chapter 55

Unit xI
Cortical and Brain Stem Control
of Motor Function

Most “voluntary” move- Motor Sensory

ments initiated by the cere- Primary
motor Somatic
bral cortex are achieved cortex
Supplementary area 1
when the cortex activates Somatic
area Legs
“patterns” of function stored association
in lower brain areas—the Feet area
cord, brain stem, basal gan- Trunk
glia, and cerebellum. These lower centers, in turn, send Arm
specific control signals to the muscles. 4 7
For a few types of movements, however, the cortex has Hand
5
almost a direct pathway to the anterior motor neurons of 6
Face

1
the cord, bypassing some motor centers on the way. This

3, 2,
is especially true for control of the fine dexterous move- Mouth
ments of the fingers and hands. This chapter and Chapter
56 explain the interplay among the different motor areas
of the brain and spinal cord to provide overall synthesis of
voluntary motor function.
Premotor
area

Motor Cortex and Corticospinal Tract Figure 55-1 Motor and somatosensory functional areas of the
cerebral cortex. The numbers 4, 5, 6, and 7 are Brodmann’s cortical
areas, as explained in Chapter 47.
Figure 55-1 shows the functional areas of the cerebral
cortex. Anterior to the central cortical sulcus, occupying
Figure 55-1 lists the approximate topographical repre-
approximately the posterior one third of the frontal lobes,
sentations of the different muscle areas of the body in the
is the motor cortex. Posterior to the central sulcus is the
primary motor cortex, beginning with the face and mouth
somatosensory cortex (an area discussed in detail in earlier
region near the sylvian fissure; the arm and hand area, in
chapters), which feeds the motor cortex many of the sig-
the midportion of the primary motor cortex; the trunk,
nals that initiate motor activities.
near the apex of the brain; and the leg and foot areas, in the
The motor cortex itself is divided into three subareas,
part of the primary motor cortex that dips into the longi-
each of which has its own topographical representation of
tudinal fissure. This topographical organization is demon-
muscle groups and specific motor functions: (1) the pri-
strated even more graphically in Figure 55-2, which shows
mary motor cortex, (2) the premotor area, and (3) the sup-
the degrees of representation of the different muscle areas
plementary motor area.
as mapped by Penfield and Rasmussen. This mapping was
done by electrically stimulating the different areas of the
Primary Motor Cortex motor cortex in human beings who were undergoing neu-
The primary motor cortex, shown in Figure 55-1, lies in rosurgical operations. Note that more than one half of the
the first convolution of the frontal lobes anterior to the entire primary motor cortex is concerned with control-
central sulcus. It begins laterally in the sylvian fissure, ling the muscles of the hands and the muscles of speech.
spreads superiorly to the uppermost portion of the brain, Point stimulation in these hand and speech motor areas on
and then dips deep into the longitudinal fissure. (This rare occasion causes contraction of a single muscle; most
area is the same as area 4 in Brodmann’s classification of often, stimulation contracts a group of muscles instead. To
the brain cortical areas, shown in Figure 47-5.) express this in another way, excitation of a single motor

667
Unit XI The Nervous System: C. Motor and Integrative Neurophysiology

cortex neuron usually excites a specific movement rather A special class of neurons called mirror neurons becomes
than one specific muscle. To do this, it excites a “pattern” active when a person performs a specific motor task or
of separate muscles, each of which contributes its own when he or she observes the same task performed by others.
direction and strength of muscle movement. Thus, the activity of these neurons “mirrors” the behavior
of another person as though the observer was performing
Premotor Area the specific motor task. Mirror neurons are located in the
The premotor area, also shown in Figure 55-1, lies 1 to 3 premotor cortex and the inferior parietal cortex (and per-
centimeters anterior to the primary motor cortex, extend- haps in other regions of the brain) and were first discovered
ing inferiorly into the sylvian fissure and superiorly into in monkeys. However, brain imaging studies indicate that
the longitudinal fissure, where it abuts the supplemen- these neurons are also present in humans and may serve
tary motor area, which has functions similar to those of the same functions as observed in monkeys—to transform
the premotor area. The topographical organization of the sensory representations of acts that are heard or seen into
premotor cortex is roughly the same as that of the primary motor representations of these acts. Many neurophysi-
motor cortex, with the mouth and face areas located most ologists believe that these mirror neurons may be impor-
laterally; as one moves upward, the hand, arm, trunk, and tant for understanding the actions of other people and for
leg areas are encountered. learning new skills by imitation. Thus, the premotor cortex,
Nerve signals generated in the premotor area cause basal ganglia, thalamus, and primary motor cortex consti-
much more complex “patterns” of movement than the tute a complex overall system for the control of complex
discrete patterns generated in the primary motor cortex. patterns of coordinated muscle activity.
For instance, the pattern may be to position the shoul-
ders and arms so that the hands are properly oriented Supplementary Motor Area
to perform specific tasks. To achieve these results, the The supplementary motor area has yet another topo-
most anterior part of the premotor area first develops a graphical organization for the control of motor function.
“motor image” of the total muscle movement that is to be It lies mainly in the longitudinal fissure but extends a few
performed. Then, in the posterior premotor cortex, this centimeters onto the superior frontal cortex. Contractions
image excites each successive pattern of muscle activity elicited by stimulating this area are often bilateral rather
required to achieve the image. This posterior part of the than unilateral. For instance, stimulation frequently leads
premotor cortex sends its signals either directly to the pri- to bilateral grasping movements of both hands simulta-
mary motor cortex to excite specific muscles or, often, by neously; these movements are perhaps rudiments of the
way of the basal ganglia and thalamus back to the primary hand functions required for climbing. In general, this
motor cortex. area functions in concert with the premotor area to pro-
vide body-wide attitudinal movements, fixation move-
ments of the different segments of the body, positional
movements of the head and eyes, and so forth, as back-
ground for the finer motor control of the arms and hands
by the premotor area and primary motor cortex.

