Proprioception ( ) is the

sense A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of Stimulus (physiology), stimuli. Although, in some cultures, five human senses were traditio ...

of self-movement, force, and body position. Proprioception is mediated by proprioceptors, a type of

sensory receptor Sensory neurons, also known as afferent neurons, are neurons in the nervous system, that convert a specific type of stimulus, via their receptors, into action potentials or graded receptor potentials. This process is called sensory transduc ...

, located within

muscle Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...

tendon A tendon or sinew is a tough band of fibrous connective tissue, dense fibrous connective tissue that connects skeletal muscle, muscle to bone. It sends the mechanical forces of muscle contraction to the skeletal system, while withstanding tensi ...

s, and

joint A joint or articulation (or articular surface) is the connection made between bones, ossicles, or other hard structures in the body which link an animal's skeletal system into a functional whole.Saladin, Ken. Anatomy & Physiology. 7th ed. McGraw- ...

s. 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

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

, where they are integrated with information from other

sensory systems The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons (including the sensory receptor cells), neural pathways, and parts of the brain involved i ...

, such as the visual system and the

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

, 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.

System overview

In vertebrates, limb movement and velocity (muscle length and the rate of change) are encoded by one group of sensory neurons (

type Ia sensory fiber A type Ia sensory fiber, or a primary afferent fiber, is a type of afferent nerve fiber. It is the sensory fiber of a stretch receptor called the muscle spindle found in muscles, which constantly monitors the rate at which a muscle stretch chan ...

) and another type encode static muscle length ( group II neurons). These two types of sensory neurons compose

muscle spindles Muscle spindles are stretch receptors within the body of a skeletal muscle that primarily detect changes in the length of the muscle. They convey length information to the central nervous system via afferent nerve fibers. This information can be ...

. There is a similar division of encoding in invertebrates; different subgroups of neurons of the

chordotonal organ Chordotonal organs are stretch receptor organs found only in insects and crustaceans. They are located at most joints and are made up of clusters of scolopidia that either directly or indirectly connect two joints and sense their movements relativ ...

encode limb position and velocity. To determine the load on a limb, vertebrates use sensory neurons in the Golgi tendon organs: 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 campaniform sensilla. These proprioceptors are active when a limb experiences resistance. A third role for proprioceptors is to determine when a joint is at a specific position. In vertebrates, this is accomplished by

Ruffini ending The bulbous corpuscle, Ruffini ending or Ruffini corpuscle is a slowly adapting mechanoreceptor A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are locat ...

s and

Pacinian corpuscles The Pacinian corpuscle (also lamellar corpuscle, or Vater–Pacini corpuscle) is a low-threshold mechanoreceptor responsive to vibration or pressure, found in the skin and other internal organs. In the skin it is one of the four main types of cuta ...

. These proprioceptors are activated when the joint is at a threshold position, usually at the extremes of joint position. Invertebrates use

hair plate Hair plates are a type of Proprioception, proprioceptor found in the folds of insect joints. They consist of a cluster of hairs, in which each hair is innervated by a single Mechanosensation, mechanosensory neuron. Functionally, hair plates operate ...

s 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.

Reflexes

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

mechanosensation Mechanosensation is the transduction of mechanical stimuli into neural signals. Mechanosensation provides the basis for the senses of light touch, hearing, proprioception, and pain. Mechanoreceptors found in the skin, called cutaneous mechanorecept ...

circuits are important for flexibly maintaining posture and balance, especially during locomotion. For example, consider the

stretch reflex The stretch reflex (myotatic reflex), or more accurately ''muscle stretch reflex'', is a muscle contraction in response to stretching a muscle. The function of the reflex is generally thought to be maintaining the muscle at a constant length but ...

, in which stretch across a muscle is detected by a sensory receptor (e.g.,

muscle spindle Muscle spindles are stretch receptors within the body of a skeletal muscle that primarily detect changes in the length of the muscle. They convey length information to the central nervous system via afferent nerve fibers. This information can be ...

, chordotonal neurons), 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.

Conscious and nonconscious

In humans, a distinction is made between ''conscious'' proprioception and ''nonconscious'' proprioception: * Conscious proprioception is communicated by the

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

to the

cerebrum The cerebrum (: cerebra), telencephalon or endbrain is the largest part of the brain, containing the cerebral cortex (of the two cerebral hemispheres) as well as several subcortical structures, including the hippocampus, basal ganglia, and olfac ...

