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Proprioception

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Proprioception (/ˌproʊpri.oʊˈsɛpʃən, -ə-/<a href="#cite_note-2">[2]</a><a href="#cite_note-3">[3]</a> PROH-pree-oh-SEP-shən, -⁠ə-) is the <a href="Sense" title="Sense">sense</a> of self-movement, force, and body position.<a href="#cite_note-:3-1">[1]</a><a href="#cite_note-4">[4]</a>

Proprioception is mediated by proprioceptors, a type of <a href="Sensory_receptor" class="mw-redirect" title="Sensory receptor">sensory receptor</a>, located within <a href="Muscle" title="Muscle">muscles</a>, <a href="Tendon" title="Tendon">tendons</a>, and <a href="Joint" title="Joint">joints</a>.<a href="#cite_note-:3-1">[1]</a> Most animals possess multiple subtypes of proprioceptors, which detect distinct kinesthetic parameters, such as joint position, movement, and load. Although all mobile animals possess proprioceptors, the structure of the sensory organs can vary across species.

Proprioceptive signals are transmitted to the <a href="Central_nervous_system" title="Central nervous system">central nervous system</a>, where they are integrated with information from other <a href="Sensory_nervous_system" title="Sensory nervous system">sensory systems</a>, such as <a href="Visual_perception" title="Visual perception">the visual system</a> and the <a href="Vestibular_system" title="Vestibular system">vestibular system</a>, to create an overall representation of body position, movement, and acceleration. In many animals, sensory feedback from proprioceptors is essential for stabilizing body posture and coordinating body movement.

<div class="mw-heading mw-heading2"><h2 id="System_overview">System overview</h2></div>

In vertebrates, limb movement and velocity (muscle length and the rate of change) are encoded by one group of sensory neurons (<a href="Type_Ia_sensory_fiber" title="Type Ia sensory fiber">type Ia sensory fiber</a>) and another type encode static muscle length (<a href="Type_II_sensory_fiber" title="Type II sensory fiber">group II neurons</a>).<a href="#cite_note-5">[5]</a> These two types of sensory neurons compose <a href="Muscle_spindles" class="mw-redirect" title="Muscle spindles">muscle spindles</a>. There is a similar division of encoding in invertebrates; different subgroups of neurons of the <a href="Chordotonal_organ" title="Chordotonal organ">chordotonal organ</a><a href="#cite_note-6">[6]</a> encode limb position and velocity.

To determine the load on a limb, vertebrates use sensory neurons in the Golgi tendon organs:<a href="#cite_note-7">[7]</a> type Ib afferents. These proprioceptors are activated at given muscle forces, which indicate the resistance that muscle is experiencing. Similarly, invertebrates have a mechanism to determine limb load: the <a href="Campaniform_sensilla" title="Campaniform sensilla">campaniform sensilla</a>.<a href="#cite_note-8">[8]</a> These proprioceptors are active when a limb experiences resistance.<a href="#cite_note-:3-1">[1]</a>

A third role for proprioceptors is to determine when a joint is at a specific position. In vertebrates, this is accomplished by <a href="Ruffini_ending" class="mw-redirect" title="Ruffini ending">Ruffini endings</a> and <a href="Pacinian_corpuscles" class="mw-redirect" title="Pacinian corpuscles">Pacinian corpuscles</a>. These proprioceptors are activated when the joint is at a threshold position, usually at the extremes of joint position. Invertebrates use <a href="Hair_plate" title="Hair plate">hair plates</a><a href="#cite_note-9">[9]</a> to accomplish this; a field of bristles located within joints that detects the relative movement of limb segments through the deflection of the associated cuticular hairs.

<div class="mw-heading mw-heading3"><h3 id="Reflexes">Reflexes</h3></div>

The sense of proprioception is ubiquitous across mobile animals and is essential for the motor coordination of the body. Proprioceptors can form reflex circuits with motor neurons to provide rapid feedback about body and limb position. These <a href="Mechanosensation" title="Mechanosensation">mechanosensation</a> circuits are important for flexibly maintaining posture and balance, especially during locomotion. For example, consider the <a href="Stretch_reflex" title="Stretch reflex">stretch reflex</a>, in which stretch across a muscle is detected by a sensory receptor (e.g., <a href="Muscle_spindle" title="Muscle spindle">muscle spindle</a>, <a href="Chordotonal_organ" title="Chordotonal organ">chordotonal neurons</a>), which activates a motor neuron to induce muscle contraction and oppose the stretch. During locomotion, sensory neurons can reverse their activity when stretched, to promote rather than oppose movement.<a href="#cite_note-10">[10]</a><a href="#cite_note-11">[11]</a>

<div class="mw-heading mw-heading3"><h3 id="Conscious_and_nonconscious">Conscious and nonconscious</h3></div>

In humans, a distinction is made between conscious proprioception and nonconscious proprioception:

<ul><li>Conscious proprioception is communicated by the <a href="Dorsal_column-medial_lemniscus_pathway" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">dorsal column-medial lemniscus pathway</a> to the <a href="Cerebrum" title="Cerebrum">cerebrum</a>.<a href="#cite_note-isbn0-7817-2829-0-12">[12]</a></li>

<li>Nonconscious proprioception is communicated primarily via the <a href="Dorsal_spinocerebellar_tract" class="mw-redirect" title="Dorsal spinocerebellar tract">dorsal spinocerebellar tract</a><a href="#cite_note-titleChapter_7A:_Somatosensory_Systems-13">[13]</a> and <a href="Ventral_spinocerebellar_tract" class="mw-redirect" title="Ventral spinocerebellar tract">ventral spinocerebellar tract</a>,<a href="#cite_note-14">[14]</a> to the <a href="Cerebellum" title="Cerebellum">cerebellum</a>.</li>

<li>A nonconscious reaction is seen in the human proprioceptive reflex, or <a href="Righting_reflex" title="Righting reflex">righting reflex</a>—in the event that the body tilts in any direction, the person will cock their head back to level the eyes against the horizon.<a href="#cite_note-15">[15]</a> This is seen even in infants as soon as they gain control of their neck muscles. This control comes from the <a href="Cerebellum" title="Cerebellum">cerebellum</a>, the part of the brain affecting balance.</li></ul>

<div class="mw-heading mw-heading2"><h2 id="Physiology">Physiology</h2></div>

Proprioception is mediated by mechanically sensitive proprioceptor neurons distributed throughout an animal's body. Most vertebrates possess three basic types of proprioceptors: <a href="Muscle_spindle" title="Muscle spindle">muscle spindles</a>, which are embedded in <a href="Skeletal_muscle" title="Skeletal muscle">skeletal muscles</a>, <a href="Golgi_tendon_organ" title="Golgi tendon organ">Golgi tendon organs</a>, which lie at the interface of muscles and tendons, and joint receptors, which are <a href="Mechanosensation#Cutaneous_Mechanoreceptors" title="Mechanosensation">low-threshold mechanoreceptors</a> embedded in <a href="Joint_capsule" title="Joint capsule">joint capsules</a>. Many invertebrates, such as insects, also possess three basic proprioceptor types with analogous functional properties: <a href="Chordotonal_organ" title="Chordotonal organ">chordotonal neurons</a>, <a href="Campaniform_sensilla" title="Campaniform sensilla">campaniform sensilla</a>, and <a href="Hair_plate" title="Hair plate">hair plates</a>.<a href="#cite_note-:3-1">[1]</a>

The initiation of proprioception is the activation of a proprioceptor in the periphery.<a href="#cite_note-16">[16]</a> The proprioceptive sense is believed to be composed of information from <a href="Sensory_neuron" title="Sensory neuron">sensory neurons</a> located in the <a href="Labyrinth_(inner_ear)" class="mw-redirect" title="Labyrinth (inner ear)">inner ear</a> (motion and orientation) and in the <a href="Stretch_receptor" title="Stretch receptor">stretch receptors</a> located in the <a href="Muscle_spindle" title="Muscle spindle">muscles</a> and the joint-supporting ligaments (stance). There are specific nerve receptors for this form of perception termed "proprioceptors", just as there are specific receptors for pressure, light, temperature, sound, and other sensory experiences. Proprioceptors are sometimes known as <a href="Adequate_stimulus" title="Adequate stimulus">adequate stimuli</a> receptors.

