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Kinesthetic Senses

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ABSTRACT

The kinesthetic senses are the senses of position and movement of the body, senses we are aware of only on introspection. A method used to study kinesthesia is muscle vibration, which engages afferents of muscle spindles to trigger illusions of movement and changed position. When vibrating elbow flexors, it generates sensations of forearm extension, when vibrating extensors, sensations of forearm flexion. Vibrating the elbow joint produces no illusion. Vibrating flexors and extensors together at the same frequency also produces no illusion, because what is perceived is the signal difference between antagonist muscles of each arm and between arms. The size of the illusion depends on how the muscle has been conditioned beforehand, due to a property of muscle called thixotropy. When measuring the illusion, blindfolded subjects may carry out a matching or pointing task. In pointing, signals from muscle spindles are less important than in matching. Afferent signals from kinesthetic receptors project to areas of somatosensory cortex to generate sensations of detection and location. This is referred to the body model, which provides information about size and shape of body parts. Kinesthesia, together with vision and touch, is associated with the sense of body ownership. All three can combine or each, on its own, can generate ownership. Related is the sense of agency, the sense of being responsible for one's own actions. In recent times, much progress has been made using neuroimaging techniques to identify the various areas of the brain likely to be responsible for generating these sensations. © 2017 American Physiological Society. Compr Physiol 8:1157‐1183, 2018.

