Proprioception 1143
Proprioception
J L Taylor, Prince of Wales Medical Research Institute,
Sydney, NSW, Australia
ã 2009 Elsevier Ltd. All rights reserved.
Introduction
Bastian, in 1887, described ‘kinaesthesia’ as ‘‘the sense
of movement’’ whereby ‘‘we are made acquainted
with the position and movements of our limbs, we
are enabled to discriminate between different degrees
of ‘resistance’ and ‘weight,’ and by means of it the
brain also derives much unconscious guidance in the
performance of movements generally.’’ The term ‘pro-
prioception’ is used synonymously with kinesthesia.
It was first used by Sherrington in 1906 and can
be thought of as referring to the sensations of the
body. Although proprioception is a sense like vision,
hearing, touch, taste, and smell, these more commonly
recognized sensory systems give us information about
the outside world, whereas proprioceptors are the
sensory receptors which tell us what the body itself is
doing. As Bastian suggested, kinesthesia encompasses
a complex of sensations which include a sense of joint
movement, a sense of joint position, a sense of muscle
force or tension, and a sense of effort. Sensations
derived from the vestibular apparatus are also some-
times included in proprioception, but these signals of
the position and movement of the head relative to the
external world will not be discussed here.
The sense of joint movement (movement sense) is
the process by which we perceive movements of parts
of the body relative to one another. It includes detec-
tion of movements and perception of their direction,
velocity, distance, and timing. Both passive move-
ments, which are imposed by an external force, and
active movements, generated by the muscles, are per-
ceived. The sense of joint position (position sense) is
the process by which we perceive the current positions
of parts of the body relative to one another. Again,
relative positions can be perceived when the body parts
are at rest and when they are held in position by muscle
contraction. The sense of force or tension is the process
by which we perceive forces generated through the
muscles, whereas the sense of effort gives a perception
of the strength of muscle contraction relative to the
total strength of the muscle. Both of these signals are
used in perception of applied forces, and in judging
heaviness. All of these sensations are used separately
and together to give knowledge of the body’s own
actions and its interactions with the environment.
Thus, proprioceptive sensations provide feedback of
the consequences of motor output and are vital to the
control of voluntary movement.
Sensory Receptors Contributing to
Proprioception
Sensory receptors in the muscles, joints, and skin are
all involved in proprioception. In muscle, the major
receptors for proprioception are muscle spindles and
Golgi tendon organs. Muscle spindles are complex
receptors which lie in parallel with the muscle fibers.
They are fluid-filled fusiform (spindle-shaped) cap-
sules of connective tissue with bundles of small (intra-
fusal) muscle fibers inside them. On the intrafusal
fibers are two kinds of sensory nerve endings: the
spiral endings of the primary muscle spindle (Ia) affer-
ents and the flower-spray endings of the secondary
muscle spindle (II) afferents. Both classes of muscle
spindle endings fire with stretch of the muscle and so
give a signal of muscle length. The primary ending has
the more dynamic response. It fires a burst at the
beginning of a stretch and signals velocity and accel-
eration as well as muscle length. Muscle spindles also
have their own motor nerve supply to the intrafusal
muscle fibers. As contraction of the intrafusal fibers
stretches the sensory endings, muscle spindle firing
can result from motor output as well as from muscle
stretch. Thus, interpretation of muscle spindle signals
during active muscle contractions is complex.
Golgi tendon organs are situated at the musculo-
tendinous junction. They consist of several strands of
collagen attached to the tendon at one end and to a
number (3–25) of muscle fibers at the other end. The
strands of collagen are covered by a connective tissue
capsule and are innervated by a Ib sensory neuron.
Although tendon organs can fire in response to pas-
sive stretch of the whole muscle, they are much more
sensitive to active forces generated by the specific
muscle fibers that are attached to them.