Members of the <a href="Transient_receptor_potential" class="mw-redirect" title="Transient receptor potential">transient receptor potential</a> family of <a href="Ion_channel" title="Ion channel">ion channels</a> have been found to be important for proprioception in <a href="Drosophila_melanogaster" title="Drosophila melanogaster">fruit flies</a>,<a href="#cite_note-17">[17]</a> <a href="Caenorhabditis_elegans" title="Caenorhabditis elegans">nematode worms</a>,<a href="#cite_note-18">[18]</a> <a href="African_clawed_frog" title="African clawed frog">African clawed frogs</a>,<a href="#cite_note-19">[19]</a> and <a href="Zebrafish" title="Zebrafish">zebrafish</a>.<a href="#cite_note-20">[20]</a> <a href="PIEZO2" title="PIEZO2">PIEZO2</a>, a nonselective cation channel, has been shown to underlie the mechanosensitivity of proprioceptors in mice.<a href="#cite_note-21">[21]</a> Humans with loss-of-function mutations in the PIEZO2 gene exhibit specific deficits in joint proprioception,<a href="#cite_note-piezoChannels-23">[a]</a> as well as vibration and touch discrimination, suggesting that the PIEZO2 channel is essential for mechanosensitivity in some proprioceptors and low-threshold mechanoreceptors.<a href="#cite_note-24">[23]</a>

Although it was known that finger kinesthesia relies on skin sensation, recent research has found that kinesthesia-based <a href="Haptic_perception" title="Haptic perception">haptic perception</a> relies strongly on the forces experienced during touch.<a href="#cite_note-25">[24]</a> This research allows the creation of "virtual", illusory haptic shapes with different perceived qualities.<a href="#cite_note-26">[25]</a>

<div class="mw-heading mw-heading3"><h3 id="Central_pattern_generators">Central pattern generators</h3></div>

Central pattern generators are groups of neurons in the spinal cord that are responsible for generating stereotyped movement. It has been shown that in cats, rhythmic activation patterns are still observed following removal of sensory afferents and removal of the brain.,<a href="#cite_note-27">[26]</a> indicating that there is neural pattern generation in the spinal cord independent of descending signals from the brain and sensory information. It is currently understood that the spinal cord receives sensory input from proprioceptive organs and descending commands from the brain, integrates these signals, and sends activation signals to muscle through alpha motoneurons and fusimotor signals through gamma motoneurons in a coordinated and rhythmic fashion.

<div class="mw-heading mw-heading3"><h3 id="Muscle_spindles">Muscle spindles</h3></div>

The <a href="Muscle_spindle" title="Muscle spindle">muscle spindle</a> is a proprioceptive organ that lies embedded in the muscle. It consists of bag- and chain-type fibers, which correspond to dynamic and static responses, respectively. Spindles relay information through primary (Group Ia) and secondary (Group II) sensory afferents, with the primary afferent attached at the nucleus of the spindle and the secondary afferent attached at the end of the spindle. Spindles are conventionally thought of as encoding muscle length, velocity, and acceleration, however there is evidence to suggest that they respond to the force and yank (the first time-derivative of force) exerted on intrafusal muscle. Spindles are also composed of bag- and chain-type fibers, with dynamic and static stretch responses, respectively.<a href="#cite_note-:1a-28">[27]</a><a href="#cite_note-:2a-29">[28]</a><a href="#cite_note-ReferenceA-30">[29]</a>

Key features of muscle spindle firing responses include initial bursts, history-dependence, and rate relaxation. Initial bursts occur at the onset of stretch and only last a very short time. History dependence refers to how the response of muscle spindles is affected by past stretch inputs. Rate relaxation refers to how the firing rate of muscle spindles decreases over time when held at a constant length.<a href="#cite_note-:1a-28">[27]</a><a href="#cite_note-:2a-29">[28]</a>

<div class="mw-heading mw-heading3"><h3 id="Golgi_tendon_organs">Golgi tendon organs</h3></div>

The <a href="Golgi_tendon_organ" title="Golgi tendon organ">Golgi tendon organ</a> (GTO) is a proprioceptive organ that lies at the muscle-tendon junction. GTOs relay information through group Ib afferents, and encode active muscle force. As they are connected at one end to motor units, individual GTOs only relay information on a few fibers. At the same time, GTOs exhibit self-adaptation, in which GTO response decreases after prior activation, and cross-adaptation, in which GTO activity is modulated by prior activation of another GTO. Similar to muscle spindles, GTO firing is characterized by a heightened response at the onset of activity (dynamic response) and gradual relaxation to a resting firing rate (static response).<a href="#cite_note-31">[30]</a><a href="#cite_note-32">[31]</a>

<div class="mw-heading mw-heading3"><h3 id="Fusimotor_system">Fusimotor system</h3></div>

While muscle spindles relay information via primary afferents, they receive descending efferent signals from the spinal cord via <a href="Gamma_motor_neuron" title="Gamma motor neuron">gamma motoneurons</a>. This gamma innervation modulates the sensitivity of muscle spindle afferents to stretch. In cat studies, muscle spindle afferent firing rates with gamma fusimotor innervation were shown to be approximately equal to the sum of the gamma motoneuron firing rate and muscle spindle firing rate with no gamma innervation. In these same studies, gamma activity was shown to be correlated with joint angle during locomotion, indicating that fusimotor activity is periodically modulated during locomotion. Similar to muscle spindles, gamma motoneurons are also categorized according to static and dynamic response properties.<a href="#cite_note-33">[32]</a><a href="#cite_note-34">[33]</a>

<div class="mw-heading mw-heading3"><h3 id="Motor_control">Motor control</h3></div>

In motor control, proprioceptors provide critical feedback to the central nervous system. Muscle spindles relay information regarding muscle stretch, Golgi tendon organs relay information regarding tendon force, and gamma motoneurons modulate muscle spindle feedback. Afferent signals from spindles and tendon organs are integrated in the spinal cord, which then output muscle activation commands to muscle via alpha motoneurons. Because muscle spindles and tendon organs exhibit burst-like activity in response to rapid stretch, they play a vital role in reflexive perturbation responses. In a simulation study, it has been shown that the controllability of a limb in response to a perturbation is significantly increased when utilizing muscle spindle and tendon organ feedback in conjunction.<a href="#cite_note-35">[34]</a> However, proprioceptive feedback is also critical in controlling steady movements. In one study, de-afferented mice were unable to walk as quickly as the control group, and showed some reduced activity in extensor muscles.<a href="#cite_note-36">[35]</a> It's also been shown in cats that disruption of feedback from muscle spindles impairs inter-joint coordination during ramp descent tasks.<a href="#cite_note-37">[36]</a> In a study on people with amputations, those with a higher degree of proprioceptive feedback from muscle spindles were able to better control the movement of a virtual limb.<a href="#cite_note-:0a-38">[37]</a>

<div class="mw-heading mw-heading2"><h2 id="Anatomy">Anatomy</h2></div>

Proprioception of the head stems from the muscles innervated by the <a href="Trigeminal_nerve" title="Trigeminal nerve">trigeminal nerve</a>, where the <a href="General_somatic_afferent_fibers" class="mw-redirect" title="General somatic afferent fibers">general somatic afferent fibers</a> pass without synapsing in the <a href="Trigeminal_ganglion" title="Trigeminal ganglion">trigeminal ganglion</a> (first-order sensory neuron), reaching the mesencephalic tract and the <a href="Mesencephalic_nucleus_of_trigeminal_nerve" title="Mesencephalic nucleus of trigeminal nerve">mesencephalic nucleus of trigeminal nerve</a>.<a href="#cite_note-Arslan2014-39">[38]</a> Proprioception of limbs often occurs due to receptors in <a href="Connective_tissue" title="Connective tissue">connective tissue</a> near joints.<a href="#cite_note-40">[39]</a>

<div class="mw-heading mw-heading2"><h2 id="Function">Function</h2></div>

<div class="mw-heading mw-heading3"><h3 id="Stability">Stability</h3></div>

An important role for proprioception is to allow an animal to stabilize itself against perturbations.<a href="#cite_note-MageeZachazewskiQuillen2008-41">[40]</a> For instance, for a person to walk or stand upright, they must continuously monitor their posture and adjust muscle activity as needed to provide balance. Similarly, when walking on unfamiliar terrain, or even tripping, the person must adjust the output of their muscles quickly based on estimated limb position and velocity. <a href="#Reflexes">Proprioceptor reflex circuits</a> are thought to play an important role to allow fast and unconscious execution of these behaviors. To make control of these behaviors efficient, proprioceptors are also thought to regulate reciprocal inhibition in muscles, leading to <a href="Anatomical_terms_of_muscle#Agonist-antagonist_pairs" title="Anatomical terms of muscle">agonist-antagonist muscle pairs</a>.<a href="#cite_note-proske-42">[41]</a>

<div class="mw-heading mw-heading3"><h3 id="Planning_and_refining_movements">Planning and refining movements</h3></div>

