Muscle spindle

Muscle spindles are stretch receptors within the body of a 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, by activating motor neurons via the stretch reflex to resist muscle stretch.

The muscle spindle has both sensory and motor components.

Muscle spindle
Muscle spindle model
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 motoneurons activate the intrafusal muscle fibres, changing the resting firing rate and stretch-sensitivity of the afferents. [a]
Details
Part ofMuscle
SystemMusculoskeletal
Identifiers
Latinfusus neuromuscularis
MeSHD009470
THH3.11.06.0.00018
FMA83607
Anatomical terminology

Structure

Muscle spindles are found within the belly of muscles, between extrafusal muscle fibers.[b] The specialised fibers that constitute the muscle spindle are known as intrafusal fibers (as they are present within the spindle), to distinguish themselves from the fibres of the muscle itself which are called extrafusal fibers. Muscle spindles have a capsule of connective tissue, and run parallel to the extrafusal muscle fibers.[c]

Composition

Muscle spindles are composed of three to twelve muscle fibers, of which there are three types: dynamic nuclear bag fibers (bag1 fibers), static nuclear bag fibers (bag2 fibers), nuclear chain fibers and afferent nerve fibers.

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

Sensory fibres spiral around the intrafusal muscle fibres, ending near the middle of each fibre. These fibres, primary type Ia sensory fibers and secondary type II sensory fibers, send information by stretch-sensitive ion-channels of the axons.

The motor part of the spindle is provided by motor neurons: up to a dozen gamma motor neurons and one or two beta motor neurons, collectively called fusimotor neurons. 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.

  • The static axons innervate the chain or bag2 fibers. They increase the firing rate of Ia and II afferents at a given muscle length (see schematic of fusimotor action below).
  • The dynamic axons innervate the bag1 intrafusal muscle fibers. They increase the stretch-sensitivity of the Ia afferents by stiffening the bag1 intrafusal fibers.

Efferent nerve fibers of gamma motoneurons 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.[1]

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 motoneurons 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 motoneurons during normal movement because they have very small axons. Several theories have been proposed, based on recordings from spindle afferents.

  • 1) Alpha-gamma coactivation. Here it is posited that gamma motoneurons are activated in parallel with alpha motoneurons to maintain the firing of spindle afferents when the extrafusal muscles shorten.[2]
  • 2) Fusimotor set: Gamma motoneurons are activated according to the novelty or difficulty of a task. Whereas static gamma motoneurons are continuously active during routine movements such as locomotion, dynamic gamma motoneoruns tend to be activated more during difficult tasks, increasing Ia stretch-sensitivity.[3]
  • 3) Fusimotor template of intended movement. Static gamma activity is a "temporal template" of the expected shortening and lengthening of the receptor-bearing muscle. Dynamic gamma activity turns on and off abruptly, sensitizing spindle afferents to the onset of muscle lengthening and departures from the intended movement trajectory.[4]

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 motoneurons and interneurons to the Ia and II afferent signals.[5]

Additional images

MuscleSpindle

Muscle spindle

Muskelspindel3

Gamma fiber

Muskelspindel4

1A fiber

Muskelspindel5

Alpha fiber

Fusimotor action

schematic of fusimotor action

See also

Notes

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

References

  1. ^ Hulliger M (1984). "The mammalian muscle spindle and its central control". Rev. Physiol. Biochem. Pharmacol. 101: 1–110. doi:10.1007/bfb0027694. PMID 6240757.
  2. ^ Vallbo AB, al-Falahe NA (February 1990). "Human muscle spindle response in a motor learning task". J. Physiol. 421: 553–68. doi:10.1113/jphysiol.1990.sp017961. PMC 1190101. PMID 2140862.
  3. ^ Prochazka, A. (1996). "Proprioceptive feedback and movement regulation". In Rowell, L.; Sheperd, J.T. Exercise: Regulation and Integration of Multiple Systems. Handbook of physiology. New York: American Physiological Society. pp. 89–127. ISBN 0195091744.
  4. ^ Taylor A, Durbaba R, Ellaway PH, Rawlinson S (March 2006). "Static and dynamic gamma-motor output to ankle flexor muscles during locomotion in the decerebrate cat". J. Physiol. 571 (Pt 3): 711–23. doi:10.1113/jphysiol.2005.101634. PMC 1805796. PMID 16423858.
  5. ^ Heckmann CJ, Gorassini MA, Bennett DJ (February 2005). "Persistent inward currents in motoneuron dendrites: implications for motor output". Muscle Nerve. 31 (2): 135–56. doi:10.1002/mus.20261. PMID 15736297.

