The somatosensory system (also somatosensory nervous system) is a complex system of nerve cells that responds to changes to the surface or internal state of the body. Nerve cells called "sensory receptors" (including thermoreceptors, mechanoreceptors, chemoreceptors and nociceptors) send signals along a chain of nerve cells to the spinal cord where they may be processed by other nerve cells and then relayed to the brain for further processing. Sensory receptors are found in many parts of the body including the skin, epithelial tissues, skeletal muscles, bones and joints, internal organs, and the cardiovascular system.
Somatic senses are sometimes referred to as somesthetic senses, with the understanding that somesthesis includes the sense of touch, proprioception (sense of position and movement), and (depending on usage) also haptic perception.
The mapping of the body surfaces in the brain is called a homunculus and plays a fundamental role in the creation of body image. This brain-surface ("cortical") map is not immutable, however. Dramatic shifts can occur in response to stroke or injury.
- 1 Structure
- 2 System overview
- 3 Clinical significance
- 4 Society and culture
- 5 See also
- 6 References
- 7 Further reading
- 8 External links
The somatosensory system is spread through all major parts of a mammal's body (and other vertebrates). It consists both of sensory receptors and sensory (afferent) neurons in the periphery (skin, muscle and organs for example), to deeper neurons within the central nervous system.
General somatosensory pathway
A somatosensory pathway will typically have three long neurons: primary, secondary, and tertiary (or first, second, and third).
- The first neuron always has its cell body in the dorsal root ganglion of the spinal nerve (if sensation is in parts of the head or neck not covered by the cervical nerves, it will be the trigeminal nerve ganglia or the ganglia of other sensory cranial nerves).
- The second 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.
- In the case of touch and certain types of pain, the third neuron has its cell body in the VPN of the thalamus and ends in the postcentral gyrus of the parietal lobe.
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. 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.
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). 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 is the pathway responsible for the sending of fine touch information to the cerebral cortex 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) 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 actually coded differently than other sensory information. Intensity of affective touch is still encoded in the primary somatosensory cortex, but 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. 
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 Meissner's corpuscles, 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.
The 4 skin receptors each respond to different stimuli for short or long periods.
Merkel 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 a small receptive field (extremely detailed info) they are used in areas like fingertips the most; they are not covered (shelled) and thus respond to pressures over long periods.
Meissner’s 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. 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 reacts only to sudden stimuli so pressures like clothes that are always compressing their shape are quickly ignored.
Ruffini 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 ruffini (slow-response) are myelinated; the rest (fast-response) are not. Merkel, pacinian, ruffini and meissner are all activated upon pressures that squish their shape causing an action potential.
All afferent touch/vibration info ascends the spinal cord via the posterior (dorsal) column-medial lemniscus pathway via gracilis (T7 and below) or cuteatus (T6 and above).
Cuteatus 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.
The postcentral gyrus includes the primary somatosensory cortex (Brodmann area 3), and BA2. They are 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 somato info, it projects texture info to BA1 and shape + size info 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 + shape info.
Area S2 processes light touch, pain, visceral sensation, and tactile attention.
The insular cortex 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.
A somatosensory deficiency may be caused by a peripheral neuropathy involving peripheral nerves of the somatosensory system.
Certain types of seizures are associated with the somatosensory system. Cortical injury may lead to loss of thermal sensation or the ability to discriminate pain. An aura involving thermal and painful sensations is a phenomenon known to precede the onset of an epileptic seizure or focal seizure. Another type of seizure, called a sensory Jacksonian seizure involves an abnormal, localizable, cutaneous sensation but does not have apparent stimulus. This sensation may progress along a limb or to adjacent cutaneous body areas, reflecting abnormal neuronal firing in the postcentral gyrus where an epileptic discharge is propagated. These episodes in which patients are consciously aware during a seizure have been useful for identifying problems associated with the somatosensory cortex. Patients can describe the nature of the seizure and how they feel during it.
The absence of proprioception or two-point tactile discrimination on one side of the body suggests injury to the contralateral side of the primary somatosensory cortex. However, depending on the extent of the injury, damage can range in loss of proprioception of an individual limb or the entire body. A deficit known as cortical astereognosis of the receptive type describes an inability to make use of tactile sensory information for identifying objects placed in the hand. For example, if this type of injury effects the hand region in the primary somatosensory cortex for one cerebral hemisphere, a patient with closed eyes cannot perceive the position of the fingers on the contralateral hand and will not be able to identify objects such as keys or a cell phone if they are placed into that hand.
Evaluation of any suspected disease of the somatosensory system is included in a neurological examination of the peripheral nervous system
Clinically, Tuning Fork of 128 Hz used For Vibration Testing.