When planning complex movements such as reaching or <a href="Personal_grooming" title="Personal grooming">grooming</a>, an animal must consider the current position and velocity of its limb and use that information to adjust dynamics to target a final position. If the animal's estimate of its limb's initial position is wrong, then a deficiency in the movement can result. Furthermore, proprioception is crucial in refining the movement if it deviates from the trajectory.<a href="#cite_note-proske-42">[41]</a><a href="#cite_note-43">[42]</a>

<div class="mw-heading mw-heading2"><h2 id="Development">Development</h2></div>

In adult fruit flies, each proprioceptor class arises from a specific <a href="Cell_lineage" title="Cell lineage">cell lineage</a> (i.e. each chordotonal neuron is from the chordotonal neuron lineage, although multiple lineages give rise to sensory bristles). After the last cell division, proprioceptors send out axons toward the central nervous system and are guided by hormonal gradients to reach stereotyped synapses.<a href="#cite_note-44">[43]</a>

The mechanisms underlying <a href="Axon_guidance" title="Axon guidance">axon guidance</a> are similar across invertebrates and vertebrates.<a href="#cite_note-45">[44]</a>

In mammals with longer gestation periods, <a href="Muscle_spindles" class="mw-redirect" title="Muscle spindles">muscle spindles</a> are fully formed at birth. Muscle spindles continue to grow throughout post-natal development as muscles grow.<a href="#cite_note-pmid9074933-46">[45]</a>

<div class="mw-heading mw-heading2"><h2 id="Mathematical_models">Mathematical models</h2></div>

Proprioceptors transfer the mechanical state of the body into patterns of neural activity. This transfer can be modeled mathematically, for example to better understand the internal workings of a proprioceptor<a href="#cite_note-:4-47">[46]</a><a href="#cite_note-:5-48">[47]</a><a href="#cite_note-:6-49">[48]</a> or to provide more realistic feedback in neuromechanical simulations.<a href="#cite_note-50">[49]</a><a href="#cite_note-51">[50]</a>

Various proprioceptor models of complexity have been developed. They range from simple phenomenological models to complex structural models, in which the mathematical elements correspond to anatomical features of the proprioceptor. The focus has been on <a href="Muscle_spindle" title="Muscle spindle">muscle spindles</a>,<a href="#cite_note-:4-47">[46]</a><a href="#cite_note-:5-48">[47]</a><a href="#cite_note-:6-49">[48]</a><a href="#cite_note-52">[51]</a> but <a href="Golgi_tendon_organ" title="Golgi tendon organ">Golgi tendon organs</a><a href="#cite_note-53">[52]</a><a href="#cite_note-:7-54">[53]</a> and insects' <a href="Hair_plate" title="Hair plate">hair plates</a><a href="#cite_note-55">[54]</a> have been modeled too.

<div class="mw-heading mw-heading3"><h3 id="Muscle_spindles_2">Muscle spindles</h3></div>

Poppele and Bowman <a href="#cite_note-56">[55]</a> used <a href="Linear_time-invariant_system" title="Linear time-invariant system">linear system theory</a> to model mammalian muscle spindles Ia and II afferents. They obtained a set of de-afferented muscle spindles, measured their response to a series of sinusoidal and step function stretches, and fit a transfer function to the spike rate. They found that the following <a href="Transfer_function" title="Transfer function">Laplace transfer function</a> describes the firing rate responses of the <a href="Type_Ia_sensory_fiber" title="Type Ia sensory fiber">primary sensory fibers</a> for a change in length:

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<annotation encoding="application/x-tex">{\displaystyle H(s)=K_{1}{\frac {s(s+0.44)(s+11.3)(s+44)}{(s+0.04)(s+0.816)}}}</annotation>

</semantics>

</math><img src="./4f8ffebf188b0ca52787a0db5f0f9b47e35f17a3.svg" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.671ex; width:40.829ex; height:6.509ex;" alt="{\displaystyle H(s)=K_{1}{\frac {s(s+0.44)(s+11.3)(s+44)}{(s+0.04)(s+0.816)}}}" loading="lazy">

The following equation describes the response of <a href="Type_II_sensory_fiber" title="Type II sensory fiber">secondary sensory fibers</a>:

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<annotation encoding="application/x-tex">{\displaystyle H(s)=K_{2}{\frac {(s+0.44)(s+11.3)}{s+0.816}}}</annotation>

</semantics>

</math><img src="./b1cefbeced1efda302d574db3e0bfccf96198809.svg" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.005ex; width:31.674ex; height:5.843ex;" alt="{\displaystyle H(s)=K_{2}{\frac {(s+0.44)(s+11.3)}{s+0.816}}}" loading="lazy">

More recently, Blum et al.<a href="#cite_note-57">[56]</a> showed that the muscle spindle firing rate is modeled better as tracking the force of the muscle, rather than the length. Furthermore, muscle spindle firing rates show history dependence which cannot be modeled by a linear time-invariant system model.

<div class="mw-heading mw-heading3"><h3 id="Golgi_tendon_organs_2">Golgi tendon organs</h3></div>

Houk and Simon <a href="#cite_note-:7-54">[53]</a> provided one of the first mathematical models of a Golgi tendon organ receptor, modeling the firing rate of the receptor as a function of the muscle tension force. Just as for muscle spindles, they find that, as the receptors respond linearly to sine waves of different frequencies and has little variance in response over time to the same stimulus, Golgi tendon organ receptors may be modeled as linear time-invariant systems. Specifically, they find that the firing rate of a Golgi tendon organ receptor may be modeled as a sum of 3 decaying exponentials:

</mstyle>

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<annotation encoding="application/x-tex">{\displaystyle r(t)=K[1+A\exp(-at)+B\exp(-bt)+C\exp(-ct)]u(t)}</annotation>

</semantics>

</math><img src="./e10f5a80eebc1deaa42680f0d6a415eae38941ea.svg" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:57.515ex; height:2.843ex;" alt="{\displaystyle r(t)=K[1+A\exp(-at)+B\exp(-bt)+C\exp(-ct)]u(t)}" loading="lazy">

where <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle r(t)}">

</mstyle>

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<annotation encoding="application/x-tex">{\displaystyle r(t)}</annotation>

</semantics>

</math><img src="./ec653b463c709d42cb85133595bf0da29801f6e5.svg" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:3.698ex; height:2.843ex;" alt="{\displaystyle r(t)}" loading="lazy"> is the firing rate and <math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle u(t)}">

</mstyle>

</mrow>

<annotation encoding="application/x-tex">{\displaystyle u(t)}</annotation>

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</math><img src="./b375df3b65d282f8715835dc91ccb22f46993959.svg" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:3.979ex; height:2.843ex;" alt="{\displaystyle u(t)}" loading="lazy"> is a step function of force.

The corresponding Laplace transfer function for this system is:

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<annotation encoding="application/x-tex">{\displaystyle H(s)=K\left(1+{\frac {As}{s+a}}+{\frac {Bs}{s+b}}+{\frac {Cs}{s+c}}\right)}</annotation>

</semantics>

</math><img src="./04129a81bda63a30fd79b3ba6e4358e177f7d617.svg" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.505ex; width:41.155ex; height:6.176ex;" alt="{\displaystyle H(s)=K\left(1+{\frac {As}{s+a}}+{\frac {Bs}{s+b}}+{\frac {Cs}{s+c}}\right)}" loading="lazy">

For a <a href="Soleus_muscle" title="Soleus muscle">soleus</a> receptor, Houk and Simon obtain average values of K=57 pulses/sec/kg, A=0.31, a=0.22 sec−1, B=0.4, b=2.17 sec−1, C=2.5, c=36 sec−1 .

When modeling a cat stretch reflex, Lin and Crago<a href="#cite_note-58">[57]</a> improved upon this model by adding a logarithmic nonlinearity before the Houk and Simon model and a <a href="Rectifier_(neural_networks)" title="Rectifier (neural networks)">threshold nonlinearity</a> after.

<div class="mw-heading mw-heading2"><h2 id="Impairment">Impairment</h2></div>

Proprioceptive feedback is also linked to motor deficits in Parkinson's disease and cerebral palsy. People with cerebral palsy often suffer from spasticity due to hyperreflexia.<a href="#cite_note-59">[58]</a> A common clinical test of spasticity is the pendulum test, in which the subject remains seated and the relaxed leg is dropped from horizontal. In individuals with spasticity, the leg comes to rest much more quickly due to increased reflexive muscle contraction.