Some Specialized Areas of Motor Control

Hip
Knee
Trunk
Shoulder
Elbow
Wrist

Found in the Human Motor Cortex

Hand

Mi Ring inger
ex fing r
Indddle finge
Th fing er

A few highly specialized motor regions of the human cere-

um er
le f
An oes

ck bral cortex (shown in Figure 55-3) control specific motor

Litt
kle

Ne ow
T

B r functions. These regions have been localized either by elec-

ll trical stimulation or by noting the loss of motor function
eba
a n d y
e
Eye
lid Face when destructive lesions occur in specific cortical areas.
Lips Some of the more important regions are the following.
Vocalization

Jaw Broca’s Area and Speech. Figure 55-3 shows a

Tongu premotor area labeled “word formation” lying immediately
Swa e
llowin
g anterior to the primary motor cortex and immediately
Mastication above the sylvian fissure. This region is called Broca’s
Salivation
area. Damage to it does not prevent a person from
vocalizing but makes it impossible for the person to speak
whole words rather than uncoordinated utterances or an
occasional simple word such as “no” or “yes.” A closely
Figure 55-2 Degree of representation of the different mus-
associated cortical area also causes appropriate respiratory
cles of the body in the motor cortex. (Redrawn from Penfield
W, Rasmussen T: The Cerebral Cortex of Man: A Clinical Study of function, so respiratory activation of the vocal cords can
Localization of Function. New York: Hafner, 1968.) occur simultaneously with the movements of the mouth

668
 Chapter 55 Cortical and Brain Stem Control of Motor Function

Supplemental
and premotor Primary Corticospinal (Pyramidal) Tract
areas motor
cortex The most important output pathway from the motor cor-
tex is the corticospinal tract, also called the pyramidal

Legps
s
t
Fee
i
tract, shown in Figure 55-4. The corticospinal tract origi-

rs H
Fin Arm nk
ge s
Tru

Unit xI
Hand skills
nates about 30 percent from the primary motor cortex,

ck
Head rotation

Ne
b
Lips Th
um 30 percent from the premotor and supplementary motor
Contralateral
eye movements
Vocalization
Jaw Choice areas, and 40 percent from the somatosensory areas pos-
of words
Tongue
Swallowing terior to the central sulcus.
Chewing
Eye After leaving the cortex, it passes through the posterior
fixation
limb of the internal capsule (between the caudate nucleus
and the putamen of the basal ganglia) and then down-
Word formation
(Broca’s area) ward through the brain stem, forming the pyramids of the
medulla. The majority of the pyramidal fibers then cross
in the lower medulla to the opposite side and descend into
Figure 55-3 Representation of the different muscles of the body the lateral corticospinal tracts of the cord, finally termi-
in the motor cortex and location of other cortical areas respon- nating principally on the interneurons in the intermediate
sible for specific types of motor movements. regions of the cord gray matter; a few terminate on sensory
relay neurons in the dorsal horn, and a very few terminate
directly on the anterior motor neurons that cause muscle
and tongue during speech. Thus, the premotor neuronal contraction.
activities related to speech are highly complex. A few of the fibers do not cross to the opposite side
in the medulla but pass ipsilaterally down the cord in the
ventral corticospinal tracts. Many, if not most, of these
“Voluntary” Eye Movement Field. In the premotor
fibers eventually cross to the opposite side of the cord
area immediately above Broca’s area is a locus for
either in the neck or in the upper thoracic region. These
controlling voluntary eye movements. Damage to this
fibers may be concerned with control of bilateral postural
area prevents a person from voluntarily moving the eyes
movements by the supplementary motor cortex.
toward different objects. Instead, the eyes tend to lock
The most impressive fibers in the pyramidal tract
involuntarily onto specific objects, an effect controlled
are a population of large myelinated fibers with a mean
by signals from the occipital visual cortex, as explained
diameter of 16 micrometers. These fibers originate
in Chapter 51. This frontal area also controls eyelid
from giant pyramidal cells, called Betz cells, that are
movements such as blinking.
found only in the primary motor cortex. The Betz cells
are about 60 micrometers in diameter, and their fibers
Head Rotation Area. Slightly higher in the motor transmit nerve impulses to the spinal cord at a veloc-
association area, electrical stimulation elicits head rotation. ity of about 70 m/sec, the most rapid rate of transmis-
This area is closely associated with the eye movement sion of any signals from the brain to the cord. There
field; it directs the head toward different objects. are about 34,000 of these large Betz cell fibers in each
corticospinal tract. The total number of fibers in each
corticospinal tract is more than 1 million, so these large
Area for Hand Skills. In the premotor area
fibers represent only 3 percent of the total. The other 97
immediately anterior to the primary motor cortex for
percent are mainly fibers smaller than 4 micrometers in
the hands and fingers is a region that is important for
diameter that conduct background tonic signals to the
“hand skills.” That is, when tumors or other lesions
motor areas of the cord.
cause destruction in this area, hand movements become
uncoordinated and nonpurposeful, a condition called Other Fiber Pathways from the Motor Cortex. The motor
motor apraxia. cortex gives rise to large numbers of additional, mainly small,
fibers that go to deep regions in the cerebrum and brain
stem, including the following:
Transmission of Signals from the Motor
Cortex to the Muscles 1. The axons from the giant Betz cells send short collater-
als back to the cortex itself. These collaterals are believed
Motor signals are transmitted directly from the cortex to to inhibit adjacent regions of the cortex when the Betz
the spinal cord through the corticospinal tract and indi- cells discharge, thereby “sharpening” the boundaries of
rectly through multiple accessory pathways that involve the excitatory signal.
the basal ganglia, cerebellum, and various nuclei of the 2. A large number of fibers pass from the motor cortex into
brain stem. In general, the direct pathways are concerned the caudate nucleus and putamen. From there, additional
more with discrete and detailed movements, especially of pathways extend into the brain stem and spinal cord, as
the distal segments of the limbs, particularly the hands discussed in the next chapter, mainly to control body pos-
and fingers. tural muscle contractions.

