Somatosensory system

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Touch is a crucial means of receiving information. This photo shows tactile markings identifying stairs for visually impaired people. Tactile markings stairs for visually impaired.jpg
Touch is a crucial means of receiving information. This photo shows tactile markings identifying stairs for visually impaired people.

The somatosensory system, or somatic sensory system is a subset of the sensory nervous system. It has two subdivisions, one for the detection of mechanosensory information related to touch, and the other for the nociception detection of pain and temperature. [1] The main functions of the somatosensory system are the perception of external stimuli, the perception of internal stimuli, and the regulation of body position and balance (proprioception). [2]

Contents

Mechanosensory information includes that of light touch, vibration, pressure and tension in the skin. Much of this information belongs to the sense of touch which is a general somatic sense in contrast to the special senses of sight, smell, taste, hearing and balance. [3]

Nociceptory information is that received from pain and temperature that is deemed as harmful (noxious). Thermoreceptors relay temperature information in normal circumstances. [1] Nociceptors are specialised receptors for signals of pain. [4]

The sense of touch in perceiving the environment uses special sensory receptors in the skin called cutaneous receptors. They include mechanoreceptors such as tactile corpuscles that relay information about pressure and vibration; nociceptors, and thermoreceptors for temperature perception. [5]

Stimulation of the receptors activate peripheral sensory neurons that convey signals to the spinal cord that may drive a responsive reflex, and may also be conveyed to the brain for conscious perception. Somatosensory information from the face and head enter the brain via cranial nerves such as the trigeminal nerve.

The neural pathways that go to the brain are structured such that information about the location of the physical stimulus is preserved. In this way, neighboring neurons in the somatosensory cortex represent nearby locations on the skin or in the body, creating a map or sensory homunculus.

Touch communication

Tactile signing

Tactile signing is a common means of communication used by people with deafblindness. It is based on a sign language or another system of manual communication.

Emotion communication

Humans can communicate specific emotions through touch alone including anger, fear, disgust, love, gratitude, and sympathy via touch at much-better-than-chance levels. [6]

Overview

This diagram linearly (unless otherwise mentioned) tracks the projections of all known structures that allow for touch to their relevant endpoints in the human brain. Comprehensive List of Relevant Pathways for the Somatosensory System.png
This diagram linearly (unless otherwise mentioned) tracks the projections of all known structures that allow for touch to their relevant endpoints in the human brain.

Sensory receptors

The two different types of mechanoreceptor in the skin are termed low-threshold mechanoreceptors, and high threshold mechanoreceptors. The four mechanoreceptors in glabrous skin are low-threshold that respond to harmless stimuli. They are innervated by four different afferent fibers. High-threshold mechanoreceptors, respond to harmful stimuli. [7]

Merkel cell nerve endings are found in the basal epidermis and hair follicles; they react to low vibrations (5–15  Hz) and deep static touch such as shapes and edges. Due to having a small receptive field (extremely detailed information), they are used in areas like fingertips the most; they are not covered (shelled) and thus respond to pressures over long periods.

Tactile corpuscles react to moderate vibration (10–50 Hz) and light touch. They are located in the dermal papillae; due to their reactivity, they are primarily located in fingertips and lips. They respond in quick action potentials, unlike Merkel nerve endings. They are responsible for the ability to read Braille and feel gentle stimuli.

Pacinian corpuscles determine gross touch and distinguish rough and soft substances. They react in quick action potentials, especially to vibrations around 250 Hz (even up to centimeters away). They are the most sensitive to vibrations and have large receptor fields. Pacinian corpuscles react only to sudden stimuli so pressures like clothes that are always compressing their shape are quickly ignored. They have also been implicated in detecting the location of touch sensations on handheld tools. [8]

Bulbous corpuscles react slowly and respond to sustained skin stretch. They are responsible for the feeling of object slippage and play a major role in the kinesthetic sense and control of finger position and movement. Merkel and bulbous cells - slow-response - are myelinated; the rest - fast-response - are not. All of these receptors are activated upon pressures that distort their shape causing an action potential. [9] [10] [11] [12]

