# Nociception

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How an organism receives and responds to painful stimuli

In [physiology](/source/Physiology), **nociception** /ˌnəʊsɪˈsɛpʃ(ə)n/, also **nocioception** (from [Latin](/source/Latin_language) *nocere* 'to [harm](/source/Harm)/hurt'), is the [sensory nervous system](/source/Somatosensory_system)'s process of encoding [noxious stimuli](/source/Noxious_stimulus). It deals with a series of events and processes required for an organism to receive a [painful](/source/Pain) stimulus, convert it to a molecular signal, and recognize and characterize the signal to trigger an appropriate defensive response.

In nociception, intense chemical (e.g., [capsaicin](/source/Capsaicin) present in [chili pepper](/source/Chili_pepper) or [cayenne pepper](/source/Cayenne_pepper)), mechanical (e.g., cutting, crushing), or thermal (heat and cold) stimulation of [sensory neurons](/source/Sensory_neuron) called [nociceptors](/source/Nociceptor) produces a signal that travels along a chain of [nerve fibers](/source/Nerve_fiber) to the [brain](/source/Brain).[1] Nociception triggers a variety of physiological and behavioral responses to protect the organism against an aggression, and usually results in a subjective experience, or [perception](/source/Perception), of pain in [sentient](/source/Sentience) beings.[2]

## Detection of noxious stimuli

Mechanism of nociception via [sensory afferents](/source/Sensory_afferents)

Potentially damaging mechanical, thermal, and chemical stimuli are detected by nerve endings called nociceptors, which are found in the [skin](/source/Skin), on internal surfaces such as the [periosteum](/source/Periosteum), [joint](/source/Joint) surfaces, and in some internal [organs](/source/Organ_(anatomy)). Some nociceptors are unspecialized [free nerve endings](/source/Free_nerve_ending) that have their cell bodies outside the [spinal column](/source/Spinal_column) in [dorsal root ganglia](/source/Dorsal_root_ganglia).[3] Others are specialised structures in the skin such as nociceptive [Schwann cells](/source/Schwann_cells).[4] Nociceptors are categorized according to the [axons](/source/Axon) which travel from the receptors to the [spinal cord](/source/Spinal_cord) or brain. After nerve injury, it is possible for touch fibers that normally carry non-noxious stimuli to be perceived as noxious.[5]

Nociceptive pain consists of an adaptive alarm system.[6] Nociceptors have a certain threshold; that is, they require a minimum intensity of stimulation before they trigger a signal. Once this threshold is reached, a signal is passed along the neuron's axon into the spinal cord.

Nociceptive threshold testing deliberately applies a noxious stimulus to a human or animal subject to study pain. In animals, the technique is often used to study the efficacy of [analgesic drugs](/source/Analgesic_drugs) and to establish dosing levels and periods of effect. After establishing a baseline, the drug under test is given, and the elevation in threshold is recorded at specified times. The threshold should return to the baseline (pretreatment) value when the drug wears off. In some conditions, the excitation of pain fibers increases as the pain stimulus continues, leading to a condition called [hyperalgesia](/source/Hyperalgesia).

## Theory

Main article: [Pain theories](/source/Pain_theories)

## Consequences

Nociception can also cause generalized [autonomic responses](/source/Autonomic_nervous_system) before or without reaching consciousness to cause [pallor](/source/Pallor), [sweating](/source/Sweating), [tachycardia](/source/Tachycardia), [hypertension](/source/Hypertension), [lightheadedness](/source/Lightheadedness), [nausea](/source/Nausea), and [fainting](/source/Fainting).[7]

## System overview

This diagram linearly (unless otherwise mentioned) tracks the projections of all known structures that allow for pain, proprioception, thermoception, and chemoception to their relevant endpoints in the human brain. Click to enlarge.

This overview discusses [proprioception](/source/Proprioception), [thermoception](/source/Thermoception), chemoception, and nociception, as they are all integrally connected.

### Mechanical

See also: [Neuropathic pain](/source/Neuropathic_pain)

Proprioception is determined by using standard mechanoreceptors (especially [ruffini corpuscles](/source/Ruffini_corpuscle) (stretch) and [transient receptor potential channels](/source/Transient_receptor_potential_channel) (TRP channels). Proprioception is completely covered within the [somatosensory system](/source/Somatosensory_system), as the brain processes them together.

Thermoception refers to stimuli of moderate temperatures 24–28 °C (75–82 °F), as anything beyond that range is considered pain and moderated by nociceptors. TRP and potassium channels [TRPM (1-8), TRPV (1-6), TRAAK, and TREK] each respond to different temperatures (among other stimuli), which create action potentials in nerves that join the mechano (touch) system in the posterolateral tract. Thermoception, like proprioception, is then covered by the somatosensory system.[8][9][10][11][12]

TRP channels that detect noxious stimuli (mechanical, thermal, and chemical pain) relay that information to nociceptors that generate an action potential. Mechanical TRP channels react to depression of their cells (like touch), thermal TRPs change shape in different temperatures, and chemical TRPs act like [taste buds](/source/Taste_bud), signalling if their receptors bond to certain elements/chemicals.

