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Chapter 3: Motor Cortex

James Knierim, Ph.D., Department of Neuroscience, The Johns Hopkins University

Last Review 20 Oct 2020

3.1 Introduction
The previous chapters discussed the lower levels of the motor hierarchy (the spinal cord and brainstem), which are involved in the low-level, “nuts and
bolts” processing that controls the activity of individual muscles. Individual alpha motor neurons control the force exerted by a particular muscle, and
spinal circuits can control sophisticated and complex behaviors such as walking and reflex actions. The types of movements controlled by these
circuits are not initiated consciously, however. Voluntary movements require the participation of the third and fourth levels of the hierarchy: the motor
cortex and the association cortex. These areas of the cerebral cortex plan voluntary actions, coordinate sequences of movements, make decisions
about proper behavioral strategies and choices, evaluate the appropriateness of a particular action given the current behavioral or environmental
context, and relay commands to the appropriate sets of lower motor neurons to execute the desired actions.

3.2 Motor Cortex Comprises the Primary Motor Cortex, Premotor Cortex, and Supplementary Motor Area

c 2000 UTHS CH

Primary mo to r c o rte x
Pre mo to r c o rte x
S upple me ntal mo to r are a

Figure 3.1
Motor cortex areas (lateral, dorsal, and medial
views). The primary motor cortex is located
immediately anterior to the central sulcus.
Select from the boxes in the center to see larger
views.

The motor cortex comprises three different areas of the frontal lobe, immediately anterior to the central sulcus. These areas are the primary motor
cortex (Brodmann’s area 4), the premotor cortex, and the supplementary motor area (Figure 3.1). Electrical stimulation of these areas elicits
movements of particular body parts. The primary motor cortex, or M1, is located on the precentral gyrus and on the anterior paracentral lobule on
the medial surface of the brain. Of the three motor cortex areas, stimulation of the primary motor cortex requires the least amount of electrical current
to elicit a movement. Low levels of brief stimulation typically elicit simple movements of individual body parts. Stimulation of premotor cortex or the
supplementary motor area requires higher levels of current to elicit movements, and often results in more complex movements than stimulation of
primary motor cortex. Stimulation for longer time periods (500 msec) in monkeys results in the movement of a particular body part to a stereotyped
posture or position, regardless of the initial starting point of the body part (Figure 3.2). Thus, the premotor cortex and supplementary motor areas
appear to be higher level areas that encode complex patterns of motor output and that select appropriate motor plans to achieve desired end results.

c 2000 UTHS CH STIMULATE 1

STIMULATE 2

Figure 3.2
Electrical stimulation of premotor cortex of a monkey for 500 msec produces movement to stereotyped postures depending on
the location of the stimulating electrode. Stimulation of site one (click STIMULATE 1) causes the monkey to bring its arm in front
of its eyes, regardless of the starting location of the arm, as if the monkey were producing a defensive posture. Stimulation of
site two (click STIMULATE 2) causes the monkey to bring its arm to its mouth and open the mouth, regardless of the starting
location of the arm, as if it were bringing a piece of food to its mouth (Graziano et al., 2002).

Like the somatosensory cortex of the postcentral gyrus, the primary motor cortex is somatotopically organized (Figure 3.3). Stimulation of the anterior
paracentral lobule elicits movements of the contralateral leg. As the stimulating electrode is moved across the precentral gyrus from dorsomedial to
ventrolateral, movements are elicited progressively from the torso, arm, hand, and face (most laterally). The representations of body parts that perform
precise, delicate movements, such as the hands and face, are disproportionately large compared to the representations of body parts that perform only
coarse, unrefined movements, such as the trunk or legs. The premotor cortex and supplementary motor area also contain somatotopic maps.

Figure 3.3
Somatotopic representation of motor outputs in motor
cortex.

One might predict that the motor cortex “homunculus” arises because neurons that control individual muscles are clustered together in the cortex.
That is, all of the neurons that control the biceps muscle may be located together, and all of the neurons that control the triceps may be clustered
nearby, and the neurons that control the soleus muscle may be clustered in a region further removed. Electrophysiological recordings have shown that
this is not the case, however. Movements of individual muscles are correlated with activity from widespread parts of the primary motor cortex. Similarly,
stimulation of small regions of primary motor cortex elicits movements that require the activity of numerous muscles. Thus, the primary motor cortex
homunculus does not represent the activity of individual muscles. Rather, it apparently represents the movements of individual body parts, which often
require the coordinated activity of large groups of muscles throughout the body.