Somatosensory cortex

Gray's Anatomy, figure 759: the sensory tract, showing the pathway (blue) up the spinal cord, through the somatosensory thalamus, to S1 (Brodmann areas 3, 1, and 2), S2, and BA7 Gray759.png
Gray's Anatomy, figure 759: the sensory tract, showing the pathway (blue) up the spinal cord, through the somatosensory thalamus, to S1 (Brodmann areas 3, 1, and 2), S2, and BA7
Gray's Anatomy, figure 717: detail showing path adjacent to the insular cortex (marked insula in this figure), adjacent to S1, S2, and BA7 Gray717.png
Gray's Anatomy, figure 717: detail showing path adjacent to the insular cortex (marked insula in this figure), adjacent to S1, S2, and BA7

The postcentral gyrus is in the parietal lobe and its cortex is the primary somatosensory cortex (Brodmann areas 3, 2 and 1) collectively referred to as S1.

BA3 receives the densest projections from the thalamus. BA3a is involved with the sense of relative position of neighboring body parts and amount of effort being used during movement. BA3b is responsible for distributing somatosensory information, it projects texture information to BA1 and shape and size information to BA2.

Region S2 (secondary somatosensory cortex) divides into Area S2 and parietal ventral area. Area S2 is involved with specific touch perception and is thus integrally linked with the amygdala and hippocampus to encode and reinforce memories.

Parietal ventral area is the somatosensory relay to the premotor cortex and somatosensory memory hub, BA5.

BA5 is the topographically organized somato memory field and association area.

BA1 processes texture info while BA2 processes size and shape information.

Area S2 processes light touch, pain, visceral sensation, and tactile attention.

S1 processes the remaining info (crude touch, pain, temperature). [13] [14] [15]

BA7 integrates visual and proprioceptive info to locate objects in space. [16] [17]

The insular cortex (insula) plays a role in the sense of bodily-ownership, bodily self-awareness, and perception. Insula also plays a role in conveying info about sensual touch, pain, temperature, itch, and local oxygen status. Insula is a highly connected relay and thus is involved in numerous functions.

Structure

The somatosensory system is spread through all major parts of the vertebrate body. It consists both of sensory receptors and sensory neurons in the periphery (skin, muscle and organs for example), to deeper neurons within the central nervous system. [3]

General somatosensory pathway

All afferent touch/vibration info ascends the spinal cord via the dorsal column-medial lemniscus pathway via gracilis (T7 and below) or cuneatus (T6 and above). Cuneatus sends signals to the cochlear nucleus indirectly via spinal grey matter, this info is used in determining if a perceived sound is just villi noise/irritation. All fibers cross (left becomes right) in the medulla.

A somatosensory pathway will typically have three neurons: [18] first-order, second-order, and third-order. [19]

  1. The first-order neuron is a type of pseudounipolar neuron and always has its cell body in the dorsal root ganglion of the spinal nerve with a peripheral axon innervating touch mechanoreceptors and a central axon synapsing on the second-order neuron. If the somatosensory pathway is in parts of the head or neck not covered by the cervical nerves, the first-order neuron will be the trigeminal nerve ganglia or the ganglia of other sensory cranial nerves).
  2. The second-order neuron has its cell body either in the spinal cord or in the brainstem. This neuron's ascending axons will cross (decussate) to the opposite side either in the spinal cord or in the brainstem.
  3. In the case of touch and certain types of pain, the third-order neuron has its cell body in the ventral posterior nucleus of the thalamus and ends in the postcentral gyrus of the parietal lobe in the primary somatosensory cortex (or S1).

Photoreceptors, similar to those found in the retina of the eye, detect potentially damaging ultraviolet radiation (ultraviolet A specifically), inducing increased production of melanin by melanocytes. [20] Thus tanning potentially offers the skin rapid protection from DNA damage and sunburn caused by ultraviolet radiation (DNA damage caused by ultraviolet B). However, whether this offers protection is debatable, because the amount of melanin released by this process is modest in comparison to the amounts released in response to DNA damage caused by ultraviolet B radiation. [20]

Tactile feedback

Touch can result in many different physiological reactions such as laughing at being tickled. Tickle.jpg
Touch can result in many different physiological reactions such as laughing at being tickled.

