Tactile markings stairs for visually impaired

physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical ...

, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch ( haptic perception), as well as temperature ( thermoception), body position ( proprioception), and

pain Pain is a distressing feeling often caused by intense or damaging stimuli. The International Association for the Study of Pain defines pain as "an unpleasant sensory and emotional experience associated with, or resembling that associated with, ...

. It is a subset of the sensory nervous system, which also represents

visual The visual system comprises the sensory organ (the eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) which gives organisms the sense of sight (the ...

, auditory,

olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...

, and gustatory stimuli. Somatosensation begins when mechano- and thermosensitive structures in the skin or internal organs sense physical stimuli such as pressure on the skin (see mechanotransduction,

nociception Nociception (also nocioception, from Latin ''nocere'' 'to harm or hurt') is the sensory nervous system's process of encoding noxious stimuli. It deals with a series of events and processes required for an organism to receive a painful stimulus, co ...

). Activation of these structures, or receptors, leads to activation of peripheral sensory neurons that convey signals to the

spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the spi ...

as patterns of

action potential An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...

s. Sensory information is then processed locally in the spinal cord to drive reflexes, and is also conveyed to the brain for conscious perception of touch and proprioception. Note, somatosensory information from the face and head enters the brain through peripheral sensory neurons in the

cranial nerve Cranial nerves are the nerves that emerge directly from the brain (including the brainstem), of which there are conventionally considered twelve pairs. Cranial nerves relay information between the brain and parts of the body, primarily to and ...

s, such as the

trigeminal nerve In neuroanatomy, the trigeminal nerve ( lit. ''triplet'' nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for sensation in the face and motor functions such as biting and chewin ...

. 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 cerebral cortex in the brain represent nearby locations on the skin or in the body, creating a map, also called a

cortical homunculus A cortical homunculus () is a distorted representation of the human body, based on a neurological "map" of the areas and proportions of the human brain dedicated to processing motor functions, or sensory functions, for different parts of the bo ...

System overview

Comprehensive List of Relevant Pathways for the Somatosensory System

Sensory receptors

The four

mechanoreceptor 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, ...

s in the

skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...

each respond to different stimuli for short or long periods.

Merkel cell Merkel cells, also known as Merkel-Ranvier cells or tactile epithelial cells, are oval-shaped mechanoreceptors essential for light touch sensation and found in the skin of vertebrates. They are abundant in highly sensitive skin like that of the f ...

nerve endings are found in the basal

epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...

and

hair follicles The hair follicle is an organ found in mammalian skin. It resides in the dermal layer of the skin and is made up of 20 different cell types, each with distinct functions. The hair follicle regulates hair growth via a complex interaction between h ...

; 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 corpuscle 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. ...

s 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.

Bulbous corpuscle The Bulbous corpuscle or Ruffini ending or Ruffini corpuscle is a slowly adapting mechanoreceptor located in the cutaneous tissue between the dermal papillae and the hypodermis. It is named after Angelo Ruffini Angelo Ruffini (Pretare of Arquata ...

s 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 Proprioception ( ), also referred to as kinaesthesia (or kinesthesia), is the sense of self-movement, force, and body position. It is sometimes described as the "sixth sense". Proprioception is mediated by proprioceptors, mechanosensory neurons ...

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 squish their shape causing an action potential.

Somatosensory cortex

The postcentral gyrus includes the

primary somatosensory cortex 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 paralle ...

( Brodmann areas 3, 2 and 1) collectively referred to as S1. BA3 receives the densest projections from the

thalamus The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...

. 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 The human secondary somatosensory cortex (S2, SII) is a region of cortex in the parietal operculum on the ceiling of the lateral sulcus. Region S2 was first described by Adrian in 1940, who found that feeling in cats' feet was not only represente ...

) 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). BA7 integrates visual and proprioceptive info to locate objects in space. 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 Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...

body. It consists both of

sensory receptor 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 potentials. This process is called sensory transduction. The cell ...

s and

sensory neuron 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 potentials. This process is called sensory transduction. The cell ...

s in the periphery (skin, muscle and organs for example), to deeper neurons within the

central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all par ...

General somatosensory pathway

All afferent touch/vibration info ascends the spinal cord via the

dorsal column-medial lemniscus pathway Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal co ...

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: first-order, second-order, and third-order. # The first-order neuron is a type of pseudounipolar neuron and always has its

cell body The soma (pl. ''somata'' or ''somas''), perikaryon (pl. ''perikarya''), neurocyton, or cell body is the bulbous, non-process portion of a neuron or other brain cell type, containing the cell nucleus. The word 'soma' comes from the Greek '' σῶμ ...

in the

dorsal root ganglion A dorsal root ganglion (or spinal ganglion; also known as a posterior root ganglion) is a cluster of neurons (a ganglion) in a dorsal root of a spinal nerve. The cell bodies of sensory neurons known as first-order neurons are located in the dorsa ...

of the

spinal nerve A spinal nerve is a mixed nerve, which carries motor, sensory, and autonomic signals between the spinal cord and the body. In the human body there are 31 pairs of spinal nerves, one on each side of the vertebral column. These are grouped into th ...

with a peripheral axon innervating touch

s 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). # The second-order neuron has its

either in the spinal cord or in the brainstem. This neuron's ascending

axons An axon (from Greek ἄξων ''áxōn'', axis), or nerve fiber (or nerve fibre: see American and British English spelling differences#-re, -er, spelling differences), is a long, slender projection of a nerve cell, or neuron, in vertebrates, th ...

will cross (

decussate Decussation is used in biological contexts to describe a crossing (due to the shape of the Roman numeral for ten, an uppercase 'X' (), ). In Latin anatomical terms, the form is used, e.g. . Similarly, the anatomical term chiasma is named aft ...