3.3 Cortical Afferents and Efferents

The motor cortex exerts its influence over muscles by a variety of descending routes (Figure 3.4). Some of the descending pathways reviewed in the
last chapter can be influenced by motor cortex output. Thus, in addition to the direct cortical innervation of alpha motor neurons via the corticospinal
tract, the following cortical efferent pathways influence the remaining descending tracts:

1. the corticorubral tract allows cortex to modulate the rubrospinal tract

2. the corticotectal tract allows cortex to modulate the tectospinal tract
3. the corticoreticular tract allows cortex to modulate the reticulospinal tracts

c 2000 UTHS CH

Figure 3.4
Parallel pathways from the motor cortex allow the cortical
motor areas to influence the processing of all descending
motor tracts and side loops of the motor system.
Mouse over the pathways for more information.

The cortex can also influence the processing of the side loops of the motor hierarchy. The corticostriate tract innervates the caudate nucleus and
putamen of the basal ganglia. The corticopontine tract and cortico-olivary tract innervate important inputs to the cerebellum. Finally, cortical areas can
influence other cortical areas, directly via corticocortical pathways and indirectly via the corticothalamic pathways (Figure 3.5). Most of these pathways
are bi-directional. Thus, motor cortex receives input from other cortical areas, directly and indirectly through the thalamus, and it receives input from the
cerebellum and basal ganglia, always through the thalamus.

P os te rior pa rie ta l corte x S oma to

cor

M
S MA c
P re motor
corte x

CM VL VA

Ba s a l
Ce
ga nglia

Figure 3.5
Major connections of motor cortex. The cross-section on the left is a schematic version
of an idealized brain section that contains the major structures of the motor system
hierarchy for illustrative purposes; no actual brain section would contain all of these
structures. Move the cursor over each box on the right to highlight the inputs (blue) and
outputs (red) of each region.

3.4 Motor Cortex Cytoarchitecture

Like all parts of the neocortex, the primary motor cortex is made of six layers (Figure 3.6). Unlike primary sensory areas, primary motor cortex is
agranular cortex; that is, it does not have a cell-packed granular layer (layer 4). Instead, the most distinctive layer of primary motor cortex is its
descending output layer (Layer 5), which contains the giant Betz cells. These pyramidal cells and other projection neurons of the primary motor cortex
make up ~30% of the fibers in the corticospinal tract. The rest of the fibers come from the premotor cortex and the supplementary motor area (~30%),
the somatosensory cortex (~30%), and the posterior parietal cortex (~10%).

Figure 3.6
Pyramidal and non-pyramidal
neurons in motor cortex. The
cerebral cortex is organized into
six layers. These layers contain
different proportions of the two
main classes of cortical
neurons, pyramidal and non-
pyramidal cells. Pyramidal cells
send long axons down the
spinal cord and are the major
output neurons. They are
abundant in layer 5. Non-
pyramidal cells have axons
which terminate locally.

3.5 Encoding of Movement by Motor Cortex

Primary Motor Cortex

As discussed above, the primary motor cortex does not generally control individual muscles directly, but rather appears to control individual
movements or sequences of movements that require the activity of multiple muscle groups. Alpha motor neurons in the spinal cord, in turn, encode the
force of contraction of groups of muscle fibers using the rate code and the size principle. Thus, in accordance with the concept of hierarchical
organization of the motor system, the information represented by motor cortex is a higher level of abstraction than the information represented by
spinal motor neurons.

What is encoded by the neurons in primary motor cortex? Clues have come from recording the activity of these neurons as experimental animals
perform different motor tasks. In general, primary motor cortex encodes the parameters that define individual movements or simple movement
sequences.

1. Primary motor cortex neurons fire 5-100 msec before the onset of a movement. Thus, rather than firing as the result of muscle activity,
these neurons are involved in relaying motor commands to the alpha motor neurons that eventually cause the appropriate muscles to contract.
2. Primary motor cortex encodes the force of a movement. The amount of force required to raise the arm from one location to another is much
greater if one is holding a bowling ball than if one is holding a balloon. Many neurons in primary motor cortex encode the amount of force that is
necessary to make such a movement (Figure 3.7). Note the distinction between movement force and muscle force. Whereas a minority of
primary motor cortex neurons encodes individual muscle force, a larger number encodes the amount of force necessary for a particular
movement, regardless of which individual muscles are used. Alpha motor neurons, in turn, translate the commands of the motor cortex neurons
and control the amount of force generated by individual muscles to accomplish that movement, under the principles of the rate code and the size
principle.