. * Nonconscious proprioception is communicated primarily via the

dorsal spinocerebellar tract The spinocerebellar tracts are nerve tracts originating in the spinal cord and terminating in the same side ( ipsilateral) of the cerebellum. The two main tracts are the dorsal spinocerebellar tract, and the ventral spinocerebellar tract. Both of ...

and

ventral spinocerebellar tract The spinocerebellar tracts are nerve tracts originating in the spinal cord and terminating in the same side ( ipsilateral) of the cerebellum. The two main tracts are the dorsal spinocerebellar tract, and the ventral spinocerebellar tract. Both of ...

, to the

cerebellum The cerebellum (: cerebella or cerebellums; Latin for 'little brain') is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as it or eve ...

. * A nonconscious reaction is seen in the human proprioceptive reflex, or

righting reflex The righting reflex, also known as the labyrinthine righting reflex, or the Cervico-collic reflex; is a reflex that corrects the orientation of the body when it is taken out of its normal upright position. It is initiated by the vestibular system, ...

—in the event that the body tilts in any direction, the person will cock their head back to level the eyes against the horizon. This is seen even in infants as soon as they gain control of their neck muscles. This control comes from the

, the part of the brain affecting balance.

Mechanisms

Proprioception is mediated by mechanically sensitive proprioceptor neurons distributed throughout an animal's body. Most vertebrates possess three basic types of proprioceptors:

s, which are embedded in

skeletal muscle Skeletal muscle (commonly referred to as muscle) is one of the three types of vertebrate muscle tissue, the others being cardiac muscle and smooth muscle. They are part of the somatic nervous system, voluntary muscular system and typically are a ...

Golgi tendon organ The Golgi tendon organ (GTO) (also called Golgi organ, tendon organ, neurotendinous organ or neurotendinous spindle) is a proprioceptor – a type of sensory receptor that senses changes in muscle tension. It lies at the interface between a mus ...

s, which lie at the interface of muscles and tendons, and joint receptors, which are low-threshold mechanoreceptors embedded in

joint capsule In anatomy, a joint capsule or articular capsule is an envelope surrounding a synovial joint.chordotonal neurons, campaniform sensilla, and

s. The initiation of proprioception is the activation of a proprioceptor in the periphery. The proprioceptive sense is believed to be composed of information from

sensory neuron Sensory neurons, also known as afferent neurons, are neurons in the nervous system, that convert a specific type of stimulus, via their receptors, into action potentials or graded receptor potentials. This process is called sensory transduc ...

s located in the

inner ear The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...

(motion and orientation) and in the

stretch receptor Stretch receptors are mechanoreceptors responsive to distention of various organs and muscles, and are neurologically linked to the Medulla oblongata, medulla in the brain stem via Afferent nerve fiber, afferent nerve fibers. Examples include stre ...

s located in the

muscles Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...

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 adequate stimuli receptors. Members of the

transient receptor potential Transient receptor potential channels (TRP channels) are a group of ion channels located mostly on the plasma membrane of numerous animal cell types. Most of these are grouped into two broad groups: Group 1 includes TRPC ( "C" for canonical), TRP ...

family of

ion channel Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by Gating (electrophysiol ...

s have been found to be important for proprioception in fruit flies, nematode worms,

African clawed frog The African clawed frog (''Xenopus laevis''), also known as simply xenopus, African clawed toad, African claw-toed frog or the ''platanna'') is a species of African Aquatic animal, aquatic frog of the family Pipidae. Its name is derived from the ...

s, and

zebrafish The zebrafish (''Danio rerio'') is a species of freshwater ray-finned fish belonging to the family Danionidae of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (an ...

PIEZO2 Piezo-type mechanosensitive ion channel component 2 is a protein that in humans is encoded by the ''PIEZO2'' gene. It has a homotrimeric structure, with three blades curving into a nano-dome, with a diameter of 28 nanometers. Function Piezos ...

, a nonselective cation channel, has been shown to underlie the mechanosensitivity of proprioceptors in mice. Humans with loss-of-function mutations in the ''PIEZO2'' gene exhibit specific deficits in joint proprioception, as well as vibration and touch discrimination, suggesting that the PIEZO2 channel is essential for mechanosensitivity in some proprioceptors and low-threshold mechanoreceptors. Although it was known that finger kinesthesia relies on skin sensation, recent research has found that kinesthesia-based

haptic perception Haptic perception ( "palpable", ''haptikόs'' "suitable for touch") means literally the ability "to grasp something", and is also known as stereognosis. Perception in this case is achieved through the active exploration of surfaces and objects by a ...

relies strongly on the forces experienced during touch. This research allows the creation of "virtual", illusory haptic shapes with different perceived qualities.