In the joints, sensory endings are located in the joint
capsule and ligaments. Ruffini endings are slowly
adapting afferents which are sensitive to stretch of
the joint capsule. They fire most at the ends of joint
range, but a few afferents can fire across the midrange
of joint movement. Larger tendon-organlike recep-
tors are found in the ligaments. These also respond
to stretch but have a higher threshold compared to
the joint capsule receptors.
In the skin, the receptors most directly associated
with proprioception are the slowly adapting type II
(SAII) receptors. These fire in response to stretch of
the skin in specific directions. They can signal changes
1144 Proprioception

in joint angle when the skin is stretched over one side
of a joint and compressed on the other side.
Neural signals from all of these sensory receptors
are carried via large-diameter myelinated sensory
neurons with cell bodies in the dorsal root ganglia
and projections which enter the spinal cord through
the dorsal roots. Cutaneous afferents from both the
upper and the lower limbs ascend directly to the
medulla, where they synapse. Muscle afferents from
the upper limb follow the same route whereas those
from the lower limb have an intermediary synapse
in Clarke’s column in the spinal cord. From the med-
ulla, proprioceptive signals cross the midline to the
contralateral thalamus and from there project to
the cortex.
Sensations of Movement and Position
Position sense and movement sense give perceptions
about the relative locations and movements of parts
of the body. These can be perceptions of angle or
movement at a single joint (e.g., bend of the knee),
about many joints (e.g., perception of the distance
between the thumb and a fingertip), or even of posi-
tion or movement that does not involve joints (e.g., of
facial expressions or the direction of the eyes). The
sense of movement and sense of position can be diffi-
cult to separate, as movement is a change of position
and each position is reached through movement.
However, it is possible to impose movements that
are too slow to feel and yet, when displacement is
great enough, a new position is perceived. This sug-
gests that there is a sense of position separate from
movement.
Testing Movement Sense
The sensitivity of movement sense is often tested by
determining the smallest imposed movement that can
be detected with the muscles relaxed. The size of this
threshold movement depends on velocity, with faster
movements detected more easily than slower move-
ments (Figure 1). With velocities of >1 s 1, very small
movements of 0.1–0.5 can be detected at limb joints
and for rotation of the neck and trunk. With slower
velocities, larger movements of 1–3 are required.
With very slow velocities (2 min 1), study partici-
pants no longer report any sensation of movement.
At the interphalangeal joints of the fingers and toes,
much larger angular movements are needed for detec-
tion. For the big toe, detection thresholds of 20 at
1 s 1 are reported. It is not clear how active muscle
contraction changes the detection of imposed move-
ments, as both impairment and improvement have
been reported. The strength of contraction and the
task performed may be important.
Perception of the velocity of movement is another
facet of movement sense that has been tested. Dis-
crimination between velocities of imposed movement
about a joint is better for slower velocities. For move-
ments at 15 s 1, individuals perceive differences of
2–4 s 1, whereas for movements at 50 s 1, differ-
ences of 10 s 1 are detected. Individuals can actively
match the velocity of an imposed movement to within
5 s 1 over velocities from 15 to 80 s 1.
Testing Position Sense
Position sense also has a number of facets that can
be tested in different ways. As noted, movements
can be imposed so slowly that individuals do not

7
Shoulder
6
Elbow
Detection threshold (°)

5
MCP
4
DIP
3 Hip

2 Knee

Ankle
1
Toe
0
0.01 0.1 1.0 10 100
Angular velocity (° s−1)
Figure 1 The magnitudes of movement required for detection of movements of different velocities are plotted for joints of the upper limb –
shoulder, elbow, distal interphalangeal joint (DIP) and metacarpophalangeal joint (MCP) of the finger – and for joints of the lower limb – hip,
knee, ankle, and the interphalangeal joint of the big toe (toe). Movement detection is better at faster velocities. If velocities are slow enough
(0.1 s 1), detection is independent of velocity. Adapted from Refshauge KM, Chan R, Taylor JL, et al. (1995) Detection of movements
imposed on human hip, knee, ankle and toe joints. Journal of Physiology 488: 231–241, with permission.