External links

Ankle

The ankle, or the talocrural region, is the region where the foot and the leg meet. The ankle includes three joints: the ankle joint proper or talocrural joint, the subtalar joint, and the inferior tibiofibular joint. The movements produced at this joint are dorsiflexion and plantarflexion of the foot. In common usage, the term ankle refers exclusively to the ankle region. In medical terminology, "ankle" (without qualifiers) can refer broadly to the region or specifically to the talocrural joint.The main bones of the ankle region are the talus (in the foot), and the tibia and fibula (in the leg). The talocrural joint is a synovial hinge joint that connects the distal ends of the tibia and fibula in the lower limb with the proximal end of the talus. The articulation between the tibia and the talus bears more weight than that between the smaller fibula and the talus.

Extrafusal muscle fiber

Extrafusal muscle fibers are the skeletal standard 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 muscle tissue and are attached to bone by fibrous tissue extensions (tendons).

Each alpha motor neuron and the extrafusal muscle fibers innervated by it make up a motor unit. The connection between the alpha motor neuron and the extrafusal muscle fiber is a neuromuscular junction, where the neuron's signal, the action potential, is transduced to the muscle fiber by the neurotransmitter acetylcholine.

Extrafusal muscle fibers are not to be confused with intrafusal muscle fibers, which are innervated by sensory nerve endings in central noncontractile parts and by gamma motor neurons in contractile ends and thus serve as a sensory proprioceptor.

Extrafusal muscle fibers can be generated in vitro (in a dish) from pluripotent stem cells through directed differentiation. This allows study of their formation and physiology.

Gamma motor neuron

A gamma motor neuron (γ motor neuron), also called gamma motoneuron, 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.The presence of myelination in gamma motor neurons allows a conduction velocity of 4 to 24 meters per second, significantly faster than with non-myelinated axons but slower than in alpha motor neurons.

Ganglion

A ganglion is a nerve cell cluster or a group of nerve cell bodies located in the autonomic nervous system and sensory system. Ganglia house the cell bodies of afferent nerves (input nerve fibers) and efferent nerves (output/motor nerve fibers), or axons.

A pseudoganglion looks like a ganglion, but only has nerve fibers and has no nerve cell bodies.

Group A nerve fiber

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.The other classification is a sensory grouping that uses the terms type Ia and type Ib, type II, type III, and type IV, sensory fibers.

H-reflex

The H-reflex (or Hoffmann's reflex) is a reflectory reaction of muscles after electrical stimulation of sensory fibers (Ia afferents stemming from muscle spindles) in their innervating nerves (for example, those located behind the knee). The H-reflex test is performed using an electric stimulator, which gives usually a square-wave current of short duration and small amplitude (higher stimulations might involve alpha fibers, causing an F-wave, compromising the results), and an EMG set, to record the muscle response. That response is usually a clear wave, called H-wave, 28-35 ms after the stimulus, not to be confused with an F-wave. An M-wave, an early response, occurs 3-6 ms after the onset of stimulation. The H and F-waves are later responses. As the stimulus increases, the amplitude of the F-wave increases only slightly, and the H-wave decreases, and at supramaximal stimulus, the H-wave will disappear. The M-wave does the opposite of the H-wave. As the stimulus increases the M-wave increases. There is a point of minimal stimulus where the M-wave is absent and the H-wave is maximal.