Anatomy

Proprioception of the head stems from the muscles innervated by the

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

, where the

general somatic afferent fibers The general somatic afferent fibers (GSA or somatic sensory fibers) are afferent fibers that arise from neurons in sensory ganglia and are found in all the spinal nerves, except occasionally the first cervical. General somatic afferents conduct im ...

pass without synapsing in the

trigeminal ganglion The trigeminal ganglion (also known as: Gasserian ganglion, semilunar ganglion, or Gasser's ganglion) is the sensory ganglion of each trigeminal nerve (CN V). The trigeminal ganglion is located within the trigeminal cave (Meckel's cave), a cav ...

(first-order sensory neuron), reaching the mesencephalic tract and the

mesencephalic nucleus of trigeminal nerve The mesencephalic nucleus of trigeminal nerve is one of the sensory nuclei of the Trigeminal nerve, trigeminal nerve (cranial nerve V). It is located in the brainstem. It receives Proprioception, proprioceptive sensory information from the muscles ...

. Proprioception of limbs often occurs due to receptors in

connective tissue Connective tissue is one of the four primary types of animal tissue, a group of cells that are similar in structure, along with epithelial tissue, muscle tissue, and nervous tissue. It develops mostly from the mesenchyme, derived from the mesod ...

near joints.

Function

Stability

An important role for proprioception is to allow an animal to stabilize itself against perturbations. 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. Proprioceptor reflex circuits 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 agonist-antagonist muscle pairs.

Planning and refining movements

When planning complex movements such as reaching or grooming, 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.

Development

In adult fruit flies, each proprioceptor class arises from a specific

cell lineage Cell lineage denotes the developmental history of a tissue or organ from the fertilized egg. This is based on the tracking of an organism's cellular ancestry due to the cell divisions and relocation as time progresses. This starts with the origina ...

(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. The mechanisms underlying axon guidance are similar across invertebrates and vertebrates. In mammals with longer gestation periods,

are fully formed at birth. Muscle spindles continue to grow throughout post-natal development as muscles grow.

Mathematical models

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 or to provide more realistic feedback in neuromechanical simulations. 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

s, but

s and insects'

s have been modeled too.

Muscle spindles

Poppele and Bowman used linear system theory 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 Laplace transfer function describes the firing rate responses of the primary sensory fibers for a change in length:

H(s) = K_1 \frac

The following equation describes the response of secondary sensory fibers:

H(s) = K_2 \frac

More recently, Blum et al. 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.

Golgi tendon organs

Houk and Simon 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:

r(t) = K + A\exp(-at) + B \exp(-b t) + C \exp(-c t) u(t)

where

r(t)

is the firing rate and

u(t)

is a step function of force. The corresponding Laplace transfer function for this system is:

H(s) = K\left(1 + \frac  + \frac + \frac \right)

For a soleus 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 improved upon this model by adding a logarithmic nonlinearity before the Houk and Simon model and a threshold nonlinearity after.

Impairment

Chronic

Proprioception, a sense vital for rapid and proper body coordination, can be permanently lost or impaired as a result of genetic conditions, disease, viral infections, and injuries. For instance, patients with joint hypermobility or Ehlers–Danlos syndromes, genetic conditions that result in weak connective tissue throughout the body, have chronic impairments to proprioception.

Autism spectrum disorder Autism, also known as autism spectrum disorder (ASD), is a neurodevelopmental disorder characterized by differences or difficulties in social communication and interaction, a preference for predictability and routine, sensory processing di ...

and

Parkinson's disease Parkinson's disease (PD), or simply Parkinson's, is a neurodegenerative disease primarily of the central nervous system, affecting both motor system, motor and non-motor systems. Symptoms typically develop gradually and non-motor issues become ...

can also cause chronic disorder of proprioception. 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". 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

phantom limb syndrome A phantom limb is the sensation that an amputation, amputated or missing limb (anatomy), limb is still attached. It is a chronic condition that is often resistant to treatment. When the cut ends of sensory fibres are stimulated during thigh movem ...

. 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

phantom limb A phantom limb is the sensation that an amputated or missing limb is still attached. It is a chronic condition that is often resistant to treatment. When the cut ends of sensory fibres are stimulated during thigh movements, the patient feels as ...

