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Motor neuron
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A motor neuron (or motoneuron or efferent neuron[1]) is a neuron whose cell body is located in the
motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or
outside of the spinal cord to directly or indirectly control effector organs, mainly muscles and
glands.[2] There are two types of motor neuron – upper motor neurons and lower motor neurons.
Axons from upper motor neurons synapse onto interneurons in the spinal cord and occasionally
directly onto lower motor neurons.[3] The axons from the lower motor neurons are efferent nerve fibers
that carry signals from the spinal cord to the effectors.[4] Types of lower motor neurons are alpha
motor neurons, beta motor neurons, and gamma motor neurons.

A single motor neuron may innervate many

Motor neurons
muscle fibres and a muscle fibre can undergo
many action potentials in the time taken for a
single muscle twitch. Innervation takes place at
a neuromuscular junction and twitches can
become superimposed as a result of
summation or a tetanic contraction. Individual
twitches can become indistinguishable, and
tension rises smoothly eventually reaching a
plateau.[5]
Micrograph of the hypoglossal nucleus showing
Although the word "motor neuron" suggests
motor neurons with their characteristic coarse Nissl
that there is a single kind of neuron that substance ("tigroid" cytoplasm). H&E-LFB stain.
controls movement, this is not the case.
Indeed, upper and lower motor neurons—which Details
differ greatly in their origins, synapse
Location Ventral horn of the
locations, routes, neurotransmitters, and lesion
spinal cord, some cranial
characteristics—are included in the same nerve nuclei
classification as "motor neurons." Essentially,
motor neurons, also known as motoneurons, Shape Projection neuron
are made up of a variety of intricate, finely
Function Excitatory projection (to
tuned circuits found throughout the body that
NMJ)
innervate effector muscles and glands to
enable both voluntary and involuntary motions. Neurotransmitter UMN to LMN: glutamate;
Two motor neurons come together to form a LMN to NMJ: ACh
two-neuron circuit. While lower motor neurons
Presynaptic Primary motor cortex via
start in the spinal cord and go to innervate
connections the Corticospinal tract
muscles and glands all throughout the body,
upper motor neurons originate in the cerebral Postsynaptic Muscle fibers and other
cortex and travel to the brain stem or spinal connections neurons
cord. It is essential to comprehend the
Identifiers
distinctions between upper and lower motor
neurons as well as the routes they follow in MeSH D009046
order to effectively detect these neuronal
injuries and localise the lesions. [6] NeuroLex ID nifext_103

TA98 A14.2.00.021
Contents
TA2 6131
Development
FMA 83617
Anatomy and physiology
Upper motor neurons
Anatomical terms of neuroanatomy
Nerve tracts [edit on Wikidata]

Lower motor neurons

Somatic motor neurons

Special visceral motor neurons

General visceral motor neurons

Neuromuscular junctions

Synaptic input to motor neurons

See also

References

Sources

Development

Motor neurons begin to develop early in embryonic development, and motor function continues to
develop well into childhood.[7] In the neural tube cells are specified to either the rostral-caudal axis or
ventral-dorsal axis. The axons of motor neurons begin to appear in the fourth week of development
from the ventral region of the ventral-dorsal axis (the basal plate).[8] This homeodomain is known as
the motor neural progenitor domain (pMN). Transcription factors here include Pax6, OLIG2, Nkx-6.1,
and Nkx-6.2, which are regulated by sonic hedgehog (Shh). The OLIG2 gene being the most important
due to its role in promoting Ngn2 expression, a gene that causes cell cycle exiting as well as promoting
further transcription factors associated with motor neuron development.[9]

Further specification of motor neurons occurs when retinoic acid, fibroblast growth factor, Wnts, and
TGFb, are integrated into the various Hox transcription factors. There are 13 Hox transcription factors
and along with the signals, determine whether a motor neuron will be more rostral or caudal in
character. In the spinal column, Hox 4-11 sort motor neurons to one of the five motor columns.[9]

Motor columns of spinal cord [10]

Motor column Location in spinal cord Target

Median motor
Present entire length Axial muscles
column

Hypaxial motor Body wall

Thoracic region
column muscles

Preganglionic motor Sympathetic

Thoracic region
column ganglion

Lateral motor Brachial and lumbar region (both regions are further divided into Muscles of the
column medial and lateral domains) limbs

Phrenic motor
Cervical region Diaphragm[11]
column

Anatomy and physiology

Upper
motor
neurons

Upper motor
neurons originate
in the motor
cortex located in
the precentral
gyrus. The cells
that make up the
primary motor Spinal cord tracts
cortex are Betz
cells, which are giant pyramidal cells. The axons of these cells
descend from the cortex to form the corticospinal tract.[12]
Corticomotorneurons project from the primary cortex directly
onto motor neurons in the ventral horn of the spinal cord.[13][14]
Their axons synapse on the spinal motor neurons of multiple
muscles as well as on spinal interneurons.[13][14] They are unique
to primates and it has been suggested that their function is the
adaptive control of the hands including the relatively independent
Location of lower motor neurons in spinal
control of individual fingers.[14][15] Corticomotorneurons have so cord
far only been found in the primary motor cortex and not in
secondary motor areas.[14]