H-reflex is analogous to the mechanically induced spinal stretch reflex (for example, knee jerk reflex). "The primary difference between the H-reflex and the spinal stretch reflex is that the H-reflex bypasses the muscle spindle, and, therefore, is a valuable tool in assessing modulation of monosynaptic reflex activity in the spinal cord." Although stretch reflex gives just qualitative information about muscle spindles and reflex arc activity; if the purpose of the test to compare performances from different subjects, H-reflex should be used. In that case, in fact, latencies (ms) and amplitudes (mV) of H-wave can be compared.

H-reflex amplitudes measured by EMG are shown to decrease significantly with applied pressure such as massage and tapping to the cited muscle. The amount of decrease seems to be dependent on the force of the pressure, with higher pressures resulting in lower H-reflex amplitudes. H-reflex levels return to baseline immediately after pressure is released except in high pressure cases which had baseline levels returned within the first 10 seconds.After about 5 days in zero gravity, for instance in orbit around Earth, the h-reflex diminishes significantly. It is generally assumed that this is due to a marked reduction in the excitability of the spinal cord in zero gravity. Once back on Earth, a marked recovery occurs during the first day, but it can take up to 10 days to return to normal. The H-reflex was the first medical experiment completed on the International Space Station.

Hypotonia

Hypotonia, commonly known as floppy baby syndrome, is a state of low muscle tone (the amount of tension or resistance to stretch in a muscle), often involving reduced muscle strength. Hypotonia is not a specific medical disorder, but a potential manifestation of many different diseases and disorders that affect motor nerve control by the brain or muscle strength. Hypotonia is resistance to passive movement, whereas muscle weakness results in impaired active movement. Central hypotonia originates from the central nervous system, while peripheral hypotonia is related to problems within the spinal cord, peripheral nerves and/or skeletal muscles. Recognizing hypotonia, even in early infancy, is usually relatively straightforward, but diagnosing the underlying cause can be difficult and often unsuccessful. The long-term effects of hypotonia on a child's development and later life depend primarily on the severity of the muscle weakness and the nature of the cause. Some disorders have a specific treatment but the principal treatment for most hypotonia of idiopathic or neurologic cause is physical therapy, occupational therapy for remediation, and/or music therapy.

Hypotonia is thought to be associated with the disruption of afferent input from stretch receptors and/or lack of the cerebellum’s facilitatory efferent influence on the fusimotor system, the system that innervates intrafusal muscle fibers thereby controlling muscle spindle sensitivity. On examination a diminished resistance to passive movement will be noted and muscles may feel abnormally soft and limp on palpation. Diminished deep tendon reflexes also may be noted. Hypotonia is a condition that can be helped with early intervention.

Intrafusal muscle fiber

Intrafusal muscle fibers are skeletal muscle fibers that serve as specialized sensory organs (proprioceptors) that detect the amount and rate of change in length of a muscle. They constitute the muscle spindle and are innervated by two axons, one sensory and one motor. Gamma effrents from small multipolar cells from anterior gray coloumn innervate it. These form a part of neuromuscular spindles. Intrafusal muscle fibers are walled off from the rest of the muscle by a collagen sheath. This sheath has a spindle or "fusiform" shape, hence the name "intrafusal".

There are two types of intrafusal muscle fibers: nuclear bag and nuclear chain fibers. They bear two types of sensory ending, known as annulospiral and flower-spray endings. Both ends of these fibers contract but the central region only stretches and does not contract.

They are innervated by gamma motor neurons and beta motor neurons.

It is by the sensory information from these two intrafusal fiber types that an individual is able to judge the position of their muscle, and the rate at which it is changing.

Intrafusal muscle fibers are not to be confused with extrafusal muscle fibers, which contract, generating skeletal movement and are innervated by alpha motor neurons.