669
Unit XI The Nervous System: C. Motor and Integrative Neurophysiology

Incoming Sensory Fiber Pathways to the Motor Cortex

Motor cortex
The functions of the motor cortex are controlled mainly by
nerve signals from the somatosensory system but also, to
some degree, from other sensory systems such as hearing
and vision. Once the sensory information is received, the
motor cortex operates in association with the basal ganglia
and cerebellum to excite an appropriate course of motor
action. The more important incoming fiber pathways to the
Posterior limb of internal
capsule motor cortex are the following:
1. Subcortical fibers from adjacent regions of the cerebral
cortex, especially from (a) the somatosensory areas of the
parietal cortex, (b) the adjacent areas of the frontal cortex
Genu of corpus callosum anterior to the motor cortex, and (c) the visual and audi-
tory cortices.
2. Subcortical fibers that arrive through the corpus callo-
sum from the opposite cerebral hemisphere. These fibers
connect corresponding areas of the cortices in the two
Basis pedunculi of
mesencephalon sides of the brain.
3. Somatosensory fibers that arrive directly from the ven-
trobasal complex of the thalamus. These relay mainly
cutaneous tactile signals and joint and muscle signals
from the peripheral body.
Longitudinal fascicles
4. Tracts from the ventrolateral and ventroanterior nuclei of
of pons
the thalamus, which in turn receive signals from the cerebel-
lum and basal ganglia. These tracts provide signals that are
necessary for coordination among the motor control func-
tions of the motor cortex, basal ganglia, and cerebellum.
5. Fibers from the intralaminar nuclei of the thalamus.
These fibers control the general level of excitability of
Pyramid of medulla the motor cortex in the same way they control the gen-
oblongata eral level of excitability of most other regions of the cere-
bral cortex.
Lateral corticospinal tract
Ventral corticospinal tract Red Nucleus Serves as an Alternative Pathway
for Transmitting Cortical Signals to the Spinal
Figure 55-4 Corticospinal (pyramidal) tract. (Modified from Cord
Ranson SW, Clark SL: Anatomy of the Nervous System. Philadelphia:
WB Saunders, 1959.) The red nucleus, located in the mesencephalon, func-
tions in close association with the corticospinal tract.
As shown in Figure 55-5, it receives a large number of
direct fibers from the primary motor cortex through the
corticorubral tract, as well as branching fibers from the
3. A moderate number of motor fibers pass to red nuclei
of the midbrain. From these, additional fibers pass down corticospinal tract as it passes through the mesencepha-
the cord through the rubrospinal tract. lon. These fibers synapse in the lower portion of the red
nucleus, the magnocellular portion, which contains large
4. A moderate number of motor fibers deviate into the retic-
ular substance and vestibular nuclei of the brain stem; neurons similar in size to the Betz cells in the motor cor-
from there, signals go to the cord by way of reticulospinal tex. These large neurons then give rise to the rubrospinal
and vestibulospinal tracts, and others go to the cerebellum tract, which crosses to the opposite side in the lower brain
by way of reticulocerebellar and vestibulocerebellar tracts. stem and follows a course immediately adjacent and ante-
5. A tremendous number of motor fibers synapse in the rior to the corticospinal tract into the lateral columns of
pontile nuclei, which give rise to the pontocerebellar the spinal cord.
fibers, carrying signals into the cerebellar hemispheres. The rubrospinal fibers terminate mostly on the
6. Collaterals also terminate in the inferior olivary nuclei, interneurons of the intermediate areas of the cord gray
and from there, secondary olivocerebellar fibers transmit matter, along with the corticospinal fibers, but some of
signals to multiple areas of the cerebellum. the rubrospinal fibers terminate directly on anterior
Thus, the basal ganglia, brain stem, and cerebellum all motor neurons, along with some corticospinal fibers. The
receive strong motor signals from the corticospinal system red nucleus also has close connections with the cerebel-
every time a signal is transmitted down the spinal cord to lum, similar to the connections between the motor cortex
cause a motor activity. and the cerebellum.

670
 Chapter 55 Cortical and Brain Stem Control of Motor Function

Motor of the direct corticospinal-pyramidal system. These include

Corticorubral
cortex tract pathways through the basal ganglia, the reticular forma-
tion of the brain stem, the vestibular nuclei, and often the
red nuclei. This is such an all-inclusive and diverse group
of motor control areas that it is difficult to ascribe spe-

Unit xI
cific neurophysiologic functions to the so-called extrapy-
ramidal system as a whole. In fact, the pyramidal and
extrapyramidal systems are extensively interconnected and
interact to control movement. For these reasons, the term
“extrapyramidal” is being used less often both clinically and
physiologically.

Red nucleus
Excitation of the Spinal Cord Motor Control Areas
Interpositus
nucleus by the Primary Motor Cortex and Red Nucleus
Reticular formation Dentate Vertical Columnar Arrangement of the Neurons in
nucleus
Rubrospinal tract the Motor Cortex. In Chapters 47 and 51, we pointed
Cerebellum out that the cells in the somatosensory cortex and visual
cortex are organized in vertical columns of cells. In like
manner, the cells of the motor cortex are organized in
vertical columns a fraction of a millimeter in diameter,
Figure 55-5 Corticorubrospinal pathway for motor control, show-
ing also the relation of this pathway to the cerebellum. with thousands of neurons in each column.
Each column of cells functions as a unit, usually stim-
Function of the Corticorubrospinal System. The ulating a group of synergistic muscles, but sometimes
magnocellular portion of the red nucleus has a stimulating just a single muscle. Also, each column has
somatographic representation of all the muscles of the six distinct layers of cells, as is true throughout nearly
body, as is true of the motor cortex. Therefore, stimulation all the cerebral cortex. The pyramidal cells that give rise
of a single point in this portion of the red nucleus causes to the corticospinal fibers all lie in the fifth layer of cells
contraction of either a single muscle or a small group of from the cortical surface. Conversely, the input signals
muscles. However, the fineness of representation of the all enter by way of layers 2 through 4. And the sixth layer
different muscles is far less developed than in the motor gives rise mainly to fibers that communicate with other
cortex. This is especially true in human beings, who have regions of the cerebral cortex itself.
relatively small red nuclei. Function of Each Column of Neurons. The neurons
of each column operate as an integrative processing
The corticorubrospinal pathway serves as an accessory system, using information from multiple input sources
route for transmission of relatively discrete signals from to determine the output response from the column. In
the motor cortex to the spinal cord. When the corticospi- addition, each column can function as an amplifying
nal fibers are destroyed but the corticorubrospinal path- system to stimulate large numbers of pyramidal fibers
way is intact, discrete movements can still occur, except to the same muscle or to synergistic muscles simulta-
that the movements for fine control of the fingers and neously. This is important because stimulation of a sin-
hands are considerably impaired. Wrist movements are gle pyramidal cell can seldom excite a muscle. Usually,
still functional, which is not the case when the corticoru- 50 to 100 pyramidal cells need to be excited simultane-
brospinal pathway is also blocked. ously or in rapid succession to achieve definitive muscle
Therefore, the pathway through the red nucleus to the contraction.
spinal cord is associated with the corticospinal system. Dynamic and Static Signals Are Transmitted by the
Further, the rubrospinal tract lies in the lateral columns of Pyramidal Neurons. If a strong signal is sent to a muscle
the spinal cord, along with the corticospinal tract, and ter- to cause initial rapid contraction, then a much weaker
minates on the interneurons and motor neurons that con- continuing signal can maintain the contraction for long
trol the more distal muscles of the limbs. Therefore, the periods thereafter. This is the usual manner in which
corticospinal and rubrospinal tracts together are called excitation is provided to cause muscle contractions. To
the lateral motor system of the cord, in contradistinction do this, each column of cells excites two populations
to a vestibuloreticulospinal system, which lies mainly of pyramidal cell neurons, one called dynamic neurons
medially in the cord and is called the medial motor system and the other static neurons. The dynamic neurons are
of the cord, as discussed later in this chapter. excited at a high rate for a short period at the beginning
of a contraction, causing the initial rapid development
“Extrapyramidal” System of force. Then the static neurons fire at a much slower
The term extrapyramidal motor system is widely used in rate, but they continue firing at this slow rate to main-
clinical circles to denote all those portions of the brain and tain the force of contraction as long as the contraction is
brain stem that contribute to motor control but are not part required.