The tactile feedback from proprioception is derived from the proprioceptors in the skin, muscles, and joints. [21]

Balance

The receptor for the sense of balance resides in the vestibular system in the ear (for the three-dimensional orientation of the head, and by inference, the rest of the body). Balance is also mediated by the kinesthetic reflex fed by proprioception (which senses the relative location of the rest of the body to the head). [22] In addition, proprioception estimates the location of objects which are sensed by the visual system (which provides confirmation of the place of those objects relative to the body), as input to the mechanical reflexes of the body.

Fine touch and crude touch

The cortical homunculus, a map of somatosensory areas of the brain, was devised by Wilder Penfield. Sensory Homunculus.png
The cortical homunculus, a map of somatosensory areas of the brain, was devised by Wilder Penfield.

Fine touch (or discriminative touch) is a sensory modality that allows a subject to sense and localize touch. The form of touch where localization is not possible is known as crude touch. The dorsal column–medial lemniscus pathway is the pathway responsible for the sending of fine touch information to the cerebral cortex of the brain.

Crude touch (non-discriminating) is a sensory modality that allows the subject to sense that something has touched them, without being able to localize where they were touched (contrasting "fine touch"). Its fibres are carried in the spinothalamic tract, unlike the fine touch, which is carried in the dorsal column. [23] As fine touch normally works in parallel to crude touch, a person will be able to localize touch until fibres carrying fine touch (in the dorsal column–medial lemniscus pathway) have been disrupted. Then the subject will feel the touch, but be unable to identify where they were touched.

Neural processing of social touch

The somatosensory cortex encodes incoming sensory information from receptors all over the body. Affective touch is a type of sensory information that elicits an emotional reaction and is usually social in nature, such as a physical human touch. This type of information is actually coded differently than other sensory information. Intensity of affective touch is still encoded in the primary somatosensory cortex and is processed in a similar way to emotions invoked by sight and sound, as exemplified by the increase of adrenaline caused by the social touch of a loved one, as opposed to the physical inability to touch someone you do not love.

Meanwhile, the feeling of pleasantness associated with affective touch activates the anterior cingulate cortex more than the primary somatosensory cortex. Functional magnetic resonance imaging (fMRI) data shows that increased blood-oxygen-level contrast (BOLD) signal in the anterior cingulate cortex as well as the prefrontal cortex is highly correlated with pleasantness scores of an affective touch. Inhibitory transcranial magnetic stimulation (TMS) of the primary somatosensory cortex inhibits the perception of affective touch intensity, but not affective touch pleasantness. Therefore, the S1 is not directly involved in processing socially affective touch pleasantness, but still plays a role in discriminating touch location and intensity. [23]

Tactile interaction is important amongst some animals. Usually, tactile contact between two animals occurs through stroking, licking, or grooming. These behaviours are essential for the individual's social healthcare, as in the hypothalamus they induce the release of oxytocin, a hormone that decreases stress and anxiety and increases social bonding between animals. [24] [ clarification needed ]

More precisely, the consistency of oxytocin neuron activation in rats stroked by humans has been observed, especially in the caudal paraventricular nucleus. [25] It was found that this affiliative relationship induced by tactile contact is common no matter the relationship between the two individuals (mother-infant, male-female, human-animal). It has also been discovered that the level of oxytocin release through this behaviour correlates with the time course of social interaction as longer stroking induced a greater release of the hormone. [26]

The importance of somatosensory stimulation in social animals such as primates has also been observed. Grooming is part of the social interaction primates exert on their conspecifics. This interaction is required between individuals to maintain the affiliative relationship within the group, avoid internal conflict and increase group bonding. [27] However, such social interaction requires the recognition of every member in the group. As such, it has been observed that the size of the neocortex is positively correlated with the size of the group, reflecting a limit to the number of recognizable members amongst which grooming can occur. [27] Furthermore, the time course of grooming is related to vulnerability due to predation to which animals are exposed to whilst performing such social interaction. The relationship between tactile interaction, stress reduction and social bonding depends on the evaluation of risks that occur during conducting such behaviours in the wild life, and further research is required to unveil the connection between tactile caring and fitness level.