perceive them. With these very slow displacements,
changes of position of 2–5 of the ankle, shoulder,
and finger are discriminated.
The ability to match joint positions is tested in a
variety of ways. The simplest is for the person to
move one limb to match the angle of a joint of the
contralateral limb which was moved passively and is
supported in position. Under such conditions, indivi-
duals can match joint angle within 5–10 . Some var-
iants on this test include repositioning the same limb
rather than matching with the other limb, using a
pointer or indicator rather than using a matching
limb, and giving a verbal estimate of angular position.
Other variations focus on changes in the limb being
matched – for example, varying the time between
arrival at the position and matching, having the per-
son move actively to position, or hold the position
actively. When muscle activity is used to maintain
position, accuracy is improved with errors of 2 .
Another aspect of position sense is the ability to
perceive the instantaneous position of the arm during
an ongoing movement. During an imposed movement
at the elbow at velocities of 70–200 s 1, individuals
can indicate with an accuracy of 1 when the arm
passes through a particular joint angle. This contrasts
with the poorer performance in detection of impercep-
tibly slow movements, and in position-matching tasks.
Evidence for the Mechanisms Underlying
Movement and Position Sense
It is generally accepted that sensory inputs from mus-
cle, skin, and joints all contribute to movement and
position sense, although belief in the relative impor-
tance of each contribution has varied over the past
century. The evidence that supports the contribution
from each type of afferent includes the findings that
(1) the afferents fire in response to joint movement or
joint position, (2) activation of the afferents leads to
the perception of joint movement or altered joint
position, and (3) elimination of the firing of the affer-
ents impairs the ability to perceive joint movement or
position.
Joint receptors The activity of joint afferents has
been recorded from awake people through micro-
neurography, in which a fine needle can record the
activity from single nerve fibers. In general, the affer-
ents tend to fire at the ends of joint range and do not
give an unambiguous signal of joint position or the
direction of movement. However, occasional affer-
ents do fire throughout the whole range in a way
related to position, and this may be sufficient to give
a useful signal. With microneurography it is possible
to stimulate a single nerve fiber electrically. For
30% of joint afferents, this stimulation is perceived
Proprioception 1145
as small joint movements. With other afferents, indi-
viduals report pressure in the joint. When joint affer-
ents are not active, either during testing after total
joint replacements or with injection of local anes-
thetic into the joint, there is little impairment. Thus,
although joint afferents contribute to perception
of joint movement and position, other inputs can
compensate for their loss.
Skin receptors Recording from skin afferents shows
that the slowly adapting afferents which respond to
stretch of the skin can fire with changes in joint angle.
For example, afferents with receptive fields over the
back of the hand increase firing with flexion of the
metacarpophalangeal joints. Thus, these afferents can
signal joint movement and position. Activation of
skin afferents can produce perceptions of movement
of the fingers and limb joints. If the skin is stretched
on one side of a joint by pulling on tape stuck to the
skin, many people report feeling movements about
the joint. The illusion is enhanced if the skin on the
other side of the joint is compressed at the same time.
When input from the skin is removed by local anes-
thetic, perceptions of movement and position are
impaired but not abolished at finger joints. However,
impairment does not occur with anesthesia of a sleeve
of skin around the knee. Thus, sensory input from the
skin contributes to movement and position sense. It
may be easier to compensate for its loss at the larger
limb joints than in the hand.
Muscle spindles The muscle afferents associated
with movement and position sense are those from
the muscle spindles. These fire when the muscle spin-
dle is stretched during lengthening of the muscle.