Figures 3.7A, 3.7B, and 3.7C

Motor cortex encodes the force necessary to make a movement. (Evarts 1968)

PLAY
No lo ad Exte ns io n mo to r ne uro n

Figure 3.7A. When there is little load, a

motor neuron in primary motor cortex
that controls an extension of the wrist
fires when the wrist extends. A motor
Fle xio n mo to r ne uro n neuron that controls wrist flexion does
not change its low rate of activity. Note
that the extension motor neuron begins
to fire spikes before the onset of the
movement.
(e le c tro de in text goes here
primary
mo to r c o rte x)
c 2000 UTHS CH

PLAY
Exte ns o r lo ad Exte ns io n mo to r ne uro n

Figure 3.7B. When a 5 lb. load is placed

on the left pulley, more force must be
used to initially hold the weight steady
Fle xio n mo to r ne uro n and then lift it. The extension motor
neuron in primary motor cortex fires
more strongly to produce the greater
force.
text goes here
5 (e le c tro de in
lbs primary
mo to r c o rte x)
c 2000 UTHS CH

PLAY
Fle xo r lo ad Exte ns io n mo to r ne uro n Figure 3.7C. When a 5 lb. load is placed
on the right pulley, the load is on the
flexor. Thus, primary motor cortex
neurons for flexion are activated to keep
the weight stable. When the wrist
extends, the neurons are quieter, as the
force of the movement is actually
Fle xio n mo to r ne uro n produced by the weight itself. (Note that
motor cortex encodes the force of a
movement, such as wrist extension or
more complicated, multi-joint
movements. The force of individual
5 (e le c tro de in muscles is encoded by alpha motor
lbs primary neurons in the spinal cord and brain
mo to r c o rte x) stem.)
c 2000 UTHS CH text goes here

3. Primary motor cortex encodes the direction of movement. Many neurons in the primary motor cortex are selective for a particular direction
of movement. For example, one cell may fire strongly when the hand is moved to the left, whereas it will be inhibited when the hand is moved to
the right (Figure 3.8).

Clic k o n e ac h c 2000 UTHS CH

dire c tio nal c o o rdinate
0
90
0
135
0
45

0 0
180 0

0
225 315
0

0
270

Figure 3.8
Directional tuning of motor cortex neurons. The
cell fires maximally when the hand is moved in
the 135º or 180º directions, moderately when the
hand moves in the 90º and 225º directions, and is
silent when the hand moves in the opposite
directions (0º, 45º, 270º, and 315º) (Georgopoulos
et al., 1982).

4. Primary motor cortex encodes the extent of movement. The firing of some neurons is correlated with the distance of a movement. A monkey
was trained to move its arm to different target locations that varied in direction and distance from the center. The firing of many neurons was
correlated with the direction of movement (as in Point 3), whereas the firing of other neurons was correlated with the distance of the movement.
Interestingly, some neurons were correlated with the interaction of a particular distance and direction; that is, they were correlated with a
particular target position.
5. Primary motor cortex neurons encode the speed of movement. Almost all targeted movements follow a typical bell-shaped curve of velocity
as a function of distance (Figure 3.9). For example, when the hand moves an object such as a coffee cup from one location to another (the
target), the hand accelerates during the first half of the movement, reaches a peak velocity approximately halfway to the target, and then
decelerates until it reaches the target. The firing rate of some primary motor cortex neurons in monkeys correlates with this bell-shaped speed
profile, demonstrating that information about movement speed is contained in the spike trains of these neurons.

PLAY
Mo ve me nt s pe e d

c 2000 UTHS CH s tart midpo int e nd

Figure 3.9
Velocity profile of targeted movements.

Premotor Cortex

The premotor cortex sends axons to the primary motor cortex as well as to the spinal cord directly. It performs more complex, task-related processing
than primary motor cortex. Stimulation of premotor areas in the monkey at a high level of current produces more complex postures than stimulation of
the primary motor cortex. The premotor cortex appears to be involved in the selection of appropriate motor plans for voluntary movements, whereas
the primary motor cortex is involved in the execution of these voluntary movements.