Studies show a correlation between touching a soft or hard object and how a person thinks or even makes decisions. [28] Further, between the firmness of a touch and the evoking of gender stereotyping. [29]

Tactile memories as part of haptic memory, are organized somatotopically, following the organization of the somatosensory cortex.

Individual variation

A variety of studies have measured and investigated the causes for differences between individuals in the sense of fine touch. One well-studied area is passive tactile spatial acuity, the ability to resolve the fine spatial details of an object pressed against the stationary skin. A variety of methods have been used to measure passive tactile spatial acuity, perhaps the most rigorous being the grating orientation task. [30] In this task subjects identify the orientation of a grooved surface presented in two different orientations, [31] which can be applied manually or with automated equipment. [32] Many studies have shown a decline in passive tactile spatial acuity with age; [33] [34] [35] the reasons for this decline are unknown, but may include loss of tactile receptors during normal aging. Remarkably, index finger passive tactile spatial acuity is better among adults with smaller index fingertips; [36] this effect of finger size has been shown to underlie the better passive tactile spatial acuity of women, on average, compared to men. [36] The density of tactile corpuscles, a type of mechanoreceptor that detects low-frequency vibrations, is greater in smaller fingers; [37] the same may hold for Merkel cells, which detect the static indentations important for fine spatial acuity. [36] Among children of the same age, those with smaller fingers also tend to have better tactile acuity. [38] Many studies have shown that passive tactile spatial acuity is enhanced among blind individuals compared to sighted individuals of the same age, [35] [39] [40] [41] [42] possibly because of cross modal plasticity in the cerebral cortex of blind individuals. Perhaps also due to cortical plasticity, individuals who have been blind since birth reportedly consolidate tactile information more rapidly than sighted people. [43]

Clinical significance

A somatosensory deficiency may be caused by a peripheral neuropathy involving peripheral nerves of the somatosensory system. This may present as numbness or paresthesia.

Society and culture

Haptic technology can provide touch sensation in virtual and real environments. [44] In the field of speech therapy, tactile feedback can be used to treat speech disorders.[ citation needed ]

Affectionate touch is present in everyday life and can take multiple forms. These actions, however, seem to carry specific functions even though the evolutionary benefit from such a wide range of behaviours is not entirely understood. Researchers investigated the expression patterns and characteristics of 8 different affectionate touch actions - embracing, holding, kissing, leaning, petting, squeezing, stroking, and tickling - in a self-report study. [45] It was found that the affectionate touch has distinct target areas on the body, different associated affect, comfort-value, and expression frequency based on the type of touch action that is performed.

Besides the rather obvious sensory consequences of touch, it can also affect higher-level aspects of cognition such as social judgements and decision-making. This effect might arise due to a physical-to-mental scaffolding process in early development, whereby sensorimotor experiences are linked to the emergence of conceptual knowledge. [46] Such links might be maintained throughout life, and so touching an object may cue the physical sensation to its related conceptual processing. Indeed, it was found that different physical properties - weight, texture, and hardness - of a touched object can influence social judgement and decision-making. [47] For example, participants described a passage of a social interaction to be harsher when they touched a hard wooden block instead of a soft blanket prior to the task. Building on these findings, the ability of touch to have an unconscious influence on such higher-order thoughts may provide a novel tool for marketing and communication strategies.

See also

Related Research Articles

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

The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons, neural pathways, and parts of the brain involved in sensory perception and interoception. Commonly recognized sensory systems are those for vision, hearing, touch, taste, smell, balance and visceral sensation. Sense organs are transducers that convert data from the outer physical world to the realm of the mind where people interpret the information, creating their perception of the world around them.