Recording in humans and in animals shows that mus-
cle spindles can signal joint movement and position
during passive movements. The Ia afferents signal
velocity (with a position component) whereas the
group II afferents primarily signal position. Muscle
spindle activity increases with muscle lengthening,
but most spindles fall silent during passive shortening
of the muscle. Thus, muscles on both sides of a joint
are needed to give adequate signals of movement and
position. Stimulation of a single muscle spindle affer-
ent causes no sensation, but when a population of
muscle spindles is activated by vibration applied
over the tendon of the muscle, joint movement and
altered joint position are perceived. Illusory move-
ments are felt in the direction consistent with length-
ening of the vibrated muscle. Higher frequency
vibration (100 Hz) generates a strong illusion of
movement with a velocity related to the frequency
of vibration. This illusion is attributed to discharge
of Ia afferents. With lower frequency vibration, the
1146 Proprioception
movement illusion is diminished but an illusion of
altered position remains. This is attributed to group II
afferent activity. The anatomy of muscle attachments
to the phalanges allows the muscles to be disengaged
functionally from the distal interphalangeal joint of
the middle finger by positioning the hand with the
middle finger fully flexed at the proximal inter-
phalangeal joint and all the other fingers extended.
Because no muscles act at the distal interphalangeal
joint in this posture, movement and position sense can
be tested without muscle spindle activity. The absence
of muscle spindle firing impairs, but does not abolish,
detection of joint movement and position.
During active movements, signals from joint and
skin afferents are the same as during passive move-
ments. Muscle spindle activity in the antagonist (non-
contracting) muscle should also be unchanged. In
contrast, the firing of muscle spindles in the active
muscle depends on fusimotor drive, in addition to
changes in muscle length. In humans, fusimotor
drive to the muscle fibers inside the muscle spindles
usually occurs concurrently with voluntary activation
of the muscle. Therefore, during an isometric contrac-
tion, spindle firing increases from fusimotor drive
even though the muscle is not stretched. During a
shortening contraction, spindles will keep firing if
the shortening is slow, but will fall silent if the short-
ening velocity of the muscle exceeds the ability of the
intrafusal fibers to contract quickly. During a length-
ening contraction, both fusimotor drive and muscle
stretch will add to spindle discharge. It has been
postulated that a signal related to fusimotor output
must be used by the brain to interpret muscle spindle
input in terms of joint movement and position during
voluntary contraction.
Corollary discharge A final mechanism that may
give rise to sensations of movement or position is a
central signal related to motor output. Such a signal is
known as a corollary discharge. This idea is well
established for the eyes. Because the muscles of the
eyes always have to move the same load, a set amount
of drive to the muscles should always result in the
same eye movement. That is, you should know which
way your eyes are pointing because you told them to
go there. Individuals’ reports that the visual world
moves when passive movements are imposed on the
eyes are consistent with this proposal. In contrast,
there is no fixed relationship between motor com-
mand and the resultant movement or position for
the limb muscles, because they work under ever-
changing loads. However, a recent study, in which
individuals experienced illusions of altered joint
position when they tried to move a paralyzed and
anesthetized hand, suggests that a signal related to
motor command may also have a role in position
sense in the limbs.
Sensations of Force and Effort
The sense of muscle force or tension and the sense of
effort are difficult to separate. One provides a percep-
tion of the absolute force being generated by the
muscle. The other gives a perception of the propor-
tion of total muscle strength being used. Both contrib-
ute to judgments of applied forces and of heaviness.
The mechanisms underlying the sensations differ. The
sense of muscle force is attributed to sensory input
whereas the sense of effort is attributed to a central
signal related to motor command.
Testing Force and Effort
Sensations of force are often tested by asking persons
to make judgments about the weights of objects. The
heavier an object, the more muscle force is required to
move or lift it. People can discriminate between
weights which differ by about 6%. This ability is
independent of the muscle group which is used to
support the weight. When individuals give numerical
estimates of the magnitude of force or weight, per-
ceived force doubles if exerted force is increased by
50%. However, if they lift a weight with one muscle
group and then choose a matching weight with the
contralateral muscles, the relationship between
the two weights is linear, although lower weights
(<40% maximal force) are overestimated and higher
ones (>80% maximal force) are underestimated. The
linearity of weight-matching suggests that the rela-
tionships between force production and perception
are similar in the two muscle groups. People can
match isometric forces in the same way as weights,
with an average error of 4% of maximal force.