Society and culture
- Cell signalling
- Molecular cellular cognition
- Haptic communication
- Haptic perception
- Muscle spindle
- Phantom limb
- Secondary somatosensory cortex
- Somatosensory Rehabilitation of Pain
- Special senses
- Supramarginal gyrus
- Tactile illusion
- Vibratese, method of communication through touch
- Tactile imaging
- O. Franzen, R. Johansson, L. Terenius (1996) Somesthesis and the Neurobiology of the Somatosensory Cortex
- Robles-De-La-Torre G (2006). "The Importance of the Sense of Touch in Virtual and Real Environments" (PDF). IEEE Multimedia. 13 (3): 24–30. doi:10.1109/MMUL.2006.69.
- Saladin KS. Anatomy and Physiology 3rd edd. 2004. McGraw-Hill, New York.
- Zukerman, Wendy. "Skin 'sees' the light to protect against sunshine". newscientist.com. New Scientist. Retrieved 2015-01-22.
- Proske U, Gandevia SC."The proprioceptive senses: their roles in signaling body shape, body position and movement, and muscle force." Physiol Rev. 2012 Oct;92(4):1651-97. doi: 10.1152/physrev.00048.2011
- Proske & Gandevia 2009 "The kinaesthetic senses" p.4144 "The three sources for limb position and movement representation are tactile, visual, and proprioceptive."
- "Encoding of Touch Intensity But Not Pleasantness in Human Primary Somatosensory Cortex". PMID 27225773.
- . PMID 27225773. Missing or empty
- Van Boven, R. W.; Johnson, K. O. (1 December 1994). "The limit of tactile spatial resolution in humans: Grating orientation discrimination at the lip, tongue, and finger". Neurology. 44 (12): 2361–2361. doi:10.1212/WNL.44.12.2361.
- Craig JC. (1999). "Grating orientation as a measure of tactile spatial acuity.". Somatosens Mot Res. 16 (3): 197–206. doi:10.1080/08990229970456. PMID 10527368.
- Goldreich, D; Wong, M; Peters, RM; Kanics, IM (Jun 3, 2009). "A Tactile Automated Passive-Finger Stimulator (TAPS).". Journal of visualized experiments (28). doi:10.3791/1374. PMID 19578327.
- Stevens, JC; Alvarez-Reeves, M; Dipietro, L; Mack, GW; Green, BG (2003). "Decline of tactile acuity in aging: a study of body site, blood flow, and lifetime habits of smoking and physical activity.". Somatosensory & motor research. 20 (3–4): 271–9. doi:10.1080/08990220310001622997. PMID 14675966.
- Manning, Hélène; Tremblay, FranÇois (2006). "Age differences in tactile pattern recognition at the fingertip". Somatosensory & Motor Research. 23 (3–4): 147–155. doi:10.1080/08990220601093460.
- Goldreich, D; Kanics, IM (2003). "Tactile acuity is enhanced in blindness.". Journal of Neuroscience. 23 (8): 3439–45. PMID 12716952.
- Peters, RM; Hackeman, E; Goldreich, D (2009). "Diminutive digits discern delicate details: fingertip size and the sex difference in tactile spatial acuity.". Journal of Neuroscience. 29 (50): 15756–61. doi:10.1523/JNEUROSCI.3684-09.2009. PMID 20016091.
- Dillon, YK; Haynes, J; Henneberg, M (2001). "The relationship of the number of Meissner's corpuscles to dermatoglyphic characters and finger size.". Journal of Anatomy. 199 (Pt 5): 577–84. doi:10.1046/j.1469-7580.2001.19950577.x. PMID 11760888.
- Peters, RM; Goldreich, D (2013). "Tactile spatial acuity in childhood: effects of age and fingertip size.". PLOS ONE. 8 (12): e84650. doi:10.1371/journal.pone.0084650. PMC . PMID 24454612.
- Stevens, Joseph C.; Foulke, Emerson; Patterson, Matthew Q. (1996). "Tactile acuity, aging, and braille reading in long-term blindness.". Journal of Experimental Psychology: Applied. 2 (2): 91–106. doi:10.1037/1076-898X.2.2.91.
- Van Boven, RW; Hamilton, RH; Kauffman, T; Keenan, JP; Pascual-Leone, A (2000). "Tactile spatial resolution in blind braille readers.". Neurology. 54 (12): 2230–6. doi:10.1212/wnl.54.12.2230. PMID 10881245.
- Goldreich, D; Kanics, IM (2006). "Performance of blind and sighted humans on a tactile grating detection task.". Perception & psychophysics. 68 (8): 1363–71. doi:10.3758/bf03193735. PMID 17378422.
- Wong, M; Gnanakumaran, V; Goldreich, D (2011). "Tactile spatial acuity enhancement in blindness: evidence for experience-dependent mechanisms.". Journal of Neuroscience. 31 (19): 7028–37. doi:10.1523/JNEUROSCI.6461-10.2011. PMID 21562264.