<span class="mw-page-title-main">Parietal lobe</span> Part of the brain responsible for sensory input and some language processing

The parietal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The parietal lobe is positioned above the temporal lobe and behind the frontal lobe and central sulcus.

Stimulus modality, also called sensory modality, is one aspect of a stimulus or what is perceived after a stimulus. For example, the temperature modality is registered after heat or cold stimulate a receptor. Some sensory modalities include: light, sound, temperature, taste, pressure, and smell. The type and location of the sensory receptor activated by the stimulus plays the primary role in coding the sensation. All sensory modalities work together to heighten stimuli sensation when necessary.

A cutaneous receptor is the type of sensory receptor found in the skin. They are a part of the somatosensory system. Cutaneous receptors include mechanoreceptors, nociceptors (pain), and thermoreceptors (temperature).

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">Dorsal column–medial lemniscus pathway</span> Sensory spinal pathway

The dorsal column–medial lemniscus pathway (DCML) (also known as the posterior column-medial lemniscus pathway is the major 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 this information to the somatosensory cortex of the postcentral gyrus in the parietal lobe of the brain. The pathway receives information from sensory receptors throughout the body, and carries this in the gracile fasciculus and the cuneate fasciculus, tracts that make up the white matter dorsal columns of the spinal cord. At the level of the medulla oblongata, the fibers of the tracts decussate and are 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">Tactile corpuscle</span> Type of mechanoreceptor that detects light touch

Tactile corpuscles or Meissner's corpuscles are a type of mechanoreceptor discovered by anatomist Georg Meissner (1829–1905) and Rudolf Wagner. This corpuscle is a type of nerve ending in the skin that is responsible for sensitivity to pressure. In particular, they have their highest sensitivity when sensing vibrations between 10 and 50 hertz. They are rapidly adaptive receptors. They are most concentrated in thick hairless skin, especially at the finger pads.

<span class="mw-page-title-main">Sensory neuron</span> Nerve cell that converts environmental stimuli into corresponding internal stimuli

Sensory neurons, also known as afferent neurons, are neurons in the nervous system, that convert a specific type of stimulus, via their receptors, into action potentials or graded receptor potentials. This process is called sensory transduction. The cell bodies of the sensory neurons are located in the dorsal root ganglia of the spinal cord.

<span class="mw-page-title-main">Primary somatosensory cortex</span> Region of the brain which processes touch

In neuroanatomy, the primary somatosensory cortex is located in the postcentral gyrus of the brain's parietal lobe, and is part of the somatosensory system. It was initially defined from surface stimulation studies of Wilder Penfield, and parallel surface potential studies of Bard, Woolsey, and Marshall. Although initially defined to be roughly the same as Brodmann areas 3, 1 and 2, more recent work by Kaas has suggested that for homogeny with other sensory fields only area 3 should be referred to as "primary somatosensory cortex", as it receives the bulk of the thalamocortical projections from the sensory input fields.

<span class="mw-page-title-main">Pacinian corpuscle</span> Type of mechanoreceptor cell in hairless mammals

The Pacinian corpuscle, lamellar corpuscle or Vater-Pacini corpuscle is one of the four major types of cutaneous receptor a type of mechanoreceptor found in mammalian skin. This type of mechanoreceptor is found in both hairy, and hairless skin, organs, joints, and attached to the periosteum of bone. Pacinian corpuscles are primarily responsible for sensitivity to vibration, and deep pressure. A few are also sensitive to quasi-static or low frequency pressure stimuli. Most of them respond only to sudden disturbances and are especially sensitive to vibration of a few hundreds hertz. The vibrational role may be used for detecting surface texture, such as rough and smooth. Most of the Pacinian corpuscles act as rapidly adapting mechanoreceptors. Groups of corpuscles respond to pressure changes, such as on grasping or releasing an object.

