Motor neuron

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Motor neurons
Medulla oblongata - posterior - cn xii - very high mag.jpg
Micrograph of the hypoglossal nucleus showing motor neurons with their characteristic coarse Nissl substance ("tigroid" cytoplasm). H&E-LFB stain.
Details
Location Ventral horn of the spinal cord, some cranial nerve nuclei
ShapeProjection neuron
FunctionExcitatory projection (to NMJ)
Neurotransmitter UMN to LMN: glutamate; LMN to NMJ: ACh
Presynaptic connections Primary motor cortex via the Corticospinal tract
Postsynaptic connections Muscle fibers and other neurons
Identifiers
MeSH D009046
NeuroLex ID nifext_103
TA98 A14.2.00.021
TA2 6131
FMA 83617
Anatomical terms of neuroanatomy

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.

Contents

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. 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]

Although the word "motor neuron" suggests that there is a single kind of neuron that controls movement, this is not the case. Indeed, upper and lower motor neurons—which differ greatly in their origins, synapse locations, routes, neurotransmitters, and lesion characteristics—are included in the same classification as "motor neurons." Essentially, motor neurons, also known as motoneurons, are made up of a variety of intricate, finely tuned circuits found throughout the body that innervate effector muscles and glands to enable both voluntary and involuntary motions. Two motor neurons come together to form a two-neuron circuit. While lower motor neurons start in the spinal cord and go to innervate muscles and glands all throughout the body, upper motor neurons originate in the cerebral cortex and travel to the brain stem or spinal cord. It is essential to comprehend the distinctions between upper and lower motor neurons as well as the routes they follow in order to effectively detect these neuronal injuries and localise the lesions. [6]

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 columnLocation in spinal cordTarget
Median motor columnPresent entire lengthAxial muscles
Hypaxial motor columnThoracic regionBody wall muscles
Preganglionic motor columnThoracic regionSympathetic ganglion
Lateral motor columnBrachial and lumbar region (both regions are further divided into medial and lateral domains)Muscles of the limbs
Phrenic motor columnCervical regionDiaphragm [11]

Anatomy and physiology

Spinal cord tracts Spinal cord tracts - English.svg
Spinal cord tracts
Location of lower motor neurons in spinal cord Polio spinal diagram-en.svg
Location of lower motor neurons in spinal cord

Upper motor neurons

Upper motor neurons originate in the motor cortex located in the precentral gyrus. The cells that make up the primary motor 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 control of individual fingers. [14] [15] Corticomotorneurons have so 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]

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

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 NSPositionNeurotransmitter
Somaticn/a Acetylcholine
ParasympatheticPreganglionicAcetylcholine
ParasympatheticGanglionicAcetylcholine
SympatheticPreganglionicAcetylcholine
SympatheticGanglionic 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

Related Research Articles

<span class="mw-page-title-main">Nerve</span> Enclosed, cable-like bundle of axons in the peripheral nervous system

A nerve is an enclosed, cable-like bundle of nerve fibers in the peripheral nervous system.

<span class="mw-page-title-main">Motor nerve</span> Nerve located in the central nervous system

A motor nerve is a nerve that transmits motor signals from the central nervous system (CNS) to the muscles of the body. This is different from the motor neuron, which includes a cell body and branching of dendrites, while the nerve is made up of a bundle of axons. Motor nerves act as efferent nerves which carry information out from the CNS to muscles, as opposed to afferent nerves, which transfer signals from sensory receptors in the periphery to the CNS. Efferent nerves can also connect to glands or other organs/issues instead of muscles. In addition, there are nerves that serve as both sensory and motor nerves called mixed nerves.

<span class="mw-page-title-main">Somatic nervous system</span> Part of the peripheral nervous system

The somatic nervous system (SNS), or voluntary nervous system is the part of the peripheral nervous system associated with the voluntary control of body movements via skeletal muscles.

<span class="mw-page-title-main">Muscle spindle</span> Innervated muscle structure involved in reflex actions and proprioception

Muscle spindles are stretch receptors within the body of a skeletal muscle that primarily detect changes in the length of the muscle. They convey length information to the central nervous system via afferent nerve fibers. This information can be processed by the brain as proprioception. The responses of muscle spindles to changes in length also play an important role in regulating the contraction of muscles, for example, by activating motor neurons via the stretch reflex to resist muscle stretch.

<span class="mw-page-title-main">Afferent nerve fiber</span> Axonal projections that arrive at a particular brain region

Afferent nerve fibers are axons of sensory neurons that carry sensory information from sensory receptors to the central nervous system. Many afferent projections arrive at a particular brain region.