Computational models have shown that results from pendulum tests in children with spastic cerebral palsy are explained by increased muscle tone, short-range stiffness, and increased stretch reflex responses due to increased muscle force feedback.<a href="#cite_note-60">[59]</a> Pendulum test results are also dependent on prior motion, indicating that muscle spindle feedback is a large component of spastic movement due to the history-dependent behavior of muscle spindles.<a href="#cite_note-61">[60]</a> Increased proprioceptive feedback has also explained properties of gait in children with spastic cerebral palsy<a href="#cite_note-ReferenceA-30">[29]</a>

In addition to functional impairments, proprioceptive deficits are linked to compensatory adaptations in the central nervous system. In the study on people with amputations mentioned previously, those with a lower degree of proprioception showed stronger connectivity between their visual and motor cortices, which is interpreted as a greater reliance on visual feedback to coordinate movement. Those with higher degrees of proprioception also showed higher connectivity between brain regions associated with sensorimotor feedback and sensory integration.<a href="#cite_note-:0a-38">[37]</a>

<div class="mw-heading mw-heading3"><h3 id="Chronic">Chronic</h3></div>

Proprioception, a sense vital for rapid and proper body coordination,<a href="#cite_note-62">[61]</a> can be permanently lost or impaired as a result of genetic conditions, disease, viral infections, and injuries. For instance, patients with joint hypermobility or <a href="Ehlers%E2%80%93Danlos_syndromes" class="mw-redirect" title="Ehlers–Danlos syndromes">Ehlers–Danlos syndromes</a>, genetic conditions that result in weak connective tissue throughout the body, have chronic impairments to proprioception.<a href="#cite_note-Castori_2012-63">[62]</a> <a href="Autism_spectrum" class="mw-redirect" title="Autism spectrum">Autism spectrum disorder</a><a href="#cite_note-64">[63]</a> and <a href="Parkinson's_disease" title="Parkinson's disease">Parkinson's disease</a> can also cause chronic disorder of proprioception.<a href="#cite_note-pmid19592360-65">[64]</a> In regards to Parkinson's disease, it remains unclear whether the proprioceptive-related decline in motor function occurs due to disrupted proprioceptors in the periphery or signaling in the spinal cord or brain.

In rare cases, viral infections result in a loss of proprioception. Ian Waterman and Charles Freed are two such people that lost their sense of proprioception from the neck down from supposed viral infections (i.e. gastric flu and a rare viral infection). After losing their sense of proprioception, Ian and Charles could move their lower body, but could not coordinate their movements. However, both individuals regained some control of their limbs and body by consciously planning their movements and relying solely on visual feedback. Interestingly, both individuals can still sense pain and temperature, indicating that they specifically lost proprioceptive feedback, but not tactile and nociceptive feedback. The impact of losing the sense of proprioception on daily life is perfectly illustrated when Ian Waterman stated, "What is an active brain without mobility".<a href="#cite_note-66">[65]</a><a href="#cite_note-67">[66]</a>

Proprioception is also permanently lost in people who lose a limb or body part through injury or amputation. After the removal of a limb, people may have a confused sense of that limb's existence on their body, known as <a href="Phantom_limb" title="Phantom limb">phantom limb syndrome</a>. Phantom sensations can occur as passive proprioceptive sensations of the limb's presence, or more active sensations such as perceived movement, pressure, pain, itching, or temperature. There are a variety of theories concerning the etiology of <a href="Phantom_limb" title="Phantom limb">phantom limb</a> sensations and experience. One is the concept of "proprioceptive memory", which argues that the brain retains a memory of specific limb positions and that after amputation there is a conflict between the visual system, which actually sees that the limb is missing, and the memory system which remembers the limb as a functioning part of the body.<a href="#cite_note-68">[67]</a> Phantom sensations and phantom pain may also occur after the removal of body parts other than the limbs, such as after amputation of the breast, extraction of a tooth (phantom tooth pain), or removal of an eye (<a href="Phantom_eye_syndrome" title="Phantom eye syndrome">phantom eye syndrome</a>).

There is a decline in the sense of proprioception with <a href="Ageing" title="Ageing">ageing</a>. This can often result in chronic lower back pain, and be the cause of falls in the elderly.<a href="#cite_note-Sakai-69">[68]</a>

<div class="mw-heading mw-heading3"><h3 id="Acute">Acute</h3></div>

Proprioception is occasionally impaired spontaneously, especially when one is tired. Similar effects can be felt during the <a href="Hypnagogia#Other_sensations" title="Hypnagogia">hypnagogic state of consciousness</a>, during the onset of sleep. One's body may feel too large or too small, or parts of the body may feel distorted in size. Similar effects can sometimes occur during <a href="Epilepsy" title="Epilepsy">epilepsy</a> or <a href="Migraine" title="Migraine">migraine</a> <a href="Aura_(symptom)" title="Aura (symptom)">auras</a>. These effects are presumed to arise from abnormal stimulation of the part of the <a href="Parietal_cortex" class="mw-redirect" title="Parietal cortex">parietal cortex</a> of the <a href="Brain" title="Brain">brain</a> involved with integrating information from different parts of the body.<a href="#cite_note-70">[69]</a> Proprioceptive illusions can also be induced, such as the "Pinocchio illusion", the illusion that one's nose is growing longer. <a href="#cite_note-g625-71">[70]</a>

Temporary impairment of proprioception has also been known to occur from an overdose of <a href="Vitamin_B6" title="Vitamin B6">vitamin B6</a> (pyridoxine and pyridoxamine). This is due to a reversible neuropathy.<a href="#cite_note-72">[71]</a> Most of the impaired function returns to normal shortly after the amount of the vitamin in the body returns to a level that is closer to that of the physiological norm. Impairment can also be caused by <a href="Cytotoxicity" title="Cytotoxicity">cytotoxic</a> factors such as <a href="Chemotherapy" title="Chemotherapy">chemotherapy</a>.

It has been proposed that even common <a href="Tinnitus" title="Tinnitus">tinnitus</a> and the attendant hearing frequency-gaps masked by the perceived sounds may cause erroneous proprioceptive information to the balance and comprehension centers of the brain, precipitating mild confusion. <a href="#cite_note-a306-73">[72]</a>

Temporary loss or impairment of proprioception may happen periodically during growth, mostly during adolescence. Growth that might also influence this would be large increases or drops in bodyweight/size due to fluctuations of fat (<a href="Liposuction" title="Liposuction">liposuction</a>, rapid <a href="Weight_loss" title="Weight loss">fat loss</a> or gain) and/or muscle content (<a href="Bodybuilding" title="Bodybuilding">bodybuilding</a>, <a href="Anabolic_steroid" title="Anabolic steroid">anabolic steroids</a>, <a href="Catabolism" title="Catabolism">catabolisis</a>/<a href="Starvation" title="Starvation">starvation</a>).<a href="#cite_note-74">[73]</a> It can also occur in those that gain new levels of <a href="Flexibility_(anatomy)" title="Flexibility (anatomy)">flexibility</a>, <a href="Stretching" title="Stretching">stretching</a>, and <a href="Contortion" title="Contortion">contortion</a>. A limb's being in a new range of motion never experienced (or at least, not for a long time since youth perhaps) can disrupt one's sense of location of that limb. Possible experiences include suddenly feeling that feet or legs are missing from one's mental self-image; needing to look down at one's limbs to be sure they are still there; and falling down while walking, especially when attention is focused upon something other than the act of walking.

<div class="mw-heading mw-heading3"><h3 id="Diagnosis">Diagnosis</h3></div>

Impaired proprioception may be diagnosed through a series of tests, each focusing on a different functional aspect of proprioception.

The <a href="Romberg's_test" title="Romberg's test">Romberg's test</a> is often used to assess balance. The subject must stand with feet together and eyes closed without support for 30 seconds. If the subject loses balance and falls, it is an indicator for impaired proprioception.

For evaluating proprioception's contribution to motor control, a common protocol is joint position matching.<a href="#cite_note-75">[74]</a> The patient is blindfolded while a joint is moved to a specific angle for a given period of time and then returned to neutral. The subject is then asked to move the joint back to the specified angle. Recent investigations have shown that hand dominance, participant age, active versus passive matching, and presentation time of the angle can all affect performance on joint position matching tasks.<a href="#cite_note-76">[75]</a><a href="#cite_note-77">[76]</a>

For passive sensing of joint angles, recent studies have found that experiments to probe psychophysical thresholds produce more precise estimates of proprioceptive discrimination than the joint position matching task.<a href="#cite_note-78">[77]</a> In these experiments, the subject holds on to an object (such as an armrest) that moves and stops at different positions. The subject must discriminate whether one position is closer to the body than another. From the subject's choices, the tester may determine the subject's discrimination thresholds.

Proprioception is tested by American <a href="Police_officer" title="Police officer">police officers</a> using the <a href="Field_sobriety_testing" title="Field sobriety testing">field sobriety testing</a> to check for <a href="Health_effects_of_alcohol" title="Health effects of alcohol">alcohol intoxication</a>. The subject is required to touch his or her nose with eyes closed; people with normal proprioception may make an error of no more than 20 mm (0.79 in), while people with impaired proprioception (a symptom of moderate to severe alcohol intoxication) fail this test due to difficulty locating their limbs in space relative to their noses.