671
Unit XI The Nervous System: C. Motor and Integrative Neurophysiology

The neurons of the red nucleus have similar dynamic

Sensory neurons
and static characteristics, except that a greater percent-
age of dynamic neurons is in the red nucleus and a greater Propriospinal tract
percentage of static neurons is in the primary motor cor- Interneurons
tex. This may be related to the fact that the red nucleus
Corticospinal tract
is closely allied with the cerebellum, and the cerebellum from pyramidal cells
plays an important role in rapid initiation of muscle con- of cortex
traction, as explained in the next chapter. Rubrospinal tract
Reticulospinal tract
Somatosensory Feedback to the Motor Cortex Helps
Anterior motor
Control the Precision of Muscle Contraction neuron
When nerve signals from the motor cortex cause a mus- Motor nerve
cle to contract, somatosensory signals return all the way Tectospinal and
from the activated region of the body to the neurons in the reticulospinal tracts
motor cortex that are initiating the action. Most of these Vestibulospinal and
somatosensory signals arise in (1) the muscle spindles, reticulospinal tracts
(2) the tendon organs of the muscle tendons, or (3) the Figure 55-6 Convergence of different motor control pathways on
tactile receptors of the skin overlying the muscles. These the anterior motor neurons.
somatic signals often cause positive feedback enhance-
ment of the muscle contraction in the following ways: provide certain specific reflex patterns of movement in
In the case of the muscle spindles, if the fusimotor mus- response to sensory nerve stimulation. Many of these same
cle fibers in the spindles contract more than the large patterns are also important when the cord’s anterior motor
skeletal muscle fibers contract, the central portions of neurons are excited by signals from the brain. For example,
the spindles become stretched and, therefore, excited. the stretch reflex is functional at all times, helping to damp
Signals from these spindles then return rapidly to the any oscillations of the motor movements initiated from
pyramidal cells in the motor cortex to signal them that the brain, and probably also providing at least part of the
the large muscle fibers have not contracted enough. motive power required to cause muscle contractions when
The pyramidal cells further excite the muscle, helping the intrafusal fibers of the muscle spindles contract more
its contraction to catch up with the contraction of the than the large skeletal muscle fibers do, thus eliciting reflex
muscle spindles. In the case of the tactile receptors, if “servo-assist” stimulation of the muscle, in addition to the
the muscle contraction causes compression of the skin direct stimulation by the corticospinal fibers.
against an object, such as compression of the fingers Also, when a brain signal excites a muscle, it is usually
around an object being grasped, the signals from the unnecessary to transmit an inverse signal to relax the antag-
skin receptors can, if necessary, cause further excitation onist muscle at the same time; this is achieved by the recipro-
of the muscles and, therefore, increase the tightness of cal innervation circuit that is always present in the cord for
the hand grasp. coordinating the function of antagonistic pairs of muscles.
Finally, other cord reflex mechanisms, such as with-
Stimulation of the Spinal Motor Neurons drawal, stepping and walking, scratching, and postural
Figure 55-6 shows a cross section of a spinal cord mechanisms, can each be activated by “command” signals
segment demonstrating (1) multiple motor and senso- from the brain. Thus, simple command signals from the
rimotor control tracts entering the cord segment and (2) brain can initiate many normal motor activities, particularly
a representative anterior motor neuron in the middle of for such functions as walking and attaining different pos-
the anterior horn gray matter. The corticospinal tract tural attitudes of the body.
and the rubrospinal tract lie in the dorsal portions of the
lateral white columns. Their fibers terminate mainly on Effect of Lesions in the Motor Cortex or in the Corticospinal
interneurons in the intermediate area of the cord gray Pathway—The “Stroke”
matter. The motor control system can be damaged by the common
abnormality called a “stroke.” This is caused by either a rup-
In the cervical enlargement of the cord where the hands
tured blood vessel that hemorrhages into the brain or by
and fingers are represented, large numbers of both corti-
thrombosis of one of the major arteries supplying the brain.
cospinal and rubrospinal fibers also terminate directly on In either case, the result is loss of blood supply to the cor-
the anterior motor neurons, thus allowing a direct route tex or to the corticospinal tract where it passes through the
from the brain to activate muscle contraction. This is in internal capsule between the caudate nucleus and the puta-
keeping with the fact that the primary motor cortex has men. Also, experiments have been performed in animals to
an extremely high degree of representation for fine con- selectively remove different parts of the motor cortex.
trol of hand, finger, and thumb actions. Removal of the Primary Motor Cortex (Area
Patterns of Movement Elicited by Spinal Cord Pyramidalis). Removal of a portion of the primary motor
Centers. From Chapter 54, recall that the spinal cord can cortex—the area that contains the giant Betz pyramidal

672
 Chapter 55 Cortical and Brain Stem Control of Motor Function

c­ ells—causes varying degrees of paralysis of the represented Support of the Body Against Gravity—Roles
muscles. If the sublying caudate nucleus and adjacent premo- of the Reticular and Vestibular Nuclei
tor and supplementary motor areas are not damaged, gross
postural and limb “fixation” movements can still occur, but Figure 55-7 shows the locations of the reticular and ves-
there is loss of voluntary control of discrete movements of the tibular nuclei in the brain stem.