<span class="mw-page-title-main">Grey column</span>

The grey column refers to a somewhat ridge-shaped mass of grey matter in the spinal cord. This presents as three columns: the anterior grey column, the posterior grey column, and the lateral grey column, all of which are visible in cross-section of the spinal cord.

<span class="mw-page-title-main">Reflex arc</span> Neural pathway which controls a reflex

A reflex arc is a neural pathway that controls a reflex. In vertebrates, most sensory neurons do not pass directly into the brain, but synapse in the spinal cord. This allows for faster reflex actions to occur by activating spinal motor neurons without the delay of routing signals through the brain. The brain will receive the input while the reflex is being carried out and the analysis of the signal takes place after the reflex action.

<span class="mw-page-title-main">Pyramidal tracts</span> Include both the corticobulbar tract and the corticospinal tract

The pyramidal tracts include both the corticobulbar tract and the corticospinal tract. These are aggregations of efferent nerve fibers from the upper motor neurons that travel from the cerebral cortex and terminate either in the brainstem (corticobulbar) or spinal cord (corticospinal) and are involved in the control of motor functions of the body.

<span class="mw-page-title-main">Dorsal column–medial lemniscus pathway</span> Sensory spinal pathway

The dorsal column–medial lemniscus pathway (DCML) is a sensory pathway of the central nervous system that conveys sensations of fine touch, vibration, two-point discrimination, and proprioception from the skin and joints. It transmits information from the body to the primary somatosensory cortex in the postcentral gyrus of the parietal lobe of the brain. The pathway receives information from sensory receptors throughout the body, and carries this in nerve tracts in the white matter of the dorsal column of the spinal cord to the medulla, where it is continued in the medial lemniscus, on to the thalamus and relayed from there through the internal capsule and transmitted to the somatosensory cortex. The name dorsal-column medial lemniscus comes from the two structures that carry the sensory information: the dorsal columns of the spinal cord, and the medial lemniscus in the brainstem.

<span class="mw-page-title-main">Nuclear chain fiber</span> Specialized sensory organ within a muscle

A nuclear chain fiber is a specialized sensory organ contained within a muscle. Nuclear chain fibers are intrafusal fibers that, along with nuclear bag fibers, make up the muscle spindle responsible for the detection of changes in muscle length.

<span class="mw-page-title-main">Type Ia sensory fiber</span> Type of afferent nerve fiber

A type Ia sensory fiber, or a primary afferent fiber is a type of afferent nerve fiber. It is the sensory fiber of a stretch receptor called the muscle spindle found in muscles, which constantly monitors the rate at which a muscle stretch changes. The information carried by type Ia fibers contributes to the sense of proprioception.

<span class="mw-page-title-main">Upper motor neuron</span> Neurons in the brain that carry signals to lower motor neurons

Upper motor neurons (UMNs) is a term introduced by William Gowers in 1886. They are found in the cerebral cortex and brainstem and carry information down to activate interneurons and lower motor neurons, which in turn directly signal muscles to contract or relax. UMNs represent the major origin point for voluntary somatic movement.

<span class="mw-page-title-main">Gamma motor neuron</span>

A gamma motor neuron, also called gamma motoneuron, or fusimotor neuron, is a type of lower motor neuron that takes part in the process of muscle contraction, and represents about 30% of (Aγ) fibers going to the muscle. Like alpha motor neurons, their cell bodies are located in the anterior grey column of the spinal cord. They receive input from the reticular formation of the pons in the brainstem. Their axons are smaller than those of the alpha motor neurons, with a diameter of only 5 μm. Unlike the alpha motor neurons, gamma motor neurons do not directly adjust the lengthening or shortening of muscles. However, their role is important in keeping muscle spindles taut, thereby allowing the continued firing of alpha neurons, leading to muscle contraction. These neurons also play a role in adjusting the sensitivity of muscle spindles.

Extrafusal muscle fibers are the standard skeletal muscle fibers that are innervated by alpha motor neurons and generate tension by contracting, thereby allowing for skeletal movement. They make up the large mass of skeletal striated muscle tissue and are attached to bone by fibrous tissue extensions (tendons).

<span class="mw-page-title-main">Alpha motor neuron</span>

Alpha (α) motor neurons (also called alpha motoneurons), are large, multipolar lower motor neurons of the brainstem and spinal cord. They innervate extrafusal muscle fibers of skeletal muscle and are directly responsible for initiating their contraction. Alpha motor neurons are distinct from gamma motor neurons, which innervate intrafusal muscle fibers of muscle spindles.

