal motor pathways (vestibulospinal,
reticulospinal, and descending propriospinal) have been traced in humans by Nathan
and his colleagues. The lateral vestibulospinal tract lies at the periphery of the cord,
where it occupies the most anterolateral portion of the anterior funiculus. The medial
vestibulospinal fibers mingle with those of the medial longitudinal
fasciculus. Reticulospinal fibers are less compact; they descend bilaterally, and most
of them come to lie just anterior to the lateral corticospinal tract. The descending
propriospinal pathway consists of a series of short fibers (one or two segments long)
lying next to the gray matter.

The somatotopic organization of the corticospinal system is of importance in clinical
work, especially in relation to certain stroke syndromes. As the descending axons
subserving limb and facial movements emerge from the cortical motor strip, they
maintain the anatomic specificity of the overlying cortex; therefore a discrete
corticalsubcortical lesion will result in a restricted weakness of the hand and arm or
the foot and leg. More caudally, the descending motor fibers converge and are
collected in the posterior limb of the internal capsule, so that even a small lesion there
may cause a pure motor hemiplegia, in which the face, arm, hand, leg, and foot are
affected to more or less the same degree (Lacunar syndromes). The axons subserving
facial movement are situated rostrally in the posterior limb of the capsule, those for
hand and arm in the central portion, and those for the foot and leg caudally (see

This topographic distribution is more or less maintained in the cerebral peduncle,
where the corticospinal fibers occupy approximately the middle of the peduncle, the
fibers destined to innervate the facial nuclei lying most medially. More caudally, in
the basis pontis (base, or ventral part of the pons), the descending motor tracts
separate into bundles that are interspersed with masses of pontocerebellar neurons and
their cerebellipetal fibers. A degree of somatotopic organization can be recognized
here as well, exemplified by selective weakness of the face and hand with dysarthria,
or of the leg, which may occur with pontine lacunar infarctions. Anatomic studies in
nonhuman primates indicate that armleg distribution of fibers in the rostral pons is
much the same as in the cerebral peduncle; in the caudal pons this distinction is less-
well defined. In humans, a lack of systematic anatomic study leaves the precise
somatotopic organization of corticospinal fibers in the pons less certain. However, a
study conducted by Marx and colleagues using sophisticated MRI mapping
techniques of patients with hemiplegia from brainstem lesions suggests that the usual
somatotopic organization breaks down in the base of the pons, and there is, instead, a
concentration of fibers innervating proximal muscles lying more dorsally and those
exciting distal parts of the limbs, more ventrally.

Another point of uncertainty has been the existence and course of fibers that descend
through the lower pons and upper medulla and then ascend again to innervate the
facial motor nucleus on the opposite side. Such a connection must exist to explain
occasional instances of facial palsy from brainstem lesions caudal to the midpons. A
discussion of the various hypothesized sites of this pathway, including a recurrent
tract (Pick bundle), can be found in the report by Terao and colleagues. They
conclude from imaging studies that corticobulbar fibers destined for the facial nucleus
descend in the ventromedial pons to the level of the upper medulla, where they
decussate and then ascend again; but there is considerable variability between
individuals in this configuration.

The descending pontine bundles, now devoid of their corticopontine fibers, reunite to
form the medullary pyramid.The brachialcrural pattern may persist in the pyramids
and is certainly reconstituted in the lateral columns of the spinal cord (Fig. 8-3), but it
should be emphasized that the topographic separation of motor fibers at cervical,
thoracic, lumbar, and sacral levels is not as discrete as usually shown in schematic
diagrams of the spinal cord.

The corticospinal tracts and other upper motor neurons terminate mainly in relation to
nerve cells in the intermediate zone of spinal gray matter (internuncial neurons), from
which motor impulses are then transmitted to the anterior horn cells. Only 10 to 20
percent of corticospinal fibers (presumably the thick, rapidly conducting axons
derived from Betz cells) establish direct synaptic connections with the large motor
neurons of the anterior horns.

The Motor, Premotor, and Supplementary Motor Cortex and the Cerebral
Control of Movement
The motor area of the cerebral cortex is defined physiologically as the region of
electrically excitable cortex from which isolated movements can be evoked by stimuli
of minimal intensity. The muscle groups of the contralateral face, arm, trunk, and leg
are represented in the primary motor cortex (area 4 in Fig. 3-3), those of the face
being in the most inferior part of the precentral gyrus on the lateral surface of the
cerebral hemisphere and those of the leg in the paracentral lobule on the medial
surface of the cerebral hemisphere. The parts of the body capable of the most delicate
movements have, in general, the largest cortical representation, as displayed in the
motor homunculus (little man, a term first suggested by Wilder Penfield) shown in
Fig. 3-4.

Area 6, the premotor area, is also electrically excitable but requires more intense
stimuli than area 4 to evoke movements. Stimulation of its caudal aspect (area 6a)
produces responses that are similar to those elicited from area 4. These responses are
probably effected by transmission of impulses from all of area 6a to area 4 (as they
cannot be obtained after ablation of area 4). Stimulation of the rostral premotor area
(area 6a) elicits more general movement patterns, predominantly of proximal limb
musculature. The latter movements are effected via pathways other than those derived
from area 4 (hence, extrapyramidal). Very strong stimuli elicit movements from a
wide area of premotor frontal and parietal cortex, and the same movements may be
obtained from several widely separated points. From this it may be assumed, as Ash
and Georgopoulus point out, that the premotor cortex includes several anatomically
distinct subregions with different afferent and efferent connections. In general, it may
be said that the motorpremotor cortex is capable of synthesizing agonist actions into
an infinite variety of finely graded, highly differentiated patterns.