<div class="mw-heading mw-heading2"><h2 id="Training">Training</h2></div>

Proprioception is what allows someone to learn to walk in complete darkness without losing balance. During the learning of any new skill, sport, or art, it is usually necessary to become familiar with some proprioceptive tasks specific to that activity. Without the appropriate integration of proprioceptive input, an artist would not be able to brush paint onto a canvas without looking at the hand as it moved the brush over the canvas; it would be impossible to drive an automobile because a motorist would not be able to steer or use the pedals while looking at the road ahead; a person could not <a href="Touch_type" class="mw-redirect" title="Touch type">touch type</a> or perform ballet; and people would not even be able to walk without watching where they put their feet.

<a href="Oliver_Sacks" title="Oliver Sacks">Oliver Sacks</a> reported the case of a young woman who lost her proprioception due to a viral infection of her <a href="Spinal_cord" title="Spinal cord">spinal cord</a>.<a href="#cite_note-79">[78]</a> At first she could not move properly at all or even control her tone of voice (as voice modulation is primarily proprioceptive). Later she relearned by using her sight (watching her feet) and <a href="Inner_ear" title="Inner ear">inner ear</a> only for movement while using hearing to judge voice modulation. She eventually acquired a stiff and slow movement and nearly normal speech, which is believed to be the best possible in the absence of this sense. She could not judge effort involved in picking up objects and would grip them painfully to be sure she did not drop them.

The proprioceptive sense can be sharpened through study of many disciplines.Standing on a <a href="Balance_board" title="Balance board">wobble board or balance board</a> is often used to retrain or increase proprioceptive abilities, particularly as <a href="Physical_therapy" title="Physical therapy">physical therapy</a> for ankle or knee injuries. <a href="Slacklining" title="Slacklining">Slacklining</a> is another method to increase proprioception.

Standing on one leg (stork standing) and various other body-position challenges are also used in such disciplines as <a href="Yoga" title="Yoga">yoga</a>, <a href="Wing_Chun" title="Wing Chun">Wing Chun</a> and <a href="Tai_chi" title="Tai chi">tai chi</a>.<a href="#cite_note-80">[79]</a> The vestibular system of the inner ear, vision and proprioception are the main three requirements for balance.<a href="#cite_note-81">[80]</a> Moreover, there are specific devices designed for proprioception training, such as the <a href="Exercise_ball" title="Exercise ball">exercise ball</a>, which works on balancing the abdominal and back muscles.

<div class="mw-heading mw-heading2"><h2 id="History_of_study">History of study</h2></div>

In 1557, the position-movement sensation was described by <a href="Julius_Caesar_Scaliger" title="Julius Caesar Scaliger">Julius Caesar Scaliger</a> as a "sense of locomotion".<a href="#cite_note-82">[81]</a>

In 1826, <a href="Charles_Bell" title="Charles Bell">Charles Bell</a> expounded the idea of a "muscle sense",<a href="#cite_note-Singh1991-83">[82]</a> which is credited as one of the first descriptions of physiologic feedback mechanisms.<a href="#cite_note-Dickinson1976-84">[83]</a> Bell's idea was that commands are carried from the brain to the muscles, and that reports on the muscle's condition would be sent in the reverse direction.

In 1847, the London neurologist <a href="Robert_Bentley_Todd" title="Robert Bentley Todd">Robert Todd</a> highlighted important differences in the anterolateral and <a href="Posterior_column" class="mw-redirect" title="Posterior column">posterior columns</a> of the spinal cord, and suggested that the latter were involved in the coordination of movement and balance.<a href="#cite_note-85">[84]</a>

At around the same time, <a href="Moritz_Heinrich_Romberg" title="Moritz Heinrich Romberg">Moritz Heinrich Romberg</a>, a Berlin neurologist, was describing unsteadiness made worse by eye closure or darkness, now known as the eponymous <a href="Romberg's_test" title="Romberg's test">Romberg's sign</a>, once synonymous with <a href="Tabes_dorsalis" title="Tabes dorsalis">tabes dorsalis</a>, that became recognised as common to all proprioceptive disorders of the legs.

In 1880, <a href="Henry_Charlton_Bastian" title="Henry Charlton Bastian">Henry Charlton Bastian</a> suggested "kinaesthesia" instead of "muscle sense" on the basis that some of the <a href="Afferent_nerve_fiber" title="Afferent nerve fiber">afferent</a> information (back to the brain) comes from other structures, including tendons, joints, and skin.<a href="#cite_note-Foster2010-86">[85]</a>

In 1889, <a href="Alfred_Goldscheider" title="Alfred Goldscheider">Alfred Goldscheider</a> suggested a classification of kinaesthesia into three types: muscle, tendon, and articular sensitivity.<a href="#cite_note-BrookhartMountcastle1984-87">[86]</a>

In 1906, the term proprio-ception (and also <a href="Interoception" title="Interoception">intero-ception</a> and extero-ception) is attested in a publication by <a href="Charles_Scott_Sherrington" title="Charles Scott Sherrington">Charles Scott Sherrington</a> involving <a href="Receptor_(biochemistry)" title="Receptor (biochemistry)">receptors</a>.<a href="#cite_note-88">[87]</a> He explains the terminology as follows:<a href="#cite_note-89">[88]</a>

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The main fields of distribution of the <a href="Receptor_(biochemistry)" title="Receptor (biochemistry)">receptor</a> organs fundamentally distinguishable seem, therefore, to be two, namely, a surface field constituted by the surface layer of the organism, and a deep field constituted by the tissues of the organism beneath the surface sheet.

[...]

the stimulations occurring in [the] deep field is that the stimuli are traceable to actions of the organism itself, and are so in much greater measure than are the stimulations of the surface field of the organism. Since in the deep field the stimuli to the receptors are delivered by the organism itself,<a href="#cite_note-proprio-90">[b]</a> the deep receptors may be termed proprio-ceptors, and the deep field a field of proprio-ception.

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Today, the "exteroceptors" are the organs that provide information originating outside the body, such as the eyes, ears, mouth, and skin. The <a href="Interoceptor" class="mw-redirect" title="Interoceptor">interoceptors</a> provide information about the internal organs, and the "proprioceptors" provide information about movement derived from muscular, tendon, and articular sources. Using Sherrington's system, physiologists and anatomists search for specialised nerve endings that transmit mechanical data on joint capsule, tendon and muscle tension (such as <a href="Golgi_tendon_organ" title="Golgi tendon organ">Golgi tendon organs</a> and <a href="Muscle_spindles" class="mw-redirect" title="Muscle spindles">muscle spindles</a>), which play a large role in proprioception.

Primary endings of muscle spindles "respond to the size of a muscle length change and its speed" and "contribute both to the sense of limb position and movement".<a href="#cite_note-Proske-91">[89]</a> Secondary endings of muscle spindles detect changes in muscle length, and thus supply information regarding only the sense of position.<a href="#cite_note-Proske-91">[89]</a> Essentially, muscle spindles are stretch receptors.<a href="#cite_note-92">[90]</a> It has been accepted that cutaneous receptors also contribute directly to proprioception by providing "accurate perceptual information about joint position and movement", and this knowledge is combined with information from the muscle spindles.<a href="#cite_note-93">[91]</a>

<div class="mw-heading mw-heading3"><h3 id="Etymology">Etymology</h3></div>

Proprioception is from <a href="Latin" title="Latin">Latin</a> <a href="https://en.wiktionary.org/wiki/proprius#Latin" class="extiw external" title="wikt:proprius">proprius</a>, meaning "one's own", "individual", and <a href="https://en.wiktionary.org/wiki/capio#Latin" class="extiw external" title="wikt:capio">capio</a>, capere, to take or grasp. Thus to grasp one's own position in space, including the position of the limbs in relation to each other and the body as a whole.

The word kinesthesia or kinæsthesia (kinesthetic sense) refers to movement sense, but has been used inconsistently to refer either to proprioception alone or to the brain's integration of proprioceptive and vestibular inputs. Kinesthesia is a modern medical term composed of elements from Greek; kinein "to set in motion; to move" (from PIE root *keie- "to set in motion") + aisthesis "perception, feeling" (from PIE root *au- "to perceive").