1. Premotor cortex neurons signal the preparation for movement. Monkeys were trained to make a particular movement in response to a
visual signal, with a variable delay between the onset of the signal and the onset of the movement (Figure 3.10). Recordings from premotor
cortex have shown that many neurons fire selectively in the delay interval, for many seconds before the onset of the movement. A particular
neuron will fire when the monkey is preparing to make a movement to the left, for example, but will be silent when the monkey is preparing to
make a movement to the right. Thus, the firing of this type of neuron does not cause the movement itself, but appears to be involved in preparing
the monkey to make the correct movement when the “Go” signal is given. This type of neuron is called a motor-set neuron, as it fires when the
monkey is preparing, or getting set, to make a movement.

c 2000 U
Pre pare rig ht c e ll PLAY

PLAY
Pre pare le ft c e ll

Figure 3.10
Premotor cortex neurons signal preparation for movement. A monkey is trained to prepare to make
a movement to the right or left depending on a cue instruction, but to delay the movement until a
“Move” signal is given (Weinrich and Wise 1982). Some neurons will fire selectively when the animal
is preparing to make a movement to the right (Play Prepare right cell). Other neurons will fire
selectively when the animal is preparing to make a movement to the left (Play Prepare left cell). Note
that the cells fire in the interval between the Prepare instruction and the Move instruction, but they
do not fire during the movement itself.

2. Premotor cortex neurons signal various sensory aspects associated with particular motor acts. Some premotor neurons fire when the
animal is performing a particular action, such as breaking a peanut (Figure 3.11). Interestingly, the same neuron fires selectively when the
animal sees another monkey or person breaking the peanut. It also fires selectively to the sound of a peanut shell being broke, even without any
visual or motor activity. These neurons are called “mirror” neurons, because they respond not only to a particular action of the monkey but also
to the sight (or sound) of another individual performing the same action. (For an interesting PBS video on mirror neurons, go to
http://www.pbs.org/wgbh/nova/sciencenow/3204/01.html.)

c rac k
A

Mo nke y bre aking pe anut.

c rac k
B

Mo nke y watc hing a pe rs o n

bre ak a pe anut.

c rac k

Mo nke y he aring s o und o f

pe anut bre aking . c 2000 UTHS CH

Figure 3.11
Mirror neuron in premotor cortex fires to the monkey’s action as well as the monkey’s perception of a
person performing the same action (Kohler et al., 2002).

3. Premotor cortex is sensitive to the behavioral context of a particular movement. The premotor cortex of human subjects was imaged with
functional MRI as they observed video of a hand grasping a cup (Figure 3.12). In one condition, the cup was full and surrounded by full plates of
food; the implication was that the person was grasping the cup to take a drink. In the other condition, the cup was empty and surrounded by dirty
dishes; the implication was that the person was grasping the cup to clear the table. In this experiment, the premotor cortex was more active
when subjects viewed the former video than the latter, even though the movements were the same. Thus, premotor cortex neurons are sensitive
to the inferred intentions of a movement, not just the movement itself, as deduce from the behavioral context in which the movement occurred.

PLAY
Gras p c up
to drink

PLAY
Gras p c up to
c le ar table

c 2000 UTHS CH

Figure 3.12
Premotor cortex activity distinguishes the same movement based on the behavioral context
of the movement (Iacoboni et al., 2005). When the subject viewed an arm moving to pick up
a cup to drink (PLAY top), the activity in premotor cortex was greater than when the subject
viewed an arm moving to pick up a cup to clear the table after a meal (PLAY bottom). Note
that the strength of activity in the cortex (denoted by the brightness of the activated cortical
region) is greater in the top than in the bottom animations.

4. Premotor cortex signals correct and incorrect actions. Human subjects were studied in an fMRI experiment as they observed video clips of
various correct and incorrect motor acts. A correct action was one in which the movement and the associated object was performed correctly,
such as setting the time on a clock. An object error was one in which the action was correct, but the object was incorrect, such as polishing a
brown shoe with black shoe polish. A movement error was one in which the object was correct, but the movement was incorrect, such as
attempting to put a coin into a piggy bank when the coin was oriented perpendicular to the coin slot. In this experiment, the premotor cortex was
activated bilaterally during the correct actions trials and the movement error trials; for the object error trials, only the premotor cortex of the left
hemisphere was activated preferentially.