Unit xI
distal segments of the limbs, especially of the hands and fin-
gers. This does not mean that the hand and finger muscles Excitatory-Inhibitory Antagonism Between
themselves cannot contract; rather, the ability to control the Pontine and Medullary Reticular Nuclei
fine movements is gone. From these observations, one can con-
The reticular nuclei are divided into two major groups: (1)
clude that the area py­ramidalis is essential for voluntary initia-
tion of finely controlled movements, especially of the hands pontine reticular nuclei, located slightly posteriorly and
and fingers. laterally in the pons and extending into the mesenceph-
Muscle Spasticity Caused by Lesions That Damage alon, and (2) medullary reticular nuclei, which extend
Large Areas Adjacent to the Motor Cortex. The through the entire medulla, lying ventrally and medially
­ rimary motor cortex normally exerts a continual tonic
p near the midline. These two sets of nuclei function mainly
stimulatory effect on the motor neurons of the spinal cord; antagonistically to each other, with the pontine exciting
when this stimulatory effect is removed, hypotonia results. the antigravity muscles and the medullary relaxing these
Most lesions of the motor cortex, especially those caused same muscles.
by a stroke, involve not only the primary motor cortex but Pontine Reticular System. The pontine reticular
also adjacent parts of the brain such as the basal ganglia. In nuclei transmit excitatory signals downward into the cord
these instances, muscle spasm almost invariably occurs in through the pontine reticulospinal tract in the anterior
the afflicted muscle areas on the opposite side of the body
column of the cord, as shown in Figure 55-8. The fibers
(because the motor pathways cross to the opposite side).
of this pathway terminate on the medial anterior motor
This spasm results mainly from damage to accessory path-
ways from the nonpyramidal portions of the motor cortex. neurons that excite the axial muscles of the body, which
These pathways normally inhibit the vestibular and reticular support the body against gravity—that is, the muscles
brain stem motor nuclei. When these nuclei cease their state of the vertebral column and the extensor muscles of the
of inhibition (i.e., are “disinhibited”), they become spontane- limbs.
ously active and cause excessive spastic tone in the involved The pontine reticular nuclei have a high degree of
muscles, as we discuss more fully later in the chapter. This natural excitability. In addition, they receive strong exci­
is the spasticity that normally accompanies a “stroke” in a tatory signals from the vestibular nuclei, as well as from
human being. deep nuclei of the cerebellum. Therefore, when the pon-
tine reticular excitatory system is unopposed by the med-
Role of the Brain Stem in Controlling ullary reticular system, it causes powerful excitation of
Motor Function antigravity muscles throughout the body, so much so that

The brain stem consists of the medulla, pons, and mes-

encephalon. In one sense, it is an extension of the spinal
cord upward into the cranial cavity because it contains
motor and sensory nuclei that perform motor and sen-
sory functions for the face and head regions in the same
way that the spinal cord performs these functions from
the neck down. But in another sense, the brain stem is Pontine reticular
its own master because it provides many special control nuclei
functions, such as the following:
1. Control of respiration
2. Control of the cardiovascular system
Vestibular nuclei
3. Partial control of gastrointestinal function
4. Control of many stereotyped movements of the
body
Medullary reticular
5. Control of equilibrium nuclei
6. Control of eye movements
Finally, the brain stem serves as a way station for “com-
mand signals” from higher neural centers. In the following
sections, we discuss the role of the brain stem in control-
ling whole-body movement and equilibrium. Especially
important for these purposes are the brain stem’s reticu- Figure 55-7 Locations of the reticular and vestibular nuclei in the
lar nuclei and vestibular nuclei. brain stem.

673
Unit XI The Nervous System: C. Motor and Integrative Neurophysiology

to signals from the vestibular apparatus. We discuss this

more fully later in the chapter.
Medullary
reticulospinal The Decerebrate Animal Develops Spastic Rigidity
tract
When the brain stem of an animal is sectioned below the
midlevel of the mesencephalon, but the pontine and medul-
lary reticular systems, as well as the vestibular system, are
left intact, the animal develops a condition called decerebrate
Lateral vestibulospinal rigidity. This rigidity does not occur in all muscles of the
tract body but does occur in the antigravity muscles—the muscles
of the neck and trunk and the extensors of the legs.
Medial vestibulospinal tract Pontine reticulospinal tract
The cause of decerebrate rigidity is blockage of normally
Figure 55-8 Vestibulospinal and reticulospinal tracts descending strong input to the medullary reticular nuclei from the cere-
in the spinal cord to excite (solid lines) or inhibit (dashed lines) the
bral cortex, the red nuclei, and the basal ganglia. Lacking this
anterior motor neurons that control the body’s axial musculature.
input, the medullary reticular inhibitor system becomes non-
functional; full overactivity of the pontine excitatory system
four-legged animals can be placed in a standing position, occurs, and rigidity develops. We shall see later that other
supporting the body against gravity without any signals causes of rigidity occur in other neuromotor diseases, espe-
from higher levels of the brain. cially lesions of the basal ganglia.
Medullary Reticular System. The medullary reticu-
lar nuclei transmit inhibitory signals to the same antigrav-
ity anterior motor neurons by way of a different tract, the
Vestibular Sensations and Maintenance
medullary reticulospinal tract, located in the lateral column
of Equilibrium
of the cord, as also shown in Figure 55-8. The medullary
reticular nuclei receive strong input collaterals from (1) the
Vestibular Apparatus
corticospinal tract, (2) the rubrospinal tract, and (3) other
motor pathways. These normally activate the medullary The vestibular apparatus, shown in Figure 55-9, is the
reticular inhibitory system to counterbalance the excitatory sensory organ for detecting sensations of equilibrium. It is
signals from the pontine reticular system, so under normal encased in a system of bony tubes and chambers located in
conditions the body muscles are not abnormally tense. the petrous portion of the temporal bone, called the bony
Yet some signals from higher areas of the brain can labyrinth. Within this system are membranous tubes and
“disinhibit” the medullary system when the brain wishes chambers called the membranous labyrinth. The mem-
to excite the pontine system to cause standing. At other branous labyrinth is the functional part of the vestibular
times, excitation of the medullary reticular system can apparatus.
inhibit antigravity muscles in certain portions of the The top of Figure 55-9 shows the membranous laby-
body to allow those portions to perform special motor rinth. It is composed mainly of the cochlea (ductus cochle-
activities. The excitatory and inhibitory reticular nuclei aris); three semicircular canals; and two large chambers,
constitute a controllable system that is manipulated by the utricle and saccule. The cochlea is the major sensory
motor signals from the cerebral cortex and elsewhere organ for hearing (see Chapter 52) and has little to do
to provide necessary background muscle contractions with equilibrium. However, the semicircular canals, the
for standing against gravity and to inhibit appropriate utricle, and the saccule are all integral parts of the equilib-
groups of muscles as needed so that other functions can rium mechanism.
be performed.
“Maculae”—Sensory Organs of the Utricle and
Saccule for Detecting Orientation of the Head with
Role of the Vestibular Nuclei to Excite Respect to Gravity. Located on the inside surface of each
the Antigravity Muscles utricle and saccule, shown in the top diagram of Figure 55-9,
All the vestibular nuclei, shown in Figure 55-7, function is a small sensory area slightly over 2 millimeters in diameter
in association with the pontine reticular nuclei to control called a macula. The macula of the utricle lies mainly in the
the antigravity muscles. The vestibular nuclei transmit horizontal plane on the inferior surface of the utricle and
strong excitatory signals to the antigravity muscles by way plays an important role in determining orientation of the
of the lateral and medial vestibulospinal tracts in the ante- head when the head is upright. Conversely, the macula of
rior columns of the spinal cord, as shown in Figure 55-8. the saccule is located mainly in a vertical plane and signals
Without this support of the vestibular nuclei, the pontine head orientation when the person is lying down.
reticular system would lose much of its excitation of the Each macula is covered by a gelatinous layer in which
axial antigravity muscles. many small calcium carbonate crystals called statoconia
The specific role of the vestibular nuclei, however, is are embedded. Also in the macula are thousands of hair
to selectively control the excitatory signals to the different cells, one of which is shown in Figure 55-10; these project
antigravity muscles to maintain equilibrium in response cilia up into the gelatinous layer. The bases and sides of