Figure 1. Figure 1. Muscle thixotropy. (A) Diagrammatic representation of a string of passive sarcomeres, part of a myofibril, containing thick and thin myofilaments. Scattered randomly along the length of the myofibril are occasional points of attachment between thick and thin filaments, the stable cross‐bridges (red circles). They are responsible for giving muscle its thixotropic property. (B) Tension changes in a passive muscle during ramp stretch and shortening. During stretch, due to the presence of attached cross‐bridges, tension rises steeply, to plateau out as the yield point is reached, and bridges begin to detach to reform at the longer length. During shortening, attached cross‐bridges exert a splinting effect on the shortening muscle, leading tension to fall steeply and remain low for the remainder of the shortening. Length and tension traces redrawn, with permission, from Hill ().
Figure 2. Figure 2. (A) Flexion conditioning of elbow muscles: The blindfolded subject flexes their arm so that the paddle supporting the forearm is at right angles to the supporting base (conditioning angle). The experimenter holds the paddle in position while the subject contracts their elbow flexors, trying to pull the paddle toward their body. After a brief, half‐maximum voluntary contraction, the subject relaxes their arm and the experimenter moves it to the test position (actual). This is perceived as more extended than the true position, as shown by the image ghost (perceived angle). (B) Extension conditioning of elbow muscles: The blindfolded subject extends their forearm to lie on the supporting base (conditioning angle). They then push down on the base, to contract their elbow extensors. After a half‐maximal contraction, the subject relaxes their arm and the experimenter moves it to the test angle (actual). This is perceived as more flexed than the true position, as shown by the image ghost (perceived angle).
Figure 3. Figure 3. Distribution of position errors at the forearm from different forms of thixotropic conditioning: The blindfolded subjects matched positions of their forearms after conditioning of elbow muscles. Left‐hand values: mean (±SEM) position errors for 14 subjects at 5 s after arm alignment. After flexion conditioning of both arms, small errors in the direction of flexion are made (blue circle); after extension conditioning, similar sized errors are made in the direction of extension (red circle). Replotted, with permission, from Ref. (). Middle values: position errors for the same 14 subjects after co‐conditioning of the reference arm (see Fig. 4) and flexion conditioning of the indicator arm (blue circle) or extension conditioning of the indicator arm (red circle). Large errors in the direction of flexion are made after flexion conditioning of the indicator arm and in the direction of extension after extension conditioning of the indicator arm. Data replotted, with permission, from Ref (). Right‐hand values: errors (±SEM) for nine subjects after co‐conditioning of the reference arm with slack introduced in the indicator arm. Slack is introduced in one of two ways; the indicator is flexion conditioned and then the arm is moved into full extension before being returned to the test angle (blue circle). Alternatively, the arm is extension conditioned, then fully flexed before being returned to the test angle (red circle). Data replotted, with permission, from Ref. ().
Figure 4. Figure 4. Test angle co‐conditioning: The experimenter moves the blindfolded subject's forearm to the test position (actual). While holding the paddle, the experimenter asks the subject first to pull it toward their body, to generate a contraction in elbow flexors, then to push it into extension to contract elbow extensors. The subject then relaxes their arm, which is perceived to adopt a position close to its true position, as shown by the ghost image (perceived angle).
Figure 5. Figure 5. Measuring position sense in matching and pointing tasks: In the matching task (A), blindfolded subjects’ arms were strapped to paddles, one arm designated the reference, the other the indicator. Forearm position was measured by potentiometers located at the hinges of the paddles, which were colinear with the elbow joint. The experimenter placed the reference arm at a test angle and the subject held it there while they moved their indicator arm into a matching position. In the pointing task (B), the two arms were separated by a screen (dashed line) which blocked the subject's view of the reference arm. The reference arm was strapped to a paddle, as before, and was placed at the test angle by the experimenter. Once the reference arm was in position, the subject pressed a lever with their other hand to move the pointer paddle to align it with the perceived position of the hidden reference arm. Data replotted, with permission, from Ref. ().
Figure 6. Figure 6. Position errors in matching and pointing tasks. (A) Two arm matching task. Left‐hand data points, both arms were flexion conditioned before the match; red circle, mean (±SEM) error for 14 subjects, blue circle, predicted error. Right‐hand data points, the arms were extension conditioned; red circle, mean measured error, blue circle, expected error. Replotted, with permission, from Ref. (). (B) Pointing to the position of a hidden arm. Left‐hand data points, the arm was flexion conditioned. Red circle, mean (±SEM) error for 14 subjects, blue circle, expected error. Right‐hand values, the arm was extension conditioned. Red circle, observed error, blue circle, expected error. Replotted, with permission, from Ref. ().
Figure 7. Figure 7. Subjects have their right hand resting on the lower table. The right (test) index finger is held in a pipe connected to a shaft on which an artificial prosthetic finger is mounted. 12 cm separates the subject's test index and the artificial finger. The vertical dotted line indicates the axis of rotation of the shaft, which was colinear with the proximal interphalangeal joint of the artificial finger and the subject's test finger. A coupling on the shaft allowed movement between the artificial finger and the subject's test finger to be either congruent or incongruent. In some studies, vision is excluded and the experimenter holds the thumb in a passive pinch grip on the artificial finger. In other studies, the experimenter moves the artificial finger and this can be seen by the subject (the left hand is not involved). Redrawn, with permission, from Ref. ().
Figure 8. Figure 8. (A) The effect of congruent movement and blocking the digital nerves of the grasping and test fingers on perceived ownership over the artificial finger. The median ± interquartile range responses to the statement “I feel that I am holding my right index finger with my left hand.” Following the digital nerve blocks median responses were higher after congruent movement than incongruent movement. (B) Perceived vertical spacing between the index fingers for congruent and incongruent movement trials after the digital nerves of the test index finger and the grasping index finger and thumb were blocked with local anesthetic. A smaller median perceived spacing was reported after congruent movement compared to incongruent movement. Results show median ± interquartile range. Replotted, with permission, from Ref. ().


Figure 1. Muscle thixotropy. (A) Diagrammatic representation of a string of passive sarcomeres, part of a myofibril, containing thick and thin myofilaments. Scattered randomly along the length of the myofibril are occasional points of attachment between thick and thin filaments, the stable cross‐bridges (red circles). They are responsible for giving muscle its thixotropic property. (B) Tension changes in a passive muscle during ramp stretch and shortening. During stretch, due to the presence of attached cross‐bridges, tension rises steeply, to plateau out as the yield point is reached, and bridges begin to detach to reform at the longer length. During shortening, attached cross‐bridges exert a splinting effect on the shortening muscle, leading tension to fall steeply and remain low for the remainder of the shortening. Length and tension traces redrawn, with permission, from Hill ().


Figure 2. (A) Flexion conditioning of elbow muscles: The blindfolded subject flexes their arm so that the paddle supporting the forearm is at right angles to the supporting base (conditioning angle). The experimenter holds the paddle in position while the subject contracts their elbow flexors, trying to pull the paddle toward their body. After a brief, half‐maximum voluntary contraction, the subject relaxes their arm and the experimenter moves it to the test position (actual). This is perceived as more extended than the true position, as shown by the image ghost (perceived angle). (B) Extension conditioning of elbow muscles: The blindfolded subject extends their forearm to lie on the supporting base (conditioning angle). They then push down on the base, to contract their elbow extensors. After a half‐maximal contraction, the subject relaxes their arm and the experimenter moves it to the test angle (actual). This is perceived as more flexed than the true position, as shown by the image ghost (perceived angle).