### Neural

- [Laminae 3-5](/source/Nucleus_proprius_of_spinal_cord) make up [nucleus proprius](/source/Nucleus_proprius) in spinal grey matter.

- [Lamina 2](/source/Rexed_lamina) makes up [substantia gelatinosa of Rolando](/source/Substantia_gelatinosa_of_Rolando), unmyelinated spinal grey matter. Substantia receives input from nucleus proprius and conveys intense, poorly localized pain.

- [Lamina 1](/source/Marginal_nucleus_of_spinal_cord) primarily project to the [parabrachial area](/source/Parabrachial_area) and [periaqueductal grey](/source/Periaqueductal_gray), which begins the suppression of pain via neural and hormonal inhibition. Lamina 1 receive input from thermoreceptors via the [posterolateral tract](/source/Posterolateral_tract). Marginal nucleus of the spinal cord are the only unsuppressible pain signals.

- The [parabrachial area](/source/Parabrachial_area) integrates taste and pain info, then relays it. Parabrachial checks if the pain is being received in normal temperatures and if the [gustatory system](/source/Gustatory_system) is active; if both are so the pain is assumed to be due to poison.

- [Ao fibers](https://en.wikipedia.org/w/index.php?title=Ao_fiber&action=edit&redlink=1) synapse on laminae 1 and 5 while [Ab](/source/Ab_fiber) synapses on 1, 3, 5, and C. [C fibers](/source/C_fiber) exclusively synapse on lamina 2.[13][14]

- The [amygdala](/source/Amygdala) and [hippocampus](/source/Hippocampus) create and encode the memory and emotion due to pain stimuli.

- The [hypothalamus](/source/Hypothalamus) signals for the release of hormones that make pain suppression more effective; some of these are sex hormones.

- [Periaqueductal grey](/source/Periaqueductal_grey) (with hypothalamic hormone aid) hormonally signals [reticular formation](/source/Reticular_formation)'s [raphe nuclei](/source/Raphe_nucleus) to produce [serotonin](/source/Serotonin) that inhibits laminae pain nuclei.[15]

- [Lateral spinothalamic tract](/source/Lateral_spinothalamic_tract) aids in localization of pain.

- [Spinoreticular](/source/Spinoreticular_tract) and [spinotectal tracts](/source/Spinotectal_tract) are merely relay tracts to the [thalamus](/source/Thalamus) that aid in the perception of pain and alertness towards it. Fibers cross over (left becomes right) via the spinal [anterior white commissure](/source/Anterior_white_commissure).

- [Lateral lemniscus](/source/Lateral_lemniscus) is the first point of integration of sound and pain information.[16]

- [Inferior colliculus](/source/Inferior_colliculus) (IC) aids in sound orienting to pain stimuli.[17]

- [Superior colliculus](/source/Superior_colliculus) receives IC's input, integrates visual orienting info, and uses the balance topographical map to orient the body to the pain stimuli.[18][19]

- [Inferior cerebellar peduncle](/source/Inferior_cerebellar_peduncle) integrates proprioceptive info and outputs to the [vestibulocerebellum](/source/Vestibulocerebellum). The peduncle is not part of the lateral-spinothalamic-tract-pathway; the medulla receives the info and passes it onto the peduncle from elsewhere (see [somatosensory system](/source/Somatosensory_system)).

- The [thalamus](/source/Thalamus) is where pain is thought to be brought into [perception](/source/Perception); it also aids in pain suppression and modulation, acting like a [bouncer](/source/Bouncer_(doorman)), allowing certain intensities through to the cerebrum and rejecting others.[20]

- The [somatosensory cortex](/source/Somatosensory_cortex) decodes nociceptor info to determine the exact location of pain and is where proprioception is brought into consciousness; inferior cerebellar peduncle is all unconscious proprioception.

- [Insula](/source/Insular_cortex) judges the intensity of the pain and provides the ability to imagine pain.[21][22]

- [Cingulate cortex](/source/Cingulate_cortex) is presumed to be the memory hub for pain.[23]

## In non-mammals

Further information: [Pain in animals](/source/Pain_in_animals), [Pain in amphibians](/source/Pain_in_amphibians), [Pain in fish](/source/Pain_in_fish), [Pain in invertebrates](/source/Pain_in_invertebrates), [Pain in crustaceans](/source/Pain_in_crustaceans), and [Pain in cephalopods](/source/Pain_in_cephalopods)

Nociception has been documented in other animals, including fish[24] and a wide range of [invertebrates](/source/Pain_in_invertebrates),[25] including [leeches](/source/Leech),[26] [nematode](/source/Nematode) worms,[27] [sea slugs](/source/Sea_slug),[28] and [fruit flies](/source/Drosophilidae).[29] As in mammals, nociceptive neurons in these species are typically characterized by responding preferentially to high temperature (40 °C (104 °F) or more), low pH, [capsaicin](/source/Capsaicin), and [tissue damage](/source/Tissue_damage).