People can also scale forces based on their own effort.
When individuals produce designated proportions of
maximum effort, they produce more force than
expected for weak efforts and less for strong efforts.
This is consistent with the overestimation and under-
estimation in matching light and heavy weights.
Evidence for the Mechanisms Underlying
Sensations of Force and Effort
The main sensory inputs that underlie the sense of
muscle force are the Golgi tendon organs. Because
each tendon organ is activated by the contraction of
only a few muscle fibers, the firing of single tendon
organs does not represent the force generated by
the whole muscle. In contrast, the response from a
population of tendon organs is a good measure of
overall force, although a dynamic component to the

firing accentuates changes in force. It is not clear
whether stimulation of tendon organs gives a percep-
tion of muscle force, although people do experience
sensations of muscle force when muscle contractions
are produced by electrical stimulation.
Sensory signals of muscle force and central signals
of effort are both directly related to the force gener-
ated by a muscle. However, when a muscle becomes
fatigued, it cannot produce as much force, and sen-
sations of muscle force and effort are dissociated. It
is a common experience to hold an object which gets
heavier and heavier as the muscles holding it get
tired. The increase in perceived heaviness despite
the object’s constant weight is an indication that
the sensation is not conveyed by the tendon organs,
which continue to give a true signal of muscle force.
In this situation, progressive fatigue of the muscle
fibers means that more of the muscle needs to be
recruited to support the object. That is, the motor
drive or command to the muscle is increased, and
the sense of effort, which is related to the proportion
of the muscle that is being used, also increases.
Thus, the increase in heaviness of objects during
fatigue suggests that the sense of effort has a large
influence on perception of weight and force. This is
confirmed when individuals match weights lifted
by two muscle groups of different strength. Heavier
weights lifted with the stronger muscles are matched
to lighter weights lifted with the weaker muscles
(Figure 2). However, the sense of effort is not
the sole contributor to perceptions of heaviness.
Although a weight supported by a fatigued muscle
feels heavier, the increase in heaviness is not directly
proportional to the reduction in force-generating
capacity of the muscle. That is, if the maximal
force of the muscle is reduced by half, the weight
does not feel twice as heavy. This suggests that some
other signal, presumably sensory input from tendon
organs, also has a role. Similar results are reported
when individuals’ muscles are weakened by partial
curarization.
Finally, in most judgments of force, tactile informa-
tion is also important. Pressure-sensitive receptors are
not truly proprioceptors, because they give information
about the body’s interaction with the environment
rather than conveying sensations about the body itself.
However, skin receptors can signal the magnitude and
direction of forces. When muscle receptors and central
signals of effort cannot contribute, such as when
objects are placed on a hand which is resting on a
table, people can discriminate changes in weight of
30%. Although discrimination is improved when
the object is supported actively, the elimination of tac-
tile input alters force sensations even during active
conditions.
Proprioception 1147
16
Matching force (N)
12
Hand
8
4 Finger
0
0 2 4 6 8 10 12
a Reference force produced by elbow (N)
20
Matching force (% MVC)
15
Hand
Finger
10
5
0
0 2 4 6 8 10
b Reference force produced by elbow (% MVC)
Figure 2 Individuals matched forces produced by elbow flexion
▪
(reference) with forces produced by a finger ( ) or by a pinch grip
of the hand (○). (a) The same force produced by the elbow was
matched with a lower force produced by the finger than by the
hand. As the finger could produce a lower maximal force, com-
pared to the hand, the difference in matching is consistent with
matching effort. (b) The same data as in panel (a) are shown
expressed as percentages of maximal voluntary force (MVC). For a
force produced by the elbow, individuals matched the same propor-
tion of muscle force for the finger and hand. Note that these weak
forces produced by the elbow (2–8% MVC) were overestimated by
both the finger and the hand, which matched with 5–15% MVC.