Figure 3. Distribution of position errors at the forearm from different forms of thixotropic conditioning: The blindfolded subjects matched positions of their forearms after conditioning of elbow muscles. Left‐hand values: mean (±SEM) position errors for 14 subjects at 5 s after arm alignment. After flexion conditioning of both arms, small errors in the direction of flexion are made (blue circle); after extension conditioning, similar sized errors are made in the direction of extension (red circle). Replotted, with permission, from Ref. (). Middle values: position errors for the same 14 subjects after co‐conditioning of the reference arm (see Fig. 4) and flexion conditioning of the indicator arm (blue circle) or extension conditioning of the indicator arm (red circle). Large errors in the direction of flexion are made after flexion conditioning of the indicator arm and in the direction of extension after extension conditioning of the indicator arm. Data replotted, with permission, from Ref (). Right‐hand values: errors (±SEM) for nine subjects after co‐conditioning of the reference arm with slack introduced in the indicator arm. Slack is introduced in one of two ways; the indicator is flexion conditioned and then the arm is moved into full extension before being returned to the test angle (blue circle). Alternatively, the arm is extension conditioned, then fully flexed before being returned to the test angle (red circle). Data replotted, with permission, from Ref. ().


Figure 4. Test angle co‐conditioning: The experimenter moves the blindfolded subject's forearm to the test position (actual). While holding the paddle, the experimenter asks the subject first to pull it toward their body, to generate a contraction in elbow flexors, then to push it into extension to contract elbow extensors. The subject then relaxes their arm, which is perceived to adopt a position close to its true position, as shown by the ghost image (perceived angle).


Figure 5. Measuring position sense in matching and pointing tasks: In the matching task (A), blindfolded subjects’ arms were strapped to paddles, one arm designated the reference, the other the indicator. Forearm position was measured by potentiometers located at the hinges of the paddles, which were colinear with the elbow joint. The experimenter placed the reference arm at a test angle and the subject held it there while they moved their indicator arm into a matching position. In the pointing task (B), the two arms were separated by a screen (dashed line) which blocked the subject's view of the reference arm. The reference arm was strapped to a paddle, as before, and was placed at the test angle by the experimenter. Once the reference arm was in position, the subject pressed a lever with their other hand to move the pointer paddle to align it with the perceived position of the hidden reference arm. Data replotted, with permission, from Ref. ().


Figure 6. Position errors in matching and pointing tasks. (A) Two arm matching task. Left‐hand data points, both arms were flexion conditioned before the match; red circle, mean (±SEM) error for 14 subjects, blue circle, predicted error. Right‐hand data points, the arms were extension conditioned; red circle, mean measured error, blue circle, expected error. Replotted, with permission, from Ref. (). (B) Pointing to the position of a hidden arm. Left‐hand data points, the arm was flexion conditioned. Red circle, mean (±SEM) error for 14 subjects, blue circle, expected error. Right‐hand values, the arm was extension conditioned. Red circle, observed error, blue circle, expected error. Replotted, with permission, from Ref. ().


Figure 7. Subjects have their right hand resting on the lower table. The right (test) index finger is held in a pipe connected to a shaft on which an artificial prosthetic finger is mounted. 12 cm separates the subject's test index and the artificial finger. The vertical dotted line indicates the axis of rotation of the shaft, which was colinear with the proximal interphalangeal joint of the artificial finger and the subject's test finger. A coupling on the shaft allowed movement between the artificial finger and the subject's test finger to be either congruent or incongruent. In some studies, vision is excluded and the experimenter holds the thumb in a passive pinch grip on the artificial finger. In other studies, the experimenter moves the artificial finger and this can be seen by the subject (the left hand is not involved). Redrawn, with permission, from Ref. ().