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 Chapter 55 Cortical and Brain Stem Control of Motor Function

Anterior
Ampullae Kinocilium
Utricle
Maculae and Stereocilia
statoconia

Unit xI
Semi-
circular Filamentous
canals attachments

Saccule Ductus
Posterior
cochlearis
Crista ampullaris Ductus endolymphaticus
MEMBRANOUS LABYRINTH

Gelatinous Statoconia
mass of
cupula Gelatinous
Hair tufts layer
Hair tufts
Hair
cells Hair cells

Nerve fibers
Nerve
fibers
Sustentacular cells Sustentacular cells Nerve fiber
CRISTA AMPULLARIS AND MACULA
Figure 55-9 Membranous labyrinth and organization of the crista
ampullaris and the macula.

Figure 55-10 Hair cell of the equilibrium apparatus and its

synapses with the vestibular nerve.
the hair cells synapse with sensory endings of the vestibu-
lar nerve.
The calcified statoconia have a specific gravity two to of positive ions. Therefore, positive ions pour into the
three times the specific gravity of the surrounding fluid cell from the surrounding endolymphatic fluid, causing
and tissues. The weight of the statoconia bends the cilia receptor membrane depolarization. Conversely, bending
in the direction of gravitational pull. the pile of stereocilia in the opposite direction (backward
to the kinocilium) reduces the tension on the attach-
Directional Sensitivity of the Hair Cells— ments; this closes the ion channels, thus causing receptor
Kinocilium. Each hair cell has 50 to 70 small cilia called hyperpolarization.
stereocilia, plus one large cilium, the kinocilium, as shown Under normal resting conditions, the nerve fibers lead-
in Figure 55-10. The kinocilium is always located to one ing from the hair cells transmit continuous nerve impulses
side, and the stereocilia become progressively shorter at a rate of about 100 per second. When the stereocilia are
toward the other side of the cell. Minute filamentous bent toward the kinocilium, the impulse traffic increases,
attachments, almost invisible even to the electron often to several hundred per second; conversely, bending
microscope, connect the tip of each stereocilium to the the cilia away from the kinocilium decreases the impulse
next longer stereocilium and, finally, to the kinocilium. traffic, often turning it off completely. Therefore, as the
Because of these attachments, when the stereocilia orientation of the head in space changes and the weight
and kinocilium bend in the direction of the kinocilium, of the statoconia bends the cilia, appropriate signals are
the filamentous attachments tug in sequence on the ste- transmitted to the brain to control equilibrium.
reocilia, pulling them outward from the cell body. This In each macula, each of the hair cells is oriented in a
opens several hundred fluid channels in the neuronal different direction so that some of the hair cells are stimu-
cell membrane around the bases of the stereocilia, and lated when the head bends forward, some are stimulated
these channels are capable of conducting large numbers when it bends backward, others are stimulated when it