Nerve tracts

Nerve tracts are bundles of axons as white matter, that carry action potentials to their effectors. In the
spinal cord these descending tracts carry impulses from different regions. These tracts also serve as
the place of origin for lower motor neurons. There are seven major descending motor tracts to be
found in the spinal cord:[16]

Lateral corticospinal tract

Rubrospinal tract

Lateral reticulospinal tract

Vestibulospinal tract

Medial reticulospinal tract

Tectospinal tract

Anterior corticospinal tract

Lower motor neurons

Lower motor neurons are those that originate in the spinal cord and directly or indirectly innervate
effector targets. The target of these neurons varies, but in the somatic nervous system the target will
be some sort of muscle fiber. There are three primary categories of lower motor neurons, which can
be further divided in sub-categories.[17]

According to their targets, motor neurons are classified into three broad categories:[18]

Somatic motor neurons

Special visceral motor neurons

General visceral motor neurons

Somatic motor neurons

Somatic motor neurons originate in the central nervous system, project their axons to skeletal
muscles[19] (such as the muscles of the limbs, abdominal, and intercostal muscles), which are involved
in locomotion. The three types of these neurons are the alpha efferent neurons, beta efferent neurons,
and gamma efferent neurons. They are called efferent to indicate the flow of information from the
central nervous system (CNS) to the periphery.

Alpha motor neurons innervate extrafusal muscle fibers, which are the main force-generating component
of a muscle. Their cell bodies are in the ventral horn of the spinal cord and they are sometimes called
ventral horn cells. A single motor neuron may synapse with 150 muscle fibers on average.[20] The motor
neuron and all of the muscle fibers to which it connects is a motor unit. Motor units are split up into 3
categories:[21]
Slow (S) motor units stimulate small muscle fibers, which contract very slowly and provide small
amounts of energy but are very resistant to fatigue, so they are used to sustain muscular contraction,
such as keeping the body upright. They gain their energy via oxidative means and hence require
oxygen. They are also called red fibers.[21]

Fast fatiguing (FF) motor units stimulate larger muscle groups, which apply large amounts of force
but fatigue very quickly. They are used for tasks that require large brief bursts of energy, such as
jumping or running. They gain their energy via glycolytic means and hence do not require oxygen.
They are called white fibers.[21]

Fast fatigue-resistant motor units stimulate moderate-sized muscles groups that do not react as fast
as the FF motor units, but can be sustained much longer (as implied by the name) and provide more
force than S motor units. These use both oxidative and glycolytic means to gain energy.[21]

In addition to voluntary skeletal muscle contraction, alpha motor neurons also contribute to muscle
tone, the continuous force generated by noncontracting muscle to oppose stretching. When a muscle
is stretched, sensory neurons within the muscle spindle detect the degree of stretch and send a signal
to the CNS. The CNS activates alpha motor neurons in the spinal cord, which cause extrafusal muscle
fibers to contract and thereby resist further stretching. This process is also called the stretch reflex.

Beta motor neurons innervate intrafusal muscle fibers of muscle spindles, with collaterals to extrafusal
fibres. There are two types of beta motor neurons: Slow Contracting- These innervate extrafusal fibers.
Fast Contracting- These innervate intrafusal fibers.[22]

Gamma motor neurons innervate intrafusal muscle fibers found within the muscle spindle. They regulate
the sensitivity of the spindle to muscle stretching. With activation of gamma neurons, intrafusal muscle
fibers contract so that only a small stretch is required to activate spindle sensory neurons and the stretch
reflex. There are two types of gamma motor neurons: Dynamic- These focus on Bag1 fibers and enhance
dynamic sensitivity. Static- These focus on Bag2 fibers and enhance stretch sensitivity.[22]

Regulatory factors of lower motor neurons

Size Principle – this relates to the soma of the motor neuron. This restricts larger neurons to receive a
larger excitatory signal in order to stimulate the muscle fibers it innervates. By reducing unnecessary
muscle fiber recruitment, the body is able to optimize energy consumption.[22]

Persistent Inward Current (PIC) – recent animal study research has shown that constant flow of ions
such as calcium and sodium through channels in the soma and dendrites influence the synaptic input.
An alternate way to think of this is that the post-synaptic neuron is being primed before receiving an
impulse.[22]

After Hyper-polarization (AHP) – A trend has been identified that shows slow motor neurons to have
more intense AHPs for a longer duration. One way to remember this is that slow muscle fibers can
contract for longer, so it makes sense that their corresponding motor neurons fire at a slower rate.[22]
Special visceral motor neurons