Figure 8. (A) The effect of congruent movement and blocking the digital nerves of the grasping and test fingers on perceived ownership over the artificial finger. The median ± interquartile range responses to the statement “I feel that I am holding my right index finger with my left hand.” Following the digital nerve blocks median responses were higher after congruent movement than incongruent movement. (B) Perceived vertical spacing between the index fingers for congruent and incongruent movement trials after the digital nerves of the test index finger and the grasping index finger and thumb were blocked with local anesthetic. A smaller median perceived spacing was reported after congruent movement compared to incongruent movement. Results show median ± interquartile range. Replotted, with permission, from Ref. ().
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Teaching Material

U. Proske, S. C. Gandevia. Kinesthetic Senses. Compr Physiol 8: 2018, 1157-1183.

Didactic Synopsis

Major Teaching Points:

  1. The two major kinesthetic senses are the sense of position of our body and the sense of movement.
  2. They can be measured in two ways:
    • Measure the size of an illusion of position and movement of our limbs by vibrating a muscle.
    • Measure changes in perceived position after contracting a muscle and stretching or shortening it.
  3. To measure the illusion, a blindfolded subject aligns one forearm with the other to indicate its location. Alternatively, the subject points to the position of one arm hidden from view.
  4. The kinesthetic senses combine with vision and touch to give us a sense of body ownership. This is linked to a sense of agency, “I am moving my arm.”
  5. Sensory and motor areas of the brain receive kinesthetic signals to tell us where our body parts are and where they are located in relation to our surroundings.

Didactic Legends

The figures—in a freely downloadable PowerPoint format—can be found on the Images tab along with the formal legends published in the article. The following legends to the same figures are written to be useful for teaching.

Figure 1 Muscle thixotropy. Teaching points: (A) The diagram shows a series of repeating units called sarcomeres. A string of sarcomeres is called a myofibril and each skeletal muscle fiber in our body is composed of bundles of myofibrils. The sarcomere is composed of thick longitudinal filaments of a protein, myosin (shown in blue), which attaches to the adjacent thin filaments of the protein actin (shown in black) by means of cross-bridges with grape-like heads. During a muscle contraction, all of these heads attach to the actin and rotate, in the process pulling the two halves of a sarcomere towards one another and generating force. At the end of a contraction, as the muscle relaxes, the myosin heads detach from the actin. Subsequently a few, randomly scattered, heads reattach (shown in red), but they do not go through the usual rotation process to generate force. They remain attached for as long as the muscle is left undisturbed. They give the muscle a resting stiffness whose value depends on whether the muscle has been previously moved or not. Such history-dependent behavior is called thixotropy. (B) These are records of the length (blue) and tension (red) changes in a passive muscle showing thixotropic behavior during stretch and shortening movements. During muscle lengthening (upward moving blue ramp on the left), passive tension rises steeply, as the attached cross-bridges (red in A) are stressed by the stretch. A point is reached at peak tension where stress is sufficient to lead to detachment of the cross-bridges from actin and tension begins to fall. When a shortening is applied to the muscle (downward blue ramp on the right), the presence of attached cross-bridges (red in A), exert a splinting effect, preventing the muscle from fully shortening and it falls slack. This pushing effect by the attached cross-bridges leads to the precipitous fall in tension.

Figure 2 Teaching points: (A) Flexion conditioning. This is a method of placing the muscles of the human forearm in a defined state with respect to the thixotropic behavior of muscle stretch receptors, the muscle spindles, which are known to signal muscle length, that is, limb position. A blindfolded subject sits at a table, which supports both forearms. The arms have been taped to paddles hinged at a point aligned with the subject's elbow joints. Potentiometers at the hinges provide signals of elbow angles (see Figure 5). The subject is asked to relax while the experimenter moves one forearm into a vertical position (conditioning angle). The subject is then asked to contract their elbow flexor muscles by trying to pull the arm towards their body, while it is held in position by the experimenter. The contraction leads to sensitization of elbow flexor muscle spindles, as a result of their thixotropic property. After the arm has relaxed, when the experimenter extends it to the test angle (actual), this stretches the muscle spindles and they generate a strong position signal. This shows up when the subject is asked to match position of the arm with their other arm, which they place into a more extended position (perceived angle) than the actual position of the arm. (B) Extension conditioning. This is a similar procedure to that in A, but the starting point is the relaxed arm lying on the table. The subject is asked to push down on the table, in the process contracting elbow extensor muscles and sensitizing their muscle spindles (conditioning angle). The experimenter then flexes the arm, moving it to the test angle (actual) and the subject is asked to indicate location of the arm by matching its position with their other arm. As a result of the sensitization of muscle spindles, the subject thinks their extensors are more stretched, the elbow more flexed than is really the case. They, therefore, adopt a too flexed matching position with the other arm (perceived angle).

