Sensory Systems/Somatosensory System

Introduction

Anatomy of the Somatosensory System

Our somatosensory system consists of sensors in the skin and sensors in our muscles, tendons, and joints. The receptors in the skin, the so called cutaneous receptors, tell us about temperature (thermoreceptors), pressure and surface texture (mechano receptors), and pain (nociceptors). The receptors in muscles and joints provide information about muscle length, muscle tension, and joint angles. (The following description is based on lecture notes from Laszlo Zaborszky, from Rutgers University.)

Cutaneous receptors

Mechanoreceptors

Receptors in the human skin: Mechanoreceptors can be free receptors or encapsulated. Examples for free receptors are the hair receptors at the roots of hairs. Encapsulated receptors are the Pacinian corpuscles and the receptors in the glabrous (hairless) skin: Meissner corpuscles, Ruffini corpuscles and Merkel’s disks.

Sensory information from Meissner corpuscles and rapidly adapting afferents leads to adjustment of grip force when objects are lifted. These afferents respond with a brief burst of action potentials when objects move a small distance during the early stages of lifting. In response to rapidly adapting afferent activity, muscle force increases reflexively until the gripped object no longer moves. Such a rapid response to a tactile stimulus is a clear indication of the role played by somatosensory neurons in motor activity.

The slowly adapting Merkel's receptors are responsible for form and texture perception. As would be expected for receptors mediating form perception, Merkel‘s receptors are present at high density in the digits and around the mouth (50/mm² of skin surface), at lower density in other glabrous surfaces, and at very low density in hairy skin. This innervations density shrinks progressively with the passage of time so that by the age of 50, the density in human digits is reduced to 10/mm². Unlike rapidly adapting axons, slowly adapting fibers respond not only to the initial indentation of skin, but also to sustained indentation up to several seconds in duration.

Activation of the rapidly adapting Pacinian corpuscles gives a feeling of vibration, while the slowly adapting Ruffini corpuscles respond to the lateral movement or stretching of skin.

	Rapidly adapting	Slowly adapting
Surface receptor / small receptive field	Hair receptor, Meissner's corpuscle: Detect an insect or a very fine vibration. Used for recognizing texture.	Merkel's receptor: Used for spatial details, e.g. a round surface edge or "an X" in brail.
Deep receptor / large receptive field	Pacinian corpuscle: "A diffuse vibration" e.g. tapping with a pencil.	Ruffini's corpuscle: "A skin stretch". Used for joint position in fingers.

Nociceptors

Nociceptors have free nerve endings. Functionally, skin nociceptors are either high-threshold mechanoreceptors or polymodal receptors. Polymodal receptors respond not only to intense mechanical stimuli, but also to heat and to noxious chemicals. These receptors respond to minute punctures of the epithelium, with a response magnitude that depends on the degree of tissue deformation. They also respond to temperatures in the range of 40–60°C, and change their response rates as a linear function of warming (in contrast with the saturating responses displayed by non-noxious thermoreceptors at high temperatures).

Pain signals can be separated into individual components, corresponding to different types of nerve fibers used for transmitting these signals. The rapidly transmitted signal, which often has high spatial resolution, is called first pain or cutaneous pricking pain. It is well localized and easily tolerated. The much slower, highly affective component is called second pain or burning pain; it is poorly localized and poorly tolerated. The third or deep pain, arising from viscera, musculature and joints, is also poorly localized, can be chronic and is often associated with referred pain.

Thermoreceptors

The thermoreceptors have free nerve endings. Interestingly, we have only two types of thermoreceptors that signal innocuous warmth and cooling respectively in our skin (however, some nociceptors are also sensitive to temperature, but capable of unambiguously signaling only noxious temperatures). The warm receptors show a maximum sensitivity at ~ 45°C, signal temperatures between 30 and 45°C, and cannot unambiguously signal temperatures higher than 45°C , and are unmyelinated. The cold receptors have their maximum sensitivity at ~ 27°C, signal temperatures above 17°C, and some consist of lightly myelinated fibers, while others are unmyelinated. Our sense of temperature comes from the comparison of the signals from the warm and cold receptors. Thermoreceptors are poor indicators of absolute temperature but are very sensitive to changes in skin temperature.

