Boundless Anatomy and

Physiology

Peripheral Nervous System

Motor Pathways

Organization of Motor Neuron

Pathways

The motor system is the part of the central

nervous system that is involved with
movement.

LEARNING OBJECTIVES

Describe the organization of motor

neuron pathways

KEY TAKEAWAYS

Key Points

The pyramidal tract, which

includes both the
corticospinal and
corticobulbar tracts, serves as
the motor pathway for upper
motor neuronal signals
coming from the cerebral
cortex and from primitive
brainstem motor nuclei.

Peripheral motor nerves carry

the motor impulses from the
spinal cord to the voluntary
muscles.

The large majority (90%) of

motor neurons cross
(decussate) to the
contralateral side of the brain
at the level of the brainstem.

Key Terms

extrapyramidal system: A
biological neural network that
is part of the motor system
that causes involuntary
movements.

corticospinal tract: The

nervous system tract that
conducts impulses from the
brain to the spinal cord. It
contains mostly motor axons
and is made up of two
separate tracts in the spinal
cord: the lateral corticospinal
tract and the anterior
corticospinal tract.

motor system: The part of the

central nervous system that is
involved with movement. It
consists of the pyramidal and
extrapyramidal systems.

cerebral cortex: The gray,

folded, outermost layer of the
cerebrum that is responsible
for higher brain processes
such as sensation, voluntary
muscle movement, thought,
reasoning, and memory.

The motor system is

the part of the
central nervous
system that is
involved with
movement. It
consists of the
pyramidal and
extrapyramidal
system.

Decussation of the
The motor pathway, pyramids: A deep
dissection, lateral view
also called the of a brainstem. The
pyramidal tract is visible
pyramidal tract or in red, and pyramidal
the corticospinal decussation is labeled
at lower right.
tract, serves as the
motor pathway for
upper motor neuronal signals coming from
the cerebral cortex and from primitive
brainstem motor nuclei. There are upper and
lower motor neurons in the corticospinal
tract.

The motor impulses originate in the giant

pyramidal cells (Betz cells) of the motor area,
i.e., the precentral gyrus of the cerebral
cortex. These are the upper motor neurons of
the corticospinal tract. The axons of these
cells pass from the cerebral cortex to the
midbrain and the medulla oblongata.
Peripheral motor nerves carry the motor
impulses from the anterior horn to the
voluntary muscles.

Cortical upper motor neurons originate from

Brodmann areas 1, 2, 3, 4, and 6, then
descend into the posterior limb of the internal
capsule, through the crus cerebri, down
through the pons, and to the medullary
pyramids, where about 90% of the axons
cross to the contralateral side at the
decussation of the pyramids. They then
descend as the lateral corticospinal tract.

These axons synapse with lower motor

neurons in the ventral horns of all levels of
the spinal cord. The remaining 10% of axons
descend on the ipsilateral side as the ventral
corticospinal tract. These axons also synapse
with lower motor neurons in the ventral
horns. Most of them will cross to the
contralateral side of the cord (via the anterior
white commissure) just before synapsing.

Brodmann areas of the brain: This drawing shows

the regions of the human cerebral cortex as
delineated by Korvinian Brodmann on the basis of
cytoarchitecture.

The midbrain nuclei include four motor tracts

that send upper motor neuronal axons down
the spinal cord to lower motor neurons.
These are the rubrospinal tract, the
vestibulospinal tract, the tectospinal tract,
and the reticulospinal tract.

The function of lower motor neurons can be

divided into two different groups: the lateral
corticospinal tract and the anterior
corticalspinal tract. The lateral tract contains
upper motor neuronal axons that synapse on
the dorsal lateral lower motor neurons, which
are involved in distal limb control.

The anterior corticospinal tract descends

ipsilaterally in the anterior column, where the
axons emerge and either synapse on
ventromedial lower motor neurons in the
ventral horn ipsilaterally or descussate at the
anterior white commissure where they
synapse on ventromedial lower motor
neurons contralaterally.

The ventromedial lower motor neurons

control the large, postural muscles of the
axial skeleton. These lower motor neurons,
unlike those of the dorsal lateral, are located
in the ventral horn throughout the spinal
cord.

Motor and descending (efferent) Sensory and ascending

pathways (red) (afferent) pathways
(blue)
Sacral

Pyramidal tracts
al
r

rvic
Lumba

cic

- Lateral corticospinal tract Dorsal Column Medial

Ce
Thora

- Anterior corticospinal tract Lemniscus System

Gracile fasciculus
al
cr

r
ba

cic
Sa

l
i ca

Cuneate fasciculus
m

ra
Lu

rv
o
Ce
Th

Ce
Extrapyramidal Tracts rvic Spinocerebellar Tracts
Sacral
Thora
Lumb

al
- Rubrospinal tract Posterior spinocerebellar tract
cic
ar

- Reticulospinal tracts Anterior spinocerebellar tract

- Olivospinal tract
- Vestibulospinal tract Anterolateral System
Lateral spinothalamic tract
Anterior spinothalamic tract

Spino-olivary fibers

Spinal cord tracts: This diagram of spinal cord tracts

shows the motor and efferent pathways in red and
the sensory and afferent pathways in blue. Included
in the diagram are the following motor pathways:
corticospinal tracts (pyramidal tract), and
extrapyramidal tracts (tectospinal tract not
delineated).