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. 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 (

phantom eye syndrome Phantom eye syndrome (PES) is a phantom pain in the eye and visual hallucinations after the removal of an eye ( enucleation, evisceration). Symptoms Many patients experience one or more phantom phenomena after the removal of the eye: *Phant ...

). There is a decline in the sense of proprioception with

ageing Ageing (or aging in American English) is the process of becoming older until death. The term refers mainly to humans, many other animals, and fungi; whereas for example, bacteria, perennial plants and some simple animals are potentially biol ...

. This can often result in chronic lower back pain, and be the cause of falls in the elderly.

Acute

Proprioception is occasionally impaired spontaneously, especially when one is tired. Similar effects can be felt during the hypnagogic state of consciousness, 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

epilepsy Epilepsy is a group of Non-communicable disease, non-communicable Neurological disorder, neurological disorders characterized by a tendency for recurrent, unprovoked Seizure, seizures. A seizure is a sudden burst of abnormal electrical activit ...

migraine Migraine (, ) is a complex neurological disorder characterized by episodes of moderate-to-severe headache, most often unilateral and generally associated with nausea, and light and sound sensitivity. Other characterizing symptoms may includ ...

auras. These effects are presumed to arise from abnormal stimulation of the part of the

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

of the

brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...

involved with integrating information from different parts of the body. Proprioceptive illusions can also be induced, such as the "Pinocchio illusion", the illusion that one's nose is growing longer. Temporary impairment of proprioception has also been known to occur from an overdose of

vitamin B6 Vitamin B6 is one of the B vitamins, and is an essential nutrient for humans. The term essential nutrient refers to a group of six chemically similar compounds, i.e., "vitamers", which can be interconverted in biological systems. Its active f ...

(pyridoxine and pyridoxamine). This is due to a reversible neuropathy. 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

cytotoxic Cytotoxicity is the quality of being toxic to cells. Examples of toxic agents are toxic metals, toxic chemicals, microbe neurotoxins, radiation particles and even specific neurotransmitters when the system is out of balance. Also some types of dr ...

factors such as

chemotherapy Chemotherapy (often abbreviated chemo, sometimes CTX and CTx) is the type of cancer treatment that uses one or more anti-cancer drugs (list of chemotherapeutic agents, chemotherapeutic agents or alkylating agents) in a standard chemotherapy re ...

. It has been proposed that even common

tinnitus Tinnitus is a condition when a person hears a ringing sound or a different variety of sound when no corresponding external sound is present and other people cannot hear it. Nearly everyone experiences faint "normal tinnitus" in a completely ...

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. 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 (

liposuction Liposuction, or simply lipo, is a type of fat-removal procedure used in plastic surgery. Evidence does not support an effect on weight beyond a couple of months and does not appear to affect obesity-related problems. In the United States, lip ...

, rapid fat loss or gain) and/or muscle content (

bodybuilding Bodybuilding is the practice of Resistance training, progressive resistance exercise to build, control, and develop one's skeletal muscle, muscles via muscle hypertrophy, hypertrophy. An individual who engages in this activity is referred to a ...

anabolic steroid Anabolic steroids, also known as anabolic-androgenic steroids (AAS), are a class of drugs that are structurally related to testosterone, the main male sex hormone, and produce effects by binding to the androgen receptor (AR). Anabolism, Anaboli ...

s, catabolisis/

starvation Starvation is a severe deficiency in caloric energy intake, below the level needed to maintain an organism's life. It is the most extreme form of malnutrition. In humans, prolonged starvation can cause permanent organ damage and eventually, de ...

). It can also occur in those that gain new levels of

flexibility Stiffness is the extent to which an object resists deformation in response to an applied force. The complementary concept is flexibility or pliability: the more flexible an object is, the less stiff it is. Calculations The stiffness, k, of a ...

stretching Stretching is a form of physical exercise in which a specific muscle or tendon (or muscle group) is deliberately expanded and flexed in order to improve the muscle's felt elasticity and achieve comfortable muscle tone. The result is a feelin ...

, and contortion. 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.

Diagnosis

Impaired proprioception may be diagnosed through a series of tests, each focusing on a different functional aspect of proprioception. The Romberg's test 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. 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. 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. 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

police officer A police officer (also called policeman or policewoman, cop, officer or constable) is a Warrant (law), warranted law employee of a police, police force. In most countries, ''police officer'' is a generic term not specifying a particular rank. ...

s using the field sobriety testing to check for

alcohol intoxication Alcohol intoxication, commonly described in higher doses as drunkenness or inebriation, and known in overdose as alcohol poisoning, is the behavior and physical effects caused by recent consumption of Alcohol (drug), alcohol. The technical ter ...