<span class="mw-page-title-main">Spinal cord</span> Long, tubular central nervous system structure in the vertebral column

The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal cord is hollow and contains a structure called central canal, which contains cerebrospinal fluid. The spinal cord is also covered by meninges and enclosed by the neural arches. Together, the brain and spinal cord make up the central nervous system (CNS).

A motor pool consists of all individual motor neurons that innervate a single muscle. Each individual muscle fiber is innervated by only one motor neuron, but one motor neuron may innervate several muscle fibers. This distinction is physiologically significant because the size of a given motor pool determines the activity of the muscle it innervates: for example, muscles responsible for finer movements are innervated by motor pools consisting of higher numbers of individual motor neurons. Motor pools are also distinguished by the different classes of motor neurons that they contain. The size, composition, and anatomical location of each motor pool is tightly controlled by complex developmental pathways.

Beta motor neurons, also called beta motoneurons, are a kind of lower motor neuron, along with alpha motor neurons and gamma motor neurons. Beta motor neurons innervate intrafusal fibers of muscle spindles with collaterals to extrafusal fibers - a type of slow twitch fiber. Also, axons of alpha, beta, and gamma motor neurons become myelinated. Moreover, these efferent neurons originate from the anterior grey column of the spinal cord and travel to skeletal muscles. However, the larger diameter alpha motor fibers require higher conduction velocity than beta and gamma.

<span class="mw-page-title-main">Spinal interneuron</span> Interneuron relaying signals between sensory and motor neurons in the spinal cord

A spinal interneuron, found in the spinal cord, relays signals between (afferent) sensory neurons, and (efferent) motor neurons. Different classes of spinal interneurons are involved in the process of sensory-motor integration. Most interneurons are found in the grey column, a region of grey matter in the spinal cord.

<span class="mw-page-title-main">Golgi tendon organ</span> Proprioceptive sensory receptor organ that senses changes in muscle tension

The Golgi tendon organ (GTO) is a proprioceptor – a type of sensory receptor that senses changes in muscle tension. It lies at the interface between a muscle and its tendon known as the musculotendinous junction also known as the myotendinous junction. It provides the sensory component of the Golgi tendon reflex.

References

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  11. ↑ Philippidou, Polyxeni; Walsh, Carolyn; Aubin, Josée; Jeannotte, Lucie; Dasen, Jeremy S. (2012). "Sustained Hox5 Gene Activity is Required for Respiratory Motor Neuron Development". Nature Neuroscience. 15 (12): 1636–1644. doi:10.1038/nn.3242. ISSN   1097-6256. PMC   3676175 . PMID   23103965.
  12. ↑ Fitzpatrick, D. (2001) The Primary Motor Cortex: Upper Motor Neurons That Initiate Complex Voluntary Movements. In D. Purves, G.J. Augustine, D. Fitzpatrick, et al. (Ed.), Neuroscience. Retrieved from "The Primary Motor Cortex: Upper Motor Neurons That Initiate Complex Voluntary Movements - Neuroscience - NCBI Bookshelf". Archived from the original on 2018-06-05. Retrieved 2017-11-30.
  13. 1 2 Mack, Sarah; Kandel, Eric R.; Jessell, Thomas M.; Schwartz, James H.; Siegelbaum, Steven A.; Hudspeth, A. J. (2013). Principles of neural science. Kandel, Eric R. (5th ed.). New York. ISBN   9780071390118. OCLC   795553723.{{cite book}}: CS1 maint: location missing publisher (link)
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  22. 1 2 3 4 5 Manuel, Marin; Zytnicki, Daniel (2011). "Alpha, Beta, and Gamma Motoneurons: Functional Diversity in the Motor System's Final Pathway". Journal of Integrative Neuroscience. 10 (3): 243–276. doi:10.1142/S0219635211002786. ISSN   0219-6352. PMID   21960303. S2CID   21582283.
  23. ↑ Azevedo, Anthony; Lesser, Ellen; Mark, Brandon; Phelps, Jasper; Elabbady, Leila; Kuroda, Sumiya; Sustar, Anne; Moussa, Anthony; Kandelwal, Avinash; Dallmann, Chris J.; Agrawal, Sweta; Lee, Su-Yee J.; Pratt, Brandon; Cook, Andrew; Skutt-Kakaria, Kyobi (2022-12-15). "Tools for comprehensive reconstruction and analysis of Drosophila motor circuits": 2022.12.15.520299. doi:10.1101/2022.12.15.520299. S2CID   254736092.{{cite journal}}: Cite journal requires |journal= (help)

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