## History of term

The term "nociception" was coined by [Charles Scott Sherrington](/source/Charles_Scott_Sherrington) to distinguish the physiological process (nervous activity) from pain (a subjective experience).[30] It is derived from the Latin verb *nocēre*, which means "to harm".

## See also

- [Electroreception](/source/Electroreception) – Biological electricity-related abilitiesPages displaying short descriptions of redirect targets

- [Mechanoreceptor](/source/Mechanoreceptor) – Sensory receptor cell responding to mechanical pressure or strain

- [Thermoception](/source/Thermoception) – Sensation and perception of temperature

- [Proprioception](/source/Proprioception) – Sense of self-movement, force, and body position

## References

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v t e Sensation and perception Processes and concepts Sensation Stimulus Sensory receptor Transduction (physiology) Sensory processing Active sensory system Perception Multimodal integration Awareness Consciousness Cognition Feeling Motion perception Qualia Human External Sensory organs Eyes Ears Inner ear Nose Mouth Skin Sensory systems Visual system (sense of vision) Auditory system (sense of hearing) Vestibular system (sense of balance) Olfactory system (sense of smell) Gustatory system (sense of taste) Somatosensory system (sense of touch) Sensory cranial and spinal nerves Optic (II) Vestibulocochlear (VIII) Olfactory (I) Facial (VII) Glossopharyngeal (IX) Trigeminal (V) Spinal Cerebral cortices Visual cortex Auditory cortex Vestibular cortex Olfactory cortex Gustatory cortex Somatosensory cortex Perceptions Visual perception (vision) Color Auditory perception (hearing) Equilibrioception (balance) Olfaction (smell) Gustation (taste or flavor) Touch mechanoreception nociception (pain) thermoception Internal Proprioception Hunger Thirst Suffocation Nausea Visceral pain Nonhuman Animal Electroreception Magnetoreception Echolocation Infrared sensing in vampire bats Infrared sensing in snakes Surface wave detection Frog hearing Toad vision Plant Photomorphogenesis Gravitropism Artificial Robotic sensing Computer vision Machine hearing Types of sensory receptors Mechanoreceptor Baroreceptor Mechanotransduction Lamellar corpuscle Tactile corpuscle Merkel nerve ending Bulbous corpuscle Campaniform sensilla Slit sensilla Stretch receptor Photoreceptor Photoreceptor cell Cone cell Rod cell ipRGC Photopigment Aureochrome Chemoreceptor Taste receptor Olfactory receptor Osmoreceptor Thermoreceptor Cilium TRP channels Nociceptor Nociceptin receptor Juxtacapillary receptor Disorders Visual Visual impairment Alice in Wonderland syndrome Amaurosis Anopsia Color blindness Diplopia Hemeralopia and Nyctalopia Optic neuropathy Oscillopsia Palinopsia Papilledema Photophobia Photopsia Polyopia Scotoma Stereoblindness Visual snow Auditory Amblyaudia Auditory agnosia Auditory hallucination Auditory verbal agnosia Cortical deafness Hearing loss Microwave auditory effect Music-specific disorders Palinopsia Spatial hearing loss Tinnitus Vestibular Vertigo BPPV Labyrinthine fistula Labyrinthitis Ménière's disease Olfactory Anosmia Dysosmia Hyperosmia Hyposmia Olfactory reference syndrome Parosmia Phantosmia Gustatory Ageusia Hypergeusia Hypogeusia Parageusia Tactile Astereognosis CMT disease Formication Hyperesthesia Hypoesthesia Paresthesia Tactile hallucination Nociception (pain) Hyperalgesia Hypoalgesia Pain dissociation Phantom pain Proprioception Asomatognosia Phantom limb syndrome Sensory ataxia Somatoparaphrenia Supernumerary phantom limb Multimodal Aura Agnosia Allochiria Derealization Hallucination HSAN Sensory processing disorder Synesthesia Biases and errors Illusion · Pareidolia Category

v t e Pain By region/system Head and neck Eye strain Headache Neck Odynophagia (swallowing) Toothache Respiratory system Sore throat Pleurodynia Musculoskeletal Arthralgia (joint) Bone pain Myalgia (muscle) Acute Delayed-onset Neurologic Neuralgia Pain asymbolia Pain disorder Paroxysmal extreme pain disorder Allodynia Chronic pain Hyperalgesia Hypoalgesia Hyperpathia Phantom pain Referred pain Congenital insensitivity to pain congenital insensitivity to pain with anhidrosis congenital insensitivity to pain with partial anhidrosis Other Pelvic pain Proctalgia Back Low back pain Measurement and testing Pain scale Cold pressor test Dolorimeter Grimace scale Hot plate test Tail flick test Visual analogue scale Pathophysiology Nociception Anterolateral system Posteromarginal nucleus Substance P Management Analgesia Anesthesia Cordotomy Pain eradication Related concepts Pain threshold Pain tolerance Suffering SOCRATES Philosophy of pain Cancer pain Drug-seeking behavior

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Adapted from the Wikipedia article [Nociception](https://en.wikipedia.org/wiki/Nociception) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Nociception?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.