. 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 , 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.

Training

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 touch type or perform ballet; and people would not even be able to walk without watching where they put their feet.

Oliver Sacks Oliver Wolf Sacks (9 July 1933 – 30 August 2015) was a British neurology, neurologist, Natural history, naturalist, historian of science, and writer. Born in London, Sacks received his medical degree in 1958 from The Queen's College, Oxford ...

reported the case of a young woman who lost her proprioception due to a viral infection of her

spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the lower brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal c ...

. 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

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 wobble board or balance board is often used to retrain or increase proprioceptive abilities, particularly as

physical therapy Physical therapy (PT), also known as physiotherapy, is a healthcare profession, as well as the care provided by physical therapists who promote, maintain, or restore health through patient education, physical intervention, disease preventio ...

for ankle or knee injuries. Slacklining 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

yoga Yoga (UK: , US: ; 'yoga' ; ) is a group of physical, mental, and spiritual practices or disciplines that originated with its own philosophy in ancient India, aimed at controlling body and mind to attain various salvation goals, as pra ...

Wing Chun Wing Chun (Cantonese) or Yong Chun (Mandarin Chinese, Mandarin) (, lit. "singing spring") is a concept-based martial art, a form of Nanquan (martial art), Southern Chinese kung fu, and a close-quarters system of self-defense. It is a martial ...

and

tai chi is a Chinese martial art. Initially developed for combat and self-defense, for most practitioners it has evolved into a sport and form of exercise. As an exercise, tai chi is performed as gentle, low-impact movement in which practitioners ...

. The vestibular system of the inner ear, vision and proprioception are the main three requirements for balance. Moreover, there are specific devices designed for proprioception training, such as the exercise ball, which works on balancing the abdominal and back muscles.

History of study

In 1557, the position-movement sensation was described by

Julius Caesar Scaliger Julius Caesar Scaliger (; 23 April or August 1484 – 21 October 1558), or Giulio Cesare della Scala, was an Italian scholar and physician, who spent a major part of his career in France. He employed the techniques and discoveries of Renaissance ...

as a "sense of locomotion". In 1826,

Charles Bell Sir Charles Bell (12 November 177428 April 1842) was a Scottish surgeon, anatomist, physiologist, neurologist, artist, and philosophical theologian. He is noted for discovering the difference between sensory nerves and motor nerves in the ...

expounded the idea of a "muscle sense", which is credited as one of the first descriptions of physiologic feedback mechanisms. 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 Robert Todd highlighted important differences in the anterolateral and posterior columns of the spinal cord, and suggested that the latter were involved in the coordination of movement and balance. At around the same time, Moritz Heinrich Romberg, a Berlin neurologist, was describing unsteadiness made worse by eye closure or darkness, now known as the eponymous

Romberg's sign Romberg's test, Romberg's sign, or the Romberg maneuver is a test used in an exam of neurological function for balance. The exam is based on the premise that a person requires at least two of the three following senses to maintain balance whil ...

, once synonymous with

tabes dorsalis Tabes dorsalis is a late consequence of neurosyphilis, characterized by the slow degeneration (specifically, demyelination) of the neural tracts primarily in the Dorsal root ganglion, dorsal root ganglia of the spinal cord (nerve root). These pati ...

, that became recognised as common to all proprioceptive disorders of the legs. In 1880,

Henry Charlton Bastian Henry Charlton Bastian (26 April 1837 in Truro, Cornwall, England – 17 November 1915 in Chesham Bois, Buckinghamshire) was an English physiologist and neurologist. Biography Bastian was born at Truro, Truro, Cornwall and graduated from Unive ...

suggested "kinaesthesia" instead of "muscle sense" on the basis that some of the afferent information (back to the brain) comes from other structures, including tendons, joints, and skin. In 1889, Alfred Goldscheider suggested a classification of kinaesthesia into three types: muscle, tendon, and articular sensitivity. In 1906, the term ''proprio-ception'' (and also '' intero-ception'' and ''extero-ception'') is attested in a publication by

Charles Scott Sherrington Sir Charles Scott Sherrington (27 November 1857 – 4 March 1952) was a British neurophysiology, neurophysiologist. His experimental research established many aspects of contemporary neuroscience, including the concept of the spinal reflex as a ...

involving

receptors Receptor may refer to: *Sensory receptor, in physiology, any neurite structure that, on receiving environmental stimuli, produces an informative nerve impulse *Receptor (biochemistry), in biochemistry, a protein molecule that receives and responds ...

