Muscle spindle

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Muscle spindle
Muscle spindle model.jpg
Mammalian muscle spindle showing typical position in a muscle (left), neuronal connections in spinal cord (middle) and expanded schematic (right). The spindle is a stretch receptor with its own motor supply consisting of several intrafusal muscle fibres. The sensory endings of a primary (group Ia) afferent and a secondary (group II) afferent coil around the non-contractile central portions of the intrafusal fibres. Gamma motor neurons activate the intrafusal muscle fibres, changing the resting firing rate and stretch-sensitivity of the afferents. [lower-alpha 1]
Details
Part of Muscle
System Musculoskeletal
Identifiers
Latin fusus neuromuscularis
MeSH D009470
TH H3.11.06.0.00018
FMA 83607
Anatomical terminology

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.

Contents

The muscle spindle has both sensory and motor components.

Structure

Muscle spindles are found within the belly of a skeletal muscle. Muscle spindles are fusiform (spindle-shaped), and the specialized fibers that make up the muscle spindle are called intrafusal muscle fibers. The regular muscle fibers outside of the spindle are called extrafusal muscle fibers. Muscle spindles have a capsule of connective tissue, and run parallel to the extrafusal muscle fibers. [lower-alpha 2]

Composition

Muscle spindles are composed of 5-14 muscle fibers, of which there are three types: dynamic nuclear bag fibers (bag1 fibers), static nuclear bag fibers (bag2 fibers), and nuclear chain fibers. [1] [2]

Light microscope photograph of a muscle spindle. HE stain. Muscle Spindle LM HE stain.jpg
Light microscope photograph of a muscle spindle. HE stain.

Primary type Ia sensory fibers (large diameter) spiral around all intrafusal muscle fibres, ending near the middle of each fibre. Secondary type II sensory fibers (medium diameter) end adjacent to the central regions of the static bag and chain fibres. [2] These fibres send information by stretch-sensitive mechanically-gated ion-channels of the axons. [3]

The motor part of the spindle is provided by motor neurons: up to a dozen gamma motor neurons also known as fusimotor neurons. [4] These activate the muscle fibres within the spindle. Gamma motor neurons supply only muscle fibres within the spindle, whereas beta motor neurons supply muscle fibres both within and outside of the spindle. Activation of the neurons causes a contraction and stiffening of the end parts of the muscle spindle muscle fibers.

Fusimotor neurons are classified as static or dynamic according to the type of muscle fibers they innervate and their effects on the responses of the Ia and II sensory neurons innervating the central, non-contractile part of the muscle spindle.

Efferent nerve fibers of gamma motor neurons also terminate in muscle spindles; they make synapses at either or both of the ends of the intrafusal muscle fibers and regulate the sensitivity of the sensory afferents, which are located in the non-contractile central (equatorial) region. [5]

Function

Stretch reflex

When a muscle is stretched, primary type Ia sensory fibers of the muscle spindle respond to both changes in muscle length and velocity and transmit this activity to the spinal cord in the form of changes in the rate of action potentials. Likewise, secondary type II sensory fibers respond to muscle length changes (but with a smaller velocity-sensitive component) and transmit this signal to the spinal cord. The Ia afferent signals are transmitted monosynaptically to many alpha motor neurons of the receptor-bearing muscle. The reflexly evoked activity in the alpha motor neurons is then transmitted via their efferent axons to the extrafusal fibers of the muscle, which generate force and thereby resist the stretch. The Ia afferent signal is also transmitted polysynaptically through interneurons (Ia inhibitory interneurons), which inhibit alpha motorneurons of antagonist muscles, causing them to relax.

Sensitivity modification

The function of the gamma motor neurons is not to supplement the force of muscle contraction provided by the extrafusal fibers, but to modify the sensitivity of the muscle spindle sensory afferents to stretch. Upon release of acetylcholine by the active gamma motor neuron, the end portions of the intrafusal muscle fibers contract, thus elongating the non-contractile central portions (see "fusimotor action" schematic below). This opens stretch-sensitive ion channels of the sensory endings, leading to an influx of sodium ions. This raises the resting potential of the endings, thereby increasing the probability of action potential firing, thus increasing the stretch-sensitivity of the muscle spindle afferents.

How does the central nervous system control gamma fusimotor neurons? It has been difficult to record from gamma motor neurons during normal movement because they have very small axons. Several theories have been proposed, based on recordings from spindle afferents.

Development

It is also believed that muscle spindles play a critical role in sensorimotor development.