Figure 3 Distribution of position errors at the forearm from different forms of thixotropic conditioning. Teaching points: The figure shows the variety of position errors that can be generated at the human forearm as a result of thixotropic conditioning of arm muscles. All measurements of errors are by blindfolded subjects matching positions of their forearms, using the apparatus described in Figure 5A. The pair of position errors shown on the left has been measured for groups of subjects whose arms had both been flexion conditioned (blue symbol) or extension conditioned (red symbol), as shown in Figure 2. After flexion conditioning, spindles in flexor muscles of both arms have been sensitized. This leads to small matching errors in the direction of elbow flexion (blue symbol). Errors are not zero because of movement related spindle activity in the indicator arm moving into the matching position. It leads the subject to declare a match too soon, with the indicator still too flexed and before the arms are truly aligned. The same reasoning is applied to the generation of small errors into extension after extension conditioning of both arms (red symbol). The middle pair of values in the figure shows the distribution of errors when the two arms are conditioned differently from one another. One arm has both its extensor and flexor muscles co-conditioned, that is, the arm is brought to the test angle and its flexors and then extensors are contracted while the arm is held fixed in position (see Figure 4). The perceived position of the arm is indicated with the other arm, which has been flexion, conditioned or extension conditioned (Figure 2). For the co-conditioned arm, because spindle signals in both antagonist muscles are similarly sensitized, this leads to a low difference signal, that is, a low position signal. The other arm used to indicate this position has been only flexion or extension conditioned and so it has a high flexor or extensor signal. When the subject is trying to match arm positions with one arm that has a high flexor signal and the other a low signal, the subject keeps their flexor conditioned arm as flexed as possible, trying to generate a low matching signal. As a result, large errors into flexion result (blue symbol). Similarly, large errors in the direction of extension (red symbol) are generated when the indicator arm is extension conditioned. The right-hand pair of data points demonstrate that muscle conditioning can be arranged to achieve position errors that lie close to zero. Here for both error values the reference arm has been brought to the test angle and its muscles co-conditioned (see Figure 4), leading to a low position signal in that arm. To match this, it is necessary to lower the position signal in the other arm as well. That is done by introducing slack in its muscle spindles. For a flexion-conditioned arm (Figure 2), after the conditioning contraction and sensitization of flexor spindles, the arm is fully extended before being returned to the matching position. Flexion of the arm after the extension means that flexor spindles are shortened and because of their thixotropic property they are unable to shorten themselves and they fall slack (Figure 1). A slack spindle has no tension on its sensory ending and so its prevailing activity falls to low levels. Now, during the match both arms have low position signals and the error (blue symbol) is small. A similar outcome can be achieved when the indicator arm is extension conditioned. Here, after the conditioning contraction in the extended position (Figure 2), the arm is brought into full flexion before being returned to the matching position. The extension movement from flexion to the matching position introduces slack in extensor spindles and their low level of activity matches that in the co-conditioned arm (red symbol).

Figure 4 Test angle co-conditioning. Teaching points: It is known that it is not the absolute level of discharge coming from muscle spindles of one muscle of the forearm that determines the forearm position signal, but the difference in signal between the two antagonists, the flexors, and extensors. Co-conditioning is a method of conditioning of the two antagonists so that both are left in a near identical state, leading to a low nett position signal coming from the arm. The blindfolded subject has their forearm placed at the test angle by the experimenter and they are asked to contract their elbow flexors by trying to pull the forearm toward the body and then their extensors by trying to push the arm away from the body while it is being held in position by the experimenter. Once the two contractions have been completed and the subject has relaxed their arm, its position is indicated with the other arm. The perceived position (perceived angle) lies close to the true position (actual).

Figure 5 Measuring position sense in matching and pointing tasks. Teaching points: (A) Matching task. Position sense at the forearm can be measured by the experimenter placing a blindfolded subject's arm (reference) at a test angle and the subject is asked to match its position by placement of their other (indicator) arm. This form of position matching is really a sensation-matching task, in which the subject moves their indicator arm until the sensations arising in it match those coming from the reference. Muscle spindles in muscles acting at the elbow are believed to provide the position signal. Because spindles exhibit thixotropic behavior, it is necessary to put both arms into a thixotropically defined state before such measurements are made. (B) Pointing task. Intuitively, we do not typically determine the position of one of our arms in space by mimicking its position with the other arm. Rather, we point to where we think the arm is located. This is a very different task to one involving sensation matching. In pointing no comparison of sensory signals between the arms is possible and the evidence suggests that in this form of position location muscle spindles are not involved. In the experimental arrangement, one arm (reference) is hidden from view behind a screen while the subject moves a paddle (pointer) to align its position with that of the reference.