) to the opposite side either in the

or in the

brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is cont ...

. # In the case of touch and certain types of pain, the third-order neuron has its

in the

ventral posterior nucleus The ventral posterior nucleus is the somato-sensory relay nucleus in thalamus of the brain. Input and output The ventral posterior nucleus receives neuronal input from the medial lemniscus, spinothalamic tracts, and trigeminothalamic tract. It p ...

of the thalamus and ends in the postcentral gyrus of the parietal lobe in the

(or S1).

Photoreceptors, similar to those found in the

retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...

of the

eye Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conv ...

, detect potentially damaging ultraviolet radiation ( ultraviolet A specifically), inducing increased production of

melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...

melanocytes Melanocytes are melanin-producing neural crest-derived cells located in the bottom layer (the stratum basale) of the skin's epidermis, the middle layer of the eye (the uvea), the inner ear, vaginal epithelium, meninges, bones, and heart. ...

. 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.

Tactile feedback

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

Balance

The receptor for the sense of balance resides in the

vestibular system The vestibular system, in vertebrates, is a sensory system that creates the sense of balance and spatial orientation for the purpose of coordinating movement with balance. Together with the cochlea, a part of the auditory system, it constitutes ...

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). 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

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

posterior column–medial lemniscus pathway Posterior may refer to: * Posterior (anatomy), the end of an organism opposite to its head ** Buttocks, as a euphemism * Posterior horn (disambiguation) * Posterior probability The posterior probability is a type of conditional probability that r ...

is the pathway responsible for the sending of fine touch information to the

cerebral cortex The cerebral cortex, also known as the cerebral mantle, is the outer layer of neural tissue of the cerebrum of the brain in humans and other mammals. The cerebral cortex mostly consists of the six-layered neocortex, with just 10% consisting of ...

of the brain. Crude touch (or non-discriminative touch) 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. As fine touch normally works in parallel to crude touch, a person will be able to localize touch until fibres carrying fine touch (

Posterior column–medial lemniscus pathway Posterior may refer to: * Posterior (anatomy), the end of an organism opposite to its head ** Buttocks, as a euphemism * Posterior horn (disambiguation) * Posterior probability The posterior probability is a type of conditional probability that r ...

) 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 don't 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 Functional magnetic resonance imaging or functional MRI (fMRI) measures brain activity by detecting changes associated with blood flow. This technique relies on the fact that cerebral blood flow and neuronal activation are coupled. When an area o ...

(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. Tactile interaction is important amongst most 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. More precisely, the consistency of oxytocin neuron activation in rats stroked by humans has been observed, especially in the caudal paraventricular nucleus. 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. 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. 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. 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.

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. In this task subjects identify the orientation of a grooved surface presented in two different orientations, which can be applied manually or with automated equipment. Many studies have shown a decline in passive tactile spatial acuity with age; 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; this effect of finger size has been shown to underlie the better passive tactile spatial acuity of women, on average, compared to men. The density of

tactile corpuscle 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. ...

s, a type of mechanoreceptor that detects low-frequency vibrations, is greater in smaller fingers; the same may hold for Merkel cells, which detect the static indentations important for fine spatial acuity. Among children of the same age, those with smaller fingers also tend to have better tactile acuity. Many studies have shown that passive tactile spatial acuity is enhanced among blind individuals compared to sighted individuals of the same age, 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.

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 Paresthesia is an abnormal sensation of the skin (tingling, pricking, chilling, burning, numbness) with no apparent physical cause. Paresthesia may be transient or chronic, and may have any of dozens of possible underlying causes. Paresthesias ar ...

Society and culture

Haptic technology can provide touch sensation in virtual and real environments. In the field of

speech therapy Speech is a human vocal communication using language. Each language uses phonetic combinations of vowel and consonant sounds that form the sound of its words (that is, all English words sound different from all French words, even if they are th ...

, tactile feedback can be used to treat speech disorders. 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. 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. 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. 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.

Notes

References

External links

*
Anatomy of Touch
Factual documentary series by

BBC Radio 4 BBC Radio 4 is a British national radio station owned and operated by the BBC that replaced the BBC Home Service in 1967. It broadcasts a wide variety of spoken-word programmes, including news, drama, comedy, science and history from the BBC' ...

. {{DEFAULTSORT:Somatosensory System