675
Unit XI The Nervous System: C. Motor and Integrative Neurophysiology

bends to one side, and so forth. Therefore, a different pat- Into the cupula are projected hundreds of cilia from
tern of excitation occurs in the macular nerve fibers for hair cells located on the ampullary crest. The kinocilia
each orientation of the head in the gravitational field. It is of these hair cells are all oriented in the same direction
this “pattern” that apprises the brain of the head’s orienta- in the cupula, and bending the cupula in that direc-
tion in space. tion causes depolarization of the hair cells, whereas
bending it in the opposite direction hyperpolarizes the
cells. Then, from the hair cells, appropriate signals are
Semicircular Ducts. The three semicircular ducts in sent by way of the vestibular nerve to apprise the cen-
each vestibular apparatus, known as the anterior, posterior, tral nervous system of a change in rotation of the head
and lateral (horizontal) semicircular ducts, are arranged at and the rate of change in each of the three planes of
right angles to one another so that they represent all three space.
planes in space. When the head is bent forward about 30
degrees, the lateral semicircular ducts are approximately
horizontal with respect to the surface of the earth; the Function of the Utricle and Saccule in the
anterior ducts are in vertical planes that project forward Maintenance of Static Equilibrium
and 45 degrees outward, whereas the posterior ducts are It is especially important that the hair cells are all oriented
in vertical planes that project backward and 45 degrees in different directions in the maculae of the utricles and
outward. saccules so that with different positions of the head, differ-
Each semicircular duct has an enlargement at one of ent hair cells become stimulated. The “patterns” of stimu-
its ends called the ampulla, and the ducts and ampulla lation of the different hair cells apprise the brain of the
are filled with a fluid called endolymph. Flow of this position of the head with respect to the pull of gravity. In
fluid through one of the ducts and through its ampulla turn, the vestibular, cerebellar, and reticular motor nerve
excites the sensory organ of the ampulla in the follow- systems of the brain excite appropriate postural muscles
ing manner: Figure 55-11 shows in each ampulla a small to maintain proper equilibrium.
crest called a crista ampullaris. On top of this crista is This utricle and saccule system functions extremely
a loose gelatinous tissue mass, the cupula. When a per- effectively for maintaining equilibrium when the head
son’s head begins to rotate in any direction, the inertia is in the near-vertical position. Indeed, a person can
of the fluid in one or more of the semicircular ducts determine as little as half a degree of dysequilib-
causes the fluid to remain stationary while the semicir- rium when the body leans from the precise upright
cular duct rotates with the head. This causes fluid to position.
flow from the duct and through the ampulla, bending
the cupula to one side, as demonstrated by the posi- Detection of Linear Acceleration by the Utricle
tion of the colored cupula in Figure 55-11. Rotation of and Saccule Maculae. When the body is suddenly
the head in the opposite direction causes the cupula to thrust forward—that is, when the body accelerates—
bend to the opposite side. the statoconia, which have greater mass inertia than the
surrounding fluid, fall backward on the hair cell cilia, and
information of dysequilibrium is sent into the nervous
centers, causing the person to feel as though he or she
were falling backward. This automatically causes the
person to lean forward until the resulting anterior shift
of the statoconia exactly equals the tendency for the
statoconia to fall backward because of the acceleration.
Cupula
At this point, the nervous system senses a state of proper
Cristae
equilibrium and leans the body forward no farther. Thus,
Ampulla ampullaris the maculae operate to maintain equilibrium during linear
acceleration in exactly the same manner as they operate
during static equilibrium.
The maculae do not operate for the detection of linear
velocity. When runners first begin to run, they must lean
far forward to keep from falling backward because of initial
acceleration, but once they have achieved running speed,
Hair cells if they were running in a vacuum, they would not have to
Nerve lean forward. When running in air, they lean forward to
maintain equilibrium only because of air resistance against
their bodies; in this instance, it is not the maculae that
make them lean but air pressure acting on pressure end-
Figure 55-11 Movement of the cupula and its embedded hairs at organs in the skin, which initiate appropriate equilibrium
the onset of rotation. adjustments to prevent falling.

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 Chapter 55 Cortical and Brain Stem Control of Motor Function

Detection of Head Rotation by the balance in the forward direction, in the side direction,
Semicircular Ducts or in the backward direction, one might ask: What is
the semicircular ducts’ function in the maintenance of
When the head suddenly begins to rotate in any direc-
equilibrium? All they detect is that the person’s head is
tion (called angular acceleration), the endolymph in the
beginning or stopping to rotate in one direction or another.

Unit xI
semicircular ducts, because of its inertia, tends to remain
Therefore, the function of the semicircular ducts is not
stationary while the semicircular ducts turn. This causes
to maintain static equilibrium or to maintain equilibrium
relative fluid flow in the ducts in the direction opposite to
during steady directional or rotational movements. Yet
head rotation.
loss of function of the semicircular ducts does cause a
Figure 55-12 shows a typical discharge signal from a
person to have poor equilibrium when attempting to
single hair cell in the crista ampullaris when an animal is
perform rapid, intricate changing body movements.
rotated for 40 seconds, demonstrating that (1) even when
The function of the semicircular ducts can be explained
the cupula is in its resting position, the hair cell emits a
by the following illustration: If a person is running forward
tonic discharge of about 100 impulses per second; (2)
rapidly and then suddenly begins to turn to one side, he or
when the animal begins to rotate, the hairs bend to one
she will fall off balance a fraction of a second later unless
side and the rate of discharge increases greatly; and (3)
appropriate corrections are made ahead of time. But the
with continued rotation, the excess discharge of the hair
maculae of the utricle and saccule cannot detect that he
cell gradually subsides back to the resting level during the
or she is off balance until after this has occurred. The
next few seconds.
semicircular ducts, however, will have already detected
The reason for this adaptation of the receptor is that
that the person is turning, and this information can easily
within the first few seconds of rotation, back resistance
apprise the central nervous system of the fact that the per-
to the flow of fluid in the semicircular duct and past the
son will fall off balance within the next fraction of a sec-
bent cupula causes the endolymph to begin rotating as
ond or so unless some anticipatory correction is made.
rapidly as the semicircular canal itself; then, in another
In other words, the semicircular duct mechanism pre-
5 to 20 seconds, the cupula slowly returns to its resting
dicts that dysequilibrium is going to occur and thereby
position in the middle of the ampulla because of its own
causes the equilibrium centers to make appropriate antic-
elastic recoil.
ipatory preventive adjustments. This helps the person
When the rotation suddenly stops, exactly opposite
maintain balance before the situation can be corrected.
effects take place: The endolymph continues to rotate
Removal of the flocculonodular lobes of the cerebel-
while the semicircular duct stops. This time, the cupula
lum prevents normal detection of semicircular duct sig-
bends in the opposite direction, causing the hair cell to
nals but has less effect on detecting macular signals. It is
stop discharging entirely. After another few seconds, the
especially interesting that the cerebellum serves as a “pre-
endolymph stops moving and the cupula gradually returns
dictive” organ for most rapid movements of the body, as
to its resting position, thus allowing hair cell discharge to
well as for those having to do with equilibrium. These
return to its normal tonic level, as shown to the right in
other functions of the cerebellum are discussed in the fol-
Figure 55-12. Thus, the semicircular duct transmits a sig-
lowing chapter.
nal of one polarity when the head begins to rotate and of
opposite polarity when it stops rotating. Vestibular Mechanisms for Stabilizing the Eyes
When a person changes his or her direction of movement
“Predictive” Function of the Semicircular Duct rapidly or even leans the head sideways, forward, or back-
System in the Maintenance of Equilibrium. Because ward, it would be impossible to maintain a stable image
the semicircular ducts do not detect that the body is off on the retinas unless the person had some automatic con-
trol mechanism to stabilize the direction of the eyes’ gaze.
In addition, the eyes would be of little use in detecting an
Rotation image unless they remained “fixed” on each object long
400
enough to gain a clear image. Fortunately, each time the
head is suddenly rotated, signals from the semicircular ducts
Impulses per second

