Proprioception

Overview
Proprioception ( PRO-pree-o-SEP-shun); from Latin proprius, meaning "one's own" and perception) is the sense of the relative position of neighbouring parts of the body. Unlike the six exteroceptive senses (sight, taste, smell, touch, hearing, and balance) by which we perceive the outside world, and interoceptive senses, by which we perceive the pain and the stretching of internal organs, proprioception is a third distinct sensory modality that provides feedback solely on the status of the body internally. It is the sense that indicates whether the body is moving with required effort, as well as where the various parts of the body are located in relation to each other. The Position-Movement sensation was originally described in 1557 by Julius Caesar Scaliger as a 'sense of locomotion'. Much later in 1826 Charles Bell expounded the idea of a 'muscle sense' and this is credited with being one of the first physiologic feedback mechanisms. Bell's idea was that commands were being carried from the brain to the muscles, and that reports on the muscle's condition would be sent in the reverse direction. Later, in 1880, Henry Charlton Bastian suggested 'kinaesthesia' instead of 'muscle sense' on the basis that some of the afferent information (back to the brain) was coming from other structures including tendon, joints, skin, and muscle. In 1889, Alfred Goldscheider suggested a classification of kinaesthesia into 3 types: muscle, tendon, and articular sensitivity. In 1906, Sherrington published a landmark work which introduced the terms 'proprioception', 'interoception', and 'exteroception'. The 'exteroceptors' being the organs responsible for information from outside the body such as the eyes, ears, mouth, and skin. The interoceptors then gave information about the internal organs, while 'proprioception' was awareness of movement derived from muscular, tendon, and articular sources. Such a system of classification has kept physiologists and anatomists searching for specialised nerve endings which transmit data on joint capsule and muscle tension (such as muscle spindles and Pacini corpuscles).

Proprioception vs. kinesthesia
Kinesthesia is another term that is often used interchangeably with proprioception. Some differentiate the kinesthetic sense from proprioception by excluding the sense of equilibrium or balance from kinesthesia. An inner ear infection, for example, might degrade the sense of balance. This would degrade the proprioceptive sense, but not the kinesthetic sense. The infected person would be able to walk, but only by using the sense of sight to maintain balance; the person would be unable to walk with eyes closed.

Proprioception and kinaesthesia are seen as interrelated and there is considerable disagreement regarding the definition of these terms. Some of this difficulty stems from Sherrington's original description of joint position sense (or the ability to determine where a particular body part exactly is in space) and kinaesthesia (or the sensation that the body part has moved) under a more general heading of proprioception. Clinical aspects of proprioception are measured in tests that measure a subject's ability to detect an externally imposed passive movement, or the ability to reposition a joint to a predetermined position. Often it is assumed that the ability of one of these aspects will be related to another; unfortunately, experimental evidence suggests there is no strong relation between these two aspects. This suggests that, while these components may well be related in a cognitive manner, they seem to be separate physiologically.

Much of the foregoing work is dependent on the notion that proprioception is essentially a feedback mechanism; that is, the body moves (or is moved) and then the information about this is returned to the brain, whereby subsequent adjustments could be made. More recent work into the mechanism of ankle sprains suggests that the role of reflexes may be more limited due to their long latencies (even at the spinal cord level) as ankle sprain events occur in perhaps 100msec or less. Accordingly, a model has been proposed to include a 'feedforward' component of proprioception where the subject will also have central information about the body's position prior to attaining it.

Kinesthesia is a key component in muscle memory and hand-eye coordination, and training can improve this sense (see blind contour drawing). The ability to swing a golf club or to catch a ball requires a finely-tuned sense of the position of the joints. This sense needs to become automatic through training to enable a person to concentrate on other aspects of performance, such as maintaining motivation or seeing where other people are.

Basis of proprioceptive sense
The proprioceptive sense is believed to be composed of information from sensory neurons located in the inner ear (motion and orientation) and in the stretch receptors located in the muscles and the joint-supporting ligaments (stance). There are specific nerve receptors for this form of perception, just as there are specific receptors for pressure, light, temperature, sound, and other sensory experiences, known as adequate stimuli receptors.

Although it was known that finger kinesthesia relies on skin sensation, recent research has found that kinesthesia-based haptic perception strongly relies on the forces experienced during touch. This research allows the creation of "virtual", illusory haptic shapes with different perceived qualities.

Law enforcement
Proprioception is tested by American police officers using the field sobriety test where 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 millimetres. People suffering from 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.

Learning
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 foot 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 once reported the case of a young woman who lost her proprioception due to a viral infection of her spinal cord. At first she was not able to 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 vestibulum (or inner ear) 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 didn't drop them.

Training
The proprioceptive sense can be sharpened through study of many disciplines. The Alexander Technique uses the study of movement to enhance kinesthetic judgment of effort and location. Juggling trains reaction time, spatial location, and efficient movement. Standing on a wobble board or balance board is often used to retrain or increase proprioception abilities, particularly as physical therapy for ankle or knee injuries. Standing on one leg (stork standing) and various other body-position challenges are also used in such disciplines as Yoga or Wing Chun. In addition, the slow, focused movements of Tai Chi practice provide an environment whereby the proprioceptive information being fed back to the brain stimulates an intense, dynamic "listening environment" to further enhance mind / body integration. Several studies have shown that the efficacy of these types of training is challenged by closing the eyes, because the eyes give invaluable feedback to establishing the moment-to-moment information of balance.

Impairment
Apparently, 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, rapid fat loss, rapid fat gain) and muscle content (bodybuilding, anabolic steroids, catabolisis/starvation). It can also occur to those who gain new levels of flexibility, stretching, 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 these: 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.

Proprioception is occasionally impaired spontaneously, especially when one is tired. One's body may appear too large or too small, or parts of the body may appear distorted in size. Similar effects can sometimes occur during epilepsy or migraine auras. These effects are presumed to arise from abnormal stimulation of the part of the parietal cortex of the brain involved with integrating information from different parts of the body.

Proprioceptive illusions can also be induced, such as the Pinocchio illusion.

The proprioceptive sense is often unnoticed because humans will adapt to a continuously-present stimulus; this is called habituation, desensitization, or adaptation. The effect is that proprioceptive sensory impressions disappear, just as a scent can disappear over time. One practical advantage of this is that unnoticed actions or sensation continue in the background while an individual's attention can move to another concern. The Alexander Technique addresses these issues.

People who have a limb amputated may still have a confused sense of that limb existence on their body, known as Phantom Limb Syndrome. 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. The etiology of the phantom limb phenomenon was disputed in 2006, but some consensus existed in favour of neurological (e.g. neural signal bleed across a preexisting sensory map, as posited by V.S. Ramachandran) over psychological explanations. 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).

Temporary impairment of proprioception has also been known to occur from an overdose of vitamin B6 (pyridoxine and pyridoxamine). Most of the impaired function returns to normal shortly after the intake of vitamins returns to normal. Impairment can also be caused by cytotoxic factors such as chemotherapy.

It has been proposed that even common Tinnitus 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.

Proprioception is permanently impaired in patients who suffer from joint hypermobility or Ehlers-Danlos Syndrome (a genetic condition that results in weak connective tissue throughout the body). It can also be permanently impaired from viral infections as reported by Sacks. The catastrophic effect of major proprioceptive loss is reviewed by Robles-De-La-Torre (2006).