Clinical significance

After stroke or spinal cord injury in humans, spastic hypertonia (spastic paralysis) often develops, whereby the stretch reflex in flexor muscles of the arms and extensor muscles of the legs is overly sensitive. This results in abnormal postures, stiffness and contractures. Hypertonia may be the result of over-sensitivity of alpha motor neurons and interneurons to the Ia and II afferent signals. [10]

Additional images

Notes

  1. Animated version: https://www.ualberta.ca/~aprochaz/research_interactive_receptor_model.html Arthur Prochazka's Lab, University of Alberta
  2. unlike Golgi tendon organs, which are oriented in series

Related Research Articles

<span class="mw-page-title-main">Motor neuron</span> Nerve cell sending impulse to muscle

A motor neuron 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. 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. The axons from the lower motor neurons are efferent nerve fibers that carry signals from the spinal cord to the effectors. Types of lower motor neurons are alpha motor neurons, beta motor neurons, and gamma motor neurons.

<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">Efferent nerve fiber</span> Axonal projections that exit a particular region

Efferent nerve fibers refer to axonal projections that exit a particular region; as opposed to afferent projections that arrive at the region. These terms have a slightly different meaning in the context of the peripheral nervous system (PNS) and central nervous system (CNS). The efferent fiber is a long process projecting far from the neuron's body that carries nerve impulses away from the central nervous system toward the peripheral effector organs. A bundle of these fibers is called an efferent nerve. The opposite direction of neural activity is afferent conduction, which carries impulses by way of the afferent nerve fibers of sensory neurons.

<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.

A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, are sent to the central nervous system.

<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.

Renshaw cells are inhibitory interneurons found in the gray matter of the spinal cord, and are associated in two ways with an alpha motor neuron.

<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.

Lower motor neurons (LMNs) are motor neurons located in either the anterior grey column, anterior nerve roots or the cranial nerve nuclei of the brainstem and cranial nerves with motor function. Many voluntary movements rely on spinal lower motor neurons, which innervate skeletal muscle fibers and act as a link between upper motor neurons and muscles. Cranial nerve lower motor neurons also control some voluntary movements of the eyes, face and tongue, and contribute to chewing, swallowing and vocalization. Damage to the lower motor neurons can lead to flaccid paralysis, absent deep tendon reflexes and muscle atrophy.

<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.

<span class="mw-page-title-main">Intrafusal muscle fiber</span> Skeletal muscle fibers

Intrafusal muscle fibers are skeletal muscle fibers that serve as specialized sensory organs (proprioceptors). They detect the amount and rate of change in length of a muscle. They constitute the muscle spindle, and are innervated by both sensory (afferent) and motor (efferent) fibers.

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">Stretch reflex</span> Muscle contraction in response to stretching

The stretch reflex, or more accurately "muscle stretch reflex", is a muscle contraction in response to stretching within the muscle. The reflex functions to maintain the muscle at a constant length. The term deep tendon reflex is often wrongfully used by many health workers and students to refer to this reflex. "Tendons have little to do with the response, other than being responsible for mechanically transmitting the sudden stretch from the reflex hammer to the muscle spindle. In addition, some muscles with stretch reflexes have no 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.

The Golgi tendon reflex (also called inverse stretch reflex, autogenic inhibition, tendon reflex) is an inhibitory effect on the muscle resulting from the muscle tension stimulating Golgi tendon organs (GTO) of the muscle, and hence it is self-induced. The reflex arc is a negative feedback mechanism preventing too much tension on the muscle and tendon. When the tension is extreme, the inhibition can be so great it overcomes the excitatory effects on the muscle's alpha motoneurons causing the muscle to suddenly relax. This reflex is also called the inverse myotatic reflex, because it is the inverse of the stretch reflex.

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.

Group A nerve fibers are one of the three classes of nerve fiber as generally classified by Erlanger and Gasser. The other two classes are the group B nerve fibers, and the group C nerve fibers. Group A are heavily myelinated, group B are moderately myelinated, and group C are unmyelinated.

<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.

Proprioception refers to the sensory information relayed from muscles, tendons, and skin that allows for the perception of the body in space. This feedback allows for more fine control of movement. In the brain, proprioceptive integration occurs in the somatosensory cortex, and motor commands are generated in the motor cortex. In the spinal cord, sensory and motor signals are integrated and modulated by motor neuron pools called central pattern generators (CPGs). At the base level, sensory input is relayed by muscle spindles in the muscle and Golgi tendon organs (GTOs) in tendons, alongside cutaneous sensors in the skin.

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