The Role of the Basal Ganglia in

Movement

The basal ganglia are responsible for

voluntary motor control, procedural learning,
and eye movement, as well as cognitive and
emotional functions.

LEARNING OBJECTIVES

Describe the role of the basal

ganglia in movement

KEY TAKEAWAYS

Key Points

The basal ganglia are studied

extensively in the context of
two disorders of the basal
ganglia: Parksinson’s disease
and Huntington’s disease.

Hemiballismus, a movement
disorder arising from neuronal
damage in the subthalamic
nucleus, presents with violent
movements of the arms and
legs.

Eye movement, a function of

the basal ganglia, is
influenced by the superior
colliculus, a region of the
brain that directs eye
movement to specific points
in space in response to
stimuli.

Basal ganglia are also thought

to play a role in motivation.

In the basal ganglia,

the majority of the neurons
uses gamma-aminobutyric
acid (GABA) as the
neurotransmitter
and have inhibitory effects on
their
targets.

Key Terms

hemiballismus: A rare
movement disorder with
involuntary flinging motions of
the extremities.

voluntary motor control: The

act of directing motion with
intent.

forebrain: The anterior part of

the brain, including the
cerebrum, thalamus, and
hypothalamus.

nucleus accumbens: A region

in the basal forebrain rostral
to the preoptic area of the
hypothalamus. This region
and the olfactory tubercle
collectively form the ventral
striatum.

Location of the Basal Ganglia

The basal ganglia (or basal nuclei) are a

group of nuclei of varied origin in the brains
of vertebrates that act as a cohesive
functional unit. They are situated at the base
of the forebrain and are strongly connected
with the cerebral cortex, thalamus, and other
brain areas.

The basal ganglia are associated with a

variety of functions, including voluntary motor
control, procedural learning relating to
routine behaviors or habits such as bruxism
and eye movements, as well as cognitive
and emotional functions.

Basal ganglia: Locations of the basal ganglia.

Action Selection

Currently popular theories hold that the basal

ganglia play a primary role in action
selection. Action selection is the decision of
which of several possible behaviors to
execute at a given time.

Experimental studies show that the basal

ganglia exert an inhibitory influence on a
number of motor systems, and that a release
of this inhibition permits a motor system to
become active. The behavior switching that
takes place within the basal ganglia is
influenced by signals from many parts of the
brain, including the prefrontal cortex, which
plays a key role in executive functions.

Movement

The greatest source of insight into the

functions of the basal ganglia has come from
the study of two neurological disorders,
Parkinson’s disease and Huntington’s
disease. For both of these disorders, the
nature of the neural damage is well-
understood and can be correlated with the
resulting symptoms.

Parkinson’s disease involves the major loss of

dopaminergic cells in the substantia nigra.
Huntington’s disease involves the massive
loss of medium spiny neurons in the striatum.

The symptoms of the two diseases are

virtually opposite: Parkinson’s disease is
characterized by a gradual loss of the ability
to initiate movement, whereas Huntington’s
disease is characterized by an inability to
prevent parts of the body from moving
unintentionally.

It is noteworthy that, although both diseases

have cognitive symptoms, especially in their
advanced stages, the most salient symptoms
relate to the ability to initiate and control
movement. Thus, both are classified primarily
as movement disorders.

A different movement disorder, called

hemiballismus, may result from damage
restricted to the subthalamic nucleus.
Hemiballismus is characterized by violent
and uncontrollable flinging movements of the
arms and legs.

Function in Eye Movement

One of the most intensively studied functions

of the basal ganglia is their role in controlling
eye movements. Eye movement is influenced
by an extensive network of brain regions that
converge on a midbrain area called the
superior colliculus (SC).

The SC is a layered structure whose layers

form two-dimensional retinotopic maps of
visual space. A bump of neural activity in the
deep layers of the SC drives eye movement
toward the corresponding point in space.

Motivation

Although the role of the basal ganglia in

motor control is clear, there are also many
indications that it is involved in the control of
behavior in a more fundamental way, at the
level of motivation. In Parkinson’s disease,
the ability to execute the components of
movement is not greatly affected, but
motivational factors such as hunger fail to
cause movements to be initiated or switched
at the proper times.

The immobility of patients with Parkingson’s

disease has sometimes been described as a
paralysis of the will. These patients have
occasionally been observed to show a
phenomenon called kinesia paradoxica, in
which a person who is otherwise immobile
responds to an emergency in a coordinated
and energetic way, then lapses back into
immobility once the emergency has passed.