Nuclear bag fiber

A nuclear bag fiber is a type of intrafusal muscle fiber that lies in the center of a muscle spindle. Each has a large number of nuclei concentrated in bags and they cause excitation of both the primary and secondary nerve fibers.

There are two kinds of bag fibers based upon contraction speed and motor innervation.

BAG2 fibers are the largest. They have no striations in middle region and swell to enclose nuclei, hence their name.

BAG1 fibers, smaller than BAG2.Both bag types extend beyond the spindle capsule.

These sense dynamic length of the muscle. They are sensitive to length & velocity.

Nuclear chain fiber

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.

There are 3–9 nuclear chain fibers per muscle spindle that are half the size of the nuclear bag fibers. Their nuclei are aligned in a chain and they excite the secondary nerve. They are static, whereas the nuclear bag fibers are dynamic in comparison. The name "nuclear chain" refers to the structure of the central region of the fiber, where the sensory axons wrap around the intrafusal fibers.

The secondary nerve association involves an efferent and afferent pathway that measure the stress and strain placed on the muscle (usually the extrafusal fibers connected from the muscle portion to a bone). The afferent pathway resembles a spring wrapping around the nuclear chain fiber and connecting to one of its ends away from the bone. Again, depending on the stress and strain the muscles sustains, this afferent and efferent coordination will measure the "stretch of the spring" and communicate the results to the central nervous system.

A similar structure attaching one end to muscle and the other end to a tendon is known as a Golgi tendon organ. However, Golgi tendon organs differ from nuclear chain and nuclear bag fibers in that they are considered in series rather than in parallel to the muscle fibers.

Reciprocal inhibition

Reciprocal inhibition describes the process of muscles on one side of a joint relaxing to accommodate contraction on the other side of that joint. In some allied health disciplines this is known as reflexive antagonism. Joints are controlled by two opposing sets of muscles, extensors and flexors, which must work in synchrony for smooth movement. When a muscle spindle is stretched and the stretch reflex is activated, the opposing muscle group must be inhibited to prevent it from working against the resulting contraction of the homonymous muscle. This inhibition is accomplished by the actions of an inhibitory interneuron in the spinal cord.

The afferent of the muscle spindle bifurcates in the spinal cord. One branch innervates the alpha motor neuron that causes the homonymous muscle to contract, producing the reflex. The other branch innervates the inhibitory interneuron, which in turn innervates the alpha motor neuron that synapses onto the opposing muscle. Because the interneuron is inhibitory, it prevents the opposing alpha motor neuron from firing, thereby reducing the contraction of the opposing muscle. Without this reciprocal inhibition, both groups of muscles might contract simultaneously and work against each other.

If opposing muscles were to contract at the same time, a muscle tear can occur. This may occur during physical activities, such as running, during which muscles that oppose each other are engaged and disengaged sequentially to produce coordinated movement. Reciprocal inhibition facilitates ease of movement and is a safeguard against injury. However, if a "misfiring" of motor neurons occurs, causing simultaneous contraction of opposing muscles, a tear can occur. For example, if the quadriceps femoris and hamstring contract simultaneously at a high intensity, the stronger muscle (traditionally the quadriceps) overpowers the weaker muscle group (hamstrings). This can result in a common muscular injury known as a pulled hamstring, more accurately called a muscle strain.

"When the central nervous system sends a message to the agonist muscle (muscle causing movement) to contract, the tension in the antagonist muscle (muscle opposing movement) is inhibited by impulses from motor neurons, and thus must simultaneously relax. This neural phenomenon is called reciprocal inhibition."

Taken from Massage Therapy Principles & Practices by Susan Salvo 1999, pg 161

Reflex arc

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. However, the brain will receive the sensory input while the reflex is being carried out and the analysis of the signal takes place after the reflex action.