Multisensory integration, also known as multimodal integration, is the study of how information from the different sensory modalities may be integrated by the nervous system. A coherent representation of objects combining modalities enables animals to have meaningful perceptual experiences. Indeed, multisensory integration is central to adaptive behavior because it allows animals to perceive a world of coherent perceptual entities. Multisensory integration also deals with how different sensory modalities interact with one another and alter each other's processing.

Sensory substitution is a change of the characteristics of one sensory modality into stimuli of another sensory modality.

In neuroanatomy, topographic map is the ordered projection of a sensory surface or an effector system to one or more structures of the central nervous system. Topographic maps can be found in all sensory systems and in many motor systems.

Pallesthesia, or vibratory sensation, is the ability to perceive vibration. This sensation, often conducted through skin and bone, is usually generated by mechanoreceptors such as Pacinian corpuscles, Merkel disk receptors, and tactile corpuscles. All of these receptors stimulate an action potential in afferent nerves found in various layers of the skin and body. The afferent neuron travels to the spinal column and then to the brain where the information is processed. Damage to the peripheral nervous system or central nervous system can result in a decline or loss of pallesthesia.

Tactile discrimination is the ability to differentiate information through the sense of touch. The somatosensory system is the nervous system pathway that is responsible for this essential survival ability used in adaptation. There are various types of tactile discrimination. One of the most well known and most researched is two-point discrimination, the ability to differentiate between two different tactile stimuli which are relatively close together. Other types of discrimination like graphesthesia and spatial discrimination also exist but are not as extensively researched. Tactile discrimination is something that can be stronger or weaker in different people and two major conditions, chronic pain and blindness, can affect it greatly. Blindness increases tactile discrimination abilities which is extremely helpful for tasks like reading braille. In contrast, chronic pain conditions, like arthritis, decrease a person's tactile discrimination. One other major application of tactile discrimination is in new prosthetics and robotics which attempt to mimic the abilities of the human hand. In this case tactile sensors function similarly to mechanoreceptors in a human hand to differentiate tactile stimuli.

<span class="mw-page-title-main">Two-point discrimination</span> Ability to discern between single and pairs of points on ones skin

Two-point discrimination (2PD) is the ability to discern that two nearby objects touching the skin are truly two distinct points, not one. It is often tested with two sharp points during a neurological examination and is assumed to reflect how finely innervated an area of skin is.

A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of stimuli. Although, in some cultures, five human senses were traditionally identified as such, many more are now recognized. Senses used by non-human organisms are even greater in variety and number. During sensation, sense organs collect various stimuli for transduction, meaning transformation into a form that can be understood by the brain. Sensation and perception are fundamental to nearly every aspect of an organism's cognition, behavior and thought.

<span class="mw-page-title-main">Cross modal plasticity</span> Reorganization of neurons in the brain to integrate the function of two or more sensory systems

Cross modal plasticity is the adaptive reorganization of neurons to integrate the function of two or more sensory systems. Cross modal plasticity is a type of neuroplasticity and often occurs after sensory deprivation due to disease or brain damage. The reorganization of the neural network is greatest following long-term sensory deprivation, such as congenital blindness or pre-lingual deafness. In these instances, cross modal plasticity can strengthen other sensory systems to compensate for the lack of vision or hearing. This strengthening is due to new connections that are formed to brain cortices that no longer receive sensory input.

Many types of sense loss occur due to a dysfunctional sensation process, whether it be ineffective receptors, nerve damage, or cerebral impairment. Unlike agnosia, these impairments are due to damages prior to the perception process.

Haptic memory is the form of sensory memory specific to touch stimuli. Haptic memory is used regularly when assessing the necessary forces for gripping and interacting with familiar objects. It may also influence one's interactions with novel objects of an apparently similar size and density. Similar to visual iconic memory, traces of haptically acquired information are short lived and prone to decay after approximately two seconds. Haptic memory is best for stimuli applied to areas of the skin that are more sensitive to touch. Haptics involves at least two subsystems; cutaneous, or everything skin related, and kinesthetic, or joint angle and the relative location of body. Haptics generally involves active, manual examination and is quite capable of processing physical traits of objects and surfaces.

References

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