300 cause the eyes to rotate in a direction equal and opposite to

Tonic
the rotation of the head. This results from reflexes transmit-
200 level of ted through the vestibular nuclei and the medial longitudi-
discharge Stop rotation nal fasciculus to the oculomotor nuclei. These reflexes are
described in Chapter 51.
100

Begin rotation Other Factors Concerned with Equilibrium

0 Neck Proprio­ceptors. The vestibular apparatus detects
0 10 20 30 40 50 60 70 80 90 the orientation and movement only of the head. Therefore, it
Seconds is essential that the nervous centers also receive appropriate
Figure 55-12 Response of a hair cell when a semicircular canal is information about the orientation of the head with respect
stimulated first by the onset of head rotation and then by stop- to the body. This information is transmitted from the pro-
ping rotation. prioceptors of the neck and body directly to the vestibular

677
Unit XI The Nervous System: C. Motor and Integrative Neurophysiology

and reticular nuclei in the brain stem and indirectly by way Fastigial Medial longitudinal
of the cerebellum. Dentate nucleus nucleus fasciculus
Among the most important proprioceptive information
needed for the maintenance of equilibrium is that transmit- Red
ted by joint receptors of the neck. When the head is leaned in nucleus
one direction by bending the neck, impulses from the neck Reticular
proprioceptors keep the signals originating in the vestibular substance
apparatus from giving the person a sense of dysequilibrium. Fastigioreticular
tract
They do this by transmitting signals that exactly oppose the
signals transmitted from the vestibular apparatus. However, Vestibular nucleus
when the entire body leans in one direction, the impulses
from the vestibular apparatus are not opposed by signals Vestibular nerve
Flocculonodular
from the neck proprioceptors; therefore, in this case, the
lobe
person does perceive a change in equilibrium status of the Vestibulospinal tract
entire body. Rubrospinal tract
Recticulospinal
Proprioceptive and Exteroceptive Information from Other
tract
Parts of the Body. Proprioceptive information from parts of
the body other than the neck is also important in the mainte- Figure 55-13 Connections of vestibular nerves through the ves-
tibular nuclei (large oval white area) with other areas of the central
nance of equilibrium. For instance, pressure sensations from
nervous system.
the footpads tell one (1) whether weight is distributed equally
between the two feet and (2) whether weight on the feet is
more forward or backward.
Exteroceptive information is especially necessary for the tation and inhibition of the many antigravity muscles, thus
maintenance of equilibrium when a person is running. The automatically controlling equilibrium.
air pressure against the front of the body signals that a force The flocculonodular lobes of the cerebellum are espe-
is opposing the body in a direction different from that caused cially concerned with dynamic equilibrium signals from the
by gravitational pull; as a result, the person leans forward to semicircular ducts. In fact, destruction of these lobes results
oppose this. in almost exactly the same clinical symptoms as destruction
Importance of Visual Information in the Maintenance of the semicircular ducts themselves. That is, severe injury
of Equilibrium. After destruction of the vestibular appara- to either the lobes or the ducts causes loss of dynamic equi-
tus, and even after loss of most proprioceptive information librium during rapid changes in direction of motion but does
from the body, a person can still use the visual mechanisms not seriously disturb equilibrium under static conditions. It
reasonably effectively for maintaining equilibrium. Even a is believed that the uvula of the cerebellum plays a similar
slight linear or rotational movement of the body instan- important role in static equilibrium.
taneously shifts the visual images on the retina, and this Signals transmitted upward in the brain stem from
information is relayed to the equilibrium centers. Some both the vestibular nuclei and the cerebellum by way of
people with bilateral destruction of the vestibular appara- the medial longitudinal fasciculus cause corrective move-
tus have almost normal equilibrium as long as their eyes ments of the eyes every time the head rotates, so the eyes
are open and all motions are performed slowly. But when remain fixed on a specific visual object. Signals also pass
moving rapidly or when the eyes are closed, equilibrium is upward (either through this same tract or through reticu-
immediately lost. lar tracts) to the cerebral cortex, terminating in a primary
cortical center for equilibrium located in the parietal lobe
deep in the sylvian fissure on the opposite side of the fis-
Neuronal Connections of the Vestibular Apparatus sure from the auditory area of the superior temporal gyrus.
with the Central Nervous System These signals apprise the psyche of the equilibrium status
of the body.
Figure 55-13 shows the connections in the hindbrain of the
vestibular nerve. Most of the vestibular nerve fibers terminate
in the brain stem in the vestibular nuclei, which are located
approximately at the junction of the medulla and the pons. Functions of Brain Stem Nuclei in Controlling
Some fibers pass directly to the brain stem reticular nuclei Subconscious, Stereotyped Movements
without synapsing and also to the cerebellar fastigial, uvular,
and flocculonodular lobe nuclei. The fibers that end in the Rarely, a baby is born without brain structures above the
brain stem vestibular nuclei synapse with second-order neu- mesencephalic region, a condition called anencephaly. Some
rons that also send fibers into the cerebellum, the vestibu- of these babies have been kept alive for many months. They
lospinal tracts, the medial longitudinal fasciculus, and other are able to perform some stereotyped movements for feed-
areas of the brain stem, particularly the reticular nuclei. ing, such as suckling, extrusion of unpleasant food from
The primary pathway for the equilibrium reflexes begins the mouth, and moving the hands to the mouth to suck the
in the vestibular nerves, where the nerves are excited by the ­fingers. In addition, they can yawn and stretch. They can cry
vestibular apparatus. The pathway then passes to the ves- and can follow objects with movements of the eyes and head.
tibular nuclei and cerebellum. Next, signals are sent into the Also, placing pressure on the upper anterior parts of their
reticular nuclei of the brain stem, as well as down the spinal legs causes them to pull to the sitting position. It is clear that
cord by way of the vestibulospinal and reticulospinal tracts. many of the stereotyped motor functions of the human being
The signals to the cord control the interplay between facili- are integrated in the brain stem.

678