Proprioceptors

The term proprioceptive or kinesthetic sense is used to refer to the perception of joint position, joint movements, and the direction and velocity of joint movement. There are numerous mechanoreceptors in the muscles, the muscle fascia, and in the dense connective tissue of joint capsules and ligaments. There are two specialized encapsulated, low-threshold mechanoreceptors: the muscle spindle and the Golgi tendon organ. Their adequate stimulus is stretching of the tissue in which they lie. Muscle spindles, joint and skin receptors all contribute to kinesthesia. Muscle spindles appear to provide their most important contribution to kinesthesia with regard to large joints, such as the hip and knee joints, whereas joint receptors and skin receptors may provide more significant contributions with regard to finger and toe joints.

Muscle Spindles

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.

Scattered throughout virtually every striated muscle in the body are long, thin, stretch receptors called muscle spindles. They are quite simple in principle, consisting of a few small muscle fibers with a capsule surrounding the middle third of the fibers. These fibers are called intrafusal fibers, in contrast to the ordinary extrafusal fibers. The ends of the intrafusal fibers are attached to extrafusal fibers, so whenever the muscle is stretched, the intrafusal fibers are also stretched. The central region of each intrafusal fiber has few myofilaments and is non-contractile, but it does have one or more sensory endings applied to it. When the muscle is stretched, the central part of the intrafusal fiber is stretched and each sensory ending fires impulses.

Numerous specializations occur in this simple basic organization, so that in fact the muscle spindle is one of the most complex receptor organs in the body. Only three of these specializations are described here; their overall effect is to make the muscle spindle adjustable and give it a dual function, part of it being particularly sensitive to the length of the muscle in a static sense and part of it being particularly sensitive to the rate at which this length changes.

Intrafusal muscle fibers are of two types. All are multinucleated, and the central, non-contractile region contains the nuclei. In one type of intrafusal fiber, the nuclei are lined up single file; these are called nuclear chain fiber. In the other type, the nuclear region is broader, and the nuclei are arranged several abreast; these are called nuclear bag fibers. There are typically two or three nuclear bag fibers per spindle and about twice that many chain fibers.
There are also two types of sensory endings in the muscle spindle. The first type, called the primary ending, is formed by a single Ia (A-alpha) fiber, supplying every intrafusal fiber in a given spindle. Each branch wraps around the central region of the intrafusal fiber, frequently in a spiral fashion, so these are sometimes called annulospiral endings. The second type of ending is formed by a few smaller nerve fibers (II or A-Beta) on both sides of the primary endings. These are the secondary endings, which are sometimes referred to as flower-spray endings because of their appearance. Primary endings are selectively sensitive to the onset of muscle stretch but discharge at a slower rate while the stretch is maintained. Secondary endings are less sensitive to the onset of stretch, but their discharge rate does not decline very much while the stretch is maintained. In other words, both primary and secondary endings signal the static length of the muscle (static sensitivity) whereas only the primary ending signals the length changes (movement) and their velocity (dynamic sensitivity). The change of firing frequency of group Ia and group II fibers can then be related to static muscle length (static phase) and to stretch and shortening of the muscle (dynamic phases).
Muscle spindles also receive a motor innervation. The large motor neurons that supply extrafusal muscle fibers are called alpha motor neurons, while the smaller ones supplying the contractile portions of intrafusal fibers are called gamma neurons. Gamma motor neurons can regulate the sensitivity of the muscle spindle so that this sensitivity can be maintained at any given muscle length.

Golgi tendon organ

Mammalian tendon organ showing typical position in a muscle (left), neuronal connections in spinal cord (middle) and expanded schematic (right). The tendon organ is a stretch receptor that signals the force developed by the muscle. The sensory endings of the Ib afferent are entwined amongst the musculotendinous strands of 10 to 20 motor units.

The Golgi tendon organ is located at the musculotendinous junction. There is no efferent innervation of the tendon organ, therefore its sensitivity cannot be controlled from the CNS. The tendon organ, in contrast to the muscle spindle, is coupled in series with the extrafusal muscle fibers. Both passive stretch and active contraction of the muscle increase the tension of the tendon and thus activate the tendon organ receptor, but active contraction produces the greatest increase. The tendon organ, consequently, can inform the CNS about the “muscle tension”. In contrast, the activity of the muscle spindle depends on the “muscle length” and not on the tension. The muscle fibers attached to one tendon organ appear to belong to several motor units. Thus the CNS is informed not only of the overall tension produced by the muscle but also of how the workload is distributed among the different motor units.

Joint receptors

The joint receptors are low-threshold mechanoreceptors and have been divided into four groups. They signal different characteristics of joint function (position, movements, direction and speed of movements). The free receptors or type 4 joint receptors are nociceptors.

Proprioceptive Signal Processing

Feedback loops for proprioceptive signals for the perception and control of limb movements. Arrows indicate excitatory connections; filled circles inhibitory connections.

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