<div class="mw-heading mw-heading2"><h2 id="Plants_and_bacteria">Plants and bacteria</h2></div>

Although they lack neurons, systems responding to stimuli (analogous to the sensory system in animals with a nervous system, which includes the proprioception) have also been described in some plants (<a href="Flowering_plant" title="Flowering plant">angiosperms</a>).<a href="#cite_note-:0-94">[92]</a><a href="#cite_note-:1-95">[93]</a> <a href="Terrestrial_plant" title="Terrestrial plant">Terrestrial plants</a> control the orientation of their <a href="Primary_growth" title="Primary growth">primary growth</a> through the sensing of several vectorial <a href="Stimulus_(physiology)" title="Stimulus (physiology)">stimuli</a> such as the light gradient or the <a href="Gravitational_acceleration" title="Gravitational acceleration">gravitational acceleration</a>. This control has been called <a href="Tropism" title="Tropism">tropism</a>. A quantitative study of <a href="Shoot_(botany)" title="Shoot (botany)">shoot</a> <a href="Gravitropism" title="Gravitropism">gravitropism</a> demonstrated that, when a plant is tilted, it cannot recover a steady erected posture under the sole driving of the sensing of its angular deflection versus gravity. An additional control through the continuous sensing of its <a href="Curvature" title="Curvature">curvature</a> by the organ and the subsequent driving an active straightening process are required.<a href="#cite_note-:0-94">[92]</a><a href="#cite_note-:1-95">[93]</a><a href="#cite_note-96">[94]</a> Being a sensing by the plant of the relative configuration of its parts, it has been called proprioception. This dual sensing and control by gravisensing and proprioception has been formalized into a unifying <a href="Mathematical_model" title="Mathematical model">mathematical model</a> simulating the complete driving of the gravitropic movement. This model has been validated on 11 species sampling the <a href="Angiosperm_Phylogeny_Group_III_system" class="mw-redirect" title="Angiosperm Phylogeny Group III system">phylogeny of land angiosperms</a>, and on organs of very contrasted sizes, ranging from the small germination of <a href="Wheat" title="Wheat">wheat</a> (<a href="Coleoptile" title="Coleoptile">coleoptile</a>) to the trunk of <a href="Populus" title="Populus">poplar trees</a>.<a href="#cite_note-:0-94">[92]</a><a href="#cite_note-:1-95">[93]</a>

Further studies have shown that the cellular mechanism of proprioception in plants involves <a href="Myosin" title="Myosin">myosin</a> and <a href="Actin" title="Actin">actin</a>, and seems to occur in specialized cells.<a href="#cite_note-97">[95]</a> Proprioception was then found to be involved in other tropisms and to be central also to the control of <a href="Nutation_(botany)" title="Nutation (botany)">nutation</a>.<a href="#cite_note-98">[96]</a>

The discovery of proprioception in plants has generated an interest in the popular science and generalist media.<a href="#cite_note-99">[97]</a><a href="#cite_note-:2-100">[98]</a> This is because this discovery questions a long-lasting a priori that we have on plants. In some cases this has led to a shift between proprioception and <a href="Self-awareness" title="Self-awareness">self-awareness</a> or <a href="Self-consciousness" title="Self-consciousness">self-consciousness</a>. There is no scientific ground for such a semantic shift. Indeed, even in animals, proprioception can be unconscious; so, it is thought to be in plants.<a href="#cite_note-:1-95">[93]</a><a href="#cite_note-:2-100">[98]</a>

Recent studies suggest that bacteria have control systems that may resemble proprioception.<a href="#cite_note-101">[99]</a>

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<ul><li><a href="Balance_disorder" title="Balance disorder">Balance disorder</a> – Physiological disturbance of perception</li>

<li><a href="Body_schema" title="Body schema">Body schema</a> – Postural model that keeps track of limb position</li>

<li><a href="Broken_escalator_phenomenon" title="Broken escalator phenomenon">Broken escalator phenomenon</a> – Illusion when stepping onto a broken escalator</li>

<li><a href="Dizziness" title="Dizziness">Dizziness</a> – Neurological condition causing impairment in spatial perception and stability</li>

<li><a href="Equilibrioception" class="mw-redirect" title="Equilibrioception">Equilibrioception</a> – Physiological sense regarding posturePages displaying short descriptions of redirect targets</li>

<li><a href="Hand%E2%80%93eye_coordination" class="mw-redirect" title="Hand–eye coordination">Hand–eye coordination</a> – Coordination between the eyes and handPages displaying short descriptions of redirect targets</li>

<li><a href="Ideomotor_phenomenon" title="Ideomotor phenomenon">Ideomotor phenomenon</a> – Concept in hypnosis and psychological research</li>

<li><a href="Illusions_of_self-motion" title="Illusions of self-motion">Illusions of self-motion</a> – Misperception of one's location or movement</li>

<li><a href="Instinctive_aiming" class="mw-redirect" title="Instinctive aiming">Instinctive aiming</a> – Shooting method where the weapon's sights are not used or relied onPages displaying short descriptions of redirect targets</li>

<li><a href="Kinaesthetics" title="Kinaesthetics">Kinaesthetics</a> – Study of body motion, and preception of motion</li>

<li><a href="Kinesthetic_learning" title="Kinesthetic learning">Kinesthetic learning</a> – Learning by physical activities</li>

<li><a href="List_of_distinct_cell_types_in_the_adult_human_body" class="mw-redirect" title="List of distinct cell types in the adult human body">List of distinct cell types in the adult human body</a></li>

<li><a href="Motion_sickness" title="Motion sickness">Motion sickness</a> – Nausea caused by motion or perceived motion</li>

<li><a href="Motor_control" title="Motor control">Motor control</a> – Regulation of movement within organisms possessing a nervous system</li>

<li><a href="Multisensory_integration" title="Multisensory integration">Multisensory integration</a> – Study of senses and nervous system</li>

<li><a href="Seasickness" class="mw-redirect" title="Seasickness">Seasickness</a> – Motion sickness occurring at sea</li>

<li><a href="Spatial_disorientation" title="Spatial disorientation">Spatial disorientation</a> – Inability of a person to correctly determine their body position in space</li>

<li><a href="Theory_of_multiple_intelligences" title="Theory of multiple intelligences">Theory of multiple intelligences</a> – Educational model of human intelligence</li>

<li><a href="Vertigo" title="Vertigo">Vertigo</a> – Type of dizziness where a person has the sensation of moving or surrounding objects moving</li></ul>

</div>

<div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2></div>

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<li id="cite_note-piezoChannels-23"><a href="#cite_ref-piezoChannels_23-0">^</a> The Piezo channel receptors play key roles in the perception of pressure, touch, and proprioception (Piezo2 receptor).<a href="#cite_note-4Oct2021-22">[22]</a>

</li>

<li id="cite_note-proprio-90"><a href="#cite_ref-proprio_90-0">^</a> in <a href="Latin" title="Latin">Latin</a>: <a href="https://en.wiktionary.org/wiki/proprius#Latin" class="extiw external" title="wikt:proprius">propriō</a>

</li>

</ol></div></div>

<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>

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</style><cite id="CITEREFTuthillAzim2018" class="citation journal cs1">Tuthill JC, Azim E (1 March 2018). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cub.2018.01.064">"Proprioception"</a>. <a href="Current_Biology" title="Current Biology">Current Biology</a>. 28 (5): R194 – R203. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2018CBio...28.R194T">2018CBio...28.R194T</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cub.2018.01.064">10.1016/j.cub.2018.01.064</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/29510103">29510103</a>.</cite>

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<li id="cite_note-101"><a href="#cite_ref-101">^</a> <cite id="CITEREFAntaniGuptaLeeRhee2021" class="citation journal cs1">Antani JD, Gupta R, Lee AH, Rhee KY, Manson MD, Lele PP (September 2021). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8440544">"Mechanosensitive recruitment of stator units promotes binding of the response regulator CheY-P to the flagellar motor"</a>. Nature Communications. 12 (1) 5442. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2021NatCo..12.5442A">2021NatCo..12.5442A</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fs41467-021-25774-2">10.1038/s41467-021-25774-2</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8440544">8440544</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/34521846">34521846</a>.</cite>

</li>

</ol></div></div>

<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>

<ul><li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=Proprioception">Proprioception</a> at the U.S. National Library of Medicine <a href="Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li></ul>

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</style><div id="Sensory_receptors112" style="font-size:114%;margin:0 4em"><a href="Sensory_neuron" title="Sensory neuron">Sensory receptors</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Somatosensory_system" title="Somatosensory system">Touch</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Mechanoreceptor" title="Mechanoreceptor">Mechanoreceptor</a></li>

<li>Vibration

<ul><li><a href="Lamellar_corpuscle" class="mw-redirect" title="Lamellar corpuscle">Lamellar corpuscle</a></li></ul></li>

<li>Light touch

<ul><li><a href="Tactile_corpuscle" title="Tactile corpuscle">Tactile corpuscle</a></li></ul></li>

<li>Pressure

<ul><li><a href="Merkel_nerve_ending" title="Merkel nerve ending">Merkel nerve ending</a></li></ul></li>