- Bhattacharjee, A; Ye, AJ; Lisak, JA; Vargas, MG; Goldreich, D (2010). "Vibrotactile masking experiments reveal accelerated somatosensory processing in congenitally blind braille readers.". Journal of Neuroscience. 30 (43): 14288–98. doi:10.1523/JNEUROSCI.1447-10.2010. PMID 20980584.
- Tillotson, Joanne. McCann, Stephanie. Kaplan’s Medical Flashcards. Apr. 02. 2013.
- Albertine, Kurt. Barron’s Anatomy Flash Cards
- "Sense of Touch - How Do We Feel? A Simple Explanation." HubPages. HubPages, 19 July 2014. Web. 27 Mar. 2016.
- Paré, Michel, and Catherine Behets. "Paucity of Presumptive Ruffini Corpuscles in the Index Finger Pad of Humans." Wiley Online Library. 10 Feb. 2003. Web. 27 Mar. 2016.
- Scheibert J, Leurent S, Prevost A, Debrégeas G (2009). "The role of fingerprints in the coding of tactile information probed with a biomimetic sensor". Science. 323 (5920): 1503–6. doi:10.1126/science.1166467. PMID 19179493.
- Biswas, Abhijit; Manivannan, M.; Srinivasan, Mandyam A. (2015). "Vibrotactile Sensitivity Threshold: Nonlinear Stochastic Mechanotransduction Model of the Pacinian Corpuscle". IEEE Transactions on Haptics. 8 (1): 102–113. doi:10.1109/TOH.2014.2369422. PMID 25398183.
- Paré, Michel, and Robert Elde. "The Meissner Corpuscle Revised: A Multiafferented Mechanoreceptor with Nociceptor Immunochemical Properties." JNeurosci. 15 Sept. 2001. Web. 27 Mar. 2016.
- Asaga, Emi, and Kenjiro Takemura. "Tactile Evaluation Based on Human Tactile Perception Mechanism." Science Direct. 01 Dec. 2013. Web. 27 Mar. 2016.
- Buccino, G., and F. Binkofski. "Action Observation Activates Premotor and Parietal Areas in a Somatotopic Manner: An FMRI Study." Wiley Online Library. Jan. 2001. Web. 27 Mar. 2016.
- Seelke, Adele M.H., and Jeffrey J. Padberg. "Topographic Maps within Brodmann's Area 5 of Macaque Monkeys." Oxford Journals. 2011. Web. 27 Mar. 2016.
- Menon, Vinod, and Lucina Q. Uddin. "Saliency, Switching, Attention and Control: A Network Model of Insula." Springer. 29 May 2010. Web. 28 Mar. 2016.
- Geyer, Stefan, and Axel Schleicher. "Areas 3a, 3b, and 1 of Human Primary Somatosensory Cortex: 1. Microstructural Organization and Interindividual Variability." Science Direct. July 1999. Web. 27 Mar. 2016.
- Disbrow, Elizabeth, and Evangelos Litinas. "Thalamocortical Connections of the Parietal Ventral Area (PV) and the Second Somatosensory Area (S2) in Macaque Monkeys." Cambridge Journals Online. June 2002. Web. 27 Mar. 2016.
- Augustine, James R. (2008). Human Neuroanatomy. San Diego, CA: Academic Press. p. 360. ISBN 978-0-12-068251-5.
- Gabriel Robles-De-La-Torre. "International Society for Haptics: Haptic technology, an animated explanation". Isfh.org. Retrieved 2010-02-26.
- Boron, Walter F.; Boulpaep, Emile L. (2003). Medical Physiology. Saunders. pp. 352–358. ISBN 0-7216-3256-4.
- Flanagan, J.R., Lederman, S.J. Neurobiology: Feeling bumps and holes, News and Views, Nature, 2001 Jul. 26;412(6845):389-91.
- Hayward V, Astley OR, Cruz-Hernandez M, Grant D, Robles-De-La-Torre G. Haptic interfaces and devices. Sensor Review 24(1), pp. 16–29 (2004).
- Purves, Dale (2012). Neuroscience, Fifth Edition. Sunderland, MA: Sinauer Associates, Inc. pp. 202–203. ISBN 978-0-87893-695-3.
- Robles-De-La-Torre G., Hayward V. Force Can Overcome Object Geometry In the perception of Shape Through Active Touch" Nature 412 (6845) 445-8 (2001).
- Robles-De-La-Torre G. The Importance of the Sense of Touch in Virtual and Real Environments. IEEE Multimedia 13(3), Special issue on Haptic NO User Interfaces for Multimedia Systems, pp. 24–30 (2006).
- Grunwald, M. (Ed.) Human Haptic Perception – Basics and Applications. Boston/Basel/Berlin: Birkhäuser, 2008, ISBN 978-3-7643-7611-6
- Encyclopedia of Touch Scholarpedia Expert articles