Reproduced from Experimental Brain Research, Vol. 151, 2003,
197–203, Perceptual constancy and the perceived magnitude of
muscle forces, Jones LA, figures 2 and 3, Copyright Springer. With
kind permission of Springer Science and Business Media.
Interactions of Senses
Some sensations require interactions between the pro-
prioceptive senses. For example, heaviness of an object
can only be gauged if the object can be moved. Move-
ment signals the time when the perceived force is a
measure of the object’s heaviness. The perceptions of
qualities such as stiffness, which is a combination
of force and distance, and viscosity, which combines
force and velocity, need continuous interaction of sen-
sations of movement and force. The senses of move-
ment, position, force, and effort can all contribute to
the perception of limb position under different circum-
stances. If the limb is resting passively, only position
sense is available to give a perception of joint angle.
In contrast, if the limb is supported against gravity
by active muscle contraction, then the muscle force
1148 Proprioception
needed to hold the limb in position will also vary with
joint angle. Thus, the sense of effort associated with
the level of muscle contraction, as well as signals of
muscle force from tendon organs, will be able to con-
tribute (Figure 3). Finally, if the joint position was
reached by a movement, then movement sense may
also have a role.
Interactions between proprioception and other
senses provide orientation in the environment. Per-
ception of where the body is in space relies on the
sensations of movement and position of the neck,
trunk, and limbs, combined with sensory input from
the vestibular apparatus, which signals movements of
the head in space. If the head rotates 90 to the right,
it might mean that the neck has turned, that the trunk
has turned or that the whole body has turned. Sensa-
tions of movement and position from the neck and
trunk allow us to differentiate these conditions which
give similar vestibular signals. Sensations of position
and movement of the eyes are essential for the per-
ception of the location of visual objects relative to the
head, with proprioceptive input from the neck needed
to identify the location relative to the body. Similarly,
the location of sounds relative to the body requires
sensations of movement and position of the head on
the neck.
Corollary discharge related
to motor command
Golgi tendon
organs (Ib)
Biceps
Joint receptors
Triceps
Skin receptors
Muscle spindle primary endings (Ia)
Secondary endings (II)
Figure 3 Diagram showing the proprioceptive signals available
to contribute to the perception of elbow angle (limb position) while
the forearm is supported against gravity by active contraction
of biceps brachii. Sensory signals are available from muscle spin-
dles and tendon organs in the agonist (biceps) and antagonist
(triceps) muscles. Sensory input is also available from joint recep-
tors and from skin receptors on both sides of the joint. A central
signal related to the motor output (corollary discharge) is also
available.
The contribution of proprioceptive sensations
to awareness of the size and shape of the body is
demonstrated by illusions that can be induced by
tendon vibration. If vibration is applied to the biceps
brachii tendon while a person touches the nose with
the finger, the nose feels like it grows. As discussed
previously, the vibration activates muscle spindles,
which give the illusion that the arm is extending at
the elbow (consistent with biceps lengthening). How-
ever, tactile sensations insist that the finger is still
touching the nose. The brain’s resolution to this
dilemma is that the nose is getting longer. Thus, pro-
prioceptive signals influence the body image.
Changes in Proprioception with
Pathology
Proprioceptive function changes with normal events
such as fatigue as well as with injury and disease. As
discussed, muscle fatigue alters judgment of force and
heaviness. It can also impair the detection of imposed
movement and judgment of limb position when the
limb is actively held against gravity.
Musculoskeletal injuries such as ankle sprains and
knee ligament injuries are associated with impaired
movement and position sense at the injured joint.
