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Motor Pathways - Boundless Anatomy and Physiology
Motor Pathways - Boundless Anatomy and Physiology
Motor Pathways
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The motor system is the part of the central nervous system that is in-
volved with movement.
LEARNING OBJECTIVES
KEY TAKEAWAYS
Key Points
Key Terms
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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 commis-
sure) 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 neu-
ronal 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.
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The ventromedial lower motor neurons control the large, postural mus-
cles of the axial skeleton. These lower motor neurons, unlike those of
the dorsal lateral, are located in the ventral horn throughout the spinal
cord.
al
Pyramidal tracts
r
rvic
Lumba
cic
Gracile fasciculus
r
ba
Ce cic
l
Sa
ica
m
Cuneate fasciculus
a
or
rv
Th
Ce
Extrapyramidal Tracts rvic Spinocerebellar Tracts
Sacral r
Thora
Lumba c
al
- Rubrospinal tract Posterior spinocerebellar tract
ci
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 basal ganglia are responsible for voluntary motor control, proce-
dural learning, and eye movement, as well as cognitive and emotional
functions.
LEARNING OBJECTIVES
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KEY TAKEAWAYS
Key Points
targets.
Key Terms
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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.
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 influ-
ence 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
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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 result-
ing symptoms.
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).
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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 af-
fected, but motivational factors such as hunger fail to cause movements
to be initiated or switched at the proper times.
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 com-
bine to demonstrate that the dopaminergic projection from the VTA to
the NA plays a central role in the brain’s reward system.
Neurotransmitters
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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 ini-
tial excitation of the striatum, the internal dynamics of the basal ganglia
are dominated by inhibition and disinhibition.
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from cortex
back to cortex
glutamatergic
c
(excitatory) glutamatergi
(excitatory)
G
(di ABA
- d s-inh ergi
ire ibi c
ct t
pa ory)
thw
ay
Thalamus
Striatum External
globus GABAergic
pallidus G
(in ABA (inhibitory)
hib er
Internal ito gic
ry)
globus
pallidus
GABAergic Su
do (dis-dis-inhibitory) GA bth gluta
pa ala m
m - indirect pathway - (dis- BAer mi aterg
ine ind inhib gic cn
irec ito i
rg
ic t pa ry) uc c
thw leu
ay s
G
(dis ABA Substantia nigra
pars
-in ergic dopaminergic pars
hib compacta
ito reticularis
ry)
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.
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LEARNING OBJECTIVES
KEY TAKEAWAYS
Key Points
Key Terms
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It receives input from sensory systems of the spinal cord and from other
parts of the brain, including the cerebral cortex, and integrates these in-
puts to fine-tune motor activity. Because of this fine-tuning function,
damage to the cerebellum does not cause paralysis, but instead pro-
duces 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 unidirection-
ally from input to output. This feedforward mode of operation means that
the cerebellum cannot generate self-sustaining patterns of neural activ-
ity, in contrast to the cerebral cortex. However, the cerebellum can re-
ceive information from the cerebral cortex and processes this informa-
tion to send motor impulses to the skeletal muscle.
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Within this thin layer are several types of neurons with a highly regular
arrangement, the most important being Purkinje cells and granule cells.
This complex neural network gives rise to a massive signal-processing
capability, but almost all of its output is directed to a set of small, deep
cerebellar nuclei lying in the interior of the cerebellum.
Function
Marr-Albus Theory
In addition to its direct role in motor control, the cerebellum is also nec-
essary for several types of motor learning, the most notable one being
learning to adjust to changes in sensorimotor relationships.
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weak. However, each cerebellar Purkinje cell also gets input from one
single climbing fiber, which is so strong that a single climbing fiber action
potential will reliably cause a target Purkinje cell to fire a burst of action
potentials.
The basic concept of the Marr-Albus theory is that the climbing fiber
serves as a teaching signal, which induces a long-lasting change in the
strength of synchronously activated parallel fiber inputs. Observations of
long-term depression in parallel fiber inputs have provided support for
theories of this type, but their validity remains controversial.
The strongest clues to the function of the cerebellum have come from
examining the consequences of damage to it. Animals and humans with
cerebellar dysfunction show, above all, problems with motor control.
They continue to be able to generate motor activity, but it loses preci-
sion, producing erratic, uncoordinated, or incorrectly timed movements.
A standard test of cerebellar function is to reach with the tip of the finger
for a target at arm’s length. A healthy person will move the fingertip in a
rapid straight trajectory, whereas a person with cerebellar damage will
reach slowly and erratically, with many mid-course corrections.
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LEARNING OBJECTIVES
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KEY TAKEAWAYS
Key Points
Key Terms
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Cerebellar Function
Feedforward Processing
The cerebellum differs from most other parts of the brain in that the sig-
nal processing is almost entirely feedforward—that is, signals move uni-
directionally through the system from input to output, with very little re-
current internal transmission.
The small amount of recurrence that does exist consists of mutual inhibi-
tion; there are no mutually excitatory circuits. This feedforward mode of
operation means that the cerebellum, in contrast to the cerebral cortex,
cannot generate self-sustaining patterns of neural activity.
Signals enter the circuit, are processed by each stage in sequential or-
der, and then leave. As Eccles, Ito, and Szentágothai wrote,”This elimina-
tion in the design of all possibility of reverberatory chains of neuronal ex-
citation is undoubtedly a great advantage in the performance of the
cerebellum as a computer, because what the rest of the nervous system
requires from the cerebellum is presumably not some output expressing
the operation of complex reverberatory circuits in the cerebellum, but
rather a quick and clear response to the input of any particular set of
information.”
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Modularity
Different modules share input from mossy fibers and parallel fibers, but
in other respects they appear to function independently. The output of
one module does not appear to significantly influence the activity of
other modules.
Plasticity
The synapses between parallel fibers and Purkinje cells, and the
synapses between mossy fibers and deep nuclear cells, are both sus-
ceptible to modification of their strength. In a single cerebellar module,
input from as many as a billion parallel fibers converge onto a group of
less than 50 deep nuclear cells, and the influence of each parallel fiber
on those nuclear cells is adjustable. This arrangement gives tremendous
flexibility for fine-tuning the relationships between the cerebellar inputs
and outputs.
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Microzone
Zone
Purkinje cell
dendritic Parallel
trees fibers
Unfolded
cerebellar
cortex
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