<li>Stretch

<ul><li><a href="Bulbous_corpuscle" title="Bulbous corpuscle">Bulbous corpuscle</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Pain" title="Pain">Pain</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Free_nerve_ending" title="Free nerve ending">Free nerve ending</a></li>

<li><a href="Nociceptor" title="Nociceptor">Nociceptors</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Temperature" title="Temperature">Temperature</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Thermoreceptor" title="Thermoreceptor">Thermoreceptors</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Golgi_tendon_organ" title="Golgi tendon organ">Golgi organ</a></li>

<li><a href="Muscle_spindle" title="Muscle spindle">Muscle spindle</a>

<ul><li><a href="Intrafusal_muscle_fiber" title="Intrafusal muscle fiber">Intrafusal muscle fiber</a></li>

<li><a href="Nuclear_chain_fiber" title="Nuclear chain fiber">Nuclear chain fiber</a></li>

<li><a href="Nuclear_bag_fiber" title="Nuclear bag fiber">Nuclear bag fiber</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Hair_cell" title="Hair cell">Hair cells</a></li>

<li><a href="Baroreceptor" title="Baroreceptor">Baroreceptor</a></li></ul>

</div></td></tr></tbody></table></div>

<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Sensation_and_perception287" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Sensation_and_perception287" style="font-size:114%;margin:0 4em"><a href="Sense" title="Sense">Sensation</a> and <a href="Perception" title="Perception">perception</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Cognitive_process" class="mw-redirect" title="Cognitive process">Processes and concepts</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Sense" title="Sense">Sensation</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Stimulus_(physiology)" title="Stimulus (physiology)">Stimulus</a></li>

<li><a href="Sensory_neuron" title="Sensory neuron">Sensory receptor</a></li>

<li><a href="Transduction_(physiology)" title="Transduction (physiology)">Transduction (physiology)</a></li>

<li><a href="Sensory_processing" title="Sensory processing">Sensory processing</a></li>

<li><a href="Active_sensory_system" class="mw-redirect" title="Active sensory system">Active sensory system</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Perception" title="Perception">Perception</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Multimodal_integration" class="mw-redirect" title="Multimodal integration">Multimodal integration</a></li>

<li><a href="Awareness" title="Awareness">Awareness</a></li>

<li><a href="Consciousness" title="Consciousness">Consciousness</a></li>

<li><a href="Cognition" title="Cognition">Cognition</a></li>

<li><a href="Feeling" title="Feeling">Feeling</a></li>

<li><a href="Motion_perception" title="Motion perception">Motion perception</a></li>

<li><a href="Qualia" title="Qualia">Qualia</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Human" title="Human">Human</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Exteroception" class="mw-redirect" title="Exteroception">External</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Sensory_organ" class="mw-redirect" title="Sensory organ">Sensory organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<li><a href="Inner_ear" title="Inner ear">Inner ear</a></li>

<li><a href="Human_mouth" title="Human mouth">Mouth</a></li>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Sensory_system" class="mw-redirect" title="Sensory system">Sensory systems</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Visual_system" title="Visual system">Visual system (sense of vision)</a></li>

<li><a href="Auditory_system" title="Auditory system">Auditory system (sense of hearing)</a></li>

<li><a href="Vestibular_system" title="Vestibular system">Vestibular system (sense of balance)</a></li>

<li><a href="Olfactory_system" title="Olfactory system">Olfactory system (sense of smell)</a></li>

<li><a href="Gustatory_system" class="mw-redirect" title="Gustatory system">Gustatory system (sense of taste)</a></li>

<li><a href="Somatosensory_system" title="Somatosensory system">Somatosensory system (sense of touch)</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Sensory <a href="Cranial_nerves" title="Cranial nerves">cranial</a> and <a href="Spinal_nerves" class="mw-redirect" title="Spinal nerves">spinal nerves</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Optic_nerve" title="Optic nerve">Optic (II)</a></li>

<li><a href="Vestibulocochlear_nerve" title="Vestibulocochlear nerve">Vestibulocochlear (VIII)</a></li>

<li><a href="Olfactory_nerve" title="Olfactory nerve">Olfactory (I)</a></li>

<li><a href="Facial_nerve" title="Facial nerve">Facial (VII)</a></li>

<li><a href="Glossopharyngeal_nerve" title="Glossopharyngeal nerve">Glossopharyngeal (IX)</a></li>

<li><a href="Trigeminal_nerve" title="Trigeminal nerve">Trigeminal (V)</a></li>

<li><a href="Spinal_nerve" title="Spinal nerve">Spinal</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Sensory_cortex" title="Sensory cortex">Cerebral cortices</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Visual_cortex" title="Visual cortex">Visual cortex</a></li>

<li><a href="Auditory_cortex" title="Auditory cortex">Auditory cortex</a></li>

<li><a href="Vestibular_cortex" title="Vestibular cortex">Vestibular cortex</a></li>

<li><a href="Olfactory_cortex" class="mw-redirect" title="Olfactory cortex">Olfactory cortex</a></li>

<li><a href="Gustatory_cortex" title="Gustatory cortex">Gustatory cortex</a></li>

<li><a href="Somatosensory_cortex" class="mw-redirect" title="Somatosensory cortex">Somatosensory cortex</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Perceptions</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Visual_perception" title="Visual perception">Visual perception (vision)</a>

<ul><li><a href="Color" title="Color">Color</a></li></ul></li>

<li><a href="Auditory_perception" class="mw-redirect" title="Auditory perception">Auditory perception (hearing)</a></li>

<li><a href="Equilibrioception" class="mw-redirect" title="Equilibrioception">Equilibrioception (balance)</a></li>

<li><a href="Sense_of_smell" title="Sense of smell">Olfaction (smell)</a></li>

<li><a href="Taste" title="Taste">Gustation (taste or flavor)</a></li>

<li><a href="Touch" class="mw-redirect" title="Touch">Touch</a>

<ul><li><a href="Mechanoreception" class="mw-redirect" title="Mechanoreception">mechanoreception</a></li>

<li><a href="Nociception" title="Nociception">nociception (pain)</a></li>

<li><a href="Thermoception" title="Thermoception">thermoception</a></li></ul></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Interoception" title="Interoception">Internal</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul>

<li><a href="Hunger_(physiology)" title="Hunger (physiology)">Hunger</a></li>

<li><a href="Thirst" title="Thirst">Thirst</a></li>

<li><a href="Suffocation" class="mw-redirect" title="Suffocation">Suffocation</a></li>

<li><a href="Nausea" title="Nausea">Nausea</a></li>

<li><a href="Visceral_pain" title="Visceral pain">Visceral pain</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Nonhuman" class="mw-redirect" title="Nonhuman">Nonhuman</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Sense" title="Sense">Animal</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Electroreception" class="mw-redirect" title="Electroreception">Electroreception</a></li>

<li><a href="Magnetoreception" title="Magnetoreception">Magnetoreception</a></li>

<li><a href="Animal_echolocation" title="Animal echolocation">Echolocation</a></li>

<li><a href="Infrared_sensing_in_vampire_bats" title="Infrared sensing in vampire bats">Infrared sensing in vampire bats</a></li>

<li><a href="Infrared_sensing_in_snakes" title="Infrared sensing in snakes">Infrared sensing in snakes</a></li>

<li><a href="Surface_wave_detection" class="mw-redirect" title="Surface wave detection">Surface wave detection</a></li>

<li><a href="Frog_hearing_and_communication" title="Frog hearing and communication">Frog hearing</a></li>

<li><a href="Vision_in_toads" title="Vision in toads">Toad vision</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Plant_perception_(physiology)" title="Plant perception (physiology)">Plant</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Photomorphogenesis" title="Photomorphogenesis">Photomorphogenesis</a></li>

<li><a href="Gravitropism" title="Gravitropism">Gravitropism</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Machine_perception" title="Machine perception">Artificial</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Robotic_sensing" title="Robotic sensing">Robotic sensing</a></li>

<li><a href="Computer_vision" title="Computer vision">Computer vision</a></li>

<li><a href="Machine_hearing" class="mw-redirect" title="Machine hearing">Machine hearing</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types of <a href="Sensory_receptor" class="mw-redirect" title="Sensory receptor">sensory receptors</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Mechanoreceptor" title="Mechanoreceptor">Mechanoreceptor</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Baroreceptor" title="Baroreceptor">Baroreceptor</a></li>

<li><a href="Mechanotransduction" title="Mechanotransduction">Mechanotransduction</a></li>

<li><a href="Lamellar_corpuscle" class="mw-redirect" title="Lamellar corpuscle">Lamellar corpuscle</a></li>

<li><a href="Tactile_corpuscle" title="Tactile corpuscle">Tactile corpuscle</a></li>

<li><a href="Merkel_nerve_ending" title="Merkel nerve ending">Merkel nerve ending</a></li>