Similarly, impaired proprioception is reported with
arthritis and low back pain. Disturbance of proprio-
ceptive input from the neck may lead to the dizziness
and impaired balance in patients with whiplash. The
mechanism of such impairments is not clear. Possibi-
lities include changes in the periphery, including
inflammation or altered tissue compliance, which
could change receptor firing, and more central effects,
such as interaction between pain and proprioceptive
sensations.
Interruption of the neural pathway from the sensory
receptors to the brain will also result in impairment of
proprioception. This could happen in the periphery
through neural injury or neuropathy, in the spinal
cord with injury or diseases such as multiple sclerosis,
and in the brain stem or at the cortex through injury or
stroke. Very rarely, patients who have lost all large
myelinated nerve fibers have no sensory input to pro-
vide proprioceptive signals and must rely on central
signals related to muscle contraction and on vision to
perform motor tasks successfully. They are unable
to learn new motor skills.
Apart from lesions in the proprioceptive pathway
to the cortex, movement disorders (such as Parkin-
son’s disease, focal dystonia, Huntington’s chorea)
and cerebellar disorders have associated impairments
of various proprioceptive sensations. For example,
detection of passive movements and joint position is

impaired in Parkinson’s disease, whereas in cerebellar
disorders, the discrimination of velocity and duration
of movements is impaired.
Contribution of Proprioception to
Motor Control
Proprioception and motor control are so entwined
that it is often impossible to separate where the sen-
sory signals subserve a direct motor function
and where the same signals evoke proprioceptive
sensations which might then be used in control of
movement. Proprioceptive signals contribute to move-
ment at every level. Proprioceptive afferents have roles
in reflex responses at the spinal and cortical levels and
provide feedback that allows the control of all pur-
poseful movements. We can attend to some of this
feedback and report sensations related to it, but we
more commonly employ it to guide our actions.
Although we usually pay no attention to the current
position of a hand before we use it to pick something
up, we could, if asked, report the hand’s location.
Proprioception is essential to motor learning. New
patterns of movement, such as driving a car or hitting
a ball, cannot be learned in its absence. Without pro-
prioceptive sensations there is no direct link between
the motor output and its consequences.
Summary and Conclusions
Proprioception is the sense which allows perception
of the location, movement, and action of parts of the
body relative to each other. Proprioceptive sensations
are derived from combined sensory input from muscle,
joint, and skin receptors, plus central signals related
to motor output. Sensations of movement and position
and force and effort can be tested separately, but are
used in conjunction with each other during many per-
ceptual and motor tasks. Proprioception is so closely
tied to the control of voluntary movement that im-
pairments are more likely to be noticed as deficits
in the performance of motor tasks than as impaired
sensations.
See also: Evolution of Sensory Receptor Specializations in
the Glabrous Skin; Mechanoreceptors; Motor Skill
Learning; Multisensory Convergence and Integration;
Proprioception 1149
Neural Circuitry in the Somatosensory System; Neuro-
muscular Junction (NMJ): Presynaptic Stretch Effects on
Neuromuscular Transmission; Spatial Orientation: Our
Whole-Body Motion and Orientation Sense; Tactile
Texture.
Further Reading
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ensemble cutaneous input from the dorsum of the human hand.
Journal of Physiology 496: 857–871.
Cordo P, Carlton L, Bevan L, et al. (1994) Proprioceptive coordi-
nation of movement sequences: Role of velocity and position
information. Journal of Neurophysiology 71: 1848–1861.
Donaldson IM (2000) The functions of the proprioceptors of
the eye muscles. Philosophical Transactions of the Royal
Society of London, Series B. Biological Sciences 355(1404):
1685–1754.
Edin BB and Abbs JH (1991) Finger movement responses of cuta-
neous mechanoreceptors in the dorsal skin of the human hand.
Journal of Neurophysiology 65: 657–670.
Gandevia SC (1996) Kinesthesia: Roles for afferent signals and
motor commands. In: Rowell LB and Shepherd JT (eds.) Hand-
book on Integration of Motor, Circulatory, Respiratory and
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