There are two types: autonomic reflex arc (affecting inner organs) and somatic reflex arc (affecting muscles). However, autonomic reflexes sometimes involve the spinal cord and some somatic reflexes are mediated more by the brain than the spinal cord.During a somatic reflex, nerve signals travel along the following pathway:

Somatic receptors in the skin, muscles and tendons

Afferent nerve fibers carry signals from the somatic receptors to the posterior horn of the spinal cord or to the brainstem

An integrating center, the point at which the neurons that compose the gray matter of the spinal cord or brainstem synapse

Efferent nerve fibers carry motor nerve signals from the anterior horn to the muscles

Effector muscle innervated by the efferent nerve fiber carries out the response.A reflex arc, then, is the pathway followed by nerves which (a.) carry sensory information from the receptor to the spinal cord, and then (b) carry the response generated by the spinal cord to effector organ(s) during a reflex action.

Secondary sensory endings

Secondary sensory endings of the muscle spindle are composed of type II sensory fibers that terminate on nuclear chain fibers and static nuclear bag fibers, but not dynamic nuclear bag fibers. Whereas primary endings respond mostly to rate of change, secondary endings respond mostly to amount of stretch.

Spinocerebellar tract

The spinocerebellar tract is a nerve tract originating in the spinal cord and terminating in the same side (ipsilateral) of the cerebellum.

Stretch receptor

Stretch receptors are mechanoreceptors responsive to distention of various organs and muscles, and are neurologically linked to the medulla in the brain stem via afferent nerve fibers. Examples include stretch receptors in the arm and leg muscles and tendons, in the heart, in the colon wall, and in the lungs.Stretch receptors are also found around the carotid artery, where they monitor blood pressure and stimulate the release of antidiuretic hormone (ADH) from the posterior pituitary gland.

Types include:

Golgi organ

Muscle spindle, sensory receptors within the belly of a muscle, which primarily detect changes in the length of this muscle

Pulmonary stretch receptors, mechanoreceptors found in the lungs

Chordotonal organ, in insects

Stretch reflex

The stretch reflex (myotatic reflex) is a muscle contraction in response to stretching within the muscle. It is a monosynaptic reflex which provides automatic regulation of skeletal muscle length.

When a muscle lengthens, the muscle spindle is stretched and its nerve activity increases. This increases alpha motor neuron activity, causing the muscle fibers to contract and thus resist the stretching. A secondary set of neurons also causes the opposing muscle to relax. The reflex functions to maintain the muscle at a constant length.

Gamma motoneurons regulate how sensitive the stretch reflex is by tightening or relaxing the fibers within the spindle. There are several theories as to what may trigger gamma motoneurons to increase the reflex's sensitivity. For example, alpha-gamma co-activation might keep the spindles taut when a muscle is contracted, preserving stretch reflex sensitivity even as the muscle fibers become shorter. Otherwise the spindles would become slack and the reflex would cease to function.

This reflex has the shortest latency of all spinal reflexes including the Golgi tendon reflex and reflexes mediated by pain and cutaneous receptors.

Tonic vibration reflex

Tonic vibration reflex is a sustained contraction of a muscle subjected to vibration. This reflex is caused by vibratory activation of muscle spindles — muscle receptors sensitive to stretch.

Tonic vibration reflex is evoked by placing a vibrator — which in this case is typically an electrical motor with an eccentric load on its shaft — on a muscle's tendon. 30–100 Hz vibration activates receptors of the skin, tendons and, most importantly, muscle spindles. Muscle spindle discharges are sent to the spinal cord through afferent nerve fibers, where they activate monosynaptic and polysynaptic reflex arcs, causing the muscle to contract.

The effects of sustained vibratory stimulation on muscle contraction, posture and kinesthetic perceptions are much more complex than merely contraction of the muscle being vibrated.

Russian scientists Victor Gurfinkel, Mikhail Lebedev, Andrew Polyakov and Yuri Levick used vibratory stimulation to study human posture control and spectral characteristics of electromyographic (EMG) activity.

Type Ia sensory 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 found in muscles called the muscle spindle, which constantly monitors how fast a muscle stretch changes. (In other words, it monitors the velocity of the stretch).

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