The role in motivation of the limbic part of the

basal ganglia—the nucleus accumbens (NA),
ventral pallidum, and ventral tegmental area
(VTA)—is particularly well established.
Thousands of experimental studies combine
to demonstrate that the dopaminergic
projection from the VTA to the NA plays a
central role in the brain’s reward system.

Numerous things that people find rewarding,

including addictive drugs, good-tasting food,
and sex, have been shown to elicit activation
of the VTA dopamine system. Damage to the
NA or VTA can produce a state of profound
torpor.

Neurotransmitters

In most regions of the brain, the predominant

classes of neurons use glutamate as the
neurotransmitter and have excitatory effects
on their targets. In the basal ganglia,
however, the great majority of neurons uses
gamma-aminobutyric acid (GABA) as the
neurotransmitter and have inhibitory effects
on their targets.

The inputs from the cortex and thalamus to

the striatum and subthalamic nucleus are
glutamatergic, but the outputs from the
striatum, pallidum, and substantia nigra pars
reticulata all use GABA. Thus, following the
initial excitation of the striatum, the internal
dynamics of the basal ganglia are dominated
by inhibition and disinhibition.

Other neurotransmitters have important

modulatory effects. Dopamine is used by the
projection from the substantia nigra pars
compacta to the dorsal striatum and also in
the analogous projection from the ventral
tegmental area to the ventral striatum
(nucleus accumbens).

Acetylcholine also plays an important role, as

it is used both by several external inputs to
the striatum and by a group of striatal
interneurons. Although cholinergic cells
make up only a small fraction of the total
population, the striatum has one of the
highest acetylcholine concentrations of any
brain structure.

from cortex
glutamatergic
back to cortex
ic
(excitatory) glutamatergy)
(excitator

G
(di AB
- d s-inh Aerg
ire ib
ct ito ic
pa ry)
thw
ay
Thalamus

Striatum External
globus GABAergic
pallidus G
(in ABA (inhibitory)
hib e
Internal ito rgic
ry)
globus
pallidus
GABAergic Su
do
pa (dis-dis-inhibitory) GA bth glut
m (
- indirect pathway- i dis-in BAer ala ama
ndi hib gic mi t
ine
rg rec ito c n ergic
ic t pa ry) uc
thw leu
ay s
G
(di ABA Substantia nigra
s-i e pars pars
nh rgi dopaminergic
ibit c compacta
ory reticularis
)

Main circuits of the basal ganglia: This diagram

shows the main circuits of the basal ganglia. Two
coronal slices have been superimposed to include
the involved basal ganglia structures. The + and –
signs at the point of the arrows indicate whether the
pathway is excitatory or inhibitory, respectively, in
effect. Green arrows refer to excitatory
glutamatergic pathways, red arrows refer to
inhibitory GABAergic pathways and turquoise
arrows refer to dopaminergic pathways that are
excitatory on the direct pathway and inhibitory on
the indirect pathway.

Modulation of Movement by the

Cerebellum

The cerebellum is important for motor control

—specifically coordination, precision, and
timing—as well as some forms of motor
learning.

LEARNING OBJECTIVES

Describe the role of the cerebellum

in movement modulation

KEY TAKEAWAYS

Key Points

The cerebellum is a parallel

grooved structure at the
bottom of the brain containing
a highly regular cellular
arrangement of Purkinje cells,
granule cells, and other cell
types.

The cerebellum adjusts to

changes in sensorimotor
relationships, possibly
functioning as in the Marr-
Albus theory: Strong inputs
from a single climbing fiber
serve as a teaching signal to
change the strength of
impulses from the
corresponding group of
parallel fibers.

Four principles of cerebellum

function have been identified.
They include: feedforward
processing, divergence and
convergence, modularity, and
plasticity.

Key Terms

Purkinje cells: A class of

GABAergic neurons located
in the cerebellum.

mossy fibers: One of the

major inputs to the
cerebellum from sources such
as the cerebral cortex.

granule cells: These cells

receive excitatory input from
mossy fibers that originate
from pontine nuclei.

The cerebellum is a region of the brain that

plays an important role in motor control. It
may also be involved in some cognitive
functions such as attention and language,
and in regulating fear and pleasure
responses, but its movement-related
functions are the most solidly established.
The cerebellum does not initiate movement,
but it contributes to coordination, precision,
and accurate timing.

It receives input from sensory systems of the

spinal cord and from other parts of the brain,
including the cerebral cortex, and integrates
these inputs to fine-tune motor activity.
Because of this fine-tuning function, damage
to the cerebellum does not cause paralysis,
but instead produces disorders in fine
movement, equilibrium, posture, and motor
learning.

The cerebellum differs from most other parts

of the brain, especially the cerebral cortex, in
regards to the ability of signals to move
unidirectionally from input to output. This
feedforward mode of operation means that
the cerebellum cannot generate self-
sustaining patterns of neural activity, in
contrast to the cerebral cortex. However, the
cerebellum can receive information from the
cerebral cortex and processes this
information to send motor impulses to the
skeletal muscle.

Cerebellum: View of the cerebellum from above and

behind.