Figure 6 Position errors in matching and pointing tasks. Teaching points: (A) Matching task. A plot of the distribution of errors when the position of one arm is indicated by placement of the blindfolded subject's other arm into a matching position (Figure 5A). To make sure that the muscle spindles believed to be responsible for this sense are in a comparable thixotropic state in elbow muscles of both arms, both are first flexion or extension conditioned (Figure 2). The prediction for flexion conditioning is small errors into flexion, for extension conditioning small errors into extension (Figure 3). The predicted errors (blue symbols) lie close to the actually measured errors (red symbols). In other words, the distribution of position errors in a matching task is consistent with predictions based on the properties of the position sensors, the muscle spindles. (B) Pointing task. Here position errors are measured in a pointing task where the subject aligns a pointer with the perceived position of their other arm, which remains hidden from view (Figure 5B). Assuming that in position sense measured in this way muscle spindles in elbow muscles of the hidden arm provide the position signal, after flexion conditioning (Figure 2A) significant errors into extension are expected. This distribution is different from that in A mentioned earlier because only one arm is involved. The observed errors (red symbol, left) are consistent with predictions (blue symbol, left). That, however, is not the case for extension conditioning of the arm (Figure 2B). Here the prediction is for large errors into flexion (blue symbol, right) and what is observed is large errors into extension (red symbol, right), the opposite direction to expectations. The conclusion from this experiment is that the distribution of position errors in a one-arm-pointing task no longer conforms to predictions based on the known behavior of muscle spindles. The broader conclusion is that muscle spindle signals do not contribute to limb position sense measured by pointing.

Figure 7 Teaching points: The diagram shows the way in which the experiments are conducted for the study of the perceived vertical distance between the index fingers in the vertical plane, the extent to which the subjects believe that they are holding their right index finger with their left hand (a measure of body ownership). The right forearm of the subject is positioned on the lower table so that the end of the index finger is held comfortably in a padded pipe, which can only rotate around the proximal interphalangeal joint. In one study, in which the left hand is not involved, the subject cannot see an artificial index finger which is part of the upper table and which can be rotated by the experimenter. Depending on whether the coupling between the upper and lower tables is fixed or not, the movement can be directly coupled so that the right index finger moves together with the artificial finger (congruent movement) or it can be uncoupled so that the motion of the two fingers occurs together and is incongruent. The vertical distance between the right index finger and the artificial finger is 12 cm. In another study, vision was excluded (see screen) and the experimenter lightly held the subject's left index finger and thumb against an object (actually the artificial finger). Congruent or incongruent movement was delivered to the right index finger and the artificial finger being held by the left index finger and thumb.

Figure 8 Teaching points: (A) The diagram shows subjective ratings on a 7-point scale of the subjects’ belief that they were holding their right index finger with their left hand under two conditions. All measures were made after the right index finger, and left index finger and thumb (which were touching the artificial finger) were locally anaesthetized. This meant that no information from the fingers could influence judgments. Proximal muscles in the hand and forearm were not affected by this procedure and information from them could be used by the brain. When movements of the index fingers on the two hands (and the artificial finger) were moved together in a congruent way, the subjects agreed that they were holding their right index finger with their left hand. They disagreed with this statement when the movements were made in an incongruent way. (B) This diagram shows the perceived spacing in the vertical plane between the left and right index fingers in the same study as in Panel A. Again, the right index finger and the left index finger and thumb were anaesthetized. When the movements of the index fingers on the two hands (and the artificial finger) were congruent, the subjects perceived that the vertical spacing between them was only approximately 2 cm. However, this perceived vertical distance was greater when the movements were incongruent (approximately 4 cm). Again, the true perceived distance between the index fingers was 12 cm. Thus, simply grasping the finger-like object without vision being available is sufficient to shrink the perceived distance between the left and right index fingers.

 


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Uwe Proske, Simon C. Gandevia. Kinesthetic Senses. Compr Physiol 2018, 8: 1157-1183. doi: 10.1002/cphy.c170036