<li><a href="Bulbous_corpuscle" title="Bulbous corpuscle">Bulbous corpuscle</a></li>

<li><a href="Campaniform_sensilla" title="Campaniform sensilla">Campaniform sensilla</a></li>

<li><a href="Slit_sensilla" title="Slit sensilla">Slit sensilla</a></li>

<li><a href="Stretch_receptor" title="Stretch receptor">Stretch receptor</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Photoreceptor</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Photoreceptor_cell" title="Photoreceptor cell">Photoreceptor cell</a></li>

<li><a href="IpRGC" class="mw-redirect" title="IpRGC">ipRGC</a></li>

<li><a href="Photopigment" title="Photopigment">Photopigment</a></li>

<li><a href="Aureochromes" title="Aureochromes">Aureochrome</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Chemoreceptor" title="Chemoreceptor">Chemoreceptor</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Taste_receptor" title="Taste receptor">Taste receptor</a></li>

<li><a href="Olfactory_receptor" title="Olfactory receptor">Olfactory receptor</a></li>

<li><a href="Osmoreceptor" title="Osmoreceptor">Osmoreceptor</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Thermoreceptor" title="Thermoreceptor">Thermoreceptor</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Cilium#Sensing_the_extracellular_environment" title="Cilium">Cilium</a></li>

<li><a href="TRP_channels" class="mw-redirect" title="TRP channels">TRP channels</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Nociceptor" title="Nociceptor">Nociceptor</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Nociceptin_receptor#Pain_circuitry" title="Nociceptin receptor">Nociceptin receptor</a></li>

<li><a href="Juxtacapillary_receptors" title="Juxtacapillary receptors">Juxtacapillary receptor</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Disorders</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Vision_disorder" title="Vision disorder">Visual</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Visual_impairment" title="Visual impairment">Visual impairment</a></li>

<li><a href="Alice_in_Wonderland_syndrome" title="Alice in Wonderland syndrome">Alice in Wonderland syndrome</a></li>

<li><a href="Amaurosis" title="Amaurosis">Amaurosis</a></li>

<li><a href="Anopsia" title="Anopsia">Anopsia</a></li>

<li><a href="Color_blindness" title="Color blindness">Color blindness</a></li>

<li><a href="Diplopia" title="Diplopia">Diplopia</a></li>

<li><a href="Hemeralopia" title="Hemeralopia">Hemeralopia</a> and <a href="Nyctalopia" title="Nyctalopia">Nyctalopia</a></li>

<li><a href="Optic_neuropathy" title="Optic neuropathy">Optic neuropathy</a></li>

<li><a href="Oscillopsia" title="Oscillopsia">Oscillopsia</a></li>

<li><a href="Palinopsia" title="Palinopsia">Palinopsia</a></li>

<li><a href="Papilledema" title="Papilledema">Papilledema</a></li>

<li><a href="Photophobia" title="Photophobia">Photophobia</a></li>

<li><a href="Photopsia" title="Photopsia">Photopsia</a></li>

<li><a href="Cerebral_polyopia" title="Cerebral polyopia">Polyopia</a></li>

<li><a href="Scotoma" title="Scotoma">Scotoma</a></li>

<li><a href="Stereoblindness" title="Stereoblindness">Stereoblindness</a></li>

<li><a href="Visual_snow" class="mw-redirect" title="Visual snow">Visual snow</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Auditory_processing_disorder" title="Auditory processing disorder">Auditory</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Amblyaudia" title="Amblyaudia">Amblyaudia</a></li>

<li><a href="Auditory_agnosia" title="Auditory agnosia">Auditory agnosia</a></li>

<li><a href="Auditory_hallucination" title="Auditory hallucination">Auditory hallucination</a></li>

<li><a href="Auditory_verbal_agnosia" title="Auditory verbal agnosia">Auditory verbal agnosia</a></li>

<li><a href="Cortical_deafness" title="Cortical deafness">Cortical deafness</a></li>

<li><a href="Hearing_loss" title="Hearing loss">Hearing loss</a></li>

<li><a href="Microwave_auditory_effect" title="Microwave auditory effect">Microwave auditory effect</a></li>

<li><a href="Music-specific_disorders" title="Music-specific disorders">Music-specific disorders</a></li>

<li><a href="Palinopsia" title="Palinopsia">Palinopsia</a></li>

<li><a href="Spatial_hearing_loss" title="Spatial hearing loss">Spatial hearing loss</a></li>

<li><a href="Tinnitus" title="Tinnitus">Tinnitus</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Balance_disorder" title="Balance disorder">Vestibular</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Vertigo" title="Vertigo">Vertigo</a></li>

<li><a href="Labyrinthine_fistula" title="Labyrinthine fistula">Labyrinthine fistula</a></li>

<li><a href="Labyrinthitis" title="Labyrinthitis">Labyrinthitis</a></li>

<li><a href="M%C3%A9ni%C3%A8re's_disease" title="Ménière's disease">Ménière's disease</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Olfaction" class="mw-redirect" title="Olfaction">Olfactory</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Anosmia" title="Anosmia">Anosmia</a></li>

<li><a href="Dysosmia" title="Dysosmia">Dysosmia</a></li>

<li><a href="Hyperosmia" title="Hyperosmia">Hyperosmia</a></li>

<li><a href="Hyposmia" title="Hyposmia">Hyposmia</a></li>

<li><a href="Olfactory_reference_syndrome" title="Olfactory reference syndrome">Olfactory reference syndrome</a></li>

<li><a href="Parosmia" title="Parosmia">Parosmia</a></li>

<li><a href="Phantosmia" title="Phantosmia">Phantosmia</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Taste_disorder" class="mw-redirect" title="Taste disorder">Gustatory</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Ageusia" title="Ageusia">Ageusia</a></li>

<li><a href="Hypergeusia" title="Hypergeusia">Hypergeusia</a></li>

<li><a href="Hypogeusia" title="Hypogeusia">Hypogeusia</a></li>

<li><a href="Parageusia" class="mw-redirect" title="Parageusia">Parageusia</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Somatosensory_disorder" title="Somatosensory disorder">Tactile</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Astereognosis" title="Astereognosis">Astereognosis</a></li>

<li><a href="Charcot%E2%80%93Marie%E2%80%93Tooth_disease" title="Charcot–Marie–Tooth disease">CMT disease</a></li>

<li><a href="Formication" title="Formication">Formication</a></li>

<li><a href="Hyperesthesia" title="Hyperesthesia">Hyperesthesia</a></li>

<li><a href="Hypoesthesia" title="Hypoesthesia">Hypoesthesia</a></li>

<li><a href="Paresthesia" title="Paresthesia">Paresthesia</a></li>

<li><a href="Tactile_hallucination" title="Tactile hallucination">Tactile hallucination</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Nociception (pain)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Hyperalgesia" title="Hyperalgesia">Hyperalgesia</a></li>

<li><a href="Hypoalgesia" title="Hypoalgesia">Hypoalgesia</a></li>

<li><a href="Pain_dissociation" class="mw-redirect" title="Pain dissociation">Pain dissociation</a></li>

<li><a href="Phantom_pain" title="Phantom pain">Phantom pain</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Proprioception</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Asomatognosia" title="Asomatognosia">Asomatognosia</a></li>

<li><a href="Phantom_limb" title="Phantom limb">Phantom limb syndrome</a></li>

<li><a href="Somatoparaphrenia" title="Somatoparaphrenia">Somatoparaphrenia</a></li>

<li><a href="Supernumerary_phantom_limb" title="Supernumerary phantom limb">Supernumerary phantom limb</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Multimodal</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<li><a href="Agnosia" title="Agnosia">Agnosia</a></li>

<li><a href="Allochiria" title="Allochiria">Allochiria</a></li>

<li><a href="Derealization" title="Derealization">Derealization</a></li>

<li><a href="Hallucination" title="Hallucination">Hallucination</a></li>

<li><a href="Sensory_processing_disorder" title="Sensory processing disorder">Sensory processing disorder</a></li>

<li><a href="Synesthesia" title="Synesthesia">Synesthesia</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Biases and errors</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Illusion" title="Illusion">Illusion</a> · <a href="Pareidolia" title="Pareidolia">Pareidolia</a></li></ul>

</div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> Category</div></td></tr></tbody></table></div></div><div><div class="zim-footer">

This article is issued from <a class="external text" title="Last edited on 2025-07-18" href="https://en.wikipedia.org/wiki/?title=Proprioception&oldid=1301180935">Wikipedia</a>. The text is available under <a class="external text" href="https://creativecommons.org/licenses/by-sa/4.0/deed.en">Creative Commons Attribution-Share Alike 4.0</a> unless otherwise noted. Additional terms may apply for the media files.

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