Supplementary Motor Area

The supplementary motor area (SMA) is involved in programming complex sequences of movements and coordinating bilateral movements.
Whereas the premotor cortex appears to be involved in selecting motor programs based on visual stimuli or on abstract associations, the
supplementary motor area appears to be involved in selecting movements based on remembered sequences of movements.

1. SMA responds to sequences of movements and to mental rehearsal of sequences of movements (Figure 3.13). Brain activity was
measured in a PET scanner while subjects made simple and complex sequences of movement. When the movements were simple, such as a
repetitive movement of a single digit, the primary motor cortex and the primary somatosensory cortex were activated on the contralateral
hemisphere. When the subject was asked to perform a complex sequence of finger movements, the SMA was activated bilaterally, in addition to
the contralateral primary motor and somatosensory cortex activation. Finally, when the subject was asked to remain still but to mentally rehearse
the complex sequence of activity, the SMA was still active, even though the primary motor and somatosensory cortex areas were silent. Thus,
the SMA appears to be involved in bilateral movements and in the mental rehearsal of these movements.

Le ft he mis phe re Rig ht he mis phe re Le ft hand Rig ht hand

PLAY M1 and S 1

S imple
mo ve me nt

PLAY S MA M1 and S 1 S MA

Co mple x
mo ve me nt

PLAY S MA S MA

Imag inary
c o mple x
mo ve me nt

c 2000 UTHS CH

Figure 3.13
Positron emission tomography (PET) study of simple vs. complex finger movements (Roland et al.,1980). The SMA
is activated bilaterally when subjects perform complex movements, and even when they only imagine performing
the movements.

2. SMA is involved in the transformation of kinematic to dynamic information. Movements can be defined in terms of dynamics (the amount
of force necessary to make a movement) and kinematics (the distance and angles that define a particular movement in space). Many
movement plans are represented in kinematic terms (e.g., Move the hand to the left). However, the motor system must eventually translate this
to a representation based on dynamics, in order to instruct the appropriate muscles to contract with the appropriate force. Recordings from
monkeys have shown that during the preparatory delay before a monkey makes an instructed movement, some SMA neurons change their firing
correlates from a kinematic-based representation to a dynamics-based representation, suggesting that SMA plays a vital role in this
transformation.

Association Cortex

The fourth level of the motor hierarchy is the association cortex, in particular the prefrontal cortex and the posterior parietal cortex (Figure 3.14).
These brain areas are not motor areas in the strict sense. Their activity does not correlate precisely with individual motor acts, and stimulation of these
areas does not result in motor output. However, these areas are necessary to ensure that movements are adaptive to the needs of the organism and
appropriate to the behavioral context.

Figure 3.14
Association cortex.
The prefrontal cortex is highlighted on the left,
and the posterior parietal cortex is highlighted
on the right.

1. Posterior parietal cortex is involved in ensuring that movements are targeted accurately to objects in external space. This area is
involved in processing spatial relationships of objects in the world and in constructing a representation of external space that is independent of
the observer’s eye position or body position. Such representations allow a stable percept of the world that is independent of viewer orientation,
as well as the representation of desired trajectories in space that are independent of body position. Damage to the posterior parietal cortex can
result in a number of apraxias, that is, the inability to make complex, coordinated movements. For example, a patient with constructional apraxia
is unable to replicate the configuration of a set of blocks in the proper sequence, even though the patient is able to maneuver each block
individually with dexterity.
2. Prefrontal cortex is involved in the selection of appropriate actions for a particular behavioral context. It is also involved in the evaluation
of the consequences of a particular course of action. Patients with damage to the prefrontal cortex have problems in executive processing. They
make inappropriate behavioral decisions, and often cannot anticipate the likely consequences of their actions. They display impulsive behavior,
often showing an inability to delay instant gratification for a long-term larger reward.

Test Your Knowledge

Question 1 A B C D E

Betz cells are most abundant in layer...

A. IV of somatosensory cortex.

B. V of somatosensory cortex.

C. IV of motor cortex.

D. V of motor cortex.

E. III of motor cortex.

Question 2 A B C D E

A corticospinal neuron in primary motor cortex can do all of the following EXCEPT:

A. Project to multiple motor neuron pools in the spinal cord.

B. Participate in the initiation of movement.

C. Code for the amount of force of individual muscles.

D. Code for the direction of movement.

E. Code for the extent of movement.

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