. He explains the terminology as follows: Today, the "exteroceptors" are the organs that provide information originating outside the body, such as the eyes, ears, mouth, and skin. The interoceptors 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

s and

), 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". Secondary endings of muscle spindles detect changes in muscle length, and thus supply information regarding only the sense of position. Essentially, muscle spindles are stretch receptors. 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.

Etymology

Proprioception is from

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

'' proprius'', meaning "one's own", "individual", and '' capio'', ''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").

Plants and bacteria

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 (

angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit. T ...

Terrestrial plant A terrestrial plant is a plant that grows on, in or from land. Other types of plants are aquatic plant, aquatic (living in or on water), semiaquatic (living at edge or seasonally in water), epiphyte, epiphytic (living on other plants), and litho ...

s control the orientation of their

primary growth Primary growth in plants is growth that takes place from the tips of roots or shoots. It leads to lengthening of roots and stems and sets the stage for organ formation. It is distinguished from secondary growth that leads to widening. Plant growth ...

through the sensing of several vectorial stimuli such as the light gradient or the

gravitational acceleration In physics, gravitational acceleration is the acceleration of an object in free fall within a vacuum (and thus without experiencing drag (physics), drag). This is the steady gain in speed caused exclusively by gravitational attraction. All bodi ...

. This control has been called

tropism In biology, a tropism is a phenomenon indicating the growth or turning movement of an organism, usually a plant, in response to an environmental stimulus (physiology), stimulus. In tropisms, this response is dependent on the direction of the s ...

. A quantitative study of shoot

gravitropism Gravitropism (also known as geotropism) is a coordinated process of differential growth by a plant in response to gravity pulling on it. It also occurs in fungi. Gravity can be either "artificial gravity" or natural gravity. It is a general feat ...

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

curvature In mathematics, curvature is any of several strongly related concepts in geometry that intuitively measure the amount by which a curve deviates from being a straight line or by which a surface deviates from being a plane. If a curve or su ...

by the organ and the subsequent driving an active straightening process are required. 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

mathematical model A mathematical model is an abstract and concrete, abstract description of a concrete system using mathematics, mathematical concepts and language of mathematics, language. The process of developing a mathematical model is termed ''mathematical m ...

simulating the complete driving of the gravitropic movement. This model has been validated on 11 species sampling the phylogeny of land angiosperms, and on organs of very contrasted sizes, ranging from the small germination of

wheat Wheat is a group of wild and crop domestication, domesticated Poaceae, grasses of the genus ''Triticum'' (). They are Agriculture, cultivated for their cereal grains, which are staple foods around the world. Well-known Taxonomy of wheat, whe ...

(

coleoptile Coleoptile is the pointed protective sheath covering the emerging shoot in monocotyledons such as grasses in which few leaf primordia and shoot apex of monocot embryo remain enclosed. The coleoptile protects the first leaf as well as the growing ...

) to the trunk of poplar trees. Further studies have shown that the cellular mechanism of proprioception in plants involves

myosin Myosins () are a Protein family, family of motor proteins (though most often protein complexes) best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are adenosine triphosphate, ATP- ...

and

actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ...

, and seems to occur in specialized cells. Proprioception was then found to be involved in other tropisms and to be central also to the control of

nutation Nutation () is a rocking, swaying, or nodding motion in the axis of rotation of a largely axially symmetric object, such as a gyroscope, planet, or bullet in flight, or as an intended behaviour of a mechanism. In an appropriate reference fra ...

. The discovery of proprioception in plants has generated an interest in the popular science and generalist media. 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

self-awareness In philosophy of self, philosophy, self-awareness is the awareness and reflection of one's own personality or individuality, including traits, feelings, and behaviors. It is not to be confused with consciousness in the sense of qualia. While ...

self-consciousness Self-consciousness is a heightened sense of awareness of oneself. It is not to be confused with consciousness in the sense of qualia. Historically, "self-consciousness" was synonymous with " self-awareness", referring to a state of awareness th ...

. 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. Recent studies suggest that bacteria have control systems that may resemble proprioception.

Notes

References

External links

* {{Sensation and perception Articles containing video clips Sensory systems