These are also known as branchial motor neurons, which are involved in facial expression, mastication,
phonation, and swallowing. Associated cranial nerves are the oculomotor, abducens, trochlear, and
hypoglossal nerves.[18]

Branch of NS Position Neurotransmitter

Somatic n/a Acetylcholine

Parasympathetic Preganglionic Acetylcholine

Parasympathetic Ganglionic Acetylcholine

Sympathetic Preganglionic Acetylcholine

Sympathetic Ganglionic Norepinephrine*

*Except fibers to sweat glands and certain blood vessels

Motor neuron neurotransmitters

General visceral motor neurons

These motor neurons indirectly innervate cardiac muscle and smooth muscles of the viscera ( the
muscles of the arteries): they synapse onto neurons located in ganglia of the autonomic nervous
system (sympathetic and parasympathetic), located in the peripheral nervous system (PNS), which
themselves directly innervate visceral muscles (and also some gland cells).

In consequence, the motor command of skeletal and branchial muscles is monosynaptic involving only
one motor neuron, either somatic or branchial, which synapses onto the muscle. Comparatively, the
command of visceral muscles is disynaptic involving two neurons: the general visceral motor neuron,
located in the CNS, synapses onto a ganglionic neuron, located in the PNS, which synapses onto the
muscle.

All vertebrate motor neurons are cholinergic, that is, they release the neurotransmitter acetylcholine.
Parasympathetic ganglionic neurons are also cholinergic, whereas most sympathetic ganglionic
neurons are noradrenergic, that is, they release the neurotransmitter noradrenaline. (see Table)

Neuromuscular junctions

A single motor neuron may innervate many muscle fibres and a muscle fibre can undergo many action
potentials in the time taken for a single muscle twitch. As a result, if an action potential arrives before
a twitch has completed, the twitches can superimpose on one another, either through summation or a
tetanic contraction. In summation, the muscle is stimulated repetitively such that additional action
potentials coming from the somatic nervous system arrive before the end of the twitch. The twitches
thus superimpose on one another, leading to a force greater than that of a single twitch. A tetanic
contraction is caused by constant, very high frequency stimulation - the action potentials come at
such a rapid rate that individual twitches are indistinguishable, and tension rises smoothly eventually
reaching a plateau.[5]

The interface between a motor neuron and muscle fiber is a specialized synapse called the
neuromuscular junction. Upon adequate stimulation, the motor neuron releases a flood of
acetylcholine (Ach) neurotransmitters from the axon terminals from synaptic vesicles bind with the
plasma membrane. The acetylcholine molecules bind to postsynaptic receptors found within the motor
end plate. Once two acetylcholine receptors have been bound, an ion channel is opened and sodium
ions are allowed to flow into the cell. The influx of sodium into the cell causes depolarization and
triggers a muscle action potential. T tubules of the sarcolemma are then stimulated to elicit calcium
ion release from the sarcoplasmic reticulum. It is this chemical release that causes the target muscle
fiber to contract.[20]

In invertebrates, depending on the neurotransmitter released and the type of receptor it binds, the
response in the muscle fiber could be either excitatory or inhibitory. For vertebrates, however, the
response of a muscle fiber to a neurotransmitter can only be excitatory, in other words, contractile.
Muscle relaxation and inhibition of muscle contraction in vertebrates is obtained only by inhibition of
the motor neuron itself. This is how muscle relaxants work by acting on the motor neurons that
innervate muscles (by decreasing their electrophysiological activity) or on cholinergic neuromuscular
junctions, rather than on the muscles themselves.

Synaptic input to motor neurons

Motor neurons receive synaptic input from premotor neurons. Premotor neurons can be 1) spinal
interneurons that have cell bodies in the spinal cord, 2) sensory neurons that convey information from
the periphery and synapse directly onto motoneurons, 3) descending neurons that convey information
from the brain and brainstem. The synapses can be excitatory, inhibitory, electrical, or
neuromodulatory. For any given motor neuron, determining the relative contribution of different input
sources is difficult, but advances in connectomics have made it possible for fruit fly motor neurons. In
the fly, motor neurons controlling the legs and wings are found in the ventral nerve cord, homologous
to the spinal cord. Fly motor neurons vary by over 100X in the total number of input synapses.
However, each motor neuron gets similar fractions of its synapses from each premotor source: ~70%
from neurons within the VNC, ~10% from descending neurons, ~3% from sensory neurons, and ~6%
from VNC neurons that also send a process up to the brain. The remaining 10% of synapses come
from neuronal fragments that are unidentified by current image segmentation algorithms and require
additional manual segmentation to measure.[23]

See also

Betz cell

Central chromatolysis

Motor dysfunction

Motor neuron disease

Nerve

Sensory nerve

Motor nerve

Afferent nerve fiber

Efferent nerve fiber

Sensory neuron

References

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Last edited on 10 January 2024, at 20:13

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